reviewer acknowledgement http://www.koedoe.co.za open access koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping the individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely, productive peer review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: page 1 of 1 in an effort to facilitate the selection of appropriate peer reviewers for koedoe, we ask that you take a moment to update your electronic portfolio at www.koedoe.co.za for our records. this will allow us better access to your areas of interest and expertise, in order to match reviewers with submitted manuscripts. if you would like to become a reviewer, please visit the journal website and register as a reviewer. to access your details on the website, you will need to follow these steps: 1. log into the online journal at http://www. koedoe.co.za 2. in your ‘user home’ [http://www.koedoe. co.za/index.php/koedoe/ user] select ‘edit my profile’ under the heading ‘my account’ and insert all relevant details, bio statement and reviewing interest. 3. it is good practice as a reviewer to update your personal details regularly, to ensure contact with you throughout your professional term as reviewer to koedoe. please do not hesitate to contact me if you require assistance in performing this task. margo martens submissions@koedoe.co.za tel: +27 21 975 2602 fax: +27 21 975 4635 koedoe african protected area conservation and science anneli douglas antoinette p. malan brian kuhn bruce h. brockett carla staver cleo gosling clinton carbutt dave balfour david le maitre edwin muchapondwa gabriela bucini george bredenkamp ian a. russell jens oldeland jesse m. kalwij johan van tol john o’brien joop schaminée juergen dengler keenan stears laura k. blamey lucas enrico matthew mcconnachie megan griffiths navashni govender nicola stevens norbert hahn olaf weyl osborn v. ferrel pieter bester richard stirzaker rina c. grant robert buitenwerf robert guldemond sam m. ferreira shaun levick tammy robinson tercia strydom theo h.c. mostert ute schmiedel vaughan spaull yolanda pretorius 94 filelist convert a pdf file! koedoe 19: 175-176 (1976) two new bird records for the tsitsikama national parks g . a. robinson tsitsikama coastal national park p. o. stormsrivier 6)08 a comprehensive check-list of the birds that possibly could occur in the vicinity of the parks was published by skead and liversidge (1967). they refer to 210 bird species but only record 205; three species were duplicated in part 1 (forest habitats) and part 2 (sea habitats). it would appear that two species were omitted from part 1. however, one of the species recently recorded should be included in part 1 while the other belongs to part 2. on 1973.07.03 a lesser gallinule porphyrio alieni, (roberts number 209) was observed outside the boathouse at storms river mouth. this bird was captured , fed to a healthy state and eventually released at nature's valley. prior to the sighting of this bird the weather had been foul with strong south-westerly winds and rough seas and rain. the bird was in a poor state of health weak and undernourished . as this species selects marshy reeded areas to hive in and no such area exists in the immediate vicinity of storms river mouth, it was surmised that the bird had been blown in from one of these habitats west of storms river. nature's valley, nearly 40 km to the western boundary seemed to be a likely area that would attract gallinules. identification was made by comparing the dimensions of the wing, tarsus, culmen, shield width and tail length to those of the purple, american and lesser gallinules as presented by mclachlan and liversidge (1970). these are listed in table 1. more recently on 1975.07.15 another lesser gallinule was collected at storms river mouth. this bird was found shortly after the same adverse weather conditions had prevailed as the first, i.e. stormy southwesterly winds accompanied by heavy rain. the bird was a juvenile and in a weak and hungry condition, and it was therefore assumed that the bird probably could not have flown. because the storms river was in flood during the period of both sightings both birds could have come down the river which is marshy and has reeds upstream. on the 1973 .07. 10 a sea-bird in poor condition, which later died, was found on the beach at nature's valley. professor j. m. winterbottom (pen . comm.), identified the bird as a blue petrel halobaena caerlea (roberts number 20). its measurements are given in table 1. these two additional records brings the total of birds species for the parks to 207. 175 table 1 comparative measurements of gallinules and blue petrels gallinules blue petrel measurements storms river purple' american lesser .... nature's blue peu-e!· specimen purple' valley specimen wing 153mm 225-248 mm 162-181 mm 144-160mm 217mm 205-221 mm tarsus 44mm 70--92 mm 58--jj2 mm 45-56 mm 32mm 39-33 mm cu lmen 37 mm 56--71 mm 38-48 mm 35-40 mm 25mm 25-28 mm shield width ilmm 18-24 mm tail length 65mm 82-101 mm 64-mm 66--73 mm 104mm 77-91 mm r.: measurements from roberts references mclachlan, g. r. and r. liversidge. 1970. roberts birds of south africa. cape town: trustees of the s.a. bird book fund. skead, c.j. and r. liversidge. 1967. birds of the tsitsikama forest and coastal national park, 1966. koedoe 10:43-62. 176 page 1 page 2 book information book title: environmental risk management in south africa book cover author: mike mentis isbn: 9780620507523 publisher: publisher.co.za; 2010, r228.00* *book price at time of review book review information reviewer title: review of environmental risk management in south africa reviewer: harry biggs1 affiliation: 1sanparks, skukuza, south africa email: biggs@sanparks.org postal address: private bag x402, skukuza 1350, south africa how to cite this article: biggs, h., 2011, ‘review of environmental risk management in south africa’, koedoe 53(1), art. #1065, 1 page. doi:10.4102/koedoe.v53i1.1065 copyright notice: ©2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) review of environmental risk management in south africa congratulations to mike mentis on producing such a concise, clear and powerful guide and critique! i would recommend this book to any environmental manager or consultant who is seeking something beyond the largely once-off and routinised processes in use in south africa for integrated environmental management (iem) especially if they have an existing or potential interest in a more adaptive, risk-based approach. i would also hope that open-minded officials responsible for the legislation he criticises, would give the arguments put forward in the book a fair hearing. for active practitioners, the book is useful because it uses ‘risk’ as the central notion to improve on what was probably all along intended by the word ‘impact’. this is convincingly justified in the course of the book. many practitioners will have heard of, or even used some components of risk-based techniques. however, i believe that the steps and techniques that he clearly lays out, provide the necessary bridge from a more conventional eia to a much improved eia with a central emphasis on adaptation and risk analysis. these approaches can also be used in wider environmental contexts. in fact i wondered whether we are not eventually getting to the point that so-called ‘ecosystem management’ will at last, in actual practice, be seen in a stronger continuum with ‘environmental management’, the latter usually understood in the sense of dealing with consequences of ‘developments’. i always found it curious that ‘ecosystem management’ textbooks (and now slowly, ecosystem management practice, at least in some settings) shows far more uptake of the adaptive ethos. regarding the latter, he makes the point over and over again (rising to a crescendo in the last chapter) that the law’s prescriptions, although not excluding this, do very little to encourage a learning approach through iterations – the vast majority of eias and environmental management plans produce close to static blueprints. he likens this, amongst other analogies, to a general heading out to war with a full and irrevocable plan written out of his units’ movements, intentions and assumed outcomes for the next several years! a semi-independent chapter on decision-making deals with several complementary ideas which add further value to the whole risk theme, but which, again, can be used in many other settings too. these useful principles are not restricted to iem, though most of his examples are. regarding the author’s critique of the existing de facto practice of iem in south africa as an outflow of the law, the author builds a solid interconnected argument (which i won’t spoil for the reader) that a serious re-think is necessary and overdue. this includes the paranoia of omission – the underlying fear of not covering every possible risk in an eia. if the risks are sensibly prioritised by analysis (one subject of the book) and action restricted, at least initially, to those that have the best chance of delivering, then achieving clearly set environmental goals actually becomes possible. what really locks us into the ‘once-off blueprint mode’ is the fact that the environmental departments of government, who in good but misguided faith want to control everything centrally, cannot even deal with the ‘once-offs’, let alone the updates which are actually essential to make the system work properly. he suggests several possible changes to the way the country deals with this, including placing the onus on developers to actually buy into and defend their reputations based on the real underlying fundamentals of the constitution (the right to an environment not harmful to health, free from pollution and degradation) rather than seeing that they as quickly or easily as possible pass the ‘tick-box’ test of the current eia requirements. the ultimate changes he suggests need to be radical, but he suggests these could be phased in for workability. except in a few very small ways, the 2010 regulation changes (which appeared just after his book came out) under the south african national environmental management act, do not move in the directions he proposes. so all those with an interest in robust debate on this topic – this book is also for you! there are a few minor glitches (some semi-estranged footnotes and some typos/miscalculations in a key decision-tree graphic) but nothing serious enough to do much more than keep the reader on her or his feet. many messages are repeated over and over again, and the final chapter reads a bit like a sermon. but in my opinion he is close to spot on, and i hope we see practitioners using more of this approach, and ultimately that we see a revamp in south africa (he does not discuss whether other countries are any better – i fear not – and in which case it is indeed another chance for south africa to show some leadership). readers who don’t agree with him will grow by at least facing the suggestions and challenges articulated in this highly readable book with its clear examples. my suggestion is to either consider adopting this line of thinking, or to produce something better, but not to let it leave one cold. book information title: the limits to scarcity: contesting the politics of allocation book cover: book cover the limits to scarcity: contesting the politics of allocation editor: lyla mehta isbn: 978-84407-542-3 publisher: earthscan: london and washington dc; 2010, r400* *book price at time of review review title: scarcity in the time of over-consumption reviewer: wendy annecke1affiliation: 1cape research centre, south african national parks, south africa postal address: po box 216, steenberg 7947, cape town, south africa how to cite this article: annecke, w., 2012, ‘scarcity in the time of over-consumption’, koedoe 54(1), art. #1089, 2 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1089 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. scarcity in the time of over-consumption in this book review... open access resource use and the sustainability of resources constitute a significant concern for many scientists and field staff in south african national parks (sanparks). according to its mission ‘to develop, manage and promote a system of national parks that represents biodiversity and heritage assets by applying best practice, environmental justice, benefit sharing and sustainable use’, sanparks is committed to sharing the benefits of the areas it protects with the wider society. this wider society, in immediate terms, primarily incorporates tourists and the communities living on park borders. tourists require access to resources such as roads, water, energy, food and waste facilities, whereas people living on the boundaries of the parks require access to grazing, medicinal and food plants, wood fuel and bush-meat. consequently, some of the biodiversity resources that sanparks was established to protect, are under severe pressure from ‘the wider society’ in general. for example many medicinal plants are threatened by over harvesting (both legal and illegal), whilst other resources, such as water, are threatened by pollution, changes in land use and increased demand for these. this book will be of interest to conservationists who are concerned with the ethics and sustainability of resource use, as well as the status of biodiversity health in the parks. the limits to scarcity does not provide answers to the difficult issues arising with regard to resource use, but it does provide a different way of thinking about scarcity and why this occurs. in doing so it raises questions about who the primary resource users are, and encourages a shift in perspective about the rights of all users. the limits to scarcity is compiled from of a series of papers delivered at a conference on ‘scarcity and the politics of allocation’ held at the institute of development studies, at the university of sussex, in june 2005. the fourteen chapters are divided into three parts with each part preceded by an introduction and commentary by the editor, leyla mehta. the guidance she provides is useful for novice readers in the field of scarcity or for readers with limited time. each chapter explores the notion of scarcity from a different angle and includes how the notion of scarcity is constructed and whose purposes it serves. the three chapters in part i discuss why scarcity matters, part ii explores the different perspectives of scarcity within economics and part iii investigates the politics of scarcity in case studies of water, energy and food supplies. authors include well known nicholas xenos, fred luks and ben fine. the latter is better known to most south africans for his and zavareh rustomjee’s seminal work on the minerals-energy-complex. the limits to scarcity makes an important contribution to resource use debates, particularly in the light of the rampant consumer culture which is seen by most people as the only way out of the on-going global economic woes. chapter by chapter the book unpacks the dynamics of how resources are allocated in various societies: who has access to what resources, and under what circumstances. the authors are careful to acknowledge the ‘biophysical realities of falling groundwater levels, melting icecaps and declining soil fertility’ (mehta 6) whilst they dissect the politics of need and those involved in the access and allocation of resources. the book has chapters of interest to economists (such as that by fine), engineers (such as that by lankford) and to philosophers and non-economists (xenos). for those interested in social justice and equitable societies as well as those involved in development and the poverty industry, there is plenty of significant content over which to ponder. what is most disturbing about this book is the evidence demonstrating how all-pervasive neo-malthusian economics has become, even in ‘development thinking’. in the first chapter, mehta provides a historical overview about the legacy of and approaches to normalizing the ‘scarcity postulate’. this overview sets the tone for the rest of the book by demonstrating that scarcity is not necessarily a natural or universal concept. she offers the chapters that follow as evidence for her argument as well as her challenge to the inertia evident in much mainstream thinking and economics. the ensuing chapters provide ample evidence that there is enough water, energy and food for everyone in the world (fao 2000 in hilyard 2010:152). one has only to visit the landfill sites and the dump pickers industry in south africa to observe the magnitude of waste generated by society, but, as raynor (p. xviii) argues, scarcity has become a rationale for the inequitable allocation of resources: it constitutes a gate-keeping mechanism. fine (p. 82) follows this up by asking uncomfortable questions about the manner in which a society chooses to rank one person’s welfare over another; their criteria for selecting one group of users who are able to access resources easily and use them intensely, whilst another group may have only minimal access to the same resources. i would suggest that feminists have become adept at conducting analyses that evaluate access to, control over and benefits from resources, and that feminist literature could contribute to this debate and perhaps shift its dynamics. in dipak gyawali and ajaya dixit’s chapter 13, scarcity is highlighted as a modern concept driven by technological choices to meet specific socio-political needs (p. 234). the authors quote helena norberg-hodge who worked in laddakh when there was no word for ‘poor’ in that region, but on returning some ten years later, they found her former translator pleading poverty and that his country desired a western notion of development. he and his countrymen (and women) had fallen victim to the notion of wealth that emanates from material goods. there are chapters on soil fertility in africa, agriculture and hunger, and two chapters for those involved in the water sector and the shared responses to its scarcity. the editor, mehta, has herself worked extensively on issues around water scarcity and is well known in this field. bruce lankford argues, in chapter 11, that water scarcity should not only be understood as ‘volumetric imbalance to be dealt with by saving, storing and delivering more water’ but suggests a framework for supply, demand and shared responses. he highlights how the natural and social scarcity of water can be exacerbated by people and institutions that are not designed to react to the rapidly occurring variations in water scarcity, in time and space, and provides a table comparing approaches for managing the dynamic supply of water in sub-catchment areas. from this he draws a number of conclusions, that provide persuasive arguments for the adoption of his composite and complex framework. in chapter 12, jasveen jairath re-examines the meaning and explanations of water scarcity and critiques what he calls the ‘conceptual weakness and politics’ of integrated water resource management. he highlights the importance of historical conditions, suggests the kind of questions that should be asked when bringing the weak and the strong together at a table, and points out that the democratic governance of water is seldom achieved. whilst the concept of scarcity itself is thoroughly unpacked, the book is weak on recommendations about how mainstream and pervasive thinking might be changed from valuing over-consumption to valuing sufficiency. xenos argues that historically the time for appealing to the morals and ethics of a society appears to be over. such appeals did not, for example, win the friends and support that president carter had hoped for in 1979. likewise, in south africa, appealing to the adherence to ex-president mandela’s values has worn thin. the question is what could provide the impetus for change? whilst none of the authors define the term ‘enough’, some point to individuals and communities that have described it for themselves: the amish and po chi-i are mentioned in this regard. i particularly like the words of po chi-i, the retired mandarin who wrote poetry, from about 850 ad, (in thompson 2010): what i shall need are a very few things. a single rug to warm me through the winter; one meal to last me the whole day. it does not matter that my house is rather small; one cannot sleep in more than one room! it does not matter that i have not many horses; one cannot ride two horses at once! (p. 131) there are typing errors throughout the book including the sub-title of a book in the science in society series: participation and exalusion in nuclear decision-making. mehta herself thanks the science and society programme – this name is an unlikely change from the series name and there are various other errors that should have been noticed and corrected. nonetheless, these issues should not detract from a provocative and worthwhile book. article information authors: mariette marais1 antoinette swart1 affiliations: 1agricultural research council, plant protection research institute, south africa correspondence to: mariette marais postal address: private bag x134, queenswood 0121, south africa dates: received: 11 dec. 2013 accepted: 27 aug. 2014 published: 17 nov. 2014 how to cite this article: marais, m. & swart, a., 2014, ‘plant nematodes in south africa. 12. checklist of plant nematodes of the protected areas of the eastern cape province’, koedoe 56(1), art. #1220, 3 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1220 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v56i1.1220-1. copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. plant nematodes in south africa. 12. checklist of plant nematodes of the protected areas of the eastern cape province in this checklist... open access • abstract • introduction • materials and methods • results and discussion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ soil-inhabiting nematodes, including plant-parasitic nematodes, are considered to be the most abundant multicellular organisms in the soil, and of particular interest since they are an integral part of the interlocking chain of nutrient conversions. because of their abundance and relative susceptibility to both physical and chemical changes, these organisms are used as indicator organisms. the national collection of nematodes (ncn) consists of a core collection, the meloidogyne collection and the juan heyns collection, which are housed at the plant protection research institute of the agricultural research council in pretoria. vast amounts of biodiversity data are contained in ncn, and the digitising of the collection from 2007 to 2014 yielded unpublished locality information, especially datasets of plant nematodes reported from protected areas of the eastern cape. two hundred and thirty plant nematode species belonging to 36 genera were reported from the eastern cape. of these, only 80 were from protected areas, whilst 163 were from uncultivated areas (outside protected areas) and 148 from cultivated areas. ten species were described from protected areas, namely criconemoides silvicola, meloinema silvicola, ogma tuberculatum, paralongidorus cebensis, paralongidorus hanliae, scutellonema tsitsikamense, trichodorus vandenbergae, xiphinema erriae, xiphinema ornatizulu and xiphinema simplex. only m. silvicola, o. tuberculatum, p. cebensis and s. tsitsikamense were not reported from other provinces, suggesting endemism.conservation implications: the diversity of nematode fauna is not adequately protected as most nematode biodiversity in the eastern cape lies outside protected areas, with only 80 of the 230 plant-feeding nematode species in the province being reported from protected areas. introduction top ↑ the national collection of nematodes (ncn), one of the four national collections housed in the biosystematics programme of the plant protection research institute, agricultural research council, pretoria, was founded in the mid-1950s, with the first type specimens deposited in 1958. the national collection of nematodes consists of a core collection, the meloidogyne collection and the juan heyns collection. the national collection of nematodes currently consists of 180 000 specimens, of which 7140 are type specimens, mostly from southern africa, but also from such diverse localities as antarctica and the amazon.the type specimens of 510 species are currently deposited in ncn and we now know that 455 plant nematode or plant-feeding nematode species are reported from south africa. the south african plant-parasitic nematode survey (sappns) programme was initiated in 1987 to produce a comprehensive assessment of the nematode biodiversity resources of south africa. one of the four objectives of sappns is to compile an inventory of the plant nematodes of south africa (marais 2006). as part of this initiative, surveys were undertaken in various areas, including some of the protected areas of the eastern cape, in order to establish the incidence and distribution of plant nematodes in these areas (van den berg 1996). in this article, plant nematodes are seen as nematodes that feed on plants and therefore always have a tylenchoid stomatostyle, dorylaimoid odontostyle or trichodorid onchiostyle. this includes nematodes that have an ectoparasitic, endoparasitic or semi-endoparasitic life cycle (yeates et al. 1993). as recommended in the international code of zoological nomenclature, locality data and host plant data are usually published as part of species descriptions. apart from a checklist of the tsitsikamma national park (van den berg 1996), the nematode biodiversity information of the eastern cape had not been consolidated into a single publication. during the digitising of the specimens of ncn, a number of unpublished records of plant nematodes reported from protected areas in the eastern cape were discovered. materials and methods top ↑ the datasets reflected in the checklist consist of nematodes collected by various nematologists over a period of 30 years. in some cases the collection and extraction methods are not known. the standard procedure of ncn is that soil samples are taken with a garden trowel, spade or soil auger, depending on the terrain in the rhizosphere of the plant. the plants are collected at the same time and the soil and accompanying plant are sealed in plastic bags, placed in a cool box containing ice bricks and transported to the laboratory of ncn where the samples are stored in a cold room at 12 °c. the relevant collection data, including locality, habitat, substrate, moisture regime, soil type, exposure slope, aspect and host plant, are noted in the field.the nematodes are extracted from the soil using the sugar centrifugal-flotation method (jenkins 1964). specimens are killed in water by the gradual application of heat, then fixed and preserved in distilled water, 4% formaldehyde and 1% propionic acid (known as fpg), dehydrated to anhydrous glycerine and permanently mounted in anhydrous glycerine on cobb aluminium slides (hooper & evans 1993; netscher & seinhorst 1969). nematodes are extracted from plant material using the method described by kleynhans (1997), and are killed, preserved and mounted in the same way as the nematodes extracted from soil. the classification of south african plant nematodes followed here is a synthesis of the classification by maggenti et al. (1988) for tylenchina; and decraemer (1995) and duarte et al. (2010) for trichodoridae. authorities for genera regarded as valid here are hunt, luc and manzanilla-lópez (2005) for criconemoides; marais (2001) for helicotylenchus; escuer and arias (1997) for paralongidorus; brzeski (1998) for paratylenchus; and handoo (2000) for tylenchorhynchus. results and discussion top ↑ the checklist reflects specimens collected between 1968 and 1998 from seven protected areas in the eastern cape. the sappns database contains 311 records of nematodes sampled in uncultivated areas in the eastern cape, of which 98 records are from localities in protected areas (online appendix 1). the database also contains 505 records of localities sampled in cultivated areas, which include crop fields, plantations, gardens and sports fields. it was found that 80 plant nematode species were reported from protected areas (table 1), 163 species from uncultivated areas (outside protected areas) and 150 species from cultivated areas.in total, 230 (51%) of the 455 currently known plant nematode species from south africa have been recorded from the eastern cape. of these, 29 species were described from the province; 10 of these were described from the protected areas in the province: criconemoides silvicola van den berg 1996, meloinema silvicola kleynhans 1988, ogma tuberculatum van den berg 1996, paralongidorus cebensis (heyns & coomans 1989) escuer & arias 1997, paralongidorus hanliae liebenberg, heyns & swart 1993, scutellonema tsitsikamense van den berg 1976, trichodorus vandenbergae de waele & kilian 1992, xiphinema erriae hutsebaut, heyns & coomans 1988, xiphinema ornatizulu hutsebaut, heyns & coomans 1989 and xiphinema simplex hutsebaut, heyns & coomans 1989. only m. silvicola, o. tuberculatum, p. cebensis and s. tsitsikamense were not reported from other provinces, suggesting endemism. acknowledgements top ↑ we thank various colleagues and clients for collecting the samples, the taxonomists at arc-ppri and (the former) rand afrikaans university for species identification and the south african biodiversity information facility for financial assistance in order to digitise a major portion of ncn. competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions m.m. (plant protection research institute) is the coordinator of sappns, project leader of digitising the national collection and taxonomist responsible for helicotylenchus, belonolaimidae, dolichoridae, trichodoridae and the subfamily meloidogyninae, and was responsible for writing the draft concept of the manuscript. a.s. (plant protection research institute) is the taxonomist responsible for longidoridae, anguinidae, aphelenchoididae and the subfamily heteroderinae, and was responsible for the final review of the manuscript. m.m. and a.s. were both responsible for writing this article. references top ↑ brzeski, m., 1998, nematodes of tylenchina in poland and temperate europe, museum i instytut zoologii polska akademia nauk, warzawa.decraemer, w., 1995, the family trichodoridae: stubby root and virus vector nematodes, kluwer academic press, dordrecht. http://dx.doi.org/10.1007/978-94-015-8482. duarte, i.m., de almeida, m.t., brown, d.j.f., marques, i., nielsen, r. & decraemer, w., 2010, ‘phylogenetic relationships, based on ssu rdna sequences, among the didelphic genera of the family trichodoridae from portugal’, nematology 12(2), 171–180. http://dx.doi.org/10.1163/156854109x461721 escuer, m. & arias, m., 1997, ‘paralongidorus iberiss p.n. and p. monegrensiss p.n. from spain with a polytomous key to the species of the genus paralongidorus siddiqi, hooper & khan, 1963 (nematoda: longidoridae)’, fundamental and applied nematology 20(2), 135–148 viewed n.d., from http://horizon.documentation.ird.fr/exl-doc/pleins_textes/fan/010009433.pdf handoo, z., 2000, ‘a key and diagnostic compendium to the species of the genus tylenchorhynchus cobb, 1913 (nematoda: belonolaimidae)’, journal of nematology 32(1), 20–34. pmid:19270946. hooper, d.j. & evans, k., 1993, ‘extraction, identification and control of plant nematodes’, in k. evans, d.l. trudgill & j.m. webster (eds.), plant parasitic nematodes in temperate agriculture, pp. 31–59, cab international, wallingford. hunt, d.j., luc, m. & manzanilla-lópez, r.h., 2005, ‘identification, morphology and biology of plant parasitic nematodes’, in m. luc, r.a. sikora & j. bridge (eds.), plant parasitic nematodes in subtropical and tropical agriculture, 2nd edn., pp. 11–52, cabi publishing, wallingford. http://dx.doi.org/10.1079/9780851997278.0011 jenkins, w.r., 1964, ‘a rapid centrifugal-flotation technique for separating nematodes from soil’, plant disease reporter 48, 692. kleynhans, k.p.n., 1997, collecting and preserving nematodes. manual for a safrinet course in practical nematology, safrinet, the southern african (sadc) loop of bionet-international, pretoria. maggenti, a.r., luc, m., raski, d.j., fortuner, r. & geraert, e., 1988, ‘a reappraisal of tylenchina (nemata). 11. list of generic and supra-generic taxa, with their junior synonyms’, revue de nématologie 11, 177–188. marais, m., 2001, ‘a monograph of the genus helicotylenchus steiner, 1945 (nemata: hoplolaimidae)’, phd dissertation, department of entomology and nematology, university of stellenbosch. marais, m., 2006, ‘south african plant-parasitic nematode survey (sappns)’, plant protection news 67, 6. netscher, c. & seinhorst, j.w., 1969, ‘propionic acid better than acetic acid for killing nematodes’, nematologica 15, 286. van den berg, e., 1996, ‘a first list of plant-parasitic nematodes from the tsitsikamma national park, with descriptions of two new species of the subfamily criconematinae’, koedoe 39(1), 43–54. http://dx.doi.org/10.4102/koedoe.v39i1.281 yeates, g.w., bongers, t., de goede, r.g.m., freckman, d.w. & georgieva, s.s., 1993, ‘feeding habits in nematode families and genera – an outline for soil ecologists’, journal of nematology 25, 315–331. filelist convert a pdf file! supplement to koedoe. 1977: 255-259. suid-afrika se nasionale parke sisteem wat van die toekoms? sy edele h schoeman* minister van landbou republiek van suid-afrika abstract the significance of the 50th anniversary of the national parks board of trustees is highlighted and homage is paid to all those who have contributed to the development of the national parks system in the republic of south africa. the government never lightly refuses requests on sustained pressures for new national parks and establishment of a new national park near beaufort west in the karoo, is announced. mnr die voorsitter, uitgelese gaste, dames en here. u sal my toelaat om reg by die begin verskoning te maak dat sy edele minister h schoeman nie aanwesig kan wees nie weens omstandighede buite sy beheer. u sal my ook toelaat om sommer nou aan die voorsitter 'n woordjie toe te voeg van 'n belofte wat hy gemaak het en dit is toe hy ons uitgenooi het na hierdie funksie, dat ek hier kan kom en op my gemak kom sit en die tydjie in die wildtuin net kan geniet. hy het die belofte egter vergeet, ek is seker daarvan. maar, weens omstandighede buite die beheer van sy edele die minister, moet ek nou vandag hier optree soos wat dit in engels gestel word "i am deputising, true to the real meaning of what a deputy minister means". u weet minister schoeman het van tyd tot tyd gese 'n adjunk-minister is 50% van niks en ek dink hy is nie baie ver uit nie! mnr die voorsitter, u sal my dit vergun dat ek vir u gelukwens met u persoonlike verjaardag. ons weet dat dr knobel 'n besondere vindingryke persoonlikheid is. ek wil hom nie blameer en ek glo nie iemand hier in die gehoor sal hom blameer, dat hy dit so bewerkstellig het dat die nasionale parkeraad se verjaardag saamval met sy eie nic. maar ek wil dit darem net as 'n opmerking laat dat ek nie 'n enkele amptenaar ken, maak nie saak van watter status, wat so vindingryk is om te sorg dat soveel uitgelese gaste sy verjaardag bywoon nie! ek wil hom gelukwens daarmee en daarmee gaan dit ook na mev knobel. ons weet dat sy hom beslis in alles bystaan en beslis veel kleur gee aan dit wat hy doen. ons hoop dat hulle hierdie jaar wat voorle, baie sal geniet en dat u ook vee i *referaat voorgedra deur sy ed ele j j malan, adjunk-minister van landbou. 255 sukses sal he met u werk hierso as hoofdirekteur van die nasionale parkeraad. waar ek namens minister schoeman hier optree wi! ek dan ook hoop dat u sover 'n besondere vrugbare simposium gehad het en as ek geluister het na wat dr. martiny hier as opsomming aan ons voorgehou het, is ek jammer dat ek nie die hele tyd hier kon gewees het nie. dit wil my voorkom onder andere, en dan is ek heeltemal objektief, dat iemand uit ons departement landbou-tegniese dienste minstens gesorg het dat die boer se standpunt hier gestel is en dat u u voete op die grond hou. dit is vir my baie interessant toe dr. martiny gepraat het, en waar hy dit so mooi gestel het dat daar selfs van owerheidswee meer gedoen moet word, kon ek egter nie help as om daaraan te dink dat dit darem eienaardig is dat mense nie die essensie van dinge snap nie. wanneer 'n demokratiese land soos ons s'n aan mense eise stel, verwagtinge na vore bring, dan se hulle by implikasie, ons wil dit he, en ons is gewillig om daarvoor te betaal. dit is ook waar ek 'n probleem het met die regering. u weet ek het drie besware teen die regering t.w.: as jy hulle nader dan het hulle probleme in die eerste instansie met finansies, in die tweede instansie het hulle probleme met verkryging van fondse en in die derde instansie het hulle probleme met geld! dit is vir my 'n eer om vandag hier namens my minister op te tree by hierdie besondere geleentheid. dit is vir my dan ook aangenaam om sy toespraak hier aan u voor te lees. die bereiking van so 'n belangrike mylpaal as die 50ste verjaardag van die totstandkoming van die liggaam soos die raad van kuratore vir nasionale parke is ongetwyfeld 'n trotse gebeurtenis. dit getuig van bestendigheid en dat die liggaam tot groot hoogte die taak wat aan hom opgedra is suksesvol uitgevoer het. geen liggaam wat nie redelikerwys voldoen aan die eise wat aan hom gestel word, sal 50 jaar lank kan bestaan en gedy nie. op 'n dag soos hierdie is dit paslik om 'n terugblik te werp in die verlede. by so 'n terugblik is dit baie duidelik dat daar veral rede is tot groot dankbaarheid, maar ook om trots te kan wees. soos ook elders vandag by 'n ander geleentheid staan ons in groot ootmoed en erkentlikheid voor die herinnering van die historiese figuur paul kruger. aan hom kom die hulde toe vir sy grootmoedigheid en gebruiklike versiendheid om in die pionierstyd van die ontwikkeling van ons land en volk reeds 'n gebied af te sonder wat die natuur in sy ongeskonde staat bewaar en sou beskerm. hy staan natuurlik nie aileen in die opsig nie, maar aan die voorpunt van 'n lang ry mense, wat groot bydraes gel ewer het tot die totstandkoming en ontwikkeling van hierdie en ons ander nege nasionale parke. daar was lede van die regering van ons land wat van tyd tot tyd groot entoesiaste van natuurbewaring was en wat deur regeringsoptrede die ontwikkeling van die stelsel van ons nasionale parke bevorder het. ek verswyg liewer hulle name omdat dit seker is dat wanneer ek hulle probeer noem die name van sommige van hulle onregverdiglik weggelaat sal word. 256 mr chairman, we cherish the memory of many private individuals who have already passed away but who during their lifetimes were enthusiastic nature lovers and contributed to the development of our national parks by the grant of land, money and other gifts. it is gratifying to know that also the present generation has not been lacking in displaying the true participation in our company here today and who have also made many material and substantial contributions both in money and in respect of services rendered towards the further development of our national parks. it is only because i realise that they would not like to advertise the fact that they are serving the country also in this respect, that i refrain from mentioning their names. we remember the many persons who have served as members of the national parks board of trustees. some of them with distinction. it is a fact that the board has created an image of itself which is looked upon by many people with such high respect that they regard an appointment to the board as the highest achievement possible. dit is interessant om net daaraan te dink hulle se dit word oorvertel in die wandelgange van die parlement " ... that the definition of a national park is a place where people are sent when they are tired and need to relax, but appointment to the board is for those who have spent their lives there!". ons neem vandag afskeid van een van die dienende lede van die parkeraad. hy is mnr tiny faure. hy het met onderskeiding en met groot toewyding as lid van die parkeraad gedien vir nie minder as 26 jaar nie. oom tiny sou seker nog graag voortgegaan het as lid van die parkeraad as dit nie was dat hy noodwendig sy aktiwiteite wil begin inkort nie. ek se dankie aan hom vir sy lange jare van uitmuntende diens as lid van die raad en wens hom die allerbeste toe op die pad vorentoe. ons is jammer om te hoor dat mev. faure nie so wel is nie en ons wil graag dit wat ons van oom tiny se, dat hy dit aan haar sal oordra. ons is dankbaar ook teenoor haar vir wat sy gedoen het en ons wens is dat sy geheel-en-al sal hers tel. ons dink ook aan die aandeel van die amptenare van die raad oor die jare. die van hulle wat vandag nog in die diens van die parkeraad is, diegene wat die parke geskep het, waarop ons so trots is en wat aan ons volk en aan ons mense asook mense van oor die hele wereld so 'n unieke vreugde en genot verskaf. ons wil ook met groot waardering verwys na die verskillende staatsdepartemente en ander owerheidsinstansies wat oar die jare steeds uiters simpatiek opgetree het wanneer hulle met sake van die nasionale parkeraad te doen gekry het. hierdie welwillendheid het in 'n aansienlike mate die totstandkoming en ontwikkeling van ons nasionale parke moontlik gemaak. mr chairman, we regard what has been achieved in the field of nature conservation in our country with justifiable pride. nature conservation in what may be termed national parks, is rightly the function of the national parks board of trustees. national parks to my mind are 257 necessary to protect and nourish the continued existence of objects of nature for the benefit of the nation, where such conservation bears a national character and calls for action on a national scale. enthusiasts from the ranks of the board and its officials and other nature lovers are continually pressing for the creation of further national parks, or for the extension of their areas, or for improvements of the facilities in existing national parks. i wish to assure them that the government never lightly refuses these requests in this regard and i am sure the chairman will bear me out on that. hoe graag ookal die regering hoogsprioriteit sa! wil verleen aan aansoeke om ondersteuning van die nasionale parkeraad is hy nogtans soms verplig om sulke aansoeke af te wys. dit gebeur in elk geval nie dikwels nie. daarmee moet egter rekening gehou word dat aansprake op die land se bates, veral in die vorm van grond en geld geweldig is, en dat dit al hoe moeiliker word om selfs die verdienstelikste voorstelle te bevredig. dit is vir my as minister van landbou ook 'n voorreg om te kan getuig dat in die landbou, van die grootste gedeelte van die oppervlakte van ons land wat landboukundig aangewend word, die belangrikheid van bewaring van die gawes van die natuur in die vorm van landbouhulpbronne oor die jare heen al hoe meer besef word. vandag is juis hierdie aspek een van die belangrikste onderdele van die boerdery. ek maak die stelling dat geweldig baie reeds gedoen word om die gawes wat die natuur ons vaderland bied, te beskerm en te bewaar. ook is daar groot vordering gemaak met die inskerping van 'n nasionale bewustheid van die waarde van ons natuurlike hulpbronne by die publiek. terwyl ons erken dat baie meer nog gedoen moet word , hoef ons land nie skaam te wees vir wat reeds bereik is en wat tans onderneem word nie. moontlik moet ondersoek ingestel word of daar met inagneming van beskikbare fondse nie uitbreidings aan die volgende aksies bewerkstellig kan word nie . ek noem hulle: (a) ek dink dat dit noodsaaklik is om te kyk tot watter mate die bergkwagga nasionale park, die bontebok nasionale park en die tsitsikamabos en -seekus nasionale park vergroot kan word. dat dit nodig is, is seker. of dit gedoen kan word, dit moet die toekoms bepaal. (b) die stigting van nuwe nasionale parke om meeste van die volgende belangrike gebiede te bewaar: die rigtersveld; sommige voeleilande en robeilande aan ons kuste; die kosibaai-kompleks; deel van die sederberge; deel van die drakensberg eskarpement; 'n namakwaland blommepark; die bobbejaanskloof; 'n grot nasionale park en dan natuurlik 'n karoo nasionale park. dit is dan met groot vreugde dat ek kan aankondig dat die kabinet in beginsel goedkeuring verleen het aan die stigting van 'n karoo nasio258 nale park by beaufortwes. die park word moontlik gemaak deur 'n wonderlike skenking van grond op die dorpsmeent deur die munisipaliteit van beaufort-wes. die munisipaliteit word vandag hier verteenwoordig en ons se baie dankie vir die besondere bydrae deur 'n enkele munisipaliteit vir 'n skenking wat 'n groot nasionale bate gaan word. tot hierdie grond word bygevoeg grond wat bygekoop word deur gelde voorsien deur die suid-afrikaanse natuurstigting wat bykans rloo 000 beloop asook 'n bedrag uit die staatskis van rioo 000. die suidafrikaanse natuurstigting wat hier verteenwoordig word, moet spesiale vermelding kry omdat hulle dit reggekry het om gelde van die algemene publiek en veral skoolkinders vir die doe! in te same!. ek dink dit was 'n besonderse poging. ons wil die stigting en deur hulle aan almal wat bygedra het tot die karoo parkefonds baie dankie se. ons vertrou dat die wonderlike aksie wat hulle begin het, voortgesit sal word en dit die algemene publiek meer en meer sal aanspoor om grond vir nasionale parke te verkry, sodat die raad nie net na die staat hoef te kyk vir die verkryging van fondse nie. die gronde bly 'n volksbesit en dit gee my dus groot vreugde om op die 50ste bestaansjaar van die nasionale parkeraad die stigting van die lode nasionale park te mag aankondig. ek sluit af met die gedagte, mnr die voorsitter, dat die nasionale parkeraad veral kan trots wees op wat hy tot stand gebring het. die hele land kan trots wees op wat tot stand gebring is en kan spog met dit wat die raad opgebou het. ons is dankbaar daarvoor en ons vertrou dat die raad op die pad vorentoe net sovee! sukses sal he as op die pad wat reeds agter ie. mag die ideale wat vir die toekoms gekoester word, almal bewaarheid word. baie dankie. 259 page 1 page 2 page 3 page 4 page 5 filelist convert a pdf file! koedoe 19: 49-62(1976) a revised check-list of birds in the kalahari gemsbok national park m. g . l. mills kalahari gemsbok national park private bag x890 p.o. gemsbokpark 8815 abstract a more complete list of the birds in the kalahari gemsbok national park is given, including the results of three and a half years of observations. a total of 214 species have now been identified for the park, among which 75 are resident throughout the year, 37 are migrants, 14 are nomads and 88 are vagrants. introduction several lists of the birds in the kalahari gemsbok national park, republic of south africa, have been published in recent years (de villiers 1958; prozesky and haagner 1962; broekhuysen, broekhuysen, martin, martin, martin and morgan 1968; maclean 1970). with the exception of maclean (1970) these lists have been drawn up by the authors during short visits to the park. however, maclean only worked in the southern area for 19 months. this list represents three and a half years observations over the entire park, with additional records from liversidge (pen. comm.). several groups of birds, notably the rap tors, pipits, larks and warblers, are difficult to identify in the field. where any doubt exists as to the identity of a species, it has been omitted unless a specimen has been taken. systematic list an indication of the status of each species in the park is given according to the following key (after kemp 1974); r resident (found in the park throughout the year) m migrant (a species known to come to south africa on a regular annual migration) n nomad (a regular, temporary visitor to the area) v vagrant (an irregular, temporary visitor to the area) . the number of each species has been taken from mclachlan and liversidge (1966) and the nomenclature follows the check list of the birds of south africa (s.a.o.s . list committee 1969). 49 ostrich (r) volstruis 6 cape dabchick (v) kaapse duikertjie struthio camelus linnaeus podiceps ruficollis (pallas) 47 white-breasted cormorant (v) witbo rsdu i ker phalacrocorax lucidus (lichtenstein) 50 reed cormorant (v) rietduiker 54 grey heron (v) bloureier 55 black-headed heron (v) phalacrocorax ajricanus (gmelin) ardea cinerea linnaeus swartkopreier ardea melanocephala vigors & children 58 great white egret (v) groot witreier 60 yellow-billed egret (v) geelbek-witreier 61 cattle eg ret (v) bosluisvoel 62 squacco heron (v) ralreier 67 little bittern (v) woudapie 69 night heron (v) nagreier 72 hamerkop (v) egretta alba (linnaeus) egretta intermedia (wagler) ardeola ibis (linnaeus) ardeola ralloides (scopoli) ixobrychus minutus (linnaeus) nycticorax nycticorax (linnaeus) scopus umbretta gmelin 73 marabou stork (v) leptoptilos crumeniferus (lesson) maraboe 75 saddlebill (v) ephippiorhynchus senegalensis (shaw) saalbekooivaar 77 wooly -necked stork (v) ciconia episcopus (boddaert) wolnekooievaar 50 78 white-bellied stork (m) klein swart sprinkaanvoel 79 black stork (v) ciconia abdimii (lichtenstein) ciconia nigra (linnaeus) groot swart sprinkaanvoel 80 white stork (m) groot wit sprinkaanvoel 123 rock kestrel (r) rooivalkie ciconia ciconia (linnaeus) falco tinnunculus linnaeus 126 pigmy falcon (r) dwergvalkie poliohierax semitorquatus a. smith 128 black kite (m) swartwou 129 yellow-billed kite (m) geelbekwou milvus migrans (boddaert) milvus migrans parasitus (daudin) 130 black-shouldered kite (r) elanus caeruleus (desfontaines) blouvalkie 134 tawny eagle (r) kouvoel 135 steppe eagle (m) steppe-arend 139 booted eagle (m) dwergarend 141 african hawk eagle (v) afrikaanse jagarend 142 martial eagle (r) breekoparend 145 brown snake-eagle (r) bruin slangarend aquila rapax (temminck) aquila nipalensis (hodgson) aquila pennata (gmdin) aquila fasciata vieillot polemaetus bellicosus (daudin) circaetus cinereus vieillot 146 black-breasted snake-eagle (r) swartbos slangarend ciraetus pectoralis a. smith 149 fish eagle (v) visarend 51 haliaetus vocifer (daudin) 151 bataleur (r) berghaan 152 jackal buzzard (v) jakkalsvoel 154 buzzard (m) jakkalsvoel 157 ovambo sparrowhawk (v) ov ambosperwer 161 little-banded goshawk (v) klein gebande sperwer 162 gabar goshawk (r) klei n witv alki e 165 chanting goshawk (r) groot witv alk 168 pallid harrier (m) vaal paddavreter 169 black harrier (v) witkruisv alk 170 montagu's harrier (m) blou paddavreter 171 banded harrier hawk (v) gymnogene 189 african quail (n) afrikaanse kwartel terathopius ecaudatus (daudin) buteo rujofuscus (forster) buteo buteo (linnaeus) accipiter ovampensis gurney accipiter badius (gmelin) micronisus gabar (daudin) melierax musicus (daudin) circus macrourus (gmelin) circus maurus (temminck) circus pygargus (linnaeus) polyboroides radiatus (scopoli) cortumix cotumix (linnaeus) 190 harlequin quail (v) bontkwartel cotumix delegorguei delegorgue 192 crownded guinea-fowl (v) numida meleagris (linnaeus) tarentaal 196 kurrichane button-quail (n) bosveldkwartel tjie tumix sylvatica (desfontains) 52 208 purple gallinule (v) porphyrio porphyrio (linnaeus) koningriethaan 212 red-knobbed coot (v) fulica cristata gmelin bleshoender 217 kori bustard (r) otis kori burchell gompou 218 ludwig's bustard (v) otisludwigii (ruppel) ludwigse pou 219 stanley bustard (v) otis denhami children veldpou 224 red-crested korhaan (r) eupodotis rujicrista (a. smith) boskorhaan 225 black korhaan (r) afrotis ajra (linnaeus) swartkorhaan 228 african jacana (v) actophilomis africanus (gmelin) langtoon 238 three-banded sand plover (v) charadrius tricollaris vieillot drieband-strandlopertjie 240 caspian plover (m) asiastiese strandloper 242 crowned plover (r) kiewietjie 245 blacksmith plover (v) bontkiewietjie 251 curlew sandpiper (m) kro mbek-strandloper 253 little stint (m) klein strandloper 256 ruff (m) kemphaan 258 common sandpiper (m) gewone ruiter 53 charadrius asiaticus pallas vanellus coronatus (boddaen) vanellus armatus (burchell) calidris minuta (pontoppidan) calidris minuta (leislerl philomachus pugnax (linnaeus) tringa hypoleucos (linnaeus) 262 marsh sandpiper (m) tringa stagnatilis (bechstein) m 0 erasruiter 263 greenshank (m) tringa nebularia (gunnerus) groenpoot-ruiter 264 wpod sandpiper (m) tringaglareola linnaeus bosruiter 268 whimbrel (m) numenius phaeopus (linnaeus) klein wulp 270 black-winged stilt (v) himantopus himantopus (linnaeus) rooipoot-elsie 275 cape dikkop (r) burhinus capensis (lichtenstein) dikkop 276 burchell's courser (n) cursorius rufus gould bloukop-drawwertjie 277 temminck's courser (v) cursorius temminckii swainson trekdrawwertjie 278 double-banded courser (r) dubbelband-drawwertjie rhinoptilus ajricanus (temminck) 280 bronze-wing courser (v) rhinoptilus chalcopterus (heuglin) bronsvlerkdrawwertjie 304 white -winged lake tern (m) witvlerkmeerswawel childonias leucoptera (temminck) 307 namaq.ua sandgrouse (r) pterocles namaqua (gmdin) kelkiewyn 308 spotted sandgrouse (r) pterocles burchelli w. sclater sandpatrys 311 rock pigeon (v) columba guinea linnaeus bosduif 316 cape turtle dove (r) streptopelia capicola (sundevalj) tortelduif 317 laughing dove (r) rooibors-duifie streptopelia senegalensis (linnaeus) 54 318 namaqua dove (r) namakwaduifie 340 cuckoo (m) koekoek oena capensis (linnaeus) cuculus canorus linnaeus 346 great spotted cuckoo (v) clamator glandarius (linnaeus) groot gevlektekoekoek 348 jacobin cuckoo (m) clamator jacobinus (boddaert) nuwejaarsvoel 352 diederik cuckoo (m) chrysococcyx caprius (boddaert) diedrikkie 359 barn owl (r) tyto alba (scopoli) nonnetjie-uil 360 grass owl (v) tyto capensis (a. smith) grasuil 363 scops owl (v) otus scops (linnaeus) klein ooruil '''364 white-faced owl (r) otusleucotis (temminck) witwang-ooruiltjie 365 pearl-spotted owlet (r) glaucidium perlatum (vieillot) dwerg-uiltjie 368 spotted eagle owl (r) bubo ajricanus (temminck) gevlekte ooruil 369 giant eagle owl (r) bubo lacteus (temminck) reuse-ooruil 371 european nightjar (m) camprimulgus europaeus linnaeus europese naguil 372 rufous-cheeked nightjar (m) rooiwang-naguil caprimulgus rufigena a. smith 378 european swift (m) europese windswawel 380 black swift (m) swart windswawel 55 apus apus (linnaeus) apus barbatus (p. sclater) 383 white-rumped swift (v) apus caffer (lichtenstein) witkrui swi ndswa wel 385 little swift (v) apus ajjinis (gray) klein windswawel 391 white-backed mousebird (v) colius colius (linnaeus) witkruis-muisvot.l 392 red-faced mousebird (v) urocolius indicus (latham) rooiwang-muisvot.l 403 striped kingfisher (v) halcyon chelicuti (stanley) gestreepte visvanger 404 european bee-eater (m) merops apiaster linnaeus europese byvreter 411 swallow-tailed bee-eater (r) miksterbyvreter merops hirundineus (lichtenstein) 412 european roller (m) coracias garrulus linnaeus europese troupant 413 lilac-breasted roller (r) coraciascaudata linnaeus gewone troupant 415 purple roller coracias naevia daudin groot troupant 418 african hoopoe (r) upupa epops bechstein hoephoep 419 red-billed hoopoe (v) phoeniculus purpureus (miller) kakelaar 421 scimitar-bill hoopoe (r) swartbekkakelaar rhinopomastus cyanomelas (vieillot) 424 grey hornbill (v) grys neushoringvot.l tockus nasutus linnaeus 426 yellow-billed hornbill (r) tockusflavirostris (ruppel!) geelbekneushoringvot.l 432 pied barbet (r) lybius leucomelas (boddaert) bont houtkapper 56 450 cardinal woodpecker (r) dendropicosfuscescens (vieillot) kardinaal-spegt 457 monotonous lark (v) mirofrajavanica horsfield bosveldlewerkie 459 fawn-coloured lark (r) mirajra ajricanoides a. smith vaalbruin lewerkie 460 sabota lark (r) mirajrasabotaa. smith sabota-lewerkie 464 dusky lark (v) pinarocorys nigricans (sundevall) donker lewerkie 466 clapper lark (r) mirajra apiata (vieillot) klappertj i e 474 spike-heeled lark (r) vlakvoeltjie certhilauda albojasciata lafresnaye 485 grey-backed finch-lark (n) grys kaffertjie eremopterix verticalis (a. smith) 486 black-eared finch-lark (v) egte kaffertjie eremopterix australis (a. smith) 488 red-capped lark (v) rooikoplewerkie tephrocorys cinerea (gmelin) 490 pink-billed lark (v) rooibeklewerkie calandrella conirostris (sundevall) 492 stark's lark (n) starkse lewerkie 493 european swallow (m) europese swawel 495 white-throated swallow (v) calandrella slarki shelley hirundo rustica linnaeus witkeelswa wel hirundo albigularis strickland 498 pearl-breasted swallow (v) hirundo dimidiata sundevall p~relbo rsswa wel tj i e 502 larger striped swallow (v) hirundo culcullata (boddaert) groot streepswawel 57 504 cliff swallow (r) familieswawel 506 rock martin (r) kransswawel 517 fork-tailed drongo (r) m i kstertbyv an g er 519 golden oriole (m) europese wielewaal 522 pied crow (v) witborskraai 523 black crow (r) swartkraai 525 grey tit (r) piettjoutjou hirundo spilodera (sundevall) hirundo rupestris scopoli dierurus adsimilis (bechstein) oriolus oriolus (linnaeus) corvus albus (miiller) corvus eapensis lichtenstein parus ajer gmelin 531 penduline tit (r) anthoseopus minutus (shaw and nodder) grys kapokvoel 536 pied babbler (v) turdoides bieolor (jardine) wit ka tlagter 544 red-eyed bulbul (n) pyenonotus nigrieans (vieillot) rooioog-tiptol 557 groundscraper thrush (v) turduslitsitsirupa (a. smith) gevlekte lyster 561 short-toed rock thrush (v) korttoon-klipwagter montieola brevi pes (waterhouse) 564 mountain chat (v) bergtapult 566 grey-rumped sickle wing chat (v) bleekt apuit 568 capped wheatear (m) skaapwagter 570 familiar chat (r) spekvreter 58 oenanthe montieola vieillot cereomela sehlegelii (wahlberg) oenanthe pileala (gmelin) cereomela jamiliaris (stephens) 571 layard ' s chat (v) cercomela tractrac (wilkes) klein spekvreter 575 ant-eating chat (r) myrmecocichlaformicivora (vieillot) swartpiek 586 kalahari scrub robin (r) erythropygia paena a. smith wipstert 599 willow warbler (m) phylloscopus trochilus (linnaeus) hofzanger 600 yellow-bellied bush warbler (r) geelbuikbossanger eremomela icteropygialis (lafresnaye) 619 rufous-eared warbler (r) roo i 0 0 rklei njantji e 621 crombek (v) krombek 629 common fantail cisticola (n) prinia pectoralis a. smith sylvietta rujescens (vieillot) gewone veldtingtinkie cisticolajuncidis (rafinesque) 630 desert cisticola (r) cisticola aridula whiterby bleekveld-tinktinkie 650 black-chested prinia (r) priniajlavicans (vieillot) swartbors-langsterttinktinkie 654 spotted flycatcher (m) muscicapa striata (pallas) europese vlieevanger 658 tit-babbler (r) parisoma subcaerulem (vieillot) tjeriktik 661 marl co flycatcher (r) melaenomis mariquensis a . smith marl co vlieevanger 663 chat flycatcher (r) melaenomis injuscatus (a . smith) groot vlieevanger 674 pririt flycatcher (r) batispririt(vieillot) pririt-bonbontrokkie 686 cape wagtail (v) motacilla capensis linnaeus kwikkie 59 689 yellow wagtail (v) geel kwikkie 692 richard's pipit (n) gewone koester 695 buffy pipit (n) vaalkoester 706 lesser grey shrike (m) europese grys laksman 707 fiscal shrike (r) laksman 708 red-backed shrike (m) rooiruglaksman motacilla lutea (gmelin) anthus novaeseelandiae (gmelin) anthus vaalensis shelley lanius minor gmelin lanius collaris linnaeus lanius collurio linnaeus 711 crimson-breasted shrike (r) rooiborsfiskaal liniarius atrococcineus burchell 714 three-streaked tchagra (v) tchagra australis (a. smith) klein rooivlerklaksman 722 bokmakierie (r) malaconotus z.eylonus (linnaeus) 730 white-crowned shrike (v) kremetartlaksman eurocephalus anguitimens a. smith 731 brubru shrike (r) bontroklaksman 735 wattled starling (n) sprinkaanvoel 737 cape glossy starling (r) klein glansspreeu nilaus afer (latham) creatophora cinerea (menschen) lamprotornis nitens (linnaeus) 743 burchell's glossy starling (r) groot glansspreeu lamprotornis australis (a. smith) 764 dusky sunbird (v) namakwa-suikerbekkie 775 cape white-eye (v) kaapse glasogie 60 nectarinia fusca (vieillot) zosterops pallidus swainson 779 buffalo weaver (v) buffelwewer bubalomis albirostris (vieillot) 780 white-browed sparrow-weaver (r) koringvoel plocepasser mahali a. smith 783 sociable weaver (r) familievoel 784 house sparrow (r) engelse mossie 785 great sparrow (n) groot mossie 786 cape sparrow (r) mossie 787 grey-headed sparrow (r) gryskopmossie philetairus socius (latham) passer domesticus (linnaeus) passer motitensis (smith) passer melanurus (muller) passer griseus (vieillot) 789 scaly-feathered finch (r) sporopipes squamifrons (a. smith) baardmannetjie 803 masked weaver (r) ploceus velatus vieillot swartkeel-g eel vin k 805 red-billed quelea (n) quelea quelea (linnaeus) rooibekvink 808 red bishop bird (v) euplectes orix (linnaeus) rooi kaffervink 820 red-headed finch (n) amadina erythrocephala (linnaeus) rooikopmossie 840 violet-eared waxbill (v) uraeginthus granatina (linnaeus) koningrooibekkie 843 common waxbill (v) estrilda astrild (linnaeus) rooibekkie 847 shaft-tailed whydah (r) vidua regia (linnaeus) pylstert 861 blackhead canary (v) serinus alario (linnaeus) swartkop-kanarie 61 866 yellow canary (r) geelsysie 871 lark-like bunting (n) vaal streepkoppie references serinus flaviventris (swainson) emberiza impetuani a. smith. broekhuysen, g. j., m. h. broekhuysen, j. martin e. martin, r. martin and h. k. morgan. 1968. observations on the birdlife in the kalahari gemsbok national park. koedoe ii: 145-160. de villiers, j. s. 1958. a report on the bird life of the kalahari gemsbok national park. koedoe 1: 143-161. kemp, a. c. 1974. the distribution and status of the birds of the kruger national park. koedoe monograph 2. mclachlan, g. r. and r. liversidge. 1966. roberts birds oj south africa. central news agency, south africa. maclean, g. l. 1970. an analysis of the avifauna of the southern kalahari gemsbok national park. zool. afro 5(2):249-273. prozesky, o. p. m. and c. h. haagner. 1962. a check-list of the birds of the kalahari gemsbok park. koedoe 5: 171-182. s.a.o.s . list committee. 1969. check. list oj the birds of south africa. cape town: south african ornithological society. 62 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 page 11 page 12 page 13 page 14 filelist convert a pdf file! supplement to koedoe. 1977: 203-209. the conservation role of forestry in south africa d p ackerman secretary, department of forestry private bag x93 pretoria 0001 in september 1965, at the signing ce remony of a national park bill, united states presid ent johnson remarked: " we are living in th e century of change. but if future gen erations are to rem ember us more with gratitude than with sorrow, we must achieve more than just th e miracles of technology. we must also leave them a glimpse of th e world as god really mad e it, not just as it looked when we got through with it." in th e same spirit, forestry in the republic of south africa (rsa) shares the responsibility of conserving and restoring as far as possible the environm ent in which we and succeeding generations must live. the rol e which forestry plays in the economic developm ent of the rsa has long been recognised, but its other major function the stewardship if green landscapes may be overlooked by those who are obsessed with immediate material gain. apart from their task of assuring the supply of timber and other forest produce for their country, foresters hav e a duty to preserve vegetation and oth er natural ass ets on all th e land entrusted to their care which is not devoted to commercial plantations. these natural assets are not preserved purely for their own sake. conservation is for people. conservation costs money and the taxpayer who pays for it naturally wants to see where his money goes and enjoy the fruits of conservation, and for this reason there is a close tie between conservation and outdoor recreation. in som e parts of southern africa nature conservation is mainly orientated to the preservation of indigenous wildlife in gam e parks. many wild animals are dangerous, and visitors to the parks can view them only from motor vehicles or protected lookouts. there is clearly an imperative need for nature reserves in which the public can move without restriction beyond that which is necessary to pres erve the natural wildness. in europe and america forests constitute a significant proportion of nature r eserves and they are managed by foresters to preserve and increase their scientific, aesthetic and recreational valu es. the establishment offorest res erves in the rsa in which the accent is on flora rather than fauna , where visitors can walk and ride, picnic and 203 camp, has to date perhaps not received the attention it deserves because foresters have to some extent been pre-occupied with timber production. nature conservation combined with outdoor recreation and the promotion of tourism, is an aspect of land utilisation that is receiving increasing attention. in natural forests, controlled exploitation need not be excluded and open areas can even be made available to farmers for grazing in times of drought. in this way effective multiple use of the land can be achieved while still retaining public enjoyment of recreation and amenity values as an important object of management. the recreational use of state forest land in the rsa has been sporadic and unplanned for many years. there was a tendency to regard it as a thing to be discouraged. the formulation of a definite policy for facilitating and encouraging outdoor recreation in the state forests was to a large extent the outcome of a visit to the rsa in 1962 by prof c f brockman of the school of forestry and natural resources of the university of washington, who expressed the following views in reporting on his tour: " ... it is quite evident that in addition to wood production your forests , too , can become of increasing importance in outdoor recreation for the following reasons: (i) their scenic beauty and predominantly mountain settings. (ii) accessibility to motorists along a number of main roads and the possibility of opening forest roads to the public. (iii) the possibility of developing a wide variety of interests and activities for visitors in certain areas." the cons ervation role offorestry can be best appreciated if we go back to the original meaning of the word forest. it is derived from the latin ''foris'' meaning " outside" andforestis in the latin of the middle ages came to mean an unenclosed area lying outside the boundaries of villages or parks. originally, therefore, forest referred to all uncultivated or untended land, which in europe would in the main have had trees growing on it, but would also have included scrub, wasteland and what we call veld and savannah. nowadays we think of a forest as a plant community in which a fairly dense growth of trees is dominant. terms such as "high forest", "scrub forest" and " savannah" are used, but those who think of forestry primarily as a commercial undertaking are inclined to envisage a forest essentially as a regimented "plantation" with trees "lined up and lopped and drilled" to create a mechanistic, artifically-maintained sawlog factory, rather than an independently viable association of plants and animals a biocoenose adapted to the environment a forest ecosystem. the direct value of forests as a source of forest produce and the indirect benefits flowing from their amenity values, are both important, 204 but the emphasis on one or other type of benefit has varied because of differences in the socio-economic conditions of the country in the historical period concerned. in ancient times the forest was a refuge for people from their enemies and a shelter from inclement weather. it may also have harboured rogues and criminals. at a later stage many forests were reserved for hunting and pleasure. the direct value of forests as sources of timber and other products did not exceed the indirect values until the industrial revolution sharply increased the demand for timber from the end of the 18th century onwards. the continuous growth of industry since then and the pressure of expanding population are still causing increased demands for timber and other forest products, but in those areas such as western europe and the eastern united states where industrial civilisation has attained its greatest development, the accessory benifits of forest land have come to be accepted as of paramount importance. governments have realised that there is much more to forestry than the establishment and management of plantations for immediate profits, the exploitation of natural forests and the operation of sawmills, pulpmills and other wood-processing plants. the practice of forestry also includes the preservation and management of water catchments, the conservation of nature reserves in natural forests and other categories of land and the development of outdoor recreation facilities. foresters stabilise drifting sands and give expert advice on the choice of trees for woodlots, shade, shelter and ornament on farms and in parks and gardens and along streets in urban areas. the maintenance of adequate green areas is also an essential aspect of the forester's task. you may ask what the department of forestry has done in the rsa to conserve the natural vegetation. the department of forestry is probably best known to the public for the plantations of exotics, mainly pines, which it has established for the production of timber. it is perhaps not so widely realised that the department was the first body in the rsa to concern itself with conservation of the country's indigenous flora. the first step taken to protect the indigenous forests of the southern cape from the destructive exploitation then taking place, was the closure of these forests by the colonial administration of the cape in 1846 at the instance of its forest service. these forests were subsequently re-opened to controlled exploitation but with the passage of time control was intensified by successive legal enactments culminating in 1939 with the de-registration of the remaining woodcutters who were granted state pensions in lieu of their right to fell and market timber from these forests. since that time all indigenous forests belonging to the state have been managed strictly on a conservation basis. active research is being undertaken with the object of rehabilitating and consolidating these forests. legislation aimed at the acquisition of suitable privately-owned indigenous forests has also been enacted. ifforest officers had not fought with dedicated determination to preserve the indigenous forests and if 205 alternative timber resources in the form of plantations had not been established, it can be accepted that very little of these forests would remain today. provision is made in the forest act for the protection of any tree species or forest by proclamation, and a list of protected trees, which may not be felled without the authority of the minister, has been drawn up and published. several private forests in the southern cape have been likewise protected. included in the area controlled by the department are 30 nature reserves comprising some 7 500 ha, which are managed for the conservation of specific rare plant species or ecosystems. the department's main conservation operations, however, are conducted in the mountain catchment reserves which exceed a million hectares in extent. they are managed primarily for the optimum yield of clear water, but this objective is fully compatible with the conservation of the flora and the maintenance of a favourable habitat for wild life. the area of mountain ca tchments is being steadily increased by the acquisition of suitable land . an expanding staff of conservation officers which includes plant and animal ecologists, is employed on research aimed at the establishment of the most effective methods of management for maximum yield of pure, silt-free water in conjunction with protection of the fauna and flora . rare and endangered plant and animal species receive special attention. res earch is also being undertaken into the re-introduction of animals, such as eland, and other antelope into state forests where they formerly lived but have long since disappeared. special conservation research projects on state forest land are being conducted on behalf of the d epartment by universities and provincial nature conservation departmen ts. in recent years there has been a growing awareness, in the rsa as in other western countries, of the need to conserve primitive environments where people of this and future generations can study ecosystems virtually unaltered by man; where they can enjoy landscapes unscarred by roads, railways or power lines; where they can find peace and solitude; and where they can pit their own unaided resources against the forces of nature. although all demarcated forests enjoy a degree of entrenchment under the forest act of 1968, there was a need for the special preservation of selected areas which had retained their original wild character before they too would be changed by technological progress. the forest act was accordingly amended in 1971 to empower the minister, on the recommendation of the national monuments council, to set aside state forests or portions of state forests as wilderness areas for the preservation of forests and natural scenery. areas so proclaimed cannot be alienated nor can any rights be granted over them except with the approval of the senate and the house of assembly. five wilderness areas, totalling an area of nearly 185 000 ha, have already been 206 proclaimed and a number of others, making up an even larger area, are under consideration . to make conservation areas accessible to people, th e department is engaged in the d evelopment of a system of hiking trails on the lines of the appalachian trail in the eastern united states and similar trails in other countries. the south african trail with its supplementary routes is intended ultimately to stretch continuously from the cedarberg in the western cape along the mountains following the southern and eastern coastlines to terminate in the soutpansberg in the northern transvaal. the trail is not confined to state forest land and it is controlled by the national hiking way board on which state departments, provincial administrations, the national parks board and mountain and other outdoor clubs are represented. the system consists basically of hiking trails and walks. hiking trails are longer than walks and include overnight facilities such as huts, shelters and sites for tents. hiking trails are more strenuous than walks and cater for those who wish to carry a backpack or rucksack through relatively undeveloped lands with overnight shelters. walks provide leisurely outings through selected environments lasting a few hours and do not include overnight facilities. the following sections have been completed and are open for public use: the fanie botha hiking trail from near sabie to beyond graskop; the soutpansberg hiking trail along the soutpansberg eastwards from above louis trichardt; and the elephant walk in the diepwalle state forest near knysna. three other sections are due for completion by f ebruary 1977. the national parks board ' s spectacular otter hiking trail along the tsitsikama coast from stormsrivier to grootrivier will also form a link in the system. since the fanie botha hiking trail was opened in 1973, usage has increased and it is expected that there will be regular growth in the number of hikers using the already completed sections. it can be confidently expected that the time will come when a significant proportion of the active population in the more thickly populated areas will be attracted to these gateways to the un trammelled outdoors. the trails and walks can bring large numbers of the public, and particularly the impressionable younger segment, into closer contact with nature and all her wonders. this cannot fail to foster the cause of conservation in the rsa and to bring into many lives a deep appreciation of the need to preserve the green heritage which is their birthright. the department makes a major contribution to the conservation of a number of the country's veld types on its mountain catchment reserves. one of its most important functions is the permanent protection of the cape fynbos which comprises one of the richest and most divers e floras in the world. rapid agricultural and industrial development has resulted in the sacrifice of many hectares offynbos. were it not for the protection 207 by the department of its mountain catchments, the flora in these areas would be sadly depleted. the mountain catchments provide a last refuge for many of south africa's most beautiful flowering plants and several rare and endangered species. through d edicated research and farsighted conservation planning, excellent progress has been made with the rehabilitation of many endangered species . conservation offauna also has a high priority on all state forests, and good progress has been made in this direction. it is not generally realised that plantations of exotic species also provide excellent cover for game, which has increased spectacularly in numbers on many state forests. the periodic burning of firebreaks provides grazing for the game which finds sanctuary in the plantations. large sums of money are spent on the eradication of undesirable alien vegetation growing on state forest land. the problem presented by the intrusion of aggressive exotic species is daunting in its magnitude. it can be tackled successfully only by the application of co-ordinated national effort backed by adequate finance. the matter is receiving attention at the highest levels and continuous research is directed at finding cheaper and more effective methods of control. the valuable work being done by forestry bodies in the reclamation of driftsands along the south african coastline is well known. in this we are following the example of denis the first of portugal, who pioneered the reclamation of coastal dunes 500 years ago by planting them with the maritime pine. the timber from these plantations was used in shipbuilding. reclaimed driftsand has made valuable residential and industrial development possible, especially in the expansion of the cities of cape town and port elizabeth. casuarina species have also been used in the reclamation of coastal drift sands in the summer rainfall area. although progress made by the department in all aspects of nature conservation in the rsa is impressive, much still remains to be done. the scale and scope of the work is being expanded as rapidly as the recruitment of trained conservation staff and availability of funds permit. although the rsa is fortunate in having large areas of open veld and mountain, the threat posed by the encroachment of the growing population and expanding industries is very real. pollution of the atmosphere and water supplies, and indeed of our whole environment, is a danger we dare not ignore simply because it is not as immediately threatening to us as it is in some of the more intensively industrialised countries. the fact must be faced pollution is no longer a problem peculiar to the densely-populated, highly-industrialised countries. it has become a universal problem and we in the rsa must join in the fight against it now. we all have a responsibility to playa full part in exercising the stewardship if the green landscape. let us take timely and effective action to preserve the land entrusted to our care by meticulously planning the conservation of water sources, soil and vegetation, and providing 208 facilities for outdoor recreation to attract tourists as well as our own people out into the open to enjoy pure air, sparkling waters, undisturbed vegetation and protected wildlife. may i conclude by asserting that the planting of trees has a rightful place in these plans. trees are essential to satisfy the national demand for forest products but they can also beautify and conserve the south african environment. 209 page 1 page 2 page 3 page 4 page 5 page 6 page 7 filelist convert a pdf file! koedoe 19: jj3-144 (976) a survey of the mammals occurring in the golden gate highlands national park i. l. rautenbach transvaal museum p.o. box 413 pretoria 0001 abstract this paper reports on a survey of the mammals of the golden gate highlands national park, republic of south africa. fifty-seven species are mentioned, the majority recorded through material or sight records. those species which may occur in the park, as deducted from their overall distribution ranges, or from other indirect observations such as spoor or droppings, are considered as well. habitat preferences are mentioned wherever possible, and the conservation status and relocation histories of the antelope species are quoted. introduction the main object of this study is to provide a checklist of mammals for the golden gate highlands national park, republic of south africa, but stems from an interest in the taxonomy and zoogeography of the mammals of southern africa. these interests can only be successfully pursued through intensive and continuous collecting. when the late dr. n.]. van der merwe, and later his successor dr. g. de graaff, prompted myself and my predecessor mr c. g. coetzee to produce such a checklist for this park, we obligingly agreed without hesitation. this park in the northeastern orange free state on the border of lesotho, was proclaimed in 1963, primarily for its scenic beauty. six farms were dispossessed, comprising a total area of 4 792 ha of mountain sourveld (van rensburg 1968). the park constitutes the upper catchment area of the little caledon river, and varies in altitude from 1 800 m at the riverbed to 3 000 m at the peak of rhebokkop. the watershed between the orange-and vaal river systems is at the eastern boundary of the park. the national parks board of trustees, in agreeance with its conservation policy, subsequently reintroduced the game animals that long ago disappeared from the area as a result of human settlement and agricultural interests (liebenberg 1964; penzhorn 1971) . for obvious reasons the board could not reintroduce some of the 133 former residents such as lion, wild dog, hippopotamus and elephant. primarily as a result of pressing conservation problems in other national parks, very little research has thus far been conducted on the fauna and flora of the golden gate highlands national park. roberts (1969) pioneered research within the park by conducting a botanical survey. material and methods in aid of this survey, the area has been visited three times, i.e. for a ten-day period during july 1965, as well as two additional ten-day periods during may and november 1969. the species discussed in the text include not only those whose presence have been confirmed through sight or material records, but also those whose occurrence are suspected through circumstantial evidence such as general species range, or spoor and faeces encountered in the park. this latter group comprises predominantly mammals in the medium-size range which migrate or wander mostly unnoticed over great distances. specimens of only those species still presenting taxonomic problems were collected, being without exception in the smallermammal species range. specimens were procured through the normal mammal-collecting techniques, which need no elaboration here as it has been extensively described in the literature. trapping was done predominantly in those area offering the best habitat. the animals collected were prepared as standard study-material and have been incorporated in the permanent collection of the transvaal museum. results the following species have been noted to occur, or may occur in the park: erinaceusfrontalis smith, 1831 hedgehog krimpvarkie although as yet not recorded from the park, it almost certainly occurs here, since suitable habitat is available, and since the park falls within the known range of the species. it has been recorded from the nearby reitz and senekal districts (lynch 1975). myosorex varius smuts, 1832 forest shrew bos skeerbek a large series of 17 males and 21 females have been collected, mostly on the grassy slopes of bakenkop. a few specimens were also procured in dense grass on the riverbanks near wilgenhof. 134 crocidura jlavescens i. geoffroy, 1827 red musk shrew rooi skeerbek although this species has never been recorded from the o.f.s., it is almost certain to occur in the park, as it falls within the species range and affords suitable habitat. faeces, presumably of this species, was found in old rock walls, but attempts at trapping animals were unsuccessful. crocidura cyanea duvernoy, 1838 reddish-grey musk shrew rooigrys skeerbek a single specimen was procured in dense grass along a stream. amblysomus hottentotus a. smith, 1829 hottentot golden mole hotnotkruipmol four females were collected during november 1969 under willow trees on the banks of the small caledon river. at this locality this species coexists with the rodent mole (cryptomys hottentotus lesson, 1826) in the same area, albeit in separate tunnel systems. on the surface the tunnel systems of these two unrelated fossorial animals cannot be distinguished. the substrate was of a black clayish type . two of the females were reproductively active. chlorotalpa sclateri broom, 1907 sclater's golden mole sclaterse kruipmol no material has as yet been collected within the park, but three specimens in the transvaal museum collection have been collected from the n'earby clocolan district (2827dc). the characteristic shallow subsurface tracks of this species have been found in the low-lying areas of the park. pipistrellus kuhli n atterer, 1817 kuhl's pipistrelle kuhlse vlermuis recorded by lynch (j 975) from areas adjacent to the park. the species is as a consequence almost certain to occur within the boundaries of the park. eptesicus capensis a. smith, 1829 cape serotine kaapse dakvlermuis lynch (op . cit.) recorded this species within 25 km east of golden gate, and it is therefore not unlikely that it will in time be recorded from within the park. papio ursinus kerr, 1792 chacma baboon kaapse bobbejaan a fairly large troop has regularly been observed in the park on the mountain slopes and crests. 135 lepus saxatilis f. cuvier, 1823 scrub hare kolhaas a single specimen was collected during november 1969. hares of the genus lepus are not at all common in golden gate. lynch (1975) unfortunately did not verify the identification of this specimen while extracting transvaal museum records for his paper, and consequently incorrectly cited it under lepus capensis linnaeus, 1758. lepus capensis linnaeus, 1758 cape hare vlakhaas it is not impossible that this species is to be found at golden gate, but no material exists from the near vicinity of the park to substantiate such a speculation. lynch (1975) claims the presence of this species in the park and vicinity, but apart from a misidentified specimen recorded, the remainder of his records are non-material. he did not state whether the latter are from roadkills or sightings. it is of course impossible to distinguish between l. capensis and l. saxatilis in the wild state. pronolagus rupestris a. smith, 1834 smith's red hare smith se rooihaas two specimens, a male and a female, were collected in the hills behind glen reenen camp during november 1969. the female was lactating. judging from the number of toilet-sites found, so characteristic of the genus, this species is relatively abundant in the park. the measurement parameters quoted by petter ( 197 i) in his identification key are not clear, and as a consequence the key and taxonomic approach of ellerman, morrison-scott and hayman (1953) is followed here. cryptomys hottentotus lesson, 1826 common mole rat hotnot grysmol three specimens were collected in the park to date. this species occurs in the low-lying areas as well as on the higher platos and hillslopes, indicating a wider habitat tolerance than the golden moles. hystrix africaeaustralis peters, 1852 cape porcupine ystervark although this species has never been observed during collecting trips, its presence has been confirmed by several quills found in the veld, particularly at the bases of rockfaces on the lower hillslopes. pedetes capensis forster, 1778 springhare springhaas springhares have never been observed during night-hunting operations, but they almost certainly occur here. lynch (i 975) reports sight records from the near vicinity of the park . 136 graphiurus murinus desmarest, 1822 forest dormouse boswaaierstertmuis although the vernacular name of this insectivorous rodent denotes its preferred habitat, it is also to be found in rocky places, as for instance the golden gate material. all three specimens were trapped amongst rock debris at the foot of the sentinel. the only female in the series was pregnant upon collection during november; four foetuses, one in the left and three in the right uterus horn. aethomys namaquensis smith, 1834 namaqua rock rat namakwalandse klipmuis this is an abundant inhabitant of the rocky mountain slopes of golden gate. aethomys chrysophilus de winton, 1897 african bush rat afrikaanse bosrot not recorded through this project. lynch (i 97 5) claims its presence in the near vicinity of this park, which entails a 100 km westwards extension of the known range as defined by davis (1974). however, the streambanks lined with ouhout leucosidea sericea as found in the park, do not appear to offer suitable habitat when compared with the bushveld conditions selected over the rest of the species range. leggada minutoides a. smith, 1834 dwarf mouse dwergmuis a series of four was collected in the long grass near the brandwag camp. mus musculus linnaeus, 1758 house mouse huismuis a single specimen was procured in the non-white compound. praomys natalensis smith, 1834 multimammate mouse vaalveldmuis a common rodent in the denser grass oflow-lying areas. rattus rattus linnaeus, 1758 house rat huisrot several specimens were trapped in the living quarters of the bantu staff of the park . rhabdomys pumilio sparrman, 1784 striped mouse streepmuis a common rodent in the park, especially in dense tall grass. according to coetzee (in litt.), it was also found to construct its grassnests in old rockwalls up to almost a meter above ground level, which is a deviation from the normal behaviour. 137 dendromus melanotis smith, 1834 grass climbing mouse grassklimmuis a single specimen was collected in long grass along the road near the brandwag camp. dendromus mystacalis heuglin, 1863 lesser climbing mouse kleiner klimmuis four specimens were collected, all from dense tall grass either along the road , the river or the base of a hill at wilgenhof. one specimen had the remains of insects in its stomach. this is a new record for the orange free state as the result of a westwards range extention along the 29° latitude . malacothrix typica smith, 1834 large-eared mouse grootoormuis recorded by lynch (1975) in the bethlehem/kestell districts, near the park . mystromys albicaudatus smith, 1834 white-tailed rat witstertrot as yet not recorded from the park itself. it is essentially a grasslandplains species, but may in time prove to be a resident of golden gate as it has been recorded from the nearby districts (davis 1974; lynch 1975). tatera brantsi a. smith, 1834 highveld gerbil hoeveldse nagmuis several colonies were found in sandy patches oflow-iying areas. otomys irroratus brants, 1827 vlei rat vleirot a common rodent of the grassy areas near water. somewhat atypically, the species was also recorded on the lower hillslopes, especially where the ground is soggy at the bases of rocks. coetzee collected two specimens with pure-white hindquarters in the park during a museum collecting trip in 1965. according to him (in lilt.) this is the only instance of partial albinism known in this species south of zaire. note: without examining the specimen, lynch (1975) accepts as authentic the identity of an otomys specimen from golden gate, housed in the tm collection, but misidentified as o. unisulcatus. the unfortunate result is that he plots the occurrence of o. unisulcatus at golden gate, some 200 km out of range (davis 1974) in completely atypical habitat. i examined the specimen in question, and found it answers to the description of o. irroratus. consequently lynch's (1975) record of o. unisulcatus from golden gate is rejected here. 138 otomys sloggetti thomas, 1902 rock karoo rat/ice rat klip karroorot/ysrot this species certainly occurs in the park, but unfortunately no substantiating material could as yet be collected. the typical otomys faeces have, however, been found higher up on the mountain slopes in typical o. sloggetti habitat. vulpes chama a. smith, 1833 silver fox silwervos not recorded during our visits, but as this is a widely distributed species in the orange free state, individuals will undoubtedly occur in the park from time to time. canis mesomelas schreber, 177 5 black-backed jackal rooijakkals an individual was encountered during night-collecting operations. aonyx capensis schinz, 1821 cape clawless otter groototter the spoor and characteristic droppings of this species was found along the river as well as at the big dam in the game camp. lutra maculicollis lichtenstein, 1835 spotted-necked otter klein-otter the presence of this species in the park could as yet not be confirmed. since it has been reported from the nearby districts (lynch 1975), a resident population, or at least migrants through the park, is very likely. jctonyx striatus perry, 1810 striped polecat stinkmuishond individuals have been observed in the park, but as yet none could be collected. a roadkill is reported by coetzee (in litt.) on the main road near the park entrance. genetla trigrina tigrina schreber, 1776 large-spotted genet grootkolmuskejaatkat two specimens were live-trapped in the shrub along watercourses. after deliberation with c. g. coetzee and j. pringle (pers. comm.) these specimens were assigned to this subspecies, in difference to lynch (1975) who recognizes only g. genetta linnaeus, 1758 in this area and the rest of the orange free state. consequently this constitutes a new record for this province . cynictis penicillata g. cuvier, 1829 yellow mongoose geelmeerkat no specimens of this species were collected, but a colony to the east of the game camp has often been observed. 139 herpe5tes pulverulentes wagner, 1839 cape grey mongoose kleingrysmuishond certainly the most common carnivore in the park, and several were livetrapped of which two were prepared as specimens. they seem to prefer the wooded banks of the various streams flowing through the park. a female collected during november was lactating. ichneumia albicauda g. cuvier, 1829 white-tailed mongoose witstertmuishond lynch ( 1975) reports this species from areas adjacent to golden gate. as the park offers suitable habitat this animal may in time be shown to occur here as well. alilax paludinosus g. cuvier, 1777 marsh mongoose kommetjiesgatmuishond the spoor and faeces of this animal was found, but it was not collected or seen during our visits. protele5 cri5tatus spaarman, 1783 aardwolf maanhaarjakkals this species was never encountered, but spoor seen in the park was probably that of this animal. the fact that the aardwolf is widely distributed in the orange free state renders it very likely to be present in the park. felis caracal schreber, 1776 caracal rooikat spoor were found on several occasions, while a former ranger has seen a caracal in the park. felis lib,yea forster, 1780 cape wild cat vaalboskat several of these animals were seen during night collecting trips, but unfortunately none were collected, procavia capensis pallas, 1766 cape dassie klipdas three specimens were collected, of which one was heavily infected with lungworms. dassies are particularly abundant in golden gate. equus burehelli gray, 1824 burchell's zebra bontkwagga the burchell's zebra were reintroduced from the kruger national park (penzhorn 1971). the population has settled down well and is expanding (national parks board of trustees annual report 49: 197417 5). 140 phacochoerus aethiopicus pallas, i 766 warthog vlakvark reintroduced from the hluhluwe game reserve (penzhorn 197 i), but the animals have apparently not adapted very well. syncerus calfer sparrman, 1779 cape buffalo kaapse buffel reintroduced from the addo elephant national park (penzhorn 197 i) . taurotragus oryx pallas, i 766 eland reintroduced from the kalahari gemsbok national park, addo elephant national park, and the willem pretorius game reserve (o.f.s.) (penzhorn 1971 ). the population is slowly expanding (national parks board of trustees annual report 49). redunca arundinum boddaert, 1785 reedbuck rietbok reintroduced from northam (tv!.) (penzhorn 197 i), but these animals are adapting poorly to their new environment. rrduncafulvorufula at:zelius, 1815 mountain reedbuck rooiribbok reintroduced from luckhoff (o.f.s.) (penzhorn 197 i). population numbers have not increased significantly since 1971 (national parks board of trustees annual reports 46-48). alcelaphus buselaphus caama g. cuvier, 1804 cape hartebeest rooihartbees reintroduced from the addo elephant national park, kalahari gemsbok national park, willem pretorius game reserve (o.f.s.), rooipoort estates (kimberley, c.p.), and setlagodi (c.p.) (penzhorn 197 i). only three have survived since 1971 and they are not reproducing (national parks board of trustees annual reports 46-49). damaliscus dorcas phillipsi harper, 1939 blesbok reintroduced from the mountain zebra national park, willem pretorius game reserve (o.f.s.), van riebeeck nature reserve (pretoria, tv!.), kestell (o.f.s.), petrus steyn (o.f.s.), pietersburg (tv!.), and nduli national reserve (umtata, transkei) (penzhorn 197 i). the population has settled down very well and is rapidly expanding (national parks board of trustees annual reports 46-49). 141 connochaetes gnou zimmermann, 1780 black wildebeest swartwildebees reintroduced from the willem pretorius game reserve (o.f.s.), odendaalsrus (o.f.s.), and makwassie (tvl.) (penzhorn 1971). the population is viable and expanding rapidly (national parks board of trustees annual reports 46-49). antidorcas marsupialis zimmermann, 1780 springbok reintroduced from the bontebok national park, kalahari gemsbok national park, mountain zebra national park, and cookhouse (c.p.) penzhorn 197 i). the population is rapidly expanding (national parks board of trustees annual reports 46-49) . the animals were reintroduced from the ranges of two subspecies, i.e. a. m. marsupia/is from the republic and a. m. hofmeyri thomas, 1926 from the kalahari gemsbok national park. the present golden gate population therefore could elucidate the hitherto unsettled state of subspeciation in this species. ourebia ourebi zimmermann, 1783 oribi oorbietjie reintroduced from grey town (natal) (penzhorn 1971). births have been recorded (national parks board of trustees annual report 48) and the population seems to be thriving. raphicerus campestris thunberg, 1811 steenbok the present day population is endemic (penzhorn 197 i), but its status is unknown. peiea capreo/us forster, 1790 grey rhebuck vaalribbok the existing population is also endemic (penzhorn 1971), and appears to be stable although exact numbers are not known. acknowledgements i wish to express my gratitude to the national parks board of trustees for permission to collect material from the golden gate highlands national park, as well as for accommodation provided . my sincere thanks are due to mr a. van zyl, the ranger of the park, for his assistance and advice. the late dr nico van der merwe initiated the project, and later dr g. de graaff encouraged the completion of the work, and rendered advice and assistance whenever needed . this project was carried out and the results are published with the permission of the director of the transvaal museum. 142 references davis, d. h. s. 1974. the distribution of some small southern african mammals (mammalia:insectivora, rodentia). ann. transvaal mus. 29(9):135-184. ellerman,j. r., t. c. s. morrison-scott and r. w. hayman, 1953. southern african mammals 1758 to 1951: a reclassification. trustees of the british museum. liebenberg, l. c. c. 1964. die grotere soogdiere wat vroeer dae voorgekom het in die omgewing van die golden gate hooglandpark. koedoe 7 :99-104. lynch, c. d. 1975. the distribution of mammals in the orange free state, south africa. navorsinge van die nasionale museum 3(6): 109139. penzhorn, b. l. 1971. a summary of the re-introduction of ungulates into south african national parks (to 31 december 1970). koedoe 14: 145-159. petter, f. 1971. order lagomorpha. in: meester, j. and h. w. setzer (eds.) 1971. the mammals of africa: an identification manual . washington, d.c . : smithsonian institution press. roberts, b. r. 1969. the vegetation of golden gate highlands national park. koedoe 12:15-28. van rensburg, a. p. j. 1968. golden gate die geskiedenis van twee plase wat 'n nasionale park geword het. koedoe 11 :83-138. 143 page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 page 10 133.pdf page 1 filelist convert a pdf file! koedoe 19: 177-178 (1976) additions to the check-list of birds of the addo elephant national park b. l. penzhorn and p. f. van straaten addo elephant national park private bag x6027 port elizabeth 6000 in his original check-list of the birds of the addo elephant national park, liversidge (1965) recorded 120 species. in a subsequent publication six additional species were reported from the park (penzhorn and morris 1969). a further seven species are reported here, increasing the species total for the park to 133. the seven additional species recorded are listed below. the numbers denote those used by mclachlan and liversidge (1957) in roberts birds oj south africa and the nomenclature follows the check list oj the birds oj south africa (s.a.o.s. list committee 1969). 47 white-breasted cormorant phalacrocorax lucidus witborsduiker (lichtenstein) white-breasted cormorants were first reported from caesar's dam by lockhart (1968a) in july 1967. they have subsequently been recorded regularly at the dam. 52 african darter slanghalsvot.l anhinga tufa (lacepede et daudin) three darters were seen at caesar's dam by lockhart (1968b) in july 1967. 133 black eagle witkruisarend aquila verreauxi lesson a single bird was seen soaring above the elephant enclosure in june 1972. this was probably a vagrant from the suurberg range north of the park where black eagles are resident (skead 1967). 343 red-chested cuckoo cuculus solitarius stephens piet-my-vrou red-chested cuckoos have infreqeuntly been heard calling during the early summer months. 352 didric cuckoo diedrikkie chrysococcyx caprius (boddaert) didric cuckoos have frequently been heard calling during the summer months. 177 356 burchell's coucal centro pus superciliosus hemprich vleiloerie and ehrenberg these coucals have been seen among the reeds and rushes at caesar's dam and heard calling at the small dam east of the nature conservator's house. 373 fiery-necked nightjar suid-afrikaanse naguil caprimulgus pectoralis cuvier the calls of these night jars have infrequently been heard on moonlit nights. liversidge (1965) predicted that his total of 120 species of birds would be increased, "particularly in respect to birds of the habitats in and around caesar's dam". of the 13 species subsequently reported, six were recorded at the dam. references liversidge, r. 1965. the birds of the addo national park. koedoe 8: 41-67. lockhart, p. s. 1968a. new distributional data: 1. white-breasted cormorant. ostrich 39 :271. lockhart, p. s. 1968b. new distributional data: 1. african darter. ostrich 39: 2 71. mclachlan, g. r. and r. liversidge. 1957. roberts birds oj south africa. 2nd ed. cape town: trustees of the s.a. bird book fund. penzhorn, b. l. and a. k. morris. 1969.a supplementary checklist of the birds recorded in the addo elephant national park. koedoe 12: 106-10 7. s.a.o.s. list committee. 1969. check list oj the birds oj south africa. cape town: south african ornithological society. skead, c. j. 1967. ecology of birds in the eastern cape province. ostrich suppl. 7: 1-103. 178 page 1 page 2 reviewer acknowledgement http://www.koedoe.co.za open access koedoe african protected area conservation and science koedoe recognises and acknowledges the value and importance of reviewers in the publication process – not only in shaping each individual manuscript, but also in upholding the credibility and reputation of our journal. we are committed to the timely publication of all original, innovative contributions submitted for publication. as such, the identification and selection of reviewers who have expertise and interest in the topics appropriate to each manuscript are essential elements in ensuring a timely and productive peer-review process. we would like to take this opportunity to thank all reviewers who participated in shaping this volume of koedoe: page 1 of 1 in an 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task. margo van blerk submissions@koedoe.co.za tel: +27 (0)21 975 2602 fax: +27 (0)21 975 4635 abraham nkhata albert strydom andy drumm anna spenceley art pedersen cornelius van heerden elmarie slabbert engela de crom glen hvenegaard helen suich jacob mwitwa jasmine moreira jeff marion jon kohl keith bosak laura becerra lisa king marcella daye peet van der merwe rick rollins steven martin sue snyman susan moore willie boonzaaier willie coetzee willie hood yi-chung hsu yu-fai leung 86 filelist convert a pdf file! supplement to koedoe. 1977: 143-151. the status of wildlife conservation in botswana w von richter f a 0 wildlife ecologist department of wildlife, national parks and tourism pobox 131 gaberone republic of botswana the sparse human population and the general lack of surface water over most parts of the republic of botswana, which has hampered rapid expansion of agricultural activities into the less suitable areas in the past, have contributed to the fact that botswana still supports a varied and rich wildlife population. the long history of hunting by the local populae makes them understand and appreciate the concept of wildlife conservation and utilization and has assisted in general to implement a policy for rational conservation and utilization. the next decade will be decisive whether this laudable state of affairs will continue or whether the wildlife resource will be depleted and finally restricted only to formal conservation areas as it has happened in many other countries on the african continent. the government is fully aware of the significance of wildlife conservation and utilization and the necessity to integrate it into overall landuse planning. the environment botswana is a flat, landlocked country of approximately 569800 km2 , situated in the middle of the dry southern african plateau at an average altitude of 1 000 m. the majority of the country belongs to the south west arid zone . approximately two-thirds of the country is covered by a deep layer of kalahari sands of geologically recent wind blown origin. surface water is restricted to the limpopo watershed in the east, the kwando-linyanti-chobe system, which forms part of the zambezi system, in the north and the okavango river and delta in the north west. at one time both the latter have been largely infested by tsetse fly glossina morsitans, which inhibited settlement by man and his cattle; the okavango delta is still largely infested by the fly. the makgadikgadi pans complex formed in ge?logical times a huge lake, but is now a salina only filled to a depth of a few centimeters during times of exceptional high local rainfall. the kalahari covers two-thirds of the country, and is interspersed with numerous pans, round or elleptical depressions, and ancient river beds which emptied in wetter periods into 143 national parks and game reserves in botswana national parks game reserves proposed consolidations and new reserves roads 144 (1) existing national parks and game reserves 1 chobe national park (10 800 km2) est. 1961, elevated to national park 1968. 