cronje&reilley.qxd Natural mortality amoung four common ungulate species on Letaba Ranch, Limpopo Province, South Africa H.P. CRONJE, B.K. REILLY and I.D. MACFADYEN Cronje, H.P., B.K. Reilly and I.D. Macfadyen. 2002. Natural mortality amoung four common ungulate species on Letaba Ranch, Limpopo Province, South Africa. Koedoe 45(1): 79–86. Pretoria. ISSN 0075-6458. Five years of mortality data of impala Aepyceros melampus, blue wildebeest Con- nochaetes taurinus, buffalo Syncerus caffer and kudu Tragelaphus strepsiceros have been used to describe the minimum mortality profiles of the respective species in an open savanna system with the full compliment of predators. Predation is the principle cause of natural mortalities on the study site, Letaba Ranch, which is adjacent to the Kruger National Park. The principle cause of impala mortality are leopards Panthera pardus and wild dogs Lycaon pictus. Lion Panthera leo were major contributors to the mortality of wildebeest and buffalo. Anthrax Bacillus anthracis was the main cause of kudu mortality. Key words: predation, predators, impala, blue wildebeest, buffalo, kudu, Letaba Ranch. H.P. Cronje and B.K. Reilly, Department of Nature Conservation, Technikon Preto- ria,Private Bag X680, Pretoria. 0001; I.D. Macfadyen, 214 Leonie Street, Doringkloof, Pretoria 0001. ISSN 0075-6458 79 Koedoe 45/1 (2002) Introduction Predation constitutes an important feature of the biotic environments of wild ungulates (Hirst 1969) and the effect of this on small private nature reserves can lead to financial implications. The increased perception of the benefits of tourism especially if the major predators are accommodated, has led to sev- eral private undertakings applying for per- mits to accommodate lion in particular. The availability of mortality data, expressed as a financial benefit or loss relative to tourism benefits, is an important factor in determin- ing the sustainability of undertakings. Sever- al predator-prey studies have been done to describe the impact of predators on wild ungulate species (Mitchell et al. 1965; Hirst 1969; Pienaar 1969; Mills & Skenk 1992), but they were one-off studies for short peri- ods of time. The effective management of any wildlife undertaking is dependent on the constant availability of information. Mortali- ty registers updated on a weekly and annual basis is a valuable source of information par- ticularly during periods in which game cen- suses are absent. This data is a means to monitor annual mortalities thus effecting the adjustments of ungulate trends over time. The mortality profiles of impala, blue wilde- beest, buffalo and kudu are described. Data was collected and collated from field ranger observations for the period January 1990 to December 1994. Study area The Letaba Ranch Nature Reserve is situated at 23º39'S and 31º03'E and 100 km east of Tzaneen in the Limpopo Province, South Africa. The Kruger National Park borders the eastern boundary of Letaba Ranch. The 35 000 ha reserve is now totally enclosed by an electrified fence which was not the case at the time of the study. The topography of the reserve is predomi- nantly gentle undulating landscapes at alti- tudes between 340–400 m a.s.l. with the cronje&reilley.qxd 2005/12/09 11:29 Page 79 occasional hill reaching 450 m above sea level. Letaba Ranch is located in a summer rainfall region characterised by hot summers and temperate winters, with the highest and low- est daily mean maximum and minimum tem- peratures of 31.7 ºC in the summer and 10.1 ºC in the winter. The mean annual rain- fall is 487 mm. Gertenbach (1987), in his study of the vege- tation of the Kruger National Park, distin- guishes two landscapes also common to Letaba Ranch, namely Colophospermum mopane veld on sandy soils extending over the southern two-thirds of the reserve and the Letaba River rugged veld in the northern third of the reserve. The larger herbivore