Acta Theriologica

, Volume 54, Issue 4, pp 333–343 | Cite as

Heterogeneity in the density of spotted hyaenas in Hluhluwe-iMfolozi Park, South Africa

  • Jan A. Graf
  • Michael J. Somers
  • Micaela Szykman Gunther
  • Rob Slotow


Animal population sizes and trends, as well as their distributions, are essential information to the understanding and conservation of ecosystems. During this study in Hluhluwe-iMfolozi Park, South Africa, a spotted hyaena Crocuta crocuta Erxleben, 1777 (Hyaenidae) population was surveyed by attracting individuals with pre-recorded sounds. The hyaena population (excluding cubs) is substantially larger (321 individuals) than the previous estimate of 200 and this population is the second largest protected population in South Africa. Average hyaena density, at 0.357 individuals/km2, was relatively high compared to other southern African conservation areas, and range from 0 to 1.25 individuals/km2 across sampling stations. For short periods, spatial heterogeneity in density was marked at small and large spatial scales, but decreased when averaged over a longer period. This heterogeneity may be important in promoting the coexistence of other large and mobile carnivores in Hluhluwe-iMfolozi Park by creating potential dynamic competition refugia in space and time. Furthermore, heterogeneity of hyaena density at smaller scales should influence studies investigating the avoidance of hyaenas by competitively inferior carnivores.