2 mikaelelo game reserve (360 km2) est. 1974. 3 moremi wildlife reserve (3 880 km2) est. 1965, enlarged in 1976. 4 nxai pan national park (2 100 km2) est. 1971. 5 makgadikgadi pans game reserve (3 900 km2) est. 1970. 6 central kalahari game reserve (52 800 km2) est. 1957. 7 gemsbok national park (24 800 km2) est. 1932, enlarged and upgraded. 8 mabuasehube game reserve (1 800 km2) est. 1971 to national park in 1971. 9 khutse game reserve (2 500 km2) est. 1971. 10 gaborone dam national park (5 km2) est. 1975. 15 maun educational game park (5 km2) est. 1969. (2) proposed new conservation areas 11 okavango river front. 12 tsodilo hills, aha mountains, kwihabe hills. 13 waterfowl sanctuary lake ngami. 14 kwebe hills. 16 tamafupajari pans complex. 17 pataletsabe hills and mannyelanong hills. 18 mannyelanong hill game reserve (pending gazettement). 19 shashe elephant reserve. 20 lepokola hills. 21 tswapong and mabeleapodi hills. 22 shoshong hills. 23 mokwane hills. 24 notwane limpopo area. the makgadikgadi. the soil and vegetation of these pans and river beds are of a different type than the surrounding sandveld and are of vital importance to humans, wildlife and cattle for their survival in this harsh environment. rainfall is varied and erratic, confined mainly to the summer (november to april), declining from an average of 650 mm in the north east to under 200 mm in the south west. shrub and tree savanna vegetation types are dominant over most of botswana. human population is low (approximately 1 person / km2); approximately 80% of the population is, however, confined to th e areas of better soils and more reliable rainfall in the east, and the fringes of the okavango delta, leaving large tracts of the country virtually uninhabited, save the few groups of nomadic bushman. most people have a history of pastoralism, although the ownership of the national cattle hercl is very skewed , i.e. a minority owns the majority of cattle. subsistence arable agriculture is very small and mostly confined to the more suitable soils. wildlife utilization has played and still plays a significant role in the life of the people of botswana. in times of catastrophic events such as rinderpest, foot and mouth disease or drought, the rural population has turned to wildlife as a means for support. in the rural areas wildlife still is a significant factor for the provision of protein and cash, although there is a noticeable trend away from purely subsistence hunting to commercial exploitation of the resource. 145 policy of wildlife conservation the policy for wildlife conservation in botswana has to match government's four objectives for development of the country: rapid economic growth, social justice, economic independence and sustained production. rural development and the raising of the standards ofliving in the rural areas is and integral part of this policy. policies for the conservation and utilization of the wildlife resource follows therefore from the above, and they can be summarised as such: (i) the establishment of national parks and game reserves to protect and conserve a cross section of all species and habitats occurring in botswana; (ii) complete protection of rare and endangered species; (iii) the implementation of rational and realistic conservation measures outside established conservation areas as a condition for optimum u tiliza tion; (iv) to provide adequate and relatively cheap hunting for the rural population who still depend to a significant degree on wildlife for their day-to-day living; (v) to optimise the financial returns from non-resident recreational hunting; (vi) to encourage non-hunting photographic tourism to reduce the dependence on non-resident hunting as the major revenue earner; and (vii) to educate all strata of society of the value of wildlife conservation, and that its utilization should benefit all. the status of conservation in botswana currently approximately 17% of botswana enjoys national park and game reserve status, comprising four national parks and six major game reserves (fig. 1). the national parks and game reserves system covers adequately all major habitat types in botswana, although none of these constitute ecological entitles. it is the intention of the department to consolidate some of the existing national parks and game reserves to make for better and more meaningful conservation and administration. in addition a number of new sites have been identified (fig. 1) for inclusion in the national parks and game reserves system, primarily on the grounds of their scientific, cultural or scenic values. the dichotomy of conserved areas in botswana is the result of historical and political reasons, i.e. national parks may only be established on stateland or any other land donated or bequeathed for this purpose to the president through an act of parliament. the national parks act of 1967 provides total protection to the entire environment of a park. game reserves are established under the fauna conservation proclamation of 1961 by the president on advice of cabinet after 146 consultation with the local authorities. in a strict legal sense game reserves provide only protection for vertebrate animals as no mention of any other environmental features are made; in practice, however, they also provide protection for the entire ecosystem . in 1976 it was decided that the national parks act should apply to all game reserves after their elevation to national parks status, regardless whether they are situated on state or tribal land . in this way more permanency in tenure and an uniform approach to management and development of all conserved areas will be achieved . as a number of new sites will not qualify for national park status, the introduction of a new category of reserved areas is contemplated. nature reserves will be designated those areas which warrant formal conservation status for various reasons, although they do not qualify for national parks status. certain activities will be permitted as long as they are compatible with the purpose for which the reserve was established. recognising the potential conflict between conserving an ecosystem and its development for tourism, all national parks and game reserves are being zoned in the following categories: wilderness area, low, medium and high density tourist areas; each of the four categories will be managed differently, according to their classification. the fauna conservation proclamation of 1961 makes provision for the gazetting of rare and endangered species as conserved animals which may not be hunted except in the case of defence of life and property. table i lists all currently conserved animals. the white rhinoceros is the table 1 conserved animals in botswana antbear aardwolf black-footed cat brown hyena cheetah civet giraffe hippopotamus honey badger klipspringer mountain rheedbuck night-ape oribi otter pangolin puku rhinoceros (black and white) roan antelope serval vaal rhebok yellow-spotted dassie waterbuck 147 all cranes all eagles all egrets fishing owl all flamingoes hammerkop all herons all jacanas kgori bustard all pelicans secretary bird spoonbill stanley bustard all storks all vultures only species which became exterminated during this centuary in botswana and a re-introduction programme is well under way to re-establish the species. wildlife utilization in botswana has two aspects, hunting by residents and non-residents and photographic tourism. consumptive utilization of wildlife in botswana can be separated in three categories: (i) traditional or subsistence huntong carried out by the local population and accounting for approximately 85% of the legal offtake (± 15 000 animals in 1975). the financial return to local or central government from this type of hunting is small, but the significance has to be seen in the socio-economic context of a rural population; (ii) superimposed on traditional hunting is non-resident and resident recreational hunting, which accounts for approximately 83% of all direct revenue accrued from wildlife utilization; and (iii) an industry utilising the trophies, hides and skins of wild animals is well established in botswana and benefits from the hunting activities. the most recent development is the emergence of non-hunting photographic tourism centred around the national parks and game reserves, primarily in northern botswana. currently 31 tourist operators conduct tours in botswana, ranging from the more luxury type to the "do-ityourself' arrangements. for the purpose of recreational hunting, botswana is subdivided into 40 controlled hunting areas (cha) of which presently 13 are leased as concession areas to four safari companies . the remainder of the cha's are open to booking by residents as non-residents are only permitted to hunt through a safari company. annual quotas are being set by the department of wildlife, national parks and tourism in consultation with the local authorities for each controlled hunting area. residents and non-residents pay c.h.a. booking fee (rio and r50/ week/hunter respectively) while tribesmen or stateland residents hunting in their areas are exempted from paying booking fee. after purchasing a general game licence (r50 for 4 animals for non-resident and r3 for 7 animals for a resident) the hunter may obtain supplementary licences which vary between ri to rio for a resident and r5 to r200 for a non-resident. protected species (lion, leopard, sable and eland) command the same fee both for resident and non-resident (r500, r300, r250, rl20 respectively). hunting regulations for tribesmen and stateland residents are complicated and the numbers of animals made available on a licence and fees vary considerably from area to area, due to historical and political reasons, introducing a measure of discrimination between citizens of botswana, depending where they reside. recognising this fact, the department is presently drafting regulations to introduce a unified 148 hunting system, differentiating basically only between residents/citizens and non-residents. recreational hunting, conducted through safari companies, has proved to be the most lucrative and easily to administer form of wildlife utilization in botswana, and it is the intention of the department to continue this aspect of hunting, modifying it in such a way as to optimise the returns of the country and make the best possible use of the wildlife resource. recognising the need to diversify the utilization of wildlife, encouragement is given to the development of non-hunting photographic tourism. the greatest asset botswana can offer to the wildlife viewing tourist is the unspoilt wilderness of her national parks and game reserves, and botswana therefore encourages high price/ low volume tourism. no permanent and costly tourism infrastructure is developed in the conserved areas, apart from an essential road network. development of the more luxury facilities is, however, encouraged through private enterprise outside the national parks and game reserves. wildlife utilization contributed r427 144 in direct revenue to local and central government in 1974, while the annual economic contribution of the wildlife based industry amounted to r3 ,7 million after deduction of all overheads and costs. a detailed knowledge of the ecology and biology of the habitats and species is a prerequisite for rational conservation and utilization of the wildlife resource, and since its inception the department has encouraged wildlife research. owing to the lack of finance and trained manpower in botswana, most of the research has been financed and carried out by outside research workers and donors. the department has initiated most of the projects, and co-ordinates and supervises them, in order to obtain the necessary information for the proper conservation and utilization of th e wildlife resource. the research projects range from ecological surveys of important wildlife areas, the monitoring of seasonal distribution and abundance of major wildlife species, to more detailed studies of individual species for management and conservation purposes. wildlife forms an important asset to botswana and its integration in the overall development is essential if wildlife is to survive in m ea ningful population in the country. the recently introduced tribal land grazing policy calls for a major input into landuse planning on a regional and national scale. agreement has been reached for the establishing and gazetting of wildlife management areas in which wildlife utilization will take preference over any form of landuse, and detrimental and competing landuses may be prohibited. it is anticipated that some of the areas will eventually be given over to other forms of landuse, but pending detailed assessment and evaluation for their ultimate use, their productivity is conserved. the involvement of the department in the landuse planning process is assured through the various regional and central landuse planning committees. 149 the conservation of wildlife in botswana for posterity will only succeed in the long run if the human population effected by the conservation measures accepts those as valid and reasonable. the education of all strata of the population to the value of wildlife conservation, in terms of financial as well as ethical and aesthetic benefits, is therefore of a paramount importance. the department has, with the assistance of outside donors, embarked on an education programme, aiming at the various groups of the population, and using all available media, to bring greater awareness of the importance of conservation to the people. the department is hampered in the effective execution of its duties and responsibilities by the lack of appropriate funds and trained manpower. the current worldwide financial crisis does not help to alleviate the situation in the near future and it is therefore necessary to continue as best as possible with the available resources. recognising the need for more appropriate training for the field staff to increase their efficiency and morale, it is hoped to establish in the near future a training centre in botswana, while medium level staff will continue to receive training at the college for african wildlife management in tanzania. future prospects for conservation in botsw;ma botswana is fortunately not as much plagued by overpopulation and scarcety of land as many other african countries. development during the last decade has been gradual and the ecological impact on the environment has not been too severe as a result. government is aware of the fragility of most of botswana's environment and the risks involved in large scale modification to it. various legislation has been passed in the last few years relating to the conservation and wise use of the natural renewable resources. the tribal land grazing policy attempts to arrest the uncontrolled spread oflow management cattle raising at the expense of the land and introduce forms of mangement assuring sustained production, raising at the same time the standard of living of the rural population. wildlife has been recognised as a significant factor in the economy of the country, and the necessity of its integration into the overall land use planning process is adcepted. botswana has at the present stage of the development, still most options open to make rational land use decisions, insuring that wildlife will not be restricted only to conservation areas, but continue to roam freely in many parts of botswana in the future. it is not unrealistic to expect that botswana may be one of the few countries where wildlife conservation and utilization will complement rather than compete with other, more conventional forms of land use. 150 conclusion during the last 10 years botswana has developed a viable national parks and reserve system, covering 17% of the co"jlntry and giving conservation to a cross section of all major habitats. a hunting system has been developed where the rural population enjoys cheap and generous licences and still permitting a lucrative non-resident hunting industry based on safari companies. non-hunting photographic tourism is emerging as an additional form of wildlife utilization, attracted by the wilderness the visitors can experience. there is growing recognition of wildlife as an alternative and complementary form of landuse, and the regional and national landuse planning exercise attempts to integrate wildlife . considering the limited resources available to botswana for the conservation of wildlife and its rational utilization, the results of these efforts are remarkabje. botswana has reached a stage of her development, where still many options remain open for continuous and increased efforts to conserve wildlife for future generations, as an integral part of the country's devel?pment process. lsi page 1 page 2 page 3 page 4 page 5 page 6 page 7 page 8 page 9 vol. 50 no. 1 pp. 18 21koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.za original research jozua j. viljoen 1department of nature conservation tshwane university of technology south africa 2mammal research institute department of zoology and entomology university of pretoria south africa andre ganswindt section of veterinary wildlife studies faculty of veterinary science university of pretoria south africa german primate centre department of reproductive biology germany rupert palme institute of biochemistry department of natural sciences university of veterinary medicine austria hendrik c. reynecke1 johan t. du toit 2 department of wildland resources utah state university usa william r. langbauer jr pittsburgh zoo & ppg aquarium usa correspondence to: jozua j. viljoen e-mail: viljoenjj@tut.ac.za postal address: tshwane university of technology, private bag x680, pretoria, 0001, south africa abstract during the past several years, non-invasive monitoring of steroid metabolites in faeces of elephants has become an increasingly popular technique to generate more information about the causal relationship between hormones and behaviour in both living elephant species. this is important knowledge which can be used to optimise local conservation and wildlife management by finding new strategies for better elephant population management and control. in this context, however, information about an actual involvement of the hypothalamic-pituitary-adrenal axis during assumable stressful events is still limited, especially for wildlife populations. one difficulty in discovering such information is often the lack of reliable data for hormone baseline levels. therefore, the aim of this study was to determine baseline concentrations of faecal glucocorticoid metabolites that could be expected within age classes and between seasons in african elephants (loxodonta africana) in the kruger national park (knp). a total of 374 faecal samples were collected from randomly located family herds in the southern knp between may 2002 and august 2005. the samples were analysed for immunoreactive concentrations of faecal glucocorticoid metabolites using a validated enzyme immunoassay for 3α,11oxo-cortisol metabolites (3α,11oxo-cm). all samples were grouped according to the estimated age class of the subject using a field method based on bolus diameter, and regarding the ecological season collected. no significant differences in faecal 3α,11oxo-cm concentrations were found across age classes (h 3 = 7.54; p = 0.057), but the mean 3α,11oxo-cm concentration of samples collected in the dry season (n = 196) was significantly higher than in the wet season (n = 178) (u = 15206.50; p = 0.032), which indicates a possible physiological stress situation due to a decline in food quantity and quality. the information generated in this study represents a reliable data set for baseline concentrations of faecal glucocorticoid metabolites for elephants within the knp and can be used to measure the stressrelated effects of translocations, management actions and the impact of chosen land use activities. keywords: glucocorticoid metabolites, faecal, baseline, african elephant, enzyme immunoassay measurement of concentrations of faecal glucocorticoid metabolites in free-ranging african elephants within the kruger national park 18 2008 faecal glucocorticoid metabolites in african elephants original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 18 21 a free-ranging animal is in a state of stress if it is required to make abnormal or extreme adjustments in its physiology or behaviour in order to cope with adverse aspects of its environment or management (friend 1980). from the physiological perspective, several endocrine responses are, amongst others, involved during stress situations. the frontline hormones to overcome these situations are the glucocorticoids, which are indicators of adrenal activity and thus of stress (möstl & palme 2002). as a physiological mechanism, stress is not inherently negative (moberg 2000), however, prolonged high concentrations of glucocorticoids may decrease individual fitness (munck et al. 1984) and reproductive success (liptrap 1993), which may cascade into long-term behavioural changes. the assessment of adrenal endocrine function in vertebrates via blood samples was an often used tool in the past, but the procedure in itself can elicit a glucocorticoid response (möstl & palme 2002). in order to avoid such self-induced stress responses, the use of non-invasive methodologies for endocrine assessment is necessary. the collection of urine or faeces as basic material for endocrine analysis seems to be a reliable alternative to blood hormone measurements, which is demonstrated by the large number of established non-invasive methodologies for monitoring adrenal function in a variety of different mammalian species (goymann et al. 1999; wasser et al. 2000, sands & creel 2004), including elephants (brown et al. 1995, ganswindt et al. 2003). brown et al. (1995) successfully developed and validated a method to assess adrenal function non-invasively through the analysis of cortisol in urine from captive elephants, but due to the practical difficulties involved with collecting urine samples in areas with absorptive ground, this method would not be an option for regular sampling from elephants in the wild. faecal samples offer the advantage that they can easily be collected, and a few non-invasive methods have already been developed and successfully tested to enable the measurement of glucocorticoid metabolites in elephant faeces (stead et al. 2000, wasser et al. 2000, foley et al. 2001, ganswindt et al. 2003; 2005). in this respect, it could be shown that a pharmacologically induced physiological stress response, induced via administration of adrenocorticotropic hormone (acth), could be monitored in captive african elephants. although work on free-ranging african elephants has been limited, ganswindt et al. (2005) found that elevated androgen levels in free-ranging male african elephants could not be related to the age of the individual. seasonal effects however do appear to influence concentrations of faecal glucocorticoid metabolites, as foley et al. (2001) found the highest concentrations during the dry season. however, a regular sample collection at a certain time before and after a predictable stressful event is necessary to create a reliable baseline for comparison, which already underlines the importance of known baseline levels to reliably determine if any specific situation is more or less stressful than the norm. we suggest, for example, a stressful situation can be triggered in elephants by several stimuli, including environmental factors (food quality and quantity): behavioural (courtship and mating); and psychosocial causes (translocation, culling). more information on the endocrine regulation of potential stressrelated events is needed, as the effect of human and natural disturbances on elephant populations has been restricted to behavioural indicators (whyte 2003). it is still not known what constitutes a naturally occurring stressful situation for freeranging elephants, or which management actions or land use practices will result in elevated stress indicators. in this paper we provide a quantitative baseline measure of concentrations of faecal glucocorticoid metabolites, after taking age and seasonal effects into account, in order to start establishing baseline levels for elephants in the kruger national park (knp). materials and methods study area and population the study area is located in the knp and falls within the lebombo arid mountain bushveld, sweet lowveld bushveld and mixed lowveld bushveld vegetation types (low & rebelo 1996). the knp covers an area of approximately 19 000 km² and can be longitudinally divided into resistant granites in the west, succeeded by ecca shales, basalt and rhyolites in the east, that give rise to different soil types and the associated flora and fauna (venter et al. 2003). data collection was restricted to the southern knp and extended from the sabie and crocodile river thickets, sclerocarya birrea subsp. caffra/acacia nigrescens savanna, mixed combretum to the lebombo south landscapes. sampling a matriarch of a family herd (median size 12) was fitted with a vhf radio collar and this group was followed on foot so as not to bias sampling close to roads. samples were collected as soon as possible after an individual had defecated, from individuals in the collared group as well as from family herds (median size 13) located within the study area. we tried to exclude individual bias by collecting samples throughout each feeding patch used by a family herd. when there were obvious signs that a herd had detected our presence, such as a definite orientation towards or away from us, no samples were collected, in order to avoid the collection of stress-induced samples. all samples were collected between 07:00 and 11:00 am. rubber gloves were used to collect approximately 100 grams of faecal material that was then placed in a labelled plastic bag. the time lapsed between defecation and the freezing of a sample was standardised to a maximum of two hours. each sample was marked with the date of collection, a gps coordinate, and the estimated age class of the subject using a field method based on the bolus diameter as described by wimberger (2001). faecal samples were collected over the four ecological seasons as classified by zambatis (2002), namely early dry (may–july); late dry (aug–oct), early wet (nov–jan) and late wet (feb–april). this allowed us to determine, through concentrations of glucocorticoid metabolites, whether environmental factors, which change between seasons, have any possible influence on physiological stress. samples (n = 374) were collected between may 2002 and august 2005. the average annual rainfall during the study was 572 mm, which is in the known average annual rainfall range of 500–700 mm for this region, and therefore would not have unduly influenced environmentally induced feeding stress (venter et al. 2003). faecal extraction and hormone assays faecal samples were extracted according to the procedure described by merl et al. (2000). in brief, 0.5 g faeces plus 1 ml water and 4 ml methanol were vortexed for 30 minutes and centrifuged at 2000 g for 15 minutes. a quantity of 1 ml of the supernatant was mixed with 5 ml ethyl ether and 0.25 µl of a 5% nahco 3 solution, and centrifuged at 2000 g for 15 min. the aqueous phase was frozen at -20 °c overnight and then the ether was decanted and dried down under a stream of n 2 . following evaporation, the samples were reconstituted in an assay buffer and taken to assay. faecal extracts were measured for immunoreactive glucocorticoid metabolites using an enzyme immunoassay for 3α,11oxo-cortisol metabolites (3α,11oxo-cm) (möstl et al. 2002), which has previously been shown to provide reliable information on adrenocortical function in the african elephant (ganswindt et al. 2003; 2005). briefly, 50 µl aliquots of standards, quality controls, and diluted faecal extracts were pipetted in duplicate into microtiterplate wells. a total of 100 µl of biotinylated label and antiserum (raised in a rabbit against 5βandrostane-3α-ol-11-one-17-cmo) were added and the plates incubated overnight at 4 °c. following incubation, the plates 19 original research viljoen, ganswindt, palme, reynecke, du toit & langbauer jr koedoe a fr ic an p ro te ct ed a re a c on se rv at io n an d s ci en ce http://www.koedoe.co.zavol. 50 no. 1 pp. 18 21 were washed four times and 250 µl (4.2 mu) of streptavidin horseradish peroxidase conjugate added to each well. following incubation in the dark for 45 min at 4 °c, plates were washed again, before 250 µl (69.4 nmol) tetramethylbenzidine was added and plates further incubated (45 min; 4 °c). the reaction was terminated by adding 50 µl of 2 m h 2 so 4 and the absorbance measured at 450 nm (reference filter: 620 nm) with an automated plate reader. sensitivities of the assays at 90 % binding were 3.0 pg/well and intraand interassay coefficients of variation, determined by repeated measurements of high and low value quality controls ranged between 3.0 % and 12.5 %, respectively. statistical analysis the age class data, as well as the data from the ecological season, was tested for normality using the shapiro-wilks w test. the data sets were not normally distributed and subsequently subjected to non-parametric statistical methods. an anova was performed to test for the possible effect of age and season on concentrations of faecal glucocorticoid metabolite levels. the computer program statistica (stasoft. 1995) was used for all statistical analyses. all tests were two-tailed, with the αlevel of significance set at 0.05. in cases of all pair-wise multiple comparison procedures, the α-level was adjusted by applying the procedure described by holm (1979). results no significant variation in concentrations of faecal glucocorticoid metabolites (h 3 = 7.54; n = 374; p = 0.057) was found across age classes (table 1). therefore, the age classes were combined for further analysis. although no significant differences in concentrations of faecal glucocorticoid metabolites were found between the four ecological seasons (table 2), there was a statistically significant difference (u = 15206.50; p = 0.032) between the wet season (n = 178) and dry season (n = 196), after the early and late period of both the dry and wet seasons were combined (figure 1). discussion this study provides new information on baseline concentrations of faecal glucocorticoid metabolites that could be expected within age classes and between seasons in african elephants in the southern kruger national park. our results show that the variability in baseline concentrations of faecal glucocorticoid metabolites (3α,11oxo-cm levels) in african elephant faeces is dependent on seasonal changes, rather than on the age class of the subject. in the present study faecal 3α,11oxo-cm levels differ significantly between seasons, but no differences were found across age classes. although the seasonal effect found seems to be rather small, future studies using methods of faecal hormone analysis to determine the effect of a potential stressor should account for seasonal effects, especially between the wet and dry season. the fact that no significant variation in faecal 3α,11oxo-cm levels was found across age classes confirm findings by ganswindt et al. (2005), who also reported no age effects on 3α,11oxo-cm levels in a group of african elephant bulls (n = 52, age range: 18–49 years) living in the samburu and buffalo springs national reserves, kenya. in the same study, ganswindt and colleagues described a small but clear elevation in 3α,11oxo-cm levels in longitudinal hormone profiles of african elephant bulls at the end of a long dry period (ganswindt et al. 2005). a correlation between season-dependent rainfall and adrenal endocrine function (highest concentrations of faecal glucocorticoid metabolites in the dry season) was further described for female elephants by foley et al. (2001). the present study confirms a possible influence of a dry period on increased glucocorticoid excretion, because the mean concentration of faecal glucocorticoid metabolites of samples collected in the dry season was significantly higher than in the wet season. the elevation of concentrations of faecal glucocorticoid metabolites during the dry season could be an indication of increased physiological stress due to a decline in food quantity and quality. this was a suggestion already made by codron et al. (2006), who reported that the percentage of nitrogen in elephant faeces from the southern knp showed a dramatic increase from the dry to the wet season. an elevation of nitrogen in faeces is known as a useful indicator of nutritional status. we could unfortunately not investigate differences in concentrations of faecal glucocorticoid metabolites between sexes due to safety considerations for the observers (distance away from the herd), and visual impairment caused by vegetation structure. however, further studies should examine the difference in 3α,11oxo-cm levels between the sexes after intense sampling from cows and bulls within both seasons, because sex could be a co-factor for the variability in baseline concentrations of faecal glucocorticoid metabolites. greyling (2004) found that during periods of resource limitation, males showed significantly lower levels of faecal minerals, together me a n me a n ±s e me a n ±s d d ry w e t w e t a n d d ry s e a s o n 2 0 4 0 6 0 8 0 1 0 0 1 2 0 1 4 0 1 6 0 fa ec al g lu co co rti co id (n g/ g) figure 1 box-plots of grouped concentrations of immunoreactive faecal glucocorticoid metabolites (ng/g) for free-ranging family herds of elephants in knp in the dry and wet seasons from may 2002 to august 2005. table 1 n, mean, standard deviation (concentrations of faecal glucocorticoid metabolites (ng/g ww)) in the four ecological seasons (age classes separate) for free-ranging family herds of elephants in knp from may 2002 to august 2005. season adults sub-adults juveniles n 3α,11oxo-cm conc. (mean±std) n 3α,11oxo-cm conc. (mean±std) n 3α,11oxo-cm conc. (mean±std) early dry 68 85.19, 47.08 50 94.04, 44.23 28 91.29, 56.22 late dry 23 79.70, 70.68 17 88.71, 69.90 10 80.10, 46.53 early wet 39 72.62, 43.39 23 68.48, 38.71 14 101.79, 54.27 late wet 41 67.61, 39.48 36 82.25, 48.51 25 81.36, 59.68 table 2 n, mean, standard deviation (concentrations of faecal glucocorticoid metabolites (ng/g ww)) in the four ecological seasons (age classes combined) for free-ranging family herds of elephants in knp from may 2002 to august 2005. season individuals n 3α,11oxo-cm conc.(mean±std) early dry 146 89.39, 47.86 late dry 50 82.84, 65.23 early wet 76 76.74, 45.29 late wet 102 76.15, 48.26 20 faecal glucocorticoid metabolites in african elephants original research a frican p rotected a rea c onservation and s cience http://www.koedoe.co.za koedoevol. 50 no. 1 pp. 18 21 with higher levels of fibre, than adult females. these results indicate that males could be maintaining diets of poorer quality than females and consequently be nutritionally more stressed than females during the dry season. further studies should also investigate the difference in concentrations of faecal glucocorticoid metabolites between elephants from different areas, which would finally allow a comparison of mean concentrations of faecal glucocorticoid metabolites across elephant management regions within the knp (whyte et al. 2003). this would aid in creating a baseline for the entire park; that would be per season, per region and between different elephant densities. finally, this information could also be applied more universally, e.g. if a significant correlation with average monthly rainfall can be shown, or whether a direct correlation exists between increasing physiological stress and vegetation damage. conclusion a validated method (eia) was applied in assessing the level of physiological stress in free-ranging elephant herds. the method has been shown to be practical and enables long-term monitoring of ethological or environmental factors without interfering with the result. acknowledgements south african national parks is thanked for allowing fieldwork in the kruger national park. dr i. whyte and prof. d. g. a. meltzer for their assistance. we are grateful to mrs a. human for the laboratory work. we thank the field rangers of the lower sabie section for providing protection during data collection. the south african national research foundation and pittsburgh zoological society provided funding. references brown, j.l., c.m. wemmer & j. lehnhardt. 1995. urinary cortisol analysis for monitoring adrenal activity in elephants. zoo biology 14: 533–542. codron, j. j., lee-thorp, a., sponheimer, m., codron, d., grant, r.c & de ruiter, d.j. 2006. elephant (loxodonta africana) diets in kruger national park, south africa: spatial and landscape differences. journal of mammalogy 87(1): 27–34. friend, t.h. 1980. stress: what is it and how can it be quantified? int j stud anim prob 1 (6): 366–373. foley, c.a.h., papageorge,s.& wasser, s.k. 2001. non-invasive stress and reproductive measures of social and ecological pressures in free-ranging african elephants (loxodonta africana). conservation biology 15: 1134–1142. ganswindt a., heistermann, m., palme, r., borragan, s. & hodges, j.k. 2003. non-invasive assessment of adrenal function in the male african elephant (loxodonta africana) and its relation to musth. gen comp endocrinal 134: 156–166. ganswindt, a., rasmussen, h.b., heistermann, m. & hodges, j.k. 2005. the sexually active states of free-ranging male african elephants (loxodonta africana): defining musth and non-musth using endocrinology, physical signals, and behaviour. horm behav 47 (1): 83–91. goymann, w., möstl, e., van’t hof, t, east, m.l. & hofer, h. 1999. noninvasive fecal monitoring of glucocorticoids in spotted hyenas, crocuta crocuta. gen. comp. endocrinal 114: 340-348. greyling, m.d. 2004. sex and age related feeding distinctions in the feeding ecology of the african elephant, loxodonta africana. ph.d thesis, university of the witwatersrand, johannesburg. holm, s. 1979. a simple sequentially rejective multiple test procedure. scandinavian journal of statistics 6:65–70. liptrap, r.m. 1993. stress and reproduction in domestic animals. ann ny acad sci 697: 275–84. low, a. b. & rebelo, a. g. 1996. vegetation of south africa, lesotho and swaziland. department of environmental affairs & tourism, pretoria. merl, s., scherzer, s., palme, r. & möstl, e. 2000. pain causes increased concentrations of glucocorticoid metabolites in horse faeces. j equine vet sci 20: 586–90. moberg, g.p. 2000. biological response to stress: implications for animal welfare. in: moberg, g.p. & mench, j.a. 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(unpublished) zambatis, n. 2002. annual section ranger reports. knp vegetation monitoring. scientific services knp. (unpublished) 21 article information author: adrian j.f.k. craig1 charlene bissett1,2 mark d. galpin2 bryan olver2,3 pat e. hulley1 affiliations: 1department of zoology and entomology, rhodes university, south africa 2kwandwe private game reserve, grahamstown, south africa 3phinda private game reserve, hluhluwe, south africa correspondence to: adrian craig email: a.craig@ru.ac.za postal address: department of zoology and entomology, rhodes university, grahamstown 6140, south africa dates: received: 22 jul. 2010 accepted: 03 dec. 2010 published: 02 mar. 2011 how to cite this article: craig, a.j.f.k., bissett, c., galpin, m.d., olver, b. & hulley, p.t., 2011, ‘the avifauna of kwandwe private game reserve, eastern cape, south africa’, koedoe 53(1), art. #1015, 5 pages. doi:10.4102/koedoe.v53i1.1015 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) the avifauna of kwandwe private game reserve, eastern cape, south africa in this original research... open access • abstract • introduction    • study area • methods • results and discussion    • past records    • current records    • single records    • species indicative of range expansion    • seasonal visitors    • re-introductions    • conservation significance • conclusion • acknowledgements • references abstract (back to top) a protected area since 1999, kwandwe private game reserve incorporates several former farms, for which past records of bird occurrences are available. no bird species appear to have been lost from the area. between 2001 and 2005, a group of observers conducted systematic bird surveys in most months, which allowed the status (resident, migrant or irregular visitor) of most bird species to be determined. at least three species have established breeding populations in the reserve over the past 10 years. of 302 species reliably recorded to date, 182 (60.3%) appear to be resident, 46 (15.2%) are seasonal migrants and 74 (24.5%) are vagrant visitors. eight vulnerable and eight near-threatened bird species have been recorded; blue crane, kori bustard and african crowned eagle have bred in the reserve. together with other state-owned and private protected areas in this region, this reserve holds a significant portion of the inland bird species recorded from the eastern cape. conservation implications: the varied thicket vegetation types of the great fish river valley support a considerable diversity of bird species. if these habitats are managed for biodiversity conservation, they can support a large component of the terrestrial avifauna of the eastern cape region. introduction (back to top) over the past 20 years there has been a significant shift in land usage in the eastern cape. several commercial farms have been converted to game ranches either for hunting or for ecotourism (palmer 2008), leading to the formation of extensive private reserves along with the expansion of state-owned conservation areas such as the addo elephant national park. this has particularly affected the vegetation type originally described as valley bushveld by acocks (1975), and now treated as a distinct biome, albany thicket, by mucina and rutherford (2006). in this paper, we present some baseline data to establish how this management change has influenced the avifauna in one protected area in the eastern cape. kwandwe private game reserve (pgr) was established in 1999 and the first lodges opened for guests in 2001. there are bird records from different sources for three distinct periods: museum specimens (1916–1960), bird club observations (1980–1999), and surveys by reserve staff and associates (2001–2005). the present study investigated four questions. 1. what bird species occurred in the area prior to the establishment of the reserve? 2. have any species been lost since then and, if so, can we explain their disappearance? 3. have new species become established and, if so, can we account for their arrival? 4. do periodic vehicle-based surveys reflect the current species diversity? figure 1:vegetation map of kwandwe private game reserve (from bissett 2007). study area kwandwe pgr (33°09’s, 26°37’e ) lies in the great fish river valley, approximately 35 km north of grahamstown in the eastern cape, south africa. the farms purchased for the reserve had previously been used primarily for stock farming, with irrigated lands bordering on the great fish river and ostrich pens on some sections. in 2005 additional land was purchased on the northern bank of the great fish river, increasing the area of the reserve to some 20 000 ha. the perennial great fish river flows through the reserve for about 25 km and all watercourses drain towards the river (figure 1).the main vegetation types represented in the reserve are great fish noorsveld and great fish thicket, within the albany thicket biome. the kwandwe pgr preserves a significant area of great fish noorsveld (mucina & rutherford 2006). at a finer scale, bissett (2007) described 11 vegetation types in the reserve, of which five cover more than 1000 ha. these form a mosaic of thicket and bush-clump types, with acacia karoo, cussonia spicata, euclea undulata, euphorbia spp., maytenus spp., pappea capensis, portulacaria afra, searsia spp. and schotia afra as the dominant larger plant species. there are also old cultivated lands, now covered by grasses, and two large dams. recent additions to the property include some valleys with dry forest, including species typical of both valley thicket and afro-montane forest (mucina & rutherford 2006), on the north bank of the great fish river, where large trees such as harpephyllum caffrum occur. the altitude ranges from 170 m a.s.l in the great fish river valley to 600 m a.s.l. on the ridges, and mean annual rainfall for the past 7 years was 394 mm and 358 mm at brandeston and krantzdrift, respectively (bissett 2007). table 1: summary of the bird groups recorded on kwandwe private game reserve since october 2001. methods (back to top) past records examined in this study included the published literature on birds of the eastern cape, the bird collection at the albany museum, records of outings by the diaz cross bird club, and an unpublished list of birds on the farm brandeston, compiled by c.a. whittington-jones. during 2001 the staff at kwandwe pgr began a systematic survey conducted on a specific day each month for the ’birds in reserves project’ (birp), coordinated by the (then) avian demography unit at the university of cape town. birp records exist for five months in 2001, for every month in 2002, for eight months in both 2003 and 2004, and for all months in 2005. these records allowed for an assessment of the birds that are likely to be resident as opposed to vagrants. we have also included more recent records (up to october 2009). during the birp surveys we followed a standard route, with surveying starting at 06:00 in summer and at 07:00 in winter. the route was designed to traverse all the different vegetation types in the reserve and to pass close by the major water bodies. the total distance was about 35 km, which was covered in 8–10 hours. we also included other sightings by rangers during the observation period; these reports typically included only large birds such as bustards and cranes, or birds of prey. between three and eight observers participated in the surveys, depending on staff commitments on the particular day. normally an open game-viewing vehicle was used; if another vehicle was used, there was always one observer standing on the back. all observers were experienced bird watchers equipped with binoculars and all species confirmed by call were included. the bird names used follow those in roberts birds of southern africa (eds. hockey, dean & ryan 2005). only standard english names are used in the text, but both english and scientific names are given in online appendix 1. results and discussion (back to top) past records specimens from the albany museum included 68 study skins from the present reserve area (collected on the farms brandeston and the fort, which both had river frontage). all but three specimens were collected between 1920 and 1927: a long-billed crombec collected on brandeston in 1916, a dark-capped bulbul from the fort collected in 1939, and a crowned hornbill from brandeston collected in 1960. the collections were made by v.e. watson on the fort and a.t. rivett-carnac on brandeston. hewitt (1931) cited rivett-carnac as one of his bird informants and mentioned a single sighting of a white-fronted bee-eater from the great fish river valley, but gave no precise locality. despite this being a biased sample in which larger birds such as waterbirds, gamebirds and raptors are poorly represented, it does provide insight into the avifauna history of the area. of the 56 species, 55 have been recorded on the reserve in the past 5 years. the only exception is the red-headed finch, which was categorised as ‘locally nomadic’ by dean (1997) and makes occasional incursions into this region according to recent records from brandeston. seven sets of eggs collected at heatherton towers (now the visitor reception centre on kwandwe pgr) in the 1930s, also formed part of the past records. the eggs represent species that currently breed on kwandwe pgr. they are listed together with the study skins in the category ’museum’ in online appendix 1. c.j. skead visited all the farms that have now been incorporated into kwandwe pgr during his research on eastern cape birds, but no specific records are mentioned in his reviews (skead 1965, 1967). the diaz cross bird club visited the farms brandeston, krantzdrift and the fort (now incorporated into kwandwe pgr) on four occasions between 1980 and 1998, during which 117 bird species were recorded within the area of the present reserve. three of these species (glossy ibis, southern pochard and cape canary) have not yet been recorded since the establishment of the reserve. club members also submitted sightings of interest to the club newsletter and we have extracted from the ‘diaz diary’ all such reports (listed under ’diaz’ in online appendix 1). one species, the european bee-eater, was reported in march 1988 but has not been recorded from the area since. there are unconfirmed records for denham’s bustard (formerly often confused with ludwig’s bustard – see herholdt 1988), sand martin (likely erroroneous identification for the brown-throated martin, formerly the african sand martin), yellow wagtail, and thick-billed weaver (the latter both from an unreliable observer). from july 1997 to june 1999, c.a. whittington-jones made regular visits to brandeston during his study of red-billed queleas at ostrich feedlots. he recorded 136 bird species on the farm, of which only the red-headed finch (sighted in may 1998 and may 1999) has not subsequently been recorded on the reserve. since the removal of the ostrich feedlots, the red-billed quelea has become an occasional vagrant in the area. table 2: bird species recorded at kwandwe private game reserve, listed in the current red data book for south africa (ed. barnes 2000). current records to date, 302 bird species have been recorded reliably on kwandwe pgr (online appendix 1). of this total, 274 species were recorded during the 45 birp surveys included in this overview, representing more than 90% of the species recorded up to 2006. the avifauna was subdivided into broad habitat classes within which the birds could be categorised as regulars, vagrants or migrants based on the monthly distribution of records (table 1).compared with three other eastern cape conservation areas where thicket is the predominant vegetation type, kwandwe pgr has the most diverse avifauna (302 species), of which 11% have been recorded only on this reserve. some 194 species have been recorded from the original addo elephant national park area of 12 000 ha (urquhart & klages 1997), excluding the recently added colchester, zuurberg and the greater addo sections towards the coast. these include habitat types such as southern coastal forest, suurberg quartzite and suurberg shale fynbos (mucina & rutherford 2006), which are absent from the other reserves. the checklist for the great fish river reserve complex records 245 species from the area west of the great fish river, which spans the former andries vosloo kudu reserve and the sam knott nature reserve (23 000 ha combined). shamwari game reserve currently covers 22 000 ha, with 257 species having been recorded. although addo elephant national park is thus the smallest of these conservation areas and no river runs through the reserve, all four reserves lie below the inland escarpment and at a similar distance from the coast. the great fish river reserve is almost adjacent to kwandwe pgr and shamwari game reserve is less than 20 km from addo elephant national park in direct line of flight. these reserves form part of a belt of conservation areas between the coast and the more arid interior and include a significant proportion of the south african avifauna (combined total = 334 species), despite none being mentioned in the first national survey of important bird areas (ed. barnes 1998). single records several species have been recorded only once. however, in all cases there are other records for single sightings in adjoining regions of the eastern cape and in some instances the timing of vagrant sightings suggests that the same individual birds could have been observed. this group includes the african pygmy goose, ground woodpecker, african black oystercatcher (sighted after a period of onshore gales along the coast), caspian tern, tawny eagle, bateleur, baillon’s crake, greater flamingo, abdim’s stork, dusky lark, chat flycatcher, chorister robin-chat, capped wheatear, white-browed sparrow-weaver and long-tailed paradise whydah. species indicative of range expansion the goliath heron, a rarity in the eastern cape in the 1980s, is now well established along coastal rivers, down the east coast (craig 2006) and on inland dams, perhaps spreading from major impoundments along the orange river.kurrichane buttonquails were not reported from the eastern cape during the bird atlas survey (harrison et al. 1997), although there are historic records (skead 1967). several recent records at different times of the year have led to suggestions of a range expansion into the eastern cape (eds. hockey et al. 2005). birds on kwandwe pgr were most often flushed from areas of thicket–grass patch habitat mosaic. in 2001 an adult bird was seen with a single, very small chick, indicating that breeding had occurred. the white-fronted bee-eater was previously regarded as a vagrant in the eastern cape (harrison et al. 1997; skead 1967). a pair was found breeding on the sundays river at colchester in 1998 (martin 1999) and since then birds have been seen at this site annually. found nesting on kwandwe pgr in january 2002, the birds were initially thought to be summer-breeding visitors, but in recent years they have been sighted in most months and may now be considered resident. african palm swifts were not recorded south of kwazulu-natal by skead (1967) nor quickelberge (1989), who reviewed data for the transkei region. nevertheless, the first records for the eastern cape date back to the 1980s in steytlerville (ward et al. 1984). this species is now regular along the coast where suitable palms are available and has also been recorded in grahamstown. a small colony has been present on kwandwe pgr at the brandeston homestead since 2005. the double-banded courser was not found in the grahamstown region during the bird atlas surveys (harrison et al. 1997), which supports vernon’s (1982) observation that coursers are rarely sighted in the eastern cape today. however, this courser has been sighted in one section of the kwandwe reserve quite regularly, in association with other birds typical of the arid interior (e.g. the rufous-eared warbler). breeding had been recorded in 2001 and 2002. the violet-backed starling is an intra-african migrant, which has been recorded breeding in the eastern cape to the east of the great fish river. in recent summers there have been an increasing number of reports from this southern extremity of its range in south africa. the scaly-feathered finch was first reported in 2006 and is now apparently a breeding resident, but its distribution is very localised within the reserve. to date there has been a single sight record of the white-browed sparrow-weaver, but nesting birds were found on a farm south-west of kwandwe pgr recently. this species has become established on farms in the queenstown district in the past 20 years and is currently expanding its range there (k. webster, pers. comm., may 2010). it colonised the mountain zebra national park from the late 1960s onwards (craig, hulley & parker 2005; skead 1966). passing largely unnoticed, the southern grey-headed sparrow underwent a major range expansion during the last century, colonising the albany district during the 1960s (craig, every & summers-smith 1987) and reaching the south-western cape by the 1990s (harrison et al. 1997). seasonal visitors barratt’s warbler, a bird from the mountain forests of the eastern cape often moves to lower altitudes in winter (vernon 1989). this is reflected by higher winter records on the coast and occasional sightings en route (craig 1986). the warbler was present in thick cover close to a game-viewing hide for several weeks in june 2003; it can easily be overlooked in areas traversed only by vehicles. a winter record of chorister robin-chat may also represent an individual moving from montane forest towards the coast.the dusky sunbird is evidently a winter visitor to the great fish river valley, with several sightings on the reserve where it often feeds on the honey thorn (lycium oxycarpum). there were also ringing records from brandeston and resolution farms in the fort brown area prior to the establishment of the reserve and later captures at aloes on kwandwe pgr and hounslow, a farm on the western border of the reserve. re-introductions along with other reserves, kwandwe pgr has been stocked with fauna which formerly occurred in the area (skead 1987) and, more controversially, with species such as giraffe (giraffa camelopardalis), which can be considered alien to the eastern cape (castley, boshoff & kerley 2001). by contrast, only two bird species have been considered for re-introduction.the red-billed oxpecker, which had been locally extinct, was re-introduced to the eastern cape from the kruger national park in 1990, with birds released in addo elephant national park, the great fish river reserve and on a private game farm in the highlands area (craig & weaver 1990). birds have subsequently also been introduced to shamwari game reserve in 1998 and there was a second release in the great fish river reserve in 2003. breeding has been reported from addo elephant national park, the great fish river reserve and shamwari game reserve, with regular sightings from properties adjoining these sites. the birds are often seen on giraffe, one of their favourite hosts throughout southern africa. currently oxpeckers are recorded in kwandwe pgr most months and over the past 2 years there have been sightings of dark-billed juvenile birds, despite no direct evidence of breeding within the reserve. the southern ground hornbill may be introduced to kwandwe in the future if the pilot project at shamwari game reserve proves successful, although this region clearly represents the edge of the natural range of this species (hulley & craig 2007). conservation significance the avifauna of kwandwe pgr includes 16 species of conservation concern, all of which are potential breeding species in the area (table 2). the cape vulture has suffered a major decline throughout the eastern cape, with many former breeding sites now abandoned (boshoff, piper & michael 2009). only occasional vagrants are now reported in this part of the province. similarly, the tawny eagle was formerly widespread in the eastern cape and until 1981 a regular breeding species. it is suspected to have suffered heavily from indiscriminate use of poisoned baits (ed. barnes 2000). the sighting in january 2005 was the first in the grahamstown district for some 15 years (craig 2005). bateleurs have seldom been seen south of the orange river over the past 50 years and a record of an adult bird at kwandwe pgr in june 2009 was a major surprise. african crowned eagles nest on the reserve and martial eagles still nest in the district. blue cranes nest on the reserve annually, while kori bustards have nested here occasionally. these ground-nesting species are, however, vulnerable to predators, particularly the black-backed jackal (canis mesomelas). cranes and bustards frequently collide with power lines (jenkins, smallie & diamond 2010) and it is thus laudable that all power cables on this reserve are now underground. the great fish river valley appears to form a conduit both for southward penetration of species typical of the arid interior, such as the red-billed firefinch, and for inland movements of species generally restricted to the coastal belt, such as the yellow weaver (recorded in the great fish river reserve, but not yet on kwandwe pgr). the thickets also provide acceptable habitat for some forest species such as the narina trogon and the dark-backed weaver. conservation of the thicket biome is currently considered a priority, since this habitat has been extensively modified by agriculture and is regarded as important in carbon sequestration. thus, the mosaic of privately managed and state-owned reserves in this region of the eastern cape should be seen as a significant contributor to conserving south african biodiversity. conclusion (back to top) although we lack a comprehensive checklist for the region prior to the establishment of the reserve, there is no clear evidence that formerly resident bird species have been lost, while at least three species (white-fronted bee-eater, african palm swift and scaly-feathered finch) have established breeding populations since the area had become fully protected. regular vehicle-based surveys by experienced observers seem to record a high proportion of the bird species that occur within the reserve. such monitoring should form a part of future management plans, in the interest of both conserving biodiversity and highlighting additional attractions for visiting tourists. acknowledgements (back to top) we are most grateful to the management of kwandwe pgr, in particular angus sholto-douglas, for supporting this project and placing vehicles at our disposal. our thanks also go to all the rangers, trainees, and other volunteers who have taken part in the surveys. brad fike (great fish river reserve) and antony collett (shamwari game reserve) kindly provided copies of unpublished checklists of the birds of these reserves, and craig whittington-jones offered us all his dated records for brandeston. a.c. and p.h. received research funding from rhodes university. c.b. received funding for research on carnivores from kwandwe pgr during the time of the study. references (back to top) acocks, j.p.h., 1975, ‘veld types of south africa’, 2nd edn., memoirs of the botanical survey of south africa 40, 1−128. barnes, k.n. (ed.), 1998, the important bird areas of southern africa, birdlife south africa, johannesburg. barnes, k.n. (ed.), 2000, the eskom red data book of birds of south africa, lesotho and swaziland, birdlife south africa, johannesburg. bissett, c., 2007, ‘the feeding and spatial ecologies of the large carnivore guild on kwandwe private game reserve’, phd thesis, department of zoology and entomology, rhodes university. boshoff, a., piper, s. & michael, m., 2009, ‘on the distribution and breeding status of the cape griffon gyps coprotheres in the eastern cape province, south africa’, ostrich 80(2), 85−92. doi:10.2989/ostrich.2009.80.2.4.831 castley, j.g., boshoff, a.f., kerley, g.i.h., 2001, ‘compromising south africa’s natural biodiversity: inappropriate herbivore introductions’, south african journal of science 97, 344−348. craig, a., 1986, ‘forest birds on rhodes university campus’, diaz diary 155, 4−5. craig, a., 2005, ‘will the tawny eagle return to the eastern cape?’, diaz diary 33(1), 11−12. craig, a., 2006, ‘goliath herons on the eastern cape coast’, bee-eater 57(2), 36−37. craig, a., every, b. & summers-smith, d., 1987, ‘the spread of the southern greyheaded sparrow in the cape province’, annals of the cape provincial museums (natural history) 16(9), 191−200. craig, a.j.f.k., hulley, p.e. & parker, d., 2005, ‘a re-assessment of the avifauna of the mountain zebra national park’, koedoe 48, 95−113. craig, a. & weaver, a., 1990, ‘the relocation of the redbilled oxpecker’, bee-eater 41(4), 58−61. dean, w.r.j., 1997, ‘the distribution and biology of nomadic birds in the karoo, south africa’, journal of biogeography 24, 769−779. doi:10.1046/j.1365-2699.1997.00163.x harrison, j.a., allan, d.g., underhill, l.g., herremans, m., tree, a.j., parker, v., et al., 1997, the atlas of southern african birds, birdlife south africa, johannesburg. herholdt, j.j., 1988, ‘the distribution of stanley’s and ludwig’s bustards in southern africa: a review’, ostrich 59, 8−13. doi:10.1080/00306525.1988.9633917 hewitt, j., 1931, a guide to the vertebrate fauna of the eastern cape province. part 1. mammals and birds, albany museum, grahamstown. hockey, p.a.r., dean, w.r.j. & ryan, p.g. (eds.), 2005, roberts birds of southern africa, 7th edn., john voelcker bird book fund, cape town. hulley, p.e.h. & craig, a.j.f.k., 2007, ‘the status of the southern ground-hornbill in the grahamstown region, eastern cape’, ostrich 78, 89−92. doi:10.2989/ostrich.2007.78.1.13.57 jenkins, a.r., smallie, j.j. & diamond, m., 2010, ‘avian collisions with power lines: a global review of causes and mitigation with a south african perspective’, bird conservation international 20(3), 152−278. doi:10.1017/s0959270910000122 martin, p., 1999, ‘first breeding record of white-fronted bee-eaters’, bee-eater 50(2), 13. mucina, l. & rutherford, m., 2006, ‘vegetation of south africa, lesotho and swaziland’, strelitizia 19, 1−807. palmer, a., 2008, ‘a study of land-use trends and land tenure changes in the makana local municipality (albany registration district) for the period 1994 to 2008’, unpublished report to ardri, university of fort hare, east london. quickelberge, c.d., 1989, the birds of the transkei, durban natural history museum, durban. skead, c.j., 1965, ‘birds of the albany (grahamstown) district’, south african avifauna series 30, 1−46. skead, c.j., 1966, ‘white-browed sparrow-weaver, plocepasser mahali a. smith, in the eastern and north-eastern cape’, ostrich 37, 128−129. skead, c.j., 1967, ‘ecology of birds in the eastern cape province’, ostrich suppl. 7, 1−103. skead, c.j., 1987, historical mammal incidence in the cape province. vol. 2. the eastern half of the cape province, including the ciskei, transkei and east griqualand, chief directorate, nature and environmental conservation, cape town. urquhart, c. & klages, n., 1997, addo, more than just elephants, bluecliff publishing, port elizabeth. vernon, c.j., 1982, ‘coursers in the eastern cape’, bee-eater 33(3), 31−34. vernon, c.j., 1989, ‘observations on the forest birds around east london’, ostrich 14, suppl., 14, 75−84. ward, d., martin, r., martin, j. & martin, e., 1984, ‘palm swifts in the cape province’, bokmakierie 36, 107. article information authors: bruce r. ellender1,2,3 olaf l.f. weyl1,3 affiliations: 1south african institute for aquatic biodiversity, grahamstown, south africa 2department of ichthyology and fisheries science, rhodes university, south africa 3centre for invasion biology, south african institute for aquatic biodiversity, grahamstown, south africa correspondence to: bruce ellender email: bru.ellender@gmail.com postal address: private bag 1015, grahamstown 6140, south africa dates: received: 04 nov. 2014 accepted: 27 jan. 2015 published: 22 may 2015 how to cite this article: ellender, b.r. & weyl, o.l.f., 2015, ‘resilience of imperilled headwater stream fish to an unpredictable high-magnitude flood’, koedoe 57(1), art. #1258, 8 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1258 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. resilience of imperilled headwater stream fish to an unpredictable high-magnitude flood in this original research... open access • abstract • introduction • research method and design    • study site    • field surveys    • data analysis       • rainfall and flow       • fish population dynamics • results    • rainfall and flow       • fish responses • discussion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ headwater stream fish communities are increasingly becoming isolated in headwater refugia that are often cut off from other metapopulations within a river network as a result of non-native fish invasions, pollution, water abstraction and habitat degradation downstream. this range restriction and isolation therefore makes them vulnerable to extinction. understanding threats to isolated fish populations is consequently important for their conservation. following a base-flow survey, a high-magnitude flood (peak flow of 1245 m−3s−1) provided an opportunity to investigate the response of endangered eastern cape redfin pseudobarbus afer populations to a natural disturbance in the waterkloof and fernkloof streams, two relatively pristine headwater tributaries of the swartkops river system within the groendal wilderness area, eastern cape province, south africa. pseudobarbus afer had limited distributions, occupying 3 km in both the fernkloof and waterkloof streams. fish population assessments before and after the flood event indicated that there were no longitudinal trends in p. afer abundance before or after the flood, but overall abundance post-flooding in the fernkloof stream was higher. there were no noticeable changes in p. afer size structure preand post-flood. pseudobarbus afer showed resilience to a major flooding event most likely related to evolution in river systems characterised by environmental stochasticity. conservation implications: this research provides insight into the population level responses of native headwater stream fishes to unpredictable natural disturbance. of particular relevance is information on their ability to withstand natural disturbances, which provides novel information essential for their conservation and management especially as these fishes are already impacted by multiple anthropogenic stressors. introduction top ↑ headwater stream fishes in south africa are increasingly being isolated in small fragmented headwater refuges as a result of competition with and predation by non-native fishes, downstream water abstraction, pollution and habitat degradation (marr et al. 2010; tweddle et al. 2009; weyl et al. 2014). protected areas play an important role in conserving headwater fishes by preserving habitats and preventing non-native fish introductions (russell 2011). it is increasingly recognised, however, that the persistence of fishes in these environments may be dependent on dispersal between complementary habitats for reproduction, feeding, rearing and disturbance avoidance (franssen et al. 2006; labbe & fausch 2000; schlosser & angermeier 1995). isolation may therefore compromise their long-term persistence (fausch et al. 2009). understanding the vulnerability of isolated refuge populations to natural catastrophic events such as flooding is therefore important for conservation planning. headwater streams are considered particularly susceptible to floods because they have small catchments and are easily influenced by relatively minor changes in local conditions (meyer et al. 2007). unpredictable, infrequent and catastrophic floods can result in slope failures, bank erosion, substrate scouring, and loss of habitat and biota (resh et al. 1988). in fish populations, floods can result in downstream displacements (cambray 1994), reduced abundance (magalhaes, schlosser & collares-pereira 2003; matthews 1986; nislow et al. 2002; pires et al. 2008), recruitment failure (letcher & terrick 1998) and alteration in community composition (matthews 1986; nislow et al. 2002). despite the documented impacts of floods on stream fish communities (letcher & terrick 1998; matthews 1986; nislow et al. 2002; pires et al. 2008; resh et al. 1988), fishes also display long-term resilience and have been shown to return to equilibrium fairly rapidly following catastrophic flood events (dolloff, flebbe & owen 1994; matthews 1986). under natural conditions, fishes are able to recolonise disturbed stream reaches from metapopulations in undisturbed refuge habitats (dolloff et al. 1994; matthews 1986; medeiros & maltchik 2001). isolation is a concern for freshwater fish conservation in south africa, where many headwater stream fishes are endemic to individual river systems and populations are increasingly fragmented by man-made (dams, weirs) or biological (invasive fish predation) barriers (ellender, weyl & swartz 2011; tweddle et al. 2009). one such species is the mandela lineage of the endangered eastern cape redfin minnow, pseudobarbus afer (peters, 1864) (swartz & impson 2007). its natural distribution is limited to only three river systems (sundays, swartkops and baakens) in the eastern cape province of south africa (swartz, skelton & bloomer 2007). populations are heavily impacted by non-native fish predation (ellender et al. 2011) and only persist in isolated headwater refugia (ellender 2013). fortunately, a considerable portion of these headwater refugia are located in two protected areas, the addo elephant national park (sundays river system) and the groendal wilderness area (swartkops river system). whilst these protected areas provide protection from non-native fishes and habitat destruction, the impact of natural catastrophic floods on these isolated populations are unknown. this article provides insight into the population level responses of p. afer to an unpredictable high-magnitude flood event by comparing the distributions, relative abundance and population structure of p. afer in two headwater streams before and after an unpredictable high-magnitude flood. it was hypothesised that the flood would (1) result in displacement and mortality of headwater fishes, resulting in changes in abundance and distribution in headwater streams and (2) that juvenile life history stages of p. afer would be more vulnerable to flood disturbance. research method and design top ↑ study site the study was undertaken within the groendal wilderness area on two second-order, episodic headwater tributaries (fernkloof stream, waterkloof stream) of the kwa-zunga river, a major tributary of the swartkops river system, eastern cape, south africa (figure 1). as a result of their entire catchments being situated within the groendal wilderness area (a protected area proclaimed for the protection of indigenous forest and water resources), the fernkloof and waterkloof streams are relatively pristine with undammed catchments and intact riparian zones characterised by closed canopy (fernkloof: 46% ± 23% canopy cover; waterkloof: 33% ± 24% canopy cover) tropical coastal forest (acocks 1975) and mesic grassy fynbos (campbell 1985). the character of the two streams is typical for episodic headwater streams in the eastern cape. both streams have steep, high profile banks within narrow valleys. in-stream habitat was characterised by isolated pools (fernkloof [mean ± s.d.] length: 10.8 m ± 3.2 m; width: 3.6 m ± 0.9 m; depth: 0.7 m ± 0.4 m; waterkloof [mean ± s.d.] length: 11.9 m ± 3.7 m; width: 5.5 m ± 4.8 m; depth: 0.7 m ± 0.6 m) connected by subsurface flows, with bedrock, large unconsolidated boulder, cobble and pebble substrates. surface flow only connects pools for a few days after heavy sustained rainfall. figure 1: the situation and sites sampled on the fernkloof and waterkloof streams, headwater tributaries of the swartkops river system within the groendal wilderness area, eastern cape, south africa. the position of the wincanton gauging weir (m1h004) on the elands river and the weather station (m1e001) at groendal dam are also noted. field surveys ten sites on each stream were sampled before (may 2010) and two months after (august 2012) a major flooding event (june 2012) by electrofishing using a samus 725g backpack electrofisher, attached to a 12 v battery with settings standardised at the duration of 0.3 ms and a frequency of 80 hz. because flood scouring and filling changed the position of pools within the two streams in the post-flood survey, the nearest pool to the pre-flood sample site was electrofished. single-pass electrofishing was conducted from the downstream side (tail) of the pool in an upstream direction, covering the entire length of each pool. upon completion of the pass, fish were identified to species level, measured, counted and released. electrofishing data were used as an index of relative abundance and catch per unit effort (cpue) was expressed as fish m−3 using volume estimates from the habitat data collected for each sampled pool. block nets were not used as pools were primarily isolated on the surface because of the episodic nature of the sampled streams. at each sampling site, temperature, conductivity and ph were measured using a hanna hi98129 combo ph and electrical conductivity meter (hanna instruments inc., woonsocket, usa). turbidity (ntu) was measured using a hanna hi 98703 turbidimeter (hanna instruments inc., woonsocket, usa). to estimate pool volume and habitat diversity, the length (± 0.1 m) of the pool was measured, followed by equally spaced width measurements (± 0.1 m). on each width transect, three depths (± 0.1 m) were measured; the outer two were 0.2 m each from the left-hand and right-hand stream bank and the third measurement taken midstream. at each depth measurement, the habitat type was recorded. canopy cover was estimated as a percentage of total cover and bankside vegetation type was recorded. data analysis rainfall and flow long-term hydrological and meteorological data were obtained from the department of water affairs hydrology section (dwaf 2012); the locations of the rainfall and gauging stations are presented in figure 1. these data were used to illustrate rainfall, flow variability and the magnitude of the flood in the headwaters of the swartkops river system. because of an absence of long-term flow data for the fernkloof and waterkloof streams, flow data were obtained from a representative swartkops river system headwater tributary, the gauging weir at wincanton on the elands river (m1h004) for the period 06 april 1965 – 05 january 2012. rainfall data were obtained from station m1e001 at groendal dam (period: 16 february 1950 – 11 january 2006) and supplemented with data from the nearby station m1e002 at uitenhage (period: 30 november 2006 – 05 january 2012). spearman rank order correlation was used to test for a relationship between rainfall and flow in the swartkops river system headwater streams at a significance of p < 0.05. the following criteria, outlined by resh et al. (1988), were used to classify the magnitude of the flood disturbance as: any relatively discrete event in time that is characterised by a frequency, intensity, and severity outside a predictable range, and that disrupts ecosystem, community, or population structure and changes resources or the physical environment. (p. 434) these criteria were applied to peak flow data for the swartkops river system to verify the magnitude of the flood in june 2012. fish population dynamics pre-flood and post-flood comparisons were limited to a single fish species, p. afer, as they were ubiquitously distributed from the limit of fish distributions (p. afer distribution: waterkloof = 3 km; fernkloof = 3 km) in the upper reaches to the confluence with the mainstem kwa-zunga river before the flood, allowing longitudinal analyses. to investigate the impact of a major flooding event on p. afer in headwater streams, pre-flood and post-flood electrofishing catch per unit effort (cpue, fish m−3) estimates were compared within the two streams using a main-effects analysis of variance (anova) at a significance level of p ≤ 0.05. specifically, cpue was tested for longitudinal trends (upper versus lower reaches), as well as flooding impact (pre-flood versus post-flood) and an interaction between the two factors. because of complications resulting from differences in sampling months, with the pre-flood survey occurring at the end of the spawning season and the post-flood survey at the beginning of the spawning season (cambray 1994), p. afer juveniles (< 40 mm) and adults (> 40 mm) were analysed separately. to test the vulnerability of the two p. afer size classes to the flooding event, the frequency of occurrence (number of sites per stream reach where each size class was present expressed as a percentage of all sites in that reach) by stream reach (upper and lower reaches) was tested for significance by reach and pre-flooding and post-flooding using a chi-squared test of independence. all analyses were undertaken using ms excel 2007, microsoft® and statistica 10.0, statsoft®. results top ↑ rainfall and flow mean annual rainfall (mean ± s.d.) for the period 16 february 1950 – 05 january 2012 was 620.7 ± 212.3 mm yr−1. rainfall was variable, falling in an unpredictable, erratic pattern (figure 2). there was a highly significant correlation between rainfall (station m1e001 and m1e002) and peak flow (gauging weir m1h004) (spearman r = 0.389; z = 9.45; p = 0.000), highlighting the system's episodic nature. peak flows between april 1965 and may 2012 were generally low (mean ± s.d.: 30.3 m−3s−1 ± 202.3 m−3s−1) and were predominantly < 25 m−3s−1 (89%), with a further 8.7% < 500 m−3s−1. during the 47 years on record, there were only four peak flows exceeding 1000 m−3s−1 (figure 2). the flooding event with peak flows of 1245 m−3s−1 (a measure of instantaneous flow at peak discharge) in june 2012 was of a frequency, intensity and severity outside of the predictable range and resulted in changes to the physical environment, which, according to resh et al. (1988), classified the flood as a major disturbance. figure 2: monthly peak flows (black) at the wincanton gauging weir on the elands river and total monthly rainfall (grey) at groendal dam (16 february – 11 january 2006) and at uitenhage (30 november 2006 – 05 january 2012) illustrating rainfall and flow variability in the headwater tributaries of the swartkops river system (flow data are missing for the period april 1981 – september 1986). fish responses the distribution and abundance of fishes from the fernkloof and waterkloof streams pre-flood and post-flood are summarised in table 1. four species were recorded from the fernkloof and waterkloof streams, of which three were shared (native: p. afer and cape kurper sandelia capensis [cuvier, 1831]; non-native: african sharptooth catfish clarias gariepinus [burchell, 1822]) and an additional non-native, banded tilapia tilapia sparrmanii a. smith 1840 was recorded in the fernkloof stream only. pseudobarbus afer distributions were limited to a 3 km stretch in both the fernkloof and waterkloof streams. table 1: the stream zones (lower reaches, upper reaches), sites (10 sites), distribution, catch per unit effort (fish m−3), relative abundance (%) of fishes by site (%), including water-quality variables from site 1 in the lower reaches to site 10 in the upper reaches of the fernkloof and waterkloof streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area. there were no significant pre-flood and post-flood longitudinal or interactive effects (stream reach: upper and lower reaches; disturbance: pre-flood and post-flood) in cpue for p. afer in the waterkloof stream (pre-flood mean ± s.e.: 1.24 ± 0.33 fish m−3; post-flood mean ± s.e.: 2.43 ± 0.40 fish m−3) (table 2). there was, however, a significant increase in post-flood p. afer cpue in the fernkloof stream (pre-flood mean ± s.e.: 1.33 ± 0.49 fish m−3; post-flood mean ± s.e.: 6.53 ± 2.12 fish m−3) (figure 3; table 2). the frequency of occurrence and length distributions for p. afer pre-flood and post-flood are summarised in table 3 and figure 4. table 2: a summary of pre-flood and post-flood longitudinal (upper and lower stream reaches) and disturbance (pre-flood and post-flood) main-effects anova results for pseudobarbus afer cpue (fish m−3) from the waterkloof and fernkloof streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area. figure 3: pre-flood and post-flood longitudinal abundance trends for pseudobarbus afer catch per unit effort (fish m−3) from the waterkloof and fernkloof streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area, eastern cape, south africa. table 3: the frequency of occurrence (% of sites containing pseudobarbus afer) of juvenile (< 40 mm) and adult (> 40 mm) pseudobarbus afer pre-flood and post-flood separated by stream reach (lower and upper) for the fernkloof and waterkloof streams headwater tributaries of the kwa-zunga river within the groendal wilderness area. figure 4: pre-flood and post-flood length frequency distribution for pseudobarbus afer from the upper and lower reaches of the fernkloof (a = lower; c = upper) and waterkloof (b = lower; c = upper) streams, headwater tributaries of the kwa-zunga river within the groendal wilderness area, eastern cape, south africa. the frequency of occurrence for juvenile (< 40 mm) and adult p. afer was independent of stream reach and were subsequently grouped by stream (table 3). the frequency of occurrence of juvenile and adult p. afer was independent of flooding for the waterkloof (pre-flood versus post-flood: χ2 = 0.027, df = 1, p = 0.87) and fernkloof streams (pre-flood versus post-flood: χ2 = 0.044, df = 1, p = 0.83) (table 3). in the waterkloof stream, p. afer size distributions were similar between stream reaches and pre-flooding and post-flooding (pre-flood lower mean, range: 43.7, 22 mm – 74 mm fork length (fl); pre-flood upper mean, range: 40.7, 16 mm – 71 mm fl; post-flood lower mean, range: 42.8, 21 mm – 67 mm fl; post-flood upper mean, range: 47.2, 28 mm – 81 mm fl) (figure 4). in the fernkloof stream, the mean length of p. afer was larger in the lower reaches but similar before and after the flood (pre-flood lower mean, range: 53.0, 11 mm – 96 mm fl; pre-flood upper mean, range: 43.0, 24 mm – 72 mm fl; post-flood lower mean, range: 52.0, 30 mm – 87 mm fl; post-flood upper mean, range: 46.3, 21 mm – 100 mm fl). discussion top ↑ flood-related disturbances typically elicit a variety of responses from fishes in stream environments, the nature of which are related to the predictability, magnitude/intensity and duration of the event (lytle & poff 2004) as well as the specific adaptations of the fishes to the abiotic conditions of that environment (franssen et al. 2006; lytle & poff 2004). in the fernkloof and waterkloof streams, p. afer demonstrated resilience to a major flooding event. in both small streams, there were no longitudinal trends in p. afer abundance before or after the flood, but overall abundance post-flooding in the fernkloof stream was higher. this may have been the result of the inability to sample the exact sites before and after the flood as some pools were filled up by flood deposition. the nearest pool to the pre-flood site was sampled, which may have resulted in sampling a refuge pool where post-flood abundances were high as a result of collection of fishes from that stream segment. good recruitment during the 2010/2011 spawning season might also have led to the observed increase in abundance as during this period rainfall was higher than the preceding two spawning seasons. increased recruitment during periods of good flow was recorded by franssen et al. (2006) for intermittent prairie stream fishes in north-eastern kansas, usa. however, the p. afer length frequency data from the waterkloof and fernkloof streams do not support this, as size structuring before and after the flood was similar. the vulnerability of stream fishes to flood disturbance is not always clearly evident from overall abundance trends, but in some cases certain life history stages of particular species may be more vulnerable to disturbance than others (gasith & resh 1999; letcher & terrick 1998; lytle & poff 2004). in new england streams in the usa, a massive localised flood caused an age-0 year-class failure in brook trout salvelinus fontinalis (mitchill, 1815) and brown trout salmo trutta linnaeus 1758 and a large decrease in abundance of atlantic salmon salmo salar linnaeus 1758 (letcher & terrick 1998). similar results have been recorded for other species after catastrophic flood events (dolloff et al. 1994; harvey 1987; matthews 1986; nislow et al. 2002). this was not the case for p. afer as there were no significant changes in the occurrence of either juvenile or adult p. afer following the flood. both life history stages were distributed throughout both streams from their confluence with the mainstream kwa-zunga in the lower reaches, to the limit of p. afer distribution in the upper reaches. the susceptibility of juvenile fishes to flood displacement and mortality has been documented to decrease rapidly with a small increase in length (harvey 1987) and p. afer may show similar trends in changing vulnerability with size. in the nearby wit river, a headwater tributary of the gamtoos river system, cambray (1994) reported that late free embryos and early larvae drift out of the areas where they were spawned. during the current study, the flooding event took place 3–5 months after the peak spawning season of p. afer (cambray 1994), perhaps allowing young-of-the-year fish time to grow out of the most vulnerable life history stages. distribution and movement of non-native species indicated species-specific and stream-specific flood response. for example, after the same flood, ellender, woodford and weyl (2015) noted that the flood event appeared to have flushed c. gariepinus out of the fernkloof stream but facilitated its penetration into the waterkloof stream. temporal longitudinal movement was also noted in the blindekloof stream, another kwa-zunga river tributary, during a period of prolonged flow (± 5 months) following the june 2012 flood, when c. gariepinus penetrated 5 km upstream from the mainstem (ellender et al. 2015). in conclusion, the observed resilience of p. afer populations to a major flooding event is most likely related to their evolution in river systems characterised by environmental stochasticity (dolloff et al. 1994; magalhaes et al. 2003; pires et al. 2008). the long-term persistence of p. afer in relatively pristine isolated headwater refugia may therefore not be threatened by flood disturbances of the observed magnitude and intensity. the biggest long-term threat may be the flood-facilitated penetration of headwater refugia by non-native species. acknowledgements top ↑ b.r.e. received financial support from: the south africa–netherlands research programme on alternatives in development (sanpad project 10/06), the national research foundation of south africa (nrf), rhodes university, the dst/nrf centre of excellence for invasion biology and the water research commission (wrc project no. k5/1957/4, k5/2039 and k5/2261). eastern cape parks board and staff of the groendal wilderness area are thanked for access and for their logistical assistance. geraldine taylor and patrick gourley are thanked for field assistance. research was conducted following saiab animal ethics guidelines. research permits were issues by the eastern cape department of economic development and environmental affairs (dedea) and eastern cape parks. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions b.r.e. 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cape floristic region: a new beginning for the rondegat river’, fisheries 39, 270–279. http://dx.doi.org/10.1080/03632415.2014.914924 article information authors: dirk j. roux1,2 llewellyn c. foxcroft3,4 affiliations: 1south african national parks, george, south africa 2sustainability research unit, nelson mandela metropolitan university, george, south africa 3south african national parks, skukuza, south africa 4centre for invasion biology, department of botany and zoology, stellenbosch university, south africa correspondence to: dirk roux email: dirkr@sanparks.org postal address: private bag x 6531, george 6530, south africa how to cite this article: roux, d.j. & foxcroft, l.c., 2011, ‘the development and application of strategic adaptive management within south african national parks’, koedoe 53(2), art. #1049, 5 pages. doi:10.4102/koedoe.v53i2.1049 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) the development and application of strategic adaptive management within south african national parks in this editorial... open access • introduction • strategic adaptive management in sanparks    • philosophical foundation       • adaptive planning       • adaptive implementation       • adaptive evaluation • roadmap through this special issue • conclusion • acknowledgements • references introduction (back to top) adaptive management is an appealing approach to deal with inherent uncertainty in complex and interactive social-ecological systems (holling 2001; rogers 2003). in short, adaptive management is about learning-by-doing in a scientific way, adapting behaviour and overall direction as new information becomes available. it provides a structured way for improving our incomplete understanding through an iterative process of setting objectives, implementing policy decisions and evaluating the implications of their outcomes for future decision making. in essence, adaptive management is: the process of treating natural resource management as an experiment such that the practicality of trial and error is added to the rigour and explicitness of the scientific experiment, producing learning that is both relevant and valid. (meffe et al. 2002) when adaptive management is practiced, policies become hypotheses and management actions become the experiments to test those hypotheses (folke et al. 2005). first referred to as adaptive environmental assessment and management (aeam) (holling 1978; walters 1986), adaptive management has grown into an established field of research and practice (allan & stankey 2009; armitage, berkes & doubleday 2008; meffe et al. 2002; oglethorpe 2002; walters 2002). whilst there is a rich literature on the philosophical merits of adaptive management, the actual day-to-day implementation had faced many obstacles (berkes, colding & folke 2003; johnson 1999; lee 1993; mclain & lee 1996; rogers 2003; shea et al. 2002; walters 1997). yet a version of adaptive management that developed in south africa has grown to become an integral part of the thinking, planning and decision-making within south african national parks (sanparks). this version is referred to as strategic adaptive management (sam) and this special issue is dedicated to reflecting on the development and implementation of sam within sanparks and its stakeholder community over a 10-year period. in this editorial we present a brief introduction to the main components and vocabulary of sam as practiced by sanparks, as well as a roadmap through the papers that constitute the two parts of the special issue. the papers that follow in this special issue, and the references therein, will provide the reader with a rich source of literature and in-depth treatise of sam and its development within sanparks. strategic adaptive management in sanparks (back to top) philosophical foundation the century-long evolution of management practices in the kruger national park is chronicled in venter et al. (2008). an optimisation approach in the early years (c.1902–1980), as well as command-and-control (1955–1985) and laissez-faire (1985–1995) approaches were embraced before the adoption of adaptive management in 1995. the appropriateness of adaptive management for natural resource management in general, and its adoption by kruger national park, stems from a growing awareness of two critical challenges, (1) the existence of ecological complexity and social complexity and hence social-ecological complexity and (2) the existence of multiple stakeholders with diverse (and often divergent) perceptions, values and expectations. a defining characteristic of complex systems is that patterns emerge or self-organise from the local interactions between components of the system. the interactions and feedbacks between components can be ‘nonlinear’, resulting in an inherent degree of unpredictability in cause-and-effect relationships and making them ‘knowable’ only in retrospect. an emergent property, for example patterns in organism distribution, is not a property of any single agent but of the system as a whole (levin 1998, 1999; snowden & stanbridge 2004). the second challenge relates to the multistakeholder nature of common-property natural resources; even fenced-off protected areas are increasingly influenced by external social issues (venter et al. 2008). these stakeholders may subscribe to widely varying world views, based on different values and knowledge forms, with expectations that play out over different time horizons and spatial scales. under these circumstances, management should probably avoid targeting an optimal solution for ‘the (single) problem’, but should adopt an ongoing learning and negotiation process where mutual sense-making and adaptation are prioritised (pahl-wostl & hare 2004). this reality has lead some authors and practitioners to coin the term ‘adaptive co-management’, as a descriptor of adaptive management that explicitly caters for mutual learning and cooperation between stakeholders such as conservation agencies, researchers and local communities (armitage et al. 2007; borrini-feyerabend et al. 2000; olsson, folke & berkes 2004; ruitenbeek & cartier 2001). the presence of limited predictability (or a certain level of irreducible uncertainty), as well as multiple stakeholders with frequent conflicting interests, suggests that there are two fundamental conditions necessary for effective management of natural resources, (1) to learn and adapt and (2) to do so purposefully with relevant partners. sam, which was initially developed in the context of managing rivers and their catchment areas (biggs & rogers 2003; rogers & bestbier 1997; rogers & biggs 1999), provides a framework for facilitating such learning. it incorporates the iterative learning dimension of adaptive management and the mutual learning dimension of co-management. in addition, it emphasises a forward-looking dimension, hence the reference to ‘strategic’. in summary, as grant et al. (2008) point out, sam is designed to be strategic (facilitate action with foresight and purpose), adaptive (facilitate learning whilst we are doing) and participatory (facilitate engagement and empowerment of stakeholders). sam is a modular process that allows practitioners to start with any of the five broad steps in the adaptive cycle and to expand their efforts from there. the five steps can be grouped into three interrelated subprocesses or components that have taken shape in the application of sam, namely adaptive planning, adaptive implementation and adaptive evaluation (figure 1). in the following sections we introduce the basic steps of sam in more detail. figure 1: schematic summary of the steps in the strategic adaptive management process, based on the work of biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 59-80, island press, washington dc; pollard, s.r. & du toit, d.r., 2007, guidelines for strategic adaptive management – experiences from managing the rivers of the kruger national park. guidelines of unep/gef project no. gf/27-13-03-4679. ecosystems, protected areas and people project; and scholes, r.j. & mennell, k.g., 2008, ‘summary for policymakers’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 1-21, wits university press, johannesburg. adaptive planning the adaptive planning process of sam is seen increasingly as a critical condition for achieving its successful implementation (rogers pers. comm., 14 september 2010). the aim of this process is to build a sense of common purpose amongst all relevant stakeholders and to develop a collective roadmap for getting from a current (usually undesirable) reality to a more desirable social-ecological system. stakeholder inclusivity is vital to the success of an adaptive planning process. in the case of sanparks, stakeholders include park managers, scientists, government policymakers, agency managers, wildlife activists, traditional communities neighbouring parks, farmers that share catchments of rivers that flow through parks, ngos and ecotourists. successful adaptive planning depends on the facilitation of a constructive dialogue amongst these stakeholders with their diverse and often divergent values, expectations, professional norms and reward systems (rogers & breen 2003). the first step in adaptive planning is to create a common vision in which stakeholders agree on the social, technical, economic, ecological and political contexts of the system to be managed. a critical part of this visioning exercise is to reach agreement on values, or operating principles, which should guide management decision making in the future. the v-steep (values, social, technical, economic, ecological and political) framework that emerges provides an approach to describing the context as comprehensively as possible (see pollard & du toit 2007). a further part of visioning is to deliberate and reach consensus on the vital attributes of the system to be managed and their determinants. vital attributes, as perceived by stakeholders, are the distinctive and special features of the social-ecological system of concern that are the key to its management (rogers & bestbier 1997). a vision statement is formulated on the basis of this understanding of the context and values. the vision, together with the vital attributes of the system to be managed, informs the setting of objectives. a nested hierarchy of objectives starts with high-level objectives that are set, firstly, to ensure the maintenance of the identified vital attributes of the system to be managed and, secondly, to overcome the constraints and threats to meeting the vision. through a step-by-step process, these high-level objectives (which are largely based on stakeholder values) are deconstructed into a series of objectives of increasing detail until they represent measurable, scientifically credible endpoints. the result is referred to as an objectives hierarchy (figure 2). acknowledging the dynamic nature of ecosystems, the measurable targets (figure 2) describe the boundaries of the desired state (as opposed to an optimal value). these boundaries are also referred to as thresholds of potential concern (tpcs) and are essentially hypotheses of the outer limits of acceptable change. these ‘boundaries’ are acceptable changes in the attribute of interest, embedded within specific temporal and spatial scales (see foxcroft & mcgeoch 2011). as such, their validity and appropriateness remains open to challenge and tpcs are revised as understanding of the system improves (pollard & du toit 2007). tpcs are developed in collaboration between managers, scientists and field staff responsible for monitoring. the last step that forms part of the adaptive planning process is to scope or analyse various options for achieving the objectives that were derived in the previous step. importantly, this step is still conducted in cooperation with stakeholders. different options are identified, their likely consequences predicted and the acceptability of those consequences assessed. finally, a combination of management options that provide the best potential social-ecological system outcomes and learning opportunities is selected for implementation (grant et al. 2008; pollard & du toit 2007). figure 2: generic objectives hierarchy that links the vision to scientific endpoints, based on the work of pollard, s.r. & du toit, d.r., 2007, guidelines for strategic adaptive management – experiences from managing the rivers of the kruger national park. guidelines of unep/gef project no. gf/27-13-034679. ecosystems, protected areas and people project; rogers, k. & bestbier, r., 1997, development of a protocol for the definition of the desired state of riverine systems in south africa, department of environmental affairs and tourism, pretoria; and rogers, k.h. & biggs, h.c., 1999, ‘integrating indicators, endpoints and value systems in strategic management of the kruger national park’, freshwater biology 41, 439-451. doi:10.1046/j.1365-2427.1999.00441.x. adaptive implementation adaptive implementation entails incorporating the options that were selected in the previous step as part of the operating procedures and business routines of the relevant organisation(s). this requires the development of detailed action plans, allocation of the necessary resources and the implementation of those plans. a key component of the adaptive implementation of action plans is to develop monitoring protocols to describe the subject and focus of what to monitor and establish the frequency at which to do so. monitoring endpoints are linked to the measurable targets (or tpcs) in figure 2. part of the ‘new’ management procedures is to establish a forum for the regular evaluation of monitoring results against set tpcs, as well as standard procedures for dealing with tpc excedance and for capturing and sharing learning (pollard & du toit 2007). adaptive evaluation one of the main purposes of sam (and adaptive management in general) is to purposefully learn and adapt over time. therefore, it is essential that learning becomes an explicit step in the strategic adaptive management process (figure 1). however, learning should not be seen as a mere step to be taken at the end of the process, but should rather occur throughout the planning and implementation phases via a series of feedback loops. continuous evaluation and learning is facilitated by reflecting on the following questions (figure 3): • is the monitoring adequate, cost effective and feasible? • has the intended plan of operation materialised? • were the selected options appropriate? • were the predicted consequences correct and, if not, why? • were the consequences actually acceptable? • even if the predicted consequences were correct and are acceptable, are the objectives and vision being met? figure 3: the adaptive management process with feedback loops for ongoing reflective learning at multiple points during the process. roadmap through this special issue (back to top) the papers contained in this special issue were solicited to capture and share as much as possible of the experience gained by sanparks during the development and implementation of sam to date. the authors were allowed a fair degree of freedom to ensure that a broad spectrum of perspectives and diverse lessons are captured in the special issue. we refer to the papers in this issue as essays rather than research papers as some of the authors use a narrative writing style to convey their understanding of sam. the papers in the special issue can be divided into two parts. part one consists of papers of a more systemic, generic or philosophical nature, whilst the papers in part two focus largely on thresholds of potential concern and how these thresholds were derived and applied for specific target indicators. whilst each paper can be read on its own, the papers from part one and part two are meant to be complementary. part one starts off with a detailed contextual setting of the kruger national park as the main study area for the development of sam (pollard et al. 2011). the authors of this essay provide an historical overview of how changes in management paradigms eventually led to the adoption of sam as the approach of choice, as well as how this approach was pioneered in the sphere of river management. in the next essay, biggs, breen, slotow, freitag-ronaldson and hockings (2011) examine the relationship between assessment and reflection, focusing on how these processes can be used in a complementary way to catalyse learning for adaptive management. stirzaker et al. (2011) allude to the often contradicting nature of the entrained behaviour of many scientists and managers alike, as well as the behaviour required to participate effectively in an adaptive management process. they explore the shortcomings and requirements of organisations in terms of enabling adaptive management. in the fourth essay, holness and biggs (2011) address a question that is of critical importance to a conservation agency: are systematic conservation planning and adaptive management compatible processes? these authors argue that systematic conservation planning should be practiced as an intrinsic part of a broader adaptive management approach and suggest how such a marriage can be achieved. systematic monitoring is a key factor in our ability to learn and adapt. to this end, mcgeoch et al. (2011) propose a framework for biodiversity monitoring which would address the biodiversity objectives as outlined in the management plans of south african national parks. gaylard and ferreira (2011) reflect on how the process of sam itself has been adapted in response to various implementation challenges. the final essay in part one of this issue provides a transition to the part two papers. biggs, ferreira, freitag-ronaldson and grant-biggs (2011) provide a critical assessment of the usefulness of the concept of thresholds of potential concern. these authors propose a reconceptualisation of the tpc concept, based on learning over a period of one decade, to increase its utility within the sam process. part two contains a collection of 11 essays, each focusing on a specific theme. these essays aim to summarise the development of tpcs for the particular theme and assess how this fits into the broader strategic adaptive management approach. these areas have not received the same amount of attention over the last decade. therefore, whilst some papers will provide an in-depth discussion of their evolution, others will present recent developments and provide suggestions for future directions. as the pioneering work on sam in the kruger national park was conducted on river management, mcloughlin et al. (2011) take us through a journey explaining the history and rationale behind the tpc concept. furthermore, they show how the tpcs for river management were used and how they evolved as learning progressed. fire research and management in sanparks has been an ongoing process for a number of decades, with a number of different approaches being implemented, both within and across parks nationwide. van wilgen et al. (2011) discuss the changes from ‘trial and error’ through to ‘active adaptive management’, indicating the development of various forms of thresholds. another area where adaptive management approaches and tpcs have developed over a number of years is with invasive alien species. here foxcroft and mcgeoch (2011) aim to link the management actions, monitoring programme, research efforts and tpcs in order to develop functional feedback mechanisms and enable improved management and learning. whilst not discussing the development of tpcs specifically, a number of essays propose approaches to adopting an adaptive management strategy. they also suggest tpcs and aim to test the use of these in particular cases. for example, grant et al. (2011) evaluate herbivore–vegetation interactions and how determining thresholds for these can prevent unacceptable changes in desired vegetation states and patterns. mcgeoch et al. (2011) deal with approaches to monitoring the often neglected terrestrial and freshwater biodiversity. similarly, except for some specific case studies, resource use management has been given little attention until recently. in their essay, scheepers et al. (2011) provide an overall framework for applying adaptive approaches to resource use management. they illustrate this with three case studies, covering a range of approaches and timeframes, and conclude with opportunities for future expansion. as global environmental change intensifies, one of the most pressing issues that conservation and protected area managers face is ensuring the persistence of rare species (rebelo et al. 2011). however, in areas with high endemicity, and which are facing a number of potentially negative impacts, assessing which species are most in need of special attention is problematic. for example, table mountain national park has 307 threatened iucn red list (plus 208 non-least concern) and 332 endemic terrestrial plant and animal species. rebelo et al. (2011) present an approach to dealing with this challenging problem.four essays probe some underlying philosophical or technical issues, including the idea that behind all good science lies good science support. this is an essential but underrated part of the overall science management and monitoring partnership. kruger and macfadyen (2011) discuss a number of innovations that deal with these issues, from collecting, managing and automating data management, to developing systems to report back on tpcs. a challenge with implementing tpcs is determining at what stage the breach of a tpc is triggered and how the lag effects of this breach are handled. scholes and kruger (2011) present a potential approach to this, illustrating it with an example from the kruger national park. owing to the different needs and analytical approaches necessary to implementing tpcs and management across the sanparks estate, ferreira et al. (2011) use conceptual linkages between objectives, indicators, mechanisms and modulators to help identify key concerns in relation to management objectives. based on these linkages, the underlying mechanisms responsible for the management concern may be evaluated. conclusion (back to top) the dominant message that emerges from the papers in this special issue is that adaptive management is about structured learning. the authors reflect on various mechanisms that are used to make current assumptions and understanding explicit so that relevant stakeholders can learn in a structured way. these mechanisms include co-creating a desired state or vision, setting objectives, formulating thresholds of potential concern, and monitoring and evaluating the consequences of management decisions. this volume of papers represents a comprehensive documentation and reflection of this process, after applying the principles of sam across an entire conservation agency for over 10 years. acknowledgements (back to top) we thank dr richard stirzaker, prof. kevin rogers and dr harry biggs for discussions and comments during the development of this editorial. references (back to top) armitage, d., berkes, f. & doubleday, n., 2007, adaptive co-management: collaboration, learning, and multi-level governance, ubc press, vancouver. berkes, f., colding, j. & folke, c. (eds.), 2003, navigating social-ecological systems: building resilience for complexity and change, cambridge university press, cambridge. biggs, h., ferreira, s., freitag-ronaldson, s. & grant-biggs, r., 2011, ‘taking stock after a decade: does the ‘thresholds of potential concern’ concept need a socio-ecological revamp?’ koedoe 53(2), art. #1002, 9 pages. doi:10.4102/koedoe.v53i2.1002 biggs, h.c., breen, c., slotow, r., freitag, s. & hockings, m., 2011, ‘how assessment and reflection relate to more effective learning in adaptive management’, koedoe 53(2), art. #1001, 13 pages.doi:10.4102/koedoe.v53i2.1001 biggs, h.c. & rogers, k.h., 2003, ‘an adaptive system to link science, monitoring and management in practice’, in j.t. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience. ecology and management of savanna heterogeneity, pp. 59−80, island press, washington dc. borrini-feyerabend, g., farvar, m.t., nguinguiri, j.c. & ndangang, v., 2000, co-management of natural resources: organizing negotiation and learning by doing, kasparek zverlag, heidelberg. ferreira, s., deacon, a., sithole, h., bezuidenhout, h., daemane, m. & herbst, m., 2011, ‘from numbers to ecosystems and biodiversity: a mechanistic approach to monitoring’, koedoe 53(2), art. #998, 12 pages. doi:10.4102/koedoe.v53i2.998 folke, c., hahn, t., olsson, p. & norberg, j., 2005, ‘adaptive governance of social-ecological systems’, annual review of environmental resources 30, 441−473. doi:10.1146/annurev.energy.30.050504.144511 foxcroft, l.c. & mcgeoch, m., 2011, ‘implementing invasive species management in an adaptive management framework’, koedoe 53(2), art. #1006, 11 pages. doi:10.4102/koedoe.v53i2.1006 gaylard, a. & ferreira, s., 2011, ‘advances and challenges in the implementation of strategic adaptive management beyond the kruger national park – making linkages between science and biodiversity management’, koedoe 53(2), art. #1005, 8 pages. doi:10.4102/koedoe.v53i2.1005 grant, r., sherwill, t., 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swemmer, l. & vermeulen, w.j., 2011, ‘applying adaptive management in resource use in south african national parks: a case study approach’, koedoe 53(2), art. #999, 14 pages. doi:10.4102/koedoe.v53i2.999 scholes, r.j. & kruger, j.m., 2011, ‘a framework for deriving and triggering thresholds for management intervention in uncertain, varying and time-lagged systems’, koedoe 53(2), art. #987, 8 pages. doi:10.4102/koedoe.v53i2.987 scholes, r.j. & mennell, k.g., 2008, ‘summary for policymakers’, in r.j. scholes & k.g. mennell (eds.), elephant management: a scientific assessment for south africa, pp. 1−21, wits university press, johannesburg. shea, k., possingham, h.p., murdoch, w.w. & roush, r., 2002, ‘active adaptive management in insect pest and weed control: intervention with a plan for learning’, ecological applications 12(3), 927−936. doi:10.1890/1051-0761(2002)012[0927:aamiip]2.0.co;2 snowden, d. & stanbridge, p., 2004, ‘the landscape of management: creating the context for understanding social complexity’, emergence: complexity and organization 6(1–2), 140−148. stirzaker, r.j., roux, d.j. & biggs, h.c., 2011, ‘learning to bridge the gap between adaptive management and organisational culture’, koedoe 53(2), art. #1007, 6 pages. doi:10.4102/koedoe.v53i2.1007 swemmer, l.k. & taljaard, s., 2011, ‘sanparks, people and adaptive management: understanding a diverse field of practice during changing times’, koedoe 53(2), art. #1017, 7 pages. doi:10.4102/koedoe.v53i2.1017 van wilgen, b.w., govender, n., forsyth, g.g. & kraaij, t., 2011, ‘towards adaptive fire management for biodiversity conservation: experience in south african national parks’, koedoe 53(2), art. #982, 9 pages. doi:10.4102/koedoe.v53i2.982 venter, f.j., naiman, r.j., biggs, h.c. & pienaar, d.j., 2008, ‘the evolution of conservation management philosophy: science, environmental change and social adjustments in kruger national park’, ecosystems 11, 173−192. doi:10.1007/s10021-007-9116-x walters, c., 1997, ‘challenges in adaptive management of riparian and coastal ecosystems’, conservation ecology 1(2), 1, viewed n.d., from http://www.consecol.org/vol1/iss2/art1/ filelist convert a pdf file! apologies and acknowledgements ladies and gentlemen: we have reached the end of our symposium proceedings and allow me to have the final word. mr e t matenge , director of national parks of the republic of botswana, as well as mr a b bridgens, secretary-general ofsarccus (southern african regional commission for the conservation and utilization of the soil) have tended their apologies for absence. i would like to thank the cabinet (in particular the hon. min. h schoeman) for the announcement of the institution of the 10th national park. this gratitude not only comes from the officials of the national parks board, but also from the people of the entire rsa, be they white , brown or black. i also thank my board for permission to go ahead with the arrangements for this symposium. the person who had to bite the bit was dr g de graaff and with the aid of his colleagues, everything ran smoothly. these efforts are greatly appreciated. furthermore, i would like to thank the international conference organisers. their loudspeakers and translation system never let us down. we had a large group of donours, financially and otherwise. these include the rembrandt organisation, the rennies organisation, coca-cola international, the united tobacco company, mr and mrs dingler, winter minerals (nelspruit) and the total petroleum company. to these organizations and individuals our thanks are due. without becoming sentimental, it is to be emphasized that no symposium can be held without people willing to participate. i would like to thank every speaker, especially dr f vollmar, who came all the way from switzerland to open this symposium. the reaction of the audience was wonderful. in particular i would also like to thank his royal highness, prince m. dlamini (swaziland) as well as the many leaders representing the developing territories, for their esteemed presence. my thanks are also due to the south african broadcasting corporation. we hope that whatever was summarized in such an efficient way by dr martiny, that it is clear that this symposium emphasized the importance of education and carried a message directed at the youth of southern africa. r. knobel symposium chairman 260 page 1 article information authors: frederik j. venter1 navashni govender2 affiliations: 1conservation management, kruger national park, south africa 2scientific services, kruger national park, south africa correspondence to: navashni govender postal address: private bag x402, skukuza 1350, south africa dates: received: 10 dec. 2010 accepted: 21 nov. 2011 published: 14 may 2012 how to cite this article: venter, f.j. & govender, n., 2012, ‘a geomorphic and soil description of the long-term fire experiment in the kruger national park, south africa’, koedoe 54(1), art. #1037, 10 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1037 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. a geomorphic and soil description of the long-term fire experiment in the kruger national park, south africa in this short communication... open access • abstract • introduction    • study site    • experimental burn plots • methods    • assumptions • results    • pretoriuskop    • skukuza    • satara    • mopane • discussion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ in 1954, the experimental burning programme into fire research was initiated in the kruger national park (knp), south africa. it is viewed as one of the last remaining long-term landscape fire experiments in africa. throughout the more than five decades of fire treatments in the experiment, numerous surveys (expanding various spatial and temporal scales), research projects (covering biotic and abiotic components) and analyses have been conducted with the aim to assess the impacts of different fire regimes on the savannah biome. the design of the experiment intended to test the effect of season and frequency of burning on vegetation within four major landscapes in the knp. however, these effects have been partly obscured by factors not fully taken into account by the experimental design, namely, herbivory, artificial water provision and soil variation. soil variation between replicates in the same landscape, as well as within individual replicates, has raised the issue of the representivity of the trial. this paper provided a description and ranking of the experimental burning trial according to the geomorphic and soil characteristics of each plot in comparison to the surrounding landscape. conservation implications: the knp burn plots are one of the largest and longest-running fire experiments on fire ecology in african savannahs. however, studies need to consider the underlying geomorphic and soil template when designing experiments and interpreting results. this work describes the representivity of the plots across, and within, treatments. introduction top ↑ fire has long been recognised as an essential determinant of the structure and function of african savannahs (higgins et al. 2007; sankaran, ratanam & hanan 2008; scholes & walker 1993; trollope 1982) and is often used as a management tool within conservation agencies. the application of appropriate fire regimes is usually informed and adapted by the findings of ongoing research, which is undertaken on the experimental application of selected fire regimes on fixed areas (see e.g., andersen, cook & williams 2003; knapp et al. 1998), with treatments often repeatedly applied for many decades. the changing attitudes towards fire management within south african conservation areas – from the orthodox equilibrium theory (fixed rotational fire regimes) to one that embraces a dynamic savannah system based on non-equilibrium theory (patch mosaic fires over space and time) (bond & archibald 2003; mentis & bailey 1990; van wilgen et al. 2000; van wilgen, biggs & potgieter 1998) – has questioned the validity of results from historical long-term fixed fire experiments in influencing fire management policies (van wilgen et al. 2004; van wilgen, govender & biggs 2007). a recent critical review of the long-term fire experiment in bringing about changes in the fire management policy at the kruger national park (knp) found that it had little direct influence. however, the experiment has had numerous other unforeseen benefits, particularly its support and use in ongoing key research projects undertaken on the experiment to improve our understanding of fire as a key driver of savannah dynamics (van wilgen et al. 2007). the spatio-temporal scale and the underlying template of various fire experiments often brings into question the representivity of the results from these plot-based experiments when scaling up to landscape processes. the knp long-term fire experiment (biggs et al. 2003) is often criticised regarding the difficulty in separating the effects of the experimental fire treatments from other factors that might be causing the observed differences. the experimental burn plot (ebp) trial was designed with the initial aim of testing the effects of different seasons and frequencies of burning on the vegetation of the knp. these effects have been partly obscured by factors not fully taken into account by the experimental design, most importantly, herbivory, artificial water provision and soil variation (biggs et al. 2003). the response of the vegetation and animal population to the template presented by the geology (which is reflected by the soils) and the changes caused by the ecosystem drivers such as rainfall, hebivory and fire, have led to a complex patch mosaic template within the park (venter, scholes & eckhart 2003). to quantitatively assess the effect that the varying fire regimes have on the dominant vegetation communities, it was envisioned that vegetation surveys would be conducted annually, whilst animal life would be observed periodically (nel 1953). a comprehensive baseline vegetation (woody and herbaceous) survey was conducted prior to establishment of all treatments in 1954. vegetation re-surveys were undertaken in the late 1990s and early 2000s (van wilgen et al. 2007). during the 50-year lifespan of the experiment, various researchers conducted surveys (which were predominantly of vegetation) on the experiment (govender, potgieter & biggs 2003; van wilgen et al. 2007). webber (1979) was the first to investigate the soil characteristics on the skukuza plots; thereafter, other work looking at soil characteristics was undertaken by mills and fey (2004a, 2004b), whilst the majority of research pertaining to soils on the experiment focused on the nutrient content, cycling and variation in nutrient levels with different fire regimes (coetsee 2007; feig 2004; otter 1992; shackleton & scholes 2000; van wilgen et al. 2007). analysis, understanding and incorporation of the results from the experiment of the effects of fire on savannah systems have only been undertaken in the past decade. the van wilgen et al. (2007) paper outlines, in detail, the contribution that the experiment had to effect change in management actions and policy in the park. key findings on the biota studied on the experiment include the following (van wilgen et al. 2007): • woody tree and shrub density are unresponsive to fire regimes and fire is not critical for the maintenance of woody plant richness and composition. exclusion of fire promotes an increase in woody biomass, with the most marked effects on woody vegetation in the extreme treatments (annual burning, burning in summer or wet season, or long periods of fire exclusion). these effects were also greater in areas of higher rainfall. • manipulation of fire regimes is not critical for the maintenance of herbaceous plant species diversity. herbaceous composition changed little with fires in the dormant season, but changes were significant in the wet, growing season and with fire exclusion. impacts caused by fire were most marked in wetter areas and increased grazing pressure on the plots after the treatments influenced the grass diversity at drier sites. • there were noticeable effects of fire on small mammal communities, with unburnt sites supporting the most species and highest densities of small mammals. • bird species richness and composition did not respond to the different fire regimes. • no significant effect on ant species composition and diversity between the fire treatments but significant differences in ant assemblage composition were found between the extreme fire treatments (burnt vs unburnt plots). • regardless of fire treatment, nitrogen loss is replenished soon after the fire. frequent, annual burning increases soil crusting on the plots. mycorrhizal colonisation increases with root branching and fine root development decreases with decreasing fire frequency, allowing for optimal acquisition of resources under different fire frequencies. • fire intensity varies as a result of changes in fuel moisture content rather than post-fire age, particularly because of the seasonal differences in fuel moisture content that override those of fuel load. since the inception of the experiment in the 1950s, our understanding of the role that fire plays within savannah ecosystems continues to improve. therefore, current and future fire management policies implemented in the knp will continue to be supported by the best available science that promotes appropriate adaptive fire management. owing to the lack of baseline soil surveys or soil characterisation of the ebps, this paper provides a soil and geomorphic description of the experiment in order to determine which replicate or individual plots within replicates are not representative of the land types in which they are set. it is well known that differences in soil types cause significant differences in vegetation in the knp (venter 1990) and knowledge of the variation and extent of different types of soil and geomorphology on the ebps is therefore necessary, in order to make future studies on the plots more relevant and representative of the surrounding landscape. study site the knp was proclaimed in 1926, and covers approximately 1 948 528 ha within the low-lying savannahs of north-eastern south africa. the vegetation is characterised by savannahs within 37 landscapes (gertenbach 1983), with the dominant trees being knobthorn (acacia nigrescens), marula (sclerocarya birrea), leadwood (combretum imberbe), red-bush willow (combretum appiculatum), silver cluster leaf (terminalia sericea) and mopane (colophospermum mopane). mean annual rainfall varies from 350 mm in the north to over 700 mm in the south. geologically, the knp is underlain by granites in the west, whilst the eastern sector is underlain predominantly by basalt (venter 1990). the flora of the park comprises 1983 species, including more than 400 tree and shrub species and more than 220 grass species. experimental burn plots fire research formally began in the knp in 1954 with the establishment of one of the few long-term fire ecology research experiments in africa (van der schijff 1958). the experiment consisted of the application of fires at varying return intervals and seasons, as well as protection from fire, on a series of plots approximately 7 ha in area within specific vegetation types represented by four of the major ecological regions in the park (pretoriuskop: sourveld vegetation; skukuza: combretum vegetation; satara: knobthorn and marula vegetation; mopane: mopane vegetation)(figure 1). the treatments were replicated four times in each of these ecological regions. within each ecological region, four different replicates or strings were scattered within 20 km of each other. a double firebreak road surrounds the individual replicates (with the area between the firebreak roads burnt annually in autumn) to ensure protection from wild fires. figure 1: location of the experimental burn plots within the land types as described by venter (1990), in the kruger national park. the treatments originally included annual (b1) winter fires in august and biennial (b2) and triennial (b3) fires in august (winter), october (after first spring rains), december (early summer), february (late summer) and april (autumn). in 1976, further treatments to examine the effects of fires every four (b4) and six (b6) years in october were added to selected landscapes (satara and mopane). full details and history of the experimental design and application of treatments are available from biggs et al. (2003). description of the areas in which the knp’s fire experiment was replicated within is outlined in table 1. the associated fire treatments and plots numbers for each replicate is given in online appendix 1a–d. table 1: salient features of the four major vegetation types in which the kruger national park’s fire experiment was replicated. methods top ↑ a series of aerial photographs (digital, colour, shutter speed: 1/800, scale: 1:50 000, altitude: 914.4 m, aircraft speed: 185.2 km/h) taken in april 2000 and field surveys were used to map the land units with their associated soil and vegetation patterns on each of the plots. sections that are atypical of the specified norm were delineated and assigned a score according to the system described below (table 2). however, the survey was not conducted to identify, in detail, all soil types or to give detailed descriptions of vegetation within the plots. reference made to land types in this report is according to the definitions and descriptions of venter (1990). table 2: the allocated representivity scores and description of each score assigned to plots and replicates. soil differences usually cut through individual plots so that part of the plot may score 5, whilst another part may have a score of 1. the plot as a whole was scored depending on the dominant soil types found on it and how representative it is of the environmental parameters within each particular replicate was assessed visually. assumptions the following assumptions were made with regard to this survey:• when the ebps were laid out in the 1950s, the initiators attempted to locate the different replicates on the crests in order to avoid too much soil variation. this assumption is based on the layout of the replicates in both granitic and basaltic areas, where minor portions of mid-slope and foot-slope areas are represented in some of the plots. • the presence or absence of roads influenced the layout of the ebps as they were only laid out alongside existing roads. this negatively influenced the attempt to restrict the ebps to upland areas, so that certain portions of plots occur in foot-slope areas. this scoring system was used as a means to investigate the following questions: • how representative of the broad ecological region or landscape is each group of four replicates and series? • how comparable is each of the four replicates? this is to test if one or more of the replicates in a series are outliers, why it is regarded as outliers and whether it is enough of an outlier to warrant complete exclusion from analyses under certain circumstances. • how comparable is each the individual plots in one replicate? are there particular outliers based on any geological, geomorphologic or soil criteria? results top ↑ pretoriuskop the pretoriuskop replicates of the ebps are all fairly well representative of the pretoriuskop land type (venter 1990). the fayi replicate generally has more sandy and yellow soils with patches of avalon–clovelly and shallow hutton–glenrosa–kroonstad forms (soil classification working group 1991) dominant with many t. sericea trees (figure 2a). the downstream part of some of the plots on the shabeni replicate are influenced by seepage areas and duplex soils, namely deep clovelly form (soil classification working group 1991) (figure 2b). this replicate seems to have less tall grass species than the other pretoriuskop plots. the numbi replicate features deep, red, sandy loam soils with minor areas of duplex soils (usually on the foot slope of plots), shallow soils (steep areas) or clay soils associated with intrusions of dolerite or gabbro. a clayey area caused by the presence of some gabbro may occur, for example on plot 4 (fire treatments – online appendix 1a–d) of the numbi replicate (figure 2c). the kambeni replicate has been deeply incised by a stream (plot 11) but the influence of this is limited and is characterised by very shallow soils (escourt form) (soil classification working group 1991), caused by the steep slope in this area (figure 2d). table 3a and 3b illustrates the overall representivity scores for each replicate and individual plot within pretoriuskop. figure 2: description of the soils types on the pretoriuskop replicates (a) fayi (b) shabeni (c) numbi and (d) kambeni. for the complete list of soils names and abbreviations see online appendix 2. table 3a: the overall representivity scores for each replicate within pretoriuskop. table 3b: the overall representivity scores for each plot within the four replicates in pretoriuskop. skukuza huge granite rock outcrops and boulders occur on some of the biyamiti plots. the soils within these plots are usually hydromorphic and therefore slightly different from the rest of the plots. the south-western plots (plots 9–12) feature more clayey soils as a result of the presence of dolerite dykes in this area. these plots are generally quite different from the other plots in this replicate, as well as the series, as they are dominated by estcourt–swartlands–kroonstad and hutton–clovelly–fernwood forms (soil classification working group 1991) (figure 3a). the skukuza replicate of the ebps is characterised by quite large areas of sodic and duplex soils that occur on the foot slopes of parts of plots 6 and 7. these sodic soils have relatively thin a horizons (< 15 cm), which makes these areas completely different from the other areas in the plots. serious erosion within firebreaks is already evident where the firebreaks cross into sodic soils. unfortunately, the control or no burn plot (plot 7) (see online appendix 1a–d for treatments) at this replicate is one of those most seriously affected by the sodic soils, which covers more than half of the no burn treatment (figure 3b). the n’washitshaka replicate is characterised by reddish soils (glenrosa–hutton forms), but plot 6 is almost completely associated with hydromorphic or duplex soils (kroonstad–estcourt forms) (soil classification working group 1991) (figure 3c). the napi replicate also has some duplex soils (fernwood–kroonstad and hutton–clovelly–fernwood forms), as well as granite boulders or rocky outcrops (figure 3d). table 4a and 4b illustrates the overall representivity scores for each replicate and individual plot within skukuza. figure 3: description of the soils types on the skukuza replicates (a) biyamiti (b) skukuza (c) n’waswitshaka and (d) napi. for the complete list of soil names and abbreviations see online appendix 2. table 4a: the overall representivity scores for each replicate within skukuza. table 4b: the overall representivity scores for each plot within the four replicates in skukuza. satara it is assumed that these replicates were located to represent the s. birrea–a. nigrescens savannah of the central basalt plains. the soils and general vegetation of this zone are described as the satara land type by venter (1990). the satara replicate occurs on a very flat part of the basalt plains. the geology is sabie river basalt formation with dolerite dykes and the soils are mainly moderately deep shortlands–swartland soil (soil classification working group 1991), that are red structured clays. because of the flatness of this area, a number of shallow pans and associated black vertic soils occur. some thin, elongated, rocky outcrops associated with dolerite dykes also occur (figure 4a). the lindanda replicate (figure 4b) is representative of soils and vegetation of three different land types, namely satara, mavumbye and nwanetsi (venter 1990), as it is characterised by soils associated with both the sabie river basalt formation and the letaba basalt formation. it has been influenced by colluviation from the lebombo hills. this colluvium has influenced soils in the north-eastern corner of the replicate (portions of plots 5 and 6 and plot 7). the occurrence of more clayey soils along the south-western part of the replicate (plots 8, 9, 10, 11, 12 and 14) is ascribed to the occurrence of picrite basalts of the letaba basalt formation. this area is characterised by very clayey soils of the bonheim and arcadia forms. stunted a. nigrescens dominates this area and no s. birrea occurs. a drainage line through plot 2 is the cause of alluvial soils (bonheim and swartland) (soil classification working group 1991) and typical associated dense vegetation. the influence of the valley bottom and foot-slope extends about 50 m on either side of the drainage line. the area to the west of the drainage line, covered by plot 1 and part of plot 2 corresponds to a transition between the red, medium clayey soils associated with sabie river basalt and the black very clayey soils associated with letaba basalt. the woody vegetation on parts of plots 5 and 6 and all of plot 7 indicates that sodic soil conditions occur, for example sterkspruit, with acacia welwitschii as the dominant tree. some pans also occur in this area. these plots are therefore not completely representative of the lindanda replicate or the satara series. the area covered by significant portions of plot 2 and plots 3, 4, 5, 6 and 13 of the lindanda replicate is characterised by soils of the swartland and shortlands forms and features woody plants such as s. birrea, a. nigrescens, acacia tortilis, et cetera; it is comparable with the rest of the series. the western part of the replicate is associated with letaba basalt. the marheya replicate is considered to be representative of the satara land type, but a drainage line through plot 9 also influences plots 8 and 10 with darker, calcareous soils representative of very shallow mispah–glenrosa–maya forms (soil classification working group 1991) (figure 4c). the nwanetsi replicate is regarded as very representative of the satara land type, as most of the area is flat and characterised by red, structured, clay soils, which are characterised as moderately deep shortlands–swartland forms (soil classification working group 1991) (figure 4d). table 5a and 5b illustrates the overall representivity scores for each replicate and individual plot within satara. figure 4: description of the soils types on the satara replicates (a) satara (b) lindanda (c) marheya and (d) nwanetsi. for the complete list of soil names and abbreviations see online appendix 2. table 5a: the overall representivity scores for each replicate within satara. table 5b: the overall representivity scores for each plot within the four replicates in satara. mopane three of the four replicates are situated in the mopane shrub vegetation, with relatively shallow, calcareous, dark, clay soils associated with the picrite lavas of the letaba basalt formation. the dzombo replicate is regarded as the best replicate as it is very homogeneous with respect to soils and slope and is characterised by soils from deep milkwood and shallow bonheim forms (soil classification working group 1991) (figure 5a). two of the replicates – mooiplaas (figure 5b) and tsende (figure 5c) – have parts that represent slightly incised, sloping land where soils are shallower and the mopane has a tendency to grow taller. sometimes, drainage lines or depressions represent deeper and vertic clay soils, for example arcadia (soil classification working group 1991). on the shallow or rocky areas, different vegetation may be found, namely trees or shrubs such as terminalia prunioides, c. apiculatum and acacia exuvialis, as well as grasses such as heteropogon contortus and aristida spp. the nshawo replicate is located in an area where three types of parent material occur and where the soils and vegetation are therefore fairly heterogeneous over the replicate as a whole. the nshawo replicate is located on the interface of the letaba basalt and sabie river basalt formations, as well as on the edge of the shawu vlei area where very deep alluvial soils occur. soils associated with the letaba basalt formation are dark, calcareous clays, often shallow as on the western side of the replicate, whereas those associated with the sabie river basalt formation are reddish and usually not calcareous. on the reddish soils, trees or shrubs such as combretum imberbe and albizia harveyi may be dominant. on the shallow calcareous clays, mopane may dominate and on the alluvial clay deposits thickets of mopane occur (figure 5d). table 6a and 6b illustrates the overall representivity scores for each replicate and individual plot within mopane. figure 5: description of the soils types on the mopane replicates (a) dzombo (b) mooiplaas (c) tsende and (d) nshawo. for the complete list of soil names and abbreviations see online appendix 2. table 6a: the overall representivity scores for each replicate within mopane. table 6b: the overall representivity scores for each plot within the four replicates in mopane. discussion top ↑ present long-term studies, such as those in the serengeti (stronach & mcnaughton 1989), the knp (van wilgen et al. 2007), and yellowstone national park (romme et al. 2011) usually begin as short-term academic projects and are not envisioned to continue for decades. however, they progress over time as new events lead to further research, with the combined data and results leading to insights that were unintended and support additional questions and projects for further understanding (sinclair et al. 2007). the few long-term ecological experiments in south africa are crucial to understanding and interpreting the complex ecosystem behaviour involving slow and rapid changes at multiple ecosystem states which only became apparent over a period of several decades (thirgood et al. 2007). as previously outlined, the original objective of the experiment was to determine the effect of fire frequency and season on four of the major vegetation types in the knp (biggs et al. 2003) and therefore were spaced across the landscape to maximise the representivity of the landscape, particularly with reference to the underlying soils (van der schijff 1958). the results from this geomorphic and soil description of the individual plots and replicates in the experiment has indicated that not all plots within replicates and not all replicates within the landscape can be considered homogeneous treatments. it is for this reason that researchers undertaking their projects within the ebps, can, for statistical analyses, exclude any outlier replicates (based on the soil and geomorphic description) from their sampling protocol. however, it must also be noted that it is the outlier replicate (least representative of the surrounding landscape) that encompasses the variation and heterogeneity that is inherent within knp ecosystems and therefore it should not be completely ignored. finally, the knp’s long-term fire experiment has been instrumental in highlighting that ecosystems are not static and management should not aim to maintain the status quo; rather, it should allow natural change to take place within an adaptive management framework. this understanding could not have been achieved without the array of research and projects that have been undertaken on the ebps over the past five decades (van wilgen et al. 2007) acknowledgements top ↑ we thank the south african national parks for access to data and sandra macfadyen for the preparations of the figures. the role of the fire team that maintained the fire experiment (especially andre potgieter) is also gratefully acknowledged, as are the valuable inputs provided by harry biggs. comments from the two reviewers are also appreciated. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this article. authors’ contributions f.j.v. 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h., 2003, ‘the abiotic template and its associated vegetation patterns’, in j. du toit, k.h. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, new york. webber, n.w., 1979, ‘the effects of fire on soil/plant ecological relationships in the southern part of the kruger national park: a study on soil geography’, msc thesis, department of geography, university of south africa. article information authors: elizabeth h. bouchard1 lawrence e. little2 cassandra m.l. miller3 susan m. rundell4 elana m. vlodaver5 kristine maciejewski6,7 affiliations: 1department of environmental science, wheaton college, united states 2department of biology, washington university, united states 3department of biology, grinnell college, united states 4department of ecology and evolutionary biology, yale university, united states 5department of biology, bowdoin college, united states 6organisation for tropical studies, cape town, south africa 7department of biological sciences, university of cape town, south africa correspondence to: kristine maciejewski email: krismacski@gmail.com postal address: 407 rusdon park, college road, rondebosch 7701, south africa dates: received: 25 may 2015 accepted: 17 aug. 2015 published: 08 oct. 2015 how to cite this article: bouchard, e.h., little, l.e., miller, c.m.l., rundell, s.m., vlodaver, e.m. & maciejewski, k., 2015, ‘undeclared baggage: do tourists act as vectors for seed dispersal in fynbos protected areas?’, koedoe 57(1), art. #1323, 9 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1323 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. undeclared baggage: do tourists act as vectors for seed dispersal in fynbos protected areas? in this original research... open access • abstract • introduction • research method and design    • study area    • interview survey    • plant seed collection and categorisation    • alien plant survey    • data analysis • results    • seed survey and collection    • vegetation survey • discussion • conclusion • acknowledgments    • competing interests    • authors’ contributions • references • appendix 1 abstract top ↑ encroachment by alien species is the second greatest threat to biodiversity worldwide. as south africa's cape floristic region has a botanical endemism of nearly 70%, conservation efforts are a high priority. estimates suggest that alien species cost the country over r6.5 billion per year. despite significant research on alien species dispersal, the role of tourists as seed dispersers requires further exploration. to investigate the potential role tourists play in introducing alien seeds into protected areas, long-bristle brushes were used to scrape seeds off the shoes of hikers, dog walkers and cyclists, as well as the wheels of mountain bikes and dogs themselves, upon entering the silvermine nature reserve section of the table mountain national park in the western cape province, south africa. in addition, a vegetation survey was conducted. this comprised 18 transects at various distances from the recreational paths in the park, and used a prioritisation ranking system that identified the alien species of greatest concern. it was concluded that the greatest number of alien plant species could be found along dog paths, in comparison to the hiking trails and cycling trails. this corresponded to the findings that dog walkers had the highest incidence of seeds on their shoes, suggesting that tourists were possibly dispersing seeds from their gardens. alien species significantly covered more of the vegetation transects closer to the trails than they did in transects further into the matrix. because more alien species were present in areas susceptible to human disturbance, the data suggest that tourists can act as vectors for alien seed dispersal. these findings emphasise the need for active tourism management in line with the south african national parks biodiversity monitoring programme in order to prevent the introduction and spread of alien species into south africa's protected areas. conservation implications: tourism is the main source of revenue for south african national parks, and one of the organisation's principal goals is to create a tourism management policy conducive to conservation. this research explores the potential role that tourists may play in the introduction of non-native species into a protected area, thereby providing novel information that could assist managers in the sustainable management of protected areas. introduction top ↑ south africa's cape floristic region is a world-renowned hotspot for botanical biodiversity, home to some 9000 plant species, 70% of which are endemic (forest et al. 2007). however, alien species – the second greatest threat to biodiversity worldwide (richardson & van wilgen 2004) – jeopardise the long-term survival of this region's staggering diversity. an alien species can be defined as a non-native species, the introduction of which has the potential to cause health-related, economic or environmental damage (swearingen et al. 2010). in south africa alone, nearly 200 plant taxa are legally recognised as ‘alien’ (wilson et al. 2013). these species degrade ecosystems through the excessive consumption of resources, the alteration of fire regimes, and the disruption of soil stability and composition (richardson & van wilgen 2004). alien species cost south africa an average of r6.5 billion per year, a sum set to grow by an order of magnitude in the near future (wilson et al. 2013). the management of alien species is, therefore, vital to the biological and economic well-being of the country (wilson et al. 2013). given that south africa is home to 3 of the world's 25 biodiversity hotspots, managing alien species presents a particularly daunting task due to the already established vulnerability of these areas (myers et al. 2000). south africa has one of the worst problems with alien flora of any country in the world, and managers have never successfully eradicated an alien plant species nationwide (richardson & van wilgen 2004; wilson et al. 2013). one of the challenges of containment and eradication is that, while managers may successfully clear above-ground individuals from a plot, the alien seed bank can persist in the soil for many decades, encouraging reinvasion (wilson et al. 2011). accordingly, a more effective management strategy is the prevention of seed dispersal (wilson et al. 2011). the introduction of any new alien species into a protected area is against the mandate of south african national parks (sanparks), even when the potential for a particular species to become invasive is unknown (foxcroft 2009). as such, one of the key functions of the sanparks biodiversity management programme for invasive alien species is the monitoring of potential introduction pathways (mcgeoch et al. 2011). research on such pathways has expanded from such vectors such as animal transport and wind to include the role of tourists (pickering et al. 2011). tourists are often cited as playing a role in the dispersal of seeds through activities such as hiking and mountain biking. however, there is only a small body of evidence to support these claims. to date, only a single study on seed dispersal by tourists has been conducted in the entire continent of africa (pickering & mount 2010). circumstantial evidence suggests that alien species are found most commonly near paths in protected areas, and a correlation exists that links an increased visitor presence to a greater number of alien species in protected areas in southern africa (usher 1988). this emerging field has no standardised techniques for collecting seed from clothing, vehicles, animals, or equipment used by tourists, limiting the utility of comparisons between data sets (pickering & mount 2010). despite the dearth in research on the subject, it is likely that tourists do have a profound impact on the dispersal of alien seeds. hikers, for instance, have been shown to carry seeds on their clothing for an average of 13 km, and much further if their travels include a car, aeroplane, train or boat (ansong & pickering 2014a). protected areas are usually associated with minimal artificial disturbance, reducing the impact of certain other pathways, which highlights the importance of managing this particular pathway of introduction (pickering & mount 2010; usher et al. 1988). tourism and tourism-related activities comprise upwards of 80% of sanparks’ annual revenue, distinguishing it as one of the most successful protected area networks in the world (biggs et al. 2014). nonetheless, there are ecological risks associated with an increase in visitor numbers, which highlights the need for environmentally responsible tourism (cini & saayman 2014). one of the goals of sanparks’ strategic adaptive management process is to find a balance between the goals of increased ecotourism and the needs of conservation (biggs et al. 2014). achieving this goal will require greater investigation into the role that tourists play in ecosystem processes, and integrating the results of such studies into tourism management (biggs et al. 2014). a study on the role that tourists play in the dispersal of alien plants presents an opportunity to inform and improve tourism management practices in a way that protects the native flora of south africa's iconic national parks. the aim of this study is to investigate the role that three categories of tourists – hikers, dog walkers and cyclists – play in alien seed dispersal in a fynbos protected area. by sampling the shoes of visitors, dogs and bicycles for seeds, the role that visitors play in introducing seeds into a protected area is assessed, as well as the types of tourists that act as vectors for seed dispersal. at the same time, vegetation transects at different distances from trails used by these tourists illustrate the distribution of established alien species within the protected area. since roads have been shown to act as propagules for alien species in protected areas globally (pauchard & alaback 2004), a hypothesis is put forward that greater tourist disturbance closer to trails leads to a greater percentage of cover of alien species in these areas as opposed to areas further into the matrix. research method and design top ↑ study area the cape peninsula has a mediterranean climate, with an average winter rainfall of approximately 2000 mm per year (trinder-smith 2006). furthermore, the region is characterised by strong gulf winds, blowing primarily south-east to north-west. the table mountain national park covers 57 km2 and offers over 650 km of hiking trails (clarke, mackensie & merry 2013). the silvermine nature reserve section of the table mountain national park, in cape town, western cape province, south africa (18°40's, 34°07’e), is known for its diverse fynbos vegetation (figure 1). the soil is very sandy and nutrient poor, with limited phosphorus and nitrogen availability (trinder-smith 2006). this section of the park provides various trails for visitors to use, including paths for dog-walking, cycling and hiking. of the three trails considered in this study, dogs were restricted to the dog-walking trail, as were cyclists to the cycling trail. hikers, however, could walk on all three trails. figure 1: location study area within the silvermine section of the table mountain national park, western cape province, south africa. the yellow, blue and red rings represent the vegetation transects conducted along the dog-walking, hiking and cycling trails respectively. the stretches of the dog-walking and cycling paths investigated in this study were wider than the hiking path. furthermore, the hiking path utilised in this study was one of the most commonly trafficked trails in the park, which also serves as the start to many other hiking routes. interview survey data were collected for 3 days in early february 2015 – the end of the dry season – during peak visiting hours (between 08:00 and 11:00). whilst it was partly cloudy during the first 2 days of data collection, it rained heavily on the third day, resulting in a decrease in the number of participants and overall park traffic on that day. survey stations were set up at the trailhead of three different paths in silvermine: a hiking path (18°40's, 34°08’e), a cycling path (18°41's, 34°08’e) and a dog-walking path (18°40’ s, 34°08’ e). tourists entering the park were classified into three categories: hikers, dog walkers and cyclists. each participant was asked a series of questions about their activities (see appendix 1). the data were further assessed to determine tourists’ role in seed dispersal, and the introduction and potential source of alien vegetation into protected areas. plant seed collection and categorisation at the three survey stations, prior to entering the park, each participant stepped into a 40 cm × 30 cm plastic container. here, the soles of shoes were swept, using a long-bristle brush to collect any seeds the individual may have been carrying. the debris was collected into individual plastic bags for later classification. this same procedure was followed to brush off the entirety of the front tyre of each bicycle that was surveyed. the front legs and flank of each dog surveyed was brushed using a dog brush and all the debris collected. each debris sample was then examined under an olympus ch light microscope (model lsk [20w], olympus corporation of america, new hyde park, ny, usa). all of the seeds in these debris samples were counted and categorised by morph. an attempt was made to germinate the seeds in order to identify the species from which each morph had come. however, this was unsuccessful, preventing seed identification beyond the original morph categorisation. alien plant survey in order to examine the occurrence of alien plants in protected areas, eighteen 10-m transects were set up along recreational trails in silvermine. alien species were defined as those that are non-native to south africa. to compare the potential for different human activities at dispersing alien seeds, three paths were selected: a dog-walking path, a hiking path and a cycling path. six transects were set up in two groups on each trail. using a measuring tape, the first set of 10-m transects was set up 5 m from the trailhead; the second group of transects, 100 m from the trailhead (figure 2). to evaluate the relationship between the percentage of cover of plant species, and distance from the trail, three vegetation transects were conducted: one along the edge of the trail; one 1 m away; and a third, 10 m away from the trail (figure 2). figure 2: experimental design of vegetation transect survey conducted in the silvermine section of the table mountain national park in order to investigate the distribution of invasive plants. three 1-m2 quadrats, each represented by a grey square, were sampled along a 10-m line transect. this technique was repeated at 5 m (a, b and c) and 100 m (a, b and c) from the trailhead. at both of these locations, transects were established at the edge of the trail (1), 1 m into the matrix (2) and 10 m into the matrix (3). at each vegetation transect point, a 1 m2 quadrat was used to examine the vegetation cover every 5 m along each line transect, for a total of three plots. the percentage of cover for all plant species within the quadrats was estimated by identifying the known species to at least genus level and collecting samples that were later identified at a laboratory. to approximate the relative abundances of each plant, the estimated proportion of the plot covered by each particular species was done visually. to assess the immediacy and severity of the risks posed by the identified alien species, the prioritisation system was used to score each species and to rank them accordingly (robertson et al. 2003). a prioritisation score was calculated for each species based on an assessment of the impact the species poses in a range of categories. these included long-distance dispersal mechanism, invasive elsewhere, density, biodiversity, impact on water resources, and poison status. each category score was first standardised by dividing it by the potential maximum (robertson et al. 2003). these category scores were summed up and divided by the number of categories, resulting in a weighting score. species were then ranked according to the product of their total prioritisation score and total confidence interval. a score of 5 represented a species that posed the greatest threat, whilst a score of 0 demarcated no threat. data analysis to determine the proportion of seeds found per vector, and to account for sampling effort, the number of seeds collected per vector was divided by the sample size of each vector. a permanova analysis was conducted to analyse the difference between percentage of cover of alien plant species between trail types, distance from the trailhead, and distance into the matrix (primer software version 6.1.5, clarke & gorley 2006). results top ↑ seed survey and collection in order to examine their potential as vectors for seed dispersal, a survey and debris collection was conducted among 68 participants entering silvermine nature reserve. of these 68 participants, 10 were cyclists, 12 were dog walkers (with 19 dogs in total) and 46 were hikers. all of the cyclists and dog walkers lived in the cape town area. of the 46 hikers, 16 were either non-local south africans, living in johannesburg or port elizabeth, or international visitors from scotland, england or germany. fifty of the sampled visitors were not carrying any seeds that could be detected. from the remaining 18 participants, a total of 41 seeds and 17 unique seed morphs were isolated (table 1). it was not possible to identify the seeds to species level, and it therefore remains unknown if these were native or non-native seeds. table 1: the total number and morph of seeds collected from tourists (cyclists, dog walkers and hikers) entering the silvermine section of the table mountain national park, and their potential source of origin. of the participants who were carrying seeds, 16 were from the cape town area, one was visiting from england (9 seeds) and one from germany (1 seed). the english hiker carrying 9 seeds had last worn his shoes in london, and the german hiker had last worn his shoes at lion's head, a different part of the table mountain national park. no seeds were found on bicycle tyres, but seeds of two distinct morphs were found on a single cyclist's shoes (table 1). when sampling effort and population sample size were taken into account, it was discovered that a larger proportion (75%) (9 out of 12 that were sampled) of dog walkers carried seeds than any other vector, comprising 42% of seed morphs (table 1; figure 3). hikers were shown to be the second highest seed disperser type. of the hikers, 56% (27 out of 46 that were sampled) carried seeds, representing 19.6% of the total seed morphs. only 20% of the cyclists (2 out of 10 that were sampled) carried seeds on their shoes, whilst 15% of the dogs (3 out of 19 that were sampled) carried seeds (figure 3). figure 3: the proportion of seeds collected from each vector, calculated from the number of seeds divided by vector sample size, to account for sampling effort and population sample size. the black line represents the number of different seed morphs found per vector as a proportion of total seed morphs found. vegetation survey a total of 54 plots in 18 line transects along the three selected trails in silvermine were surveyed, and a total of seven invasive plant species were documented (table 2). the most frequently occurring invasive species were briza maxima (greater quaking grass, found in 17 quadrats), lagurus ovatus (hare's tail, found in 7 quadrats) and stenotaphrum secundatum (coastal buffalo grass, found in 6 quadrats) (table 2). the dog path had the greatest number of invasive plant species (n = 6), whilst the mountain bike path had the fewest invasive plant species (n = 2). furthermore, the dog-walking path had the highest average percentage of cover of invasive species (13.8%), whilst the hiking path had 9.6% and the cycling path had only 3.0% average percentage of cover. table 2: the percentage of cover of plant species identified to family or genus level, averaged from three 1-metre2 quadrats used in the vegetation transects along the various trails in the silvermine section of the table mountain national park. the permanova analysis demonstrated a significant increase in the percentage of cover of alien plant species in the plots located at the trail edge compared to further into the vegetation matrix (pseudo-f = 3.392, p < 0.005, n = 18; figure 4). there was also a significantly higher percentage of cover of alien plants along the dog-walking path than the cycling path (pseudo-t = 2.592, p < 0.05, n = 12). there was no significant difference, however, between plots taken 5 m and 100 m along the path for all trails. figure 4: the average proportional cover of alien plant species along three different paths in the silvermine section of the table mountain national park, taken at 0 m, 1 m and 10 m into the matrix and measured at 5 m and 100 m from the trailhead. the highest ranking species with regards to the threatening status of alien species identified in this study obtained a score of 2.74 (giant cane, arundo donax), whilst the lowest ranking species (l. ovatus) scored 1.56 (table 3). species that had high prioritisation and confidence scores, such as a. donax, were of most concern (confidence score = 11.16; table 3). the high ranking of this species was mainly due to its severe impact on biodiversity (giessow et al. 2011; guthrie 2007) and its negative economic impact. this was followed closely by s. secundatum (prioritisation score = 2.54, confidence score = 9.479) and b. maxima (prioritisation score = 2.22, confidence score = 9.527) (table 3). table 3: the prioritisation scores of alien plants species surveyed on the trails in the silvermine section of the table mountain national park. the scores were calculated using the alien species prioritisation system. discussion top ↑ this study demonstrates that park visitors, in the form of dog walkers, hikers and cyclists, act as seed dispersers in protected areas, corroborating existing research. one study found that humans working in a series of meadows carried seeds from this location on their clothing for an average distance of 13 km (auffret & cousins 2013). furthermore, it has been demonstrated that seeds can remain on hikers’ shoes for distances as great as 5 km (wichmann et al. 2009). the seeds of 16 taxa were found in a single study specifically investigating dispersal via shoes (clifford 1956). the vehicles considered in studies as a means of seed dispersal, however, have been exclusively motorised, and the only animals investigated have been donkeys and horses (ansong & pickering 2014a; pickering & mount 2010). the finding that the areas closest to human activity (i.e. near the edge of the trails) are also the areas that are most susceptible to invasion from non-native plants suggests a path-user-mediated mechanism for alien seed dispersal. this evidence is strengthened by a correlation between the relative importance of different potential vectors of seed dispersal and the percentage of cover of alien species on different paths. for instance, dog walkers had the highest incidence of seeds on their shoes; correspondingly, the dog-walking path had the highest percentage of cover of alien plants. likewise, cyclists carried virtually no seeds, and the cycling trail had the lowest percentage of cover of alien species. the difference in tendency for different categories of tourists for carrying seeds into the park, therefore, suggests a possible explanation for the differences in the percentage of cover between paths. whilst the sample sizes used in this study were too small to establish an unambiguous relationship between seed incidence on trail users and the percentage of cover of alien plants along these trails, it is an observation worthy of further investigation. to confirm such a relationship, future studies need to identify the species of collected seeds, particularly since the attempts in this study to propagate and identify seeds were unsuccessful. even though the non-native species identified along the sides of the trails are all classified as ‘alien’, they all play different roles in invading or threatening the environment. in this study, the alien cane species a. donax represented the largest alien risk compared to the other species, which was also found by robertson et al. (2003) where this species received a total prioritisation score of 3.08. arundo donax, a tall perennial cane native to eastern asia, has been introduced into many areas for erosion control (mariani et al. 2010). however, it has a tendency to outcompete native plant species, reduce wildlife habitat and modify river hydrology (giessow et al. 2011). invasion by a. donax alters biomass, thereby increasing the fuel load and creating a potential fire hazard (guthrie 2007). even though this species only covered a small area in the hiking trail 100 m away from the trailhead, a monitoring programme is needed to prevent further invasion. this species may easily spread to other areas since reproduction occurs almost entirely from rhizomes and stem fragments (boose & holt 1999). significantly, a number of the invasive species that were found along the paths’ edge are used in southern africa for decorative purposes, including b. maxima and l. ovatus (van oudtshoorn 2002). furthermore, pauchard and alaback (2004) found that the land-use type surrounding protected areas influences the numbers and types of plants that are able to invade protected areas. the highly developed landscape surrounding silvermine is likely rich in decorative plants, considering the large anthropocentric role in determining the ecological makeup of urban areas (aronson et al. 2014). together, these findings suggest that tourists are possibly dispersing seeds from their gardens, a source of alien species now recognised as one of the most challenging to combat (mack 2003; mack & lonsdale 2001). the spread of ornamental grasses such as b. maximus and l. ovatus has been attracting greater attention in south africa (milton 2004), and researchers have begun to investigate the threat posed to south african protected areas by ornamental plant species (foxcroft, richardson & wilson 2008). it seems likely that this same mechanism is operating in silvermine. priorities for future studies should include more comprehensive seed collection techniques. through the course of this study, it was observed that more seeds appeared to become caught on the socks of researchers rather than on the shoes. this observation suggests that clothing other than shoes could be important mechanisms for dispersal. the socks of participants were not a priority in this study because socks are typically washed between each use. it was therefore less likely that more seeds could be brought into silvermine on socks than on shoes, which was the primary focus of this study. socks may play a more important role in dispersing seeds within protected areas than in introducing seeds into those protected areas. in addition, car tyres represent a potential vector for invasive seeds that remains unexamined in the context of south african protected areas. future studies should take this into account when considering the method of seed collection – it is suggested that the clothes and vehicles of participants are sampled in addition to shoes. to limit the spread of non-native plant species into pristine ecosystems, protected area managers must take an active role in monitoring tourists in accordance with the sanparks biodiversity management programme for invasive alien species (mcgeoch et al. 2011). tourism is the main source of revenue for south african protected areas, and a major goal for sanparks is to encourage ecotourism that does not compromise the agency's commitment to conservation (biggs et al. 2014). some research on the means of minimising tourist-mediated seed dispersal has already been conducted in other countries. a study by ansong and pickering (2014b) found that the vast majority of tourists entering an australian protected area were aware of the potential capacity for unintentionally dispersing invasive plant seeds. over half of the individuals surveyed by ansong and pickering (2014b) supported greater funding toward non-native species removal, and a vast majority of respondents either supported or felt neutral towards an initiative requiring tourists to remove seeds from their clothing and equipment before entering the park (ansong & pickering 2014b). these results indicate that an educational campaign about seed dispersal by tourists could have widespread public support (ansong & pickering 2014b). primarily, these efforts should focus on the correct form of disposal for seeds attached to clothing, shoes, dogs or recreational equipment, since many tourists interviewed said they threw seeds away indiscriminately (ansong & pickering 2014b). information posters at trail heads about the dangers of alien species and the correct method for the removal of seeds could be one easy method of raising awareness and decreasing the incidence of unintentional dispersal. shoe-sweeping stations in trailhead parking lots could similarly provide a cost-effective and convenient method for tourists to remove seeds from their shoes. the implementation of such measures could help to prevent the further spread of non-native plants species into silvermine nature reserve, and elsewhere in south africa's national parks. conclusion top ↑ this research adds to the small body of existing evidence concerning the spread of invasive seeds by tourists. tourist surveys and vegetation transects conducted in the silvermine nature reserve section of the table mountain national park demonstrate a clear link between tourist disturbance – measured as distance from the path into the matrix – and percentage of cover of invasive plants. at the same time, the path used by dog walkers, who were found to have the highest load of seeds, have the highest overall percentage of cover of invasive species. these results demonstrate that tourists can act as vectors for invasive alien species dispersal, highlighting the need for active tourism management in line with sanparks’ biodiversity monitoring programme. acknowledgments top ↑ the authors would like to thank dr laurence kruger for his assistance in plant identification. they would also like to thank the organisation for tropical studies and sanparks for their roles in facilitating this project. competing interests the authors declare that they have no financial or personal relationships, which may have inappropriately influenced them in writing this article. authors’ contributions k.m. 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http://dx.doi.org/10.1098/rspb.2008.1131 wilson, j.r.u., gairifo, c., gibson, m.r., arianoutsou, m., bakar, b.b., baret, s. et al., 2011, ‘risk assessment, eradication, and biological control: global efforts to limit australian acacia invasions’, diversity distribution 17(5), 1030–1046. http://dx.doi.org/10.1111/j.1472-4642.2011.00815.x wilson, j.r.u., ivey, p., manyama, p., nänni, i., 2013, ‘a new national unit for invasive species detection, assessment and eradication planning’, south african journal of science 109(5/6), art. #0111, 13 pages. http://dx.doi.org/10.1590/sajs.2013/20120111 appendix 1 top ↑ interview questionnaire presented to participants at the entrance of visitor trails in the silvermine section of the table mountain national park where are you from? what are you doing? cycling walking/hiking/dog-walking where did you last: wear your shoes? ride your bike? walk your dog? abstract introduction rwandan culture history and forest the context of buhanga sacred forest material and methods results discussion conclusion acknowledgements references appendix 1 about the author(s) runyambo irakiza africa rice center (africarice), east and southern africa, weed sciences, dar es salaam, tanzania association pour la conservation de la nature au rwanda (acnr), kigali, rwanda minani vedaste institute of scientific and technological research, national herbarium of rwanda, huye, rwanda bizuru elias college of science and technology, university of rwanda, rwanda brigitte nyirambangutse college of science and technology, university of rwanda, rwanda nsengimana joram serge association pour la conservation de la nature au rwanda (acnr), kigali, rwanda ndimukaga marc association pour la promotion des etudes d’impacts environnementaux au rwanda (apeier), kigali, rwanda citation irakiza, r., vedaste, m., elias, b., nyirambangutse, b., serge, n.j. & marc, n., 2016, ‘assessment of traditional ecological knowledge and beliefs in the utilisation of important plant species: the case of buhanga sacred forest, rwanda’, koedoe 58(1), a1348. http://dx.doi.org/10.4102/koedoe.v58i1.1348 original research assessment of traditional ecological knowledge and beliefs in the utilisation of important plant species: the case of buhanga sacred forest, rwanda runyambo irakiza, minani vedaste, bizuru elias, brigitte nyirambangutse, nsengimana joram serge, ndimukaga marc received: 21 aug. 2015; accepted: 05 feb. 2016; published: 22 july 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract traditional ecological knowledge is an integrated part of the african people and indeed the rwandese for cultural purpose. buhanga sacred forest is a relict forest of tremendous ecological importance to rwandan society located in musanze district. the aim of this study was to assess the traditional ecological knowledge and belief in the utilisation of some important plant species for the conservation of buhanga sacred forest. ecological information about ethnomedicinal and traditional practices were collected following structured questionnaire through interview involving eight traditional healers and three focus group discussions. data were collected from the natural habitats, home gardens, farmlands and roadsides of buhanga sacred forest. a total of 45 botanical taxa belonging to 28 families were reported to be used by the local community. species such as brillantaisia cicatricosa and senna septemtrionalis were the popular species cited by traditional healers to treat human and animal diseases and ailments, respectively. the results of the study indicated that because of the cultural norms and values associated with the sacred forest, this has led to non-exploitation. the study presents key sites and plant species in which their use and belief can lead to their conservation. however, not only is it imperative to conserve traditional local knowledge for biocultural conservation motives but there is also need to train traditional healers on how to domesticate indigenous species as conservation measure because some species have become susceptible to extinction. conservation implications: highlighting indigenous species investigated in this research will provide a powerful tool for ensuring biodiversity conservation through community participation in a country of high population density in africa. some plant species that provided satisfactory local health traditions among communities surrounding buhanga can contribute as good material for further research in rwanda. introduction traditional ecological knowledge is defined as a cumulative body of knowledge, practice and belief. it is a mutual relationship between living beings (including humans) and its environment which evolves by adaptive processes and are handed down through generations by cultural transmission (berkes 1999). such knowledge has contributed to conservation of biodiversity (gadgil, berkes & folke 1993), rare species (colding 1998) and protected areas (johannes 1998) as well as to sustainable resource use (berkes 1999) from one area to another. in africa, there are important elements to take into account regarding indigenous or traditional spirituality towards the nature. the value of traditional knowledge of natural resources is widely recognised among indigenous people because it was accumulated and transferred orally across generations. according to charnley, paige and jones (2007) and berkes, colding and folke (2000), the integral success of the traditional ecological knowledge into biodiversity conservation depends on active participation of the knowledge holders, such as indigenous communities and institutions. to avoid loss of biodiversity, traditional rules play an important role when developing conservation strategies for rare species by limiting illegal activities such as selective trees felling (ormsby 2013). some sacred species constitute excellent totems. totems are often plants or animals that are a stand-in of its owner and their presence dissuade from illegal access to sacred areas. furthermore, these sacred species contribute significantly in conservation by acting as umbrella species to the overall ecosystem (ndimukaga 2009). sacred place functions under the principle of faith and fear of god and other supernatural forces. this principle requires honesty, respect of ancestors and observation of moral values (kamga-kamdem 2008). rwandan culture history and forest in many sub-saharan african societies, the traditional ecological knowledge among indigenous people is synonymous with spirituality, which contributed to sustain the sacred forests. rwanda, one of the smallest countries in the world with only 26, 338 km2, has a rich forest heritage. the value of rwandan culture is based on oral history, ecology and nature, geographical sites and by igitaramo. igitaramo constitutes a gathering of families, communities and friends with songs and dances. during pre-colonial time, rwandan society was inherently linked to forest, which hides great lessons concerning rites, ancestral customs, taboos, arts, crafts, music, dance and many more. such integral links led to the emergence of traditional medicine (karangwa 1997). the rwandese are aware of the existence of these traditional healers who provide home-based healthcare. locally, they are known as abavuzi gakondo or abapfumu. the former denotes traditional healers involved in the use of plants and the latter denotes those who are involved especially in the ritual component of traditional medicine (rwangabo 1993). but it has been said that sometimes abapfumu use leaves, roots or barks of plants to protect their clients in exchange for money (adenkule 2007). as a matter of fact, traditional ecological knowledge is still very much alive in rwanda because it contributes significantly to solve health problems. in rwanda, cutting of certain tree species is seen as a taboo as well as killing of indigenous animal such as primates, elephants, leopards, birds, etc. according to kimenyi (1989), this conservation emphasised the symbiotic and interdependence relationship between people, plants and animals. it is for these reasons that cultural values are an essential component of every society and they act as checks and balances in the management of natural resources (verschuuren et al. 2010). the context of buhanga sacred forest buhanga sacred forest is a relict forest of tremendous ecological importance to rwandan’s society for cultural purposes. it lies in volcanic belt and has one of the highest population density in rwanda and africa, with up to 500–700 people per km2 (nisr 2012; plumptre, masozera & vedder 2001). buhanga relict forest contains sacred water-spring, marshland, medicinal plants, native trees and even a last refuge for arboreal, migratory birds. for example, african pitta pitta angolensis is a migrant bird that breeds in central tanzania, malawi, southeast democratic republic of congo, eastern zambia, zimbabwe and possibly northern south africa that was sighted in buhanga sacred forest (birdlife international africa partnership e-bulletin 2008). unfortunately, reports indicate that this relict forest is facing many threats as a result of human interference on the environment in various ways, such as harvesting of firewood and agriculture (minitere & cgis-nur 2007; rema 2009). in fact, high pressure of disturbance of natural reserves in rwanda was noticed until the resettlement of refugees after the genocide during the year 1997–1998 where people both settled and farmed within many natural reserves (plumptre et al. 2001). it should be mentioned that in most cases natural resources are still viewed as limitless in the mind of the resource-poor local communities in rwanda as a source of income. however, it should be seen furthermore that without any practical actions for sustainable utilisation of so-called ‘limitless resources’ and usage of traditional ecological knowledge, there is a risk of genetic loss and lack of traditional information from generation with time. to address the issue of loss of this critical cultural forest in rwanda, the goal of this study is to document the traditional ecological knowledge and belief in the utilisation of some important plant species of buhanga sacred forest. hence, there is a need for such information and documentation with the help of local communities to develop comprehensive and useful strategies for public awareness but also to formulate recommendations for monitoring natural resources for better future (figure 1). figure 1: view of buhanga sacred forest. material and methods geo-ethnographical overview of the study area buhanga sacred forest is an archaic forest located in northern province, district of musanze, nkotsi sector and bikara cell, at the right side of the road that leads towards vunga business centre. the site is situated at approximately 114 km from kigali, the capital city of rwanda and covers an area of 33.252 ha, of which 16.0011 ha is still intact and located between 1°34’00” and 1°34’30” south latitude and 29°34’30” and 29°37’30” east longitude at an altitude ranging between 1623 m a.s.l. and 1657 m a.s.l. geo-climatically the area falls within the tropical rainforest of the volcano massif where climate is controlled by various factors such as altitude, latitude and vegetation. the climate is cool and wet and temperatures fluctuate around 20 °c during the day. most of the lands surrounding buhanga sacred forest are used for agriculture subsistence. near subsistence agriculture, coffee, tea and pyrethrum plantations are prominent land uses as cash crop generating local employment. projects funds from government and international non-governmental organisations have been used for a range of activities from environment protection (tree planting, soil erosion control and beekeeping) to limit access in the sacred forest (figure 2). figure 2: map of the study area. ethnobotanical survey our study was carried out during the period of 26–31 may, 2008. a reconnaissance survey was conducted in abadahemuka cooperative at nkotsi-bikara, musanze district. in rwanda, most of the traditional healers are grouped into cooperatives under the supervision of the ministry of health in close collaboration with the institute of scientific and technological research. the traditional healers provided the information about traditional healing practices for both humans and animals, while the elders provided information regarding rituals and beliefs surrounding the forest. the information regarding the use of medicinal plants was collected following structured questionnaire through interviews involving eight traditional healers (five men and three women) aged between 41 and 80 years. these traditional healers were recognised by the local government authorities and selected based on their reputation on herbal medicinal uses. plant species mentioned by traditional healers were visited inside the relict forest and home gardens to verify the reliability of data obtained during interviews (alexiades 1996; gerique 2006). the recorded field data include the following: collection number, plant local name, scientific name, habit, plant parts used, mode of preparation, habitat and locality (martin 1995). the information regarding the traditional practices and utilisation of buhanga sacred forest were captured through three focus discussion groups. each discussion group comprised 15 persons (10 men and 5 women) who were natives of the nkotsi-bikara village aged between 50 and 80 years; inclusion was based on a good knowledge of buhanga sacred forest history. during the discussion, each member of the group took about 5–8 min to talk about the importance of cultural knowledge and use of buhanga sacred forest. species information provided either during the interview or group discussion were recorded. when a species was physically observed in home garden and not in the forest and vice versa, the information regarding that species was collected. species recognised only through vernacular names without physical identification either in home garden or in the forest was considered unreliable and rejected. species botanical identification was carried out in the national herbarium of rwanda using voucher specimens, identification keys, field guides and the flora of rwanda (troupin 1966, 1978, 1982, 1983, 1985, 1988; and fischer & dorothee 2008) (see picture showing different sites of study area in appendix 1). data analysis validation of plant names, family and plant authority were carried out using the royal botany garden and missouri botanic garden plant names database (http://www.theplantlist.org). collected data were analysed using descriptive statistics. the determination of relative frequency of citation of reported medicinal plant species was done using the following formula: rfc (%) = (fc/n) × 100             [eqn 1] where, fc is the number of informants mentioning the use of medicinal plant species and n is the total number of informants participating in the survey. results plant diversity this study recorded 45 plant species that were used by local communities in the vicinity of buhanga sacred forest. of the 45 species, 38 plant species were categorised in 34 genera and 19 families to have medicinal value. in terms of species composition of medicinal plants, the family of asteraceae, acanthaceae and fabaceae had three species each followed by the family of chenopodiaceae, rutaceae, rhamnaceae, ranunculaceae, polygonaceae, solanaceae, menispermaceae, verbenaceae, urticaceae and cucurbitaceae with two species each. the remaining 14 families were represented by a single species each (see figure 3). regarding life form, 21 species (47%) were herbs, 8 species (18%) were trees, 10 species (22%) were climbers, 4 species (8%) were shrubs and 2 species (4%) were grasses (see figure 4). figure 3: distribution of 45 identified plant species among botanical families. figure 4: life form proportion of plant species. plant species role in ecological, social and cultural welfare results of medicinal plants collected from different habitats show that most of the plant species, about 29 species (62.22%) were harvested from natural vegetation followed by 9 species (20.00%) under cultivation in home gardens, 6 species (13.33%) collected in farming plots and 2 species (4.44%) found near the roadsides. the goods and uses of the vegetation in our study area showed that most of the plant species are used for medicinal purposes (38 plant species [74.51%]) followed by raw materials (8 species [15.69%]), social symbol (3 species [5.88%]) and 2 species (3.92%) for spiritual purpose (see figure 5 and table 1). figure 5: (a) origin of medicinal plants and (b) social welfare of plant species. table 1: plant species uses by local communities as raw material. interviewees from the three group discussions recounted stories of transgressors who had taken resources from a sacred forest and misfortunes fell on them (see table 2). for example it was said that villagers would not dare to enter the buhanga sacred forest for fear of angering some spirits in control of the forest. they believe that these spirits were able to curse them by sending a resident giant snake protective of the surrounding marshland called igishanga cya gihanga. moreover, buhanga sacred forest is a habitat for many rare species of snakes which upon sight a curse follows. calamities such as drought were also associated by such omens. table 2: name of sites and species of spiritual importance among local people. medicinal flora of the 38 plant species reported to have medicinal uses, 29 species distributed in 19 families were reported to be used against human health problems, six species belonging to six families were used against cattle diseases and three species belonging to three families were used against both human and cattle diseases. the most striking diseases recorded in 17 human health problems were the poison vomiting treated by six plant species followed by mental diseases and kilondatumbo (symptoms of kilondatumbo: painful sensation in the digestive system with presence of ulcers in the ileum and rectum) treated with five specific plant species (table 3). stomach ache was treated by three plant species while other plant species were used to treat one or two diseases (see table 3). in regard to ethnoveterinary medicine, the most remarkable diseases were tick-borne diseases (east coast fever, babesiosis and anaplasmosis) treated by four species followed by sexual dysfunction treated by two species. all other diseases were treated by one species each (see table 4). table 3: medicinal plants used against human health problems. table 4: plants used in veterinary medicine. regarding of some medicinal plants popular than others, brillantaisia cicatricosa was the most popular plant cited by 5 traditional healers for it medicinal value (relative frequency of citation [rfc] = 62.5%) followed by three plant species senna septemtrionalis, thalictrum rhynchocarpum and solanum terminale mentioned by 4 traditional healers (rfc = 50%). the species zanthoxylum sp., rumex abyssinicus, prunus africana, ranunculus multifidus, tragia brevipes, urtica massaica and ajuga alba were mentioned by 3 informants (rfc = 37.5%) while chenopodium opulifolium, cyathula cylindrica, solanum aculeastrum and solanecio mannii were mentioned by only 2 traditional healers (rfc = 25%) (see tables 3 & 4). concerning plant parts used frequently for medicinal preparation, the leaves were the most used parts followed by both leaves and tree barks in human health problem representing 81% and 6% respectively. regarding the ethnoveterinary medicine the leaves were also the most plant part used followed by both roots and tree barks and flowers representing 78% and 11% each. the majority of remedies were prepared in the form of juice from freshly collected plant parts. the juice was prepared by pounding or crushing a plant part in a wooden or stone motor and pesters. water was the most liquid substance used to dilute the prepared juice. the remedies were taken orally, accounting 84% of medicinal plants use, followed by external application applied typically on skin representing 8% and 5% on ear application except remedies from the species eucalyptus maidenii giving anally as smoke to treat haemorrhoids and fresh leaves from t. brevipes rubbing on painful join to treat rheumatism accounting both 3% (see figure 6). figure 6: proportion of plant parts used for human and cattle health problems: (a) proportion of plant parts used for human health problems; (b) proportion of plant parts used for cattle health problems. discussion taking consideration the size of buhanga sacred forest, there is a high percentage of plants being naturally harvested in the forest by local communities. the fact that more than a half of remedies were prepared from herbs in natural habitat than cultivated or home gardens indicates that the natural area of buhanga sacred forest is very important for local communities to satisfy their home-based healthcare. our finding is similar to the finding of edwards (2004) reporting that 2/3 of medicinal plants used worldwide are harvested from the natural environment. during interview, traditional healers reported that more medicinal plants were harvested previously than now in buhanga sacred forest and some species formerly forest-habiting species are now rarely encountered in the area. this is the case of the species p. africana that became extinct in the natural forest while c. cylindrica still existed occasionally due to overharvesting in the past. studies involving rwanda on medicinal plants reveal decline of these resources (stewart 2003). our research findings revealed that the poison vomiting, mental diseases and kilondatumbo in human health problems were the most cited diseases in our study area because remedies against these three predominant health problems were mostly prepared. in a situation where several drugs in one area are required, this indicates a prepotency of a particular disease (dawit & ahadu 1993). the potential of medicinal plants recorded in this study were also confirmed by other research conducted elsewhere in rwanda (dessouter 1991; kamagaju et al. 2013; kayonga & habiyaremye 1987; mbarubukeye & niang 1996; rwangabo 1993). as per the ethnoveterinary medicine represented mostly by tick-borne diseases, suggests that these diseases were very important in the area. according to byavu et al. (2000), mbarubukeye (1991) and van puyvelde et al. (1985), the east coast fever (theileriosis) played a vital role in cattle mortality in rwanda, particularly in exotic pure breed crossing with the ankole breed while anaplasmosis and babesiosis represented an enormous cost in term of time and money. also study conducted by nshimiyimana and mutandwa (2010) indicates that damages attributable to ticks borne diseases in rwanda were considered as the first economic damage of bovine production transmitting sucking blood parasites diseases such as theileriosis, anaplasmosis and babesiosis. with more than 70% of leaves use as ingredient in making of medicinal concoction indicates that leaves are more required to solve the health problem in our study area. in reference to dawit and ahadu (1993) and poffenberger et al. (1992), the normal harvesting leaves estimated of up to 50% does not significantly affect the growth of plant species or does not cause a great danger to existence of individual plant when compared to the collection of underground part, stem or whole plant from both ecological points of view and survival of species. the extinction of p. africana as reported by the healers is due to overutilisation of both its leaves and barks as ingredients and was replaced by agricultural activities. the same were reported by sayer, harcourt and collins (1992) stewart (2003) and minitere (2003). in general there was no really threat of overharvesting of plant in buhanga sacred forest as it used to be in the past because the remedies are prepared and used locally, and therefore harvested in small quantities. traditional healers reported that they do not store remedies for a prolonged period of time because when the need comes, they go out to collect plants, prepare the remedies and use it for treatment. however the importance of buhanga were known a long time ago, this sacred forest has acquired since 2005 a status of protected area as a part of the volcanoes national park from the former rwanda office of tourism and national parks (ortpn), now rwanda development board (rdb) (rema 2009). in the same context, culture and spiritual beliefs plays an important aspect in the conservation of buhanga sacred forest. recently, water springs from buhanga are still used as source of clean water for local people in the area and caves and some tree species (dracaena steudneri, erythrina abyssinica and ficus thonningii) are important for their ritual and ceremonies. conclusion this study has revealed the importance of traditional ecological knowledge in buhanga sacred forest plays a vital role in our heritage. the high plant diversity used in traditional medicine to treat different ailments was localised mostly in the natural habitat and does not really present threat of overharvesting because remedies are prepared on a need-to-need basis. also this study has shown how cultural values and beliefs are crucial tools for conservation and should therefore be incorporated in all management plans of buhanga sacred forest. in light of this research finding, we recommended that traditional healers should be trained on how to domesticate natural plant species as some of them has becoming rare or extinct in the surrounding area due to past anthropogenic activities. among others 15 species were documented and reported in our study area as priority species for preservation of stored grains for their medicinal importance uses. these include: b. cicatricose, s. septemtrionalis, t. rhynchocarpum, s. terminale, zanthoxylum sp., r. abyssinicus, p. africana, r. multifidus, t. brevipes, u. massaica, a. alba, c. opulifolium, c. cylindrica, s. aculeastrum and s. mannii. acknowledgements this study was funded by the regional non-governmental organisation (ngo) of albertine rift (arcos) in partnership with association pour la conservation de la nature au rwanda (acnr). we thank the national herbarium of rwanda for their support in species identification and other relevant botanical information, the local government authorities’ of musanze district and traditional healers, members of abadahemuka cooperative. special thanks go to ir. gloriose barebwayire from projet d’appui du développement de l’élevage bovin laitier (padebl/minagri) for excellent technical support in cattle diseases identification, mrs usambyimbabazi madeleine, a student at the national university of rwanda (nur)-botany department for her kind field assistance and mr felix m. waweru (africarice center dar es salaam) for editing and proof-reading. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions n.j.s. was the project leader, m.v., b.e. and b.n. made conceptual 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group discussion with local indigenous, (c) and some important sites and species of buhanga sacred forest including the mysterious water-spring, (d) cave (e) and dracaena steudneri (igihondohondo). figure 2-a1: remarkable places in rwanda. article information authors: m. kyle s. smith1 nerina kruger1 taryn s. murray2,3 affiliations: 1rondevlei scientific services, south african national parks, south africa 2south african institute for aquatic biodiversity, grahamstown, south africa 3department of ichthyology and fisheries science, rhodes university, south africa correspondence to: kyle smith email: kyle.smith@sanparks.org postal address: po box 176, sedgefield 6573, south africa dates: received: 02 july 2014 accepted: 08 jan. 2015 published: 01 july 2015 how to cite this article: smith, m.k.s., kruger, n. & murray, t.s., 2015, ‘aerial surveys conducted along the garden route coastline, south africa, to determine patterns in shore fishing effort’, koedoe 57(1), art. #1236, 9 pages. http://dx.doi.org/10.4102/koedoe.v57i1.1236 copyright notice: © 2015. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. aerial surveys conducted along the garden route coastline, south africa, to determine patterns in shore fishing effort in this original research... open access • abstract • introduction • research method and design    • study area    • methods    • analyses       • aerial survey data maps       • effort • results    • spatial patterns in fishing effort    • temporal patterns in fishing effort    • total angling effort • discussion    • spatio-temporal patterns in fishing activity    • non-compliance with spatial closures       • limitations of the study • conclusion • acknowledgements    • competing interests    • authors’ contributions • references abstract top ↑ coastal environments provide a wide range of leisure opportunities, including recreational fishing. understanding spatial and temporal fishing patterns is important in ensuring wise management and sustainable use. to provide information on shore angler effort and distribution, randomised aerial surveys of the garden route coast between the eastern border of the tsitsikamma marine protected area and the kaaimans river mouth in the west were undertaken between december 2008 and november 2009. a total of 15 flights were conducted, with six flights taking place over weekends, two on public holidays and the balance on normal week days. angler effort was not uniformly distributed along the coastline, and spatial analysis highlighted coastal areas both inside and outside marine protected areas that had increased angler effort. in general, fishing effort was highest around more densely populated areas and concentrated in areas with easy access. although angler counts were highly variable, the seasonality of shore angling effort showed a slight increase during autumn and winter and angling effort was significantly higher on weekends. conservation implications: data obtained during these surveys can assist management with future conservation planning exercises, whilst also guiding daily law enforcement patrols to maximise angler encounters. introduction top ↑ coastal environments provide a wide range of recreational opportunities and are highly valued amongst various user groups (james 2000). understanding recreational use patterns is often necessary for adequate coastal management (smallwood et al. 2011). in particular, the spatial and temporal distribution of activities can be used in future coastal zoning, whilst also elucidating compliance with current zonation plans. similarly, a knowledge of resource use patterns can be used by management authorities in planning and optimising when and where law enforcement patrols should take place in order to maximise outputs and minimise costs. impacts arising from recreational activities depend on several factors, such as the type of activity, the number of participants, and the resilience of different species and habitats to the pressure (davenport & davenport 2006; meyer & holland 2008). recreational fishing is a popular activity and can arguably have a large impact through over-exploitation and harvesting of select species. although it is unlikely that a single angler's catch can have a measurable impact on fish populations, the cumulative impacts of (1) the widespread nature of recreational angling, (2) the total number of anglers involved and (3) the defined habitats in which fishing occurs are important factors to consider (cooke & cowx 2004; mcphee, leadbitter & skilleter 2002). within the south african shore-based linefishery, overfishing by recreational anglers is seen as a contributing factor to changes that have occurred in the composition of species catch (bennett 1991; brouwer & buxton 2002; brouwer et al. 1997; dunlop & mann 2012) and decreases in the abundance of target species (attwood & farquhar 1999; bennett 1991; cowley, brouwer & tilney 2002). this has resulted in increased emphasis on marine protected areas (mpas) as the basis for marine conservation, an essential component for fisheries management in south africa (attwood et al. 1997). the benefits of mpas typically include the restoration of marine ecosystems and natural ecosystem functioning, increased species diversity, and natural size and age structure of protected fish populations (halpern 2003). further fishery benefits include a spillover of subadult and adult fish, and the production of eggs and larvae that can be distributed over a large area (brouwer et al. 2003; kerwath et al. 2013; tilney et al. 1996). however, the effectiveness of mpas in meeting their conservation and fishery objectives is dependent on user compliance, with illegal fishing reducing their efficacy and negatively impacting fishery benefits (arias & sutton 2013). unfortunately, quantification of non-compliance by recreational anglers is rare, in particular with spatial zonation and over large areas. during this study, aerial surveys, as an alternative method to traditional on-the-ground or water-based surveys (brouwer 1997; mann et al. 2008; pollock et al. 1994), were used to (1) test the hypothesis that shore angling effort along the garden route coastline shows spatial and temporal variation, (2) assess compliance with spatial zoning and (3) provide recommendations to improve the efficiency of law enforcement activities within this area. research method and design top ↑ study area this study was conducted along the garden route coastline of south africa between the groot river in the east and the western border of the kaaimans river mouth (figure 1). the study area consisted of predominantly rocky coastline (78%) and sandy beaches (22%) and was 248 km long. the area falls under the jurisdiction of four local municipalities, with several towns and villages scattered throughout the adjacent interior. coastal access points are plentiful. the majority of people living in the area are situated in or near the towns of george, knysna and plettenberg bay (table 1). figure 1: a map of south africa showing the location of the study area along the south coast, and the location of marine protected areas, municipal wards and towns within the study area. table 1: population numbers per ward for each of the local municipalities within the study area. three mpas are situated within the study area: the tsitsikamma mpa consists of 68 km of closed (no resource use) coastline and forms part of the garden route national park, managed by south african national parks, whilst both the robberg and goukamma mpas, managed by capenature, are open to shore angling, but no boat angling is permitted. methods monthly aerial surveys covering the survey area were conducted between december 2008 and november 2009 in order to obtain ‘instantaneous’ counts of shore angling effort. survey days and flight times were randomly selected, but depended on the weather, and pilot and observer availability. all flights were conducted during daylight hours, with the earliest flight taking place at 09:10 and the latest at 15:55. all counts were done in an east–west direction, as the observers were seated on the starboard side of the plane. the first four surveys were undertaken in a four-seater robinson helicopter, after which a four-seater cessna light aircraft was used. flight parameters, including air speed and altitude, were standardised, but depended on weather conditions and the pilot's discretion. flight times ranged between 70 min and 80 min, with an average air speed of 189 km.hr−1 for both aircraft. altitude ranged between 30 m a.s.l. and 80 m a.s.l. for the robinson and between 150 m a.s.l. and 250 m a.s.l. for the cessna. on each flight, two observers counted shore anglers and one scribe entered the data into a global positioning system-enabled pocket pc. each spotter was equipped with a pair of binoculars (10 x 42 magnification) and continuously scanned the coastline, verifying the other spotter's counts. when large groups of anglers were encountered, a second flyby was made and anglers were re-counted by both spotters. information recorded during the aerial surveys included the date, time, weather, sea conditions, number and location of anglers or boats and habitat where shore anglers were fishing (rock or sand). data were also obtained from three aerial surveys conducted in december 2008 as part of a study researching the ecology, value and management of the garden route (chalmers et al. 2009). methods were similar, with the same cessna plane and pilot being used, two observers and similar standardised flight speed and altitude. this information was used in temporal, but not spatial (coastal effort maps), analyses. analyses aerial survey data maps the southern cape coastline that falls within the study area was buffered by 1 km to create a polygon representing the area of interest. spatial mapping of angler effort was plotted using three different coastal breaks. the first method (equal breaks) split the coastline into 10 km sections. the second method compared the status (mpa or non-mpa) using the layer of the mpas of the national spatial biodiversity assessment (lombard et al. 2004). the final method used the coastal breaks described in the coastal sensitivity atlas of southern africa (jackson & lipschitz 1984). for this analysis, exposed rocky headlands and wave-cut rocky platforms were combined into one category (rocky). aerial survey point data containing angler counts were overlaid onto each coastal break layer and the density of anglers per coastal section for each of the different breaks was calculated as anglers.km−1. angler effort was depicted in graduated colours for seven natural break classes. kruskal–wallis one-way analyses of variance (anovas) were performed to determine if there was a difference in angling effort spatially for each of the three different coastal breaks, and temporally between week and weekend days and seasons. effort the unit of fishing effort chosen was angler outings, with data pooled seasonally due to unequal sampling effort. total seasonal fishing effort (eseasonal outings) was calculated using the following formulae: where ew1 and ew2 are the weekday and weekend estimates respectively, given by: where j is week days or weekends, ei is the number of anglers on the ith day, d is the number of days sampled and p is the potential number of sampling days. total annual fishing effort (etoutings) was calculated as the sum of the estimated seasonal effort (eseasonal outings) multiplied by a correction factor (2.48) to account for angler turnover. total annual fishing effort (etoutings) was then multiplied by the average fishing trip duration of anglers interviewed during two ground-based roving creel studies (smith 2012; van zyl 2011). a kruskal–wallis one-way anova was performed to determine if there was a difference in angling effort between seasons, and a mann–whitney u test was used to establish if there was a difference in angling effort between week and weekend days. results top ↑ a total of 15 aerial surveys were conducted between december 2008 and november 2009 (table 2), with 6 flights occurring during summer months, 4 in autumn, 3 during winter and 2 in spring. most flights were conducted in the morning, with 3 weekend flights having being completed during the afternoon. table 2: summary of aerial surveys conducted each month, grouped seasonally and stratified according to week or weekend day. spatial patterns in fishing effort angling effort showed a heterogeneous distribution throughout the study area and although variability in count data was high, a significant difference in angling effort occurred between mpas and non-mpas (kruskal–wallis one-way anova, df = 6, p-value < 0.05) (figure 2) and equal area breaks (kruskal–wallis one-way anova, df = 24, p-value < 0.05) (figure 3). figure 2: the south african garden route coastal section broken into equal areas (a) and the spatial distribution of total angling effort (anglers.km−1) between kaaimans river in the west and the eastern border of the tsitsikamma marine protected area at the groot river (b). up–down bars indicate the 95% confidence interval. figure 3: the south african garden route coastal section divided into marine protected areas and open areas (a) and the spatial distribution of total angling effort (anglers.km−1) within marine protected areas and open areas between the kaaimans river in the west and the eastern border of the tsitsikamma marine protected area at the groot river (b). up–down bars indicate the 95% confidence interval. the majority of anglers (72.50%) fished in open access areas outside formal conservation borders (including no-take and controlled zones) and a bonferroni post-hoc test showed that the tsitsikamma mpa had significantly less fishing effort than all other sections (p-value < 0.05) with the exception of zone 9, in which no anglers were encountered. although the tsitsikamma mpa is zoned for no-take (i.e. no consumptive use), a total of 32 illegal fishermen were counted during all the surveys, with an overall angler density of 0.03 anglers.km−1 (figure 3). outside of formal protected areas, the highest density of anglers occurred in section one (wilderness) with 1.35 anglers.km−1, followed by section 15 (plettenberg bay) and 17 (nature's valley) with overall angler densities of 0.78 anglers.km−1 and 0.58 anglers.km−1 respectively (figure 2). these three sections contributed to the overall angler densities being highest (0.73 anglers.km−1) along the larger section of coastline between the kaaimans river mouth and the western border of the goukamma mpa. this was followed by the plettenberg bay coastline (robberg to de vasselot) with 0.65 anglers.km−1 and the de vasselot section of the garden route national park with the third highest density of 0.51 anglers.km−1 (figure 3). the goukamma and robberg mpas had overall angler densities of 0.41 anglers.km−1 and 0.14 anglers.km−1 respectively (figure 3), whilst the coastal section between the goukamma and robberg mpas, consisting of large stretches of exposed rocky coastline with few access points, had a relatively low overall angler density of 0.16 anglers.km−1. angling effort occurred on both rocky and sandy substrates throughout the study area with both high densities (2.23 anglers.km−1 – 3.94 anglers.km−1) occurring over rocky (figure 4a) and sandy substrates (figure 4b). although total angler density on rocky substrates was lower (0.31 anglers.km−1) than on sandy substrates (0.47 anglers.km−1), fishing effort between substrates was not significantly different (mann–whitney u test, p-value 0.221). figure 4: the distribution of total angling effort (anglers.km−1) between the kaaimans river in the west and the western border of the no-take tsitsikamma marine protected area between rocky areas (a), sandy areas (b) and both rocky and sandy areas combined (c). temporal patterns in fishing effort although more anglers were encountered during the autumn (0.37 anglers.km−1) and winter (0.38 anglers.km−1) months (figure 5a), with fewer anglers fishing in spring (0.20 anglers.km−1) and summer (0.21 anglers.km−1), no significant difference was observed (kruskal–wallis one-way anova, df = 3, p-value 0.266). this was due to the high variability within the data, exacerbated by the low sample size. fishing pressure was significantly higher during weekends and public holidays (0.38 anglers.km−1) than on week days (0.18 anglers.km−1) (mann–whitney u test, df = 1, p-value 0.028) (figure 5b). figure 5: mean shore angling effort (anglers.km−1) (a) seasonally and (b) during week and weekend days along the south african garden route coast between december 2008 and november 2009 (public holiday counts were included as weekend days). total angling effort total annual shore angling effort was estimated at 49 812 angler outings. recent angler interviews conducted along portions of this coastline indicate that anglers spend on average between 5 hours (smith 2012) and 6 hours (van zyl 2011) fishing per outing. using these figures, a total annual fishing effort estimate was estimated to be between 226 645 angler hours and 298 872 angler hours. discussion top ↑ assessing spatio-temporal resource use patterns is important when implementing fishery and conservation management practices. this information can help with future conservation and spatial planning exercises by highlighting areas from which anglers may be displaced or, conversely, areas where no fishing activity occurs and which may act as natural spatial harvest refugia for target species (smallwood & beckley 2012). furthermore, identifying popular fishing locations can allow management authorities to maximise resource allocation and streamline daily law enforcement operations (smallwood & beckley 2012). spatio-temporal patterns in fishing activity angler distribution is unlikely to be uniformly distributed along any coastline and may be impacted by access points and infrastructure that have a clustering effect (smallwood et al. 2012). similarly, habitat preferences of target fish species can influence the spatial distribution of anglers attracted to particular sites and different species (smallwood et al. 2013). recreational fishing along the garden route coastline showed a heterogeneous spatial distribution, with some localised areas having high angler densities. these areas were generally associated with ease of access and proximity to the more heavily populated urban areas of george and plettenberg bay. a similar pattern was shown along the kwazulu-natal coast of south africa (mann et al. 2008), where angler effort was largely concentrated around access points, with increased effort along the more developed and highly populated sections of the coastline. the concentration of anglers around easy access points along south africa's coastline has more than likely been heightened by the 2002 ban (schedule 44 of the national environmental management act [act no. 107 of 1998]) on beach driving, which limits anglers’ access to the more remote sections of coastline (mann & tyldesley 2012; mann et al. 2008). in fisheries where angling effort is widely dispersed (e.g. coastlines), instantaneous estimates of total fishing effort obtained from aerial surveys are more accurate and preferable to land-based roving creel surveys (pollock et al. 1994). although no previous aerial surveys have been undertaken along this section of coastline for comparative purposes, a number of land-based roving creel surveys covering sections of the coastline have been completed, including plettenberg bay (king 2006), the goukamma mpa (van zyl 2011) and the wilderness to sedgefield coastline (smith 2012). these projects, conducted at finer spatial scales, show similar patterns and general trends to those recorded during the aerial surveys conducted in this study. greater angling effort (1.03 anglers.km−1) was recorded along the wilderness to sedgefield (smith 2012) and plettenberg bay coastlines (0.71 anglers.km−1 [king 2006]) during land-based surveys, in comparison to 0.84 anglers.km−1 and 0.69 anglers.km−1 recorded during the aerial surveys of this study. due to the generally shorter distance and slower speed of land-based creel surveys, a higher estimation of effort in comparison to aerial surveys would be expected (pollock et al. 1994). the goukamma study did not report on angler densities in terms of anglers.km−1 of coastline, but estimated total effort relating to angler hours (21 428 [van zyl 2011]), approximately half the 48 167 angler hours estimated in the wilderness study (smith 2012). this is similar to the trend seen in the effort estimations of the aerial surveys, with an overall angler density of 0.46 anglers.km−1 recorded for the goukamma mpa. the distribution and density of recreational users in general is known to be impacted by large temporal factors such as seasons and holiday periods (hingham & hinch 2002). large-scale patterns in recreational shore fishing along the south african east coast have also been shown to fluctuate seasonally, with an increase in summer and a decrease in winter (brouwer et al. 1997). both king (2006) and smith (2012) have shown that recreational shore fishing within plettenberg bay and the wilderness coastline followed this trend, attributing the increase in effort to an influx of visitors during school holiday periods. however, results from the aerial surveys are more consistent with those from goukamma (van zyl 2011) and the kwazulu-natal shore fishery (dunlop & mann 2012; mann et al. 2008), which both showed an increase in shore angling density over the winter months. two popular targeted shore angling species, galjoen (dichistius capensis) and elf (pomatomus saltatrix), show strong seasonal abundance and have closed seasons between 15 october and the last day in february, and 01 october and 30 november respectively. it is thus likely that those anglers who specifically target these species are temporarily absent from the fishery during these closed periods (mann et al. 2008; van zyl 2011). given that no previous aerial surveys have been conducted along the garden route coastline and shore-based surveys have only covered portions of the coastline, it is impossible to say whether total fishing effort is increasing or not. a decrease in shore-based angling effort along the kwazulu-natal coastline (dunlop & mann 2012) has been attributed to security concerns, declining catches, an increase in the cost of fishing and stricter linefish regulations. as catch rates within the shore fishery have decreased (brouwer & buxton 2002), the duration and frequency of fishing may have altered, which in turn would affect the turnover rate used to calculate the total estimated effort. however, the current average fishing duration of between 5 hours (smith 2012) and 6 hours (van zyl 2011) is very similar to the 5 hours calculated 15 years earlier (brouwer 1997). independent effort calculations are seen as a crucial part of the management and monitoring of the south african shore fishery (dunlop & mann 2012) and the total angling effort estimated from this study provides a baseline for future work. non-compliance with spatial closures with greater biodiversity and abundance of fish, mpas where fishing is allowed are attractive locations for recreational anglers (cooke et al. 2006; götz, cowley & winker 2008; parker, booth & mann 2013). however, non-compliance by anglers fishing in closed, no-take areas can undermine conservation and fishery management objectives. along the garden route, shore fishing is allowed in all mpas except tsitsikamma. however, illegal shore fishing (poaching) was observed on several occasions during this study within the no-take tsitsikamma mpa. although fishing levels inside the park were low compared to outside adjacent areas, illegal fishing within this important mpa is cause for concern. within the ningaloo marine park in north west australia, anglers were found to be concentrated in localised areas with easy access, and limited non-compliance was observed within no-take zones (smallwood et al. 2011). the low levels of non-compliance were attributed to a number of factors, including (1) multiple educational tools (signboards and brochures), (2) visual zonation reference points, (3) various enforcement patrols (boatand land-based) and (4) a willingness by anglers to fish at alternative sites (smallwood & beckley 2012). more research is required regarding the illegal fishing taking place within the tsitsikamma mpa to gain a greater understanding of the extent of this activity and the underlying drivers, which could then be used by management to address and resolve this issue. although the goukamma and robberg mpas are open to shore-based fishing, these areas had comparatively lower angler densities than other sections of coastline outside the formal conservation areas. fishing locations within robberg are limited due to a large seal colony on the eastern side and the exposed rocky coastline on the west, restricting fishing to a few well-known locations (king 2005). the goukamma mpa has a combination of habitat types and land-based roving creel surveys conducted within this reserve (van zyl 2011) showed that anglers were clustered around easily accessible areas, favouring rocky and mixed shores over sandy beaches. similar to the aerial survey data presented here, angler densities were greater around the buffalo bay section of the reserve, which may be due to ease of access, with anglers in the goukamma section having to walk up to 6 km to reach some fishing locations. limitations of the study surveys to provide information on spatial and temporal patterns of resource use need to be designed and implemented at appropriate scales. numerous access points, the longitudinal and rugged nature of coastlines and the size of management areas complicate survey design. aerial surveys can be an effective technique for obtaining spatial and temporal data on recreational activities, including fishing, over a broad landscape (pollock et al. 1994). such surveys are cost-effective and efficient, with minimal personnel involved and can cover large geographic areas in a relatively short time (malvestuto 1983). to avoid sampling biases, flights must be random in terms of direction, time of day and type of day (week or weekend) (pollock et al. 1994). in practice this becomes difficult, with weather conditions in particular playing an important role in determining flight days. the greatest limitation of this study was the inability to keep sampling effort equal between months, with flights conducted on both week and weekend days. subsequently, monthly estimates of angler effort and comparisons were not possible. analysing the data by season to estimate total angler effort attempts to alleviate this issue to some extent, but an unknown degree of error is being introduced and needs to be acknowledged. spatial accuracy of observed data points was improved through the use of data loggers that automatically recorded time and positional information, enabling the observers to simply record numbers and substrate. prominent landmarks were also geo-referenced prior to the aerial flights, providing a known position that could be recorded, eliminating sampling errors associated with fishing at these points. however, visibility bias and in particular difficulty in seeing anglers on rocky sections of coastline may have led to an underestimation of total angling effort. ground truthing of counts would be important for future surveys. conclusion top ↑ aerial surveys are a useful and well-established method of collecting data across large spatial scales. these data can be used in both future planning exercises and also in determining the success of current management plans and policies (e.g. non-compliance with spatial zoning). although our ability to estimate total angler effort from the current data set is limited due to unequal monthly sampling effort, results from this study showed temporal trends with seasonal fluctuations for recreational fisheries. an increase in effort over weekends and public holidays was evident, whilst broad-scale spatial patterns were similar to results from more in-depth, land-based projects. in general, fishing effort was highest around the more heavily populated areas of george and plettenberg bay and concentrated in areas with easy access. should aerial surveys be implemented again, care should be taken to ensure equal monthly effort (i.e. flights should be conducted at least bi-monthly – on one weekday and one weekend day or public holiday), with flight scheduling incorporating back-up days for bad weather and mechanical problems. the value of surveys could be further increased by collecting data on other types of coastal resource use, such as bait harvesting. the findings of this study have relevance to the management of the fishery along the garden route coastline in guiding the deployment of fisheries management field personnel, both spatially and temporally, to maximise contact with fishers and therefore improve levels of compliance. acknowledgements top ↑ funding for the flights was made available through the national marine protected area budget, allocated to the south african national parks by the department of environmental affairs. we thank the volunteers and pilots who assisted during the aerial surveys and two anonymous reviewers for their positive contributions towards this manuscript. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions m.k.s.s. (south african national parks) was responsible for the study conceptualisation, data collection, analyses, report writing and editing. n.k. (south african national parks) contributed to the study design, data collection, gis analyses, report writing and editing, and t.s.m. 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wilderness section coastal marine resource use monitoring programme’, scientific report number 05/2012, south african national parks, skukuza. superweb by space time research, n.d., viewed april 2014, from http://interactive.statssa.gov.za/superweb/login.do tilney, r.l., nelson, g., radloff, s.e. & buxton, c.d., 1996, ‘ichthyoplankton distribution and dispersal in the tsitsikamma national park marine reserve, south africa’, south african journal of marine science 17, 1–14. http://dx.doi.org/10.2989/025776196784158482 van zyl, c., 2011, ‘the use of a roving creel survey to monitor exploited coastal fish species in the goukamma marine protected area, south africa’, mtech thesis, nelson mandela metropolitan university, port elizabeth. article information authors: mamokete n.v. dingaan1,2 pieter j. du preez1 affiliations: 1department of plant sciences, university of the free state, south africa2department of soil, crop and climate sciences, university of the free state, south africa correspondence to: mamokete dingaan postal address: po box 339, bloemfontein 9300, south africa dates: received: 15 feb. 2012 accepted: 20 sept. 2012 published: 14 mar. 2013 how to cite this article: dingaan, m.n.v. & du preez, p.j., 2013, ‘grassland communities of urban open spaces in bloemfontein, free state, south africa’, koedoe 55(1), art. #1075, 8 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1075 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/ koedoe.v55i1.1075-1 copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. grassland communities of urban open spaces in bloemfontein, free state, south africa in this original research... open access • abstract • introduction • research method and design    • study area    • vegetation sampling and data analysis • results    • vegetation classification    • vegetation description       • 1. felicia muricata – themeda triandra major community       • 1.1. panicum coloratum – digitaria eriantha community       • 1.1.1. cyperus usitatus – digitaria eriantha sub-community       • 1.2. eragrostis obtusa – sporobolus fimbriatus community       • 1.2.1. panicum schinzii – themeda triandra sub-community       • 1.3. eragrostis biflora – themeda triandra community       • 2. aristida congesta – themeda triandra major community       • 2.1. trichoneura grandiglumis – rhynchosia nervosa community       • 2.1.1. conyza bonariensis – eragrostis curvula sub-community       • 2.1.2. antephora pubescens – digitaria argyrograpta sub-community       • 2.2. eragrostis trichophora – cyperus capensis community       • 2.3. hyparrhenia hirta – themeda triandra community       • 2.4. melinis repens – eragrostis lehmanniana community    • ordination • trustworthiness    • reliability    • validity • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ natural vegetation in urban environments is greatly impacted by human activities and it is in constant threat of degradation and destruction as a result of urbanisation. this vegetation, although fragmented, serves an important ecological function and needs to be properly managed and conserved. studies on urban vegetation are lacking in south africa, with only a handful having been carried out since the end of the last century. this study was initiated to identify, classify and describe the grassland communities of the urban open spaces in bloemfontein. relevés were compiled in 61 sample plots, where species present and habitat information were recorded. care was taken to restrict sample plots to vegetation in pristine condition, wherever possible, and severely degraded stands were avoided. a two-way indicator species analysis (twinspan) classification, refined by braun-blanquet procedures, revealed two distinct major communities, seven communities and four sub-communities. both detrended and canonical correspondence analyses indicated the vegetation units to be associated with soil texture and ph, although biotic factors such as overgrazing, burning and mowing also influence the composition of the vegetation. the proper management and conservation of urban open spaces requires in-depth knowledge of the spatial distribution, floristic, structural and functional compositions within the major vegetation types in this environment. the present study further contributed towards formulating ways for the proper management, utilisation and functioning of the open spaces within the bloemfontein area.conservation implications: the grassland biome of south africa is poorly conserved, mainly because of its status as an agricultural hub of the country. the preservation of natural and semi-natural forms of urban vegetation is important because such vegetation, although often disturbed and degraded, could form dispersal corridors between peri-urban and rural vegetation. introduction top ↑ bloemfontein lies within the grassland biome (rutherford & westfall 1994) and is classified by acocks (1988) as part of the dry cymbopogon – themeda veld (vt 50). here themeda triandra is ecologically the most important grass, both to wild and domestic animals, and is the most widely distributed of the climax grasses. one of the key environmental variables affecting the functioning of the grassland ecosystem globally is rainfall variability (tilman & el haddi 1992). bloemfontein is situated in a semi-arid region, where precipitation is low and highly variable (noy-meir 1973; sala & lauenroth 1982). although semi-arid regions typically receive substantial precipitation for at least a few months of the year, lack of adequate rainfall for many months may also prevail (bailey 1979).the grassland biome of south africa is an important agricultural region, especially for the intensive production of crops such as maize and wheat (department of agriculture, forestry and fisheries 2010). it is also important for extensive stock farming; mainly for dairy, beef and wool production. however, the intensive crop production and livestock grazing pressure have resulted in the destruction or degradation of large portions of pristine vegetation in the biome. in addition, natural and semi-natural vegetation is constantly impacted by urban development and, according to low and rebelo (1996), many of the natural areas in the biome have been lost as a result of urbanisation. in recognition of the poor conservation status of the biome, cohen and hugo (1986) emphasise the importance of conservation outside officially designated nature reserves, such as in urban open spaces. in the vicinity of bloemfontein, as in many areas of the biome, natural vegetation is represented by small fragmented areas. this fragmentation of previously intact natural environments may cause the potential loss of habitat for some species, the isolation of other species on natural remnants within the urban environment and even the total extinction of certain plant species (wood et al. 1994). studies of urban vegetation in south africa are lacking and only a limited number of vegetation surveys have been carried out in these areas: for example, in durban, kwazulu-natal province (roberts 1993); potchefstroom, north west province (cilliers, van wyk & bredenkamp 1999); klerksdorp, north west province (van wyk, cilliers & bredenkamp 2000); and pretoria, midrand, johannesburg and parts of the west rand, gauteng province (grobler, bredenkamp & brown 2006). the aim of this study was therefore to identify, classify and describe the grassland communities of the natural open areas in bloemfontein. research method and design top ↑ study area the study was carried out in natural open spaces within the bloemfontein metropolitan area, which include the inner city area, as well as the surrounding farms and smallholdings. the study area extends from approximately 29°00′s to 29°15′s and 26°07′e to 26°21′e, at an altitude of 1350 m a.s.l. – 1450 m a.s.l.there are various large open spaces in the area, but, for this study, special emphasis was placed on the natural and semi-natural areas. these areas are fragmented and are most prominent towards the periphery of the city. they comprise numerous hills and ridges, areas along drainage lines and uncultivated patches on smallholdings and farms. other open spaces studied were vacant residential lots and the yet-to-be-developed areas as indicated in the spatial development framework of the mangaung metropolitan municipality (2011). green belts adjacent to railway lines and main roads were also included in the study because they form part of the natural open space system within the bloemfontein area, but the open spaces in the inner city area, such as parks and sports grounds, were excluded from the study as they were fully transformed. according to the köppen climate classification system, bloemfontein falls under the bsk climate zone (schulze 1947). this is a steppe climate in which the winters are dry and the mean annual temperature is below 18.0 °c (le houérou, poppov & see 1993; schulze & mcgee 1978). maximum temperatures are recorded in december and january, averaging 30.1 °c and 30.6 °c, respectively (figure 1). below zero temperatures are often recorded for june (-1.0 °c) and july (-1.2 °c). the annual mean maximum temperature is 24.6 °c and the annual mean minimum temperature is 7.6 °c, with the average annual temperature of 16.1 °c. rain mainly falls during the summer months, often in the form of thunderstorms. the average annual rainfall is 550 mm. figure 1: long-term (1951.2000) meteorological data of the bloemfontein airport (29°06′s, 26°18′e, 1351 m a.s.l.), sourced from the south african weather service. three different land types are distinguished in the study area, namely the ca, dc and ea land types. within each of these land types, soils vary from sandy to clayey, as a result of the variation in parent material. the ca land type, occurring as ca8 and ca22 subdivisions, is a mixture of duplex and plinthic soils. the ca8 subdivision is found on the western part of the study area and consists mainly of sandy hutton–bainsvlei soils and more clayey valsrivier–swartland soils; the ca22 subdivision occurs in the south and is mainly dominated by valsrivier soils. the dc land type occurs as subdivision dc13, which is dominated by duplex soils of the valsrivier–swartland–sterkspruit forms. it is only present in small pockets in the eastern part of the study area. the ea land type is found in the central and northern parts and occurs as subdivision ea39, the most extensive in the study area. it consists mainly of sandy oakleaf soils, although a mixture of the clayey milkwood, arcadia and valsrivier soils are also prominent. the dominant soil profiles are described according to land type survey staff (1993) and the soil forms and series are used according to the soil classification working group (1991).the geologic formations of the study area belong to the karoo supergroup and consist of the tierberg formation of the ecca group and the adelaide sub-group of the beaufort group, as well as dolerite intrusions of the post-karoo age (johnson et al. 2006). vegetation sampling and data analysis relevés were compiled in 61 sample plots, each fixed at 16 m2. care was taken to restrict sample plots to vegetation in pristine condition, wherever possible, and severely degraded stands were avoided. in each sample plot, all vascular plants present were recorded and all plant nomenclature was used according to germishuizen and meyer (2003). the cover-abundance values of all species present were allocated according to the braun-blanquet scale, as given by mueller-dombois and ellenberg (1974), as well as kent and coker (1996). habitat parameters recorded included soil depth, rockiness of the soil surface and the presence of biotic influences such as trampling and grazing. a total of 25 soil samples was collected and analysed for ph, organic matter and texture. no soil samples were collected at one part of the study area, the kwaggafontein hills, as this is a conservation area and no permission was granted to collect such samples.the vegetation and habitat data were first captured into the turboveg database (hennekens 1996a) and then imported to megatab (hennekens 1996b), a visual editor for phytosociological tables, where the first approximation of the vegetation classification was obtained by the application of the two-way indicator species analysis (twinspan) (hill 1979a). further refinement of the initial classification results was subsequently conducted within the megatab programme, resulting in a phytosociological table. the relationships between the plant communities and environmental variables were determined by an ordination algorithm, detrended correspondence analysis (dca) (hill 1979b) and also by an extension of dca, the canonical correspondence analysis (cca) (ter braak & šmilauer 2009). results top ↑ vegetation classification the grassland communities of the bloemfontein area are classified as bloemfontein dry grassland by mucina et al. (2006) the vegetation is dominated by t. triandra, a companion species with a wide ecological amplitude and eragrostis lehmanniana as a typical associate. other constantly present grasses are eragrostis curvula and digitaria eriantha. the forbs oxalis depressa, nidorella resedifolia and hibiscus pusillus are of widespread occurrence, although they are hardly prominent (see online appendix). the habitats occupied by each of the communities are, by and large, distinguished by differences in soil depth, texture and ph (table 1). biotic factors, especially livestock grazing practices, also influence the composition of the communities. table 1: soil characteristics of the grassland communities of the bloemfontein area (mean values). the classification resulted in two major communities, which are grouped into seven communities and four sub-communities. the hierarchical classification of the plant communities is as follows:1. felicia muricata – themeda triandra major community 1.1. panicum coloratum – digitaria eriantha community 1.1.1. cyperus usitatus – digitaria eriantha sub-community 1.2. eragrostis obtusa – sporobolus fimbriatus community 1.2.1. panicum schinzii – themeda triandra sub-community 1.3. eragrostis biflora – themeda triandra community 2. aristida congesta – themeda triandra major community 2.1. trichoneura grandiglumis – rhynchosia nervosa community 2.1.1. conyza bonariensis – eragrostis curvula subcommunity 2.1.2. antephora pubescens – digitaria argyrograpta subcommunity 2.2. eragrostis trichophora – cyperus capensis community 2.3. hyparrhenia hirta – themeda triandra community 2.4. melinis repens – eragrostis lehmanniana community. vegetation description 1. felicia muricata – themeda triandra major community this widespread grassland community is mainly associated with low-lying, flat open plains where the soils are generally sandy clay loam, with the clay content ranging between 18.5% and 27.0% (table 1). the vegetation has a uniform representation of grasses, with forbs and rarely shrubs interspersed between them. the two climax grasses t. triandra and d. eriantha (species group n) have a characteristically high presence, indicating good veld condition. this is further indicated by the notable poor occurrence of the pioneer grass a. congesta and the sub-climax grass eragrostis superba (species group g), which are known to occur in disturbed veld. other well represented species include eragrostis chloromelas, f. muricata (species group a) and e. lehmanniana (species group n). this major community has six diagnostic species: namely, indigofera alternans, pentzia globosa, tripteris aghillana, schkuhria pinnata, e. chloromelas and f. muricata (species group a). three communities are recognised within this major community. 1.1. panicum coloratum – digitaria eriantha community this grassland community is situated on flat plains on the city margins and also on fragmented natural areas between and within residential areas. it is differentiated and dominated by the grass species p. coloratum and eragrostis plana (species group b). other equally prominent grasses are e. chloromelas (species group a), t. triandra, e. lehmanniana, aristida adscensionis and d. eriantha (species group n). this community has a low number of species, with an average of only 11 recorded per sample plot. one sub-community is recognised within the community, which is differentiated by its relatively high soil moisture. 1.1.1. cyperus usitatus – digitaria eriantha sub-community this grassland community can be found occurring on flat open plains on the southern part of the study area. it is differentiated by species from group c, namely c. usitatus and anthospermum rigidum subsp. pumilum. this sub-community is dominated entirely by grasses, with only a handful of forbs showing a rare occurrence. the most prominent grasses are e. chloromelas (species group a), p. coloratum (species group b), t. triandra, a. adscensionis and d. eriantha (species group n). the only forb with a strong presence is c. usitatus (species group c), whilst t. aghillana (species group a) and o. depressa (species group n) are constantly present, but never prominent. an average of 13 species per sample plot was recorded. 1.2. eragrostis obtusa – sporobolus fimbriatus community this community is found in pasture areas in the south of bloemfontein, as well as towards the west of the city. it is dominated entirely by grasses, most notably eragrostis obtusa and s. fimbriatus, which are also the diagnostic species (species group d). other equally dominant grasses are t. triandra and d. eriantha (species group n), with f. muricata (species group a), h. pusillus and selago densiflora (species group n) found occurring consistently within the community, but never prominently. an average of 13 species per sample plot was recorded, and one sub-community is recognised within this community. 1.2.1. panicum schinzii – themeda triandra sub-community this is a sub-community found on the grassland plains and footslopes of the gently sloping small hills of the southern part of the study area. grasses are dominant, the most prominent being e. obtusa, s. fimbriatus (species group d), p. schinzii (species group e), t. triandra, e. lehmanniana, e. curvula and d. eriantha (species group n). the only forbs of prominent occurrence are f. muricata (species group a), conyza podocephala (species group e) and helichrysum dregeanum (species group n). the diagnostic species for the sub-community are p. schinzii, argemone ochroleuca, monsonia angustifolia, c. podocephala, chloris virgata and nenax microphylla (species group e). an average of 14 species per sample plot was recorded. 1.3. eragrostis biflora – themeda triandra community this community is associated with open grasslands on the periphery of the western suburbs of bloemfontein. the diagnostic species are the shade-loving grass eragrostis biflora, the erect perennial herb cyperus obtusiflorus, the karroid shrub lycium cinereum and the highly palatable grass digitaria tricholaenoides (species group f). the grassland is dominated by i. alternans, p. globosa (species group a), e. biflora (species group f), t. triandra and e. lehmanniana (species group n). this community has the lowest species richness of all the grassland communities of the bloemfontein area, with an average of only eight species per sample plot recorded. 2. aristida congesta – themeda triandra major community this widespread community represents grasslands that are characterised by sandy loam soils. the sand content ranges between 67.1% and 85.6% (table 1). it is mainly found in the northern and western homesteads and farming areas and, to a smaller extent, on the smallholdings to the east of bloemfontein. the most dominant species in the community is t. triandra (species group n), with other grasses such as a. congesta, e. superba (species group g) and e. lehmanniana (species group n) also prominent. the diagnostic species for the community are a. congesta and e. superba (species group g), grasses which are also indicators of veld disturbance. four communities are recognised within this major grassland. 2.1. trichoneura grandiglumis – rhynchosia nervosa community this is a community situated on the western part of the study area. it occurs on the sides of the kwaggafontein hills and also on the grass plains at the tempe airfield. seven diagnostic species are identified for the community: the grasses t. grandiglumis, pogonarthria squarrosa and heteropogon contortus and the forbs r. nervosa, pollichia campestris, moraea pallida and senecio burchellii (species group h). this community is partially dominated by the highly palatable grasses e. superba (species group g) and t. triandra (species group n) and the less palatable a. congesta (species group g), t. grandiglumis, p. squarrosa, h. contortus (species group h) and e. lehmanniana (species group n). other constantly present species are r. nervosa (species group h), o. depressa, s. densiflora and h. dregeanum (species group n). two sub-communities are recognised within this grassland community. 2.1.1. conyza bonariensis – eragrostis curvula sub-community this is a sub-community occurring on the north-western part of the study area, particularly at the tempe airfield and the neighbouring skydiving ground. the perennial grasses a. congesta (species group g), t. grandiglumis (species group h), t. triandra and e. curvula (species group n) are dominant. other species of notable occurrence include r. nervosa (species group h), c. bonariensis (species group i) and s. densiflora (species group n). the diagnostic species are salvia verbenaca, aristida canescens and c. bonariensis (species group i), whilst the strong presence of e. curvula and d. eriantha (species group n) further distinguishes the sub-community from the a. pubescens – d. argyrograpta sub-community. an average of 13 species per sample plot was recorded. 2.1.2. antephora pubescens – digitaria argyrograpta sub-community this sub-community is situated at the kwaggafontein hills, a conservation area on the western part of the study area. it occurs on the gently undulating sides of the hills, as well as on the adjacent bottomland area on deep, red sandy soils, with little or no surface rock. because this vegetation unit occurs in a conservation area, it is in primary condition and protected from the effects of grazing. it also has the highest species richness, with an average of 30 species per sample plot and the highest number of diagnostic species (20). some diagnostic species include: the grasses a. pubescens, d. argyrograpta, aristida stipitata, tragus koeleroides and cynodon dactylon, the herbs gazania krebsiana, commelina africana, harpagophytum procumbens and dicoma macrocephala and the karroid shrub chrysocoma ciliata (species group j). the vegetation consists of a wide variety of grass species, of which the most dominant is a. pubescens (species group j). its abundance is regarded as an indicator of good veld condition (van oudtshoorn 1999). other prominent grasses are a. congesta and e. superba (species group g), t. grandiglumis, p. squarrosa, and h. contortus (species group h), d. argyrograpta and a. stipitata (species group j), t. triandra and e. lehmanniana (species group n), whilst p. campestris (species group h), s. densiflora and h. dregeanum (species group n) are amongst the dominant forbs. 2.2. eragrostis trichophora – cyperus capensis community this plant community occurs on the smallholdings of the eastern city margin. species from group k differentiate the community: namely, the grasses e. trichophora and eragrotis gummiflua, the sedge c. capensis, the perennial dwarf shrub selago albida and the herb crotalaria sphaerocarpa. these diagnostic species are also the most dominant, with a. congesta (species group g) being the only other spe cies of prominent occurrence. it replaces t. triandra (species group n) in some overgrazed patches. only a limited number of species is present in this community and an average of only 11 species per sample plot was recorded. 2.3. hyparrhenia hirta – themeda triandra community this is a community occurring along the roadsides at the northern margin of the city, where the soil is slightly gravelly. the tall, tufted perennial h. hirta (species group l) is the only diagnostic species and entirely dominates, sometimes growing to heights exceeding 1 m. the other prominent species in this community are the grasses t. triandra, e. lehmanniana and e. curvula (species group n). an average of 11 species per sample plot was recorded. 2.4. melinis repens – eragrostis lehmanniana community this is a slightly disturbed community occurring on roadsides on the western and north-western part of the study area. the soil along the roads where this community is found is, in general, gravelly, at times with small stones also found. this is largely a grass community, with rhus lancea (species group m) the only woody species and t. aghillana (species group a) being the only forb of notable occurrence. the most dominant grasses include m. repens and enneapogon cenchroides (species group m), t. triandra and e. lehmanniana (species group n). e. cenchroides characteristically grows in clumps and, together with m. repens, dominates in disturbed patches, especially where the soils are exceptionally deep. the diagnostic species for this community are m. repens, e. cenchroides, r. lancea and fingerhuthia africana (species group m). an average of 12 species per sample plot was recorded. ordination the dca analysis complements the classification in the online appendix and reveals distinct discontinuities between the communities (figure 2). the cca ordination of sample plots and soil variables (figure 3) illustrates that communities of the f. muricata – t. triandra major community (community 1) occur in soils with relatively high organic matter and clay content and slightly lower ph (i.e. neutral soils). communities of the a. congesta – t. triandra major community (community 2), on the other hand, are associated with more sandy soils that have less organic matter, but slightly higher ph (table 1). there is a correlation between soil texture and organic matter content; organic matter is positively correlated with the clay content (r = 0.757; p < 0.001) and negatively correlated with the sand content (r = 0.666; p < 0.001). figure 2: a detrended correspondence analysis ordination of the grassland vegetation of bloemfontein, showing the relative positions of the relevés along the first two axes (axis 1 and axis 2). figure 3: a canonical correspondence analysis biplot of 25 sample relevés and soil environment variables for the first two axes: axis 1 (horizontal) and axis 2 (vertical). the biplot of the species and soil environmental variables (figure 4) further supports the assertion that species of the f. muricata – t. triandra major community (species groups a, b, c, d, e) have a strong association with higher clay content and organic matter in the soil. species from group f, together with the character species of the communities of a. congesta – t. triandra major community (species groups g, h, i, k) are strongly associated with the soil depth and sand gradients. in contrast, the other character species of the a. congesta – t. triandra major community, namely h. hirta (species group l), m. repens, e. cenchroides and f. africana (species group m) are strongly associated with the ph gradient. these species are commonly found on gravelly soil (van oudtshoorn 1999) and, in the study area, they are often located along roadsides and in road reserves. the only woody character species, r. lancea (species group m), is also strongly associated with the ph gradient and it often prefers calcareous substrates (van wyk & van wyk 1997). figure 4: a canonical correspondence analysis biplot of 59 plant species and soil environment variables for the first two axes: axis 1 (horizontal) and axis 2 (vertical). trustworthiness top ↑ the classification of the grassland communities of bloemfontein is based on data acquired through human observations and requires consideration of the trustworthiness of the data in the contexts below. reliability the nature of this study does not require repeated surveys and therefore the authors were able to satisfactorily draw conclusions from their research findings because they believe they have utilised the most widely used and arguably the most efficient of all types of vegetation sampling techniques. a potential bias, however, can arise during the vegetation classification but this is compensated for by the use of computer programmes such as twinspan, decorana and canoco. validity the authors believe the methodologies employed were successful at achieving the objectives of the study. discussion top ↑ the classification of the grasslands of the urban open spaces in bloemfontein revealed the existence of seven identifiable plant communities and four sub-communities, all represented within two major communities. the vegetation can be broadly classified as the t. triandra – e. lehmanniana grassland, where t. triandra is the dominant sweet grass. other major grasses include e. lehmanniana, e. curvula, d. eriantha and e. chloromelas. out of a total of 119 species recorded for the study area, 64 play a diagnostic role, 10 are companion species and a further 45 are either localised or of very rare occurrence. the distribution and composition of the plant communities are linked with habitat factors such as soil depth and texture, rockiness of the soil surface and habitat disturbance (especially utilisation by livestock).the results of the soil analysis indicate a correlation between soil texture and organic matter content, whereby organic matter is positively correlated with the clay content and negatively correlated with the sand content. these correlations can be explained in two ways. firstly, soils with high clay content have a high water holding capacity and therefore high moisture content. communities in such soils are, as a result, more productive; hence the high organic matter content in soil. secondly, according to foth (1990), clay plays a role in protecting organic matter decomposition, thus contributing towards the high organic matter content in the f. muricata – t. triandra major community. communities of the a. congesta – t. triandra major community, on the other hand, occupy areas of higher sand content that are characterised by low soil moisture content as a result of low water holding capacity. these communities are therefore less productive and hence have less organic matter. the distribution of species commonly found on gravelly soil along roadsides and in road reserves, such as h. hirta, m. repens, e. cenchroides and f. africana is strongly affected by ph. the soils near and along roads are known to have higher ph levels; this is attributed to calcareous road dust and leachate (auerbach, walker & walker 1997; johnston & johnston 2004). communities occurring on pasture land in farming areas show notable signs of overgrazing and trampling. there is also a tendency of selective grazing by livestock, through which animals over-utilise the palatable grasses. this results in the replacement of nutritious perennial grasses by annual grasses and weeds of inferior value. the invasion of the veld by the unpalatable species can have serious ecological implications because most of these invaders are a threat to the indigenous species. it is therefore necessary to manage the vegetation according to ecological principles in order to maintain optimum floristic composition and diversity. furthermore, there is a common practice of mowing or burning vegetation that grows along roads and railway tracks in the vicinity of residential and commercial areas. this vegetation, though disturbed and unstable, is vital simply because most of the natural vegetation in urban areas occurs in fragmented forms as a result of urbanisation (florgård 2007). the disturbed areas could therefore form dispersal corridors between the natural fragments and, in this way, act as stepping-stones for a variety of plant species (davis & glick 1978). the preservation of these disturbed areas may, in the end, offset the biological isolation of the natural areas and, in so doing, provide ecological continuity throughout the urban environment (millard 2004; poynton & roberts 1985). conclusion top ↑ the distribution and composition of the grassland communities of the bloemfontein area are linked with habitat factors such as soil depth and texture, with the f. muricata – t. triandra major community associated with clayey soils and the a. congesta – t. triandra major community found on sandy soils. the communities are also impacted by habitat disturbance, mainly in the form of overgrazing, mowing and burning. this phenomenon is not unique to these grasslands, because large portions of pristine vegetation in the grassland biome of south africa have been degraded and destroyed due to livestock grazing pressure and the ploughing of arable land. the proper management and conservation of urban open spaces requires in-depth knowledge of the spatial distribution, floristic, structural and functional compositions within the major vegetation types in the urban environment. the present vegetation study was initiated to contribute towards formulating ways for the proper functioning and management of the open spaces within the bloemfontein metropolitan area. the benefits of conserving urban vegetation are immense; scientifically, socially, and economically. open space within urban areas has value upon urban microclimate, hydrology, biodiversity, and ecological processes (nicholson-lord 1987; platt, rowntree & muick 1994). this phytosociological study provides valuable information on the structure and composition of the vegetation of the bloemfontein metropolitan area, which can be utilised by the relevant local and national authorities for environmental planning and conservation strategies for the area. acknowledgements top ↑ the assistance of the following persons is sincerely appreciated: dr pieter le roux (university of the free state) for his help in interpreting the soils and land types of the study area, ms yvonne dessels and the soil science laboratory staff of the university of the free state for the soil analysis, as well as the two anonymous reviewers for their useful comments and suggestions. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions m.n.v.d. 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africa, struik publishers, cape town. van wyk, e., cilliers, s.s. & bredenkamp, g.j., 2000, ‘vegetation analysis of wetlands in the klerksdorp municipal area, north west province, south africa’, south african journal of botany 66, 52–62. wood, j., low, a.b., donaldson, j.s. & rebelo, a.g., 1994, ‘threats to plant species diversity through urbanisation and habitat fragmentation in the cape metropolitan area, south africa’, in b.j. huntley (ed.), botanical diversity in southern africa, pp. 259–274, creda press, cape town. pmid:7941307 article information authors: richard j. stirzaker1 dirk j. roux2 harry c. biggs3 affiliations: 1csiro land and water, canberra, australia 2south african national parks, george, south africa 3south african national parks, skukuza, south africa correspondence to: richard stirzaker email: richard.stirzaker@csiro.au postal address: po box 1666, act 2601, australia dates: received: 04 june 2010 accepted: 28 nov. 2010 published: 13 may 2011 how to cite this article: stirzaker, r.j., roux, d.j. & biggs, h.c., 2011, ‘learning to bridge the gap between adaptive management and organisational culture’, koedoe 53(2), art. #1007, 6 pages. doi:10.4102/koedoe.v53i2.1007 copyright notice: © 2011. the authors. licensee: openjournals publishing. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) learning to bridge the gap between adaptive management and organisational culture in this essay... open access • abstract • introduction • science and certainty • complex and ‘knowable’ systems • adaptive management • adaptive management and organisational culture • learning • conclusion • references abstract (back to top) adaptive management is the problem-solving approach of choice proposed for complex and multistakeholder environments, which are, at best, only partly predictable. we discuss the implications of this approach as applicable to scientists, who have to overcome certain entrained behaviour patterns in order to participate effectively in an adaptive management process. the challenge does not end there. scientists and managers soon discover that an adaptive management approach does not only challenge conventional scientific and management behaviour but also clashes with contemporary organisational culture. we explore the shortcomings and requirements of organisations with regard to enabling adaptive management. our overall conclusion relates to whether organisations are learning-centred or not. do we continue to filter out unfamiliar information which does not fit our world view and avoid situations where we might fail, or do we use new and challenging situations to reframe the question and prepare ourselves for continued learning? conservation implications: for an organisation to effectively embrace adaptive management, its mangers and scientists may first have to adapt their own beliefs regarding their respective roles. instead of seeking certainty for guiding decisions, managers and scientists should acknowledge a degree of uncertainty inherent to complex social and ecological systems and seek to learn from the patterns emerging from every decision and action. the required organisational culture is one of ongoing and purposeful learning with all relevant stakeholders. such a learning culture is often talked about but rarely practised in the organisational environment. introduction (back to top) pursuing the life sciences is as much a calling as it is a career. we are drawn to a science course at university through some combination of a fascination with the living world and a desire to use natural resources wisely. the renowned biologist edward o. wilson says that at age eighteen he saw science as the study of ants, frogs and snakes and a wonderful excuse to stay outdoors (wilson 1998). for many of us, the objective view of the world that science provides appealed more than the literature and history classes we took at school. during university-level practicals, students have to solve problems to which objective answers exist. for example, how much chitin does a crab shell contain? or, what is the identity of the bacteria in the broth? what is the change in momentum when two bodies collide? the tutor already knows the correct answer and if the experiment is performed properly, the students should get it right too. however, there may be several different interpretations to the meaning of a poem, which can all be apparently acceptable. the scientific method gives one correct result, which is independent of the observer. modern science has built this predictive capability on four essential components. firstly, one states a clear hypothesis about cause and effect that is testable by experiment. secondly, one designs an experiment that tests a prediction emanating from the hypothesis in a controlled environment. thirdly, one replicates the experiment to show that the observations are not the result of a chance event. fourthly, the work is documented and subjected to peer review before the new knowledge can serve as building blocks for further investigations. the ability to tease apart a system into its constituent components and study it systematically allows scientists to infer cause and effect. such reductionism is said to be the primary and essential activity of scientific research (wilson 1998). at the outset of this essay, we offer a very brief description of organisational culture as it relates to science, our interest in adaptive management, and why we foresee an uneasy relationship. culture consists of bundles of shared norms, which are behaviours common to a group. norms give a group a sense of cohesion and protection against undesirable change, but can simultaneously cause a group to resist new, potentially beneficial ideas (ehrlich & levin 2005). many scientists share a culture centred around the robust methodology that underpins their craft: the production of reliable information and the ability to make predictions, quantify uncertainty and expose error. scientific thinking has produced knowledge and products that have contributed enormously to economic development and social upliftment over the last 300 years. scientists not exposed to further study of the humanities, or even an introductory course on the philosophy of science, are often surprised by growing criticism of the very norms most of them regard as self-evident. even if it were valid to pass off some of these critics as postmodern deconstructionists, scientists still have to heed a call from within their own community about the need to think differently about contemporary issues facing society (lubchenko 2008; steffen, crutzen & mcneill 2007; walker & salt 2006). for example, ulanowicz (2009:93) writes that ‘it is indeed feasible to march directly into the jaws of oblivion on the tacit assumptions that support conventional science’. holling (1995:29) warns of the ‘pathology’ that emanates from scientists’ belief that they fully understand cause and effect – that success in managing one target variable in isolation leads to ‘less resilient and more vulnerable ecosystems, more rigid and unresponsive management agencies, and more dependent societies’. it is not our purpose here to provide a critique of reductionist science; suffice it to say that we sympathise with the view of ulanowicz (2009) that biology and ecology are not entirely reducible to physics. life scientists cannot fully explain the world from the bottom up and therefore are interested in finding approaches to managing ecological systems that accept and act on this understanding. moreover, since human systems interact with biological systems in diverse ways we propose that adaptive management is the problem-solving approach of choice for environments that are highly changeable, heterogeneous and often unpredictable, and usually involve multistakeholder interest. but we also anticipate that its implementation will frequently conflict with aspects of our own science culture and that of the organisations where we work. science and certainty (back to top) modern society’s knowledge of the physical sciences is so well developed in many areas that we put our total trust in its products. most of us expect an aeroplane to get us safely to our destination even if we do not understand all the intricacies involved in the process. one reason for this certainty is that we can be reasonably sure about the boundaries of the system. we believe that the engineers, pilots and air traffic controllers know which factors they need to understand well, and which they can ignore. the biological sciences present more of a challenge because the boundaries of a system are not always clear. an experiment is performed in ‘controlled’ conditions, yet it is impossible to exclude all factors extraneous to the hypothesis owing to practical constraints often associated with larger-scale experiments. not all of us can fumigate small mangrove islands to study the re-colonisation process (simberloff & wilson 1969). statistical techniques provide an objective means for identifying cause and effect in controlled experiments and authors of scientific publications know that reviewers start to object when one extrapolates findings too far from the experimental conditions under which the data were collected. the new knowledge holds only when other factors are excluded; that is, scientists’ claims are valid within certain defined boundary conditions. by speculating outside the narrow range of the measured data or the conditions in which they were collected, one can no longer claim the same certainty. in other words, our knowledge is obtained by framing or constraining the problem. what does this mean for understanding cause and effect in biological or ecological research? it is tempting to believe that the problem of boundary conditions can be overcome just by doing more experiments until all the combinations and permutations have been exhausted. this view is problematic. as already discussed, replicating experiments under controlled conditions is extremely difficult at ecosystem level, and has even less relevance for the action research of the social disciplines (rogers 2006). but the problem may lie even deeper. walters and holling (1990) state that our ecological knowledge is not only incomplete but also elusive, particularly when we consider the range of values held by different groups in society and the political constraints to action. gallopin et al. (2001) refer to a degree of ‘irreducible uncertainty’ associated with complex social–ecological systems. complex and ‘knowable’ systems (back to top) in order to test the claim of ‘irreducible uncertainty’, one needs to differentiate between different types of p roblem and be able to infer cause and effect accordingly. snowden (2002) categorises problems into those that may be difficult or complicated but are ultimately ‘knowable’ through reductionist scientific method, and problems that are complex and always characterised by an inherent degree of uncertainty. for example, we can categorise an aeroplane as a complicated but knowable machine. although it is made of thousands of different parts, the function of each in relation to another is understood. apart from the most extreme conditions, aeroplanes behave in a predictable way, and we trust them with our lives. human systems tend to be complex: the people that make up the system change and the way they relate to one another is highly context dependent and therefore not entirely predictable. snowden gives the following example: consider what happens in an organisation when a rumour of reorganisation surfaces: the complex human system starts to mutate and change in unknowable ways; new patterns form in anticipation of the event. on the other hand, if you walk up to an aircraft with a box of tools in your hand, nothing changes. (snowden 2005:105) one can accept the notion of an aeroplane being complicated but knowable, while a human system is very unpredictable, but what about biology and ecology? surely biological systems are subject to natural laws that give the system some form of deterministic behaviour? ulanowicz (2009) cautions that the idea of tight cause and effect in open systems should be set aside in preference for poppers’ view of ‘propensities’. a propensity downgrades cause and effect to a more general likelihood of one factor influencing another. more importantly, it is the combined effect of several propensities acting together on the whole system that facilitates unique and sometimes surprising behaviour of a specific system. in table 1 we distinguish between problems that are complicated but knowable and those that are complex (informed partly by snowden and boone 2007). the complexity of the problems in the right-hand column of table 1 is due to feedbacks, thresholds and, often, nonlinear interactions within the system, together with lags and cross-scale effects. such factors combine to give the system a degree of uncertainty. an example of feedback is the reinforcement of terrestrial warming as ice sheets melt, because there is less ice to reflect the sun’s energy. an example of a threshold effect is the rapid switch from savannah vegetation to woodland thicket once perennial grasses that support fire reach a critically low density (walker & meyers 2004). nonlinear interaction means that a small change in one factor can have a big effect somewhere else. when there are multiple such interactions it becomes practically impossible to keep track of all the causal relationships. table 1: some characteristics of knowable and complex problems. complex systems also have so-called emergent properties. an analogy might be ‘team spirit’. the statistical attributes of each member of the team may be known, but the joint interaction of the team sets up a dynamic that strongly affects how the team performs as a unit. in other words, there are mutually beneficial interactions between players that give the team its unique character. if we disassemble the team to study each player’s attributes in detail, the team spirit, or emergent property, disappears. adaptive management (back to top) feedbacks, thresholds, multiple nonlinear interactions, lags, chance events and emergent properties contribute to a general uncertainty about cause and effect, and, consequently, the impact of our management actions. this realisation led to the development of the field of adaptive management (holling 1978; lee 1993; walters 1986). its fundamental premise is that the puzzle of a social–ecological system can never be fully solved by studying the pieces. we have to use real-life management of the system as a whole and turn it into an experiment by asking the right questions, implementing decisions, collecting the right data and learning from the experience. the emphasis is on formulating an explicit mental model, however imperfect, and then acting accordingly by managing and monitoring to see how our understanding can be improved as we gain further insight of the system. furthermore, many of the ecological problems we face are as much a controversy over values as disputes about cause and effect. this challenges the positivist view of science, which regards science as the principal producer of reliable knowledge that should be passed on to those with a management responsibility (ziman 2000). broader society now demands that their local and experiential knowledge, as well as their values, be considered in management plans. therefore, when both the facts are uncertain and their interpretation is contested, we need an approach that can integrate knowledge from different sources and treat management activities as experiments from which we can learn. adaptive management is a way of getting around the dilemma of delaying decisions until we fully understand all the potential consequences of our actions. the act of management is itself an experiment, but clearly not in the traditional sense of controls and replication. to distinguish adaptive management from simple trial and error, considerable effort should be put into integrating existing information from different disciplines and perspectives. appropriate models should be used to frame the questions, eliminate the least likely solutions and identify the knowledge gaps (stankey, clark & bormann 2005; walters 1997). monitoring is a central issue. adaptive management needs an intellectual paper trail to show that reasoning underlies the actions – we cannot learn without this (lee 1993; venter et al. 2008). the conceptual framework, whether represented by a simple diagram or sophisticated model, should be matched by the amount of effort put into monitoring. unfortunately, monitoring is much more expensive than modelling, which can create tension between researchers and funders or managers. often the true value of monitoring will be reaped only by the next generation of scientists; a problem for those responsible for paying the bills now (walters 1997). conversely, it is the responsibility of scientists to identify variables that are most likely to be indicative of system behaviour. this may involve identification of integral measures that remove much of the ‘noise’ and variables that warn in advance of an approaching threshold (stirzaker et al. 2010). we simply cannot measure everything. according to stankey et al. (2005) the principles of adaptive management are widely acclaimed, but remain more an ideal than a demonstrated reality. one of the several reasons rogers, roux and biggs (2000) cite for this is that the new way of operating does not comply with the old organisational culture with an authoritarian structure. moreover, when investigating large multidisciplinary problems we get overloaded with information and often experience ‘turf protection’ among scientists and between scientists and managers. this raises a new question: if issues that we once saw as ‘knowable’ are in fact complex and demand a radically different problem-solving approach, do we also need to think through the ways our organisations operate? adaptive management and organisational culture (back to top) scientists responding to the challenge of living with more uncertainty can find their organisations moving in the opposite direction. the conventional view of curiosity-driven research that leads to new findings, beneficial applications and tools for the improvement of human welfare (ravetz 2004) has given way to formal methods of planning and accountability. science no longer has special status in a government’s budget. the case for investmentin science must be carefully argued with explicit costs and benefits, with timelines to show what will be delivered when and by whom. organisational culture and adaptive management are likely to clash, at least initially, on several fronts. firstly, scientists are expected to produce the knowledge that managers need to make informed decisions. a focus on inherent unpredictability seems to be undermining the foundation of this social contract. secondly, organisations spend considerable time streamlining their portfolio of work into ‘manageable’ units aligned to corporate goals, whereas adaptive management can be a messy web of relationships encompassing scientific, social and political perspectives. thirdly, adaptive management requires us to be open to learning things that may be counter to the way we normally operate. it requires a level of flexibility that challenges the way that things have always been done. as scientific organisations strain under the pressure to adopt a more overt ‘business principles’ approach, there is a greater focus on specifying outputs of programmes well in advance and minimising risk at all stages of the project. reporting on project deliverables and milestones is required on shorter timescales, which makes it easier to follow the contract than to follow up surprises. whereas accountability and risk management are obviously important, rigid management systems run counter to the nature of adaptive management. organisations certainly need to balance order with creativity, but the desire for certainty and control can overwhelm the desire to nurture the flexible learning approach required for adaptive management. according to wheatley (2005), organisations should resist the notion that there is some optimum structure that will deliver results: if a system becomes too homogenous, it becomes vulnerable to environmental shifts. if one form is dominant, and that form no longer works when the environment shifts, the entire system can collapse. … if leaders fail to encourage diverse ways of doings things, they destroy the system’s capacity to adapt. (wheatley 2005:78) if, for example, we lose the balance between flexibility and responsibility and opt for an overly planned and rigid system, we may find we spend more time interacting with the organisation and less with the real world. at the other end of the spectrum the organisation may become too chaotic when everyone follows their own plans. people do not learn from their mistakes and bad behaviour is not brought to account. both extremes are counter-productive. wheatley (2003:39) writes: ’if a system has too much order, it atrophies and dies. yet if it lives in chaos, it has no memory.’ when science is seen as just another business, goals such as improving efficiency are accepted without question. this sounds sensible at first, but there is evidence that targeting efficiency as the prime goal can destroy the very thing we are trying to manage (walker & salt 2006). rogers et al. (2000) cite the example of a ‘command-and-control’ approach to managing water resources. what starts out as efficient delivery of services easily spills over into exploitation when the focus is too narrow and agencies take too long to respond to feedbacks. cilliers (2006:109) argues that the value of organisations lies in their ability to be ’stable enough not to be buffeted around by every fluctuation, [and] … flexible enough to be able to adapt when necessary’. he advises against a culture where ’speed is linked with efficiency, and has become a virtue in itself.’ our real identity is forged when we are able to reflect adequately on our experience, and based on that reflection, to resist certain change (cilliers 2006). yet, when the pressure is on to deliver results too quickly, we are more likely to react than to reflect. learning (back to top) if an organisation is going to embrace adaptive management, it will have to learn to do things differently. stankey et al. (2005) propose that we normally learn by accumulating new facts, but our understanding moves ahead in leaps when we ask new questions and see the old facts in a new light. this is critical for adaptive management. instead of filtering out information that is unfamiliar or does not fit our world view, or avoiding situations where we might fail, we use challenging situations to reframe the question and prepare ourselves for learning new things. garvin (1993) defines a learning organisation as one ‘skilled at creating, acquiring, and transferring knowledge, and at modifying its behaviour to reflect new knowledge and insights.’ modifying behaviour is difficult and should involve three overlapping phases. • cognitive: members of the organisation are exposed to new ideas, expand their knowledge, and begin to think differently. • behavioural: employees begin to internalise new knowledge and alter their behaviour (as manifested, for example, in the use of new vocabulary). • performance improvement: changes in behaviour lead to measurable improvements in results or outcomes. thus far we have dealt with the cognitive aspects of adaptive management: the idea that we should distinguish complex systems from those that may be difficult but ultimately able to be fully understood (table 1). we have also addressed the early behavioural changes: adopting a new language that goes beyond the use of new vocabulary to determine what the new concepts really mean in a specific context and how they can inform our approach to science. the third step – applying the new concepts in the real world – is, however, the major stumbling block. of course, if there is no performance improvement the ‘new’ behaviour will be challenged and will most likely be replaced by a next wave of ideas, or the organisation will default to its old ways. learning is the mechanism through which we change our individual and collective understanding of our world. new knowledge enables us to respond differently to new circumstances and challenges. the rate and relevance of our learning will, in effect, determine our ability to respond to external changes effectively. in this sense, learning proficiency relates to what we should learn about (and what we should forget), who we should learn with, and how we should learn. to accommodate new knowledge, previous learning and beliefs sometimes have to be left behind. selectively ‘forgetting’ outdated knowledge is referred to as unlearning (becker 2005). however, unlearning may not be a straightforward or easily manageable activity. individuals (often unknowingly) protect existing knowledge by actively disregarding conflicting information (lyndon 1989). it appears that more recently acquired knowledge is easier to relinquish than knowledge that was acquired and reinforced over an extended period of time. experts may be especially susceptible to ‘trained incapacity’ (miller & morris 1999): the more someone’s knowledge is shaped by learning within a defined field, the harder it becomes to associate with knowledge that emerges from other fields. environmental issues inevitably imply the involvement of multiple stakeholders. therefore, life scientists need to be prepared to learn together from diverse sources (keen, brown & dyball 2005; pahl-wostl & hare 2004). we should not only settle for compromise but strive for a consensus that can distribute the benefits and costs of our interventions in a way that is (1) equitable and (2) within the ecological limits (rogers 2006). compromise involves trading off conflicting demands against those who hold a contrary position, creating winners and losers. the approach of seeking consensus is about moving beyond the problem and ‘developing a set of shared values that guide future decision making’ (rogers 2006). ongoing learning is uncomfortable. it is much easier to believe we already know most of what we need to know. if we feel overwhelmed by new information our dominant learning mode is reactive and we tend to reinforce pre-established knowledge and frames of reference. scientists need to perceive their working environments as safe to envisage alternative futures and to learn along new and dynamic trajectories towards such futures (senge et al. 2005). furthermore, to learn with other parties who may hold very different world views, requires us to learn with empathy and humility. to be empathetic means to consider different perspectives and assumptions, temporarily suspending our own in the process, so that we can inquire into the reasons for people’s views (senge et al. 1999). in this sense, humility means acknowledging that the knowledge base in any given field is too vast for a single person to master. even the expert’s knowledge is only a partial reflection of what is known. however, by combining one’s partial knowledge with that of others, one can, in practice, use more knowledge than one’s own (wenger 2005). conclusion (back to top) successful adaptive management in a multistakeholder context rests on three pillars, namely the ability to form a robust, shared conceptualisation, the ability to monitor key variables that will shed light upon this conceptualisation, and the ability to learn from the experience. if any of these are compromised, the structure will collapse. it is easy to be so enamoured by the conceptualisation of the problem that we fail to invest in thorough monitoring, and equally easy to keep collecting data without knowing how the knowledge will be used. the test is: are we still learning and can we document our learning journey (venter et al. 2008)? learning is never quick or easy, and involves travelling along detours and going down blind alleys. when disillusioned, scientists should avoid the trap of falling back into the old pattern of over-promising and under-delivering as they proffer ‘silver bullet’ solutions for complex problems to those who control the purse strings. similarly, organisations should resist defaulting to command-and-control systems that appear to have delivered some certainty in the past. organisations will have to find and foster the champions of adaptive learning, including the visionary activist, the respected integrator and the rebel bureaucrat (gunderson, holling & light 1995). there will need to be a shift from the view that benefits come from power or withholding information and ideas, to one where benefits come from sharing, and there are clear incentives to reflect this. learning together rather than competing against one another is absolutely central. science culture has been forged in a competitive environment – competition for best ideas to secure limited funding, competition for space in top journals, and even a league table of citation metrics that purport to show how useful our work has been. learning should involve exploring, discovering, reflecting, listening and sharing frustrations and surprises. managers, scientists and stakeholders need to see themselves as part of the same community, where benefits and risks are shared within the context of a shared vision. rogers and breen leave us with the core challenge: perhaps the most important lesson ecologists should learn is not to enter the new social theatre as ’experts’ (ludwig 2001), but as co-learners, interactive players seeking consensus on stage. for some ecologists, and for ecology as a science, this transition will certainly be difficult. we will judge success by a shift from research outputs that impress peers to outcomes that allow society to better respond to environmental challenges. 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knowledge, vintage books, new york. ziman, j., 2000, real science: what it is, and what it means, cambridge university press, cambridge. doi:10.1017/cbo9780511541391 abstract introduction material and methods results discussion acknowledgements references about the author(s) francois roux department of zoology, university of johannesburg, south africa scientific services, mpumalanga tourism and parks agency, south africa gert steyn department of zoology, university of johannesburg, south africa clinton hay department of biological sciences, university of namibia, namibia ina wagenaar department of zoology, university of johannesburg, south africa citation roux, f., steyn, g., hay, c. & wagenaar, i., 2018, ‘movement patterns and home range size of tigerfish (hydrocynus vittatus) in the incomati river system, south africa ’, koedoe 60(1), a1397. https://doi.org/10.4102/koedoe.v60i1.1397 original research movement patterns and home range size of tigerfish (hydrocynus vittatus) in the incomati river system, south africa francois roux, gert steyn, clinton hay, ina wagenaar received: 21 apr. 2016; accepted: 05 apr. 2018; published: 27 june 2018 copyright: © 2018. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract historical data suggested that the tigerfish (hydrocynus vittatus) of the incomati river migrates upstream and downstream as part of their life history. it has been suggested that this movement was a prerequisite for successful spawning in inundated floodplains in mozambique. recent advances in aquatic radio telemetry provided a reliable mechanism to monitor fish movement and increase knowledge of the ecology of tigerfish. from 04 january 2003 to 22 december 2003, 41 tigerfish in the incomati river system were fitted with radio transmitters to record movement patterns and estimate home range size. on average, each fish was tracked 72 times, and the total number of fixes was 2971 over the study period, including 1322 summer fixes and 1649 winter fixes. the mean longest distance travelled by tigerfish was 730 m (range = 75 m to 3200 m). the home range size varied between individual fish, but on average fish stayed within a defined home range of 48 846 m2. tigerfish showed high site fidelity to specific habitats within specific activity zones and movement occurred primarily within these defined zones. differences in movement pattern, longest distance travelled and home range size could not be attributed to the sex or size of the fish. no large-scale movement patterns associated with specific life history activity were observed; thus, previous reports of large-scale downstream migrations and spawning migrations appear to be invalid. the presence of weirs in the study area impedes free fish movement as these weirs create migration obstructions. conservation implications: river regulation such as damming, water abstraction, obstructive barriers and channel modification may have a detrimental impact on the survival strategy of this species. implementation of these results in a management policy will provide a reliable basis for species specific requirements such as upstream reservoir release management; minimum flow volumes required for downstream ecosystem maintenance and management and planning of structures obstructing natural flow. introduction the freshwater fish genus hydrocynus is represented by six species, all endemic to africa. they are pikelike predators, commonly termed ‘tigerfishes’ for their prominent dentition and dark lateral stripes (gery 1977). in southern africa, one of these species, hydrocynus vittatus (commonly known as tigerfish), occurs in the zambezi and okavango rivers and in the lowveld reaches of coastal systems (skelton 2001). the southern african tigerfish (h. vittatus) has a limited distribution in south africa, where it is restricted to the lowveld reaches of the limpopo river system, mainly within the kruger national park (knp), and further south in the lower reaches of the usutho and phongolo rivers (gaigher 1967). the incomati river system (south africa) is a marginal area in the distribution range of tigerfish where they occur in relatively low abundance. being essentially a lowveld species in south africa, it is intolerant to cold water and migrates downstream to lower lying reaches of these rivers during winter where water temperatures are higher and more stable (pienaar 1978; steyn et al. 1996; van loggerenberg 1983; skelton 2001). mortalities caused by a sudden drop in temperature (< 16.0 °c) related to cold water in the incomati river were reported on several occasions (deacon 1991; gagiano 1997; personal observation by authors; van loggerenberg 1983). gagiano (1997) reported mortalities in the piet grobler dam in the knp at a temperature of 14.5 °c during the winter period. the habitat and environmental conditions in the incomati river system differ considerably from the favourable conditions present in the larger northern tropical river systems such as the zambezi river. tigerfish are inhabitants of open, well-oxygenated waters such as found in the larger rivers and lakes (pienaar 1978). in contrast to the larger rivers and lakes in the north of south africa, the rivers of the knp are relatively small, highly regulated because of anthropogenic impacts and subject to extreme seasonal variations (du preez & steyn 1992; gertenbach 1991). variation and flow volumes, especially in the presence of instream damming structures such as weirs, can severely impact the ability of fish species to migrate in accordance with their life history requirement (baras & lucas 2002). furthermore, all the major rivers of the knp are subjected to high silt loads which can severely reduce dissolved oxygen concentrations of the water and may be lethal to fish (buermann et al. 1995). there has been a long history of fish mortalities in the knp caused by large amounts of suspended particles present in the water (knp annual reports 1946–1992). the negative impact of increased silt loads on the aquatic macro-invertebrate diversity in the major rivers of the knp was reported by moore and chutter (1988). sub-lethal effects of suspended solids on fish are varied and include negative impacts on reproduction, egg survival, growth, oxygen consumption, haematology, feeding and social behaviour (crouse, callahan & malaug 1981; wilber 1983). indirect effects include reduced food availability, clogging of gillrakes and filaments, reduced growth rate, reduced resistance to disease and disturbances of natural movements and migrations of fish (albaster & lloyd 1980; bruton 1985). tigerfish has a prominent ecological status as top predator, sharing the same trophic level as crocodiles in the knp riverine ecosystems. their limited presence in the knp and their vulnerability to impacts described above served as motivation for several studies since the work of gaigher (1967). in south africa, research on tigerfish concentrated on ecological aspects (gaigher 1970, 1973; 1975; gagiano 1997; van loggerenberg 1983), reproduction (steyn 1993; steyn & van vuren, 1992; steyn et al. 1996), tooth replacement (gagiano, steyn & du preez 1996), age estimation and maturity (gerber et al. 2009) and genetics (kotze et al. 1998). recent advances in aquatic radio telemetry provided a reliable means to acquire further information on the behaviour ecology of fish species and to improve our knowledge on tigerfish. despite several comprehensive studies as mentioned above, conservationists and river managers were still left with key questions on the (1) migrational requirements, (2) movement patterns and (3) ability to overcome obstructions in order to maintain functionality of a viable tigerfish population in the incomati river system. the objective of this study was to use biotelemetry to answer these key questions. material and methods description of the study area the incomati river drains parts of mpumalanga, swaziland and mozambique between the limpopo river system in the north and the phongolo river system in the south. it is economically one of the most important river basins in south africa, and it consists of three adjacent sub-basins: the komati, crocodile and sabie (darwall et al. 2009). the main river descends from the highland plateau in mpumalanga and swaziland and flows through the coastal plains of mozambique to the indian ocean just north of maputo at villa laisa. the total basin area is about 46 800 km2 of which 63% is in south africa, 5% in swaziland and 32% in mozambique. the average discharge of the incomati river basin at the estuary is about 100 m3/s to 200 m3/s, corresponding to about 3600 million m3 per year, to which south africa contributes 82%, swaziland about 13% and mozambique about 4% (darwall et al. 2009). the study area includes two rivers, namely the crocodile river and the komati river, which join to form the incomati river below the border town of komatipoort. the crocodile river flows along the boundary of the knp, and at the confluence, the border extends across the river to also include the lower reach of the komati river (figures 1 and 2). below the confluence, the incomati river can be described as a meandering river, incised into a wide sandy river bed, and in some sections, it flows through multiple bedrock channels. the river varies between 40 m and 50 m wide, with mostly large sandy pools and occasional rapids and a few riffles (roux et al. 1990). collection and tagging were done upstream and downstream of the confluence between knp and tenbosch weirs and the low-water bridge in the komati river. the choice of the collection and tagging area was motivated by the relative abundance of tigerfish in this river reach. the ability of tigerfish to overcome obstructions and their various home ranges later defined the extent of the study area. historically, tigerfish distribution data would indicate that tigerfish occur up to an altitude of 300 m in the incomati river system. gaigher (1967) previously collected tigerfish in the crocodile river gauge close to the town of nelspruit and in the komati river close to the town of tonga on the border between south africa and swaziland. consequently, the experimental design made provision for long-distance tracking in relation to historical distribution in the incomati river system. figure 1: map indicating the location of the study area in the mpumalanga province of south africa, in close proximity to mozambique. the applicable rivers are illustrated and marked with the numerous weirs. figure 2: map of the study area indicating tagging sites of all 41 radio-tagged fish. collection and handling of the species collection and handling of fish were performed in such a manner as to minimise physical and physiological stress to the specimens (spedicato, lembo & marmulla 2005). tigerfish were caught using two techniques: rod and reel with artificial lures and fly-fishing, both using barbless hooks to reduce injury to fish and to facilitate quick release, thereby reducing lactic acid stress and ensuring survival after handling and release (gerber et al. 2017). tagging of fish in total, 41 sexually mature tigerfish were tagged with radio transmitters (advanced telemetric systems inc. ats, usa, 142 mhz–144 mhz) in 2003. as the sexing of h. vittatus is relatively difficult based on external characteristics, males were only positively identified if they were ripe and running and producing semen. large females in or close to the spawning season were easily sexed as they displayed characteristic body size, form and weight (gaigher 1967; gagiano 1997; g.j. steyn pers. comm., 2003). the standard length (sl) was measured (mm), and mass (g) of each specimen collected was determined using a measuring tape and a bogagrip (scale). following capture, fish were anaesthetised with 2-phenoxyethanol (0.3 ml/l), minimising hyperactivity and stress. the radio transmitters were selected from ats models f2040, f2130 and f2010 with trailing whip antennae and were externally attached to fish with two strands of orthopaedic wire (0.65 mm diameter) below the dorsal fin following thorstad, økland and heggeberget (2001). to facilitate rapid healing of the needle wounds, the tagged fish were placed in a terramycin bath (25 mg/l water) for 10 min prior to release. the deployment of the small f2040 transmitters made it possible to tag smaller fish because of the relatively low weight of the transmitter, but remaining within the 2% rule (brown et al. 1999; peake & mckinley 1997). all radio-tagged fish were released at their respective sampling points, and staggered deployment over several months allowed for continuous data retrieval over a full year period, consequently covering all seasons (table 1; figure 2). staggered deployment was necessary because of the limited lifespan of the transmitters. table 1: individual fish collection, release and radio-tagging data (home range and longest distances). table 1 (continues...): individual fish collection, release and radio-tagging data (home range and longest distances). fish tracking procedures fish were tracked using a challenger r2100 receiver and a four-element yagi antenna (ats inc.) over a 12-month period (04 january to 22 december 2003) on average every second day, covering both summer and winter periods. care was taken to minimise behavioural side-effects by keeping a reasonable distance from tagged fish (hocutt, seibold & jesien 1994). tracking was done on foot from the banks of the river by using the homing-in technique (jick 1979). if there was any uncertainty regarding the position of the fish, the triangulation method was then applied (jick 1979). in instances where fish were lost, aerial surveys were conducted using a micro-light aircraft to relocate a specific fish. for all tracking surveys, location was determined using a handheld global positioning system receiver (garmin etrax). upon detection, the global position system (gps) coordinates of the fish’s location were noted (accuracy ± 5 m). hydrology, water quality and meteorological data flow levels in the incomati river system were determined from daily readings at the knp gauging weir in the incomati river. water temperature, ph and conductivity were recorded daily in the crocodile river, komati river and below the confluence of the two rivers (in the incomati river) using eutech portable microprocessor-based water quality instruments. meteorological data were gathered from a nearby weather station (transvaal sugar board, komatipoort), including rainfall, minimum and maximum air temperatures. data analysis two fish that moved out of the study area into mozambique shortly after tagging were excluded from the analysis. in addition, a third fish showed no movement for an extended period after tagging and was presumed dead and excluded from the analysis. descriptive statistics for the entire study period (summer and winter) were based on more than 10 fixes per fish for 38 fish. gps coordinates of the radio-tracked tigerfish were used to calculate longest distances travelled and to determine home range sizes. bi-variate gaussian or normal distribution kernel methods (seaman & powell 1996; silwerman 1986; worton 1989) were used to plot home ranges. this group of methods is part of a more general group of parametric kernel methods that employ distributions other than the normal distributions as the kernel elements which are associated with each point in the set of location data. because of the meandering nature and relatively small width and limited available habitat within the incomati river system during low flow periods at specific sites, an adaptation of the simplified minimum convex polygon (mcp) (baker 2002; creel & creel 2002; meulman & klomp 1999) was used. boundaries of home ranges were drawn using different sets of location data (planet gis). this method of using the shoreline as a boundary of the home range is a widely accepted and commonly used method in fish telemetry experiments (hocutt et al. 1994). for ease of statistical analysis, a binning algorithm was implemented in which the longest distance travelled, home range size and the radio-tagged fish were grouped in classes according to their magnitude. for longest distance travelled (økland et al. 2005), fish were organised in classes ranging from 100 m to 500 m, 501 m to 1000 m, 1001 m to 1500 m, 1501 m to 2000 m and > 2000 m travelled. the home range size were classed in groups ranging from 0 m2 to 10 000 m2, 10 001 m2 to 20 000 m2, 20 001 m2 to 50 000 m2, 50 001 m2 to 100 000 m2 and > 100 000 m2. the ibm spss statistics 18 program was used for basic and inferential statistics which include frequencies, normality, correlations and comparisons (spss 2009). ethical consideration the project proposal was approved with ethical clearance by the faculty of science, university of johannesburg and mpumalanga parks and tourism (permit number mpb 8553.). results water quality, hydrology and meteorological data mean water temperature results in the incomati river system indicate that the minimum is reached in july (18.02 °c) after which temperatures gradually increase to a mean temperature of 24 °c during october. the highest mean monthly river water temperature during this study (30.61 °c) was recorded in the crocodile river during january (figure 3). the highest mean monthly river water temperature in the komati river (30.17 °c) was recorded during february. temperatures in the incomati river, below the confluence, were influenced by both tributaries, and consequently, the highest mean monthly temperature for the incomati river (28.88 °c) was recorded during february. figure 3: graph indicating water and air temperature for the months january–december 2003 in the incomati river. for the tigerfish active summer period, november to april, the mean monthly ph values varied between 8.1 and 8.5, whereas the conductivity fluctuated between 274 µs/cm and 622 µs/cm in the incomati river. summer conductivity values were lower than winter values, but summer ph values were higher. during summer, the turbidity levels increased as a result of the higher summer flows. although not measured, turbidity was observed to be closely associated with rainfall events in the catchment during the summer period. the highest rainfall recorded was during the months of november (115.4 mm) and february (191.7 mm). the mean monthly flow (figure 4) for the winter period (may–october) in the incomati river, when tigerfish are less active, varied between 0.44 m3/s and 1.89 m3/s compared to 0.82 m3/s and 13.12 m3/s for the summer period (november–april), when tigerfish are active. the highest flow spikes were recorded during the spawning season (october–february) in the summer period (steyn 1993; steyn & van vuren 1991; steyn et al. 1996). on three occasions, flow spikes in excess of 25.00 m3/s, with the largest of 51.76 m3/s, occurred in january (figure 4). figure 4: water flow in the incomati river over the period january–december 2003. radio-tagged fish in total, 41 fish were radio-tagged with a mean length (sl) of 62.7 cm (range 47 cm – 76 cm) and a mean weight of 2418 g (range 910 g – 4990 g) (table 1; figure 2). of the 41 radio-tagged fish, 11 (26.8%) were males and 30 (73.2%) were females in a 1:3 sex ratio. for the radio-tagged males, the length (sl) varied between 47 cm and 60 cm (mean = 55.4 cm) and the weight varied between 910 g and 2040 g (mean = 1605.5 g). for the radio-tagged females, the length (sl) ranged from 57 cm to 76 cm (mean = 65.4 cm) and the weight ranged from 1810 g to 4990 g (mean = 2717 g) (table 2). table 2: summary of average fish length, number of fixes, longest distance travelled and home range. movement the total distance of the river where adult fish were captured and equipped with radio tags measured 5.2 km. after capture, tagging and the associated disturbance to a fish when released, the fish normally moved upstream or downstream and normally only returned 2 to 5 days later to the original tagging site, thereby suggesting site fidelity. the distance moved directly after tagging varied over the 2to 5-day period from 48 m to 1038 m. in total, 35 (85.4%) of the fish tagged returned to the original tagging site within the mentioned period, but 6 (14.6%) never