component common on Letaba Ranch are impala, blue wildebeest, buffalo, bushbuck, elephant Loxodonta africana, zebra Equus burchelli, giraffe Giraffa camelopardalis, kudu, sable Hippo- tragus niger, waterbuck Kobus ellipsiprym- nus, warthog Phacochoerus africanus and white rhinoceros Ceratotherium simum. Lion, leopard Panthera pardus, spotted hyaena Crocuta crocuta and black-backed jackal Canis mesomelas are the most numer- ous ungulate predators. Cheetah Acinionyx jubatus, wild dog and side-striped jackal Canis adustus are present, but not common. In terms of resource utilisation, Letaba Ranch functioned as a trophy hunting reserve until 1994 (Macfadyen pers. comm.). The rugged and rough terrain, poor roads and scrub mopane all contributed to this utilisa- tion option which had financial benefits higher than that of other eco-tourism ven- tures (Macfadyen pers. comm.). Methods To collect a large data set on the game mortalities occurring on Letaba Ranch, the rangers who patrolled the reserve on a daily basis were tasked with noting mortality data. Hirst (1969), Pienaar (1969) and Mills (1984) also used this data collec- tion technique. To standardise the classification of age classes by the rangers, examples of the various ages of the ungulates under investigation were demonstrated to the rangers. The age classes of kill data recorded were adult male (ADM), adult female (ADF), sub-adult male (SAM), sub-adult female (SAF), and juvenile (J). The cause of death was determined by the evidence of characteristic predator kill signs (that is the method by which the animals were killed and eaten), dominant tracks at the site of the kill or the visual confirmation of the predator at a carcass. In the event that the cause of death could not be determined, cause of death was recorded as unknown. The miscellaneous (M) category encom- passed infrequently recorded mortalities due to croc- odile and caracal. Natural mortalities refer to mortal- ities that were caused by predators or due to intraspe- cific interactions of a particular species. Although malnutrition is the direct cause of mortality during drought periods, the category drought was chosen to represent malnutrition. The main source of bias in the technique used to col- lect mortality data is fortuitous observations (Mitchell et al. 1965; Hirst 1969; Pienaar 1969; Mills 1984) in that for the small prey species and younger age classes, the detection of the carcass was negligible due to the rapid rate at which the carcass was devoured. Thus the data is biased to the larger ungulate and age classes. Data collection is also biased towards the dry season due to the increased visibility and detection of carcasses during this period. With cognisance of these limitations, it is believed that the data presented provides an estimate of the mortalities that can be expected on a reserve accommodating the full spectrum of predators. The agents of mortality, relative to each ungulate species, were collectively summed with the impact of each agent of mortality expressed as a proportion of the total cause of mortality. Results and discussion Of the four ungulate species, predation on impala was mostly by leopard and wild dog. Lion were responsible for the greater part of deaths among blue wildebeest, buffalo and kudu (Table 1). Over the spectrum of natural mortalities recorded, in descending order of contribution, lion had the greatest impact (27.5 %) followed by leopard (21.3 %), Anthrax (20.5 %) and wild dog with 11.8 % (Table 2). Koedoe 45/1 (2002) 80 ISSN 0075-6458 cronje&reilley.qxd 2005/12/09 11:29 Page 80 ISSN 0075-6458 81 Koedoe 45/1 (2002) Ta bl e 1 N at ur al m or ta lit y to ta ls o f f ou r un gu la te s pe ci es a nd th e pr op or tio na l m or ta lit y of e ac h ag en t o f m or ta lit y on L et ab a R an ch , L im po po P ro vi nc e Sp ec ie s n A nt hr ax C he et ah Fi gh tin g H ya en a Ja ck al L io n L eo pa