Key words

call-ins carnivore Crocuta crocuta playbacks survey 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Acocks J. P. H. 1988. Veld types of South Africa. Memoirs of the Botanical Survey of South Africa 57: 1–146.Google Scholar
  2. Bearder S. K. 1977. Feeding habits of spotted hyaenas in a woodland habitat. East African Wildlife Journal 15: 236–280.Google Scholar
  3. Bertram B. C. R. 1979. Serengeti predators and their social systems. [In: Serengeti: dynamics of an ecosystem. A. R. E. Sinclair and M. Norton-Griffiths, eds]. University of Chicago Press, Chicago: 221–248.Google Scholar
  4. Beschta R. L. 2005. Reduced cottonwood recruitment following extirpation of wolves in Yellowstone’s northern range. Ecology 86: 391–403.CrossRefGoogle Scholar
  5. Bourquin O., Vincent J. and Hitchins P. M. 1971. The vertebrates of the Hluhluwe game reserve-corridor (state land)-Umfolozi game reserve complex. Lammergeyer 14: 1–58.Google Scholar
  6. Bowler M. 1991. The implications of large predator management on commercial ranchland in Zimbabwe. MSc thesis, University of Zimbabwe, Harare: 1–87.Google Scholar
  7. Boydston E. E., Kapheim K. M., Szykman M. and Holekamp K. E. 2003a. Individual variation in space use by female spotted hyenas. Journal of Mammalogy 84: 1006–1018.CrossRefGoogle Scholar
  8. Boydston E. E., Kapheim K. M., Watts H. E., Szykman M. and Holekamp K. E. 2003b. Altered behaviour in spotted hyenas associated with increased human activity. Animal Conservation 6: 207–219.CrossRefGoogle Scholar
  9. Boydston E. E., Kapheim K. M. and Holekamp K. E. 2006. Patterns of den occupation by the spotted hyaena (Crocuta crocuta). African Journal of Ecology 44: 77–86.CrossRefGoogle Scholar
  10. Brooks P. M. and Macdonald I. A. W. 1983. The Hluhluwe-Umfolozi Reserve: An ecological case history. [In Management of large mammals in African conservation areas. R. N. Owen-Smith, ed]. Haum Education Publishers, Pretoria: 1–77.Google Scholar
  11. Carbone C., Frame L., Frame G., Malcolm J., Fanshawe J., FitzGibbon C., Schaller G., Gordon I. J., Rowcliffe J. M. and du Toit J. T. 2005. Feeding success of African wild dogs (Lycaon pictus) in the Serengeti: the effects of group size and kleptoparasitism. Journal of Zoology, London 266: 153–161.CrossRefGoogle Scholar
  12. Cooper S. M. 1989. Clan sizes of spotted hyaenas in the Savuti region of Chobe National Park, Botswana. Botswana Notes and Records 21: 121–133.Google Scholar
  13. Cooper S. M. 1991. Optimal hunting group size: the need for lions to defend their kills against loss to spotted hyaenas. African Journal of Ecology 29: 130–136.CrossRefGoogle Scholar
  14. Creel S. and Creel N. M. 1996. Limitation of African wild dogs by competition with larger carnivores. Conservation Biology 10: 526–538.CrossRefGoogle Scholar
  15. Creel S. and Creel N. M. 2002. The African wild dog: Behavior, Ecology, and Conservation. Princeton University Press, Princeton. 1–341.Google Scholar
  16. Croll D. A., Maron J. L., Estes J. A., Danner E. M. and Byrd G. V. 2005. Introduced predators transform subarctic islands from grassland to tundra. Science 307: 1959–1961.PubMedCrossRefGoogle Scholar
  17. Cromsigt J. P. G. M. 2006. Large herbivores in space: Resource partitioning among savanna grazers in a heterogeneous environment. PhD thesis, University of Groningen, Groningen: 1–146.Google Scholar
  18. Durant S. M. 1998. Competition refuges and coexistence: an example from Serengeti carnivores. Journal of Animal Ecology 67: 370–386.CrossRefGoogle Scholar
  19. Durant S. M. 2000a. Living with the enemy: avoidance of hyenas and lions by cheetahs in the Serengeti. Behavioural Ecology 11:624–6322.CrossRefGoogle Scholar
  20. Durant S. M. 2000b. Predator avoidance, breeding experience and reproductive success in endangered cheetahs, Acinonyx jubatus. Animal Behaviour 60: 121–130.PubMedCrossRefGoogle Scholar
  21. ESRI 2003. ArcMap 8.3. Redlands, California.Google Scholar
  22. Estes R. D. and Goddard J. 1967. Prey selection and hunting behavior of the African wild dog. Journal of Wildlife Management 31: 52–70.CrossRefGoogle Scholar
  23. Fanshawe J. H. and FitzGibbon C. D. 1993. Factors influencing the hunting success of an African wild dog pack. Animal Behaviour 45: 479–490.CrossRefGoogle Scholar
  24. Friedman Y. and Daly D. 2004. The Red Data Book of the Mammals of South Africa: A Conservation Assessment. Endangered Wildlife Trust, CBSG Southern Africa, Conservation Breeding Specialist Group (SSC/IUCN), Johannesburg: 1–722.Google Scholar
  25. Gasaway W. C., Mossestad K. T. and Stander P. E. 1991. Food acquisition by spotted hyaenas in Etosha National Park, Namibia: predation versus scavenging. African Journal of Ecology 29: 64–75.CrossRefGoogle Scholar
  26. Gusset M., Graf J. A. and Somers M. J. 2006. The re-introduction of endangered wild dogs into Hluhluwe-iMfolozi Park, South Africa: an update on the first 25 years. Reintroduction News 25: 31–33.Google Scholar
  27. Gusset M., Maddock A. H., Szykman M., Gunther G. J., Slotow R., Walters M. and Somers M. J. 2008. Conflicting human interests over the re-introduction of endangered wild dogs in South Africa. Biodiversity and Conservation 17: 83–101.CrossRefGoogle Scholar
  28. Hayward M. W. and Kerley G. I. H. 2008. Prey preferences and the conservation status of Africa’s large predators. South African Journal of Wildlife Research 38: 93–108.CrossRefGoogle Scholar
  29. Hayward M. W., O’Brien J., and Kerley G. I. H. 2007. Carrying capacity of large African predators: predictions and tests. Biological Conservation 139: 219–229.CrossRefGoogle Scholar
  30. Hofer H. and East M. L. 1993. The commuting system of Serengeti spotted hyaenas: how a predator copes with migratory prey. I. Social organization. Animal Behaviour 46: 547–557.CrossRefGoogle Scholar
  31. Hofer H. and East M. 1995. Population dynamics, population size, and the commuting system of Serengeti spotted hyenas. [In Serengeti II: dynamics, management, and conservation of an ecosystem. 0A. R. E. Sinclair and P. Arcese, eds]. University of Chicago Press, Chicago: 332–363.Google Scholar