returned, 3 of which moved downstream into mozambique and were not recorded again. this showed angling in the form of catch and release may be a major disturbance, but this also confirmed site fidelity of tigerfish to a specific home range. the gps coordinates of each sample or release site, tag number, type of tag and size, weight and sex of each fish are presented in table 1. over time, a movement pattern emerged for each of the 41 radio-tagged fish, and the longest distances travelled and home ranges could be determined (table 1). on average, fish were tracked 72.5 times (table 2) and the total number of fixes was 2971 for the period 04 january 2003 to 22 december 2003. some individuals were tracked up to 161 times. the maximum total of fixes (n = 161) per individual was associated with a tag life of 10 months. for the summer period (january–april, november and december 2003), there were 1322 fixes, and for the winter period (may–october 2003), there were 1649 fixes. for the summer period (or part thereof), there were 40 active radio-tagged fish, but only 32 active radio-tagged fish for the winter period (or part thereof). the mean number of fixes for females was 78.4 (n = 30) per fish with a range of 6–161. the mean number of fixes for males was 56.2 (n = 11) per fish with a range of 7–110. the reason for the lower amount of fixes for males (56.2 fixes) in comparison with females (78.4 fixes) can be ascribed to the differences in radio tag types used. as males are generally smaller than females, smaller f2040 radio tags, with a much shorter lifespan (94 days), were used to stay within the 2% rule. longest distance travelled for the statistical analysis, data were obtained from 38 radio-tagged tigerfish with more than 10 fixes. the mean longest distance travelled (n = 38) was 729.66 m (table 2) with a range from 74.5 m to 3200 m. when analysing the longest distance travelled by the different radio-tagged fish, 47.4% (18 out of the 38 fish) travelled between 100 m and 500 m, 34.2% (13 fish) between 501 m and 1000 m, 10.5% (4 fish) between 1001 m and 1500 m, 5.3% (2 fish) between 1501 m and 2000 m and 2.6% (1 fish) travelled more than 2000 m (table 2; figure 5). figure 5: clustering of longest distance travelled of radio-tagged tigerfish for the period january–december 2003. a, 0 m–500 m; b, 501 m–1000 m; c, 1001 m–1500 m; d, 1501 m–2000 m; e, > 2000 m. when distinguishing between the different sexes and longest distance travelled, 46.4% of females (13 out of 28 fish) travelled between 100 m and 500 m, 39.3% (11 fish) between 501 m and 1000 m, 7.1% (2 fish) between 1001 m and 1500 m, 3.6% (1 fish) between 1501 m and 2000 m and 3.6% (1 fish) travelled more than 2000 m. for the males, 50% (5 out of 10 fish) travelled between 100 m and 500 m, 20% (2 fish) between 501 m and 1000 m, 20% (2 fish) between 1001 m and 1500 m and 10% (1 fish) between 1501 m and 2000 m (table 2). the furthest movement recorded was 3200 m over a 3-day period. this female moved out of its known home range (18 fixes) and established a new home range approximately 3018 m upstream (table 2). for females, the mean longest distance travelled was 734.4 m (n = 28) with a range of 74.5 m to 3200 m, and for males, the mean longest distance travelled was 716.3 m (n = 10) with a range of 148.9 m to 1800 m. no significant differences were found between males and females for longest distances travelled (mann–whitney u test, mean ranking males 19.8 and females 18.6, p = 0.753). three different tigerfish movement patterns were recorded (figure 6). movement patterns were obtained from a combined effect of distance travelled and home range sizes (figure 7). although all the fish displayed some degree of site fidelity within a specific activity zone, movement pattern 1 represents fish that moved 100 m to 500 m within a well-defined home range, and movement occurred only within this specific home range. movements of fish number 8 serves as example for this type of movement pattern (figure 8). the majority (47.37%) of the radio-tagged fish displayed characteristics of movement pattern 1 (figure 6, cluster a). movement pattern 2 represents fish that displayed site fidelity for two or more areas within a larger well-defined home range, spanning a distance of 501 m to 1500 m. movements of fish number 15 serve as example for this type of movement pattern (figure 9). this group was represented by 44.7% of radio-tagged fish (figure 6, cluster b). movement pattern 3 represents fish that showed little site fidelity and would temporarily occupy small areas within a large undefined home range that spans more than 1500 m. movements of fish number 23 serve as example for this type of movement (figure 10). fish within the latter group can be seen as vagrants without established home ranges for a specific period. most of these fish were also later lost as they moved out of the study area and could not be relocated. fish in this group were large females of weight ranging between 2720 g and 3580 g and represented 7.89% of the radio-tagged fish (figure 6, cluster c). figure 6: scatter graph depicting three different movement patterns by radio-tagged tigerfish related to home range size and longest distance travelled for the study period january–december 2003. figure 7: clustering of home range sizes of radio-tagged fish. a, 0 m2–10 000 m2; b, 10 001 m2–20 000 m2; c, 20 001 m2–50 000 m2; d, 50 001 m2–100 000 m2; e, > 100 000 m2. figure 8: map of radio-tagged fish number 8 indicating home range and longest distance travelled (type 1 movement pattern). figure 9: map of radio-tagged fish number 15 indicating home range and longest distance travelled (type 2 movement pattern). figure 10: map of radio-tagged fish number 23 indicating home range and longest distance travelled (type 3 movement pattern) for a detailed account of the movement patterns and demarcation of the home ranges of each of the 41 radio-tagged fish, see roux (2013). the dots indicate individual fixes during tracking and the contours around the fixes indicate the defined home range. home range sizes the home range size varied between individual fish with 38.2% (13 fish) localising within an area between 0 m2 and 10 000 m2 (mean = 5567.95 m2) and 18.42% (7 fish) localising within an area between 10 001 m2 and 20 000 m2 (mean = 14 435.53 m2). furthermore, 18.42% (7 fish) occupied a home range area between 20 001 m2 and 50 000 m2 (mean = 31 092.2 m2), whereas 10.5% (4 fish) occupied an area between 50 001 m2 and 100 000 m2 (mean = 79 809.55 m2) and 18.42% (7 fish) utilised an area > 100 000 m2 (mean = 163 692.90 m2) (table 2; figure 7). on average, the fish (n = 38) stayed within a defined home range of 48 846.36 m2. the home range size for males and females compared favourably with a mean of 53 296.52 m2 (n = 28) and a range from 331.6 m2 to 236 496 m2 for females and a mean of 36 385.9 m2 (n = 10) and a range from 1338.8 m2 to 135 982.6 m2 for males. no statistically significant differences were found between the sexes for their home range size (mann–whitney u test, mean ranking females = 20.71 and males = 16.10, p = 0.260). migration obstructions none of the 41 tagged fish crossed the tenbosch weir. three individuals, namely numbers 7, 12 and 18, moved upstream in the crocodile river to be briefly recorded in the vicinity of this weir. the tenbosch weir has a crest height of 2 m and a fish ladder constructed at the side of the weir. this ladder is of the vertical slot type and appears to be non-functional to fish migration in general. only two radio-tagged fish (fish numbers 15 and 39) ventured into the lower komati river, close to the confluence with the crocodile river, where they were confined in a pool below the low-water bridge for a few days. they were not able to overcome this obstacle. this low-water bridge at komatipoort was constructed on a natural dolerite intrusion that stretches across the river. contrary to the above, a total number of 16 crossings, both upstream and downstream, were recorded at the knp weir. this gauging weir has a crest height of approximately 1.2 m with a well-designed fish way to facilitate fish movement at medium to high flow conditions. tagged fish with allocated numbers 1, 4, 6, 20, 27 and 37 crossed the knp weir downstream and upstream over the period january to march, whereas fish 19 crossed the knp weir downstream during july and returned upstream three days later. fish numbers 1 and 27 each crossed on three occasions, whereas fish number 20 crossed the knp weir on four occasions with only a few day intervals between upstream and downstream crossings. numerous visual observations were made of untagged tigerfish jumping over this weir over the duration of this study. successful crossing at the knp weir occurred at flow velocities between 1.78 m³/s and 16.2 m³/s (table 3). table 3: successful crossings of radio-tagged tigerfish at the kruger national park weir. discussion this study confirmed that external tagging attachment protocol (thorstad et al. 2001) was suitable for the study of tigerfish behavioural ecology through biotelemetry in that only a single mortality was recorded from the 41 radio-tagged fish. furthermore, visual observations of radio-tagged fish swimming just below the surface were made on numerous occasions and fouling of radio tags appeared to be minimal, thus having no significant effect on the swimming capabilities or movement patterns of tagged fish. after capture, tagging and the associated disturbance to a fish, it normally moved either upstream or downstream and returned 2–5 days later to the original tagging site, thereby confirming site fidelity. the distance moved directly after tagging varied over the 2to 5-day period from 48 m to 1038 m. in total, 35 of the tagged fish returned to the original tagging site within the mentioned time frame. six fish never returned; three of these moved downstream into mozambique and were lost. in general, tigerfish displayed high site fidelity to specific habitats within specific activity zones, and movement occurred primarily within these defined home ranges. the longest distance travelled by fish was during summer and early winter, when water temperatures exceeded 24 °c. these periods coincided with high water levels in the study area, which probably facilitated movement between different habitats. some degree of site fidelity of h. vittatus was also reported by økland et al. (2005) for the upper zambezi, whereas consistent fidelity to an activity core was reported by baras et al. (2002) for hydrocynus brevis in the niger river, mali. during our study, little to no movement was recorded in the winter months when water temperatures were below 24 °c. the mean lowest temperature recorded in the incomati river system of 18 °c is close to the minimum temperature range for the survival of tigerfish. during a tigerfish translocation exercise when laboratory-induced breeding was attempted, prior to successful breeding at skukuza (steyn et al. 1996), a temperature drop from 27 °c to 18 °c during a 4-hour transport period killed almost all of the fish. the mean longest distance travelled during this investigation was relatively short (729.66 m). in the zambezi, two movement patterns were distinguished where approximately 50% of the fish moved < 1000 m among tracking surveys. the remaining fish showed consistent site fidelity for periods with long-distance movements (> 1000 m) to new areas among residency periods. in the incomati river system, only 18% of the fish displayed long-distance movement > 1000 m and the longest distance was 3200 m. the longest distance travelled in the incomati river system was relatively short in comparison with the longest distance of 18.8 km travelled in the zambezi river (økland et al. 2005). irrespective of the shorter distances travelled in the incomati river system, the total unobstructed river upstream to tenbosch was not utilised by all tagged individuals and the option to migrate downstream was available but not utilised. nevertheless, three movement patterns demonstrated by incomati river system tigerfish do not describe the dependency on upstream or downstream migration behaviour expected for this species in the study area. implicit of the relative short distances travelled, they are crucial for the survival of h. vittatus in the lower incomati river system. site fidelity and restricted mean home range (48 846 m2) in comparison with the much larger home range of zambezi tigerfish (276 978 m2), supported by various historical observations of their vulnerability to environmental stressors such as low temperature, low flow and high silt loads, are indicative of a population that does not function optimally on the edge of its distribution, in accordance with the law of tolerance (odum 1971). sub-optimal functionality of another tigerfish population in the knp is also reflected in the results of gagiano (1997), during an ecological investigation on tigerfish in the olifants and letaba rivers. tigerfish of all sizes in these rivers were found to feed almost exclusively on invertebrates. this finding is in contrast with the tigerfish from other systems, where fish play a major role in their diet (jackson 1961). differences in movement patterns, longest distance travelled and home range size could not be explained by sex or the size of the fish. tigerfish show opportunistic movement patterns, and home ranges can change in size and location as a result of seasonal shifts, prey availability, habitat availability and cover as well as life history requirements. no large-scale movement pattern or specific activity-related migrations were observed. thus, reports of large-scale migrations of tigerfish downstream into mozambique during winter in the incomati river (van loggerenberg 1982) seem to be no longer relevant, probably because of their limited numbers and because of suitable habitat created by the damming of the knp weir and subsequent deeper water bodies where the temperature is more stable to find refuge during winter. there was no evidence of upstream congregation of tigerfish at the tenbosch weir or large-scale downstream crossings at the knp weir. from the pattern of crossings at the knp weir, it is inferred that some of the marked fish that successfully crossed this weir responded to the stress associated with the tagging procedure and returned later to demonstrate site fidelity. these fish were tagged either just downstream or upstream of the knp weir, followed by a fleeing response over the weir (fish numbers 1, 6, 19 and 20). some of the crossings could be associated with higher flow conditions (fish numbers 4 and 37), whereas fish number 27 probably displayed natural behaviour as the crossing occurred more than a month after tagging. irrespective of the motivation for crossing the weir, in context with the life span of the tags for above fish, these events were limited to only a few occasions during a period of several months, which again displayed site fidelity. flow volumes that varied between 1.94 m³/s and 16.22 m³/s during successful crossings suggest that the knp weir is not a restrictive barrier to tigerfish and the population is open to gene flow from mozambique. contrary to this, our results suggest that tigerfish in the crocodile river, upstream from the tenbosch weir, is isolated; consequently, the upstream population cannot be replenished after mortalities because of extreme environmental conditions such as influx of cold water, low flow and high silt loads and will most probably disappear in this part of its historical distribution range. in the komati river, upstream movement is restricted close to the confluence at the low-water bridge, consequently isolating the upstream population in the komati river which currently is heavily subjected to water abstraction and agricultural activities. the isolation of upstream tigerfish populations in the incomati river system and their vulnerability to environmental impacts emphasise the ecological significance and inclusion of this river reach into the borders of the knp as well as the functionality and importance of the knp weir. based on the knowledge gained during this study on the behaviour of tigerfish, recommendations on the instream flow requirements (ifrs) of this species need to be adopted into the ecological flow requirements for the incomati river system and setting of the ecological reserve to ensure the ecological maintenance and functioning of the instream habitats utilised by tigerfish (kleynhans & engelbrecht 2000). environmental flow allocations and maintenance of ecological requirements of aquatic ecosystems are entrenched in the national water act (no 36 of 1989) and specified as components of the ecological reserve. within the framework of resource directed measures for protection of water resources, established by the department water affairs and sanitation (dwa), the implemented ecological reserve needs to be monitored and can be adjusted to meet the targets and resource quality objectives (king, tharme & de villiers 2000). acknowledgements the authors acknowledge de beers, venetia mines and barloworld for funding this project. they thank the incomati tigerfish action group (itag), domien and bart van buynder for their logistical support and for their enthusiasm for the tigerfish species and its conservation. they also thank peter kimberg and michael mashaba for their assistance with fieldwork and data collection. 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https://doi.org/10.1111/j.1095-8649.2001.tb00174.x van loggerenberg, n.p., 1980, kunsmatige teelt van die tiervis hydrocynus vittatus castelnau, en die biologiese aspekte wat daarmee verband hou, projek tn 6/4/2/2/1/18, transvaal provinsiale vissery instituut, lydenburg. van loggerenberg, n.p., 1982, kunsmatige teelt van die tiervis hydrocynus vittatus castelnau, in transvaal, vierde projekverslag, projek tn 6/4/2/2/1/18, transvaal provinsiale vissery instituut, lydenburg. van loggerenberg, n.p., 1983, ‘conservation of tigerfish and fish farming techniques’, fauna and flora 40, 30–31. wilber, c.g., 1983, turbidity in the aquatic environment. an environmental factor in fresh and oceanic waters, cc thomas, springfield, illinois, usa. worton, b.j., 1989, ‘kernel methods for estimating the utilisation distribution in home range studies’, ecology 70, 164–168. https://doi.org/10.2307/1938423 article information authors: francesca cini1 melville saayman1 affiliations: 1tourism research in economic environs and society, north-west university, potchefstroom campus, south africa correspondence to: francesca cini postal address: private bag x6001, potchefstroom 2520, south africa dates: received: 26 mar. 2013 accepted: 03 dec. 2013 published: 24 june 2014 how to cite this article: cini, f. & saayman, m., 2014, ‘which age group spends the most in a national park?’, koedoe 56(2), art. #1158, 8 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1158 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. which age group spends the most in a national park? in this original research... open access • abstract • introduction    • literature review • research method and design    • setting    • design    • procedure       • socio-demographics       • expenditure patterns       • reasons for visiting the park    • analysis • results    • respondents' profile    • reasons for park visit    • expenditure patterns by age    • segment characterisation • ethical considerations • trustworthiness • discussion    • practical implications    • recommendations    • limitations • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ age (and its changing structure amongst the wider population) is one of the most relevant aspects required to better understand and forecast the needs, interests and associated consumption behaviours of tourists. this research used age to investigate the expenditure patterns amongst a sample of visitors to the table mountain national park (tmnp), south africa. in march 2010, visitors to the tmnp were found to differ significantly from those at other parks, as they were younger and most of them were foreigners. this study found that younger visitors (18–29 years) were higher spenders when compared to those aged 30–49 years. as parks are generally visited by older people, this study showed the economic importance of the younger market. the research also made clear implications and recommendations for park management as to how to address these findings.conservation implications: conservation is dependent on funding. one of the main sources of income is tourism and tourism related activities. this research can assist marketers and managers to target the right markets in order to be more sustainable. this research also shows the importance of environmental education at an early age in order to grow awareness and to target the right markets. introduction top ↑ the economic importance of natural protected areas is recognised at both national and international levels. since the 2003 world parks congress in durban, south africa, the view of economic development and nature conservation as opposite realities is changing. in this way, a new paradigm for protected areas was fostered: the synergy between conservation and sustainable development was forged. protected areas were seen as providers of benefits beyond boundaries – beyond their boundaries of nation-states (international union for conservation of nature [iucn] 2004b). the relationship between economic development and nature conservation became increasingly important, especially in developing countries, because conservation areas can be used as a tool for poverty alleviation (buckley 1994, 2002a; bushell & mccool 2007; bushell, staiff & eagles 2007; butler & boyd 2000; geldenhuys & saayman 2009; myburgh & saayman 1999; world commission on protected areas 1998; world tourism organization 2002; world wildlife fund 2001). practical, protected areas can provide a number of benefits to the poor. for instance, they can provide some ecosystem services including coastal protection, water purification and carbon sequestration. they also can provide some options for income by providing jobs in the tourism industry. for instance, in countries such as south africa, zimbabwe and zambia, local communities obtain income from sport hunting (child & dalal-clayton 2004; iucn 2004a). in many countries, government funding for parks is reducing in real terms (buckley 2002b; eagles, mccool & haynes 2002; mabunda & wilson 2009; saayman 2009); therefore, it is becoming increasingly important to attract markets that can generate capital that can be used for sustainable development. then, particularly for third-world countries, finance generated and secured from tourism can represent the basis for nature conservation and preservation, as well as for the eradication of economic and social inequalities. in addition, it can provide benefits such as an increase in local jobs, higher local income, higher education level of local people and equal access to quality and affordable basic services (eagles et al. 2002). as promoting the mere increase in the number of visitors to capture economic benefits might be somewhat risky in terms of the negative impacts on the natural environment (see eagles et al. 2002), a possible de-marketing strategy might be developed for targeting only those segments with the stronger propensity to spend. literature review the characteristics of consumers as demographic, behavioural (including the expenditure patterns) and psychographic are widely considered the key elements of a marketing strategy. identifying the factors linked to tourists’ motivations, preferences and behaviours is crucial to foreseeing their potential travel choices (cha, mccleary & uysal 1995; frochot & morrison 2000; kotler 1992). specifically, age (and its changing structure amongst the wider population) is one of the most relevant aspects that is considered to better understand and forecast the needs, interests and associated consumption behaviours of tourists (gibson, attle & yiannakis 1998; johns & gyimóthy 2002; jönsson & devonish 2008; moutinho 1987; oh et. al. 2004; patterson & pegg 2009; seaton 1996). in western countries, the size of the older population (55 years and older) is growing at a faster rate than the younger one and represents new target tourist markets that are healthier, affluent and have a greater desire for novelty and escape than do those of previous generations (patterson & pegg 2009; sherman & schiffman 1991). the use of age as a segmentation variable is supported by several research articles that found age to be one of the primary variables, which explained the differences in people’s travel interests preferences and behaviours. to date, several studies found age as one of the main variables in affecting even travel expenditure patterns as a type of travel behaviours (thrane & farstad 2011). for example, in a study conducted by mok and iverson (2000), age significantly characterised the different spending segments. specifically, heavy spenders were younger than 50 years old. again, jang, ismail and ham (2002) found that association amongst a sample of japanese outbound travellers. specifically, this study revealed that heavy spenders were older, had a larger party size and a longer length of stay. in addition, saayman, saayman and du plessis (2005) found differences between heavy and light spenders according to their age amongst visitors to the world cup cricket matches in south africa. specifically, heavy spenders were older than 35 years old and spent more on accommodation, restaurants and souvenirs, compared to the younger ones. in most studies, profiling the spending segments was conducted using socio-demographics such as age, level of education, annual income, place of residence and trip characteristics such as travel party size, length of stay and type of accommodation. with specific regard to the relationship between travellers’ age and their use of natural areas, for instance, a study conducted by kim, lee and klenosky (2003) amongst visitors to a national park found out that the younger group (less than 50 years) was motivated mainly by the possibility of experiencing adventure and sharing their friendship, whilst the older visitors (50 years and older) by the possibility of appreciating the natural resources and enhancing their state of health. also moscardo and green (1999) found an overall decline in activity participation associated with age, particularly for activities associated with nightlife and entertainment and outdoor, physical, nature-based activities. analysing travellers by age can also have important implications for the management of a tourist destination. types of facilities and access to infrastructures, for example, might be planned on the basis of visitors’ age. strongly associated with people’s income and expenditure patterns (down 2000), age was found as one of the main variables affecting even travel expenditure patterns (jang et al. 2004; saayman et al. 2005). however, a review of the literature revealed a dearth of studies aimed at investigating the expenditure patterns (behavioural characteristics) amongst nature-based tourists, or even of expenditure behaviours by their age. amongst the few, a study conducted by mehmetoglu (2007) investigated the relationship between the daily expenditures and the trip activities of a sample of visitors at two nature-based attractions in northern norway. heavy spenders were more likely to consider nature-based activities when compared to the light spenders who, instead, attached more importance to visiting museums and attractions. whilst motivational differences amongst the spending groups were partially supported by the study, income and age were found to significantly influence their expenditure patterns. specifically, older nature-based tourists (50 years and older) were more likely to be lighter spenders than were younger ones. another study looked at the variables that influence spending patterns and made recommendations to attract high-spending markets to the tsitsikamma national park in south africa (kruger, saayman & saayman 2010). respondents segmented by their real expenditure per person per day showed differences based on their age. two different markets were identified, (1) ‘high-spenders’, aged 35–49 years and older than 50 years, more likely to stay in chalet accommodations and to have a longer stay at the park and (2) ‘low-spenders’, aged 35–49 years old, more likely to stay in camping accommodations and have a shorter stay at the park. however, some other studies of the field did not include age as a segmentation variable (downward & lumsdon 2004) or did not detect any significant relationship between expenditure patterns concerning age (chhabra 2007; spotts & mahoney 1991). since age has been found as one of the most relevant aspects that is considered to better understand and forecast the needs, interests and associated consumption behaviours of tourists, as mentioned above, further research is required on this aspect to provide more precise information for marketers. comparing groups of visitors by age with regard to their expenditure patterns will possibly allow both known and new age lucrative markets to national parks to be explored. this will be useful when proposing strategic plans to grow visitation figures amongst those nature-based segments with the highest propensity to spend. this is particularly important for less developed countries, where a large portion of the world’s biodiversity is concentrated and where the finance generated and secured from nature-based tourism can foster and support the sustainable development of an area. based on this, the purpose of this article is to use age to investigate the expenditure patterns amongst a sample of visitors to the table mountain national park (tmnp), along with other socio-demographics and their reasons for the visit. the latter data are to assist with profiling the age groups and to match the ‘product‘s’ specificities with the public’s specific demands. research method and design top ↑ setting the tmnp is one of 22 national parks in south africa and was officially established in 1988. it stretches from signal hill in cape town to cape point at the southern end of the cape peninsula and covers 25 000 ha. this urban park is famous because of table mountain itself, its historical and cultural heritage and its world heritage status. south african national parks (sanparks), which is the custodian of national parks in south africa, indicated the promotion of the country’s nature-based tourism as one of its core pillars. without compromising sustainability, self-generated revenues from commercial operations are considered necessary to supplement government funding of conservation management. the tmnp is amongst the five top south african parks with the highest number of guests. statistics for 2011 reported 2 344 340 guests for the tmnp, followed by the kruger national park with 1 411 796 guests, the west coast national park with 201 137 guests, the tsitsikamma national park with 180 107 and the addo national park with 138 079 guests. compared to the previous year (2010), total guests-to-parks numbers increased by 3.6%. specifically, the tmnp increased by 5.6%, whilst the kruger national park increased by 1.8% (sanparks 2012). the main reason for selecting the tmnp was that this park attracts a large number of visitors and shows the highest representation of international tourists, as well as a more equal spread of respondents over different age groups, compared to other south african national parks (saayman, kruger & fouche 2009). this can be explained by the facts that tmnp is a scenic park, offering the ‘majestic mountain’ as one of the new seven wonders, together with ocean views, mountains, beaches, forests and a variety of popular recreational activities. design the study sample included day visitors to the park from 27 march to 01 april 2010. six fieldworkers (three groups of two) approached visitors whilst they were queuing at the three popular entry points to the park: the cable car to table mountain, boulders and cape point. it is difficult to have fieldworkers at all entry points as there are also open access areas; therefore, it was decided to focus on these three, as they could be considered the most important access points and tourists generally visit all these areas.those visitors who expressed willingness to take part in the research were first asked a number of screening questions to detect the expenditure patterns from only one member per family or travel group. this information avoided repeat data and biases in the actual expenditure patterns amongst the respondents. in total, 441 questionnaires were collected during the study period, of which 404 were usable after a preliminary screening. according to israel (2009), in a population of > 100 000 (n), 398 respondents (n) are seen as representative and result in a 95% level of confidence with a ± 5% sampling error. thus, 404 usable questionnaires were deemed sufficient for the purposes of this study. procedure the research used a structured self-administered questionnaire, developed on the basis of the questionnaires used by past researchers (oberholzer et al. 2009; saayman, saayman & ferreira 2009). the questionnaire consisted of three main sections, (1) socio-demographic details, (2) expenditure patterns and (3) reasons for the visit. socio-demographics taking into consideration the goal of this study, age was considered pre-eminent compared to the other socio-demographic characteristics gathered here and this factor was detected by an open-ended question: ‘year of birth’. the respondents’ ages were calculated and post-coded in groups on the basis of the strong association found between age and household finance (income and expenditure) (down 2000; mok & iverson 2000). only respondents over 18 years of age were surveyed, as they are more likely to travel by themselves and have money to spend. respondents were categorised into one of the following four age groups: 18–29 years, 30–49 years, 50–64 years and 65–74 years (see table 1). table 1: location of respondents by age group in table mountain national park. the age groups were characterised based on the expenditure patterns per day per person, together with a trip-related characteristic, reasons for the visit and other socio-demographics. two other socio-demographic details were also included in the questionnaire to assist with profiling the age groups: ‘region of residence’ and ‘education’ (‘no school’, ‘matric’, ‘degree’, ‘postgraduate’, ‘professional’ and ‘other’). the region of residence was post-coded as follows: ‘north america’, ‘australasia’, ‘europe’, ‘south africa’ and ‘other countries’. finally, a question about the awareness of being in a national park was included. a dichotomous response format (‘yes’ or ‘no’) was used. expenditure patterns the expenditure patterns section was developed based on past research (kruger et al. 2010; saayman, van der merwe & pienaar 2009). spending behaviours were captured as follows: ‘money spent for categories or services per person’ (‘entrance and conservation fee’, ‘restaurants’, ‘food’, ‘beverages’, ‘transport to the park’, ‘activities’, ‘souvenirs and jewellery’) and ‘number of people paid for’ (including the respondent). on the basis of responses to ‘money spent for categories or services’, the total expenditure was calculated. to compute the total expenditure per person, the total expenditure was divided by the number of people paid for. reasons for visiting the park one measure that was developed ad hoc for this study concerned the reasons for visiting the park and the age groups were also characterised according to this. these reasons were found to be connected to age and can be considered amongst the most important personal variables for a better understanding of the consumers’ profile (see crompton 1979; crompton & mckay 1997; iso-ahola 1999). the scale for this question was developed based on common motivations identified from the tourism literature (crompton 1979; kim, jogaratnam & noh 2006; kruger & saayman in review; richards & wilson 2003), which include, (1) relaxation and getaway, (2) knowledge, (3) family togetherness, (4) natural beauty admiration and (5) activities participation. the scale included 26 items (four to five for each motivation) and participants were asked to indicate the importance of each item using a 5-point likert scale (1 = ‘not at all important’, 2 = ‘less important’, 3 = ‘important’, 4 = ‘very important’, 5 = ‘extremely important’). the relaxation and getaway motivation included items such as, ‘to get away from routine’ and ‘to relax’. the knowledge motivation items included, ‘learn about plants and animals’ and ‘learn more about specific marine life’. the family togetherness motivation included statements such as, ‘family recreation’ and ‘benefit of children’. the natural beauty admiration motivation included items such as, ‘tmnp is world renowned’ and ‘for the scenic beauty and view’. finally, the activities participation included statements such as, ‘mountain biking’ and ‘surfing’. analysis statistical data analysis was performed using spss version 18 (2009). the four age groups were characterised using chi-square, kruskal-wallis and analysis of variance (anova) tests based on expenditure patterns, along with psychographics (reasons for visiting the park) and socio-demographics. results top ↑ respondents’ profile in the tmnp, the largest group of respondents (196, 48.5%) were 30–49 years old. this was followed by those aged 18–29 years (130, 32.2%). fewer in number were respondents aged 65–74 years (49, 12.1%) and those aged 50–64 years (29, 7.2%). during the 5-day study period, the average age of respondents was 41.3 years (s.d. = 15.69).the median of the total expenditure per respondent per day person was r125 and the majority of respondents (71.2%) said that they had paid for one or two people, including themselves. this group was followed by those respondents who paid for three or four people (22.7%). the majority of respondents were well-educated, with 37.6% holding a degree, 22.6% in professional occupations and 18.3% holding a postgraduate qualification. this result, too, is consistent with previous research (e.g. marques, reis & menezes 2010; saayman & slabbert 2004; saayman, kruger & fouche 2009; the international ecotourism society 2006). from this study, it was found that the tmnp is attracting a higher percentage of international tourists compared to local tourists. almost half (46.1%) of the visitors to the tmnp were foreigners from europe. south african tourists made up almost one-third of all respondents (28.9%), whilst north americans represented one-tenth (10.7%). in contrast, previous research at other south african national parks (saayman, kruger & fouche 2009) highlighted a higher percentage of local visitors compared to international ones. reasons for park visit a principal component analysis with oblimin rotation supported a five-factor structure, explaining 71% of the total variance: • ‘activities and adventure sports performance’ • ‘park’s attributes admiration’ • ‘knowledge and inner experience’ • ‘escape’ • ‘social interactions’. the scree plot of eigenvalues suggested that no further factors contained reliable systematic variance. satisfactory reliabilities were observed for the scale scores (0.95, 0.68, 0.84 and 0.79) with the exception of social interactions (0.51). the results are shown in table 2. table 2: principal component analysis on respondents’ motives for visiting table mountain national park. factor scores, calculated as the average of all items contributing to a specific factor, indicated the most important reasons for visiting the park were, ‘escape’ (3.49), followed by ‘park’s attributes admiration’ (3.46). less important reasons were ‘social interactions’ (3.28), ‘knowledge and inner experience’ (3.12) and ‘activities and adventure sports performance’ (2.26). these results are similar to those of studies at other south african national parks (saayman, kruger & fouche 2009), which, in most cases, found the main reason for visiting national parks was the need to escape and relax. expenditure patterns by age the kruskal-wallis test was used for continuous variables that displayed a non-normal distribution. the respondents aged 18–29 years reported the highest expenditure per person and a higher expenditure per person per day when compared to the older group aged 30–49 years. specifically, the median expenditure for the group aged 18–29 years is r160 and the median for the group aged 30–49 years is r120. the former group was above, whilst the latter was slightly below the overall median, which was r125. no significant statistical differences could be established between the groups aged 18–29 years and 30–49 years and the groups aged 50–64 years and 65–74 years, although the median of the expenditures per person was noticeably less than for the groups younger than 50 years. this is because, on the one hand, the older groups (50–64 years and 65–74 years) were fewer in numbers (29 and 49 respondents, respectively) whilst, on the other hand, few of them provided information about their expenditure patterns (10 and 15 respondents, respectively) (see table 3). table 3: expenditures per person by age group for respondents at table mountain national park. most of the travel expenditures were allocated to entrance and conservation fees (48%), whilst smaller percentages characterised restaurants (16%), transport (9%), food (8%) and activities (8%). however, no significant differences were found amongst the age groups in expenditures per person for each of these categories. segment characterisation an anova analysis on the five psychographic characteristics found significant differences for the motivational dimension ‘social interactions’ by age groups (f = 3.18, df = 3/356, p < 0.05). respondents aged 18–29 years or 30–49 years were more likely to be motivated for the visit by the possibility of being with family (or someone special) or with friends, when compared to the group aged 65–74 years. however, these results should be interpreted with scepticism because of the low reliability of the scale (see table 4). table 4: analysis of variance of reasons for the visit to table mountain national park by age group. statistically significant differences were also found for the awareness of being in a national park [χ2 (3=397) = 8.12, p < 0.05]. interestingly, although the majority of visitors amongst all age groups were aware, a higher number of younger respondents aged 18–29 years (21 out of 44) and those aged 30–49 years (20 out of 44) stated they were unaware that they were in a national park, compared to the older groups.statistically significant differences were also found in the region of residence [χ2 (9=353) = 30.1, p < 0.01]. in particular, more respondents aged 30–49 years (45.3% and 31.8%, respectively) and 18–29 years (31.8% and 32.5%, respectively) came from europe and south africa, compared to the older groups. older respondents aged 50–64 years (46.2% and 26.9%, respectively) and 65–74 years (44.2% and 30.2%, respectively) came from europe and north america (see table 5). table 5: region of residence of respondents in table mountain national park by age group. the education level of three groups also differed [χ2 (12=385) = 47.12, p < 0.01]. whilst most of the age groups held a diploma or degree, more respondents aged 65–74 years (32.6%) and 30–49 years (24.7%) were professionals, compared to those aged 18–29 years (12%) and 50–64 years (18.5%). again, more respondents aged 30–49 years (22.1%), followed by those aged 18–29 years (17.6%) and 50–64 years (14.8%) held a postgraduate qualification (see table 6). table 6: education of respondents in table mountain national park by age group. ethical considerations top ↑ respondents participated of their own free will and had the opportunity to decline participation. no information was obtained that could personally identify respondents. trustworthiness top ↑ the authors did everything in their power to ensure that the sample and the survey was conducted in a systematic and scientific manner. results of the profile confirm previous research conducted in national parks. discussion top ↑ unlike previous research, this study specifically targeted visitors to a national park on the basis of their travel expenditure patterns by age range, along with other socio-demographics and their reasons for the visit. the latter assisted with profiling the age groups and to match the ‘product’s’ specificities with the public’s specific demands. from a theoretical standpoint, the findings differ from those detected by previous research. then, too, some contradictions representing rich material to foster new research directions have been highlighted as follows. firstly, as expected by the researchers, a considerable number of younger respondents (18–29 years) (32.2%) were detected in the area and this differs from the general notion and results by other researchers that mostly older people visit national parks. the low percentages of respondents older than 50 years (50–64 years, 7.2% and 65–74 years, 12.1%) were therefore unexpected, compared to previous findings (beh & bruyere 2007; galloway 2002; kibicho 2006; marques et al. 2010; kruger et al. 2010; saayman, kruger & fouche 2009; the international ecotourism society 2006). secondly, the finding related to spending behaviour highlights the presence of a new and potentially lucrative market when compared to those detected by previous research (kruger et al. 2010; mehmetoglu 2007); that is, a younger segment of respondents aged 18–29 years. this group was characterised by the highest expenditure per person and a higher expenditure per person when compared to those aged 30–49 years. then, younger respondents seem to represent a rather lucrative market, which would be a crucial variable for the maintenance and the sustainable growth of the area and is in line with the worldwide trends of youth tourism. the youth tourist sector, which accounts for 20% of the international tourism market, is growing faster than most other travel segments, with a global volume growth of 3% – 5% a year and is valued at approximately $165 billion per year. estimates show that youth and student travel market will reach 300 million arrivals by 2020 and represent $320 billion in market values (world youth student and educational travel confederation 2011). furthermore, the reasons for visiting a national park showed some differences in terms of age groups. specifically, significant differences were found for the motivational dimension ‘social interactions’: respondents aged 18–29 years and those aged 30–49 years revealed themselves to be more likely to be motivated by a possibility to spend time with family or friends, compared to the oldest group aged 65–74 years. unlike past research (e.g. moscardo & green 1999) that found differences concerning the activities participated in by respondents’ age, no statistically significant differences were found here in regard to the motivational dimension ‘activities and adventure sports performance’ by age group. in addition, unlike past research (kim, lee & klenosky 2003), age groups reported similarities for most of the motivational factors detected. finally, it is worth noting that amongst those respondents who were not aware of being in a national park, the majority were younger than 50 years old (i.e. in the 18–29 years and 30–49 years age groups). in brief, visitors participating in the research and aged 18–29 years were the second largest group and the heaviest spending segment per person. they were more motivated to visit for the day, when compared to respondents aged 65–74 years, because of the possibility of spending time with family (or someone special) and friends, they were less aware of being in a national park when compared to respondents aged 65–74 years, they mainly held a postgraduate qualification and largely came from europe or south africa. those aged 30–49 were the largest group but the lowest spending segment per person; the rest of their profile is similar to that of the youngest group. no statistically significant results were detected for the older groups concerning their expenditure patterns. practical implications based on the findings of this research, some practical implications for the sustainable development and financing of the tmnp have been outlined. the promotion of a mere increase in the number of visitors to capture economic benefits is to be avoided because it may have a cost to the natural environment (see eagles et al. 2002). therefore, a de-marketing strategy is considered necessary, but directed at the segment that showed the strongest propensity to spend (aged 18–29 years) and whose growth will help sustain the park. however, because the younger market at national parks is generally less present and known compared to the older segments, marketing to the younger groups should be conducted by first investigating the impacts of their activities. this relates also to park visitor management capacity. by contrast, no marketing strategy should be considered for the next oldest age respondents (aged 30–49 years) who were found to be lower-spenders and whose numbers are already significant and stable in this particular national park, as well as in worldwide national parks (e.g. saayman, kruger & fouche 2009). recommendations the following are some ideas for a specific marketing strategy. increasing the frequency of visits and attracting the younger market aged 18–29 years could be achieved by offering, for instance, a youth loyalty card, especially to those visitors residing in south africa, of whom there are considerable numbers amongst the younger respondents. hosting youth days might be another successful strategy to attract this market. in addition, a marketing strategy could include social-related content, as these respondents were found to be more likely to be motivated by the possibility of spending time with family and friends. for example, picnic and braai areas, such as the buffels bay and bordjiesdrif, or the tokai picnic and braai area and its wide space for large groups, can be promoted. as an urban park offering several activities and adventure sports, a marketing strategy could be used also to target additional younger segments such as the ‘harder eco-tourists’ identified by weaver and lawton (2002) or the ‘self-centred visitors’ and the ‘occasional visitors’ identified by marques et al. (2010), who reported a high propensity to take part in these types of activities. finally, as mostly younger participants seemed to be unaware they were in a national park, the management should provide more information about this. educational programmes could represent another marketing tool for promoting repeated visits to the park and, in general, to other national parks. in fact, several research findings have highlighted how practical experiences in outdoor settings, such as hands-on conservation activities and on-site educational programmes at an early age, shape and promote positive interests and associated behaviours towards the environment, thus ensuring nature conservation and preservation (mcduff & jacobson 2000; palmer 1993; sward 1996; voordouw 1987). limitations a limitation of the current research is the difficulty in generalising these results to a wider population of the park. to test the general applicability of these findings to other samples, this study should be replicated. secondly, previous research highlighted the importance of trip-related characteristics (e.g. type of accommodation, length of stay) for the spending segments characterisation. however, these variables could not be included in the present research because the tmnp, compared to other south african national parks, has a high percentage of day visitors and low percentage of repeat day visitors and overnight visitors (saayman, kruger & fouche 2009). this may be due, in part, to the fact that the majority of the visitors across all ages were foreigners from overseas visiting the tmnp as one of the stops on their tour. therefore, the tmnp management should do more, in terms of marketing plans, to target the domestic visitor market, for example, by providing discounts and/or loyalty cards to south africans. thirdly, unlike most past research that found motivational differences by age, here the characterisation of the age groups may be considered as limited because just one factor of five motivational factors significantly differentiated the age groups. future research is required to shed more light on this point. fourthly, although past research highlighted the importance of the activities participated in, in terms of increasing expenditure patterns (mehmetoglu 2007), this study was unable to find any differences in terms of the activities as reasons for the visit by the age (spending) segments. further research should again be conducted to shed more light on this point. finally, this study was unable to find any differences amongst the age groups in terms of expenditures per person for each of the categories considered (e.g. restaurants, food, etc.). however, it would be interesting to know what younger groups spent the extra money on. future research should be conducted to shed more light on this point. conclusion top ↑ this research adds to existing literature by shedding more light on the expenditure patterns (behaviour) of a sample of visitors to the tmnp, grouped by age range, and contributes to the topic of visitor behaviours to nature-based attractions. the respondents’ profile was found to differ significantly from the profiles of visitors to other parks, as the tmnp respondents are younger and most of them are foreigners. as parks are generally visited by older people, this study highlights the importance of a younger market. future research in national parks should pay particular attention to this market, which was found as an emerging but very relevant economic market and is anticipated as playing a key role in supporting sustainable lifestyles in the future (coetzee & saayman 2009; council for game and wildlife conservation 2003). acknowledgements top ↑ the authors want to acknowledge the following people and institutions. firstly, sanparks for funding and allowing the research, especially, mr glenn phillips and bheki zwane. secondly, the national research foundation for funding this research. thirdly, all the respondents for completing the questionnaires and, lastly, the reviewers for their valuable comments. competing interests the authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article. authors’ contributions m.s. 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ecotourist market segmentation in the gold coast hinterland of australia’, journal of travel research 40, 270–280. world commission on protected areas, 1998, economic values of protected areas: guidelines for protected area managers, joint publication by task force on economic benefits of protected areas of the world commission on protected areas and economics service unit of international union for the conservation of nature, gland. world tourism organization, 2002, sustainable development of ecotourism: a compilation of good practices, wto, madrid. world wildlife fund international, 2001, guidelines for community-based development, wwfi, ledbury. world youth student and educational travel confederation, 2011, the power of youth travel, viewed 21 february 2014, from http://wysetc.org/publications/power-of-youth-travel/ article information authors: robert buitenwerf1 andrew kulmatiski2 steven i. higgins3,4 affiliations: 1institut für physische geographie, goethe universität frankfurt, germany 2department of plants, soils and climate and the ecology center, utah state university, united states 3department of botany, university of otago, new zealand 4biodiversity and climate research centre, senckenberg gesellschaft für naturforschung, germany correspondence to: robert buitenwerf postal address: altenhöferallee 1, 60438 frankfurt am main, germany dates: received: 19 mar. 2014 accepted: 04 aug. 2014 published: 10 nov. 2014 how to cite this article: buitenwerf, r., kulmatiski, a. & higgins, s.i., 2014, ‘soil water retention curves for the major soil types of the kruger national park’, koedoe 56(1), art. #1228, 9 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1228 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. soil water retention curves for the major soil types of the kruger national park in this short communication... open access • abstract • introduction • methods    • study site    • approach    • water retention curves    • soil texture • results • discussion • acknowledgements    • competing interests    • authors’ contributions • references • appendix 1 • appendix 2 abstract top ↑ soil water potential is crucial to plant transpiration and thus to carbon cycling and biosphere–atmosphere interactions, yet it is difficult to measure in the field. volumetric and gravimetric water contents are easy and cheap to measure in the field, but can be a poor proxy of plant-available water. soil water content can be transformed to water potential using soil moisture retention curves. we provide empirically derived soil moisture retention curves for seven soil types in the kruger national park, south africa. site-specific curves produced excellent estimates of soil water potential from soil water content values. curves from soils derived from the same geological substrate were similar, potentially allowing for the use of one curve for basalt soils and another for granite soils. it is anticipated that this dataset will help hydrologists and ecophysiologists understand water dynamics, carbon cycling and biosphere–atmosphere interactions under current and changing climatic conditions in the region. introduction top ↑ soil moisture is a key driver of plant productivity in many ecosystems, since water-stressed plants close their stomata to curb water loss, resulting in reduced carbon dioxide (co2) assimilation rates and, thus, growth (lambers et al. 2008). in order to understand water availability, ecologists, agronomists and land managers often use measurements of gravimetric (g of water g-1 soil) or volumetric (cm3 of water cm-3 soil) soil water to estimate water availability to plants. these measurements can easily be made by a wide array of sensors or simply by weighing and drying soil samples. the problem with these measurements is that they do not necessarily provide information about whether or not water is available to plants. this is because surface tension can bind large amounts of water to soils with a high internal surface area (i.e. clays) at such large negative pressures that plants cannot oppose them and are therefore unable to take up the water (vogel 2012). the negative pressure with which water is bound to the soil is the soil water potential. in non-saline soils, the relationship between soil water content and soil water potential largely depends on texture. clay particles have a large surface area relative to their volume and therefore have the ability to bind large amounts of water. as a result, a clay soil with, for example, 10% moisture, may have a highly negative water potential, making water uptake impossible for most plants. sand particles are larger and more spherical and therefore have a lower surface-to-volume ratio. consequently, a sandy soil with 10% moisture may have a water potential that is close to zero, allowing for easier water uptake by plants. temperate crops can transpire water through their stomata, and can therefore photosynthesize, down to a soil water potential of about -1.5 mpa. whilst this is called the permanent wilting point, some plants in arid systems are able to transpire water at a soil water potential as low as -5 mpa (baldocchi et al. 2004; rodriquez-iturbe & porporato 2004). ecologists, vegetation modellers and those modelling biosphere–atmosphere interactions therefore need accurate estimates of soil water potential in the range from -5 mpa to 0 mpa to predict and explain stomatal responses, carbon dynamics, and water and energy budgets. pedo-transfer functions transform variables that are easy and cheap to measure into more informative variables that are too difficult or expensive to measure directly (bouma 1989). one such function is the soil water retention curve, which transforms soil water content into soil water potential. the water retention curve is often estimated from information on soil texture, but can be determined precisely by measuring both variables simultaneously on samples under controlled conditions. here we provide empirically derived soil water characteristic curves for seven soils representing the major soil types of the kruger national park (knp). we test whether the same curve can be used for soils with a similar texture. methods top ↑ study site the kruger national park is situated in the north-east of south africa between 30.9–32.0 °e and 22.3–25.5 °s. the park receives between 450 mm and 750 mm yr-1 of rain, most of which falls between october and march. average temperatures are approximately 25 °c in summer and 20 °c in winter. most of knp is underlain by either basaltic rock that weathers into clay-rich soils or granitic rock that weathers into sandy soils. both of these dominant parent materials are old: the basaltic rock was formed ~200 ma, whilst the granite was formed ~2050 ma (venter et al. 2003). seven dominant soil types from across the park (venter et al. 2003) were sampled (figure 1; table 1). the sampled soil types cover approximately 65% of knp. figure 1: dominant soil types of the kruger national park following venter et al. (2003), and the sampling locations of this study. table 1: coordinates of the sampling sites and description of soils at those sites, following venter et al. (2003). the sand, silt and clay fractions are the mean of all samples from a site in percentages ± the standard error. approach using a soil auger, samples were collected at three depths: at the top of the profile (0 cm – 10 cm), at the bottom (110 cm or degraded bedrock, whichever was shallower) and in the middle (table 2). soils were air dried and sieved in a 2 mm-aperture sieve to remove large roots and rocks. the proportion of rock with dimensions greater than 2 mm was generally negligible (< 3%). table 2: parameter estimates, standard deviations of the parameter estimates, and goodness of fit for the power functions fitted as water retention curves for soils in the kruger national park. water retention curves soil moisture retention curves were derived empirically using an instrument that exploits the chilled-mirror technique (wp4t, decagon devices inc., pullman). a soil sample is inserted into the device and equilibrated with the headspace of a sealed chamber, so that the water potential of the air in the chamber is the same as the water potential of the sample. the point of condensation on a mirror is detected by shining a beam of light onto the mirror and recording its reflectance with a light sensor. at the point of condensation, the temperature of the mirror is recorded, allowing the water potential of the air, and therefore of the sample, to be calculated. the instrument measures the water potential with an accuracy of 0.05 mpa over the range of -0.1 mpa – -0.5 mpa and an accuracy of 0.1 mpa over the range of -0.5 mpa – -300 mpa. therefore, it adequately covers the range relevant to plant growth. the water content of the sample at the time of measuring water potential is determined gravimetrically (i.e. it is weighed and reweighed after oven-drying at 60 °c to a constant weight). since the relationship between water potential and water content is often hysteretic (affected by the initial state of the system), we constructed soil moisture retention curves from both drying and wetting soils. soil from each sampling location and depth was subdivided into 15 samples of 5 g – 6 g each. these samples were used to construct a single drying curve and a single wetting curve for each depth at each sampling location. for the drying curve, the samples were initially saturated with distilled water, sealed, and left to equilibrate overnight. water potential and moisture content were then measured repeatedly as the samples air dried. a typical curve was completed within a day. for the wetting curve, incremental amounts of moisture were added to initially dry samples to achieve a range of saturation levels. samples were then sealed and left to equilibrate overnight, after which the water potential and moisture content were measured. empirical models for the water retention curve are typically written to be solved for water content (see assouline et al. [1998] for an overview). we fitted two widely used models, proposed by van genuchten (1980), and fredlund and xing (1994), which were rewritten to solve for water potential given a known water content using non-linear regression in r (r core team 2013). we compared the performance of these models to simpler power and exponential functions, which were fitted by maximising the likelihood using iterative methods (plummer 2013). these functions were fitted to both the entire measured range of water potentials (0 mpa – -100 mpa) and to a subset that is relevant to plant growth (0 mpa – -8 mpa). to assess the variability amongst sites of the same geology, we compared site-specific curves to curves for which the data were aggregated by geology. this was done for four levels of water content (0.03 g g-1, 0.05 g g-1, 0.1 g g-1 and 0.15 g g-1). for this analysis, soils from punda, which are derived from quartzite (venter et al. 2003), were included with granite-derived soils as they had a very similar texture. soil texture particle size affects the total surface area for a given mass or volume of soil, and therefore the shape of the water retention curve. to quantify sand, silt and clay fractions, the hydrometer method (bouyoucos 1962) was used. for each sample, between 40 g and 50 g of soil was dispersed with an electrical mixer in 100 ml of a 5% sodium hexametaphosphate ([napo3]6) solution, in order to break down clay aggregates. this mixture was allowed to soak overnight and mixed again the following morning before being transferred to a 1 l cylinder filled up to 1 l with distilled water. sediment was dispersed with a plunger and hydrometer readings were taken after 40 s and 6 h 52 min. fractions were calculated as: where h is the hydrometer reading at t seconds after inserting the hydrometer, and w is the sample weight in grams. hydrometer readings were corrected for the added (napo3)6 by subtracting the hydrometer reading of a cylinder with 100 ml (napo3)6 solution filled up to 1 l with distilled water. the temperature was measured at the time of each hydrometer reading and 0.4 g l-1 of solute concentration was added to the hydrometer reading for every degree c above 20 °c ( http://uwlab.soils.wisc.edu/files/procedures/particle_size.pdf). results top ↑ as expected, soils from the basalt substrate had a much higher clay content than soils from the granitic substrate (table 1; table 2). the water potential and water content data is given in online appendix 1. the fredlund and xing model (1994) appeared to fit the data well, with r2 values > 0.99 for most curves. however, the confidence intervals on the parameter estimates could not be estimated using standard techniques, suggesting the parameters were not identifiable. in other words, multiple combinations of parameter values can result in an identical fit. this outcome is likely to be a symptom of over-parameterisation of the model. the van genuchten model (1980) did not converge well, also hinting at over-parameterisation. therefore only results from the simpler power and exponential functions were reported. the best fit to individual curves in the -0.5 mpa – -8 mpa range was provided by a power function of the form (table 2; appendix 1): where ψ is water potential in mpa, θ is gravimetric water content in g g-1, and a and c are estimated coefficients. for a given water content, water potentials from drying curves were generally lower than for wetting curves (appendix 1). when combining data from all sites per geological substrate, the residuals of the power function fits showed systematic bias. an exponential model to describe the data aggregated by geology was therefore used. the exponential model fits these grouped data reasonably well in the -0.5 mpa – -8 mpa range: basalt r2 = 0.59; granite r2 = 0.66 (figure 2; see appendix 2 for 0 mpa – -100 mpa range). it should be noted that r2 values should be interpreted with caution in non-linear regression, and that the standard error of the regression (ser) is a better measure of goodness of fit: basalt ser = 1.42 mpa; granite ser = 1.13 mpa. the equation to convert soil water content to water potential for basaltic (clayey) soils in the -0.5 mpa – -8 mpa range is: for granitic (sandy) soils in the -0.5 mpa – -8 mpa range, the equation is: to assess the accuracy of these geology-specific curves, we compared the range in water potential for four water contents (0.03, 0.05, 0.10 and 0.15), as predicted by the site-specific models to the predicted water potential, using the geology-specific curve (figure 3). the geology-specific curves predicted water potentials close to the median value of site-specific curves, suggesting that they might be useful for modelling applications. figure 2: the relationship between soil moisture and water potential in the range relevant to plant growth, for (a) basalt-derived and (b) granite-derived soils in the kruger national park. note that the pressures on the y-axis are negative pressures. the water retention curves were fitted using an exponential model. figure 3: comparing site-specific and geology-specific curves for (a) basalt and (b) granite. each box shows the range of predicted water potentials from all site-specific curves per geological substrate for a given water content. the solid horizontal line in the box denotes the median of predicted values. the horizontal dotted lines indicate the wilting range (-1.5 mpa – -5 mpa). the * symbols show the predicted values from the geology-specific curves. the water potentials at water contents of 0.03 g g-1 and 0.05 g g-1 in basaltic soils are not shown as they are < -25 mpa and not relevant for plant physiological processes. discussion top ↑ ecophysiological and hydrological interpretation of soil water content – an affordable measure of soil moisture – requires the use of pedo-transfer functions that transform water content into water potential. we generated such water retention curves with a wp4t instrument (decagon devices inc., pullman) using empirical data, and provided curves for common soil types in knp, south africa. a power function provided a good fit for individual samples, and, depending on the required accuracy, a single exponential function per geological substrate may be used. at low water contents (i.e. ~5% for granitic and ~15% for basaltic soils), water potentials estimated from our pedo-transfer functions begin to vary widely between sites, reflecting differences in clay content. thus, researchers interested in precise estimates of soil water potentials in dry soils should use our site-specific functions. however, these water potentials are below the permanent wilting range (-1.5 mpa – -5 mpa; figure 3) and represent small volumes of soil water. researchers using the equations provided here should take care to avoid several potential problems. first, our analyses relied on the chilled-mirror technique. this method is highly accurate in the dry range (-0.5 mpa – -300 mpa), but is less accurate in the wet range of soils (-0.001 mpa – -0.5 mpa). it should be noted that some crop species may become water limited at -0.03 mpa (rodriquez-iturbe & porporato 2004). second, the functions presented here are based on gravimetric soil water measurements, whilst some field sensors measure soil moisture volumetrically. to use the functions provided in this study, volumetric soil moisture should be divided by the bulk density of the soil. bulk density in knp varies with texture, spatially, and with soil depth (wigley et al. 2013). when converting gravimetric or volumetric contents to water potentials, researchers should be aware of the potential role of coarse rock fragments. a soil sensor that provides volumetric water content in a rocky soil will underestimate soil water potentials because the volumetric sensor reports the water content of a volume comprised of both rock and soil. in conclusion, the presented soil water retention curves will improve estimates of plant-available water from measurements of volumetric or gravimetric soil moisture in knp and surrounding areas with similar geological substrates. acknowledgements top ↑ the authors would like to thank the management of the kruger national park for enabling this study. ben wigley kindly made bulk density data available; tercia strydom made equipment available; alexander zizka, julia fischer and jenny toivio assisted with data collection; and stephen doucette-riise and his team collected soil samples. this study was funded by the deutsche forschungsgemeinschaft (dfg). competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions r.b. (goethe universität frankfurt) collected and analysed data, and wrote the article. a.k. (utah state university) conceived the study and wrote the article. s.i.h. (university of otago) analysed data and wrote the article. references top ↑ assouline, s., tessier, d. & bruand, a., 1998, ‘a conceptual model of the soil water retention curve’, water resources research 34(2), 223–231. http://dx.doi.org/10.1029/97wr03039 baldocchi, d.d., xu, l. & kiang, n., 2004, ‘how plant functional-type, weather, seasonal drought, and soil physical properties alter water and energy fluxes of an oak–grass savanna and an annual grassland’, agricultural and forest meteorology 123(1 & 2), 13–39. http://dx.doi.org/10.1016/j.agrformet.2003.11.006 bouma, j., 1989, ‘using soil survey data for quantitative land evaluation’, in b.a. stewart (ed.), advances in soil science, pp. 177–213, springer-verlag, new york. http://dx.doi.org/10.1007/978-1-4612-3532-3_4 bouyoucos, g.j., 1962, ‘hydrometer method improved for making particle size analyses of soils 1’, agronomy journal 54(5), 464–465. http://dx.doi.org/10.2134/agronj1962.00021962005400050028x fredlund, d. & xing, a., 1994, ‘equations for the soil-water characteristic curve’, canadian geotechnical journal 31(3), 521–532. http://dx.doi.org/10.1139/t94-061 lambers, h., chapin iii, f.s. & pons, t.l., 2008, plant physiological ecology, springer, new york. http://dx.doi.org/10.1007/978-0-387-78341-3 plummer, m., 2013, jags – just another gibbs sampler, viewed 2013, from http://mcmc-jags.sourceforge.net/ r core team, 2013, r: a language and environment for statistical computing, r foundation for statistical computing, vienna, austria, viewed 2013, from http://www.r-project.org/ rodriquez-iturbe, i. & porporato, a., 2004, ecohydrology of water-controlled ecosystems: soil moisture and plant dynamics, cambridge university press, cambridge. van genuchten, m.t., 1980, ‘a closed-form equation for predicting the hydraulic conductivity of unsaturated soils’, soil science society of america journal 44(5), 892–898. http://dx.doi.org/10.2136/sssaj1980.03615995004400050002x venter, f.j., scholes, r.j. & eckhardt, h.c., 2003, ‘the abiotic template and its associated vegetation pattern’, in j.t. du toit, k.v. rogers & h.c. biggs (eds.), the kruger experience: ecology and management of savanna heterogeneity, pp. 83–129, island press, washington dc. vogel, s., 2012, the life of a leaf, university of chicago press, chicago. http://dx.doi.org/10.7208/chicago/9780226859422.001.0001 wigley, b.j., coetsee, c., hartshorn, a.s. & bond, w.j, 2013, ‘what do ecologists miss by not digging deep enough? insights and methodological guidelines for assessing soil fertility status in ecological studies’, acta oecologica 51, 17–27. http://dx.doi.org/10.1016/j.actao.2013.05.007 appendix 1 top ↑ figure a1: the relationship between soil moisture and water potential in the range relevant to plant growth for each of the sampled soils in the kruger national park, letaba, lower sabie, phalaborwa, pretoriuskop, punda, satara, skukuza. the pressures on the y-axis are negative pressures. the water retention curves were fitted using a power function. the parameter values for the curves are given in table 2. figure a1 (continues...): the relationship between soil moisture and water potential in the range relevant to plant growth for each of the sampled soils in the kruger national park, letaba, lower sabie, phalaborwa, pretoriuskop, punda, satara, skukuza. the pressures on the y-axis are negative pressures. the water retention curves were fitted using a power function. the parameter values for the curves are given in table 2. appendix 2 top ↑ figure a2: the relationship between soil moisture and water potential in the 0 mpa – -100 mpa range for (a) basalt-derived and (b) granite-derived soils in the kruger national park. the pressures on the y-axis are negative pressures. the water retention curves were fitted using an exponential model. zietsman.qxd dune vegetation and coastal thicket plant communities in threatened limestone fynbos of andrew’s field and tsaba-tsaba nature reserve, struisbaai, western cape m.m. zietsman and g.j. bredenkamp zietsman, m.m. and g.j. bredenkamp. 2006. dune vegetation and coastal thicket plant communities in threatened limestone fynbos of andrew’s field and tsaba-tsaba nature reserve, struisbaai, western cape. koedoe 49(1): 33–47. pretoria. issn 00756458. the coastal thicket and dune vegetation of andrew’s field and tsaba-tsaba nature reserve was classified using braun-blanquet procedures and twinspan. the vegetation was sampled using 74 randomly stratified sample plots. the floristic composition, cover-abundance of each species, and several environmental variables were recorded in each sample plot. six plant communities were identified, namely, rhus glauca euclea racemosa low to tall closed thicket community; chrysanthemoides monilifera solanum africanum low closed dune shrub community; chrysanthemoides monilifera ehrharta villosa var. maxima low to high closed dune shrub community; ehrharta villosa var. maxima low to short closed dune grassland community; ammophila arenaria low to short closed dune grassland community; and arcthotheca populifolia thinopyrum distichum low to short open beach community. these were subdivided into eight subcommunities and four variants. all communities, sub-communities and variants were described and ecologically interpreted. the distribution of the communities, sub-communities and variants can mainly be ascribed to differences in landform, rockiness of the soil surface the degree of protection / exposure of the vegetation to the dominating winds of the area. key words: coastal thicket, conservation area, endemic plants, limestone fynbos, phytosociology, plant communities, twinspan, western cape m.m. zietsman and g.j. bredenkamp, african vegetation and plant diversity research centre, department of botany, university of pretoria, pretoria, 0002 republic of south africa. issn 0075-6458 33 koedoe 49/1 (2006) introduction a sound knowledge of the ecology of the area is an essential prerequisite for the establishment of efficient wildlife management programmes and compilation of conservation policies for any area (edwards 1972). vegetation science has been applied in the fields of nature conservation for years, but recent developments relate to the application of plant ecological knowledge to environmental management (scheepers 1983). bredenkamp & brown (2001) emphasise the use of plant communities as a reliable basis for any ecological planning and management. to obtain knowledge of the ecology of the study area, a study of the vegetation of the area should be made at the plant community level of organisation (bredenkamp & brown 2001). studying the vegetation of an area allows the identification, description and classification of plant communities. on the basis of the plant communities as vegetation units, management units can be delineated. vegetation studies also allow the identification of ecologically sensitive areas, bush encroached areas or areas infested with alien plants, degraded areas, habitats of rare or endangered plant species, and habitats of specific animals. the andrew’s field and tsaba-tsaba reserve is situated in the bredasdorp/riversdale centre of endemism (cowling 1992). this centre of endemism refers to a welldefined group of plants, confined to the limezietsman.qxd 2006/04/17 08:48 pm page 33 stone of the bredasdorp formation and associated colluvial deposits (heydenrych 1994). according to hilton-taylor & le roux (1989) limestone fynbos is one of the most threatened vegetation types in the cape floristic region. factors threatening limestone fynbos vegetation include alien plants (considered the biggest problem), land clearing, resort development, inappropriate fire management and over-harvesting of flowers (heydenrych 1994). thicket encroachment into the adjacent limestone fynbos and renosterveld is therefore a serious threat to the species-rich fynbos with many endemic and red data species. the species richness of coastal thicket in the reserve is very low compared to those of the adjacent limestone fynbos and renosterveld communities (zietsman & bredenkamp in press) and the true thicket species like rhus glauca and euclea racemosa are neither rare nor endemic. regarding the importance of the conservation of limestone fynbos in areas where the thicket communities occur, the effective management of the area becomes all the more important, and fully justifies a detailed vegetation study of the coastal thicket. study area the reserve is situated in the bredasdorp district, western cape, between struisbaai north in the south and de mond state forest in the north. it is bordered in the west by the bredasdorp/struisbaai road, and in the south by the coastline. the study area is situated close to the coastal town of struisbaai. the andrew’s field (129 ha) and tsaba-tsaba nature reserve is approximately 979 ha in extent. approximately one third of the study area consists of the coastal vegetation, which is dealt with in this report. the remainder of the study area consists of inland plains and hills, described by zietsman & bredenkamp (in press). the dominant substrate in the study area is the bredasdorp group (malan et al. 1994). two formations of the bredasdorp group namely strandveld and waenhuiskrans formations are found in the study area (malan et al. 1994). beach and terrace deposits, not formally named as a formation, but forming part of the bredasdorp group, are found in the form of roll-stones on the coastal beach (malan et al. 1994). previous changes in sea level resulted in the presence of roll-stones of table mountain sandstone. the strandveld formation is restricted to the coast. it consists of white to light-grey dune sands with a high percentage shell fragments. partial cementing of sands is a result of the high calcium carbonate content in the sand. the lithology of this formation is described as white dune sand, strand sand with finely divided shell and alluvial stones (malan et al. 1994). the waenhuiskrans formation forms outcrops adjacent to the current coastline. the waenhuiskrans stratiotype is 12.4 m deep, is locally overlain with 1 m thick calcretes and consists of medium-grained cross-layered calcarenite with well-rounded quartz and a few glauconite grains. large-scale aeolic cross-layers are characteristic of the unit. the lithology of this formation is described as partially calcified dune sand (malan et al. 1994). the study area is situated on the agulhas plain, a coastal lowland. the majority of the area falls below the 10 m contour (jefferey 1996). coastal dunes and dune plains are typical in this area, with the highest point on a ridge 31 m above sea level. the soil can be described as shallow sandy soil, overlying limestone, or shallow to deep sandy soil, overlying clay, silt and gravel. four different soil forms have been distinguished in the area (macvicar 1991): coega: orthic a on hard bank carbonate horizon; family marydale: lime containing a-horizon. this soil form is found on the limestone hills and shallow-soil limestone plain. immerpan: melanic a on hard bank carbonate horizon; family kalkpan: lime containing a-horizon. this soil form is found on the proteoid dominated limestone plain koedoe 49/1 (2006) 34 issn 0075-6458 zietsman.qxd 2006/04/17 08:48 pm page 34 where the soil is deep. brandvlei: orthic a on soft carbonate horizon; family kolke: signs of wetness in carbonate horizon. the marsh area is characterised by this soil form. namib: orthic a on regic sand; family beachwood: containing lime within 1500 mm from soil surface. this soil form is found on the dune plain and the deep-sand plains. according to tinley (1985) two distinct sand characteristics occur in dune fields. bare dune sands, yellow in colour, absorbs all rain and consequently there is no surface runoff. grey sands, stained by humus and covered with woody vegetation, have a water repellent layer near the surface, beneath the litter, causing a massive surface runoff when heavy rains follow a dry period (tinley 1985). the main factors affecting the climate are the contrasting sea surface temperatures of the two major ocean currents and the inshore circulation (tinley 1985). according to the geographical division of heydorn & tinley (1980), the reserve is situated on the south coast, which is a transitional zone between tropical and temperate waters. the study area is situated in the transitional zone between the winter-rainfall region in the west and the non-seasonal rainfall region in the east (mustart et al. 1997). the south coast has a warm temperate climate with allseasons and bimodal equinoctial rainfall (strydom 1992). the average annual precipitation of the area is 444 mm. precipitation is mostly rain, but it also occurs in the form of fog. the maximum precipitation is in june, and the minimum during february and december. according to heydorn & tinley (1980) the study area is situated in a low rainfall region, within an arid belt protruding into the interior from the coast. the low rainfall appears to be the result of cold inshore waters that inhibit shoreline rains. a walter climatic diagram was compiled (fig. 1) from data obtained from the agulhas weather station. the average annual maximum and minimum temperatures for the area are 6 ºc and 13.3 ºc respectively. the maximum temperature for the area was 36.1 ºc, obtained in february. the minimum temperature for the area is 3.9 ºc, obtained in june (zietsman & bredenkamp in press). the wind along the south coast is bidirectional. southeasterly winds alternate with issn 0075-6458 35 koedoe 49/1 (2006) fig. 1. a walter climatic diagram for andrew’s field and tsaba-tsaba nature reserve. a = altitude, b = mean annual temperature, c = mean annual rainfall, d = mean daily minimum (coldest month), e = mean daily maximum (hottest month) zietsman.qxd 2006/04/17 08:48 pm page 35 northwesterly and southwesterly winds (heydorn & tinley 1980). methods relevés were compiled in 74 stratified sample plots, placed in relatively homogeneous areas, representative of particular plant communities that occur in a mosaic distribution pattern. a plot size of 10 m² was used, and considered as large enough to ensure that all species of regular occurrence in the stand are present in the sample plot. sample plots were placed in such a way as to ensure that each plot adequately represents the structure of the particular vegetation (werger 1974). a list of plant species found in each sample plot was compiled. the cover-abundance of each species in the sample plot was assessed, using the braun-blanquet cover-abundance scale (werger 1974). cover of the height classes of the different strata (edwards 1983) was estimated and the vegetation was structurally classified according to the edwards (1983) structural classification system. the following habitat characteristics were recorded in each sample plot: altitude, topographical position, exposure to wind (seaward or towards the land), slope angle, slope direction, geology, soil, percentage rock cover and biotic influence. six main different habitat types are found in the study area, namely dune plain, renosterveld plain, limestone plain, limestone hills, dunes and beach. these were recorded in each sample plot. to obtain a first approximation of the plant communities of the area, relevés were classified using twinspan (hill 1979), and these results were then refined by application of the classical braunblanquet methodology (behr & bredenkamp 1988). results classification the results obtained from the classification are presented in a phytosociological table (table 1). in naming the plant communities, diagnostic and/or dominant species were used in combination with the edwards (1983) structural classification. 1. rhus glauca euclea racemosa low to tall closed thicket community 1.1 rhus lucida euclea racemosa low to short closed thicket sub-community 1.2 pterocelastrus tricuspidatus euclea racemosa short to tall closed thicket sub-community 1.2.1 olea exasperata euclea racemosa short to high closed thicket variant 1.2.2 carissa bispinosa euclea racemosa short to tall closed thicket variant 1.3 acmadenia obtusata euclea racemosa short to tall closed thicket subcommunity 1.4 helichrysum dasyanthum euclea racemosa low to short closed thicket sub-community 1.5 acacia cyclops euclea racemosa high to tall closed thicket sub-community 1.6 thamnocortus insignis euclea racemosa high to tall closed thicket subcommunity 1.6.1 elytropappus rhinocerotus euclea racemosa short to tall closed thicket variant 1.6.2 leucadendron coniferum euclea racemosa high to tall closed thicket variant 2. chrysanthemoides monilifera solanum africanum low closed dune shrub community 3. chrysanthemoides monilifera ehrharta villosa var. maxima low to high closed dune shrub community 3.1 chrysanthemoides monilifera rhus crenata short to high dune shrub subcommunity 3.2 chrysanthemoides monilifera morella cordifolia low to short closed shrub sub-community 4. ehrharta villosa var. maxima low to short closed dune grassland community 5. ammophila arenaria low to short closed dune grassland community 6. arcthotheca populifolia thinopyrum distichum low to short open beach community the cover and height of various strata of the main plant communities are given in table 2. the hierarchical classification and associated environmental interpretation of the plant communities is given in fig. 2. the community numbers in fig. 2 correspond with the plant community numbers used in the descriptions in the text. 1. rhus glauca euclea racemosa low to tall closed thicket community the community is found close to the sea on the dune plains, limestone plains, renosterkoedoe 49/1 (2006) 36 issn 0075-6458 zietsman.qxd 2006/04/17 08:48 pm page 36 issn 0075-6458 37 koedoe 49/1 (2006) dune plains limestone plains rhenoster plains dune slopes slopes and peaks of limestone hills flat dune plain sandy shallow limestone plain dune plains deep to flat limestone plains sandy footslopes, slopes and shoulders of limestone hills sheltered, steep north-facing slopes of back-dune deep limestone plains peaks, foot-slopes of limestone hills flat limestone plains peaks and shoulders of limesone hills rhenosterveld plain limestone hills foot-slopes dune slopes sheltered from south-eastern wind dune slopes parallel to south-eastern wind dune plain deep sandy slopes and shoulders of limestone hills limestone plains acacia cyclops encroachment in community 1 rhenosterveld plains limestone hills, footslopes very steep dry, warm dune slopes coastal dunes and dune troughs south-facing slopes, sparsely vegetated dunes northern steep dry mid-slopes of dunes beach and gravel plain frontal dune slopes 1.1 1.2.1 1.2.2 1.3 1.4 1.5 1.6.1 1.6.2 2 3.1 3.2 4 5 6 fig. 2. hierarchical classification and associated environmental characteristics of the coastal thicket, dunes and strand of andrew’s field and tsaba-tsaba nature reserve. zietsman.qxd 2006/04/17 08:48 pm page 37 ta bl e 1 p hy to so ci ol og ic al ta bl e of c oa st al th ic ke t, du ne s an d st ra nd o f a nd re w 's f ie ld a nd t sa ba -t sa ba n at ur e r es er ve zietsman.qxd 2006/04/17 08:48 pm page 38 ta bl e 1 (c on tin ue rd ) zietsman.qxd 2006/04/17 08:48 pm page 39 ta bl e 1 (c on tin ue rd ) zietsman.qxd 2006/04/17 08:48 pm page 40 issn 0075-6458 41 koedoe 49/1 (2006) veld plains, northern slope of back-dunes, and on the slopes and peaks of limestone hills. these areas have a low cover of small irregular white-grey limestone pebbles. the rhus glauca euclea racemosa short to tall closed thicket community consists of scattered, dense groups of broad-leaved shrubs. these groups have high shrubs near the centre and lower shrubs near the periphery. the structure of the vegetation is given in table 2. the shrub layer is most prominent, while restioid and grass layers cover 10 % or less. forbs and sedges have a very low cover. the shrubs euclea racemosa, rhus glauca and rhus laevigata (species group a, table 1) are diagnostic and dominant in this community. chrysanthemoides monilifera and metalasia muricata (species group q) are also prominent in the community. in the absence of fire, these thickets tend to encroach into adjacent threatened limestone fynbos and rhenosterveld communities which have several endemic and red data species (cowling & richardson 1995; heydenrych 1994; zietsman & bredenkamp in press). six sub-communities, two of which have two variants, were identified: 1.1 rhus lucida euclea racemosa low to short closed thicket sub-community this sub-community is found on the flat to undulating dune plain, namib soil form and limestone plain, brandvlei and coega soil forms (fig. 2). the vegetation forms scattered groups of low to short broad-leaved shrubs. rhus lucida (species group b, table 1) is the only diagnostic species for this sub-community. the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a, table 1) are the dominants in this sub-community. the shrubby helichrysum dasyanthum (species group g) and passerina paleacea (species group i) are locally prominent in the subcommunity. an average of only four species per relevé was recorded in this sub-community. 1.2 pterocelastrus tricuspidatus euclea racemosa short to tall closed thicket sub-community the sub-community is found on the namib soil form on the dune plain and on the immerpan and coega soil forms on sandy foot-slopes and shoulders of limestone hills (fig. 2). table 2 percentage cover (c ) (%) and height (h)(m) of the strata of the main plant communities community 1 2 3 4 5 6 stratum c h c h c h c h c h c h tall shrub 14 2.5 6 2.5 6 2.5 high shrub 20 1.5 5 1.5 10 1.5 10 1 short shrub 28 0.75 5 0.75 29 0.75 low shrub 15 0.4 51 0.15 16 0.3 forb 3 0.3 1 0.3 2 0.3 10 0.5 short restoid 4 0.75 0.3 0.2 low restoid 7.5 0.4 sedge 1 0.15 0.5 0.15 high grass 0.5 1.5 3.5 1.5 short grass 0.5 0.5 0.6 0.75 2 0.75 25 0.75 44 0.7 5 0.5 low grass 10 0.3 2 0.3 15 0.3 2 0.3 zietsman.qxd 2006/04/17 08:48 pm page 41 koedoe 49/1 (2006) 42 issn 0075-6458 the vegetation forms scattered groups of short to tall broad-leaved shrubs, with some lower shrubs at the periphery. species group c (table 1) is diagnostic for this sub-community, with the shrubby trees robsonodendron maritimum and pterocelastrus tricuspidatus dominant diagnostic species. the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a), are also very prominent in the sub-community. 1.2.1 olea exasperata euclea racemosa short to high closed thicket variant the variant occurs on the slopes and peaks of the limestone hills, and on the sandy, deep-soil limestone plains (fig. 2). species group d (table 1) is diagnostic, with olea exasperata and osyris compressa prominent diagnostic species. the diagnostic species of the community and sub-community, namely the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a), robsonodendron maritimum and pterocelastrus tricuspidatus (species group c) are dominant in the variant. an average of 12 species per relevé was recorded for this variant. 1.2.2. carissa bispinosa short to tall closed thicket variant this variant is restricted to the flat limestone plains and the shoulders and peaks of limestone hills (fig. 2). this soil is of the immerpan and coega forms. species group e (table 1) is diagnostic with the shrubs carissa bispinosa and sideroxylon inerme the most prominent diagnostic species. the shrubs euclea racemosa, rhus glauca (species group a), robsonodendron maritimum and pterocelastrus tricuspidatus (species group c) are dominant whereas the alien shrub acacia cyclops (species group h) is also locally prominent. an average of 13 species per relevé was recorded for this variant. 1.3 acmadenia obtusata euclea racemosa short to tall closed thicket sub-community the sub-community is restricted to a small patch close to the sea, in the central part of the reserve, on a steep (26° to 45°), sheltered northern slope of a back-dune. the soil is of the namib form and no rocks are found here. only a single relevé represents this sub-community. the vegetation forms scattered groups of short to tall closed broad-leaved shrubs, with some lower shrubs at the periphery. species group f (table 1) is diagnostic, with the shrub acmadenia obtusata, the forbs arctopus echinatus and zaluzianskya villosa and the grass festuca scabra, as diagnostic species. the shrubs euclea racemosa (species group a), otholobium bracteolatum (species group g), rhus crenata (species group n) and passerina rigida (species group p) are dominant. eighteen species were recoded in the single releve. 1.4 helichrysum dasyanthum euclea racemosa low to short closed thicket sub-community this sub-community is widespread on the dune plain, namib soil form, the sandy shallow-soil limestone plain and sandy slopes of limestone hills, immerpan and coega soil forms (fig. 2). the vegetation forms scattered groups of low to short broad-leaved shrubs, with some high shrubs at the centre. otholobium bracteolatum and helichrysum dasyanthum (species group g, table 1) are the diagnostic species for this sub-community. the shrubs euclea racemosa, and rhus laevigata var. laevigata for. coangoa (species group a) are dominant, while rhus glauca (species group a), pterocelastrus tricuspidatus (species group c), passerina paleacea (species group i), chrysanthezietsman.qxd 2006/04/17 08:48 pm page 42 issn 0075-6458 43 koedoe 49/1 (2006) moides monilifera (species group q) are also locally prominent. the sub-community contains an average of only six species per relevé. 1.5 acacia cyclops euclea racemosa high to tall closed thicket sub-community locally the alien acacia cyclops encroached into the rhus glauca euclea racemosa short to tall closed thicket community. acacia cyclops, the shrubby tree protea obtusifolia, and the shrub passerina galpinii are the diagostic species for this sub-community (species group h, table 1). acacia cyclops and the shrubs euclea racemosa, rhus glauca and rhus laevigata var. laevigata for. coangoa (species group a), and metalasia muricata (species group q) are dominant in the sub-community. the sub-community contains an average of seven species per relevé. 1.6 thamnochortus insignis euclea racemosa high totall closed thicket sub-community the sub-community is found on the renosterveld plain, brandvlei soil form and on sandy foot-slopes of limestone hills, namib soil form (fig. 2). the vegetation forms scattered groups of high to tall broad-leaved shrubs, with some low and short shrubs at the periphery. low forbs and some high, low and short restioids are also present in the sub-community. species group i (table 1) is diagnostic for this sub-community with the restioid thamnochortus insignis and the shrubs helichrysum patulum, passerina paleacea, metalasia densa the most prominent diagnostic species. the shrubs euclea racemosa, rhus glauca (species group a), chrysanthemoides monilifera and metalasia muricata (species group q), are also prominent in the subcommunity. two variants are recognised within this subcommunity: 1.6.1 elytropappus rhinocerotus short to tall closed thicket variant the variant is found at an altitude of 3 m on the renosterveld plain, brandvlei soil form (fig. 2). the vegetation scattered groups of short to tall broad-leaved shrubs, with some lower shrubs at the periphery, as well as some high restioids. the diagnostic species are listed in species group k (table 1), which includes the shrubs elytropappus rhinocerotis, leucadendron linifolium, oedera uniflora, lycium afrum, rhus laevigata var. villosa, asparagus lignosus, euryops hebecarpus, and the forbs limonium scabrum var. scabrum, hermannia althaeifolia and pelargonium myrrhifolium. the shrubs euclea racemosa and rhus glauca (species group a) and the restioid thamnochortus insignis (species group i) are dominant whereas the shrubs helichrysum patulum, passerina paleacea (species group i), chrysanthemoides monilifer and metalasia muricata (species group q) are locally prominent. the variant has an average of 22 species per relevé. 1.6.2 leucadendron coniferum high to tall closed thicket variant the variant is found on the sandy foot-slopes of limestone hills, namib soil form (fig. 2). the vegetation forms scattered groups of high to tall closed broad-leaved shrubs, with some low to short shrubs at the periphery, as well as some high and low restioids. a large number of species given in species group k (table 1) are diagnostic for this variant, the most prominent being the shrubs eriocephalus kingesii, leucadendron coniferum, protea susannae, maytenus procumbens, agathosma serpyllacea, agathosma collina, phylica stipularis, trigogyne repens and tephrosia capensis and the restioid chondopetalum microcarpum. the shrubs euclea racemosa and rhus glauca (species group a), and the restioid thamnochortus zietsman.qxd 2006/04/17 08:48 pm page 43 koedoe 49/1 (2006) 44 issn 0075-6458 insignis (species group i) are dominant, while the shrubs olea exasperata (species group d), cassine peragua (species group i), solanum africanum (species group l), chrysanthemoides monilifera (species group q) are locally prominent. the variant is rich in species and has an average of 43 species per relevé. 2. chrysanthemoides monilifera solanum africanum low closed shrub dune community the sparsely vegetated community is found in the shifting sand dune area adjacent to the sea. it occurs in a mosaic distribution pattern with communities 3, 4 and 5. the community is situated at an altitude of 1–20 m, on steep gradients. this vegetation is restricted to the drier northerly mid-slopes of coastal dunes, facing inland (fig. 2). the soil is of the namib form and no rocks are present. the vegetation consists of low prostrate shrubs, with a few short, high and tall shrubs also present. the low shrub stratum is prominent with an average cover of 51 % (table 2). the low shrub solanum africanum (species group l, table 1) is not only diagnostic but also dominant in the community. the shrub chrysanthemoides monilifera (species group q) is also locally prominent. the community contains an average of only four species per relevé. 3. chrysanthemoides monilifera ehrharta villosa var. maxima low to high closed dune shrub community the community is found in the sparsely vegetated shifting dune area adjacent to the sea, in a mosaic distribution pattern with communities 2, 4 and 5. it is also present in a mosaic distribution pattern with communities 2 and 4 in the more densely vegetated sand dune area at the seashore and directly behind to the shifting sand dune area (fig. 2). no rocks are present in this habitat and the soil is of the namib form. the vegetation consists of short and low shrubs, with some short, high and tall shrubs as well as some low forbs, and low and short grasses present (table 2). the shrub stoebe cinerea and the grass ehrharta villosa var. maxima (species group m, table 1), are diagnostic for the community and chrysanthemoides monilifera (species group q) is the dominant species. the shrubs solanum africanum (species group l), stoebe cinerea (species group m), morella cordifolia (species group o), passerina rigida (species group p) and metalasia muricata (species group q) are locally prominent species. the community has an average of ten species per relevé. 3.1 chrysanthemoides monilifera rhus crenata short to high dune shrub sub-community the sub-community is found on north-facing coastal dune slopes with steep gradients of 26°–45°, sheltered from the south-eastern wind carrying salt spray. no rocks are found here (fig. 2). the vegetation consists of short to high shrubs, with a few tall shrubs. low forbs, low restios, low sedges, and low, short and high grasses have low cover values. the shrub rhus crenata (species group n, table 1) is the only diagnostic species. passerina rigida (species group p), chrysanthemoides monilifera and metalasia muricata (species group q) are dominant whereas the shrubs solanum africanum (species group l), stoebe cinerea (species group m), morella cordifolia (species group o), and the grass ehrharta villosa var. maxima (species group m) are also locally prominent in the sub-community. the sub-community contains an average of 12 species per relevé. 3.2 chrysanthemoides monilifera morella cordifolia low to short closed dune shrub sub-community the steep (16–>45°) southerly-facing midslopes dunes where this sub-community occurs are larger, more exposed areas than zietsman.qxd 2006/04/17 08:48 pm page 44 issn 0075-6458 45 koedoe 49/1 (2006) those where sub-community 3.1 occurs. the vegetation is exposed to sea mist (fig. 2). the leaf structure and orientation of the most dominant plant species, morella cordifolia, enhances the ability of the plant to catch moisture from sea fog. the vegetation consists of low to short shrubs, with some high and tall shrubs, as well as some low forbs, low restios, low sedges, and low, short and high grass. the shrubs morella cordifolia, lessertia frutescens (species group o, table 1), are dominant diagnostic species, and chrysanthemoides monilifera (species group q is also dominant. the shrubs stoebe cinerea (species group m), passerina rigida (species group p) and metalasia muricata (species group q), and the grass ehrharta villosa var. maxima (species group m) are locally prominent. the sub-community contains an average of 11 species per relevé. 