rd W ild d og D ro ug ht M is ce lla ne ou s U nk ow n % % % % % % % % % % % A ep yc er os m el am pu s 49 1 2, 2 4, 7 1 8, 3 1 7, 3 41 22 6, 7 1, 2 5, 9 C on no ch ae te s ta ur in us 10 1 - 1 - 4 - 73 ,3 11 2 1 2 5, 9 Sy nc er us c af fe r 16 2 14 ,2 - - 2, 5 - 71 ,6 - - 6, 8 - 5 Tr ag el ap hu s s tr ep si ce ro s 25 0 69 1, 2 0, 8 1, 2 - 20 1, 6 3, 2 - 0, 4 2, 8 Ta bl e 2 C um ul at iv e pr op or tio na l m or ta lit y of e ac h ag en t o f m or ta lit y re la tiv e to th e to ta l nu m be r of m or ta lit ie s re co rd ed , L et ab a R an ch , L im po po P ro vi nc e n A nt hr ax C he et ah Fi gh tin g H ya en a Ja ck al L io n L eo pa rd W ild d og D ro ug ht M is ce lla ne ou s U nk ow n % % % % % % % % % % % 10 04 20 ,5 2, 7 0, 7 5, 2 0, 5 27 ,5 21 ,3 11 ,8 4, 5 0, 9 4, 5 cronje&reilley.qxd 2005/12/09 11:29 Page 81 Impala Impala were vulnerable to all six predators (Table 1). Although all predators prey on impala to a greater or lesser extent (Skinner & Smithers 1990), in this study leopard had the greatest impact (41 %). Hirst (1969), Pienaar (1969) and Le Roux & Skinner (1989) found that leopard are the main predators of impala in the Timbavati (20.7 %), Kruger National Park (34.3 %) and Londolozi Game Reserve (51 %), respec- tively. Wild dog had the second highest impact (22 %) in contrast to the 9.5 % reported by Pienaar (1969) for the Kruger National Park. This may be attributed to the fact that Letaba Ranch is smaller in size and fully fenced, causing predation in the herbi- vore component to be higher. Hyaena (8.3 %) and lion (7.3 %) had a smaller impact on impala. Cheetah (4.7 %) had the lowest impact of all the predators. Cheetah were also reported to be of less importance in the Kruger National Park, where they con- tributed a 5.8 % mortality of all carcasses recorded (Pienaar 1969). However, in areas where cheetah are in high density and where inter-specific competition is minimal, mor- tality of impala by cheetah would be expect- ed to be higher. Cheetah are regularly sub- jected to inter-specific competition (Pienaar 1969; Skinner & Smithers 1990). Miscella- neous mortalities amounted to 1.2 %, occa- sioned by five crocodile kills and one caracal kill. Anthrax was a minor factor, resulting in mean of 2.2 % over the study period, with 9.32 % deaths in 1990, the year of the out- break. The percentage mortality of impala per predator within each age class is shown in Table 3. Adult female impala had the highest mortality relative to other sex and age class- es by lion (47.2 %), leopard (39.7 %) and wild dog (49.1 %). For hyaena, adult male and adult female age classes had the same mortality rates. Cheetah seemed to concen- trate on the male component with a higher mortality rate (43.5 %) within the sub-adult male age class. Cheetah generally prey on stragglers on the fringes of the main impala herds (Skinner & Smithers 1990) which are often of the male gender. Jackals caused the highest mortality (80 %) of juvenile impala. Newborn lambs of the smaller antelope species are prone to the attention of jackals during the lambing season (Pienaar 1969). Blue Wildebeest Blue wildebeest were preyed on by five dif- ferent predators. The main predators were lion (73.3 %) followed by leopard (11 %) and hyaena (4 %) as indicated in Table 1. Pienaar (1969), Hirst (1969) and Mills & Skenk (1992) also found that lion were the major cause of wildebeest mortality. The data in Table 4 shows a high kill ratio for the adult component, with a slightly higher per- centage of females (45.9 %) in relation to the male mortality of 40.5 %, resulting from lion. Leopard prey mostly on young wilde- beest (Bourliére 1963), which is supported Koedoe 45/1 (2002) 82 ISSN 0075-6458 Table 3 Proportional predation mortality of impala Aepyceros melampus relative to age and sex class, Letaba Ranch, Limpopo Province Age-Sex Class N Cheetah(%) Hyaena(%) Jackal(%) Lion(%) Leopard(%) Wilddog(%) Adult