4. ehrharta villosa var. maxima low to short closed dune grassland community the community is found on steep, exposed southern slopes of sparsely vegetated stable sand dunes, or the more mobile shifting sand dunes adjacent to the sea. (fig. 2). no rocks are present in the namib soil form. the community is found in a mosaic distribution pattern with communities 2, 3 and 5 and also in a mosaic distribution pattern with community 6 in the coastal beach area, adjacent to the sea. the vegetation is low to short dune grassland, with a few high grasses. (table 2). the grass ehrharta villosa var. maxima (species group m, table ) is the sole diagnostic and dominant species in the community. no other species occur here. 5. ammophila arenaria low to short closed dune grassland community the community occurs on the steep, dry and warm north-facing slopes of sparsely vegetated shifting sand dunes, in a mosaic distribution pattern with communities 2, 3 and 4 (fig. 2). the vegetation is low to short dune grassland. the shrub stratum has an average height of 1 m and an average canopy cover of 10 % and the grass stratum has an average height of 0.7 m and an average canopy cover of 44 % (table 2). the exotic grass ammophila arenaria (species group r, table 1) is the diagnostic and dominant species in the community. other species present are lessertia frutescens and senecio arenarius. the community contains an average of only three species per relevé. 6. arctotheca populifolia thinopyrum distichum short to low open beach community the community is situated on the beach and south-facing slopes of low fore dunes, directly adjacent to the sea, and also on gravel plains behind the low fore dunes (fig. 2). there are no rocks on the beach and dunes and 30–50 % rock cover on the gravel plain. the rocks on the gravel plain are mostly sandstone and quartzite roll-stones, deposited by the sea while the lea level was higher. the vegetation consists of short to low open beach grassveld with low forbs (table 2). the forb arctotheca populifolia and the grass thinopyrum distichum (species group s, table 2) are diagnostic and are the only species found in this community. the community contains an average of only two species per relevé. discussion the use of twinspan and the application of braun-blanquet procedures for refinement, were successful to classify the plant communities of this vegetation type. six plant communities, with eight sub-communities and four variants, were identified. the plant communities could all be related to specific environmental conditions and are therefore zietsman.qxd 2006/04/17 08:48 pm page 45 koedoe 49/1 (2006) 46 issn 0075-6458 floristically and ecologically distinguishable and interpretable. the thicket community (community 1) forms part of the dune thicket vegetation type of the thicket biome (low & rebelo 1998), whereas the other communities form part of the fynbos biome (low & rebelo). the thicket was mostly found in a mosaic with the vegetation of the inland plains and hills (zietsman & bredenkamp in press). the thicket community does not only differ from the fynbos communities concerning species composition and structure, but also in conservation importance. according to low & rebelo (1998) 14.49 % of dune thicket is currently being conserved, as opposed to the 1.42 % of south and southwest coast renosterveld and the 13.84 % of limestone fynbos (low & rebelo 1998). according to cowling & richardson (1995), if fire could be excluded from fynbos for a century or two, many of the landscapes would become densely populated with just a few species of thicket shrubs and trees. inland thicket development is therefore a feature of vegetation that has not been subjected to fire for some time. the species richness of this thicket is very low compared to those of fynbos and renosterveld (zietsman & bredenkamp in press) and the true thicket species like rhus glauca and euclea racemosa, are neither rare nor endemic. the limestone communities and the renosterveld community, within which the thicket patches are found, have high conservation priority. renosterveld has a very high conservation priority due to the small portion left, mainly because of agricultural land clearing (low & rebelo 1998; zietsman & bredenkamp in press). limestone fynbos is one of the most threatened vegetation types in the cape floristic region (hilton-taylor & le roux 1989). there are 110 plant species endemic to limestone outcrops. limestone fynbos also contains many species that are classified as rare (heydenrych 1994). thicket forming should be seen as a threat to the natural fynbos and renosterveld vegetation, and should not be encouraged above the conservation of the other fynbos and renosterveld vegetation. the presence of sub-community 1.5 (acacia cyclops euclea racemosa high totall closed thicket sub-community) is a reason for concern. acacia cyclops occurs scattered in various communities, fairly well distributed throughout andrew’s field and tsabatsaba nature reserve, but the larger stands found locally represent sub-community 1.5. all areas containing acacia cyclops, should be managed by implementing regular prescribed burning, and alien-invasive plant control, ideally by means of the cut-and-burn method (cowling 1992), but the current woodcutting method (promoting job provision), can also be followed. the conservation of limestone and renosterveld vegetation should be treated as a high priority. fire should be kept out of the dune area, because, according to tinley (1985) fire might cause an increase in dune instability. fire should also be kept out of areas that are heavily infested with the cape dune molerat. according to tinley (1985) fire would lead to an undesired increase in burrowing by cape dune molerats to obtain sufficient food (roots). therefore, as extensive dune molerat burrowing is already a problem in some areas in the reserve, fire should be kept out of those areas. other areas should be burned using a 15-year block-burn cycle to avoid the possible substitution of limestone and renosterveld vegetation by dense thickets. a single red data species (solanum africanum) is found quite abundantly in the dune area (communities 2 and 3). the presence of limestone endemic species in the dune area can be explained by the fact that both the dune sands and the limestone soil are alkaline. the shifting sand dune area should be protected from disturbances, like trampling and vehicle tracks. protection from disturbances should also provide sufficient protection to the red data and limestone endemic species. community 9 is dominated by ammophila arenaria, an exotic grass which is an european weed, commonly found on coastal dunes (goldblatt & manning 2000). although community 6 contains no rare or endemic species, the vegetation unit should zietsman.qxd 2006/04/17 08:48 pm page 46 issn 0075-6458 47 koedoe 49/1 (2006) be protected due to its uniqueness, and due to the fact that this gravel-plain is the breeding site of the rare damara tern sterna balaenarum (jeffery 1996). andrew’s field and tsaba-tsaba nature reserve is an area of great conservation significance. this area comprises an important part of the natural heritage found at the southern tip of africa, and should be protected and conserved for future generations. the resulting classification provides a useful tool, not only for the management of the plant communities of andrew’s field and tsaba-tsaba nature reserve, but also for similar vegetation areas, found in the surrounding region. acknowledgements mrs. f. siebert is thanked for her assistance during data processing. references behr, c.m. & g.j. bredenkamp. 1988. a phytosociological classification of thevegetation of the witwatersrand national botanic garden. south african journal of botany 54(6): 525–533. bredenkamp, g.j. & l.r. brown. 2001. vegetation – a reliable ecological basis for environmental planning. urban greenfile nov-dec 2001: 38–39. cowling, r.m. 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(south african national scientific programmes report; no. 109.) werger, m.j.a. 1974. on concepts and techniques applied in the zürich-montpellier method of vegetation survey. bothalia 11: 309–323. zietsman, m.m. & g.j. bredenkamp (in press). threatened limestone fynbos plant communities of andrew’s field and tsaba-tsaba nature reserve. bothalia. zietsman.qxd 2006/04/17 08:48 pm page 47 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjdffile false /createjobticket false /defaultrenderingintent /default /detectblends true /colorconversionstrategy /leavecolorunchanged /dothumbnails false /embedallfonts true 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setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice article information authors: benjamin j. wigley1,2 herve fritz2 corli coetsee1 william j. bond3 affiliations: 1school of natural resource management, nelson mandela metropolitan university, george campus, south africa2umr cnrs 5558 lbbe, university claude bernard lyon 1, france 3department of biological sciences, university of cape town, south africa correspondence to: benjamin wigley postal address: private bag x6531, george 6530, south africa dates: received: 07 may 2013 accepted: 18 sept. 2013 published: 11 mar. 2014 how to cite this article: wigley, b.j., fritz, h., coetsee, c. & bond, w.j., 2014, ‘herbivores shape woody plant communities in the kruger national park: lessons from three long-term exclosures’, koedoe 56(1), art. #1165, 12 pages. http://dx.doi.org/10.4102/ koedoe.v56i1.1165 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. herbivores shape woody plant communities in the kruger national park: lessons from three long-term exclosures in this original research... open access • abstract • introduction • research method and design    • study sites    • community composition    • trait sampling    • soil sampling    • statistical analyses • results    • community composition data    • population structure    • leaf traits of unique inside species versus common outside species    • chemical defence    • soil nutrients    • multivariate analyses • discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ the role of grazers in determining vegetation community compositions and structuring plant communities is well recognised in grassy systems. the role of browsers in affecting savanna woody plant communities is less clear. we used three long-term exclosures in the kruger national park to determine the effect of browsers on species compositions and population structures of woody communities. species assemblages, plant traits relating to browsing and soil nutrients were compared inside and outside of the exclosures. our results showed that browsers directly impact plant species distributions, densities and population structures by actively selecting for species with traits which make them desirable to browsers. species with high leaf nitrogen, low total phenolic content and low acid detergent lignin appeared to be favoured by herbivores and therefore tend to be rare outside of the exclosures. this study also suggested that browsers have important indirect effects on savanna functioning, as the reduction of woody cover can result in less litter of lower quality, which in turn can result in lower soil fertility. however, the magnitude of browser effects appeared to depend on inherent soil fertility and climate. conservation implications: browsers were shown to have significant impacts on plant communities. they have noticeable effects on local species diversity and population structure, as well as soil nutrients. these impacts are shown to be related to the underlying geology and climate. the effects of browsers on woody communities were shown to be greater in low rainfall, fertile areas compared to high rainfall, infertile soils. introduction top ↑ the role of herbivory in structuring plant communities and determining community compositions is well recognised (see reviews by augustine & mcnaughton 1998; huntly 1991). for example, native ungulate browsers have been found to have major impacts on shrub dynamics in east african savannas (augustine & mcnaughton 2004). different feeding behaviours and food preferences of browser species have major impacts on south african (owen-smith & cooper 1987) and west african savanna woody communities (jachmann & croes 1991). these impacts are often attributed to selective feeding related to food quality (owen-smith & cooper 1987). as in savannas (e.g. staver et al. 2009), herbivores have been found to regulate recruitment rate and species composition of trees in temperate forests (ammer 1996; kriebitzsch et al. 2000; van hees, kuiters & slim 1996) and grassland (anderson & briske 1995).many of the studies on browser–plant interactions in savannas have tended to focus on interactions between browsers and specific woody species. for example, coetzee et al. (2008), gadd (2002) and helm and witkowski (2012) looked at browser interactions with sclerocarya birrea, whilst fornara and du toit (2007) documented the response of acacia nigrescens to ungulate browsing. these two species are important browse species in southern african savannas. as savanna woody species tend to be highly variable in form and function, these studies usually do not provide a better understanding of savanna dynamics at the community level. augustine and mcnaughton (2004) emphasised the need for replicated experiments that assess the effects of native browsers on shrub dynamics in african savannas, as our understanding of the importance of browser effects on woody plant dynamics remains unclear. a better understanding of how browsers and woody plants interact at the community level and how these interactions affect savanna dynamics will be invaluable in managing browser populations. the existence of three long-term herbivore exclosures in the kruger national park, south africa, provides a good opportunity to empirically determine the role of browsers in determining woody species composition and population structures of woody communities growing at the three study sites. previous studies have shown evidence for taller canopies and higher tree and shrub densities inside exclosures at nhlangwini (asner et al. 2009; levick et al. 2010; trollope et al. 1998), makhohlola (levick et al. 2009; trollope et al. 1998) and n’waxitshumbe (asner et al. 2009; levick & rogers 2008; trollope et al. 1998). in these studies, the analyses were generally based on remote sensing and ignored species identity. differences in woody community compositions and plant traits between exclosure treatments are therefore still largely undocumented. differences in species compositions between treatments would be highly informative as the species found inside but not outside, are likely to be vulnerable to herbivory and therefore no longer found outside of the exclosures or found at reduced densities. the traits of these species can be used to provide us with clues as to what makes them vulnerable to herbivory. for example, díaz et al. (2007) found that at the global scale there is a spectrum of plant responses to herbivory with high levels of herbivory favouring annual over perennial plants, short plants over tall plants, prostrate over erect plants and stoloniferous plants over those with tussock architecture. this article aims to determine how browsers and plant traits interact and how these interactions affect woody plant population dynamics and community assemblages. we aim to determine if there are differences in species compositions and key plant functional traits between the woody communities found growing inside and outside the three long-term exclosures in the kruger national park (knp). plant traits are important in determining both the type of and degree of vertebrate herbivory, whilst they also can reflect plant responses to herbivory. we measured a number of plant traits that are likely to impact and respond to browsing; these included leaf nitrogen (n) and phosphorus (p), leaf carbon–nitrogen ratio (c:n), specific leaf area (sla), leaf size, leaf dry matter content (ldmc), leaf tensile strength, bite size index (bsi) and levels of secondary compounds. high leaf n and leaf p are associated with higher nutritional quality for herbivores (cornelissen et al. 2003). leaves with lower c:n ratios are much more attractive to herbivores as these leaves have higher n content (cornelissen et al. 2003). the sla of a species is usually a good correlate of its potential growth rate. lower sla values tend to correspond with relatively high investments in defences, both structural and chemical, and long leaf life span, all of which affect nutritional value (cornelissen et al. 2003). environmental nutrient stress and disturbance, as well as phylogenetic factors, play a role in determining leaf size (cornelissen et al. 2003). smaller leaves with higher sla are typically more attractive to herbivores. leaves with high ldmc values tend to be tough and therefore more resistant to disturbances such as herbivory or wind than leaves with low ldmc. species with low ldmc are usually associated with productive and highly disturbed areas (cornelissen et al. 2003). leaf tensile strength is a good indicator of the relative carbon investment in structural protection of the photosynthetic tissue. physically stronger leaves are better protected from abiotic (winds, storms) and biotic mechanical damage (herbivory). higher physical strength usually results in longer leaf span; however, this is usually combined with other physical and chemical defences such spines and tannins (cornelissen et al. 2003). bsi is a measure of how easily leaves can be browsed by a simulated browser (a human), whilst stem specific density broadly trades off against relative growth rate and stem defences against pathogens, herbivores and mechanical damage (cornelissen et al. 2003). condensed tannins and polyphenols are organic n-free chemical defences that bind with protein, reducing n availability to herbivores, thereby decreasing preference by mammal herbivores (bergvall & leimar 2005; haslam 1988; owen-smith 2005). the concentrations of the cell wall constituents (neutral detergent fibre [ndf], acid detergent fibre [adf] and acid detergent lignin [adl]) determine the digestibility of leaf material for herbivores. detergent analysis has been used to compare fibre digestibility across different plant types (e.g. see codron et al. 2007 and references therein). the following questions are specifically addressed using the three exclosures. firstly, has the long-term exclusion (> 40 years) of herbivores led to differences in woody plant species compositions at the three exclosures? secondly, do the structures, densities and abundances of the woody species populations differ inside and outside of the exclosures? thirdly, are there differences in plant leaf and stem traits between the species only found inside versus common species found both inside and outside? fourthly, has the long-term removal of herbivores from the exclosures had any effects on soil nutrient cycling? fifthly, do the exclosures help us to predict under what conditions herbivores play an important role in determining the structure and composition of woody plant communities in savannas? research method and design top ↑ study sites three long-term herbivore exclosures (> 40 years old) exist in the knp. these include the nhlangwini exclosure near pretoriuskop, the makhohlola exclosure near crocodile bridge and the n’waxitshumbe exclosure on the northern plains (table 1). at each exclosure two treatments were sampled, one inside the exclosure and one outside the exclosure at close proximity. table 1: site names, locations and descriptions of the exclosures in the kruger national park where sampling took place. the 220.0 ha nhlangwini exclosure was established in 1973 and occurs in the south-western portion of the knp near pretoriuskop rest camp. this is the wettest part of knp with mean average precipitation close to 750 mm. the exclosure is situated within the broad-leaved bushveld vegetation type that occurs on sandy soils derived from granitic rocks, common species include terminalia sericea and s. birrea (venter, scholes & eckhardt 2003). mean fire return interval is similar inside and out, with fire occurring on average every 3.6 years inside (asner et al. 2009) and less than every 4 years outside (smit et al. 2012). the makhohlola exclosure is a 2.4 ha exclosure constructed in the early 1970s. it is located in the south-eastern corner of the knp, just north of crocodile bridge. it occurs in the s. birrea–a. nigrescens savanna type (venter et al. 2003). the site occurs on soils derived from rocks of basaltic origin with mean annual rainfall of c. 600 mm. the mean fire return interval in the makhohlola exclosure (4.5 years) was slightly longer than outside (3.2 years), until 2004 when a new fire experiment was set up (see levick et al. 2009). since 2004, half of the exclosure has been protected from fire, whilst the other half is burnt in conjunction with the surrounding landscape (levick et al. 2009). from 2004 to the time of measurements in march 2011, the burn treatment and surrounding areas were burnt twice: in 2006 and 2010. the n’waxitshumbe exclosure is situated in the arid north-eastern section of the knp and was established in 1967. the original n’waxitshumbe exclosure was 230.0 ha, with an additional 72.0 ha added in 1986 (asner et al. 2009). mean annual precipitation is close to 400 mm and the soils here are derived from rocks of basaltic origin. the vegetation type is classified as colophospermum mopane shrubveld growing in broad-leaved bushveld (venter et al. 2003). the mean fire return interval for the n’waxitshumbe exclosure is approximately 4 years (asner et al. 2009), whilst smit et al. (2012) showed that the area surrounding the n’waxitshumbe exclosure has a mean fire return interval of less than 4 years. the faunal assemblage in the park includes all the major mammal species typical of the region. common browser species include kudu tragelaphus strepsiceros (pallas 1766), giraffe giraffa camelopardalis (linnaeus 1758), black rhinoceros diceros bicornis (linnaeus 1758), steenbok raphicerus campestris (thunberg 1811), grey duiker sylvicapra grimmia (linnaeus 1758) and bushbuck tragelaphus scriptus (pallas 1766). common mixed-feeders include impala aepyceros melampus (lichtenstein 1812) and the african elephant loxodonta africana (linnaeus 1758). community composition the distribution, abundance and density of the species growing both inside and outside of each of the exclosures was determined by walking four 100 m long by 4 m wide transects for each treatment at each site (1600 m2) in march 2011. the four transects in each treatment were taken parallel to each other and 50 m apart. inside and outside pairs were situated in as close a proximity as fences and firebreaks permitted. care was also taken to ensure that the paired transects were on similar positions along catenal gradients. at nhlangwini, the paired transects were taken perpendicular to the northern fence of the exclosure and parallel to the drainage in the area, outside transects were taken to the north-west of the exclosure. at makhohlola, the paired transects were taken perpendicular to the western fence and perpendicular to the drainage at the site, outside transects were taken to the north of the exclosure. owing to the limited size of the makhohlola exclosure (100 m × 240 m) two of the four transects were taken in the no-burn part of the fire experiment (since 2004) and two were taken in the part that burns in conjunction with the surrounding landscape. at n’waxitshumbe, the paired transects were taken perpendicular to the southern fence in the south-western portion of the exclosure and parallel to the drainage in that area. species identity was recorded for each woody plant encountered in each transect using a personal digital assistant (pda) with cybertracker software. the abundance data were used to create an inside abundance index, whereby the proportion of a species found in the inside compared to the outside treatment was calculated. this index was calculated by dividing the number of individuals of each species found inside by the total number of individuals found both inside and outside. thus a value of 1 indicates the species was only found inside, whilst a value of 0 indicates the species was only found outside. the inside abundance index was then plotted against a suite of leaf traits to test whether species growing on the inside had high values of these traits. this index can be used as a proxy to determine which plant species are preferentially selected for or ‘favoured’ by browsers relative to their abundance. trait sampling the dominant woody plant species making up 80% – 90% of the woody biomass inside and outside the three exclosures in the knp were determined (table 2). fourteen different plant traits relating to the leaves and stems were sampled for each species. all trait data (except bsi) were measured and recorded according to the methods explained in cornelissen et al. (2003). for each species, leaf material was collected from five individuals and analysed for n, p and carbon (c) content using a leco truspec cn analyser (leco corporation, st. joseph, mi). sla and average leaf area were measured using four healthy and complete sun-exposed leaves from five individuals (20 leaves per species). the bsi of a species is the total dry weight of the leaves removed from two human bites taken from five individuals (total of ten bites). an attempt was made to remove the maximum amount of leaf material with each bite. the bsi for all species was measured by the same person in order to control for potential differences between individual recorders. total condensed tannins were calculated according to the methods described by hattas and julkunen-tiitto (2012), whilst total polyphenols (tp) were calculated according to the methods described by hattas et al. (2005). cell wall constituents, that is, ndf and adf, were determined in an ankom fibre analyser (ankom technology corp., fairport, ny), whereas adl was determined in accordance with the acid detergent lignin in beakers method. ash was determined by incinerating the filter bag containing plant residue in a muffle furnace at 525 °c for 3 h. all cell wall constituents were determined consecutively as outlined in the ankom technology procedures (ankom technology 2010). table 2: dominant woody species accounting for > 80% of standing biomass for inside and outside treatments at the three kruger national park exclosures. soil sampling five replicate soil samples were collected both inside and outside of each exclosure at three depths (0 cm – 10 cm, 10 cm – 20 cm and 40 cm – 50 cm) using a soil auger. care was taken to sample away from the canopies of trees. soil bulk density measurements were taken at these same depths. the methods used for soil sampling and analyses are described in detail by wigley et al. (2013). statistical analyses all analyses were performed using r (r project 2012). chi-square tests were used to test for differences in tree densities between treatments. non-parametric wilcoxon tests were used to test for differences in trait measurements as the low replication resulting from the low species diversity at each site precluded the use of statistical tests founded on the normal distribution. analyses of variances were used to test for differences in soil nutrients between treatments using pooled data from all three depths, as the conditions for homoscedasticity were not violated (fligner test: p > 0.05; conover, johnson & johnson 1981). principle components analyses (pca) were used to explore the relationships between treatments, sites and measured plant traits. the function ‘pcasignificance’ available in the biodiversityr package (kindt & coe 2005) in r was used to test for significant pca axes using the broken-stick distribution. results top ↑ community composition data at the nhlangwini exclosure, there were a number of differences between the two communities. the four most common species for both treatments were t. sericea, dichrostachys cinerea, s. birrea and acacia gerrardii. however, t. sericea was found to be much more abundant (nearly 80% of all trees) inside compared to outside (c. 35%) of the exclosure (figure 1). dichrostachys cinerea showed the opposite pattern, with a higher relative abundance outside (30%) compared to inside (15%) the exclosure (figure 1). sclerocarya birrea and a. gerrardii showed similar abundances between treatments (figure 1). both treatments had a few unique species, with strychnos madagascariensis, catunaregam spinosa and searsia leptodictya only found inside, whilst antidesma venosum, combretum hereroense, albizia harveyi, philenoptera violacea and piliostigma thonningii were only found outside. species diversity was found to be higher on the outside of the exclosure, with a total of 16 species compared to 12 species inside. figure 1: the relative abundance of the species found inside and outside the three exclosures namely nhlangwini (a) inside and (b) outside, makhohlola (c) inside and (d) outside and n’waxitshumbe (e) inside and (f) outside, in the kruger national park. there were distinct differences in woody communities growing inside and outside of the makhohlola exclosure (figure 1). acacia nigrescens was found to be more abundant inside the exclosure (32%) but was still relatively abundant outside (22%). flueggea virosa was common inside (15%) but rare outside (4%). gymnosporia senegalensis was found to have a much higher abundance outside (33%) compared to inside (12%). lannea schweinfurthii and s. birrea had lower abundances outside (7% and 1%, respectively) compared to inside (12% and 7%, respectively). dichrostachys cinerea and a. harveyi were relatively common outside (13% and 9%, respectively) but rare inside (3% for both). dalbergia melanoxylon, ehretia rigida and euclea undulata were only found inside the exclosure, making species diversity higher on the inside (13 species) than outside (10 species). there were also distinct differences in the woody communities growing inside and outside of the n’waxitshumbe exclosure. at this site, c. mopane was found to be the most common species both inside (43%) and outside (40%, figure 1). the second most common species inside the exclosure (37%), d. melanoxylon, was not found outside of the exclosure. combretum imberbe and p. violacea were more common outside (14% for both) compared to inside (7% and 1%, respectively). sclerocarya birrea and g. senegalensis were relatively rare both inside and outside of the exclosures. dalbergia melanoxylon, grewia monticola and ozoroa obovata were unique to inside, whilst d. cinerea, a. harveyi and a. nigrescens were only found outside of the exclosure. thus species diversity was the same inside and outside of the exclosure, but species composition differed. population structure total woody plant densities were significantly higher for the inside treatments at all three sites (figure 2). these large differences were determined by the most common species growing at each site. the size class distributions for the two most common species growing at the nhlangwini exclosure (t. sericea and d. cinerea) show very different patterns for each treatment. the size class distribution of t. sericea inside the exclosure shows an inverse j-shaped curve, with many plants in the smaller size classes less than 3 m and fewer plants in the large size classes. the size class distribution for outside the exclosure shows much lower numbers of plants in the smaller size classes and fewer plants in the very large size classes (figure 3). a different pattern was evident for d. cinerea, with most plants falling into size classes between 2 m and 4 m inside the exclosure, whilst plants on the outside were more evenly distributed in a greater range of size classes (figure 3). figure 2: total tree densities for all woody plants greater than 0.5 m in height inside and outside the three long-term herbivore exclosures in the kruger national park. figure 3: size class distributions of terminalia sericea populations growing (a) inside and (b) outside and dichrostachys cinerea populations growing (c) inside and (d) outside of the nhlangwini exclosure in the kruger national park. at makhohlola, major differences were evident between treatments for each of the four most common species (figure 4). the size class distributions of l. schweinfurthii and a. nigrescens showed an approximately inverse j-shaped curve for the populations growing inside the exclosures but not for the outside populations. the inside populations of these two species both had a much higher number of individuals in the smaller (< 2 m) size classes than the outside populations (figure 4). the outside population of g. senegalensis was more uniform in height (1 m – 2 m) than the inside population (0 m – 3 m). there were many more and taller f. virosa plants growing on the inside of the exclosure compared to the outside (figure 4). figure 4: size class distributions of lannea schweinfurthii populations growing (a) inside and (b) outside, gymnosporia senegalensis populations growing (c) inside and (d) outside, flueggea virosa populations growing (e) inside and (f) outside and acacia nigrescens populations growing (g) inside and (h) outside of the makhohlola exclosure in the kruger national park. at n’waxitshumbe, the two most common species c. mopane and c. imberbe had similar size class distributions in each treatment (figure 5). however, there were higher densities in each size class for the inside treatment for c. mopane, whilst c. imberbe had similar numbers in each size class for both populations. the most striking difference was found for d. melanoxylon, where a population spanning the full range of size classes was found inside the exclosure, whilst not one individual was found outside of the exclosure in the sampling area (figure 5). figure 5: size class distributions of dalbergia melanoxylon populations growing (a) inside, combretum imberbe populations growing (b) inside and (c) outside and colophospermum mopane populations growing (d) inside and (e) outside of the n’waxitshumbe exclosure. no dalbergia melanoxylon plants were found growing on the outside of the exclosure in the kruger national park. leaf traits of unique inside species versus common outside species there were no significant differences (p > 0.05; wilcoxon test) in leaf nutrient concentrations – n, c, c:n, p, calcium (ca), magnesium (mg), sodium (na) and potassium (k) – between each of the inside and outside treatments (table 3). there were, however, some consistent trends; the lack of statistically significant differences was likely a result of the small sample sizes resulting from low woody species diversity at the sites. mean leaf n (%) was noticeably higher for the inside treatment at all three sites (table 3). the higher n and similar c concentrations inside of the nhlangwini exclosure resulted in a lower mean c:n ratio, whilst the other two sites had similar c:n ratios (table 3). leaf p (%) was similar between treatments at all three sites (table 3). table 3: mean (± s.e.) values of the measured leaf traits for the inside and outside treatments of three exclosures in the kruger national park. there were no significant differences (p > 0.05; wilcoxon test) in sla, leaf size, bsi and tensile strength between treatments. mean sla was consistently higher for species restricted to the inside treatments (table 3). average leaf area, which was highly correlated to bsi, were both higher inside the exclosures at the two nutrient rich sites on basalts (makhohlola and n’waxitshumbe), whilst the opposite pattern was true for the nhlangwini site which is situated on nutrient poor granite derived soils (table 3). mean leaf tensile strength was higher outside exclosures at the two basalt sites, but no difference was evident between treatments at the granite site (table 3).the pooled data showing the inside abundance index of each species plotted against leaf traits and bsi showed a weak positive relationship between the inside abundance index and leaf n and p (figures 6a and 6b). almost no relationship was evident between the inside abundance index and sla and bsi (figures 6c and 6d; p > 0.05 for all correlations). figure 6: data for, (a) leaf nitrogen, (b) leaf phosphorus, (c) specific leaf area and (d) bite size index plotted against the proportion of species found on the inside compared to the outside of three exclosures in the kruger national park. a value of 1 indicates that the species was only found inside, whilst a value of 0 indicates the species was only found outside the exclosures. chemical defence no significant differences (p > 0.05; wilcoxon test) in total phenolics, condensed tannins and fibre were evident between treatments. however, an interesting pattern emerged; mean leaf phenolic concentrations were consistently lower for species restricted to the inside treatments compared to those in the outside treatment, with the biggest difference evident at the nhlangwini site (table 3). mean condensed tannin concentrations showed no consistent patterns between treatments and sites (table 3). the three types of fibre analyses showed different patterns. ndf, which includes hemicellulose, cellulose and lignin, was slightly lower for the outside treatments at makhohlola and n’waxitshumbe, whilst the opposite pattern was found at nhlangwini (table 3). adf, which includes cellulose and lignin, showed no consistent patterns (table 3). adl, which only includes lignin, was consistently lower for the inside treatments at all three sites (table 3). soil nutrients although the pooled data for all depths of soil n and p showed a pattern of higher values for the three inside treatments, the only statistically significant differences were evident at n’waxitshumbe (p < 0.05; figures 7a and 7c). soil c and k also showed a pattern of being consistently higher for the inside treatments at makhohlola and n’waxitshumbe, with no differences evident at nhlangwini (figures 7b–7d). similar patterns were evident for n, c and p when total nutrient stocks were calculated using soil bulk density data (data not shown). figure 7: mean (± s.e.) for, (a) soil nitrogen, (b) carbon, (c) phosphorus and (d) potassium concentrations inside and outside of three exclosures in the kruger national park. plotted means are calculated from five replicates taken at three depths. multivariate analyses the pca performed on the data using 13 traits relating to the leaves and stem (figure 8) showed that the first two principal components together explained 44.0% of the total variation amongst species. the most important traits separating species along the first pca axis were ndf, adl and tp. the most important traits separating species along the second pca axis were leaf n, leaf c:n and stem density (figure 8). the pca showed that the species from the inside tended to more variable along pca axis 2 (higher leaf n and stem density), whilst the species from the outside tend to be more variable along pca axis 1 (higher fibre and tp values). the variance explained by the first pca axis was found to be insignificant, as the percentage of explained variance (24.3%) was not larger than the corresponding percentage of variance of the broken-stick distribution (24.5%), whilst the second pca axis was significant as the percentage of explained variance (20.3%) was larger than the corresponding percentage of variance of the broken-stick (16.8%) distribution (kindt & coe 2005). figure 8: principal components analysis of trait measurements from 20 species sampled inside and outside of three exclosures in the kruger national park. discussion top ↑ clear differences were found in both the species composition and population structure of the woody plant communities growing inside and outside of three herbivore exclosures in the knp. although the three exclosures were situated in very different vegetation types, some similarities exist in species’ responses to low herbivory. some species were frequently encountered both inside and outside of the exclosures (e.g. t. sericea and d. cinerea at nhlangwini, a. nigrescens and g. senegalensis at makhohlola, c. mopane at n’waxitshumbe). smaller size classes tended to be better represented inside, and larger size classes outside, for species which were found in large numbers both inside and outside of the exclosures. there were some exceptions though; g. senegalensis for instance, had similar population structures both inside and out. this could be because this species is not a preferred browsing species as a result of the high condensed tannin and adl concentrations in the leaves (see shrader et al. 2011). we argue that savanna woody species are either well adapted to browsing (such as d. cinerea), well defended (such as g. senegalensis) or both (such as a. nigrescens). for example, fornara and du toit (2007) showed that heavily browsed a. nigrescens trees develop tolerance traits, such as high regrowth rates and extensive branching, as well as resistance traits, such as close thorn spacing, suggesting that it is both well adapted and well defended. in fact, a meta-analysis by carmona, lajeunesse and johnson (2011) has shown that secondary metabolites are less important in plant defence when compared to other defensive plant traits such as physical resistance, gross morphology and growth rates. some species were found more frequently outside of the exclosures, such as d. cinerea, which was found in higher numbers outside of the exclosure at nhlangwini, and a. harveyi at all exclosures. these species are most likely stimulated by disturbance, for example d. cinerea shrubs rootsucker, and expand significantly when stimulated by fire and browsing (wakeling & bond 2007). more frequently, species were only found inside the exclosures (e.g. d. melanoxylon, g. monticola and o. obovata at n’waxitshumbe). dalbergia melanoxylon had the highest leaf n of all the species in this study and we assume that the species found only on the inside of the exclosures are mostly very palatable and targeted by browser species outside of the exclosures. dalbergia melanoxylon was found outside of the n’waxitshumbe exclosure in a previous study, but in lower lying areas than at our study area (levick & rogers 2008). it is uncertain whether d. melanoxylon has since disappeared outside the exclosure or whether this species may be preferentially targeted by browsers at specific positions along the catena. why do species become locally extinct outside of exclosures? impala and kudu are highly selective when foraging in nutrient-poor broadleaf savannas (cooper & owen-smith 1985; owen-smith & cooper 1987). this selectivity is attributed to differences amongst species in the level of defensive chemical compounds. alternatively, these species may also be predisposed to pollarding, uprooting and ringbarking and coppice poorly in response to elephant damage (o’connor, goodman & clegg 2007). a number of studies have documented the local extinction of plant species resulting from elephant impact (e.g. boundja & midgley 2010; o’connor et al. 2007 and references therein; penzhorn, robbertse & olivier 1974). several studies have shown that other herbivores can lead to the local extirpation of plant species. bond and loffel (2001) found that acacia davyi was no longer found in areas accessible to giraffe in ithala game reserve. total tree densities were significantly higher for the inside treatments at all three sites, usually with more individuals falling into the smaller size classes inside of the exclosures. these findings are in strong agreement with previous studies from the same study sites. trollope et al. (1998), levick and rogers (2008) and asner et al. (2009) reported higher tree densities and different population structures between treatments. moncrieff et al. (2011) reported different tree allometries between trees growing inside and outside of the same herbivore exclosures in the knp, with heavily browsed trees shorter for a given stem diameter. our results are therefore in agreement with previous studies and provide strong evidence that herbivory has a major impact on savanna woody plant diversity, species population structure and overall woody plant densities. the next step was to determine if there were differences between treatments for some of the key plant functional traits of each woody community. there was a trend in leaf n concentration, with higher mean n for the species unique to the inside treatments compared to the species found growing outside. no obvious patterns or trends were evident for leaf c:n or p. leaf structural traits, sla, average leaf size and bsi, which are all correlated, showed similar trends with higher values inside the exclosures at the two fertile sites and lower values inside compared to outside at the infertile site. the opposite pattern was evident for tensile strength where the leaves were generally tougher on the outside at the two fertile sites with no difference at the infertile site. apart from leaf n, total phenolic content and lignin content were the only traits that showed consistent patterns between treatments with higher values for species outside of the exclosures. increases of carbon-based structural metabolites with decreases in leaf n have been shown with high browsing intensity in previous work (bryant et al. 1991; wessels, van der waal & de boer 2007). an alternative theory for increases in lignin and leaf n inside the exclosures may be that herbivores preferentially select species with higher leaf n and that species with lower total phenolic and lignin content are also targeted by herbivores. in general, the removal of browsers led to higher soil nutrients, the response was the greatest at the two basalt sites. at n’waxitshumbe differences in soil n and p were higher with browsers absent, whilst soil c and k were also higher inside of the exclosures. the same pattern was evident at makhohlola, whilst the nutrient-poor site (nhlangwini) showed almost no differences between treatments for all measured soil nutrients. although herbivores have been found to increase soil nutrients (e.g. augustine & mcnaughton 2006; frank 2008), ritchie, tilman and knops (1998) found that herbivores can actually depress soil n by indirectly decelerating n cycling through decreasing the abundance of plant species with n-rich tissues. the increased soil fertility inside of the exclosures could possibly be explained by differing litter dynamics. the expectation would be that leaf litter with higher n and lower fibre content would result in faster cycling and improved soil fertility (e.g. reich et al. 2001). furthermore, the higher density of woody plants would increase the amount of leaf litter reaching the soil. thus the higher soil nutrients found inside of the exclosures could possibly be a result of the indirect effect of herbivores, as their removal resulted in higher tree densities. similarly, fire indirectly affects nitrogen cycling in the knp by decreasing woody densities (coetsee, bond & february 2010). the similar soil nutrient concentrations found inside and outside of the nhlangwini exclosure can most likely be attributed to the overall low litter quality and lower densities of herbivores resulting from the nutrient poor granitic soils at this site. conclusion top ↑ this study has demonstrated the important role browsers play in savanna dynamics. browsers directly impact species distributions, densities and population structures by actively selecting for species with favourable traits; this study suggests that species with high leaf n, low total phenolic content and low adl are favoured. forty years of no browsing impact was insufficient to allow the recruitment of the dominant species into larger size classes. this supports the idea that cohorts of trees recruit simultaneously, they would however need a break from both herbivory and fire. this study also suggests that browsers have important indirect effects on savanna functioning through their impact on soil nutrient cycling. the magnitude of these indirect effects on soil nutrient cycling appears to depend on inherent soil fertility and climate. the effect of browsers on both vegetation and soils was highest at the low rainfall site with high soil fertility, with a somewhat lesser effect at the higher rainfall basalt site, with almost no effect at the infertile high rainfall granite site. this work highlights the importance of herbivore exclosures in protected areas and some of the challenges faced by researchers when studying browser effects on savanna ecosystems because of the paucity of replicated long-term herbivory exclosures. future work should identify other possible sites in savannas from around the globe that could be used to corroborate the findings from this study. acknowledgements top ↑ this work formed part of the phd thesis of b.j. wigley and was made possible thanks to funding from a bdi-ped grant from the french national centre for scientific research, as well as the andrew w. mellon foundation. the authors are hugely grateful to: sanparks, kruger national park scientific services for allowing the study and logistical support, and scientific services game guards for protection in the field. thanks to two anonymous reviewers for their helpful comments which helped improve this manuscript. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions b.j.w. 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141–147. http://dx.doi.org/10.2989/ajrfs.2007.24.3.4.297 wigley, b.j., coetsee, c., hartshorn, a.s. & bond, w.j., 2013, ‘what do ecologists miss by not digging deep enough? insights and methodological guidelines for assessing soil fertility status in ecological studies’, acta oecologica 51, 17–27. http://dx.doi.org/10.1016/j.actao.2013.05.007 abstract introduction research method and design results discussion conclusion acknowledgements references appendix 1: methods and materials used for soil analyses appendix 2: braun–blanquet scale for vegetation cover appendix 3: ant species lists for various data sets appendix 4: tables illustrating sampling completeness calculated for the rehabilitated and control sites appendix 5: list of indicator values about the author(s) samantha-leigh jamison department of plant science, university of pretoria, south africa mark robertson centre for invasion biology, department or zoology & entomology, university of pretoria, south africa ian engelbrecht department of zoology and entomology, university of pretoria, south africa peter hawkes department of zoology and entomology, university of pretoria, south africa afribugs cc, pretoria, south africa citation jamison, s-l., robertson, m., engelbrecht, i. & hawkes, p., 2016, ‘an assessment of rehabilitation success in an african grassland using ants as bioindicators’, koedoe 58(1), a1383. http://dx.doi.org/10.4102/koedoe.v58i1.1383 original research an assessment of rehabilitation success in an african grassland using ants as bioindicators samantha-leigh jamison, mark robertson, ian engelbrecht, peter hawkes received: 02 feb. 2016; accepted: 23 june 2016; published: 29 sept. 2016 copyright: © 2016. the author(s). licensee: aosis. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. abstract many studies that evaluate rehabilitation make use of invertebrate bioindicators. invertebrates, especially ants, make useful indicators as they are sensitive to environmental change. we compared ant assemblages in rehabilitated and control sites in the rietvlei nature reserve, a protected area important for grassland conservation in south africa. pitfall traps were used to sample ant assemblages at six control sites and six rehabilitated sites. in addition, environmental and vegetation surveys were conducted at each site. we found that the ant assemblages differed significantly between the control and rehabilitated sites, although there was considerable overlap; the control sites supported a greater species density and higher abundance of ants than the rehabilitated sites. in total, 36 ant species were collected (control sites: 34 species; rehabilitated sites: 26 species). the environmental survey revealed that percentages of bare ground and coarse sand, as well as soil ph, differed significantly between the control and rehabilitated sites. the control and rehabilitated sites also supported significantly different plant assemblages. three indicator ant species were identified for the control sites: crematogaster rectinota, crematogaster amita and monomorium fastidium. no indicator species were identified for the rehabilitated sites. these results suggest that recovery from the previous agricultural use of the area is still incomplete and highlights the lack of research examining the success of rehabilitation in the grassland biome. conservation implications: the present study illustrates the need for further research on rehabilitation techniques utilised in the grassland biome. this is of value as the remainder of south african grasslands are considered critically endangered. introduction rehabilitation and the need for bioindicators increasing human impacts on the environment have resulted in widespread degradation of ecosystems (hobbs & norton 1996). as a result, there is a global drive to rehabilitate terrestrial and freshwater environments (lake 2001). typically, rehabilitation projects monitor aspects such as vegetation structure, species diversity and richness as well as nutrient cycling (ruiz-jaén & aide 2005; van aarde & smit 1997). a number of rehabilitation studies make use of biological indicators. these indicators provide a means to monitor and assess the state of an ecosystem over time (cairns, mccormick & niederlehner 1993; morellet et al. 2007). however, in order to be practical, these bioindicators need to meet certain criteria. they need to be sensitive enough to provide an early warning in the event of a disturbance, they should have a wide spatial distribution and they must provide a continuous assessment for several disturbance factors (noss 1990). such indicators can aid us in understanding the rehabilitation of previously disturbed land and prove valuable for future management projects. ants as bioindicators invertebrates are potentially valuable indicators for monitoring the success of rehabilitation (andersen & sparling 1997). in particular, ants (hymenoptera: formicidae) have received much attention as ecological bioindicators. they are widespread, abundant, have a manageable species richness and perform many keystone functions and thus have the potential to be used for the assessment of rehabilitation (fagan et al. 2010; van hamburg et al. 2004). they are important ecosystem engineers that are responsible for turning soil, seed dispersal, forming mutualistic relationships and are among the top predators of other invertebrates (hölldobler & wilson 1990; lach, parr & abott 2010). as they are colonial, their response to disturbance may be very different from other invertebrates that are more susceptible to disturbance events (lach et al. 2010). the removal of individuals, resulting from mortality caused by disturbance, may not lead to the eradication of the colony from the habitat. thus, the response of ants to disturbance differs from that of other terrestrial plants and animals that may become locally extinct after a disturbance event (andersen 2000; lach et al. 2010). in response to disturbance, such as agriculture or mining activities, ant communities may show changes in species composition and interspecific interactions, as well as loss of taxonomic diversity and variation in the provision of ecosystem services (lach et al. 2010). ants have been used as bioindicators in several projects involving ecosystem management, land rehabilitation and monitoring the degree of land degradation, as well as investigating the conservation status of various zones of land (majer 1983). they have been successfully used as bioindicators of mine site rehabilitation in northern australia (andersen, hoffman & somes 2003) and on rehabilitated ash dams associated with coal-fired power stations around the highveld of south africa (van hamburg et al. 2004). they have also been used to indicate pesticide contamination in cotton-growing areas in australia (weir 1978). ants thus provide an important indication of the biotic and abiotic state of an ecosystem and ultimately reveal the impacts that various environmental changes can have on an ecosystem (hodkinson & jackons 2005). the importance of grasslands grasslands are regions of global importance and provide a series of important ecosystem services. these include the purification of water, flood attenuation and nutrient cycling, as well as carbon sequestration and storage (south african national biodiversity institute [sanbi] 2013). the south african grassland biome covers nearly 30% of the country’s land surface and is home to a large majority of the country’s population and several endangered and endemic species (neke & du plessis 2004; sanbi 2013). increasing urbanisation and development has resulted in a substantial amount of land degradation in the south african grassland biome, resulting in its classification as critically endangered (neke & du plessis 2004). despite this, grasslands throughout the country have received little conservation protection (neke & du plessis 2004). despite the necessity to rehabilitate grasslands following disturbances such as mining or agriculture, there appears to be limited knowledge on grassland rehabilitation (zaloumis & bond 2010). as rehabilitation may take many years, there is a need to quantify the extent to which ecosystems (such as grasslands) have been rehabilitated (legg & nagy 2006). bioindicators such as ants can fulfil this purpose. however, to date surprisingly little research has been done in south africa on ant diversity in grasslands and how ant communities resemble each other following a disturbance. this study investigated the success of the rehabilitation measures applied in an old agricultural land area on rietvlei nature reserve. we compared ant assemblages in areas where rehabilitation measures were applied and in nearby untransformed, natural grassland areas as an indication of the effectiveness of rehabilitation. we also aimed to identify whether any indicator species were present in the rehabilitating and untransformed (control) sites of the reserve. research method and design study area the study took place on the southern region of rietvlei nature reserve (between 25°50’s, 28°15’e and 25°56’s, 28°19’e), a protected area important for grassland conservation in south africa. the reserve covers an area of approximately 3870 ha. it is situated in close proximity to urban centres. the climate of the area is characterised by warm, wet summers that are followed by cold, dry winters, when frost is prevalent. summer temperatures may reach highs of 34 °c, which are contrasted by the low winter temperatures that can fall to –2 °c (marais 2004). the area has a mean annual rainfall of approximately 724 mm (1970–1999; marais 2004). the plant composition of the reserve is typical of the grassland biome; the vegetation type of the area is described as rand highveld grassland (mucina & rutherford 2006). around 23 years ago, the reserve extended its southern boundary and incorporated land that was previously used for agriculture. rehabilitation measures were employed on the land in an attempt to improve species diversity and ecological function. the rehabilitated areas were ploughed and sown with a mix of indigenous grass seeds (r. marais 2014, pers. comm., rietvlei nature reserve, pretoria). procedure ant sampling sampling was carried out at 12 sites, 6 located in the rehabilitated area and 6 in untransformed (control) areas of the reserve. all sites were at a minimum distance of 200 m from old land boundaries to avoid edge effects and 300 m from other sampling sites to ensure independent sampling. at each site, 10 plastic pitfall traps (55 mm in diameter and 70 mm deep) were arranged in two rows of five, positioned 10 m apart. the study was designed in such a manner so as to avoid or minimise the possibility of pseudoreplication that may have been introduced by having a single larger array with many traps and treating each array (rather than each trap) as a replicate. the traps were placed in the soil with the rim flush with the soil surface and partially filled with a mixture of water and propylene glycol. this non-toxic solution ensures the successful preservation of the captured specimens. a rain cover (20-cm plastic lid with four wire legs) was placed above each trap to avoid flooding. the traps were left in place for 3 days from 08 to 11 april 2014. the ant specimens found in each trap were removed and identified to species level where possible and their abundances recorded. ants were identified to genus using bolton (1994) and then to species where possible, using the most recent available keys to the genera concerned: for agraulomyrmex, prins (1983); anoplolepis, prins (1982); crematogaster, arnold (1920); lepisiota, arnold (1920); leptogenys, bolton (1975); meranoplus, bolton (1981); monomorium, bolton (1987); technomyrmex, bolton (2007) and tetramorium, bolton (1980). for some genera, such as camponotus, carebara, dorylus, pheidole, plagiolepis and solenopsis, no keys are presently available for the afrotropical fauna; for these genera, as well as for species in other genera that could not be identified using the available keys, specimens were identified by comparison with reference material in the afribugs collection (afrc). where no formal name could be assigned, a morphospecies code matched to voucher material in the afrc collection was applied. these codes are globally unique and images of most are available on antweb (http://www.antweb.org) to allow cross-referencing between studies. all identifications were performed by s.l. jamison under the guidance of p.g. hawkes. a specimen of each of the species identified was mounted and labelled to produce a reference collection, which is housed at the department of zoology & entomology, university of pretoria. environmental factors estimates of the percentage bare ground, rock, grass, shrub cover and height of the tallest plant were recorded in five 1-m2 quadrats at each site. these quadrats were randomly selected from the immediate area around each of the sites. the quadrats were placed within a 10-m perimeter around the pitfall traps in order to establish the best estimate of environmental factors within each of the respective sites. the braun–blanquet cover-abundance scale was used to quantify the cover of all plant species in three 1-m2 quadrats at each site. a disk pasture meter was used to obtain measures of above-ground vegetation biomass. a total of 20 disk pasture meter height measurements were recorded at random at each site. we refer to these height measurements as a biomass index from here onwards. a soil auger was used to collect a soil sample from the first 20 cm of soil at each of the sites. soil was randomly sampled within a 10-m perimeter around the pitfall traps. three soil samples were randomly taken from each of the 12 sites. the three soil samples collected at each site were then combined and used as one sample to get an indication of the average soil conditions at each site. the percentage coarse sand, sand, silt and clay content of the soil as well as the soil ph and soil organic matter were determined using standard methods (appendix 1). all soil analyses were conducted at the university of pretoria in the soil sciences department. analysis species diversity indices for the ant data were generated using primer 5.2.0 software package (clarke & warwick 2001). the indices included the pielou (pielou 1969) and shannon–weiner diversity indices (shannon 2001). these diversity indices were compared between the rehabilitated and control sites with welch two-sample t-tests (welch 1947). furthermore, welch two-sample t-tests were used to test for significant differences between the number of ant species and the number of individuals found between rehabilitated and control sites. species richness in the two sites was compared using rarefaction curves with 95% confidence intervals derived from unconditional variance estimates (colwell et al. 2012). richness estimators were generated using estimate-s (v 9.1.0; colwell 2013). we used three non-parametric species richness estimators and one parametric estimator in our analysis to correct for sampling bias. the non-parametric estimators were the incidence-based coverage estimator (ice), the chao 2 abundance-based richness estimator (chao 2) and the second-order jackknife richness estimator (jack 2). we used the michaelis–menton mean (mmmean) as the parametric estimator. we chose this suite of estimators as estimator performance varies depending on a range of factors, and no single estimator is universally accepted as superior to others (smith & jones 2005). sampling was considered adequate if the sample-based rarefaction curves and the curves of the species richness estimators converged closely at the highest observed values (cardoso 2009; cardoso et al. 2009; longino, coddington & colwell 2002; magurran 2004). observed richness as a percentage of estimated richness was used as a measure of inventory completeness (jiménez-valverde et al. 2010). the mean sampling completeness was calculated for the control and rehabilitated sites by averaging the completeness calculated for each of the different species richness estimators. species richness was compared between the rehabilitated and control sites by plotting rarefaction curves with their 95% confidence intervals. if the intervals overlapped, the differences between the treatments were taken to be not significant (colwell et al. 2012). an analysis of similarity (anosim) and non-metric multidimensional scaling (nmds) were performed to compare the ant assemblages in the rehabilitated and control areas. the similarity matrix was calculated using a bray–curtis similarity measure and a fourth-root transformation in primer 5.2.0 (clarke & warwick 2001). the fourth-root transformation reduces the contribution of very abundant species (french & major 2001). anosim generates a global r statistic that provides an indication of average dissimilarity between the assemblages being compared. values closer to 1 indicate distinct differences, while values closer to 0 indicate high levels of similarity in species composition. to assess the contribution of different species to the differences between the rehabilitated and control sites, the similarity percentages for species contributions were applied to the data (simper, primer 5.2.0). a fourth-root transformation was again used to reduce the influence of dominant species. characteristic ant species (indicator species) were identified for each of the sites with the use of the indicator values method (dufrêne & legendre 1997) in the package labdsv (roberts 2014), run in r 3.1.1 (r development core team 2011). an analysis of the relative indicator values (indval) was performed to determine the specificity (uniqueness to specific sites) and fidelity (rate of recurrence within each site) of each species for a particular site and treatment (e.g. rehabilitating or control). species that were found to have a value greater than 70%, that is, species that occur predominantly in a particular habitat and occur frequently in that habitat were considered as reliable indicator species for the different sampling units. rank abundance curves were also generated and allowed comparison of species composition between control and rehabilitated sites. the values for percentage of bare ground, rock, grass, shrub and height of tallest plant were compared between control and rehabilitated sites using welch’s two-sample t-tests, which were also used to compare the percentages of coarse sand, sand, silt and clay, as well as soil ph and soil organic matter. anosim and nmds were performed to compare the plant assemblages in the rehabilitated and control sites. the similarity matrix was calculated using a bray–curtis similarity measure and a fourth-root transformation in primer 5.2.0 (clarke & warwick 2001). the braun–blanquet scale was rescaled to range from 1 to 7 for the purpose of the anosim and nmds (table 1-a2). results ants in total, 3206 individuals representing 36 species were collected during the study (table 1-a3). the control sites had a significantly higher species density (t = 6.618, p < 0.001) and abundance (t = 4.674, p < 0.001) than the rehabilitated sites (figure 1). additionally, the control sites had a significantly higher pielou evenness index (t = 3.125, p < 0.05) and shannon–weiner index (t = 6.789, p < 0.001) than the rehabilitated sites (figure 1). in total, 34 ant species were collected from the control sites, whereas only 26 species were collected from the rehabilitated sites. of the 36 species collected in this study, 24 species (66.7%) were present in both the control and rehabilitated sites. in total, 10 species (27.8%) were unique to the control sites, whereas only 2 species (5.6%) were unique to the rehabilitated sites (table 1-a3). figure 1: box plots of ant diversity in the control and rehabilitated sites on rietvlei nature reserve, showing (a) the number of species, (b) the number of individuals, (c) the pielou evenness (j’) and (d) the shannon–weiner index (h’). the estimated species richness (sest) for the ant samples collected from the control and rehabilitated sites did not converge closely with the richness estimators (figure 1-a1). the mean sampling completeness for the rehabilitated sites was found to be 65.70%, whereas the average sampling completeness for the control sites was 70.31% (from table 1-a4 to table 5-a4). however, as it is unlikely that we will ever reach an asymptotic endpoint, particularly for ants and other invertebrates (gotelli et al. 2014), this level of sampling completeness is likely sufficient to draw conclusions about grassland rehabilitation. the confidence intervals on the rarefaction curves for the rehabilitated and control sites overlapped, indicating that there were no significant differences in species richness (figure 2). figure 2: rarefaction curves with 95% confidence intervals comparing species richness in the control and rehabilitated (rehab) sites. species richness should be compared for equal numbers of species in each treatment. anosim indicated that ant assemblages differed significantly between the treatments (global r = 0.507, p = 0.002). the nmds plot revealed that there was some separation between the control and rehabilitated sites, although some overlap is still evident (figure 3). the replicates representing the control sites (c) are clustered together at the top of the plot indicating smaller assemblage differences between these replicates than for the rehabilitated sites. the replicates representing four of the rehabilitated sites (1r, 3r, 5r and 6r) are clustered together in the lower right quadrant of the plot. they are clustered less tightly than the control sites at the top of the plot. this indicates fairly large assemblage differences between the replicates within the rehabilitated sites compared with those of the control sites. in particular, sites 1r and 6r are plotted in close proximity to the control sites indicating that the ant assemblages at these rehabilitated sites are similar to those of the control sites. sites 2r and 4r are isolated from the clusters on either side of the plot indicating that the ant assemblages found at these sites are different from those found at the other sites of both treatment types. it is interesting that 2c is located near the rehabilitated sites within the nmds plot. the potential impact of human-mediated activities could have influenced the habitat at this site. however, we have no information regarding the use of the area before the establishment of the reserve and cannot give a fully informed explanation. figure 3: non-metric multidimensional scaling plot indicating the similarity of ant assemblages among replicates of control and rehabilitated sites. the assemblage differences are further supported by the rank abundance curves observed for the different treatments (figure 4). these rank abundance curves show only species that were represented by more than five individuals. the three most abundant species found in the control sites were monomorium albopilosum emery (641 individuals), meranoplus peringueyi emery (209 individuals) and tetramorium bothae forel (193 individuals). in comparison, the three most abundant species found in the rehabilitated sites included m. albopilosum (411 individuals), tetramorium setuliferum emery (278 individuals) and pheidole megacephala fabricius (177 individuals). although many of the highly abundant species were present in both the rehabilitated and control sites, the control sites had more unique species than the rehabilitated sites (control: 10 unique species, rehabilitated: 2 unique species; table 1-a3). the criteria needed to identify indicator species (indval > 70%) were fulfilled for three species present in the control sites (table 1-a5). these species include crematogaster rectinota forel (indval = 0.932, p = 0.004), crematogaster amita forel (indval = 1, p = 0.003) and monomorium fastidium bolton (indval = 0.808, p = 0.031). no indicator species were identified for the rehabilitated sites. the results from the rank abundance curves are further supported by the simper results, which found c. amita to have the greatest contribution to the separation of the control and rehabilitated sites (6.31% of total abundance; 51 individuals in total for the control sites), followed by m. peringueyi (5.67%; 209 individuals in control sites) and lastly m. fastidium (5.53%; 167 individuals across control and rehabilitated sites). although m. albopilosum had a total of 1052 individuals for both the control and rehabilitated sites, the overall contribution of this species to the separation of sites was only 4.27%. the contribution of the other species was greater because of the difference in average abundance of the species between the sites. the mean abundance of m. albopilosum was 106.83 in the control sites and 68.50 in the rehabilitated sites. figure 4: rank abundance plot for the rehabilitated and control grassland sites on rietvlei nature reserve. environmental variables the mean percentage of bare ground was significantly higher in the rehabilitated than the control sites (t = -3.193, p = 0.003; figure 1-a2). there were no significant differences in the percentage cover of grass, rock, height of tallest plant and biomass index between the rehabilitated and control sites. the ph of the soil (t = -2.754, p = 0.021) and the percentage coarse sand (t = 2.275, p = 0.0495) differed significantly between the rehabilitated and control sites (figure 1-a3). however, the t-tests revealed that there was no significant difference in soil organic matter and percentage of sand, silt and clay content between the rehabilitated and control sites. vegetation assemblages anosim indicated that the plant assemblages differed significantly between the treatments (global r = 0.712, p = 0.001). the global r statistic indicates fairly large assemblage differences between the control and rehabilitated sites. the nmds plot illustrates that there is a clear distinction between the control and rehabilitated sites (figure 1-a4). there is a larger amount of variation in the plant communities between the various control sites than there is between the rehabilitated sites. it can be seen that the plant community of the control and rehabilitated sites is still very different, 23 years after rehabilitation. discussion this study found that the control sites on the rietvlei nature reserve had significantly higher ant species density and abundance than the rehabilitated sites. in total, 34 ant species were collected from the control sites, whereas only 26 species were collected from the rehabilitated sites. moreover, the control sites had a significantly higher pielou evenness index and shannon–weiner diversity index than the rehabilitated sites. however, species richness was not significantly different between control and rehabilitated sites. there was a significant difference in the ant assemblages between the rehabilitated and control sites, although some convergence was evident. influence of dominant ant species of the 36 ant species collected in this study, 66.7% were present in both the control and rehabilitated sites (i.e. 24 species shared). however, a large portion of the difference between ant assemblages of the rehabilitated and control sites was because of the higher abundance of individuals present in the control sites (control: 1954 individuals, rehabilitated: 1252 individuals). the difference between ant assemblages may thus be because of a combination of the abundance of individuals and the composition of species present in the control and rehabilitated sites. in particular, two ant species (t. setuliferum and p. megacephala) had higher abundances in the rehabilitated sites than in the control sites, though neither species was identified as an indicator. pheidole megacephala and several species of tetramorium and monomorium are said to be characteristic of disturbed areas (andersen 2003; samways, caldwell & osborn 1996). furthermore, dominant ant species are known to affect the ant assemblages present in a region as well as influence species coexistence (majer, delabie & smith 1994; samways et al. 1996). samways et al. (1996) attributed the lower abundance of species recorded in the disturbed sites in their study to the presence of p. megacephala. consequently, the high abundance of an aggressive species such as p. megacephala may explain the lower abundance of other ant species recorded in the rehabilitated sites in this study. environmental variables many of the environmental variables investigated in this study did not differ between the control and rehabilitated sites. the sites were similar in terms of total grass cover, other cover, rock cover, height of tallest plant and several soil properties (e.g. percentage sand, silt and clay content as well as soil organic matter). as a result, it could be said that the old agricultural land on the reserve has started to recover. despite this, the percentage bare ground, soil ph and the percentage coarse sand differed significantly between the rehabilitated and control sites. this is important as such factors could influence the ant assemblages present in the sites. for instance, soil type is known to have an effect on the ant species present in a specific area (andersen 1993; koen & breytenbach 1988; lindsey & skinner 2001). unfortunately, the exact habitat requirements of ants are not well known (lindsey & skinner 2001). however, the little research that has been done on this topic has revealed that ant assemblages are influenced by a number of habitat variables; soil moisture, soil type, the geology of an area, plant structural complexity and leaf litter cover (andersen 1993; koen & breytenbach 1988; lindsey & skinner 2001). many rehabilitation studies that have compared ant assemblages on rehabilitated and reference sites have established that the vegetation between the two compared treatments is very different. for example, van hamburg et al. (2004) noted that the vegetation composition on rehabilitated ash dams and nearby natural grasslands was distinct. consequently, these differences in plant composition result in different ant species assemblages as well as differences in ant species diversity (van hamburg et al. 2004). also, a study by zaloumis and bond (2010) found differences between rehabilitated and control sites in a south african grassland; rehabilitated sites supported a much smaller community of resprouting forb species than the control sites. this is important as forb species make up much of the diversity of south african grasslands. the poor ability of forb species to reestablish after disturbance is attributed to their poor dispersal ability and lack of propagating sources (dickson & busby 2008; kardol et al. 2008; zaloumis & bond 2010). several environmental factors such as soil properties can alter the direction of succession as well as the plant species present (zaloumis & bond 2010). also, the addition of nutrients such as nitrogen from fertilisers can influence the spread and species richness of weeds and alien plants, which has negative consequences for the native flora of the region (zaloumis & bond 2010). thus, a thorough understanding of the region’s environmental variables and their impact on plant species is of importance, as ants are dependent on vegetation structure (andersen 1995). the failure of successful rehabilitation of the plant community in grasslands could consequentially affect the ant assemblages present within the area. this is important as the present study found a significant difference between the plant assemblages of the control and rehabilitated sites on rietvlei nature reserve. this difference in plant assemblages could possibly explain the difference in ant assemblages on the reserve because of factors such as substrate and vegetation have direct effects on the colonisation ability of different ant species (van hamburg et al. 2004). as a result, the differences in plant assemblages and specific environmental factors may explain the variation between ant assemblages of control and rehabilitated sites. similarly, as ants are considered ecosystem engineers and often cause changes to soil, they too could be a contributing factor affecting the plant species composition within the area. as a result, ants could be driving much of the rehabilitation in the reserve old lands; however, more research is required before a conclusive decision can be made. indicator species three indicator species were identified, all of which were characteristic of the control sites. the indicator species were c. rectinota, c. amita and m. fastidium. no indicator species were identified for the rehabilitated sites. specialist ant predators are said to be highly sensitive to disturbance and thus are rarely recorded in disturbed habitats (andersen et al. 2003; hoffmann & andersen 2003). included in the list of specialist predators (andersen 2000) is the genus leptogenys, in the tribe ponerini. only one species of this genus, leptogenys intermedia emery, was recorded during this study. however, the fact that this species was recorded in both the rehabilitated and control sites suggests that the rehabilitated sites have started acquiring the vegetation structure, compostition and arthropod fauna that is necessary to support the specialised diet of these predators (andersen 2000). this is noteworthy as many authors have found specialist predators to be among the last species to colonise rehabilitating areas (majer & beeston 1996; hoffman & andersen 2003). this is interesting as not much is known regarding the potential effects of these species on their environment and thus requires further research. conclusion our results show that the ant assemblages on rehabilitated areas of the reserve are significantly different from those of surrounding undisturbed areas, although some convergence is evident. these differences are likely to be explained by a combination of factors, including the presence of certain dominant ant species, differences in plant species composition and differences in environmental factors (including percentage bare ground, percentage coarse sand and soil ph). a conclusive statement regarding the success of rehabilitation efforts on the reserve is not possible given the lack of background information on recovery rates and compositional changes, but our study shows that rehabilitation of grasslands would benefit from a greater understanding of ant diversity and the factors that are responsible for driving this diversity. further studies are needed to investigate the functional roles of particular ant species, especially those that are considered to be good indicator species. acknowledgements we are grateful to everyone who assisted with this project; chantal ferreira and jonathan fisher are thanked for their assistance with the ant identifications. our gratitude is also extended to the rietvlei nature reserve and its staff members, especially riaan marais, who made this study possible. furthermore, wesley daniels, chantal ferreira, michael potgieter, andré van tonder, cavin shivambu and paul akkermans are gratefully acknowledged for their assistance in the field. our thanks are extended to the soil sciences department of the university of pretoria who allowed us to use their facilities when conducting the soil analyses. we would also like to thank the anonymous reviewers whose comments helped to improve the manuscript. we are grateful for funding received from the national research foundation. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. authors’ contributions s-l.j., m.r., i.e. and p.h. conceived and designed the study. s-l.j., m.r. and i.e. performed fieldwork. s-l.j. and p.h. performed ant identifications. s-l.j., i.e. and m.r. performed statistical analyses; 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determined with the use of a ph meter (accurate to 0.05 ph units). initially, twelve 50-cm3 beakers were labelled and weighted on a balance (accurate to 0.1 g) and their weights recorded. a 10-g sample of each of the 12 soil samples was placed into the respectively labelled beakers. an automatic dispenser was used to dispense 25 cm3 of de-ionised water into each of the beakers. the beakers were shaken for approximately 5 s and allowed to stand for 60 min. each beaker was then shaken before the electrodes of the ph meter were placed into the soil sample. the results obtained are reported as ph (h2o). the sand, silt and clay contents of the soil were determined with the use of the particle size distribution method. twelve beakers (250 cm3 capacity) were weighed and their weights recorded. each beaker received 50 g of soil from a respective site. to each beaker of soil, 10 cm3 of calgon dispersing solution (a mixture that contains sodium hexametaphosphate [napo4] and sodium carbonate [na2co3] and de-ionised water) was added. the contents of a beaker were placed into a dispersion cup and filled to approximately 150 cm3 with de-ionised water. the sample was then mixed for 5 min with the use of an electric mixer (10 000 rpm – 13 000 rpm). the sand fraction of the soil was removed by washing the dispersed sample on a 0.053-mm-size sieve. this sieve would ensure that only silt and clay could pass through the funnel into a 1-dm3 cylinder. the washing process was continued until the percolate was clear. the sieve was then removed from the cylinder, and the sand was transferred into a labelled beaker. this process was repeated for all 12 soil samples. the beakers containing the separated sand were dried in a drying oven at 105 °c to constant mass. the samples were then weighed and the masses of the sand (excluding the weight of the beaker) were recorded. the mass of the sand fraction was given as a. following this, the cylinders containing the silt and clay were filled to the 1-dm3 mark. each of the cylinders was stirred with a hand stirrer for a period of 30 s. a ‘blank’ was prepared by adding 10 cm3 calgon to a 1-dm3 cylinder of water. the cylinders were left undisturbed for a total period of 6 hours 30 ± 5 min. once this time had elapsed, a standard hydrometer (with bouyoucos scale in grams per litre, ranging from –5 to 60) was slowly inserted into the suspension of each cylinder and the recording (c) was taken. the hydrometer was placed into each suspension carefully in order not to mix the solution. at this time, a temperature reading was taken. the particle sizes were calculated as follows: a = mass of sand fraction (g) b = hydrometer reading of blank c = hydrometer reading of sample m = mass (g) of soil used sand fraction: clay fraction: to calculate the clay reading, the blank hydrometer reading was subtracted from the sample reading (c − b) and the percentage clay under the appropriate temperature was determined using a particle size distribution table (where e is the reading obtained from the table). silt fraction: the soil organic matter was determined by weighing 12 labelled beakers (50 cm3) and recording the masses of each. an approximate mass of between 10 g and 20 g of each soil sample was weighed and placed into the relevant beakers. the beakers were placed in a drying oven at 105 °c and left overnight to ensure that any moisture in the samples was removed. when removed from the drying oven, the samples were weighed and transferred to a furnace (550 °c) and left for a period of 16 hours. the beakers were then weighed and the mass of the soil organic matter determined. this percentage was then subtracted from the previously determined sand fraction to produce a new adjusted sand fraction. figure 1-a1: sample-based rarefaction curves indicating the number of species (srar), ice mean (incidence-based coverage estimator), chao 2 mean (abundance-based richness estimator), jack 2 mean (second-order jackknife richness estimator) and the michaelis–menton mean (mmmeans) richness estimators of ants in (a) control sites and (b) rehabilitated sites of the rietvlei nature reserve. appendix 2: braun–blanquet scale for vegetation cover table 1-a2: plant morphospecies abundance quantified with the use of the braun–blanquet abundance scale and the altered scale that was used during the analysis of data. figure 1-a2: box plots indicating the various vegetation indices recorded in 1 m × 1 m quadrats, (a) percentage bare ground, (b) percentage grass cover, (c) percentage of other cover (forbs, shrubs and herbs), (d) percentage rock cover, (e) height (cm) of the tallest plant in each of the quadrats for each of the sites and (f) height of the biomass recorded from the disk pasture meter. appendix 3: ant species lists for various data sets specimens were identified to species level where possible. specimens that could not be identified to species level were identified to genus and assigned to numbered morphospecies. table 1-a3: the number of individuals per species, listed per subfamily, which were collected in total for rehabilitated and control sites on the rietvlei nature reserve, south africa. figure 1-a3: box plots indicating the properties of the soil collected from the different treatments: (a) percentage soil organic matter, (b) percentage sand, (c) percentage silt, (d) percentage clay, (e) ph and (f) percentage coarse sand. appendix 4: tables illustrating sampling completeness calculated for the rehabilitated and control sites table 1-a4: richness estimator values as obtained from estimate-s (v 9.1.0; colwell 2013). table 2-a4: richness estimator values as obtained from estimate-s (v 9.1.0; colwell 2013). table 3-a4: the reliability of each inventory was calculated for the control sites. table 4-a4: the reliability of each inventory was calculated for the rehabilitated sites. table 5-a4: overall sampling completeness. figure 1-a4: non-metric multidimensional scaling plot indicating the similarity of plant assemblages among replicates of control and rehabilitated sites. appendix 5: list of indicator values table 1-a5: indicator values generated in rstudio. species fulfilling the criteria (indval > 70%, p < 0.05) are identified as indicator species. article information authors: ben j. strohbach1 josephat t. kutuahuripa2 affiliations: 1school of natural resources and spatial sciences, polytechnic of namibia, namibia 2plant production research, ministry of agriculture, water and forestry, namibia correspondence to: ben strohbach email: bstrohbach @polytechnic.edu.na postal address: private bag 13388, windhoek, namibia dates: received: 30 oct. 2012 accepted: 26 aug. 2014 published: 28 nov. 2014 how to cite this article: strohbach, b.j. & kutuahuripa, j.t., 2014, ‘vegetation of the eastern communal conservancies in namibia: ii. environmental drivers’, koedoe 56(1), art. #1117, 12 pages. http://dx.doi.org/10.4102/koedoe.v56i1.1117 note: additional supporting information may be found in the online version of this article as an online appendix: http://dx.doi.org/10.4102/koedoe.v56i1.1117-1. copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. vegetation of the eastern communal conservancies in namibia: ii. environmental drivers in this original research... open access • abstract • introduction • research method and design    • study area    • procedure       • available soil moisture       • climatic data       • environmental gradients • results • discussion    • climatic gradients versus changes in soil types    • plant adaptations to environmental drivers    • shortcomings in data analysis • conclusion • acknowledgements    • competing interests    • authors’ contributions • references • footnotes abstract top ↑ the eastern communal conservancies are situated along the western fringe of the kalahari basin. under a very short rainfall gradient, the vegetation abruptly changes from microphyllous acacia-dominated savannas to mesophyll savannas, dominated by terminalia sericea and combretum spp. we hypothesise that this is caused by changes in soil moisture availability brought about by changes in soil texture from loamy soils to deep sands (the ‘inverse texture effect’). for this analysis, we used vegetation and soils data derived from a recognisance survey of the natural resources of the study area. as the sites in the soil and vegetation surveys did not overlap, it was decided to use only synoptic data for the plant associations in the analysis. non-metric multidimesional scaling ordination was utilised as ordination technique of the vegetation data and various environmental parameters, including soil texture, soil hydraulic parameters, climatic and fire regime parameters, were overlaid as biplots onto the resulting graph, as were various plant functional attributes particularly related to climatic conditions. the main environmental gradient identified within the study area is the rainfall gradient. this relatively short gradient, however, does not explain the marked change in vegetation observed within the study area. this change is attributed to the change in soil type, in particular, the soil texture and the associated soil hydraulic parameters of the soil. this gradient is closely correlated to leaf size, explaining the change from microphyll savannas to mesophyll savannas along the change from loamy to sandy soils. one of the lesser understood mechanisms for the survival of these mesophyll plants on sandy soils seems to be a deep root system, which is actively involved in water redistribution within the soil profile – by hydraulic lift, inverse hydraulic lift and stem flow. conservation implications: understanding these mechanisms will greatly assist in understanding savanna dynamics. with the threat of global climate change, we postulate that the vegetation will gradually change from the present mesophyll to a microphyll savanna, but that the grass sward will probably not develop very well. shrub and tree removal (‘bush harvesting’) is likely to speed up the desertification process within this area. introduction top ↑ in the first article of this series (strohbach 2014), the vegetation of eight communal conservancies, as well as two farming areas in east-central namibia, are described. these conservancies are situated within the camelthorn savanna in the south and the tree savanna and woodlands described by giess (1998) in the north. here, under similar climatic conditions, the vegetation abruptly changes from a microphyll, acacia-dominated savanna to a mesophyll savanna dominated by terminalia sericea and various combretum species, as the soils change from relative fine-grained soils of the central plateau to deep, coarse sands of the kalahari basin (leser 1972). on an aridity gradient stretching over roughly 1100 km from north-northwest to south-southwest within the kalahari desert of botswana (on uniformly sandy soils), macrophyllous trees are gradually replaced by microphyllous trees as the rainfall decreases and the aridity increases (skarpe 1996). this rainfall gradient is estimated to range from 600 mm in the north-northeast to less than 250 mm in the south-southwest. on a similar aridity gradient from the lesotho highlands to the west coast of southern africa (with a rainfall gradient between 522 mm and 41 mm per annum), macrophyll plants were replaced by microphyll plants within a riverine forest as aridity increased (werger & ellenbroek 1978). in the same study, an increasing number of plants with sclerophyllous leaves (as adaptations to arid conditions) were found in the more arid environments. more abrupt changes in the composition of moist savannas of central mozambique (ranging from grasslands to savannas, to woodlands, forests and swamp forests) have, in detail, been demonstrated by tinley (1982) to be related to differences in soil profiles, especially with regard to soil texture, soil depth and the presence and depth of constricting layers within the profile, despite a uniform climate. however, little is known on the causes for the abrupt change from microphyll to macrophyll savannas in the arid western kalahari basin region. water availability is the main driving force of vegetation in arid environments (noy-meir 1973; tinley 1982; walter & breckle 1984). tree roots are known to utilise an expansive soil volume to source nutrients and water. record depths of 60 m and 68 m have been documented for acacia erioloba and boscia albitrunca, respectively, in the kalahari (canadell et al. 1996). pterocarpus angolensis tree roots reach a depth of between 15 m and 20 m (mwitwa, munthali & van wyk 2008), whilst baphia massaiensis subsp. obovata has been reported to have a rooting depth of about 2.5 m – 2.7 m (savory 1963). terminalia sericea, one of the most abundant species in the study area, is known to have a shallow root system less than 1 m deep under arid conditions (350 mm mean annual rainfall) (hipondoka & versfeld 2006; hipondoka et al. 2003). under more mesic conditions (450 mm mean annual rainfall), however, this species tends to retain its tap root, reaching beyond the explored depth of 1.2 m. similar findings are known also from the banksia woodlands in australia, where some deep-rooted species obtain their water from a depth of up to 9 m. other evidence from australian eucalyptus savannas suggest that deep-rooting species are better capable of surviving drought conditions than shallow-rooting species (fensham & fairfax 2009). it is generally assumed that fine-textured soils have greater porosity, and thus a greater water-holding capacity, than coarse-textured (sandy) soils (brady & weil 2002). finer soils have a larger pore volume, albeit smaller pores, allowing more water to be stored in the profile. added to this is the fact that water flows faster through larger pores – a sandy soil loses water to deep soil layers more rapidly than a loamy or clay soil. in arid and semi-arid areas, however, the inverse texture effect is observed – vegetation is lusher and better developed, often with a higher net primary productivity, on sandy soils than on finer clay or silt soils (austin et al. 2004; noy-meir 1973). this is a result of the fact that soil water is held by adhesion to the fine-grained particles and is, in effect, not available to plants. in sandy soils, water infiltrates to deeper soil layers during the rainy season, out of reach of fast-growing, shallow-rooted plants such as grasses. during the dry season, this deep soil moisture reservoir becomes available to deep-rooted plants (brady & weil 2002; noy-meir 1973; walter & breckle 1984). an in-depth analysis of this phenomenon has been provided by laio, porporato, fernandez-illescas and rodriguez-iturbe (2001), laio, porporato, ridolfi and rodriguez-iturbe (2001), porporato et al. (2001) and rodriguez-iturbe et al. (2001). with this article, we want to illustrate the effects of environmental factors influencing the formation and distribution of various vegetation associations found within the study area. in particular, we hypothesise that the ‘sudden’ or marked change from microphyll acacia-dominated savannas to mesophyll savannas (terminalia sericea, combretum spp.) in the eastern communal areas is the result of a combination of increased rainfall and especially increased water availability in the deep, sandy soils of the kalahari. research method and design top ↑ study area the omaheke [‘great sand desert’] forms the western fringe of the kalahari basin (mendelsohn & el obeid 2002, 2003; mendelsohn et al. 2002) (figure 1). the central, major part of the landscape is formed by an extensive sand plateau, sloping from the southern, western and north-western fringe towards the central east. the topography of the kalahari sand plains is flat to almost flat. these kalahari sands, with weak to little development in the profiles, are classified as haplic or ferralic arenosols.1 these soils are generally deep to very deep and well drained. they have a low agricultural potential as they are leached, have a low water-holding capacity and a low nutrient status (coetzee 2003; mendelsohn et al. 2002; strohbach et al. 2004). figure 1: the study area (indicated in bold lines along the east-central border of namibia) in relation to the kalahari sand basin in southern africa (grey shading). within this sandy plain, occasional dune fields occur (de pauw et al. 1998; thomas 1984). the sand dunes reach a height of up to 20 m, forming an undulating to rolling landscape. the dune crest, dune slopes and foot of the dunes consist of ferralic arenosols with haplic arenosols mostly on the lower footslopes of the dunes. soils on the interdunal depressions and isolated small pans of the kalahari vary in depth from moderately deep to very deep sandy soils. the two major soil types that occur within this soil-mapping unit are petric calcisols (table 1) and haplic arenosols. isolated pans have shallow haplic calcisols with calcrete horizons surfacing and some gleying was occasionally found (strohbach et al. 2004) (figure 2). figure 2: dominant landscape units within the study area, indicating also the dominant soils for these landscape units (compare with table 1). table 1: vegetation associations and their habitats from the study area, as extracted from global information system layers. the sand plateau is drained by a number of deeply incised drainage channels, commonly referred to as omiramba (singular: omuramba) (king 1963; strohbach 2008). these fossil drainage lines were formed during the wetter phase predominating the early to mid-pleistocene age. the depths and widths of the omiramba depend on the prevailing winds and water flows experienced throughout the ages. very often, calcrete outcrops are exposed at the edges of the major omiramba, especially the otjozondjou, epukiro, eiseb and rietfontein omiramba. the major soil types of the omiramba are arenic or calcic fluvisols, occasionally associated with petric or haplic calcisols (strohbach et al. 2004) (table 1) (figure 2). along the south-western fringes of the kalahari sand basin, but also south and east of gam, the sand cover is shallow, often overlaying calcretes, with occasional calcrete pans and reworked whitish, loose, unstructured kalahari sands. the topography is flat to almost flat. typical here are haplic calcisols, whilst in the far north-east some eutric cambisols (table 1) have also been found (strohbach et al. 2004) (figure 2). floodplains occur between the klein omatako omuramba and the omuramba omatako and stretch up to okanguindi village, south-east of okakarara. they are subjected to repeated seasonal flooding as a result of the relatively low relief. the continuous erosion of the surface horizon by water, results in the formation of many pans and watercourses throughout the area. owing to continuous reshaping and removal of the parent material and topsoil, the soils of this mapping unit are much younger in terms of development and age. the dominant soils of this mapping unit, arenic fluvisols (table 1), are weakly developed and poorly structured (figure 2). soils within the lower positions are subjected to flooding and show signs of mottling caused by poor drainage and water-logging. in the far north-west of the study area (north-west of otjituuo towards grootfontein), but also north of gam, is a plateau with limestone related to the karstveld. hard and soft calcrete and/or limestone outcrops are exposed at the soil surface. the dominant soils are lithic and molic leptosols, with some skeletic leptosols being associated (strohbach et al. 2004) (figure 2). the climate is relatively uniform, with a rainfall gradient ranging, on average, from just under 400 mm in the south to just under 500 mm in the north, annually (figure 1). the variation in rainfall (cv) is about 60% for the entire study area (botha 1996; mendelsohn et al. 2002; namibian agricultural resources information system [naris] 2001). this typical summer rainfall area has a dry winter and a dry and hot spring season (figure 3). frost is not unheard of in winter, with between 5 days of frost in the north and 30 days or more in the south (mendelsohn et al. 2002; southern african science service centre for climate change and adaptive land use [sasscal] weathernet 2012) (figure 3a–3c). the average relative humidity drops to below 10% during the dry months, but rises to 70% – 80% during the rainfall months (mendelsohn et al. 2002). figure 3: climate diagrams for the stations, (a) tsumkwe and (b) sonop (both to the north of the study area) and (c) sandveld to the south of the study area. regular veld fires are known to occur within the study area (le roux 2011; mendelsohn & el obeid 2002). the fire return period ranges between less than 2 years and 10 years and longer, with especially the north-eastern part of the study area being burned very frequently. as a combination of a variety of factors, including the fire return period, the seasonality of fires, et cetera, le roux (2011) determined a ‘fire regime’ for various areas in namibia. most of the study area is classed as having a ‘mild’ to ‘moderate’ fire regime. procedure at the same time as the vegetation survey in april 2004, a survey of the major landscape types and associated soil types was conducted. in addition to describing the landscapes based on field observations and topographic map interpretation, soil profiles were exposed by digging soil pits on representative sites throughout the study area (strohbach et al. 2004). however, because of the fairly labour-intensive sampling procedure, only 71 sample pits were dug. very few of these were at the same localities as the vegetation sample plots (strohbach 2014). this means that for this particular study, whilst also relying on existing geographic information system (gis) sources (le roux 2011; mendelsohn & el obeid 2002; mendelsohn et al. 2002; naris 2001), only extrapolated environmental data were available. it thus made sense to use synoptic composition data of the associations and subassociations for all analyses, rather than the individual relevé data, for comparison to the available environmental data; see the summarised data used in table 1. for all analyses, the panicum gilvum–marsilea vlei community was omitted as a potential outlier, also because only one relevé was collected in this community. analytical soil data from the soil profile descriptions were augmented with standard soil parameters developed for the world soil reference base (batjes 2002). available soil moisture soil hydrological parameters, specifically saturated hydraulic conductivity, saturation matric potential and porosity, can be directly linked to the soil texture, that is, the percentage of sand, silt and clay in the soil (cosby et al. 1984). we followed the convention of these authors and used the composition of the midpoint of the soil-textural class on the soil texture triangle. we used the soil characteristics of the dominant soil horizon in the case of soils with widely varying soil properties in their horizons; that is, we would ignore a shallow, sandy a-horizon and rather use the characteristics of the deeper, loamy b-horizon. soil hydrological parameters were derived from the textural classes, using data provided by cosby et al. (1984) and laio, porporato, ridolfi and rodriguez-iturbe (2001). no attempt was made to estimate the actual soil matric potential based on annual rainfall, soil texture and soil depth, because, in many cases (especially within the various arenosols), the depth of the soil profile is unknown, but it could reach depths of up to 250 m or deeper (mendelsohn & el obeid 2002). climatic data data on mean annual rainfall and frost days were extracted by overlaying plot positions onto the relevant gis layers. all other relevant climatic data (cv of rainfall, average minimum temperatures, relative humidity in both wet and dry seasons, etc.) were too uniform across the study area to warrant their consideration as significant environmental gradients (botha 1996; mendelsohn et al. 2002; naris 2001). the extracted data were averaged for the associations and these averages used in the analysis. in a similar way, the fire return period and fire regime (based on the data of le roux 2011) were extracted for plots and summarised for the associations. environmental gradients in order to demonstrate the main environmental drivers, the vegetation data were processed in two ways: by means of a non-metric multidimensional scaling (nms) ordination using pc-ord 6 (mccune, grace & urban 2002), as well as a classification of plant functional attributes using vegclass (gillison 2002; gillison & carpenter 1997). reciprocal averaging (hill 1973) was not used because of a severe arch effect. as the data set was relatively small, the use of nms, using the sørensen distance measure, was regarded as more feasible. the ordination procedure was completed by using pc-ord 5 (mccune et al. 2002) and repeated twice. the first run was based on a random starting configuration with 50 real data runs and 249 runs on randomised data. after 107 iterations a final solution was reached, with a stress of 7.196 and a final instability of 0.00001. the results were saved and used as base configuration for the second run. this time, only 10 runs with real data were allowed, also requesting a varimax rotation of the results. the environmental data were overlain as a biplot, to indicate the environmental gradients of interest. a second biplot was created, using plant functional attributes (see below) as variables. as water availability is hypothesised to be the main driving force in the composition of the vegetation, plant functional attributes (pfas) as related to water availability were utilised to illuminate these gradients. the pfas were chosen according to criteria described by gillison and carpenter (1997), in order that they might be used in the vegclass software for analysis. in the context of this study, all pinnately compound leaves have been classified as ‘filicoid’ (‘filicoid’ leaves are, according to the description of gillison & carpenter 1997, fern-like leaves). the pfas were identified from a variety of sources, including the tree atlas of namibia (curtis & mannheimer 2005), the field guide to trees and shrubs of namibia (mannheimer & curtis 2009), the field guide to the plants of northern botswana (heath & heath 2009), grasses of namibia (müller 2007), the prodromus zur flora von swa (merxmüller 1966) and a number of internet sites such as jstor plant science (http://plants.jstor.org), flora of zimbabwe (http://www.zimbabweflora.co.zw/index.php) and photo guide to plants of southern africa (http://www.southernafricanplants.net/index.php). the data were captured in vegclass, ensuring that individual species had the same functional attributes across syntaxa. this is contrary to the recommendations of gillison and carpenter (1997), but necessary as these pfas were collected in retrospect; furthermore, this is acceptable as these pfas represent the typical attributes of the species within the broad forest savanna and woodland of the northern kalahari (sensu giess 1998). the distance between all associations, based on changing pfa combinations, was calculated using the wald-wolfowitz distance measure as recommended by gillison and carpenter (1997). as the software lacks an option to reduce the resulting distance matrix to two dimensions, the resulting distances were plotted using the distance measures from association 14 (burkeo africanae–pterocarpetum angolensis) and association 1 (acacio fleckii–terminalietum prunioidis) and as axes 1 and 2, respectively. biplots of both the habitat and key pfa data were overlain onto this graph, again to elucidate the gradients. in all cases, the biplots were limited to those environmental variables or pfas with an r2 > 0.3. as a total of 31 pfas were identified (resulting in 123 combinations as plant functional types), it was decided to use only key pfas in biplot overlays onto ordination diagrams. these key pfas included the following: leaf size, leaf and photosynthetic adaptations, growth form (sensu raunkiaer – mueller-dombois & ellenberg 1974) and the number of perennial plants (opposing the growth form ‘therophytes’, which are annual plants). the leaf sizes have been combined as follows: microphyll leaves are all leaf sizes ≤ microphyll, sensu gillison and carpenter (1997), or ≤ 2025 mm2. macrophyll leaves are all leaf sizes ≥ notophyll, sensu gillison and carpenter (1997), or ≥ 2025 mm2. these key pfa data are represented in table 2, whilst the detailed pfa data are presented in online appendix 2. table 2: simplified key plant functional attributes, expressed as a percentage of all species found in the relevant associations. once the main environmental drivers were determined, the significance of these in affecting the compositional and plant functional changes between various vegetation associations was determined using a multiple linear regression analysis. for this, two runs of the procedure were completed, using the percentage occurrence of macrophyll and microphyll plants in each association as dependent variables and the mean annual rainfall, mean number of frost days, mean soil depth as well as soil texture, expressed as mean percentage sand, silt and clay, as independent variables. soil ph and electric conductivity was not used in this calculation, as these factors were assumed to be closely related to the soil texture. the f-value for variables to be entered in the regression was set to 0.5 for forward selection. for the calculation, the software statistica (2013) was used. results top ↑ the nms ordination results and the vegclass results are depicted in figure 3 and figure 4, respectively. the nms results represent 91.5% of the variation between the various associations, with axis 1 representing 52.8% and axis 2 representing 38.7%. figure 4: non-metric multidimensional scaling results of the synoptic vegetation data, depicting, (a) biplot overlaid with the habitat factors and (b) biplot overlaid with the key plant functional attributes. rainfall, soil depth and sandy soils are positively correlated with ordination axis 1, whilst the clay and silt content, combined with increasing ph, are negatively correlated with this axis (figure 4a). as expected, hydraulic conductivity is co-correlated with sandy soils along the main axis, whilst soil porosity, soil matric potential and soil pore volume are closely associated with the finer textured soils. these tendencies are partially confirmed by the vegclass results, with the soil texture also influencing the distribution of associations in relation to association 14 (burkeo africanae–pterocarpetum angolensis) (figure 5a). however, the soil hydraulic parameters do not seem to be of importance in the vegclass scatter plot. figure 5: scatter diagrams of the distances between various vegetation associations based on their plant functional attributes (using the distances from associations 14 and 1 on the x-axis and y-axis respectively), depicting, (a) biplot overlaid by the habitat data and (b) biplot overlaid with the key plant functional attributes. the main gradient manifests itself in a gradual change from mesophyll to microphyll plants – the mesophyll plants, as expected, being associated with the deep, sandy soils under higher rainfall regimes, whilst the microphyll plants are more restricted to the loamy soils (figure 4b). together with the microphylls, the loamy soils support a strong annual flora (therophytes) and plants with succulent leaves, whereas the sandy soils support macrophyll plants, perennials, especially phanerophytes, but also plants with filicoid leaves and deciduous plants. exactly the same combinations of pfas are found to correlate with the main gradient in the vegclass scatter diagram (figure 5b). the two results differ in the reaction to the second axis – within the nms ordination, cryptophytes (e.g. geophytes, geoxylic suffrutices sensu white 1976) are negatively correlated to axis 2, whilst chamaphytes (dwarf shrubs) react to the secondary axis of the vegclass results. no indication of a sensitivity to frost could be identified, whilst sclerophyll leaves are only very weakly correlated with the secondary axis of the nms (r2 < 0.2). fire regime did feature in the vegclass scatter plot. the fire regime in the study area generally varies between mild to moderate (le roux 2011) and does not show any tendency on the ordination diagram (figure 4a). it is more likely that with certain associations (especially the widespread combreto collini–terminalietum sericeae on the kalahari sand plains), fire is primarily responsible for the structure and finer compositional changes (figure 6). this, however, needs to be studied in greater depth. figure 6: fires have a severe structure-altering effect on bushlands and shrublands of especially the combreto collini–terminalietum sericeae typicum (subassociation 12.3), as seen from, (a) unburned bushland and (b) vegetation reduced to a low shrubland after a severe fire. the results of the multiple linear regressions indicate that, for both microphyll and macrophyll plants, rainfall is a significant driver in the occurrence of the plants. soil depth and soil texture (sand and clay content) were only insignificantly associated with this change in composition, whilst other factors did not obtain the necessary f-value to be taken up in the regression (table 3). table 3: multiple linear regression results for the dependent variables ‘macrophyll plants’ and ‘microphyll plants’, respectively. discussion top ↑ climatic gradients versus changes in soil types rainfall has, by far, the strongest and most consistent effect on the vegetation in arid to semi-arid climates. this also holds true for the vegetation within the present study area, which was proven to be the only significant driver in the change from microphyll to macrophyll vegetation (table 3). our present gradient is roughly 200 km long, with a rainfall gradient of less than 150 mm. skarpe (1996) described a similar change in vegetation as it occurs across botswana along a rainfall gradient over 1100 km on uniformly sandy soils; she unfortunately does not provide rainfall differences in her paper. from the geographical description, it is assumed to be a gradient from bokspits in far south-west botswana (177 mm mean annual precipitation – bhattachan et al. 2012) to the four-country corner with zimbabwe, namibia and zambia in north-east botswana. for nearby katima mulilo in north-east namibia, a long-term mean annual precipitation of 758 mm has been recorded (namibia meteorological services 1997). from this, a rainfall gradient of approximately 580 mm is assumed for skarpe's study. her main finding is that broad-leafed trees and grasses are positively correlated with increasing precipitation, whilst fine-leafed phanerophytic species and chamaphytic species were positively correlated with increasing aridity along the gradient. whereas skarpe (1996) describes a gradual change in vegetation over a rather lengthy gradient, the change from microphyll-dominant vegetation (associations a1 – a11) to macrophyll-dominant vegetation (associations a12 – a14) within the study area is rather abrupt over a very short geographical distance (cf. hüttich et al. 2009; strohbach 2014). these dramatic changes occur over virtually no rainfall gradient at all and are related to the change in soil type, from a fine-textured, loamy soil to the coarse-textured, deep sands of the kalahari basin, as suggested by leser (1972). tinley (1982) describes similar dramatic changes resulting from changes in soil texture for mozambique. these changes are caused by the difference in water availability as a result of differences in soil moisture relationships (in particular, a higher soil matric potential and higher porosity, but also higher adhesion, in loamy soils versus a high hydraulic conductivity in sandy soils), causing the inverse texture effect (figure 4) (austin et al. 2004; noy-meir 1973; porporato et al. 2001). the gradual change between the various subassociations of the combreto collini–terminalietum sericeae (a12) and the burekeo africanae–pterocarpetum angolensis (a14) from south to north happens on similar sandy soils and can be compared to the changes in pfas across a climatic gradient described by skarpe (1996). plant adaptations to environmental drivers owing to the higher hydraulic conductivity of sandy soils, rainfall infiltrates faster, but, at the same time, water percolates to deep soil layers. this means that water is available in the upper soil layers for a comparatively short period during the rainy season, whereas deeper soil layers (below 2 m) act as a reservoir for deeper-rooted plants (b. strohbach, unpublished data). therefore, deep-rootedness is a prerequisite for the survival of phanerophytes growing on such sandy soils. this holds true for most of these species (hipondoka & versfeld 2006; hipondopka et al. 2003; mwita et al. 2008; savory 1963; see also timberlake & calvert 1993). deep-rooted geoxylic suffrutices, for example entada arenaria and dichapetalum cymosum, better known from moister savannas and woodlands of the sudano-zambezian ecoregions (500 mm – 700 mm mean annual rainfall) (white 1976), thus also succeed in growing in this environment. the latter species (gifblaar or magou) is problematic throughout the study area, where it can increase rapidly in density by taking advantage of disturbances caused by high numbers of cattle (bester 1989). another adaptation to the temporary high availability of water in sandy soils during the rainy season (because of high infiltration rate) can be seen in stem succulents, specifically various commiphora species (figure 4b). these are able to photosynthesise, also during the dry season, by means of a photosynthetic layer under the translucent bark – an adaptation to xeric environments (referred to as ‘cortic photosynthesis’ by gillison & carpenter 1997). whether pinnately compound leaves (‘filicoid leaves’) point to any specific adaptation, or are rather a coincidence related to the species composition, could not be established. the grass sward of the sandy soils is dominated by the annual species aristida stipitata and megaloprotachne albescens. most grass roots tend to be limited to the upper 40 cm to 60 cm of the soil profile; they are therefore unable to access water from deeper water resources (hipondoka et al. 2003). with this annual character, the grass sward utilises a strategy of growing rapidly during the rainy season to exploit available resources to the fullest in the shortest possible time, but does not store water in underground organs or access the water that is available in deeper soil layers during adverse seasons. the few perennial grass tufts either have hairy leaves (e.g. digitaria seriata), or are extremely fine-leaved (e.g. stipagrostis uniplumis var. uniplumis) as an adaptation to this harsh environment. digitaria seriata is also often found within bush clumps, making use of both shade and increased water infiltration as a result of stem flow under these canopies. similar patterns were found in other arid environments (martinez-meza & whitford 1996; pressland 1973). perennial grasses within the kalahari are also known to utilise inverse hydraulic lift to channel water down into the soil profile (to a depth of between 50 cm and 100 cm). this is postulated as a potential mechanism (not only by grasses, but also by trees) to create an own ‘reservoir’ of soil water for use during the dry season (schulze et al. 1998). most of the phanerophytes growing on sandy soils are deciduous, shedding their leaves during the dry winter and hot, dry spring months (table 2, own observations). this is a typical adaptation to avoid water loss (fensham & fairfax 2009; seghieri, floret & pontanier 1995). many phanerophytes, especially terminalia sericea and various combretum spp., have hairy leaves, an adaptation typically associated with psammophil plants (warming 1909). within this study, deciduous plants were not limited to the sandy soils. many species growing on loamy soils exhibit the same adaptation; however, several species are distinctly evergreen (e.g. boscia albitrunca) or at least semi-deciduous, that is, shedding their leaves just before the next leaf flush (e.g. acacia erioloba, acacia hebeclada subsp. hebeclada). these evergreen to semi-deciduous species are also deep-rooted (canadell et al. 1996; moustakas et al. 2006; schulze et al. 1998), enabling them to obtain water from deep soil layers during the dry season. overall, however, the tendency is for the leaves to become distinctly smaller on the more loamy soils, an indication that the soil moisture availability is far lower on loamy soils than on sandy soils. an extreme case of edaphic aridity is presented by the shallow calcrete pans of the eragrostio echinochloideae–eriocephaloetum luederitziani (a4). in addition to the very shallow profile, the soil is characterised by a high skeletal content, a high ph (> 8) and a relatively high electric conductivity (> 100 µs/cm). moreover, the low soil volume can only store limited quantities of water, resulting in this water being saturated with ions, which means that it is not readily available to plants. the nature of these soils is manifested in the vegetation structure; the association being dominated by dwarf shrubs and, furthermore, within this association, 14% of all species had sclerophyll leaves (compared to an average of 6% – 9% for all other associations) and a relatively high microphyll leaf component (89%) (table 2). although fire is known to have a severe influence on the vegetation composition in the kavango and zambezi regions (geldenhuys 1977; le roux 2011; mendelsohn & el obeid 2003; mendelsohn & roberts 1997; strohbach & petersen 2007), only structural alterations were evident during the present study (figure 6). it is thus unlikely that fire is a driving factor in the discrimination of vegetation associations within the study area. shortcomings in data analysis a major shortcoming of this study is the fact that only synoptical data (both for vegetation composition as well as for environmental variables) could be employed. because of this, the clear soil gradients illustrated by the ordination techniques were not conclusive in the multiple linear regression. future studies concerning such moisture relations should take this problem into consideration. conclusion top ↑ the study supports the hypothesis that the broad-leafed savanna of the kalahari is limited in distribution by the deep sands, as a result of better soil moisture conditions. the improved soil moisture conditions in the kalahari sands are a result of the inverse texture effect (noy-meir 1973). in combination with increased rainfall to the north, this even allows tall trees to grow and replace the dominating shrublands with woodlands, as is evident from the transition between the combreto collini–terminalietum sericeae (a12) and the burkeo africanae–pterocarpetum angolensis (a14) – roughly coinciding with the 500 mm rainfall isohyet. deep-rootedness of phanerophytic species is one of the major adaptations of psammophil plants to utilise available water resources, but also to recharge and redistribute soil moisture resources in the profile (burgess et al. 1998; kizito et al. 2012; martinez-meza & whitford 1996; schulze et al. 1998). a better understanding of these mechanisms is essential if one wishes to gain a clear grasp of savanna dynamics. furthermore, predictions regarding the effects of global climate change on the vegetation in the kalahari environment can be improved by understanding the strategies employed by different plant species in the utilisation of water. it is predicted that, over the next 50 years, rainfall will decrease over namibia, resulting in less available grazing for extensive livestock production (midgley et al. 2005). in another study, it is predicted that the sands of the kalahari basin will be remobilised during the next 50 years, resulting in largely unvegetated mobile dunes, as a result of global climate change (thomas, knight & wiggs 2005). we postulate that climate change will manifest itself initially through a change in composition, from the present mesophyll savanna to a microphyll (acacia-dominated) savanna, very similar to that of the present camelthorn savanna sensu giess (1998). the camelthorn savanna forms part of the greater kalahari landscape on sandy soils, between about 250 mm and 400 mm mean annual rainfall, to the south of the present study area. it is, however, doubtful whether the grass sward will, under conditions of prolonged drought, develop to the present, relatively productive state. an added danger to productive savannas is the ongoing practice of harvesting terminalia sericea for fencing poles. already planned are projects to harvest ‘invader’ bush within the study area, to be used as fuel for power generation. such large-scale removal of shrubs and small trees from the already poor landscape will inevitably lead to reduced water infiltration into the soil (because of reduced stem flow) and reduced water redistribution within the soil profile, in turn leading to less favourable conditions for grasses. the net effect will be an accelerated desertification through a ‘man-made drought’. acknowledgements top ↑ thanks are due to ms marianne strohbach for assistance with the vegetation survey, as well as to mr heiner mouton for helping with the soil survey. the soil samples were analysed by the agriculture laboratory, directorate of agricultural research and training, in windhoek. this project was co-funded by the global environment facility through the desert margins programme and the government of namibia, through the recurrent budget of the directorate of agricultural research and training. competing interests the authors declare that they have no financial or personal relationships which may have inappropriately influenced them in writing this article. 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1993, ‘preliminary root atlas for zimbabwe and zambia’, the zimbabwe bulletin of forestry research 10, forestry commission, harare. tinley, k.l., 1982, ‘the influence of soil moisture balance on ecosystem patterns in southern africa’, in b.j. huntley & b.h. walker (eds.), ecology of tropical savannas, ecological studies ser. 42, pp. 175–192, springer verlag, berlin. http://dx.doi.org/10.1007/978-3-642-68786-0_9 walter, h. & breckle, s.-w., 1984, spezielle ökologie der tropischen und subtropischen zonen, ökologie der erde [special ecology of the tropical and subtropical zones, ecology of the earth], vol. 2, gustav fischer verlag, stuttgart. walter, h., harnickell, e. & mueller-dombois, d., 1975, climate-diagram maps 1–36, springer verlag, berlin. warming, e., 1909, oecology of plants. an introduction to the study of plant-communities, oxford university press, london. werger, m.j.a. & ellenbroek, g.a., 1978, ‘leaf size and leaf consistence of a riverine forest formation along a climatic gradient’, oecologia 34, 297–308. http://dx.doi.org/10.1007/bf00344908 white, f., 1976, ‘the underground forests of africa: a preliminary review’, gardens’ bulletin singapore 29, 57–71. footnotes top ↑ 1.soil definitions follow the world soils reference base (food and agricultural organization & un educational, scientific and cultural organization 1988). a full description of the soil types found is presented in online appendix 1. filelist convert a pdf file! koedoe 19: 63-66 (1976) a note on mountain reedbuck redunca fulvorufula fulvorufula afzelius in the kruger national park l. r. irby caesar kleberg research programme in wildlife ecology department oj wildlife and fisheries sciences texas a & m university college station texas 7784) abstract this survey was conducted to obtain information on the mountain reedbuck population in the kruger national park for comparison with the population in the loskop dam nature reserve, transvaal. during 93 hours of search effort, 36 mountain reedbuck sightings were recorded. all of the sightings were in the malelane hills on slopes above the kwa machiyaliwane drainage. introduction mountain reedbuck in the kruger national park (knp) are found only in the malelane hills and along the lebombo mountain. the population is estimated to number 240 (pienaar 1969). the status of this species in the park is considered precarious by many people, but too little is known of its population structure and distribution to determine whether the population is declining or merely occupying a limited area of suitable habitat (smuts, pers. comm.). during february and september 1970, 19 days were spent in the park surveying the mountain reedbuck population in the malelane hills. the object of the survey was to obtain information for comparison with the mountain reedbuck population in the loskop dam nature reserve located near groblersdal, transvaal. this paper is a condensation of observations made during the survey. methods mountain reedbuck were located in four ways: 1. scanning hillsides with 7 x 35 binoculars and a 60 x spotting scope; 2. waiting by watering sites at dawn and sunset; 3. walking ridgelines accompanied by a game scout; 4. driving firebreak roads. results only six individual mountain reedbuck were sighted during 52,5 63 hours of observation in february (table 1). all sightings were in the hills along the kwa machiyaliwane drainage even though a comparable amount of time was spent in the vicinity of khandizwe hill and along the emcwebeni drainage. table 1 relative intensity of sampling and success using four techniques for sighting mountain reedbuck in the kruger national park february september observation animals observation animals time sighted time sighted technique (h) (h) spotting scope and binoculars 18,6 0 2,1 3 driving firebreaks 15,0 4 3,9 0 waterhole counts 3,0 0 4,3 0 foot transects 15,9 3 30,0 27 efforts in september were more successful (table 1). of 30 animals sighted, at least 25 were not resightings. group size ranged from one to six. table 2 gives group size, composition, and general habitat information for the september sightings. all of the animals were located on hills above the kwa machiyaliwane drainage in combretum woodland. most of the groups were seen on west-facing slopes on a burned area east of the kwa machiyaliwane streambed. none were more than 5 km from the matjulwana windpump. discussion the low number of sightings recorded in february was probably due to visibility restrictions imposed by the fully-leafed vegetation. several of the sightings in september would not have been made if the trees were not bare. the presence of juveniles in september (table 2) indicates that mountain reedbuck are reproducing in the park and that lambs are surviving until the end of the dry season. although the small sample size precludes any conclusions on the population trends of this antelope in the park, the lamb to ewe ratio in the knp is greater than the lamb to ewe ratio of the relatively stable loskop population for the same period (table 3). 64 table 2 group size, composition, and site characteristics rif mountain reedbuck sighted in the kruger national park during september 1970 habitat group composition characteristics time 1515 0840 1012 1655 0705 0810 0830 0855 0710 group size male female imm. (>1 a) (>1 a) «1 a) 1 3 2 6 1 (whistle only) 3 5 4 1 2 2 2 2 1 2 1 table 3 unaged 4 1 1 1 slope h t. on recent face slope burn w 3/4 yes sw 3/4 yes s 3/4 yes w 1/4 yes w 3/4 yes ? 1/4 yes sw 2/3 yes nw 4/5 yes e 3/4 no comparison rif mountain reedbuck sightings in the loskop dam nature reserve during august-september 1970 with sightings in the kruger national park during september 1970* lamb: ewe n males females lambs unaged ratio (> 1 a) ( > 1 a) ( < 1 a) (lambs per 100 females) loskop 50 14 25 6 5 24 kruger 26 3 10 5 8 50 * groups sighted in intensive study areas at loskop have been excluded. the 24 km 2 section of the kwa machiyaliwane drainage surveyed yielded a minimum count of 25 mountain reedbuck. exclusion of 3 km 2 of flats associated with the main streamcourse gave a minimum density of 1,2 mountain reedbuck per km 2 of slope habitat. a similar treatment of sightings at loskop during june-august 1973 (irby 1973), gave a minimum count of 205 individuals on 49 km 2 for a minimum density of 4,2 per km 2 of slope habitat. while the two counts are not directly 65 comparable, since the search intensity was greater at loskop, the data indicate that the kwa machiyaliwane sub-population is less dense than that of loskop. the absence of sightings around khandizwe hill and along the emcwebeni drainage may indicate a decline in the population, and a subsequent decrease in the range occupied, or only insufficient effort expended to locate individuals in a very sparse population. new growth on burned grass clumps in the kwa machiyaliwane area could have concentrated mountain reedbuck in that region. a mass migration from the emcwebeni drainage to this burn seems unlikely since comparable burns, with no mountain reedbuck, were observed in the emcwebeni drainage, and mountain reedbuck are territorial. acknowledgements i wish to thank dr. u. de v. pienaar and the national parks board of trustees for allowing me to work in the kruger national park and the staff of the park for the guidance and assistance provided during my stay. dr. s. l. beasom deserves thanks for reviewing this manuscript. funding for this study was provided by the mammal research institute, university of pretoria; the caesar kleberg research program in wildlife ecology, texas a & m university; and the national science foundation, washington, d.c. material presented in this paper will be used for partial fulfillment of the requirements for a ph.d. degree from texas a & m university. this manuscript is texas agricultural experiment station technical article no. ta 12059. references irby, l. r. 1973. mountain reedbuck in the transvaal. unpublished annual report of the caesar kleberg research program in wildlife ecology. 108-115. pienaar, u. de v. 1969. predator-prey relationships amongst the larger mammals of the kruger national park. koedoe 12: lo8-176. 66 page 1 page 2 page 3 page 4 article information author: susan snyman1,2 affiliations: 1environmental economics policy research unit, university of cape town, south africa2wilderness safaris, johannesburg, south africa correspondence to: susan snyman postal address: po box 5219, rivonia 2128, south africa dates: received: 01 feb. 2013 accepted: 30 nov. 2013 published: 24 june 2014 how to cite this article: snyman, s., 2014, ‘assessment of the main factors impacting community members’ attitudes towards tourism and protected areas in six southern african countries’, koedoe 56(2), art. #1139, 12 pages. http://dx.doi.org/10.4102/ koedoe.v56i2.1139 copyright notice: © 2014. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. assessment of the main factors impacting community members’ attitudes towards tourism and protected areas in six southern african countries in this original research... open access • abstract • introduction    • conceptual framework and literature review • research method and design • results    • reasons given for the importance of conservation    • probit models for tourism and protected area attitudes • ethical considerations • trustworthiness    • reliability    • validity • discussion • conclusion • acknowledgements    • competing interests • references abstract top ↑ in southern africa, many early conservation efforts from the late 1800s and early 1900s either displaced local communities or restricted their access to natural resources. this naturally affected community attitudes towards protected areas and efforts were later made to rectify growing tensions. in the last few decades of the 20th century, these efforts led to conservation and ecotourism models that increasingly included communities in the decision-making and benefit-sharing process in order to garner their support. although the results of these policies were mixed, it is clear that the future success of conservation and, consequently, ecotourism in many areas will depend on the attitudes and behaviour of communities living in or adjacent to protected areas. managing and understanding community expectations and attitudes under varying socio-economic circumstances will lead to more efficient, equitable and sustainable community-based conservation and ecotourism models. this study was based on 1400 community interview schedules conducted in botswana, malawi, namibia, south africa, zambia and zimbabwe, allowing for an accurate comparison of attitudes across countries, protected areas and communities. the results highlighted important demographic and socio-economic factors to consider in terms of understanding the attitudes of those living in and around protected areas. suggestions were put forward for managing community relationships and garnering long-term support for protected areas and ecotourism.conservation implications: it was observed that, in general, community members living in or adjacent to conservation areas in southern africa have an understanding and appreciation of the importance of conservation. formal education was found to positively impact attitudes and human–wildlife conflict negatively impacted attitudes, highlighting important policy focus areas. introduction top ↑ it is clear that sustainable african conservation has to include local populations (hoon 2004; musumali, larsen & kaltenborn 2007). many have argued that if communities receive benefits from protected areas (pas) and ecotourism they will tend to hold positive attitudes towards conservation as a land use and to care for the natural resources in their area (currie 2001; emptaz-collomb 2009; hulme & murphree 2001; mcneely, in groom & harris 2008; wang & pfister 2008; waylen et al. 2009). ecotourism allows local communities to receive benefits from conservation, both directly, through wages and salaries, and indirectly, as suppliers of goods and services. numerous authors have argued that community support is critical to the long-term success of ecotourism operations and their associated pas (alexander 2000; allendorf et al. 2006; chandralal 2010; gillingham & lee 2003; makindi 2010; nepal 2002; newmark et al. 1994; sifuna 2010). an analysis of community attitudes towards pas across southern african would highlight the determinants of community support. such information may encourage governments and ecotourism operators to focus on these factors in policy decisions. although pas and ecotourism are interlinked, community attitudes towards them may be sensitive to the direct costs and benefits of ecotourism, particularly employment and human–wildlife conflict. managing community expectations of ecotourism requires an understanding of the factors driving their attitudes. education and awareness-raising programmes also benefit from an understanding of community attitudes (allendorf et al. 2006; chidakel 2011; sifuna 2010; simelane, kerley & knight 2006). as emerton (1999) pointed out, benefit distribution is a necessary, but not sufficient, condition for communities to engage in wildlife conservation (snyman 2012a). the understanding of local residents’ attitudes, by the managers of pas and eco-lodges, will naturally affect their interactions with the community and it is hoped that this understanding will allow more focused planning in pas and associated ecotourism operations. the specific objectives of the present study were to, (1) enrich the tourism and pa literature with an analysis of factors impacting rural community members’ attitudes to tourism and pas across six southern african countries and (2) provide useful policy options and practices for pa and ecotourism managers. from these objectives three research questions emerged: • what demographic and socio-economic factors drive rural community members’ attitudes towards tourism and pas? • how do rural community members’ attitudes differ across six southern african countries? • what policy implications emerge from an understanding of factors impacting community members’ attitudes? conceptual framework and literature review following allport and mcdougall and munro (both in wang & pfister 2008:85) we shall treat ‘attitudes’ as ‘respondents’ particular feelings and perceptions towards the stated questions relating to tourism and/or conservation’. wang and pfister (2008) found that examinations of attitudes towards tourism showed that these are influenced by people’s values and personality and are therefore slow to change. they are also influenced by factors invisible to outsiders (allendorf et al., in waylen et al. 2009) making them difficult to observe or understand.in addressing individual community members’ attitudes, social exchange theory assumes that potential beneficial outcomes will create positive attitudes towards tourism (andereck et al. 2005; teye, sönmez & sirakaya 2002). the theory postulates that individuals perceiving net benefits from an exchange are likely to view it positively and those perceiving net costs are likely to view it negatively (snyman 2012a). this approach is supported by numerous studies which showed that residents dependent on tourism for support, or who perceive it offering net personal benefits, tend to view its impacts more positively than others (brunt & courtney and child, in groom & harris 2008; various sources, in andereck et al. 2005:1061; shibia 2010; walpole & goodwin 2001; wang & pfister 2008). the fact that people with lower household incomes are often less supportive of pas and tourism is largely explained by needs theories. such theories argue that an individual’s basic needs are attended to first before higher needs such as supporting community, conservation or tourism initiatives (doyal & gough and maslow, in emptaz-collomb 2009). emptaz-collomb (2009) stresses that it would therefore be prudent for companies or individuals engaged in such initiatives to assist in improving the lives of local people through improving access to and the quality of education, health, transport and communication. in the long run, this would create a more supportive environment for tourism and conservation projects and ensure their sustainability. government, non-governmental organisations (ngos) and the private sector can all play a role in this through infrastructural and other development projects. the importance of understanding attitudes lies in their theoretical connection with behaviours (manfredo et al., in lepp & holland 2006). the links between attitudes and behaviours are, however, not automatic (scanlon & kull 2009). even though nunkoo and ramkissoon (2011) found that residents in their mauritian study engaged in behaviours congruent with their attitudes, this will not always be the case. although past research has shown that many communities hold positive attitudes towards conservation (alexander 2000; currie 2001; de boer & baquete 1998; mehta & heinen 2001; sekhar 2003; weladji, moe & vedeld 2003) and ecotourism development (chandralal 2010; lepp 2007; mehta & kellert 1998), there are a number of different factors affecting attitudes. some studies (gadd 2005; lepp & holland 2006; rodriguez 2008) have also found that communities, for varying reasons, may hold negative attitudes towards conservation. some studies have also not found a correlation between economic benefits from tourism and positive attitudes to conservation (stem et al. 2003; walpole & goodwin 2001) or suggested that economic benefits alone were insufficient to encourage conservation (stem et al. 2003; stronza & pégas 2008). stem et al. (2003) and stronza and gordillo (2008:450) found that non-economic benefits, such as new skills, broader experiences in managing projects and people, ideas exchange, expanded circles of contacts, empowerment and support for community efforts, could also influence attitudes towards pas and tourism. these non-economic benefits are, however, often more difficult to measure and assess. creating and maintaining positive attitudes towards pas is especially important when other mechanisms for changing behaviour, such as regulation, are inappropriate or ineffective (waylen et al. 2009). over the years, regulation has often failed to ensure conservation success and the resultant need for a more appropriate and effective means of ensuring conservation has arisen. in summary, studying community members’ attitudes towards tourism and pas is important for a number of reasons (snyman 2012a): • it can disclose whether or not negative attitudes exist towards a pa and/or tourism operation, which, in some cases, may help to explain behaviour (lepp & holland, in anthony 2007). • it can inform policymakers and managers about which factors influence attitudes and this can assist with prioritising avenues for action (anthony 2007; browne-nuñez & jonker 2008). • it can reveal opportunities for improving relationships and outreach programmes with communities living adjacent to pas (anthony 2007). • it can assist in developing appropriate benefit-sharing and cost-minimising programmes for communities based on their attitudes and experiences. • it can highlight areas important for education and training programmes. overall, there is widespread consensus that community attitudes matter and these attitudes may vary over time and be influenced by various factors (e.g. changing incomes, land management arrangements, etc.) (andereck & vogt 2000; anthony 2007; gillingham & lee 1999). factors such as the level of education, recent experiences and degree of politicisation may also play a role. in an attempt to capture the relevant issues, this article looks at the impact of a number of socio-economic variables on community attitudes across different countries. despite the many attitude studies that have been conducted, they are not always comparable as they have been conducted under different circumstances and with different measurement tools. this study was however conducted consistently over six study countries allowing for a more accurate comparison of attitudes across countries, pas, tourism operations and communities. research method and design top ↑ in this study (adapted from snyman 2012a, 2013) extensive socio-economic interviews were conducted in over 30 rural communities, covering more than 16 different ethnic groups and an average of approximately 25% of households in each study village (ranging from 10% to 84%). the selection of study sites was dictated by the presence of a community–ecotourism relationship or partnership, or because the community lived in or adjacent to the conservation or pa where the ecotourism operation was located, or a combination of these (in this study all ecotourism operations were owned or managed by wilderness safaris, see http://www.wilderness-safaris.com). conservation areas need not be government owned, but can include areas owned and/or managed by communities, private individuals, ngos or companies (makindi 2010). the common feature of the areas covered in this study is that all have been specifically set aside for conservation. at one extreme, this involved the total relocation of local people formerly living in the area (e.g. the makuleke community in south africa); at the other extreme are conservation areas in which people still live and have access to the natural resources (e.g. the namibian conservancies). few of the study sites had accurate or recent maps of households, dictating other means of sample size selection. sample sizes varied with the size of the community being surveyed. an attempt was made to interview at least 10% of households in all adjacent communities. logistically, however, this was not always possible. if the survey area was small, then the majority of the households in the area were interviewed. if the survey area was large, then for logistical reasons, a subset of villages and households was selected. household numbers were obtained from the latest census figures and/or from the respective headman, chief or community trust. households were selected randomly, either by walking through the village interviewing every second household (or the next household where someone was available) or interviewing a household member in a public area, for example, a local shop or meeting area. in total, 1400 community interviews were conducted in rural villages either within or adjacent to pas (see table 1). attempt was made to interview the household head, but if they were not available, then either the spouse, or next oldest person in the household was interviewed. table 1: details of communities surveyed in each country. as the concept of community is central to the analysis in this article the term needs to be clarified. for the purposes of this article, we follow borrini’s (as cited in borrini-feyerabend, kothari & oviedo 2004:9) description of it as ‘a human group sharing a territory and involved in different but related aspects of livelihoods – such as managing natural resources, producing knowledge and culture, and developing technologies and practices’, whilst a local community is a group who interact regularly or who influence one another’s daily lives. in this study, such local communities may be mobile, permanently settled or semi-nomadic such as the himba people of north-west namibia. the communities are found living either within or adjacent to the pa, or in some cases, having left the pa, are now living further afield (e.g. the makuleke community which is located two hours’ drive from pafuri camp, but is still impacted by it). all communities described in this article are either directly or indirectly affected by the conservation and ecotourism strategies in their area, whilst their activities, in turn, impact nearby pas and ecotourism operations. the camps associated with the study communities were all ecotourism camps in terms of their location and the activities offered. defining ecotourism and its relationship to conservation is also contextually important at this stage. in summarising the literature, de witt, van der merwe and saayman (2011:1139) suggest that the key principles of ecotourism are that it should foster a genuine interest in nature, contribute to conservation, respect and conserve local culture, make non-consumptive use of natural resources, yield benefits to the local community and create tourist awareness of conservation and local community issues. based on this definition, ecotourism in this article is taken to include activities which are nature-based and culture-based, sustainable, promote conservation and provide benefits to local people in the area. it is therefore not simply tourism that is based on the sale of access to an interesting natural area, but tourism that also provides benefits to local communities (snyman 2013). the interviews were conducted by both male and female interviewers and local translators were used in circumstances where the respondent could not speak or understand english. the interview schedules contained questions relating to demographics, social welfare and living standards, education, employment patterns, income and expenses, health and safety and attitudes toward tourism and conservation. this article focuses on the sections relating to attitudes. each interview was conducted verbally, with the interviewer completing the questionnaire during the interview. each interview took approximately 20 min – 45 min depending on the respondent’s educational level and whether or not translation was required. the interview schedule consisted of a structured set of questions, with the majority being close-ended and some having the option for further explanation. communities associated with a single ecotourism enterprise, wilderness safaris (ws), were surveyed in this study because it was the only ecotourism company that had parallel ecotourism operations, operating according to a standard policy framework, over the six anglophone countries in the region. the use of a single company made for ease of comparison because the head office imposes a consistent management style over its different camps in southern africa. the company itself wished to quantify the impact of its ecotourism operations on rural communities and gave the author access to its camps and staff and to the communities with whom they engage. the process followed does, however, mean that some caveats attach to this research: • although the camps and communities were diverse (with varying land management systems, ethnic groups and tourism camp price ranges), as only one ecotourism operator was included in the analysis, there could be limitations to the generalisability of the research. • the author was employed by ws to conduct a study on the impact of the company’s activities on rural communities. this study was, however, performed as an independent researcher looking to discover the realities of ecotourism and community development and was in no way influenced by the company. • local residents would have associated interviewers with ws because of the use of ws vehicles in some areas and through the introduction process. this may have biased responses to questions about ws. it is however impossible to predict the direction of the bias a priori; some respondents may have been strategically negative in order to ensure changes or positive in order to win favour with the private sector operator in the area (allendorf et al. 2006). the results showed both positive and negative responses in all areas and many respondents were clearly comfortable expressing negative responses. • the presence of the researcher during the administration of the interview schedule may have influenced some respondents and their answers to attitudinal questions regarding tourism and conservation. the bulk of the interview schedule was, however, socio-economic in nature and this should not have proven problematic. there remained the risk of strategic bias. when it was felt that this was occurring, the author re-iterated to the respondent that the interview was anonymous and honest answers were required. all data collected were analysed using spss version 12 (2004) and stata version 10.2 (2008), as well as a combination of descriptive statistics, t-tests and probit models. results top ↑ table 2 summarises the respondents’ demographic details in order to provide context to the other results. these results are both aggregated across the southern african region and, in some cases, disaggregated by country to show specific country nuances. table 3 shows the results for the tourism and conservation attitudes by location. overall, the zambian community respondents held the most positive attitudes to tourism and conservation. this is most likely because there were few employment opportunities in the area other than ecotourism or with the zambian wildlife authority. table 2: summary of respondents' demographic characteristics. table 3: attitudes towards tourism and conservation by sampled communities. the importance of tourism and conservation in these remote, rural areas is highlighted by the percentage of respondents (45%) who had family members working in tourism and/or conservation. this result does, however, need to be interpreted with caution in terms of total employment impact. there is a risk of double-counting, as many village respondents are related and a number of separate households may have been referring to the same employed family member. in all, 87% of community respondents felt that conservation was important, despite 83% of them having problems with wild animals. reasons given for the importance of conservation respondents who felt that conservation was important were asked for their reasons. a number of different rationales emerged (see table 4). many respondents said that it was important for tourism. other reasons included: for their children and/or the future and to be able to use the natural resources in the future for food, firewood, et cetera. some respondents said because the ‘trees bring rain and/or prevent wind’. some respondents said that conservation was important for tourism, but not for people’s crops or livestock and others said that it was important to conserve trees and plants, but not animals because they were dangerous. a number of respondents gave more than one reason and others said that they knew it was important (often because they had been told at school), but did not know why. in some areas, respondents said it was important because the government said so. table 4: main reasons given for the importance of conservation by location. probit models for tourism and protected area attitudes probit models were run for each of the attitude questions. the demographic variables in the probit were chosen as a result of a descriptive statistical analysis and an extensive literature review of factors found to impact community attitudes. table 5 presents the probit results, reporting marginal effects for each attitude question. table 5: probit results reporting marginal effects for each attitude question. although factors affected attitudes differently, there was a trend observed in terms of the impact of education on attitudes, with respondents reporting higher levels of formal education being more positive. this was statistically significant in four of the questions. those who were more educated also had fewer problems with wild animals.statistically, demographic variables had varying impacts on attitudes. age was however, surprisingly, positively related, suggesting that older respondents were more positive about tourism and pas than younger respondents. with respect to age of the household head however, this relationship was largely negative, with respondents with older household heads being less positive about tourism and pas. areas with higher population densities were shown to be less positive about tourism and pas. distance had mixed impacts on attitudes, as did the number of years that the tourism camp had been operating. respondents with more household income sources tended to be more positive about tourism having a positive change in their village, as well as about conservation being important. a consistent issue raised in all study communities was that of wild animals interfering with households’ livelihoods and, in some cases, personal safety. one female respondent in malawi told of a night she had been terrified inside her house with her children, whilst a bull elephant tried to push the house over to reach her maize stored inside. this highlights the real threat wildlife can pose to rural households and their livelihoods. table 6 shows that across all communities surveyed, 83% of respondents had problems with wild animals at home. table 6: percentage of respondents who had problems with wild animals by location. elephants were the most frequently mentioned animals (55%) that caused problems, followed by lions (28%). other animals mentioned included cheetah, hippo, leopard, hyaena, baboon and jackal. in general, human–wildlife conflict (hwc) resulted in less positive attitudes towards ecotourism in the study areas. respondents who had family employed in tourism or conservation were generally observed to be more positive towards tourism and pas and had fewer problems with wild animals. having family employed in tourism or conservation positively impacted four attitude questions and could be because unemployment and poverty levels are high; having a family member employed in tourism or conservation therefore has a significant impact on a household and therefore on attitudes. the country variable was significant in the ‘reducing poverty’ question only, indicating that, in general, communities across southern africa did not differ significantly in their attitudes towards tourism and pas. ethical considerations top ↑ the interview schedule was approved by the university of cape town’s ethics committee. all respondents were told the reason for the study and signed a written consent form prior to the interviewer beginning the interview. respondents were told that the interview schedules were confidential and their participation in answering all questions was voluntary. this resulted in some questions not being answered. non-response to questions did not cluster on particular questions, as no particular question had a greater non-response rate than any other question. trustworthiness top ↑ although every effort was made to ensure the interviews and data analysis were conducted correctly, certain assumptions and limitations of the research necessarily arise, which should be factored into the data analysis and interpretation. reliability cross-cultural research can have limitations in terms of respondents reacting or answering differently because of the presence of an expatriate working in a developing country that was previously colonised (bruyere, beh & lelengula 2009). it was hoped that the use of local translators (who received informal training from the author) would help minimise this limitation. the author and other interviewers (where not local) also made efforts to learn some of the local languages and customs, to greet respondents in their own language and to help them feel relaxed. de boer and baquete (1998) warned that formal questionnaires become a drawback when people are unwilling to express negative opinions or attitudes to a third party, in particular where interviewees are reluctant to confess to illegal exploitation practices, such as snaring or collecting plants in a restricted area. questionnaires are, however, a cost-effective method of research. this needs to be kept in mind when analysing the data collected on opinions and attitudes to tourism and conservation in the present study. that negative attitudes were expressed by some respondents suggests that they did not feel constrained. in order to minimise bias and inaccurate information, respondents were also informed at the beginning of the interview that it was anonymous, was part of a research study, and that their responses would be aggregated and impossible to identify in the larger study. interviews run a number of other risks including the researcher leading the respondent, variation in the delivery of the survey between interviewers, respondent anticipation or desire to please the researcher and discrepancies between what people report and what they actually do or feel (de boer & baquete 1998; gadd 2005). despite this, surveying attitudes and quantifying them is necessary if one is to compare attitudes towards conservation and ecotourism in different regions or within the same region over time (gadd 2005). every effort was made to keep the interviews uniform and to ask questions in such a manner as to reduce bias or at least keep it consistent. in order to render any existing bias relatively constant, the author conducted over 1000 of the interviews herself. eight other interviewers assisted across the six countries. validity the sample size was not standardised in all study countries because of logistical constraints in some areas where communities were large. these differences in the percentage of the community interviewed could result in some issues relating to external validity. it was however felt that all sample sizes were sufficient in the areas surveyed and no new information was found after a certain percentage (approximately 100 households) of the community had been interviewed. the zambian sample is included in the analysis because, despite being small, it was felt to be relevant and representative of the area where the interviews were conducted. inferences drawn from the zambian results should however be viewed with caution. discussion top ↑ the present study analysed the impact of various demographic and socio-economic variables on attitudes of community members in six southern african countries and, as in past studies, (allendorf et al. 2006; simelane et al. 2006) found mixed results relating to various factors. individual’s attitudes, although generally long term, can change over time and also vary between and within different communities and countries. there are a number of factors that cannot be controlled or manipulated, so it is important that policy is aimed at those factors that can, to some extent, be influenced – for example, education, land ownership, et cetera. in general, community members in this study felt that tourism creates employment and can help reduce poverty. if tourism-related jobs were to disappear in an area (for example in the zambian study area where tourism is one of few employers), it is unlikely that people would feel as positive about tourism and pas in the future. this highlights the importance individuals attach to tourism in terms of employment, household income, et cetera. it is critical that this is managed in an appropriate manner to ensure that expectations are met and that there is not large-scale disappointment on the part of communities. this is highlighted by the fact that in communities where tourism had been operational for longer, respondents were less positive about conservation than those areas where tourism was new. allendorf (2007) highlighted that although there has been little attention given to the importance of non-economic benefits that residents may value in developing countries, studies indicate that people do value pas for non-economic reasons, such as ecosystem services, for the benefit of future generations and the conservation of wildlife. this was supported by the findings in the present study. although many respondents felt that conservation was important because of the income to be derived from it through employment and tourism, many felt that it was important for their children and future generations, as well as for the wood, thatch and food it provides. as found in the present study, various other studies have found differing results relating to the impact of demographic variables on attitudes towards tourism and conservation (akyeampong 2011; allendorf et al. 2006; baral & heinen 2007; currie 2001; de boer & baquete 1998; gadd 2005; gillingham & lee 1999; kideghesho, røskaft & kaltenborn 2007; larson 2010; mbaiwa & stronza 2011; mehta & heinen 2001; sarker & røskaft 2010; sekhar 2003; shibia 2010; stem et al. 2003; tessema et al. 2007; teye et al. 2002; weladji et al. 2003). it appears, therefore, that it is difficult to use demographic variables to predict attitudes. there are, however, some areas of commonality in terms of demographics that can be used as potential predictors. other studies (anthony 2007; currie 2001) have largely found that younger respondents hold more positive attitudes towards tourism and pas. this may be because many younger people have more education than older people, are less reliant on natural resources and therefore less affected by a lack of access to them and/or have alternative livelihoods that reduce the risk that they face. age was significant only in the ‘conservation important’ attitude, with a positive coefficient, indicating that as age increases, there was an increase in the predicted probability of having a positive attitude. this contradicts past research relating to the impact of age. the number of children was only significant in the ‘problem animal’ attitude question, with a positive relationship. this could be because respondents with more children perceived more problems with wild animals, as any conflict would negatively affect them as they had more mouths to feed. communities living in or adjacent to pas frequently incur direct costs associated with living next to wildlife, such as damage to crops, loss of livestock and, occasionally, loss of human life. other costs associated with wildlife include the additional direct cost of guarding crops and livestock by paying someone to look after them or the opportunity cost of protecting them by giving up one’s time which could have been put to a more productive use. this often results in a disruption to children’s schooling as they are kept out of school to guard household fields (pers. obs. author, december 2009). logic implies that those who are negatively impacted by wildlife are likely to have less positive attitudes towards conservation and consequently ecotourism. this is detrimental to the long-term success of pas and ecotourism and therefore requires attempts to mitigate hwc in rural areas. in countries where there was a high percentage of respondents who had problems with wild animals combined with less positive attitudes (e.g. malawi), it is even more important that the surrounding communities receive benefits from ecotourism and conservation, or measures to mitigate hwc. hill (2004) argues that people who believe that they do not have control over a conflict situation are often more likely to inflate their perceptions of risk and dublin and hoare (2004:274) emphasise that it is often the potential for suffering large losses, especially at harvest time, that is a major factor influencing rural communities’ attitudes, rather than actual losses. this could explain the high incidence of hwc expressed in the surveys in the present study. the close-knit community life of many rural african villages could also result in an exaggeration of hwc based on the perceptions of residents, rather than the actual number of incidences (romañach, lindsey & woodroffe 2007; woodroffe, thirgood & rabinowitz 2005). in general, respondents who said they had problems with wild animals were found to have less positive attitudes towards tourism and conservation. walpole and thouless (2005:130) highlight that tourism will only improve tolerance towards wildlife where the benefits of tourism actually reach those bearing the costs of living with wildlife and where the local communities can understand and act upon the linkages between tourism benefits and wildlife conservation. those bearing the costs are frequently not the same people receiving the benefits and this needs to be taken into consideration for long-term sustainability (walpole & thouless 2005). based on past studies (groom & harris 2008; mehta & kellert 1998), it was assumed that those with higher monthly household incomes would have more positive attitudes towards tourism and pas. this assumption is based on the fact the wealthier households have the ‘luxury’ of being able to either enjoy the tourism, pas and/or are not as greatly affected by the negative impacts of conservation, for example hwc, loss of access to natural resources, et cetera. income had mixed effects on attitudes in this study and suggests that household wealth is not always a driver of positive attitudes. of importance to policymakers is that groom and harris (2008:250) found in their kenyan study that although financial incentives from wildlife can improve community attitudes towards wildlife and conservation, the actual distribution of benefits is more important in shaping attitudes. this is supported by the results from the torra conservancy in namibia, where respondents’ attitudes were still (i.e. in 2009) influenced by a dividend payout in 2003 (see snyman 2012a). the lack of consistency of the overall importance of household income as a determinant of attitudes in the probit models suggests that income, as an incentive, is not the only incentive that matters to these communities. past studies have found that increased education impacted positively on attitudes towards tourism and pas (see chidakel 2011; larson 2010; shibia 2010; tessema et al. 2007; teye et al. 2002). teye et al. (2002), in their ghanan study, surmised that a more positive attitude towards tourism by more educated people could be explained by the fact that much has been written in english regarding the benefits of tourism in both print and electronic media, therefore educated people would be more familiar or aware of the potential benefits than those with less education. overall in the present study, formal education was found to have a statistically significant positive impact on individual’s attitudes to tourism and pas, highlighting the importance of formal education in terms of garnering support from rural communities. the benefits of formal education are numerous in terms of long-term poverty reduction, as well as biodiversity conservation (see snyman 2012c). one would surmise that if a respondent had a family member, or in the case of the community respondents, was themselves, employed in tourism and/or conservation, then they would have more positive attitudes towards tourism and pas. this is premised on the fact that they, or their family members, would be receiving direct, tangible benefits from tourism and/or pas and would therefore be more positive towards it. this premise was supported by the results in this study. in fact, having a family member employed in tourism or conservation was one of the most significant variables in a number of the analyses. this illustrates that it is not necessarily required that someone has to be receiving direct benefits themselves to be aware of the benefits that can result from tourism and pas. conclusion top ↑ it is frequently the vulnerability of poorer households and the risks they face that leads to less positive attitudes towards tourism and pas. as discussed, the costs that communities have to bear are often high. if there are no concomitant benefits associated with these costs, then it is unsurprising that households would hold negative attitudes towards pas and the associated tourism operations in the area.not only are there numerous direct factors affecting people’s attitudes to tourism and pas, they may also be influenced by costs and benefits that accrue to others, including those in different households (emerton, in sandbrook & adams 2012). this would explain why some households, who are not directly benefiting from, or negatively affected by, tourism or pas, still hold certain attitudes towards it. a factor that was shown to affect attitudes to conservation and that can be managed in order to ensure future positive attitudes was the length of operation of the ecotourism camp in the area. in general, the longer the camp had been open, the more negative respondents were about conservation; although, they interestingly tended to be more positive about tourism. this does not bode well for the long-term success of pas and needs to be addressed. the main reason for this is likely because of unmet expectations, unfulfilled promises and high levels of hwc. the management of community expectations and benefit distribution before an ecotourism camp starts operating, as well as during the operational phase, including realistic goal setting, is important to the long-term success of ecotourism in pas. it will help to reduce negative attitudes that can result from unmet expectations and the consequent dissatisfaction of communities living adjacent to pas. formal employment of any kind was shown to have a largely positive impact on attitudes as it allows a diversification of household livelihoods and reduces the risk they face. the important point to note is that there are few formal employment opportunities in these remote rural areas, other than in ecotourism. the results of this article highlight the diverse array of factors affecting people’s attitudes towards ecotourism and pas. monetary benefits from ecotourism alone will not serve to improve local people’s attitudes towards ecotourism and pas, as there are a number of factors shaping attitudes. these include receipt of tangible, as well as intangible benefits, demographic factors, local economic situation, past beliefs, cultural beliefs, land ownership systems, population density and the diversity of livelihood strategies available to households in the area. the more livelihood strategies people have available to them, the less dependent they are exclusively on natural resources for survival and livelihoods. it is premised that people will then value conservation and pas more. dependence on ecotourism as the sole livelihood can also be risky and vulnerable to external shocks. ideally, ecotourism therefore needs to be part of a diverse livelihood portfolio. conservation, for most rural africans, is an investment for present and future value, with the main goal being the maintenance or enhancement of their livelihoods (hulme & murphree 2001). ecotourism can therefore assist communities in earning much-needed income, as well as assist them in conserving their natural resources in order to maintain or enhance other livelihoods. such a scenario would however only apply in areas where communities have access to the natural resources in the area where the ecotourism camp operates (snyman 2013). communities who are excluded from pas will require further incentives to conserve natural resources, as they will not be benefitting directly from their conservation. global benefits to be derived from ecosystem services are also important and communities can play a role in this through watershed protection, preventing deforestation, et cetera. the importance of education in terms of obtaining employment, as well as more positive attitudes is important in terms of the sustainability of tourism and conservation as land uses. increasing access to formal education, improving education infrastructure, as well as implementing awareness and education campaigns in communities can serve to increase people’s knowledge and awareness of pas and tourism (snyman 2013). this, in turn, may result in a greater willingness to accept the costs of living with wildlife and more positive attitudes towards pas and tourism in the area. positive attitudes do not however necessarily suggest that behaviours will also promote conservation and tourism. poor rural households face many economic and time constraints that can prevent them from supporting conservation (snyman 2013). parry and campbell (in emptaz-collomb 2009:101) suggest that improving the living conditions and social welfare of rural people is therefore an important part of any conservation strategy. ecotourism can play an important role here, through donations towards community development projects, tourism-related infrastructure developments, directly through wages and salaries and indirectly as suppliers of goods and services. in summary, some implications for management drawn from this research include: • in areas where government owns the land (in this study, national parks) and there is no contractual community involvement (e.g. malawi, zambia and zimbabwe), there have to be benefits, both tangible and intangible, received by the community, as well as a mitigation of the negative impacts associated with conservation (i.e. hwc). outreach programmes, introduced by the private sector tourism operator, in communities abutting the park could include educational programmes as well as social welfare projects. such programmes would serve to link pas and tourism directly to benefits (snyman 2012b). • government, ngos or the private sector need to raise awareness relating to ecotourism, conservation and pas. ecotourism operators can play an important role in this through environmental talks and conservation and tourism awareness-raising days in communities, as well as offering environmental lessons and game drives to community school children, as many have never been inside the pa adjacent to their homes (snyman 2012b). • there must be a clear, structured process of setting and managing expectations prior to an ecotourism operator starting in an area, as well as through the operational phases. • overall, it is not only important to maximise benefits to communities, there needs to be a concomitant process of minimising costs, as often there are more who will bear the costs than there are those who will benefit from the conservation and ecotourism in the area (snyman 2012a). • alternative livelihoods (such as ecotourism employment) may assist in steering households away from absolute dependence on the consumptive use of natural resources for survival, which could, in turn, promote biodiversity conservation and long-term sustainable use, as well as positive attitudes (snyman 2012b). • formal education is important and has been shown to influence attitudes. improved educational infrastructure and improved access to education (e.g. scholarship programmes) should therefore be a priority in rural areas. managing community expectations, through an understanding of community members’ attitudes under varying socio-economic conditions, will lead to more efficient, equitable and sustainable community-based conservation and tourism models. also important for long-term sustainability is that, over and above this understanding, one needs to know what factors directly affect behaviours. future research should focus on monitoring behaviours related to pas, for example collection of natural resources, engagements with pa and ecotourism staff, et cetera. positive community behavioural changes towards pas and natural resources will ensure their long-term sustainability. a focus on formal education, improving social welfare and increasing the local linkages from ecotourism operations will go a long way in improving local community attitudes towards tourism and pas. acknowledgements top ↑ competing interests as stated above, the author was employed by ecotourism group, wilderness safaris, to conduct a study on the impact of the company’s activities on rural communities. however, the study itself was 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conflicts in a changing world’, in r. woodroffe, s. thirgood & a. rabinowitz (eds.), people and wildlife: conflict or coexistence?, pp. 388–405, the zoological society of london and cambridge university press, cambridge. article information authors: stoffel p. bester1,2 ronell r. klopper3,4 hester m. steyn1 hugo bezuidenhout5,6 affiliations: 1national herbarium, south african national biodiversity institute, south africa2school of environmental sciences and development, north-west university, south africa 3biosystematics research & biodiversity collections division, south african national biodiversity institute, south africa 4department of plant science, university of pretoria, south africa 5scientific services, south african national parks, south africa 6applied behavioural ecology and ecosystem research unit, university of south africa, south africa correspondence to: hester steyn postal address: private bag x101, pretoria 0001, south africa dates: received: 13 oct. 2011 accepted: 07 aug. 2012 published: 12 nov. 2012 how to cite this article: bester, s.p., klopper, r.r., steyn, h.m. & bezuidenhout, h., 2012, ‘new plant records for tankwa karoo national park, south africa’, koedoe 54(1), art. #1066, 9 pages. http://dx.doi.org/10.4102/ koedoe.v54i1.1066 copyright notice: © 2012. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. new plant records for tankwa karoo national park, south africa in this checklist... open access • abstract • introduction • research method and design • ethical considerations • results and discussion • conclusion • acknowledgements    • competing interests    • authors' contributions • references • appendix 1: detailed information and relevant specimens for new plant records from tankwa karoo national park • apocynaceae    • quaqua acutiloba (n.e.br.) bruyns    • quaqua parviflora (masson) bruyns subsp. gracilis (c.a.lückh.) bruyns    • stapelia arenosa c.a.lückh.    • stapelia surrecta n.e.br.    • tridentea parvipuncta (n.e.br.) l.c.leach subsp. truncata (c.a.lückh.) bruyns    • tromotriche thudichumii (pillans) l.c.leach • asphodelaceae: alooideae    • aloe microstigma salm-dyck    • aloe falcata baker    • aloe variegata l. • hyacinthaceae    • lachenalia ameliae w.f.barker • iridaceae    • moraea tanquana goldblatt & j.c.manning • scrophulariaceae    • nemesia suaveolens k.e.steiner • pteridophyta: sinopteridaceae    • cheilanthes parviloba (sw.) sw.    • cheilanthes induta kunze    • cheilanthes namaquensis (baker) schelpe & n.c.anthony abstract top ↑ the tankwa karoo national park has been enlarged from 27 064 ha to 143 600 ha. this whole area is severely under-collected for plants in general and therefore it was an obvious target for the south african national parks (sanparks) programme, a component of the pretoria national herbarium (pre) plant collecting programme. this programme not only aims to survey national parks that have been poorly surveyed, but also inadequately known taxa, unique habitats, remote and inaccessible areas and plant species flowering at irregular times, especially after events such as fire or unusual timing of, or high, rainfall. general collecting in the tankwa karoo national park has already led to the description of two new taxa, from two families. it furthermore resulted in new distribution records for the park and for the northern cape province. these are reported on here.conservation implications: although the tankwa karoo national park falls within the succulent karoo biome (a biodiversity hotspot of international importance), information on its plant diversity is insufficient because it is an under-collected area. results of this study will guide conservation and supply occurrence and distribution data required to compile management plans for the park. introduction top ↑ the tankwa karoo national park (tknp) is a relatively new addition to south africa’s treasure of national parks and was only declared a national park in 1986 (rubin 1998). the original park consisted of eight farms amounting to 27 064 ha, which comprised mainly long gentle slopes to low-lying plains and isolated hills (rubin 1998). by 2011 the park was extended to 143 600 ha.the south africa national parks (sanparks) plant collection programme is a facet of the larger national herbarium (pre) plant collecting programme, where priority is given to collecting in national parks that contain under-collected areas (less than 200 species known within a quarter degree squared [qds] grid according to the south african national biodiversity institute [sanbi]’s plant database pre computerised information system [precis]). this programme officially started in 2005 with collecting efforts in the tankwa karoo and namaqua national parks. the work within these parks not only targets the under-collected areas, but also inadequately known taxa, unique habitats, remote and inaccessible areas and plant species flowering at irregular times, especially after events such as fire or unusually high rainfall. the aim of such surveys in under-collected areas, such as the tknp, is to enhance our knowledge and general understanding of these conservation areas in terms of their floristic component. general collecting in these areas will enable us to compile comprehensive species checklists for the areas that will, in turn, inform decisions regarding the conservation and monitoring of these conservation areas and the plants which they contain. research method and design top ↑ tankwa karoo national park is situated in the southern section of the succulent karoo biome (mucina & rutherford 2006) and stretches from the plains in the south-west to the escarpment of the roggeveld mountains in the east (figure 1). for details on the vegetation of the tankwa karoo and the tknp in particular the reader is referred to rubin (1998 [for major plant communities]), van der merwe, van rooyen and van rooyen (2008a [fynbos related vegetation]) and van der merwe, van rooyen and van rooyen (2008b [succulent karoo vegetation]). figure 1: map of tankwa karoo national park showing the original boundaries of the park at its inception in 1986, the expanded park as in 2005, and its current size (2011). all the qds grids in the park are regarded as under-collected, and therefore the whole park was targeted for general collecting. this entails collecting herbarium specimens of all plants that are flowering or fruiting at the various collecting sites that were chosen. collecting sites were selected to represent the variety of habitat types, for example hill slopes, plains and so forth, found within the specific qds. collecting trips targeted different areas of the park and were carried out at various times of the year (july–october), to reflect the floristic component of the park as best as possible.a total of seven collecting trips were conducted within the park in 2005–2010 with collecting efforts concentrated in the six qdss that comprised the park in 2005. the collecting was carried out by 10 collectors (all staff of the national herbarium in pretoria), including the first three authors. most trips included two or three collectors at a time, with one large trip where six collectors were involved. information accompanying each specimen includes the standard information recorded on herbarium collecting labels, such as locality, habitat, substrate, lithology, exposure, aspect and slope. global positioning system (gps) readings were taken, but these mostly represent a few general readings for a collecting site as a whole, and are not necessarily specific for each individual plant collected. fertile herbarium specimens were collected and identified mainly at the national herbarium in pretoria (pre). cuttings were taken from sterile stapeliads and these were grown at the pretoria national botanical garden’s nursery until flowering. flowers were then added to the herbarium sheets with sterile stems. specimens are housed at pre and duplicates, when available, were distributed to the kimberley south african national parks herbarium (ksan), the compton herbarium in cape town (nbg), and mcgregor museum herbarium in kimberley (kmg). herbarium acronyms follow holmgren, holmgren and barnett (1990). specimen information was incorporated into the main database, the pretoria national herbarium (pre) computerised information system (precis), to allow analysis of distribution data. new specimen records were compared to existing records in precis. where a first record for a taxon in tknp was indicated, the existing checklist for the area (rubin 1998) was consulted to verify the novelty of the record. all relevant specimens acting as vouchers of the new records reported on here, are cited under each taxon in appendix 1. the citation of these voucher specimens are listed in the following format: province.– grid latitude and longitude (grid name): locality description where the specimen was collected (–qds indicator), date, collector & collector’s number (acronym of herbaria where the specimen and its duplicates are housed) ethical considerations top ↑ ethical consideration has been given to the collection of plant specimens by not collecting indiscriminately and taking care not to damage plants. results and discussion top ↑ during recent field trips targeting the tankwa karoo region and especially the tknp, approximately 1500 specimens were collected by staff from the national herbarium (pre). this substantially increased our knowledge of the floristics of the region: at the start of the programme the lowest number of species per qds was 16 (3219ba); this number was raised to 108 (3219ba) by 2011 (table 1). table 1: number of species per quarter degree square grid cell according to the pretoria national herbarium computerised information system data. many significant collections were made as part of this programme: new distribution records were added to the pre collection, for example bulbine triebneri dinter (first record for the northern cape, klopper et al. 2008); first records for the park were collected, for example dioscorea elephantipes (l'hér.) engl. (collected on the gannaga pass); a new species (moraea tanquana goldblatt & j.c.manning) was described from leeuberg (goldblatt & manning 2009) and a new monotypic genus in the apiaceae (scaraboides magee & b.-e. van wyk) was described from elandsberg (magee et al. 2009).further significant range extensions and new park and provincial records from within the park are provided (table 2) with additional information on each species (appendix 1). table 2: family, taxon and status of new plant distribution records from tankwa karoo national park. in the checklist of species for the tknp (rubin 1998) only two stapeliads were listed, namely hoodia gordonii (masson) sweet ex decne. and a stapelia species. stapeliads, in general, are poorly represented in herbaria because it is usually difficult to locate these plants. many are obscured and hidden amongst nurturing plants that provide shade and shelter or they may grow in inaccessible habitats. furthermore, because of their collector’s value, many plants are collected and kept in living collections without making representative herbarium specimens. the correct preservation of herbarium specimens of stapeliads is also perceived to be difficult. cuttings collected in the park were grown until they flowered which made positive identification possible – the list was extended to include six more taxa (table 2).no aloes (asphodelaceae) or ferns (pteridophyta) were represented on the previous checklist (rubin 1998) and general collecting led to three taxa being added from each of these plant groups (table 2). although records of aloe microstigma salm-dyck from the calvinia area has been known since the 1920s, this species has not been collected in the tknp until 2006 when a few individuals of this aloe were found growing on the gannaga pass. apart from moraea tanquana (vulnerable) and nemesia suaveolens k.e.steiner (rare), the only other taxon with a red list status of conservation concern is lachenalia ameliae w.f.barker (near threatened) (l. von staden pers. comm., february 2010). following the new records and range extensions resulting from this study, the extent of occurrence could be re-determined and the statuses of these species were revised. all other taxa reported on here are listed in the latest red list (raimondo et al. 2009) as least concern, and these new records have no influence on their status. figure 2: images of selected species reported on: (a) quaqua acutiloba, (b) stapelia surrecta, (c) tridentea parvipuncta subsp. truncata, (d) tromotriche thudichumii, (e) aloe microstigma growing on the gannaga pass, (f) lachenalia ameliae growing on gravel plain, (g) moraea tanquana and (h) nemesia suaveolens. conclusion top ↑ the general collecting carried out thus far in the tknp has increased our knowledge of the flora of a severely under-collected and unique habitat significantly. knowledge of the full distribution of taxa, especially taxa with limited distribution ranges, is imperative in helping us fully understand these plant species; for instance, without this information the assessment of red data list status of such taxa cannot be carried out accurately. data generated by the sanparks programme in tknp will be used for better informed decisions regarding the conservation of this unique area and also will assist sanparks with management of these plant species. the data are essential in conservation and monitoring actions within the park and will be taken up in the park management plan and conservation development framework. furthermore, it is useful in terms of ecotourism to show visitors which species occur in the park and to highlight the importance of the park in the conservation of these plants. the occurrence of rare and threatened plants in the tknp will also focus the attention of researchers working on these groups on the park and will thus stimulate further research in the area. acknowledgements top ↑ we would like to thank our colleagues for helpful comments and suggestions on this manuscript. cape nature and northern cape nature conservation provided collecting permits, and lize von staden assessed the threat status of species. we also acknowledge with appreciation the two anonymous referees for suggesting improvements to the manuscript. furthermore, we would like to thank sanparks for permission to work in the arid parks with a special mention of conrad strauss, letsie coetzee and the staff of tankwa karoo national park. competing interests the authors declare that they have no financial or personal relationship(s) which may have inappropriately influenced them in writing this paper. authors' contributions s.p.b. (south african national biodiversity institute), r.r.k. (south african national biodiversity institute) and h.m.s. (south african national biodiversity institute) were equally responsible for data collection and analyses used in this study, as well as compiling the manuscript. h.b. (south african national parks) provided the team with logistical support for field work and made valuable contributing comments and improvements to the manuscript. references top ↑ brown, n.e., 1909, ‘stapelia surrecta’, in w.t. thiselton-dyer (eds.), flora capensis, vol. 4(1), pp. 970–971, reeve, london.bruyns, p.v., 2005, stapeliads of southern africa and madagascar, vol. ii, umdaus press, hatfield. crouch, n.r., klopper, r.r., burrows, j.e. & burrows, s.m., 2011, ferns of southern africa: a comprehensive guide, struik, cape town. goldblatt, p. & manning, j.m., 2009, ‘new species of moraea (iridaceae: iridoideae), with range extensions and miscellaneous notes for southern african species’, bothalia 39, 1–10. holmgren, p.k., holmgren, n.h. & barnett, l.c., 1990, index herbariorum, part 1: the herbaria of the world, 8th edn., new york botanical garden, new york. klopper, r.r., klopper, a.w., baijnath, h. & smith, g.f., 2008, ‘bulbine triebneri, an earlier name for bulbine alba, as well as additional and new localities in eastern and northern cape, south africa’, bothalia 38, 67–69. klopper, r.r. & smith, g.f., 2007, ‘the genus aloe (asphodelaceae: alooideae) in namaqualand, south africa’, haseltonia 13, 38–51. http://dx.doi.org/10.2985/1070-0048(2007)13[38:tgaaai]2.0.co;2 klopper, r.r. & smith, g.f., 2010, ‘the genus aloe l. (asphodelaceae: alooideae) in the eastern cape province of south africa’, haseltonia 16, 16–53. http://dx.doi.org/10.2985/1070-0048-16.1.16 leach, l.c., 1985, a revision of stapelia l. (asclepiadaceae), excelsa taxonomic series no. 3, aloe books, johannesburg. magee, a.r., van wyk, b.-e., tilney, p.m. & downie, s.r., 2009, ‘generic delimitations and relationships of the cape genera capnophyllum, dasispermum, and sonderina, the north african genera krubera and stoibrax, and a new monotypic genus of the subfamily apioideae (apiaceae)’, systematic botany 34(3), 580–594. http://dx.doi.org/10.1600/036364409789271218 mucina, l. & rutherford, m.c. (eds.), 2006, the vegetation of south africa, lesotho and swaziland, strelitzia 19, south african national biodiversity institute, pretoria. raimondo, d., von staden, l., foden, w., victor, j.e., helme, n.a., turner, r.c. et al. (eds.), 2009, red list of south african plants, strelitzia 25, south african national biodiversity institute, pretoria. rubin, f., 1998, ‘the physical environment and major plant communities of the tankwa karoo national park’, koedoe 41, 61–94. steiner, k., 2009, ‘two new species of nemesia (schrophulariaceae) from arid areas of the northern cape, south africa’, bothalia 39, 67–72. van der merwe, h., van rooyen, m.w. & van rooyen, n., 2008a, ‘vegetation of the hantam-tanqua-roggeveld subregion, south africa. part 1: fynbos biome related vegetation’, koedoe 50(1), 61–71. van der merwe, h., van rooyen, m.w. & van rooyen, n., 2008b, ‘vegetation of the hantam-tanqua-roggeveld subregion, south africa. part 2: succulent karoo biome related vegetation’, koedoe 50(1), 160–183. appendix 1: detailed information and relevant specimens for new plant records from tankwa karoo national park apocynaceae top ↑ quaqua acutiloba (n.e.br.) bruyns this species was discovered by robert templeman in 1898 and described by n.e. brown in 1909 (bruyns 2005). quaqua acutiloba has a relatively wide distribution range, occurring from south of laingsburg on the edge of the western escarpment northwards to the vicinity of aus in namibia (figure a-1). in the tknp this species is associated with rocky and gravelly plains and foot slopes of hills, but also grows on the lower plateaux (for instance the elandsberg plateau). these plants are usually associated with various species of ruschia which, in most cases, act as nurturing plants to this species (bruyns 2005). the stems can become ± 2 cm in diameter (usually 4-angled) and up to 300 mm long. the plants form clumps of up to 0.3 m in diameter. the flowers are about 1 cm in diameter and are extremely variable in the expression of colour that ranges from plain yellow-green to dark maroon with various degrees of maroon-spotted on yellow-green background forms. the flowers are open, rotate, sometimes with a shallow basal cup (figure 2a).relevant specimens: northern cape.–3219 (wuppertal): tknp. plato above elandsberg (–bb), 07 august 2007, s.p. bester 7792 (pre); on the road between potkleiberg and pramberg. south-western foot slopes and flats at potkleiberg. farm: folmoesfontein (–bd), 05 august 2007, s.p. bester 7731 (pre), s.p. bester 7732 (pre). quaqua parviflora (masson) bruyns subsp. gracilis (c.a.lückh.) bruyns the material from which lückhoff described this taxon was collected between calvinia and pakhuis pass (bruyns 2005). compared to the previous species the flowers in quaqua parviflora subsp. gracilis are much smaller and the stems generally much thinner. the flowers are characteristic with long narrow segments that are constricted more or less in the middle, ending in a thickened apical part. the apical section is usually homogenous in colour in contrast to the rest of the corolla that are maroon-spotted on a cream background. based mainly on this apical swelling and somewhat longer segments of the corolla, it is possible to distinguish it from the typical subspecies. this taxon has a much more restricted distribution than the typical species, occurring from the ceres karoo north-westwards to the west of garies (figure a-1). plants are usually confined to the lower foot slopes of mountainous areas along the escarpment in stony and rocky soils. figure a-1: distribution of quaqua acutiloba, quaqua parviflora subsp. gracilis and tridentea parvipuncta subsp. truncata. relevant specimens: northern cape.–3220 (sutherland): tknp. langkloof gorge (–aa), 07 august 2006, s.p. bester 7139 (pre).in the tknp quaqua and stapelia can be distinguished from each other in the sterile state based on the hairiness of the stems. in quaqua the stems are always glabrous compared to stapelia where it is velvety pubescent (there are, however, some exceptions in stapelia that are also glabrous, but these do not occur in tknp). stapelia arenosa c.a.lückh. joseph archer was the first to collect this species which came from the northern cederberg. it was later described by c.a. lückhoff (bruyns 2005). in the distribution range of this species it is encountered only sporadically. it has been recorded from the kamiesberg southwards to the karoo poort in the ceres karoo (leach 1985) (no specimens in the database). in august 2007 it was collected in the elandsberg area (figure a-2). the plants form few-flowered inflorescences with short pedicels causing the flowers to be held quite close to the stems. with repeated formation of flowers in inflorescences the stem may become irregularly thickened. as the stems develop, flowers may occur all along the stem, from its base to the tips. the stems are diffuse and form lax clumps. figure a-2: distribution of stapelia arenosa, stapelia surrecta and tromotriche thudichumii. relevant specimens: northern cape.–3219 (wuppertal): tknp. southern slopes of elandsberg, (–bb), 06 august 2007, s.p. bester 7749a (ksan, pre); plato above elandsberg, (–bb), 07 august 2007, s.p. bester 7795b (kmg, ksan, nbg, pre). stapelia surrecta n.e.br. this species was first collected by rudolph marloth and later described by n.e. brown in flora capensis (1909). stapelia surrecta (figure 2b) is one of few species in the genus that carries flowers with long pedicels concentrated at the tips of the stems (another example is some forms of stapelia kwebensis). this species has a restricted range and is only known from the ceres karoo. it occurs from karoo poort in the south to bloukrans pass in the north (figure a-2). in the tknp plants were collected in rocky and sandy soils in the langkloof, but according to bruyns (2005) it is restricted to shale and tillite. this species can only be distinguished from stapelia arenosa when flowering because the vegetative stems are indistinguishable.relevant specimens: northern cape.–3220 (sutherland): tknp. langkloof gorge (–aa), 07 august 2006, s.p. bester 7161a (ksan, pre); 7163a (pre). tridentea parvipuncta (n.e.br.) l.c.leach subsp. truncata (c.a.lückh.) bruyns the subspecies was originally discovered by james lückhoff in 1931 and described by c.a. lückhoff in 1937 (bruyns 2005). it is distinguished from the typical subspecies in that the outer corona lobes are truncate and emarginate and not deeply bifid with two lobes diverging as in the typical subspecies (figure 2c). tridentea parvipuncta subsp. truncata are found from the western section of karoo poort to botterkloof (figure a-1). it is separated from the typical subspecies which is found south of merweville westwards.relevant specimens: northern cape.–3219 (wuppertal): tknp. on the road between potkleiberg and pramberg. south-western foot slopes and flats at potkleiberg. farm: folmoesfontein (–bd), 07 november 2007, s.p. bester 7730a (kmg, ksan, pre). tromotriche thudichumii (pillans) l.c.leach although this taxon was already discovered in 1937 and re-collected in 1940 by marthinus malherbe, it was only described some 20 years later by n.s. pillans in 1959 (bruyns 2005). despite a manuscript name that was given to it in the bolus herbarium (stapelia malherbei), it was eventually named after jacques thudichum (curator of the karoo botanical garden at the time), who found a number of these plants in cultivation at the garden which he could not identify. the plants grow in rocky and gravelly soils on the flats of the tanqua karoo and roggeveld escarpment (bruyns 2005), mainly in the ceres karoo (north of karoo poort) and calvinia districts (figure a-2). characteristic of this species is the corolla that is so strongly reflexed that the flowers usually seem to be clasping the stem (figure 2d).relevant specimens: northern cape.–3219 (wuppertal): tknp. southern slopes of elandsberg (–bb), 06 august 2007, s.p. bester 7749 (kmg, ksan, pre); plato above elandsberg, (–bb), 07 august 2007, s.p. bester 7795a (kmg, ksan, pre); on the road between potkleiberg and pramberg. the south-western foot slopes and flats at potkleiberg. farm: folmoesfontein (–bd), 05 august 2007, s.p. bester 7730 (pre). asphodelaceae: alooideae top ↑ aloe microstigma salm-dyck aloe microstigma occurs fairly widespread in the western parts of the eastern cape, the central areas of the western cape and just into the northern cape, with a disjunction to south-western namibia (klopper & smith 2010). although records from the calvinia area has been known since the 1920s, this species has not been collected in the tknp until 2006 when a few individuals of this aloe were found growing on steep slopes in tankwa escarpment shrubland (mucina & rutherford 2006) on the gannaga pass. this population represents the yellow flowering form of a. microstigma, previously known as aloe brunthallerii a.berger ex cammerl. and is one of the northernmost records of this colour form (figure 2e and figure a-3). figure a-3: distribution of aloe microstigma and lachenalia ameliae. relevant specimens: northern cape.–3220 (sutherland): tknp, top of gannaga pass, farm kleinfontein 1027, near the viewpoint, (–aa), 05 august 2006, r.r. klopper 321 (ksan, pre); 10 july 2010, r.r. klopper & a.w. klopper 336 & 337 (pre). aloe falcata baker aloe falcata is known to occur from the richtersveld south to the klawer area and inland to around calvinia (klopper & smith 2007). it was first observed on a rocky ridge in the tanqua karoo vegetation type (mucina & rutherford 2006) during a general collecting trip in tknp in august 2006, but was not in flower and no material was collected. the significance of this population was only realised afterwards: it is the most south-eastern record known for this aloe to date. subsequently, during a specialised collecting trip focussing on aloes in 2010, a herbarium record from this population was collected (figure a-4). figure a-4: distribution of aloe falcata. relevant specimens: northern cape.–3220 (sutherland): tknp, farm roodewerf 1091, houtkraalkloof, (–aa), 10 july 2010, r.r. klopper & a.w. klopper 339 (pre). aloe variegata l. this very widespread and small aloe, with its cryptically mottled and camouflaged leaves (klopper & smith 2007, 2010), is very inconspicuous when not in flower. it is thus not surprising that it has not been collected in the tknp before 2006, even though it occurs at several localities within the park, all within the tanqua karoo vegetation type of the rainshadow valley karoo bioregion (mucina & rutherford 2006) (figure a-5). figure a-5: distribution of aloe variegata and cheilanthes namaquensis. relevant specimens: northern cape.