Male 143 26,1 41,5 - 30,6 38,2 30,6 Adult Female 171 21,7 41.5 - 47,2 39,7 49,1 Sub-adult Male 58 43,5 9,8 20,0 13,9 12,1 13,0 Sub-adult Female 33 8,7 7,3 - 8,3 9,0 6,5 Juvenile 7 - - 80 - 1,0 0,9 cronje&reilley.qxd 2005/12/09 11:29 Page 82 by the results of this study (Table 4). Juve- niles had the highest mortality rate (63.6 %), followed by the sub-adult males (27.3 %) and with the lowest, the sub-adult females (9.5 %). Kills by wild dog, hyaena and chee- tah were low at 2 %, 4 % and 1 % respec- tively. Buffalo As with blue wildebeest, lion predation is also the principle cause of buffalo mortality (71,6 %), which is borne out by the findings of Pienaar (1969) and Mitchell et al. (1965) in the Kruger and Kafue national parks respectively. According to Bourliere (1963) adult buffalo regularly fall prey to lion. The adult female and sub-adult male buffalo age classes had the highest representation in lion kills viz. 35.4 % and 25.4 %, respectively (Table 5). Hyaena had a minimal (2.5 %) effect on the mortality of buffalo (Table 1). Adult buffalo may be too large a prey to be captured by hyaena, especially if hunted singly (Cooper 1990). Hyaena mostly select medium-sized prey but being opportunistic, would take almost anything they can over- power (Cooper 1990). This is supported by the data in Table 5 where it is noted that hyaena tend to concentrate on the younger buffalo component where the opportunity arises. Over the study period, anthrax was responsible for 14.2 % of all deaths. During 1990, out of a total of 31 mortalities record- ed, anthrax was responsible for 23 (74.2 %) of those mortalities. Buffalo are infected with anthrax by grazing on vegetation onto which blowflies have regurgitated after feed- ing on a carcass contaminated with anthrax as well as being infected with anthrax at waterholes (Bothma 1996). The principle source of anthrax infection in buffalo in the Kruger National Park is waterholes (De Vos pers. comm.) and is also assumed to be the case on the Letaba Ranch Game Reserve. In 5 % of the sample, the cause of death was ISSN 0075-6458 83 Koedoe 45/1 (2002) Table 4 Proportional predation mortality of blue wildebeest Connocheatus taurinus relative to each age and sex class, Letaba Ranch, Limpopo Province Age-Sex Class N Cheetah(%) Hyaena(%) Jackal(%) Lion(%) Leopard(%) Wild dog(%) Adult Male 32 - 50,0 - 40,5 - - Adult Female 36 - 50,0 - 45,9 - - Sub-adult Male 4 - 0 - 1,4 27,3 - Sub-adult Female 8 - - - 9,5 9,1 - Juvenile 12 100 - - 2,7 63,6 100 Table 5 Proportional predation mortality of buffalo Syncerus caffer relative to each age and sex class, Letaba Ranch, Limpopo Province Age-Sex Class N Cheetah(%) Hyaena(%) Jackal(%) Lion(%) Leopard(%) Wilddog(%) Adult Male 16 - - - 13,8 - - Adult Female 41 - - - 35,3 - - Sub-adult Male 30 - - - 25,9 - - Sub-adult Female 14 - - - 12,1 - - Juvenile 19 - 100 - 12,9 - - cronje&reilley.qxd 2005/12/09 11:29 Page 83 unknown. Two calving-related deaths (1.2 %) were recorded. Kudu Kudu were subject to high (20 %) lion- induced mortality as corroborated by Hirst (1969), with wild dog second highest (3.2 %) (Table 1). In the northern and central districts of the Kruger National Park, Pienaar (1969) found that, amongst others, kudu was one of the species heavily preyed upon by wild dog. During the study, both male and female adult animals had equal mortalities (Table 6). Leopard, hyaena and cheetah were the lowest contributors to kudu mortality. Leopard and cheetah were responsible for the mortality within younger age groups (sub-adult and juvenile) as was also noted by Pienaar (1969) in the Kruger National Park. Hyaena preyed on both adult and sub-adult females (Table 6). Anthrax caused the great- est mortality (69 %) of kudu during the study period. In 1990, of 204 deaths recorded, 171 (84 %) were anthrax related. Of all ungu- lates, kudu are particularly susceptible to anthrax (Dekker pers. comm.). Kudu are infected in a similar fashion to buffalo (Pien- aar 1961; Braack & De Vos 1990). One kudu was taken by a crocodile (0.7 %). Annual and seasonal mortality variation The mean annual natural mortalities for impala, blue wildebeest, buffalo and kudu are given in Table 8. Of particular note, are those of buffalo (31 ± 37.37) and kudu (50 ± 75.38). The cause of the wide data dispersion for kudu was due to the anthrax outbreak during 1990. That of the buffalo is attributed to excessively high lion predation during 1992. The factors contributing to this high mortality were the stress of buffalo after a capture operation, which had an adverse affect on the herd structure, as well as the occurrence of a drought during 1992 (Mac- fadyen pers. comm.). Re-computation of the dispersion of the data in the absence of the outliers (trimmed mean) show the mortalities for buffalo and kudu at the 95% confidence Koedoe 45/1 (2002) 84 ISSN 0075-6458 Table 6 Proportional predation mortality of kudu Tragelaphus strepsiceros relative to each age and sex class Letaba Ranch, Limpopo Province Age-Sex Class n Cheetah(%) Hyaena(%) Jackal(%) Lion(%) Leopard(%) Wild dog(%) Adult Male 22 - - - 40,0 - 25,0 Adult Female 24 - 66,7 - 40,0 25,0 Sub-adult Male 6 - - - 12,0 - - Sub-adult Female 10 33,3 33,3 - 8,0 75,0 12,5 Juvenile 6 66,7 - - - 25,0 37,5 Table 7 The seasonal distribution of natural mortalities on Letaba Ranch, Limpopo Province Species n Wet season Dry season Jan - Mar/Oct - Dec (%) Mar - Sept (%) Aepyceros melampus 491 37,5 62,5 Connocheatus taurinus 101 47,5 52,5 Syncerus caffer 162 42 58 Tragelaphus strepsiceros 250 11,6 88,4 cronje&reilley.qxd 2005/12/09 11:29 Page 84 limit to be 15 (± 9.92) and 15.8 (± 8.6) respectively. The majority of all deaths recorded occurred during the dry season for all four ungulates (Table 7). Conclusion The juvenile components of all four species are underrepresented in the data in this study, attributed to the rapid rate at which these ani- mals are disposed by predators (Mitchell et al.1965; Hirst 1969). Migratory animals have a lower mortality in contrast to sedentary animals in national parks that are large enough to allow for the natural migrations of gregarious ungulates (Mills & Skenk 1992). From this can be con- cluded that migratory species in restricted areas will be prone to greater mortality, as is the case of Letaba Ranch. In restricted areas where predators (especially lion), are accom- modated and the smaller the area, the greater the number of mortalities that can be expect- ed. Anthrax has the potential to adversely effect ungulate species to a greater or lesser extent and must be taken into account. The capture and removal of game is a man- agement action commonly used to regulate ungulate populations and is a prime income generator. On areas accommodating preda- tors, particularly lions, cognizance must be taken of the potential post capture increase in predation which may lead to further losses. The data in this study represents the mini- mum number of mortalities for impala, blue wildebeest, buffalo and kudu. It is impossi- ble to record the exact number of mortalities occurring in areas where predators are active due to differing hunting techniques, rates at which prey is consumed, remains left and failure of managers to detect carcasses par- ticularly during the wet season when visibil- ity is impaired. Mortality data should be used in the devel- opment of population models that incorpo- rate game count data to derive more accurate data on population trends. Annual game cen- suses are becoming more irregular due to financial constraints. A population model based on population trends (from long-term species monitoring) and incorporating annu- al mortality data (natural and unnatural) pro- viding estimates of populations during inter- vening periods, should be a research priority. Models of this nature would