–3219 (wuppertal): tknp, hill east of leeuberg, (–bb), 04 august 2006, h.m. steyn 888 (ksan, pre); grasberg south 1103, (–bc), 16 august 2008, h.m. steyn 1457 (pre); foot of slope west of potklysberg, (–bd), 05 august 2007, a.c. mudau 254 (pre). 3220 (sutherland): tknp, border of farms manus zyn dam and roodewerf 1019, remhoogte, (–aa), 08 august 2006, r.r. klopper 361 (ksan, pre); maansedam, on the plain surrounding the house, south of the house, (–aa), 06 august 2006, h.m. steyn 941 (ksan, pre). hyacinthaceae top ↑ lachenalia ameliae w.f.barker lachenalia ameliae was thought to be endemic to the western cape, occurring mainly in the montagu and touws river areas. the northernmost specimen was collected in 1921 at gansfontein in the ceres karoo (marloth 10479, pre). during august 2006, 2008 and 2011 four populations were found in three adjacent grids in the north-western parts of tknp. these populations comprise several plants growing on shale-derived soils on gravel plains in the tanqua karoo vegetation type (mucina & rutherford 2006), at altitudes between 352 m and 407 m. this represents a northward extension of the known distribution range by ± 50 km and a new record for the northern cape (figure 2f and figure a-3). because of the new records, the extent of occurrence changed from 4000 km² to 17 696 km², but this is still within the near threatened (nt) range and the status remains unchanged (l. von staden pers. comm., february 2012).relevant specimens: northern cape.–3219 (wuppertal): tknp. sandy wash west of leeuberg, (–bb), 03 august 2006, h.m. steyn 863b (pre); tknp, between varsfontein and prambergfontein, (–bb), 17 august 2008, h.m. steyn 1459 (ksan, pre); tknp, grasberg south 1103, (–bc), 16 august 2008, h.m. steyn 1452 (ksan, pre); tknp, track between luiperdskop and varsfontein, (–bd), 15 august 2011, h.m. steyn 1864 (pre). iridaceae top ↑ moraea tanquana goldblatt & j.c.manning as a result of general collecting, a previously unknown, blue flowering moraea species was collected in tknp in august 2006 (figure 2g). it was subsequently described as m. tanquana (goldblatt & manning 2009). when described, this species was only known from the type locality, but during follow-up field trips to different parts of the park, three more populations were found. in august 2008, a population was found on the lower north-eastern slopes of leeuberg (no specimens were collected), as well as on a plateau to the southeast of grasberg. in august 2009, yet another population on the eastern side of the same plateau was discovered. all three populations were found in tanqua karoo vegetation (mucina & rutherford 2006) on dolerite-derived stony or rocky soil. the populations were estimated to consist of between 80 and 120 plants each and the maximum range extension is between 13 km and 15 km (figure a-6). a preliminary threatened status of vu d1 (vulnerable) was assigned to this species based on the localised distribution and relatively small population sizes (l. von staden pers. comm., february 2010). figure a-6: distribution of moraea tanquana and nemesia suaveolens. relevant specimens: type.–northern cape, 3219 (wuppertal): tknp, small koppie east of leeuberg, (–bb), 04 august 2006, h.m. steyn 872 (nbg, holo.; ksan, pre, iso.).northern cape.–3219 (wuppertal): grasberg south 1103, (–bc), 16 august 2008, h.m. steyn 1431 (pre); hill west of track between varsfontein and luiperdskop, (–bd), 06 august 2009, h.m. steyn 1524 (pre). scrophulariaceae top ↑ nemesia suaveolens k.e.steiner a number of nemesia specimens with yellow and purple or pink flowers were collected (figure 2h). this species was usually found on gravelly sand in the tanqua karoo vegetation type (mucina & rutherford 2006). at the time, these specimens could not be matched to any nemesia species at pre and they were only recently identified as n. suaveolens k.e.steiner. this is an annual species known from a very limited area of the central tanqua karoo. the flowers have a straight spur and a pleasant spicy fragrance (steiner 2009). with this new information the national threatened status of this species was determined as rare, because it has a restricted range (eoo 478 km2) and is potentially vulnerable to grazing and population fluctuations in response to rainfall. subpopulations, however, are large and a large proportion of the population is protected in the tknp (l. von staden pers. comm., february 2010).the known distribution of this species is shown in figure a-6. the original localities are included in this map, not only to highlight the new records, but to rectify the original map (steiner 2009:70). in the original map all the collections were plotted in 3219 (–bc), although specimens were listed from (–ba), (–bc), (–bd), (–da) and 3220 (–ac) (steiner 2009). relevant specimens: northern cape.–3219 (wuppertal): tknp, western end of park. dry riverbed, (–ba), 02 august 2007, s.p. bester 7687 (pre); tknp, between varsfontein and the gate at the ceres-calvinia road, (–ba), h.m. steyn 1434 (pre); calvinia district, tknp, gravel plains west of leeuberg, (–bb), 03 august 2006, b. sachse 31b (pre); calvinia district, tknp, biesjesfontein (–bb), 15 august 2011, s.p. bester 10713 (pre); tknp, grasberg south 1103, (–bc), h.m. steyn 1442 (pre); calvinia district, tknp, luiperdskop farm se of the koppie (–bd), 15 august 2011, s.p. bester 10712 (pre). western cape.–3220 (sutherland): along the road between paulshoek and oudebaaskraal (–ac), 05 august 2009, m. koekemoer 3752 (pre); ceres district, tknp, between roodewerf and oudebaaskraal, platfontein farm (–ac), 13 august 2011, s.p. bester 10706 (pre). pteridophyta: sinopteridaceae top ↑ cheilanthes parviloba (sw.) sw. this fern is widespread from the western cape, through the eastern cape to the south of kwazulu-natal, the free state, lesotho, north west and limpopo, with a disjunction to central namibia, and also occurs in southern zimbabwe (crouch et al. 2011). it was first collected in the tknp in 2006 in the tankwa escarpment shrubland (mucina & rutherford 2006) near the top of gannaga pass. it had not been collected in the northern cape previously; thus this collection represents a range extension of ± 150 km from the nearest known record in the western cape (figure a-7). figure a-7: distribution of cheilanthes parviloba. relevant specimens: northern cape.–3220 (sutherland): tknp, farm kleinfontein 1027. plateau at top of gannaga pass, at the top edge of the pass, (–aa), 05 august 2006, r.r. klopper 319 (ksan, pre). cheilanthes induta kunze cheilanthes induta is a fairly distinctive species, characterised by long, reddish hairs on the stipe, rachis and abaxial lamina surface, as well as its unusual convoluted stipe (crouch et al. 2011). it occurs widespread in the western cape and adjacent parts of the northern cape, as far north as springbok, and the eastern cape, extending into the isolated mountains of the great karoo (crouch et al. 2011). it was first collected in the tknp in 2007 in the roggeveld shale renosterveld (mucina & rutherford 2006) (figure a-8). figure a-8: distribution of cheilanthes induta. relevant specimens: northern cape.–3220 (sutherland): tknp, klipfontein farm, (–aa), 14 september 2007, m.s. mothogoane 736 (pre). cheilanthes namaquensis (baker) schelpe & n.c.anthony cheilanthes namaquensis is a rare fern that occurs from southern namibia, through namaqualand in the northern cape, into the western cape (crouch et al. 2011). it is easily confused with cheilanthes deltoidea kunze subsp. deltoidea that has a very similar, although slightly wider, distribution range. cheilanthes namaquensis can be distinguished by its narrower fronds with basal pinnae that are not basiscopically developed and its rachis that is only winged in the apical portion of the frond and not winged throughout, as in the broadly triangular fronds of c. deltoidea with its basiscopically developed basal pinnae (crouch et al. 2011). this fern was first collected in tknp in 2006 in tankwa escarpment shrubland (mucina & rutherford 2006) (figure a-5).relevant specimens: northern cape.–3220 (sutherland): tknp, farm langekloof 60. langkloof, ± 1.5 km – 2 km north of malansgat river, (–aa), 07 august 2006, r.r. klopper 353 (ksan, pre). article information authors: leslie r. brown1 pieter j. du preez2 hugo bezuidenhout1,3 george j. bredenkamp4 theo h.c. mostert5 nacelle b. collins6 affiliations: 1applied behavioural ecology and ecosystem research unit, university of south africa, south africa2department of plant sciences, university of the free state, south africa 3south african national parks scientific services, hadison park, kimberley, south africa 4department of plant science, university of pretoria, south africa 5department of botany, university of zululand, south africa 6free state department of economic development, tourism & environmental affairs, bloemfontein, south africa correspondence to: leslie brown postal address: private bag x6, florida 1710, south africa dates: received: 25 july 2012 accepted: 29 apr. 2013 published: 23 july 2013 how to cite this article: brown, l.r., du preez, p.j., bezuidenhout, h., bredenkamp, g.j., mostert, t.h.c. & collins, n.b., 2013, ‘guidelines for phytosociological classifications and descriptions of vegetation in southern africa’, koedoe 55(1), art. #1103, 10 pages. http://dx.doi.org/10.4102/ koedoe.v55i1.1103 note: additional supporting information may be found in the online version of this article as an online appendix 1 http://dx.doi.org/10.4102/ koedoe.v55i1.1103-1 and online appendix 2 http://dx.doi.org/10.4102/ koedoe.v55i1.1103-2. copyright notice: © 2013. the authors. licensee: aosis openjournals. this is an open access article distributed under the terms of the creative commons attribution license, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. guidelines for phytosociological classifications and descriptions of vegetation in southern africa in this checklist... open access • abstract • introduction • purpose of this article • history of phytosociology in south africa • recommended minimum requirements for phytosociological studies in south africa    • approach    • principles    • field survey methods    • data analysis       • vegetation classification       • environmental gradient analysis or ordinations       • other statistical analyses, calculations and comments       • tables       • vegetation maps    • classification and description of plant communities       • classification of plant communities       • naming of plant communities       • description of plant communities       • syntaxonomic descriptions       • recommendations for concluding remarks • concluding remarks and recommendations • acknowledgements    • competing interests    • authors' contributions • references abstract top ↑ changes in the environment are first observed in changes in the vegetation. vegetation survey, classification and mapping form the basis on which informed and scientifically defendable decisions on the environment can be taken. the classification and mapping of vegetation is one of the most widely used tools for interpreting complex ecosystems. by identifying different plant communities we are essentially identifying different ecosystems at a particular hierarchical level. phytosociologists in europe have been involved in such studies following, in particular, the braun-blanquet approach since the early 1900s. in south africa, such studies were undertaken on a limited basis from the early 1970s and have since then steadily increased. the surveying of the enormous diversity of south african vegetation is one of the objectives of phytosociological studies. the demand for such data has steadily increased over the past few years to guide conservation policies, biodiversity studies and ecosystem management. in south africa, numerous publications on the vegetation of conservation and other areas in the different biomes have been produced over the last few decades. however, vegetation scientists in south africa experience unique problems. the purpose of this article is therefore to provide an overview of the history and the specific focus of phytosociological studies in south africa and to recommend minimum requirements and methods to be followed when conducting such studies. it is believed that the incorporation of these requirements will result in scientifically justifiable research of high quality by phytosociologists in south africa.conservation implications: effective conservation cannot be obtained without a thorough knowledge of the ecosystems present in an area. consistent vegetation classifications and descriptions form the basis of conservation and monitoring exercises to maintain biodiversity. the incorporation of these guidelines and requirements will facilitate quality phytosociological research in south africa. introduction top ↑ one of the earliest examples of an informal description of southern african vegetation dates back to the late 1400s. in december 1497, vasco da gama sailed from mossel bay in an easterly direction, ensuring that he always had sight of the terrestrial land. interestingly he made no mention of the different wild animals, but instead he continually referred to the vegetation they observed on the land (skead 2011). since then people have used vegetation to assist with finding their way in and around the african continent. vegetation is the most physical representation of the environment (kent 2012). any spatial and temporal changes in habitats are first observed in the vegetation. as a result, the mapping and description of vegetation has either informally or formally played an important role as a tool to classify and interpret different ecosystems. these assessments led to the development of the discipline of vegetation science (plant ecology), and its various fields of specialisation, of which synecology (the classification, description and mapping of vegetation, succession and dynamics) (barbour, burk & pitts 1987) is an important field. synecology stems from observations of the great variety of form and structure of plants that repeat themselves in similar environmental conditions. vegetation scientists started to explore the world to describe and map the diverse plant life of the planet. in the early 20th century most vegetation scientists in europe were occupied with the phytosociological (phyto = plant; sociology = groupings of species) classification, description and mapping of the continents’ vegetation. professor v. westhoff once commented that phytosociology is the science of recognising and identifying the stepping stones in vegetation within the overwhelming swamp of variation (bredenkamp 1982; mostert 2006). currently, vegetation classification and mapping is one of the most widely used tools to assist in the interpretation of complex ecosystems and to simplify the spatial and temporal complexity of these ecosystems (doing 1970; mucina & rutherford 2006). vegetation also fulfils an important function in that it provides food and shelter for wildlife (mucina & rutherford 2006; skead 2011). modern scientific society uses phytosociology either to form the background of scientific studies on animals (e.g. henzi et al. 2011; hirst 1975; pasternak et al. 2013) or to define different plant communities, which could be unique in the sense of rare or endangered plant species (e.g. du preez & brown 2011; edge, cilliers & terblanche 2008; gotze et al. 2008; janecke, du preez & venter 2003). results of phytosociological studies should be the foundation for informed decisions on wildlife management and nature conservation. phytosociology not only serves as the backbone of ecosystem studies but also forms part of the planning for monitoring plant species or communities and rare or endangered ecosystems. vegetation is a collective term for all the plant communities in the same way that flora is a collective term for all the plant species. we describe plant communities, not ‘vegetation communities’. with climate change, phytosociological studies will become more important because, in most cases, only vegetation data are available to use for comparisons. data collected according to accepted quantitative methods, can be used to give detailed information about plant species abundance and also a description of the structure of the vegetation. such data can also be used for gradient analyses, measuring plant species diversity, the study of successional changes and measuring plant production of different ecosystems. vegetation classification is also useful to assist in making informed decisions on the habitat that is available for wildlife, as well as making informed decisions on fire policy and programmes aimed at clearing alien plant species. detailed vegetation classification, mapping and description also form the basis from which informed and scientifically defendable decisions can be taken for infrastructure and other development in an area. it also assists with impact assessment for development purposes. when the expansion of conservation areas is considered, the results of phytosociological studies should be used to assist with planning of these expansions to guide conservation decisions about important, scarce or rare plant communities. at an appropriate scale (association level), plant communities can thus be used as surrogates for ecosystem delineations with very high accuracy. this ability of vegetation to act as a surrogate for ecosystem descriptions or delineations is seated in the fact that vegetation is a highly visible and measurable biological manifestation of all the other environmental factors shaping and driving a specific ecosystem (barbour et al. 1987; kent 2012). phytosociology is used in many different research fields in nature conservation, (e.g. mammalogy, ornithology, herpetology, entomology, geology, soil science, landscape ecology, limnology, etc.) to describe the habitat and give the reader an idea of the (expected) species composition and vegetation structure. it is therefore important that vegetation description studies should be as comprehensive as possible. with the current pressure on the environment, and especially natural vegetation, as a result of mining and agricultural activities. as well as urban and rural development, the need for proper planning cannot be underestimated. although the need for mining or agricultural development sometimes supersedes conservation importance, vegetation classification and description is important to identify ecologically sensitive areas. these studies are currently compulsory in south africa in terms of the national environmental management: biodiversity act (act no. 10 of 2004) (south african government 2004). by identifying different plant communities, we are essentially identifying different ecosystems at a particular hierarchical level. plant communities and their associated vegetation maps are therefore regarded as reliable surrogates for the demarcation of macro-ecosystems. describing, monitoring and managing the surrogate (plant community) are the first steps in effectively monitoring and managing the entire ecosystem, without trying to understand and manage the bewildering detail of all the different components and interactions of the ecosystem. vegetation surveys, classification and mapping provide a framework for understanding the differences between ecosystems (chytrý, schaminee & schwabe 2011). although this approach to ecosystem management is not fail safe or without its shortcomings, it provides ecologists with a sensible, tangible tool and first approximation for the management of ecosystem patterns and processes. purpose of this article top ↑ phytosociological research has a long history. however, vegetation scientists in south africa are experiencing various problems that do not occur elsewhere in the world (see later). the purpose of this article is to provide an overview of the history of phytosociological studies in south africa and to recommend broad guidelines on the minimum requirements and methods to be followed when conducting vegetation classification, description, habitat interpretation studies and mapping following the braun-blanquet approach. history of phytosociology in south africa top ↑ we attempt to present a brief history of the development of phytosociology in south africa using a few examples for explanation, rather than giving a complete review of all the phytosociological research conducted or publications that have appeared on south african vegetation during the last century. most of the earlier national and regional studies were mainly of a non-formal descriptive nature (e.g. bayer 1955; bews 1918; dyer 1937; edwards 1967; killick 1963; louw 1951; muir 1929; pole evans 1922), often providing only species lists of a particular area. acocks (1953) classified the south african vegetation into 70 veld types and 75 variations based on comparison of floristics from stand data. he recorded the abundances of all species. consequently, many south african vegetation scientists used a more flexible approach. statistical numerical classification methods were applied as a first approximation of the plant communities in a particular area, but with the option to ‘refine’ the classification by applying zurich–montpellier methods. relevés could be moved to other clusters (if deemed necessary) by considering more factors (especially total floristic composition, diagnostic species and habitat interpretation) than only those used by the particular numerical algorithm (bredenkamp 1982). application of the twinspan classification algorithm (hill 1979) made a major contribution to obtain more objectivity and repeatability in the classification whilst simultaneously retaining the advantages of a phytosociological table. this provided not only the hierarchical classification and total floristic composition of each plant community at different levels, but also a valuable overview of species cover and abundance, constancy, fidelity and the habitat. the phytosociological table thus became a necessity in every phytosociological study. as computer software became more readily available, the turboveg programme was developed to serve as a standardised format for the storage and management of vegetation data (hennekens 1996; hennekens & schaminee 2001). apart from the pioneering studies of werger (1973) and van der meulen (1979), which were considered to be regional studies, the majority of the earlier studies by south african researchers were more local in extent. the goals of sustainable utilisation linked with effective conservation cannot be achieved without a thorough knowledge of the ecology of a particular area (edwards 1972; werger 1974). it was therefore recommended that conservation policies and environmental management plans should be based on this knowledge (edwards 1972). this resulted in many phytosociological projects initiated in nature conservation areas (e.g. national parks, provincial nature reserves or privately owned and managed game reserves), which resulted in numerous publications on the vegetation of conservation areas in the different biomes of south africa. selected examples include: • grassland: bloem (1988), bredenkamp (1975), coetzee (1972), kay, bredenkamp and theron (1993) and swanepoel (2006). • savanna: bezuidenhout (1994), brown (1997), gertenbach (1978), mostert (2006), stalmans and peel (2010), van rooyen (1983) and van staden (2002). • nama-karoo: brown and bezuidenhout (2000), cleaver, brown and bredenkamp (2005), rubin and palmer (1996), van der walt (1980) and werger and coetzee (1977). • succulent karoo: jurgens (2004) and le roux (1984). • fynbos: boucher (1997), mcdonald (1988), zietsman (2003) and zietsman and bredenkamp (2006). • forest: geldenhuys and murray (1993), grobler (2009) and matthews et al. (2001). • thicket: palmer (1981). • azonal units: collins (2011), du preez and brown (2011) and pretorius (2012). • additional regional studies: bezuidenhout (1993), du preez (1991), eckhardt (1993), fuls (1993), hoare (1997) and kooij (1990). recommended minimum requirements for phytosociological studies in south africa top ↑ to achieve the goal of syntaxonomic synthesis of southern african vegetation and to attain internationally acceptable standards for local studies in, for example, national parks, nature reserves, private game farms and other conservation areas, we strongly recommend the following minimum requirements for phytosociological studies in southern africa. approach we recommend that the zurich–montpellier (braun-blanquet) school of total floristic compositions (braun-blanquet 1932; kent 2012; mueller-dombois & ellenberg 1974; werger 1973; westhoff & van der maarel 1978) should be followed. the main benefit of this approach is that much of the world’s vegetation has been and is continued to be surveyed and classified according to a relatively uniform protocol (chytrý et al. 2011). principles • sample plot placement is not subjective as originally proposed by braun-blanquet (1932), but placed in a stratified random manner within floristically uniform units because it is accepted that each of these units represents a single plant community.• the vegetation in a sample plot must be representative of a single plant community. therefore, a minimum plot size for the particular plant community is used. • the sample plot must be uniform with regard to its biophysical make-up. independent of the scale of the study, floristically homogeneous units that can physically be identified and managed in the field must be used to define plant communities. large scale-related heterogeneous sample plots are avoided as they will result in mixed relevés containing species from different plant communities and a range of biophysical features. these mixed relevés are not according to the basic zurich–montpellier principle of sampling within homogeneous units. mixed relevés are difficult to interpret ecologically and to assign to a specific recognisable plant community. the classification of mixed relevés groups unrelated communities together and may ignore smaller and sometimes threatened or endangered ecosystems. • ideally, all species present in the sample plot should be identified and recorded during the time of survey. owing to seasonality and natural impacts (e.g. grazing, fire, irregular rainfall, erosion, flooding, droughts, etc.), especially in arid and semi-arid areas, some plant species are not easily identifiable (werger 1973). in addition, the rich floristic diversity of south africa and the many cryptic species result in taxonomical problems owing to several undescribed and often unidentifiable species (e.g. some mesembs, orchids, geophytes and annuals). field survey methods the following field survey methods are recommended:• stratified random placement should be determined during the desktop phase whilst preparing for fieldwork. stratification of the vegetation can be performed by means of a geographic information system (gis) using a number of environmental data overlays in combination with satellite and aerial imagery. applicable spatial environmental data overlays include landscapes, land types, terrain units, topography, altitude, climate, geology, soil characteristics, surface rock cover, land use, land cover, or any other relevant spatial data set. overlaying these data sets onto available aerial and satellite imagery will aid in the accurate delineation of relatively homogeneous vegetation units. placement of sample plots within each homogeneous vegetation-cum-habitat unit should be random. in the field, however, the randomly determined location of each sampling plot should be critically evaluated according to the first rule of the zurich–montpellier sampling method (placement of the sampling plot should be within a homogeneous vegetation patch representative of the perceived plant community). if the sampling plot does not fall within a homogeneous representative vegetation stand, it should be moved to the nearest locality that does fulfil this criterion. • a minimum plot size should be determined based on species–area curves produced for each physiognomic–physiographic unit sampled (kent 2012). it is important to note that when gathering data to determine such species–area curves, the first rule of the braun-blanquet method (placement of the nested set of progressively larger plots within a homogeneous vegetation stand, representative of the perceived plant community) should be observed very strictly. alternatively, the researcher may use plot sizes similar to those used in other successful braun-blanquet surveys in similar physiognomic–physiographic units. the researcher should refer to available literature and clearly state the aspects of similarity between the vegetation described with the various plant communities occurring in the study area. the plot sizes given in table 1 may be used as a guideline for the relevant vegetation to be sampled. • for formal syntaxonomic classification, weber, moravec and theurillat (2000) states that only one relevé is required for a formal description of a syntaxon (plant community) although a minimum of ten plots are recommended. to comply with statistical requirements in local phytosociological studies, a minimum of three sampling plots for each of the stratified physiognomic–physiographic units is recommended. exceptions to this suggestion may be in the case of azonal vegetation covering so little of the study area that the placement of more than two plots may lead to pseudo-replication. the required minimum number of plots will depend largely on the size of the plant community in question, being less in smaller plant communities. the description of plant communities based on one or two relevés should be strongly discouraged and such communities should be regarded as doubtful units (weber et al. 2000). • apart from compiling all existing environmental data during the preparation for fieldwork (desktop gis), as many as possible of the following environmental parameters should be measured and determined in the field at each sampling plot during the actual fieldwork phase (recommended but not limited to): geology, surface rock cover, land type, terrain form, soil depth, soil form, soil texture, soil structure, altitude, coordinates, aspect and slope. • soil samples should preferably be collected at the average rooting depth utilised by the vegetation that is being studied. where the soil is classified, the diagnostic horizon from which the sample was taken should be noted. • south africa’s vegetation is highly dependent on rainfall, which is, in most cases, seasonal and erratic. it is recommended that a survey be conducted during the optimal growth period. the period of field survey should be stated clearly. • cover–abundance values for each species recorded within a sample plot should be estimated using one of the many compatible cover–abundance scales used in current phytosociological studies (e.g. modified braun-blanquet cover scale, domin cover scale, plant-number scale) (kent 2012; westfall et al. 1996). in current southern african surveys, the modified braun-blanquet cover–abundance scale is preferred for vegetation classification studies. table 1: suggested minimal area values (m2) for various plant communities. data analysis vegetation classification • it is recommended that phytosociological data are captured in a format that can be imported into programs such as juice (tichý & holt 2006). • algorithms assume that the data have a normal distribution. data should be tested for normality (e.g. using pc-ord [mccune & mefford 2006]) and transformed if the assumption is not satisfied. • either divisive or agglomerative clustering can be used for classification. when using divisive clustering, the modified twinspan (two-way-indicator species analysis [hill 1979]) algorithm proposed by roleček et al. (2009) as contained in juice (tichý 2002) is recommended. unlike the original version, the modified twinspan algorithm does not enforce a dichotomy of classification but instead, at each step, divides only the most heterogeneous cluster of the previous hierarchical level. thus, the application of the modified twinspan algorithm results in vegetation units of similar internal heterogeneity. pseudospecies cut levels used in the classification must be indicated (hotanen 1990). pseudospecies cut levels should consider the scale used during data collection (e.g. where the modified braun-blanquet scale was used, the following pseudospecies cut levels are recommended: 0, 5, 15, 25, 50, 75). • for fidelity measures the use of the phi coefficient of association (chytrý et al. 2002) of each species should be calculated to determine their fidelity to each plant community as an indication of their suitability as diagnostic species. equalisation of relevé cluster sizes (tichý & chytrý 2006) is recommended before calculating the phi coefficient. one can also weigh the relative importance of common and rare species by changing the equalised size of the site groups. this can easily be achieved in the juice program (tichý 2002). it is further recommended that fisher’s exact test be performed simultaneously with the calculation of the phi coefficient to exclude species with non-significant fidelity from the groups of diagnostic species. • description of clusters must include diagnostic, constant and dominant species. it is recommended that these are derived using juice, with 75, 60 and 50 recommended for lower threshold values and 80, 80 and 60 for upper threshold values for diagnostic, constant and dominant species, respectively. in addition, the jaccard and/or sorensen values should be reported. • manual rearrangement of species after automated clustering is permissible. however, rearrangement of clusters and relevés is strongly discouraged. if relevés or clusters were rearranged, it must be accompanied by a detailed justification. the dendrogram should still reflect the original clustering. the modified twinspan classification technique has been described and recommended here owing to its wide use in classification research. it is, however, important to note that other classification tools also exist. optimclass is one of the latest methods used to determine optimal partition between the different data sets of a study area and can also be computed in the juice program (tichý et al. 2010). this method is based on species with a high fidelity, which can subsequently be used as diagnostic species. environmental gradient analysis or ordinations • should the researcher wish to illustrate floristic gradients within and between plant communities or link these gradients with habitat variables, a suitable gradient analysis algorithm should be used, for example canoco, pc-ord and primer.• detrending should be used only where ordination without detrending produces uninterpretable results (e.g. where the objects are clustered together). • it is recommended that papers report on the outcome of procedures followed to determine whether the most important environmental variables were measured (for correspondence analysis and detrended correspondence analysis). it is also recommended that if the number of environmental variables was reduced, the amount of explanatory power lost should be reported. • the decision for the underlying model used during ordination (linear or unimodal) should be noted or explained. other statistical analyses, calculations and comments we encourage further analyses of the data using sound statistical methods to investigate some of the following aspects, where relevant:• diversity • richness • evenness • medicinal and economic value • endemism • conservation status and value • production, grazing or browsing value. tables diagnostic species (see later) are used to arrange plant communities (associations) into a hierarchical classification and are presented in a phytosociological or a synoptic table. the table is the hallmark of the phytosociological study and the interpretation of the environment and its different plant communities. the table is the manifestation of a reliable and accurate research method. the arrangement of the different species groups within the table is therefore of utmost importance (westhoff & van der maarel 1980; werger 1974):• phytosociological table: the phytososiological table should be arranged such that the diagnostic species group for the plant community (association) is above the diagnostic groups of the sub-associations and variants. the species groups are labelled alphabetically with an indication of diagnostic groups for easy reference. species commonly shared amongst communities, probably representing alliances, orders and classes, are normally placed towards the bottom of the table. an example of a typical phytosociological table is included in online appendix 1. • synoptic table: once plant communities have been identified in the phytosociological table, a synoptic table can be produced to summarise the data for each plant community (kent 2012). in a synoptic table, each plant community is summarised in a single column containing values such as fidelity and constancy for each species. an example of a typical synoptic table is included as online appendix 2. one or both of these tables must be used, as long as they contribute to a better understanding of the plant communities identified. it should, however, be noted that phi-coefficient values, used for the synoptic table, are determined from absence/presence data and not cover–abundance data. it therefore means that pseudospecies are not used in the determination of diagnostic species. thus, one cannot solely use these results in the description of the communities, as dominant species with low phi-coefficient values will be omitted from the table. it is therefore important that the researchers use the phytosociological table to describe the dominant and co-dominant species of the community. vegetation maps effective knowledge of the different plant communities and their potential to provide habitats for animals are essential to make scientifically based management decisions for natural areas. the spatial representation of the different plant communities is therefore important. in the past, technologies to provide accurate and detailed information on the location and distribution of plant communities did not exist. owing to technological advances floristically based classifications provide excellent information for the construction of accurate high-resolution vegetation maps (clegg & o’connor 2012; dias, elias & nunes 2004). vegetation maps of natural areas have become indispensable for managers of natural areas. these maps not only indicate the location, distribution and abundance of the different plant communities, together with rare and endemic species, but are also used to monitor changes in cover, structure and composition of the vegetation. it is therefore strongly recommended that vegetation classification and description manuscripts of conservation areas are all accompanied by vegetation maps. classification and description of plant communities plant species names used should follow the latest comprehensive south african plant species list (i.e. currently germishuizen & meyer [2003], with online updates from plants of south africa [posa], which can be accessed via the sanbi website, or the african plants database [version 3.3.4] at http://www.ville-ge.ch/musinfo/bd/cjb/africa). if the authors are of the opinion that the names of certain plant taxa are outdated or inappropriate, the use of newer names for those should be indicated clearly and referenced from the relevant published taxonomic literature.when describing plant communities in normal local habitat classifications (not formal syntaxonomy) the following is recommended. classification of plant communities the classification of different plant communities are based on total floristic composition, although they are recognised by their diagnostic species (character species and differential species). character species are species that are mostly restricted to a specific plant community. they thus characterise the community by their occurrence in one community and by being absent or less frequent in other communities. sub-communities are, in many cases, characterised by the presence or absence of certain species and these are referred to as differential species. these species are used mostly to define the lower syntaxa (westhoff & van der maarel 1980). all diagnostic species identified using ‘objective’ statistics (phi coefficient) should be evaluated with regard to their ‘robustness’ as reliable and predictable indicators of a given plant community. the fewer relevés used to describe a plant community, the more subjective the interpretation of diagnostic species will be. naming of plant communities according to weber et al. (2000) names are only labels to assist in the classification of plant communities and, as such, they will never be wholly adequate. it is, however, more important to understand what is meant by a name than to find one that is characteristic in every respect (weber et al. 2000). it is therefore important that basic rules are followed when naming plant communities to avoid confusion and to enable consistency. the following is therefore proposed:• plant community names are assigned following the same guidelines as presented in the international code of phytosociological nomenclature (weber et al. 2000) for formal syntaxonomical classification, but do not use the specified taxon epithets. according to this rule, the dominant plant name or the one that dominates the structure is second (weber et al. 2000:749 [article 10b]). the first name can be a diagnostic or a co-dominant species. a sub-community will start with the community name followed by a characteristic or dominant species for that sub-community (weber et al. 2000:749 [article 13]). variants can have only one name, usually referring to a diagnostic or dominant species for that variant. only an en-dash (–) is acceptable between the named taxa. example: plant community 1: themeda triandra–acacia karroo community; sub-community 1.1: themeda triandra–acacia karroo–diospyros lycioides sub-community and variant 1.1.1: selaginella dregei variant. • plant community names can have physiognomic descriptors at the end, but the style should be applied consistently. various combinations can be used, with, for example, only the major communities having descriptors. the descriptors are part of the name and should start with capital letters. environmental attributes are not used in the plant community name. example: themeda triandra–acacia karroo woodland or themeda triandra–acacia karroo short open woodland. • in the naming of plant communities, subjective emphasis may be placed on perennial (‘non-fleeting’) species that can be identified and found reliably during most years (not only during unusually good rainfall cycles). such robust species should preferably be used in the names of plant communities whilst short-lived annuals should be avoided. • it is also recommended that existing names for similar communities in related vegetation are retained to prevent several different names for similar communities. this should also assist when formal syntaxonomic classification of vegetation is undertaken. an attempt should therefore be made to avoid unnecessary synonomy. this implies that the researcher must be aware of the relevant literature. referees of papers should strive to identify such synonomy. description of plant communities although a plant community is known by its dominant species, its total floristic composition is characteristic where some species have a greater diagnostic value than others (beard 1980). it is recommended that the description of a plant community follows the standard format, namely starting with the locality and habitat (e.g. geology, land type, soil, topography, rock cover, altitude, erosion). this should be followed by the diagnostic species, which can be either listed or referred to in a table (preferred). the description is continued by listing the prominent (high-cover or abundance) and conspicuous species, their cover, growth form or any other relevant information pertaining to the community that would be useful in identifying and understanding the dynamics within the community. it is very useful to include colour photographs that illustrate typical examples of each plant community. it is highly recommended that the different communities are mapped using gis technology and that the map is included in the paper. syntaxonomic descriptions the need for formal syntaxonomic studies is recognised. smaller and localised data sets are, however, not suitable for formal syntaxonomic descriptions. these data sets need to be combined with other compatible data sets for a particular bioregion (mucina & rutherford 2006) to cover the variation of the various associations present. formal syntaxonomic classification and descriptions are discussed in detail by weber et al. (2000) and their recommendations should be followed for this purpose. recommendations for concluding remarks it has been found that readers of scientific papers pay most attention to the first and last paragraphs of a discussion (wenderoth 2012). it is therefore important that the first paragraph summarises the main findings of the research. in the last paragraph, the significance of the findings should be clearly explained.the researchers can decide whether they want to have separate discussion and conclusion sections (preferred by most journals). however, it is important that the discussion should not repeat what has already been stated in the plant community descriptions; rather, the results (i.e. plant community descriptions) should be discussed in relation to the original objective or problem stated at the beginning of the paper. the results are new knowledge and should be compared critically with similar vegetation studies elsewhere. similarities and differences should be highlighted and possible reasons given and discussed. it is important to refer continually to the results of the study and not to discuss unrelated aspects. other important aspects that can be discussed include endemic species, rare and endangered species, medicinal species, production, veld condition, environmental gradients within and between communities, and biodiversity of different plant communities. to emphasise the importance of phytosociological studies we cannot merely list the different plant communities and sub-communities anymore. therefore, an effort should be made in the discussion to discuss the conservation or biodiversity value and practical implications for the different ecosystems. concluding with applied recommendations increases the possibility that readers from other disciplines would find these studies worthwhile and interesting. it is, however, important that all recommendations be based on sound scientific theory. the conclusion should give a precise statement of the main findings and what they mean. the findings should also be mentioned within the context of previous findings. implications of the study’s findings, shortcomings of the study and further research or research questions should be mentioned. concluding remarks and recommendations top ↑ this article intended to provide broad guidelines for local phytosociological studies in conservation areas in southern africa. we believe that by developing guidelines for vegetation classification and descriptions in the southern african context, the quality and value of papers published will increase whilst, at the same time, preventing confusion between vegetation scientists in southern africa. although older datasets and studies are still recognised and valid, the use of modern analytical software and techniques is strongly encouraged. this will also align southern african vegetation studies with current international trends in vegetation science. local phytosociological studies in south africa are essential for efficient wildlife management programmes and conservation policies for ecosystems and biodiversity within national parks, nature reserves, private game farms and other natural areas. vegetation classification and descriptions provide information to interpret spatial variation between species as well as an understanding of vegetation–environment relationships (clegg & o’connor 2012). in addition, the predicted impacts of climate change on the environment makes the description of the vegetation of an area even more important, as it offers the only record of the current state of the environment. thus, classifications and descriptions offer baseline information for assessing the potential changes in the environment that may result from climate change from a plant and animal perspective (clegg & o’connor 2012). these data sets also play an important role in and contribute largely to formal syntaxonomic studies (luther-mosebach et al. 2012). this is an important reason for recommending a more standardised approach for local phytosociological studies, as it will produce compatible data sets that will assist with compiling a much-needed formal syntaxonomic classification for southern africa. for future development and growth of phytosociology in south africa, the curation of datasets and literature of all phytosociological research should be administered centrally (e.g. by the south african national biodiversity institute or the south african environmental observation network). the organisation should set up, facilitate and manage a system that will enable all researchers in this field to submit and extract electronic data and a list of publications and other literature for particular regions. such an information system will provide researchers with relevant literature and data, which could prevent unnecessary duplication of plant community names or research, and will encourage better communication and collaboration between researchers. ideally, south africa should move closer to a system such as synbiosys europe (http://www.synbiosys.alterra.nl/synbiosyseu) and synbiosys netherlands (schaminée, hennekens & ozinga 2012) to act as a national database. although south africa is currently developing synbiosys fynbos (http://www.synbiosysfynbos.org/program.html) and synbiosys kruger, these are only on a regional scale. acknowledgements top ↑ competing interests the authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article. authors’ contributions l.r.b. 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20, 11–51. http://dx.doi.org/10.4102/koedoe.v20i1.929 westfall, r.h., van staden, j.m., panagos, m.d., breytenbach, p.j.j. & greeff, a., 1996, ‘scale related vegetation sampling’, arc-range and forage institute, pretoria. westhoff, v. & van der maarel, e., 1978, ‘the braun-blanquet approach’, in r.h. whittaker (ed.), classification of plant communities, pp. 289–399, w junk, the hague. http://dx.doi.org/10.1007/978-94-009-9183-5_9 westhoff, v. & van der maarel, e., 1980, ‘the braun-blanquet approach’, in r.h. whittaker (ed.), classification of plant communities, pp. 287–399, w junk, the hague. zietsman, m.m., 2003, ‘phytosociological study of andrews field and tsaba-tsaba nature reserve, bredasdorp district, western cape’, msc dissertation, department of botany, university of pretoria. zietsman, m.m. & bredenkamp, g.j., 2006, ‘dune vegetation and coastal thicket communities in threatened limestone fynbos of andrews field and tsaba-tsaba nature reserve struisbaai, western cape’, koedoe 49, 33–47. http://dx.doi.org/10.4102/koedoe.v49i1.110 article information author: jane carruthers1 affiliation: 1department of history, university of south africa, south africa correspondence to: jane carruthers email: carruej@unisa.ac.za postal address: po box 392, unisa 0003, south africa dates: received: 05 oct. 2010 accepted: 07 mar. 2011 published: 30 june 2011 how to cite this article: carruthers, j., 2011, ‘pilanesberg national park, north west province, south africa: uniting economic development with ecological design – a history, 1960s to 1984’, koedoe 53(1), art. #1028, 10 pages. doi:10.4102/koedoe.v53i1.1028 copyright notice: © 2011. the authors. licensee: aosis openjournals. this work is licensed under the creative commons attribution license. issn: 0075-6458 (print) issn: 2071-0791 (online) pilanesberg national park, north west province, south africa: uniting economic development with ecological design – a history, 1960s to 1984 in this original research... open access • abstract • introduction • pilanesberg landscape and early politics • origins of the pilanesberg national park – the 1960s and 1970s • pilanesberg national park 1978–1983 • the process of revision 1983–1984 • conclusion • acknowledgements • references abstract (back to top) in the late 1970s, a ground-breaking project began in the pilanesberg district in what is now the north west province of south africa to create a wildlife conservation and eco-tourism venture from degraded marginal farmland in an aesthetically attractive extinct volcanic crater. the establishment of this national park was innovative in a number of respects, including a partnership between landscape and ecological designers, local community development and participation, regional tourist satisfaction, trophy hunting, environmental education, ecological restoration, and wildlife conservation and management. this paper briefly explored the park’s early history, explaining its landscape, its early peopling and historical land use. the narrative then concentrated on the first five years of the park’s existence, from its inception in 1977, under the aegis of agricor, bophuthatswana’s rural development agency, to 1984, when responsibility for the park was given over to bophuthatswana national parks, a parastatal agency, and a new era began. the article contended that 1984 is an appropriate date on which to conclude the early history of the pilanesberg national park (pnp) because it was then that the experimental phase of the park ended: its infrastructure was sufficiently developed to offer a satisfactory visitor experience, the management plan was revised, its bureaucratic structures were consolidated and an attitude survey amongst the local community was undertaken. embedding the originating period of the pnp in its historical, political and socio-economic context, the paper foregrounded those elements in the park’s beginnings that were new in the southern african protected area arena. thus, elements that relate to socio-politics, landscape and ecological design and restoration, and early relations with neighbouring communities were emphasised. this paper has been written by an historian and is therefore conceptual and historical, conforming in language and structure to the humanities style (environmental history). it relies on published and unpublished literature and oral information and the critical evaluation of these sources. conservation implications: the pioneering example of the pnp as a protected area is relevant to the field of conservation science because, as human population densities increase, as the tourism sector develops, as marginal farmland becomes available for new uses, and as it becomes important to include neighbouring communities in conservation activities, a study of this park’s early history and socio-political and economic context may be of assistance in the development of similar projects elsewhere in south africa and beyond. introduction (back to top) conservation managers and wildlife biologists in southern africa are familiar with the fact that national parks and other protected areas are often advertised and marketed as ‘unspoilt nature’, although they are, in fact, manipulated to meet objectives such as tourist satisfaction, carrying capacity, pasture and biodiversity management together with a variety of other key goals that may change over time. it is also true that many protected areas are far from being ‘pristine wilderness’ unaffected by past human activities, but are the consequence of ‘fortress conservation’ (brockington 2002) and the people (or their descendants) who were removed, often forcibly, in the interests of wildlife conservation have grievances that play out in the political arena. because of the human dimension involved in land use and present management, protected areas are not neutral spaces or landscapes without history; they are definitively shaped by their pasts.the pilanesberg national park (pnp), situated in what is today the north west province of south africa, was established in 1977. at the time, the area formed part of the western transvaal (a province of south africa until 1994) in an african reserve that was about to transform from an apartheid bantustan consisting of a number of ‘tswana homelands’ into bophuthatswana, nominally an independent state, an enclave within white south africa. it is an unusual national park from both points of view mentioned earlier. this particular area did not appear ‘natural’ at the time of its foundation. it was fully recognised to be property that had been heavily utilised and altered by many groups of people over the preceding centuries. it was deliberately and carefully redesigned as a national park, being restored ecologically from farmland and into which a wide variety of indigenous animals were reintroduced. once the white farmers had been expropriated (as will be explained at a later stage), the african people who returned to live in the pilanesberg were not to be forcibly removed as had been the case elsewhere; rather, their consent was sought and they were promised that the establishment of the national park would not be to their detriment but to their economic advantage. indeed, the very rationale of the pnp was that it was to become an engine of regional economic development. pilanesberg landscape and early politics (back to top) some 50 km north of the town of rustenburg lies the root zone of an extinct volcano – roughly circular, some 572 km2 (c.50 000 ha) in extent and measuring between 23 km and 28 km in diameter. it appears as a complex series of eroded rings of low mountains and hills that rise approximately 300 m – 600 m above the surrounding land. there is one perennial river and a number of freshwater and saline springs; however, the largest permanent body of water is mankwe dam (covering an area of approximately 2 km2), which was constructed by white farmers in the late 1950s. the climate is benign; the average rainfall is 600 mm – 700 mm per year, although there are regular droughts (farrell, van riet & tinley 1978; mccarthy & rubidge 2005; mucina & rutherford 2006). in terms of vegetation, the pilanesberg is significant because it is a transition zone between the arid savanna and the moist savanna biome. owing to the complex substrate, there is a wide variety of landscapes and habitats for both plants and animals (farrell et al. 1978; mucina & rutherford 2006). this landscape within the crater is aesthetically attractive and was the subject of comment by many 19th century travellers and explorers, amongst them thomas baines, who painted the ring of hills in 1869 as he journeyed into what is now botswana.the region had been inhabited continuously by tswana-speaking people probably for many centuries. at the time of permanent white settlement in the mid-19th century, the bakgatla bakgafela clan lived in the pilanesberg area under their chief pilane (d. 1850), who gave his name to the modern district. according to makgala (2009) and mbenga (1996), this community can also be referred to as the ‘kgafela-kgatla’, ‘bakgatla-baga-kgafela’ or ‘bakgatla-ba-ga-kgafela’ – meaning the ‘kgatla people of kafela’ in various forms of the tswana language. kgafela was the kgosi (chief) who originally gave his name to the community. the current head of the clan is kgosi kgafela ii, who resides in mochudi, botswana, whilst kgosi nyalala pilane leads the group at saulspoort. the bakgatla was one of the few groups that did not resist the arrival of the boers but became their allies, assisting them in exploits of war and ivory-hunting. their association with white settlers and access to firearms enhanced the bakgatla powerbase and enabled them to increase their cattle herds and vanquish some of their local rivals and enemies. in later decades, however, the bakgatla lands were commandeered and carved up into settler farms on which the bakgatla became rent-paying or labour tenants. many of the community settled at saulspoort (breutz 1953; makgala 2009; manson & mbenga 2009; mbenga 1996, 1997; mbenga & morton 1997; morton 1992, 1995; schapera 1953). in 1913, the natives land act (act no. 27 of 1913; union of south africa) confined black south africans to very limited areas of the country and, in 1936, the native trust and land act (act no. 18 of 1936; union of south africa) attempted to provide more land for africans by designating ‘released areas’ that were to be purchased by the south african native trust and added to the african reserves. the pilanesberg was one of these released areas and the white-occupied farms were very slowly expropriated until the exercise was complete in the early 1960s. the bakgatla were thus allowed to return to their ancestral land. in 1961, the growing severity and oppression of apartheid politics affected the pilanesberg directly when it became a designated ‘tswana homeland’. during the 1970s, these various tswana homelands were consolidated into a number of islands within white south africa and became the ‘independent nation’ of bophuthatswana in 1977. regional politics were fraught. tidimane pilane, kgosi of the bakgatla, and lucas mangope, head of the bahurutshe clan, were rivals, pilane supporting the african national congress (anc) and mangope the apartheid state (butler, rotberg & adams 1977; jones 1999; lawrence & manson 1994). origins of the pilanesberg national park – the 1960s and 1970s (back to top) the principles underlying wildlife and conservation management in southern africa were changing during the 1960s and 1970s (carruthers 2007a, 2007b, 2008). in terms of philosophy, the idea of utilising wildlife sustainably by cropping and translocation gained ground in parts of africa, replacing an older tradition of strict preservation. at the same time, there were technical improvements in the transporting and immobilisation of wild animals that led to the easier movement and sale of wildlife (dasmann 1959, 1964; dasmann & mossman 1960, 1961; eltringham 1984; johnson et al. 2008; mossman & mossman 1976). in conjunction with bophuthatswana politics, these developments were relevant to the formation of the pnp. indeed, the park could not have come into being in a pre-translocation and pre-game sales era.apartheid social planning gained momentum during the 1960s and there were government initiatives to make the bantustans more self-sufficient economically and thus able to sustain a larger number of black africans outside of ‘white’ south africa, with a view to separating the homelands permanently from the other ‘white’ parts of the country. in 1969, there was a recommendation from ‘apartheid’s social engineers’ – a ‘potchefstroom-based team of “development experts” commissioned to find ways of enhancing the economic viability of an “independent state”’ (van onselen 1996:477) – that the crater be made into a recreation resort and nature reserve. however, for reasons that are unclear, nothing came of the idea at the time, but it was raised again in 1973. the following year mangope established a feasibility study (brett 1989; johnson et al. 2008; magome & collinson 1998). the matter received a boost when the southern sun hotel group – which, through managing director sol kerzner, had close ties with the bophuthatswana president and his government – finalised a plan to build a casino and hotel resort in the pilanesberg (to be named ‘sun city’) that would bring revenue into the region. at a time of strict petrol rationing and thus the curtailment of long-distance motor car travelling, it was expected that a game reserve adjacent to the hotel would provide an added attraction for tourists from johannesburg and pretoria, who would flock to sun city for the kinds of entertainment not available in white south africa, such as multiracial mingling, soft pornography and gambling. having first considered the location of mankwe dam for the hotel, the facility was relocated to its present site and construction began in 1978 (bureau for economic research re bantu development [south africa] 1978; boonzaaier pers. comm., 01 march 2010). after gaining independence, bophuthatswana established a number of organs of state. one of these was a parastatal development body tasked with promoting rural self-sufficiency. named the agricultural development corporation (agricor), this body fell under the bophuthatswana department of agriculture and was to play a decisive role in the establishment of the pnp. through its managing director, david beuster, agricor raised the funds for the game reserve and, despite the fact that many bakgatla people and livestock lived in and used the crater, and that it contained numerous farm houses, roads, dams and fences, the pilanesberg was formally proclaimed a nature reserve in 1977. it is worth emphasising that agricor, a parastatal body specifically responsible for economic and community development in the rural sector, was given the administration of this future national park rather than the department of nature conservation, because it was regarded as a rural improvement project and not a nature conservation exercise. what was audacious for the period was that beuster and mangope employed landscape architects to design this game reserve adjacent to sun city. the firm that was instructed to act as consultants to draw up a management plan was farrell and van riet, landscape architects and ecological planners, then a recently established pretoria-based company, and it was instructed to act as consultants and to draw up a management plan. by the time he established the partnership of farrell and van riet, willem van riet was a leader in the field of landscape architecture in south africa and he was primarily responsible for linking landscape architecture with ecological planning in the country. van riet had initially qualified as an architect at the university of cape town but thereafter, from 1972 to 1975, he had benefited from studying at the university of pennsylvania under ian mcharg, the renowned landscape architect and author of design with nature (1971), a book that is widely regarded as one of the most influential of the 20th century (schnadelbach 2001). in his autobiography, mcharg explained that the genesis of design with nature lay in a meeting between himself, russell train, the president of the conservation foundation and ray dasmann, a noted ecologist and the foundation’s chief scientist. apparently, train said, ‘ian, ray and i have decided that the time has come for a book on ecology and planning’ and mcharg agreed to write it (mcharg 1996). train and dasmann were correct: the book was perfectly timed and widely used and quoted. in a quest for life (1996), mcharg describes landscape architecture as a discipline very close to nature and its preservation and he was particularly keen to encourage planning that was appropriate to specific environments. mcharg sought out trained architects such as van riet for his postgraduate landscape architecture programme, providing not only a stimulating academic environment but substantial financial subsidies (mcharg 1981). in this way, and through van riet, ideas around ecologically apt planning and design from the usa made their way into southern africa. sharon kingsland argues that the science of nature reserve design emerged in tandem with the interdisciplinary field of conservation biology. she explains that the basic rationale for such design is to protect biodiversity, using ideas from island biogeography, prioritising conservation of the indigenous species of plants and animals of the area, and employing operational research and mathematical techniques for linear programming (kingsland 2002a, 2002b). if this is the norm, then the pilanesberg was highly unusual because the biodiversity had been totally compromised by farming activities and there were extremely few remaining indigenous plants and animals – certainly large mammals had become locally extinct. the creation of the pilanesberg involved little conservation biology and focused, at first, entirely on ecological restoration and landscape design. the national park emerged from the collaboration between van riet and ken tinley, a young ecologist who had also been inspired by mcharg. as a university student, tinley advocated mcharg’s design with nature to his contemporaries (huntley 2010). tinley was one of an emerging new generation of wildlife ecologists in south africa (many of whom were educated at the university of natal). in partnership with van riet, he worked on a number of nature reserves, particularly in the ‘homelands’, where there was scope for new ideas because the reserves in these localities were not in the control of the various philosophically and bureaucratically entrenched provincial nature conservation authorities or the national parks board. these included locations in pondoland (including mkambati), in maputaland and in the gorongosa national park in mozambique, as well as private game reserves (farrell & van riet landscape architects and ecological planners 1975; tinley 1978; tinley 2010; van riet 2010). van riet and tinley were employed to design the pnp and they presented their report in 1978. what they suggested was somewhat revolutionary in the context of southern african national park and game reserve planning and it marked a strong contrast to the fortress conservation and wildlife management practices that then held sway. entitled ‘pilanesberg national park: planning and management proposals for department of agriculture, republic of bophuthatswana’ (farrell et al. 1978), this is an important document and worth summarising in some detail. the report began with what was a provocative premise in an era of fortress conservation: that any conservation measure would ultimately be futile unless wildlife and nature could deliver tangible, visible benefits to humans within a particular socio-economic and geographical milieu. in other words, the survival of wildlife in africa was dependent on rural african people. van riet and tinley (farrell et al. 1978) argued that protected areas should not be viewed in isolation, but in their regional ecological and economic contexts as productive primary (ecological services) and secondary (tourism, education and wealth-creation) landscapes. the report paid particular attention to wildlife as a source of protein, as well as of traditional medicine and other natural products that might be sustainably harvested by local people, together with wildlife tourism being a source of employment and income. these principles were in sharp contrast with those espoused by, for example, the national parks board (now sanparks) that were focused on settler values that emphasised white, middle class tourist recreation and created places in which: lessons in tidiness, adherence to and acceptance of rules and regulations … [were taught] [where] people can be disciplined not to litter, not to pick flowers … [and that generate] tranquillity … so desperately needed in a world where people are caught up in the tensions of city life. (knobel 1979:233) in contradistinction to this viewpoint that urban visitors and romantic ideas of wilderness were the focus of nature conservation initiatives, tinley (1979) had expounded the philosophy that the regional context of any conservation project was decisive. he strongly believed that the needs and aspirations of rural people were paramount and argued on the basis that many conservation departments throughout africa: have based their activities on the dogma that tourism and wildlife conservation are two sides of the same coin. thus staff and funds are used mostly for catering and tourist facilities and the natural areas become filled with urban nuclei to justify the existence of parks. in this way conservation departments continue to be directly responsible for despoiling the last wild places for which they are custodians. (tinley 1979:33) the pilanesberg was to be different and the aim was to reflect a new vision of conservation practice in africa. not only did they introduce the radical idea of using national parks sustainably as engines of regional development sensitive to local community needs, van riet and tinley also introduced a novel concept of planning and design (farrell et al. 1978). after surveying the geomorphology and other aspects of the landscape, habitats and vegetation cover, their report proposed that in order to maximise wildlife viewing in the small area and utilise it to the best advantage, all major tourism facilities should be located on the boundaries of the park, thus preserving the interior of the crater from unsightly camps, restaurants and other amenities. this peripheral development was different from other national parks and game reserves which had normally sited major visitor accommodation within the protected area itself. moreover, using the internal watersheds as ecological borders, van riet and tinley suggested dividing the crater into seven (later reduced to five) distinct activity zones. there would be zones for trails, hunting, visitors, wilderness, special use, multiple use and peripheral development. there would be no roads or amenities in the wilderness and special use zones and a buffer zone would separate these two from the resource utilisation areas, the rest camps and the intensive use areas. there were specific recommendations for planning and using each of these zones so that activities would harmonise with each other (farrell et al. 1978). the pilanesberg project was highly unusual in that van riet and tinley had a free hand and a flexible institutional, bureaucratic and policy environment within in which to work. they were not burdened with an entrenched public service, hidebound politicians, an historical legacy of preservation philosophy, or outdated or ill-sited roads and other tourist amenities (child 2008). moreover, the pilanesberg crater, with its rings of hills that hid the plains and human developments beyond from view, was the ideal topography in which to recreate a natural-looking environment. whilst at the start there were cultivated lands, evidence of stock grazing and farmsteads, alien vegetation, roads, and so on, once these – and the people – were removed, the crater presented an almost clean slate for design. a list of appropriate mammals that should be introduced was provided in the section on ‘management proposals’ (farrell et al. 1978) and appropriate herd sizes given. also departing from the then accepted norm in managing protected areas, van riet and tinley made provision for trophy hunting and prioritised environmental education. in short, the report designed a 50 000 ha national park, literally from the bottom up. pilanesberg national park 1978–1983 (back to top) in the opening chapter of their report, van riet and tinley had made strong statements about the holistic philosophy of sustainable national parks and the role of local people within them, but, on the whole, they focused on planning the future ecological management of the pnp (farrell et al. 1978). it appears that they assumed that politicians and sociologists would take care of the human and community dimensions of the enterprise. unlike other national parks and protected areas in south africa that had involved forced removals, the idea for the pnp was that the bakgatla would participate in decisions about the new national park, vacate the crater area voluntarily, contribute to its social and economic planning, and to its management thereafter. in this regard, the fact that the new bophuthatswana ‘nation’ planned a national park was important in terms of nation-building, and the pilanesberg was to be the public demonstration of these ‘civilised’, modern and international values (carruthers 1997). early in 1979, negotiations were completed between van riet and tidimane pilane and, in turn, between pilane and the other bakgatla chiefs (keenan 1984). the bakgatla, through their kgosi, agreed to surrender their grazing and land rights in the crater. just how ‘voluntary’ this agreement was, has later been hotly contested in a land restitution claim instigated by the bakgatla who aver that they were strongly coerced by the ‘strong-arm tactics’ employed by mangope and his officials (mbenga 2011) who were determined to steamroller the game reserve through for the benefit of sun city and its supporters. in any event, the bakgatla did not control the entire pilanesberg crater. the national park consisted of freehold land of 8500 ha, obtained directly from the resident bakgatla (viz. schaapkraal, welgeval and portions of legkraal, koedoesfontein, kruidfontein, saulspoort, rooderand and doornpoort), whilst some 4500 ha came from a newly arrived group, the bakubung, namely wydhoek and portions of ledig and koedoesfontein. in addition, 1000 ha was obtained from private owners and the rest – the majority of the property – consisted of 46 000 ha of state land that had been expropriated from white people by the department of bantu affairs (as explained earlier) to augment the paucity of land allocated to africans (keenan 1984:14). the bakgatla apparently acquiesced in their relocation from the pilanesberg crater on the basis that they would be allocated two nearby state farms to replace their lost communal grazing and that they would be fully compensated for the land and structures that were required by the national park. in addition, they would be recompensed for the full costs of removal and also retain the right to enter the reserve in order to visit graves and to collect firewood, thatching material and medicinal plants. moreover, they were promised an (unspecified) portion of entry ticket sales and tidimane pilane was to be appointed onto the national park’s governing and management board of trustees, thus ensuring a bakgatla a voice in a form of joint management (magome & collinson 1998; makgala 2009). it was unfortunate that these agreements were both informal and verbal and thus neither effectively witnessed nor formally contractual (van riet 2010). subsequently, it emerged that pilane, a political opponent as has been previously explained, had been threatened by mangope with eviction from other state land if he did not agree to the pnp proposal (keenan 1984:16–17). moreover, in later years, complaints surfaced that pilane, who was not a universally popular leader, had not adequately consulted with the rest of the community (makgala 2009:33−335) and thus did not speak for everyone. today, there is considerable sophistication in all quarters when consulting communities affected by national parks and other protected areas, but at the time of the pnp’s establishment there were no protocols to follow and no mechanisms for predicting or resolving disputes or difficulties that might arise between the various parties involved. there were no such examples to follow in the 1970s and in the absence of experience on the part of van riet and the bakgatla – and in the context of high apartheid and mangope’s dictatorship – many details were left vague or unrecorded. one needs to recall, however, that the bakgatla were not the only people affected by the establishment of the pnp. whilst tidimane pilane, as a traditional kgosi, was in ostensible control of bakgatla ‘communal land’, the farm welgeval inside the crater was inhabited by a community of long standing in the area, who owned part of the property in their own right. they agreed to relocation, provided they were fully compensated and this agreement was formally documented. it is on the basis of this written evidence that the welgeval community has subsequently been awarded a land restitution claim on this farm and the land has been leased back to the pnp (manson & mbenga 2009). many other local people were also not consulted, presumably either because they were considered to be fractured groups without leaders to give them voice, or perhaps because they lived on the borders of the pilanesberg rather than within it, or, even, perhaps because they were mangope dissidents and were thus ignored. thus many people were dissatisfied with, and disadvantaged by, these arrangements. for example, the farm ledig (south of the pilanesberg) was occupied by the bakubung, part of a disunited tswana-speaking group that had been forcibly removed from the outskirts of the small town of boons where they had formed a ‘black spot’ within white south africa and were therefore obliged to relocate in the late 1960s. they used the crater for grazing their cattle and goats. apparently, as far as these people were concerned, ‘care was taken to ensure that all talk of the project was kept away from the bakubung notables and strictly confined to official circles’ (van onselen 1996: 477). soon, without warning, there were reports of ‘a giant game fence snaking across the pilanesberg’, and excluding them and their livestock (van onselen 1996: 498). there were also large numbers of non-tswana nguni-speakers, many of whom had also been forced into a homeland from urban areas or white farms, who also had to make use of the pilanesberg for their survival. it seems evident from oral sources and later comments that, whilst negotiations and participation had occurred at top political levels, the views of ordinary people had neither been sought nor taken into account. this particularly included those who were not part of the formal ‘tribal’ structures of the district, newcomers and outsiders and who perhaps would pay the heaviest price in terms of losing access to land and livelihood opportunities (keenan 1984:39–43). owing to its bold conception and future plan, the pilanesberg project received considerable local and international publicity. local people were negative about the creation of the pnp because it impacted directly on their lives, whilst many scientists and conservationists were critical of its ambitious ‘operation genesis’ – the mass reintroduction of many species of wildlife. added to the difficulties (and the adverse publicity) was that development and administration did not proceed smoothly. the personnel was generally incompetent and many officials of agricor lacked experience and, for this and other reasons, they were either removed from their posts or encouraged to resign (brett 1989:112). it was only in october 1980 that jeremy anderson, whose doctoral research in zululand and subsequent study on lion management in the umfolozi game reserve (natal) had gained him a reputation as a capable and knowledgeable wildlife manager, was employed by beuster as director. anderson – with his scientific expertise, energy, enthusiasm and familiarity with the new scientific thinking emanating from east africa – was tailor-made for the job (boonzaaier pers. comm., 01 march 2010). anderson was joined by willem boonzaaier, previously employed in the private sector, as chief administrative officer to handle the financial side of the operation. between them, they appointed qualified and appropriate staff, including ecologists roger collinson and peter goodman in 1981. those involved in these early days recall the magnetism of being associated with what was then an experimental nature reserve, of working in a multiracial environment, and of encountering the dominant personalities and interesting characters who sought to put pilanesberg on its feet (owen-smith, magome & grossman pers. comm., 01 january 2011). the success of the pnp was predicated on that of sun city, and the resort prospered to the extent that it was expanded in 1981 and again in 1984. the close friendship between kerzner and mangope meant that the bophuthatswana political elite in the mangope government were extremely supportive of the sun city development, as was the south african regime; however, tidimane pilane was less enthusiastic for a number of reasons. firstly, he was a member of the opposition party, an anc supporter with a different vision for the country’s future, and he disagreed with many of the policies of the mangope government. secondly, he believed that sun city offered no benefit to local people because employment in the resort went to outsiders to the district (makgala 2009:322–323), a grievance shared by the bakubung. thirdly, tourist revenue, one of the major reasons why the bakgatla had agreed to vacate the crater in favour of a national park, was minimal, despite the promises that visitors from sun city would flock to the reserve, and no financial gain came their way. the pnp was also criticised because wildlife introductions had not gone as smoothly as planned. owing to the fact that some species were being given away freely by south african conservation bodies that had problems of over-stocking, whilst others were inexpensive to acquire, the pilanesberg became stocked with an incorrect balance of wild animals. this resulted in some habitats being inappropriately modified because of over-grazing, leading to even further reduction in biodiversity and condemnation from local and international scientists (van aarde 2010). for reasons of veterinary and disease control, it was difficult to obtain appropriate species because wildlife movements were curtailed from places in which cattle diseases were endemic. wildlife therefore had to be sourced from disease-free populations a long distance away, such as namibia or the eastern cape, and transport was thus extremely costly. additionally, because adult elephant males can pose a danger during the capture and transport process, fewer of them were translocated in comparison with females and young and thus subsequent breeding success was low and herd sizes and composition were skewed (garaï et al. 2004; hancock 1983). the mixing of gene pools was also a matter of scientific concern. some introductions were made before the fencing was complete and animals were kept in a holding camp that was too small, which resulted in many animal deaths once the grazing inside the camp was depleted (collinson & anderson 1984:169–70). these introductions of large mammals were extremely costly and the bophuthatswana government was not able to fund them. the money came from the south african wildlife foundation (sawf), founded in 1968 by afrikaner business magnate anton rupert, who was a trustee of south africa’s national parks board and who had close ties with the national party. in 1979, rupert presented the pnp as a project to the world wildlife fund international and the international union for the conservation of nature and received approval to fund it (schwarzenbach 2010). given rupert’s personal belief in cultural and linguistic ethnicity, assistance to the ‘nation’ of bophuthatswana through the sawf was entirely in character. indeed, bophuthatswana may have held special significance for the rupert family, for his wife, huberte (neé goote), came from the western transvaal and she spent some years at derdepoort on the botswana border (domisse 2005:353). as can be appreciated from the aforementioned explanations, the early years of the pnp presented huge challenges to those involved. there was no existing institutional policy framework – either scientific or bureaucratic – within which to operate. whilst this meant freedom from precedent, it also meant that the enterprise moved slowly. owing to the limited number of rangers and labourers that could be employed, the process of restoring a natural environment – a ‘mammoth task’ that hancock (1984) referred to as ‘renaturalisation’ – was extremely slow. anderson (pers. comm., 23 february 2010) recalled that in terms of basic infrastructure virtually nothing in existed and a perimeter fence, off-loading ramps, translocation stations, bomas, feedlots and pens had all to be constructed by the small staff. more than 1000 km of internal farm fencing had to be removed, as had the many solid concrete cattle dips, farm reservoirs and windmills, about 30 large farmhouses and over 100 smaller houses, outbuildings and huts. these were bulldozed and the rubble was used to fill large dongas (hancock 1984). borrow pits, landscape scars and old lands had to be rehabilitated and general farming detritus (e.g. old vehicles and heavy, rusted implements) cleared. invasive alien plants were abundant, not only jointed cactus (opuntia spp.), but huge old trees, especially australian eucalypts – all these had to go, some being felled, others poisoned (anderson pers. comm., 23 february 2010; hancock 1984). in addition, there was no living or office accommodation for game guards, management or labourer staff or for stores. prefabricated buildings were erected to meet some of these needs but, for a few years, all management personnel and their families lived in caravans. workshops and vehicle maintenance points were also needed. the construction of permanent buildings, game-viewing hides and visitor amenities such as camps and entrance gates proceeded slowly, as did re-siting of roads to make them suitable for game drives (old straight farm roads had to be obliterated). the local staff was unskilled and thus training, education and mentorship had to be provided. at this time there was no regular telephone communication in the park and only radio phones could be used – a scarce and expensive resource (anderson pers. comm., 23 february 2010). veld and wildlife monitoring systems also had to begin anew and proceed in tandem with applied management tools such as a burning regime. jules turnbull-kemp, a senior game ranger recruited by agricor from rhodesia, and who later became warden of the pnp, was responsible for receiving the wildlife introductions. at the time, there were no formal studies to assist with determining the ability of different species to survive or thrive on old farmlands and only by observation and experience did it emerge how animal populations coped with, and altered with, the recovery and restoration of the habitat (turnbull-kemp pers. comm., 01 march 2010). because large-scale models (such as the kruger national park) could not be applied to a small area like the pnp, fire policy also had to be determined from scratch. an innovation in terms of management philosophy was that stocking was carried out at a high rate (i.e. many animals at one time) so that the take-off rate (reducing numbers through hunting) was optimised as soon as possible (turnbull-kemp pers. comm., 01 march 2010). not surprisingly given the dearth of wildlife and abundant unsightly evidence of former fields, houses and roads in the crater, tourism did not take off quickly, indeed tourists were not encouraged for a number of years until a ‘satisfactory game viewing experience’ could be guaranteed (hancock 1984). anderson had doubted that gate revenue from day visitors (which was to be the major source of income for the bakgatla) was ever likely to produce any substantial income (boonzaaier pers. comm., 01 march 2010) and so took the decision to introduce trophy hunting to generate some immediate income. because the pnp was an agricor initiative, and was the only project of this nature in its stable, the administrative and financial arrangements were as independent as – and could be as experimental as – those of the conservation management. as administrative officer, boonzaaier adapted commercial systems to the park’s requirements. there was no model to follow: the park had to be up and running as quickly and profitably as possible. the first question was: where was an initial income to come from? in this regard, wildlife management and administration were able to dovetail. anderson’s idea of revenue-producing trophy hunting could only take place if there were surplus animals to shoot. to determine the optimum stocking rate that would be needed to manipulate species numbers to obtain the best returns, anderson, collinson and boonzaaier designed a complex model to determine how many (and which) species were required to profit most from game sales, hunting, meat production or tourist viewing. wildlife populations were therefore predicated on formulas that demonstrated the best return on investment, per land unit (anderson pers. comm., 23 february 2010; boonzaaier pers. comm., 01 march 2010). the pnp’s management was innovative because, instead of a few wild animals of various species being introduced and then allowed slowly to build up their numbers, large populations were introduced at the start and thus numbers increased very quickly, providing a surplus after only a year or two in the case of some species (anderson 1986). very careful records of net production versus utilisation were maintained. these calculations were novel because they were being made for the first time in a protected area: wildlife was being taken into account as a financial asset, not merely a ‘nature conservation’ ethical good. just as cattle and game farmers entered their herds into their accounting books and measured the profit from them, so too did this national park (boonzaaier pers. comm., 01 march 2010; a more detailed history of wildlife management and conservation science in the pnp will be the focus of a later paper by the present author.) however, all these developments took time. moreover, the early 1980s saw one of the subcontinent’s worst droughts of the century. this meant that the rehabilitation and restoration of the pnp grasslands and vegetation took far longer than anticipated; it also meant that the displaced bakgatla and others were short of grazing on the farms to which they had been relocated and many looked longingly at the recovering (albeit slowly) veld in the pnp that had been free of grazing cattle for a few years (manson & mbenga 2009). during the drought, mobile pnp animals wrought havoc on properties outside the reserve. baboons, in particular, climbed over the perimeter fences and ravaged the maize fields of neighbouring black communities in ledig and elsewhere (van onselen 1996:510). the process of revision 1983–1984 (back to top) by 1984, despite obstacles and slow progress, it could be said that the pnp was maturing and that it had come to a stage when an overall review to guide its future strategic direction was required. in that year, a new management plan was devised, the bophuthatswana national parks board was founded and a community relations survey was conducted. considerable experience had been gained by park management during the five years since the park’s opening and the pnp began to meet some of its objectives. as hancock (1984) described in response to those ‘wondering exactly what, if anything, has been happening in the pilanesberg’, during this time the park had been fenced, the landscape rehabilitated, buildings razed and obliterated, tourist roads constructed, wildlife introduced, two visitor camps constructed and foot safaris and trophy hunting operated satisfactorily. it seemed that the experimental phase was ending and that consolidation and review was needed (collinson & goodman 1982). tinley was not involved in the re-planning process as he had left south africa by that time. the new report, ‘a five year development plan for pilanesberg national park as requested by the bophuthatswana government and the bophuthatswana national parks board, september 1983’, was authored by willem boonzaaier, roger collinson and willem van riet. because the construction of tourist facilities within the national park had been so slow and the project had been so costly, these managers feared that the investment of the previous five years in management, rehabilitation and wildlife introductions might be wasted unless clearer objectives were re-established. the primary goal of the pnp was stated to be to ‘maintain and where necessary create an ecosystem comprising a biota of as wide a variety of indigenous plant and animal species’ as possible (boonzaaier, collinson & van riet 1983). the secondary objective was defined as ‘to utilise the area and its natural resources in ways that will yield the greatest benefits to bophuthatswana and its people, both now and in the future’ (boonzaaier et al. 1983). the multiple zoning of the initial plan had proved to be too complex to manage effectively. boonzaaier, collinson and van riet thus recommended re-zoning the park into two overarching types of areas, a ‘managed natural area’ and a ‘natural environment recreation area,’ each subdivided into zones. within a ‘managed natural area’ (i.e. well within the park’s boundaries), accommodation would be limited and cater for very small groups, with the only permissible activities being walking on designated trails and trophy hunting. the ‘natural environment recreation area’ would be devoted to general visitor and multiple uses and would include peripheral development at the manyane, bakubung and bakgatla gates. however, these plans would be extremely expensive and the park would probably continue to run at a loss whilst development proceeded through various phases (boonzaaier et al. 1983). the year 1984 was also significant because the overall managerial and bureaucratic structures of the park were altered when a national parks and wildlife management board for bophuthatswana was created along the lines of a parastatal to manage the pnp. this new structure, which was formalised in 1987 with the national parks act (act no. 24 of 1987; bophuthatswana government), resulted from the merger of the division of nature conservation of the department of agriculture and forestry and agricor, thus ending their somewhat competitive and even acrimonious relationship. the new national parks organisation, nicknamed bopparks, was to be managed by an appointed board. this move was significant because it meant that the pnp was now freed from its roots in a rural development organisation and was provided with a more conventional home within a parks board. this was to impact on its later trajectory. the need to review, and if necessary to change, the objectives and management of the pilanesberg in 1984 may also have been related to political and economic factors inherent in the bophuthatswana state. at this time bophuthatswana was economically stressed and, in fact, was experiencing ‘an acute financial crisis’ (jones 1999, 2001). there was also growing political dissension and even violence as a rupture developed between the democratic and inclusionist policies of anc supporters in bophuthatswana, including tidimane pilane, and the mangope faction, with its ideal of an ethnic tswana nation in an artificially segregated south africa (jones 1999). within the broader south (and southern) african political landscape, violence, revolutionary activities, harsh repression and military intervention were endemic and an atmosphere of tension was the order of the day. many of the flashpoints thereof were in the so-called ‘independent bantu states’. in this atmosphere of political turmoil it was clear that relations between the pilanesberg’s managers and the bakgatla had deteriorated. the matter was aggravated by the fact that tidimane pilane was sidelined when he was not appointed to the bopparks board as he should have been in terms of the ‘agreement’ with van riet (magome & collinson 1998; makgala 2009:321). this affront marginalised and offended the bakgatla, who were anc allies and thus opposed to mangope’s regime. together with the fact that no monetary compensation was accumulating for the community (which is what they had been promised) because there were few visitors and thus little by way of gate fees, relations between the pnp and the bakgatla were tense because of ‘broken promises’. perhaps the initial undertakings of beneficiation had been over-generous, but the bophuthatswana government had apparently reneged on agreements about land and financial compensation to people the state regarded as political opponents. in terms of the breakdown in communication between neighbours and the pnp, magome and collinson (1998) believe that it owed much to the heavy demand for very rapid development and effective wildlife and administrative management, which meant that park authorities had little time to devote to nurturing community relationships. in order to identify and address issues of concern, it was decided to conduct a formal survey of the attitudes of the local people to the pnp. in 1984, jeremy keenan, a sociologist then employed at the university of the witwatersrand (and who did not disguise his anti-bophuthatswana views), was tasked to report on community relations (keenan 1984:5). keenan and his researchers uncovered seriously negative perceptions of the national park at many levels. there was discontent over the verbal initial arrangements regarding the evacuation of the crater and inadequate financial and property compensation, particularly as cattle-rustling and other theft of property occurred during the removals. the bakgatla perceived the administration of bophuthatswana and its officials as ‘dictatorial and deceitful’ (keenan 1984) and they alleged that farms intended for compensation had been given away to government ministers and mangope cronies. to demonstrate their discontent, they had decided that they wanted to take back their land in the pnp and, to this end, had begun a court action (keenan 1984). the fact that tidimane pilane was a leading figure in opposition politics exacerbated the situation further. people involved at the time believe that the pnp was used to score political points in these oppositional politics at a time of unrest in south africa, creating divisions even amongst bopparks staff members (boonzaaier pers. comm., 01 march 2010). however, despite the grievances and the misunderstandings about the function of a national park that were reflected, keenan’s report also indicated that there was some local support for the pnp and for its educational outreach programme in particular (keenan 1984:67–74). because the report was leaked to the media by keenan himself, it attracted a great deal of attention that resulted in the managers of the pilanesberg being caught in the middle of the fracas between the government and the bakgatla (magome & collinson 1998). conclusion (back to top) it is now nearly 35 years since the establishment of the pnp and if the argument here has been that circumstances had shifted so much in the first five years of its existence that a review was necessary, then how much more has the context changed in the 30 succeeding years. not only did bophuthatswana itself undergo violent revolutions in 1988 and 1994, which saw lucas mangope and all that he had stood for overthrown, but the republic of south africa itself underwent a peaceful democratic revolution in 1994 that totally transformed the political environment of the country. this had enormous repercussions through every organ of government, as well as impacting greatly on the socio-economic environment, including in the conservation arena. in the post-1994 ‘new’ south africa, the management philosophy, objectives and style have altered considerably in both the pnp and in other game reserves in what is now north west province.nature conservation – although some argue that tourism income and economic beneficiation determines policy more than biodiversity conservation (johnson et al. 2009) – has expanded in the north west province and it has retained the use of ‘national park’ even though the region is no longer a separate ‘nation’. whilst governance of the protected areas in north west has shifted with the changing responsibilities of various provincial departments, thanks to the original mission of the pnp to assist rural development and upliftment, natural resource management as an income-generating, employment-creating and capacity-building exercise remains a high priority in the region. despite the political uncertainties, the pnp had become successful by 1991 and the bophuthatswana government then took over marginal farmland on the botswana border, reclaiming and stocking it in the same way as pnp to create the upmarket madikwe game reserve. after 1994, and the establishment of the north west parks and tourism board (note the inclusion of tourism in the name of this body) plans were to form a substantial heritage corridor that would link the protected areas. events subsequent to 1984 were extremely important and warrant further study. moreover, the lessons learnt around ecological restoration and wildlife and conservation management require careful research and evaluation, as do their influence in other parts of south africa. nonetheless, it is instructive to reflect on how the creation and management of the pilanesberg national park during its early years introduced a number of fresh developments into south african natural resource design, management and conservation, some of which have become more important in the current protected area estate. whilst not adopted universally, some of the experimental aspects of the pilanesberg, including peripheral development, the consumptive use of wildlife in protected areas, the provision of a variety of visitor accommodation, community engagement and local empowerment, reclamation of farmland, translocation of wildlife, trained african senior personnel, environmental education, and a commercialisation and concession policy, have now become a part of modern conservation practice in both state-owned and private protected areas. research relating to how the innovations in landscape design, wildlife management and community issues later influenced developments in other protected areas is currently being undertaken by the present author and it is anticipated that another, more focused publication in this regard will result. acknowledgements (back to top) i would like to thank the following for their generous assistance in providing inspiration, guidance, information, documentation, discussion and for correcting factual and editorial errors: jeremy anderson, willem boonzaaier, bruce brockett, vincent carruthers, roger collinson, david grossman, brian huntley, cynthia kemp, hector magome, bernard mbenga, eugene moll, archie mossman, sue mossman, norman owen-smith, alexis schwarzenbach, ken tinley, rudi van aarde and willem van riet. financial support from the university of south africa, the national research foundation, and the garden and landscape studies department of harvard university is also gratefully acknowledged. adapted from a paper delivered at the 2010 dumbarton oaks garden and landscape studies symposium, 'designing wildlife habitats' (14 and 15 may, 2010), and included in the forthcoming volume wildlife habitats (2013) published by dumbarton oaks research library and collection. references (back to top) anderson, j.l., 1986, ‘restoring a wilderness: the reintroduction of wildlife to an african national 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