ensure continu- ous data regarding ungulate trends. Count data combined with population dynamics data provide better decision support than count driven trends alone. Predator prey models also require accurate data on mortal- ities, particularly where predators are to be reintroduced to conservation areas in the sub-region. Very often wildlife managers look to expen- sive and sophisticated procedures to gather data on game populations. This analysis from daily summaries in a managers diary, from good ground coverage by field rangers, show that “primitive” methods are invalu- able in gaining insight into the status of species in protected areas. In the light of ISSN 0075-6458 85 Koedoe 45/1 (2002) Table 8 Annual variation in the natural mortalities of four ungulate species on Letaba Ranch, Limpopo Province Species n Mean anuual mortality Aepyceros melampus 491 98,2 (± 44,12) Connocheatus taurinus 101 20,2 (± 5,2) Syncerus caffer 162 31 (± 37,37) Tragelaphus strepsiceros 250 50,2 (± 75,38) cronje&reilley.qxd 2005/12/09 11:29 Page 85 increasing costs of monitoring, declining budgets and expertise in conservation agen- cies, such inexpensive opportunistic data gathering must be considered in wildlife management undertakings in southern Africa. The effectiveness of this approach is, however, dependent on the number of field staff employed for this purpose relative to the surface area that will require regular sur- veillance. Ascertaining the competence of field staff in accurately recording mortality and training them where necessary is of paramount importance for the collection of mortality data. Acknowledgements The first author would like to thank Ionka Du Preez for her contiuous support and assitance throughout the preparation of this manuscript. Gratitiude is also expressed to the referees that provided valuble input and added value to the paper. References BOURLIÉRE, F. 1963. Specific feeding habits of African carnivores. African Wildlife 17(1): 21–27. BRAAK, L.E.O. & V. DE VOS. 1990. Feeding habits and flight range of blow-flies (Chrysomyia spp) in relation to anthrax transmission in the Kruger National Park, South Africa. Onderstepoort Journal of Veterinary Research 57: 141–142. CHOQUETTE, L.P.E. & E. BROUGHTON. 1981. Anthrax Pp. 289–295. In: DAVIES, J.W., L.H. KARSTAD & D.O. TRAINER (ed). Infectious diseases of wild mammals. Ames, Iowa: Iowa State University Press. COOPER, M.S. 1990. The hunting behaviour of spot- ted hyaenas (Crocuta crocuta) in a region con- taining both sedentary and migratory popula- tions of herbivores. African Journal of Ecology 28: 131–141. GERTENBACH, W.P.D. 1987. 'n Ekologiese studie van die suidelikste mopanieveld in die Nationale Krugerwildtuin. D.Sc. verhandeling, Univer- siteit van Pretoria, Pretoria. HIRST, S.M. 1969. Populations in a Transvaal Lowveld Nature Reserve. Zoologica Africana 4(2): 199–230. LE ROUX, P.G. & J.D. SKINNER. 1989. A note on the ecology of the leopard (Panthera pardus Lin- naeus) in the Londolozi Game Reserve, South Africa. African Journal of Ecology 27: 167–171. MILLS, M.G.L. 1984. Prey selection and feeding habits of the large carnivores in the southern Kalahari. Koedoe (Suppl.): 281–296. MITCHELL, B.L., J.B. SHENTON & J.C.M. UYS. 1965. Predation on large mammals in the Kafue National Park, Zambia. Zoologica Africana 1: 297–318. MILLS, M.G.L. & T.M. SKENK. 1992. Predator-prey relationships: the impact of lion predation on wildebeest and zebra populations. Journal of Animal Ecology 61: 693–702. PIENAAR, U. DE V. 1961. The outbreak of anthrax amongst game animals in the Kruger National Park. Koedoe 4: 4–17. PIENAAR, U. DE V. 1969. Predator-prey relationships amongst the larger mammals of the Kruger National Park. Koedoe 12: 108–187. SKINNER, J.D. & R.H.N. SMITHERS. 1990. The mam- mals of the southern African subregion. Pretoria: University of Pretoria. 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