  32. Hofer H. and Mills G. 1998. Population size, threats and conservation status of hyaenas. [In Hyaenas: status survey and conservation action plan. G. Mills and H. Hofer, eds]. IUCN, Gland: 64–79.Google Scholar
  33. Holt R. D. and Polis G. A. 1997. A theoretical framework for intraguild predation. The American Naturalist 149: 745–764.CrossRefGoogle Scholar
  34. Höner O. P., Wachter B., East M. L., Runyoro V. A. and Hofer H. 2005. The effect of prey abundance and foraging tactics on the population dynamics of a social, terrestrial carnivore, the spotted hyena. Oikos 108: 544–554.CrossRefGoogle Scholar
  35. Hunter L. T. B. 1998. The behavioural ecology of reintroduced lions and cheetahs in the Phinda Resource Reserve, KwaZulu-Natal, South Africa. PhD thesis, University of Pretoria, Pretoria. 1–206.Google Scholar
  36. Infield M. 1988. Attitudes of a rural community towards conservation and a local conservation area in Natal, South Africa. Biological Conservation 45: 21–46.CrossRefGoogle Scholar
  37. Kolowski J. M. 2007. Anthropogenic influences on the behavior of large carnivores in the northern Serengeti ecosystem. PhD thesis, Michigan State University, East Lansing. 1–234.Google Scholar
  38. Kolowski J. M. and Holekamp K. E. 2009. Ecological and anthropogenic influences on space use by spotted hyaenas. Journal of Zoology, London 277: 23–36.CrossRefGoogle Scholar
  39. Kruuk H. 1972. The spotted hyena: a study of predation and social behavior. University of Chicago Press, Chicago: 1–335.Google Scholar
  40. Kruuk H. and Turner M. 1967. Comparative notes on predation by lion, leopard, cheetah and wild dog in the Serengeti area, east Africa. Mammalia 31: 1–27.CrossRefGoogle Scholar
  41. Laurenson M. K. 1994. High juvenile mortality in cheetahs (Acinonyx jubatus) and its consequences for maternal care. Journal of Zoology, London 234(3): 387–408.CrossRefGoogle Scholar
  42. Laurenson M. K. 1995. Implications of high offspring mortality for cheetah population dynamics. [In Serengeti II: dynamics, management, and conservation of an ecosystem. A. R. E. Sinclair and P. Arcese, eds]. University of Chicago Press, Chicago: 332–363.Google Scholar
  43. Leigh K. A. 2005. The Ecology and Conservation Biology of the Endangered African Wild Dog (Lycaon pictus), in the Lower Zambezi, Zambia. PhD thesis, University of Sydney, Sydney: 1–220.Google Scholar
  44. Maddock A. 1999. Wild dog demography in Hluhluwe-Umfolozi Park, South Africa. Conservation Biology 13: 412–417.CrossRefGoogle Scholar
  45. Maddox T. M. 2003. The ecology of cheetahs and other large carnivores in a pastoralist-dominated buffer zone. PhD thesis, University College and Institute of Zoology, London: 1–372.Google Scholar
  46. Messier F. 1994. Ungulate population models with predation: a case study with the North American moose. Ecology 75: 478–488.CrossRefGoogle Scholar
  47. Mills M. G. L. 1985. Hyaena survey of Kruger National Park: August–October 1984. IUCN Hyaena Specialist Group Newsletter 2: 15–25.Google Scholar
  48. Mills M. G. L. 1990. Kalahari hyaenas: comparative behavioural ecology of two species. Unwin Hyman, London: 1–304.Google Scholar
  49. Mills M. G. L. and Funston P. 2003. Large carnivores and savanna heterogeneity. In: The Kruger experience: ecology and management of savanna heterogeneity. [J. T. du Toit, K. H. Rogers and H. C. Biggs, eds]. Island Press, Washington: 370–388.Google Scholar
  50. Mills M. G. L. and Gorman M. L. 1997. Factors affecting the density and distribution of wild dogs in the Kruger National Park. Conservation Biology 11: 1397–1406.CrossRefGoogle Scholar
  51. Mills M. G. L., Juritz J. M. and Zucchini W. 2001. Estimating the size of spotted hyaena (Crocuta crocuta) populations through playback recordings allowing for non-response. Animal Conservation 4: 335–343.CrossRefGoogle Scholar
  52. Ogutu J. O. and Dublin H. T. 1998. The response of lions and spotted hyaenas to sound playbacks as a technique for estimating population size. African Journal of Ecology 36: 83–95.CrossRefGoogle Scholar
  53. Ogutu J. O., Bhola N. and Reid R. 2005. The effects of pastoralism and protection on the density and distribution of carnivores and their prey in the Mara ecosystem of Kenya. Journal of Zoology, London 265: 281–293.CrossRefGoogle Scholar
  54. Owen-Smith N. 2004. Functional heterogeneity in resources within landscapes and herbivore population dynamics. Landscape Ecology 19: 761–771.CrossRefGoogle Scholar
  55. Palomares F. and Caro T. M. 1999. Interspecific killing among mammalian carnivores. The American Naturalist 153: 492–508.CrossRefGoogle Scholar
  56. Polis G. A., Myers C. A. and Holt R. D. 1989. The ecology and evolution of intraguild predation: potential competitors that eat each other. Annual Review of Ecology and Systematics 20: 297–330.CrossRefGoogle Scholar
  57. Pringle J. A. 1977. The distribution of mammals in Natal. Part 2. Carnivora. Annuals of the Natal Museum 23: 93–115.Google Scholar
  58. R Development Core Team 2007. R: A Language and Environment for Statistical Computing, version 2.5.0. R Foundation for Statistical Computing, Vienna.Google Scholar
  59. Ripple W. J. and Beschta R. L. 2004. Wolves and the ecology of fear: can predation risk structure ecosystems? Bioscience 54: 755–766.CrossRefGoogle Scholar
  60. Ripple W. J., Larsen E. J., Renkin R. A. and Smith D. W. 2001. Trophic cascades among wolves, elk and aspen on Yellowstone National Park’s northern range. Biological Conservation 102: 227–234.CrossRefGoogle Scholar
  61. Rowe-Rowe D. T. 1992. The carnivores of Natal. Natal Parks Board, Pietermaritzburg: 1–32.Google Scholar
  62. Saleni P., Gusset M., Graf J. A., Szykman M., Walters M. and Somers M. J. 2007. Refuges in time: temporal avoidance of interference competition in endangered wild dogs Lycaon pictus. Canid News 10.2. (available from Scholar
  63. Salnicki J. 2004. The home range area dynamics of spotted hyaenas (Crocuta crocuta) in the woodland habitat of Hwange National Park, Zimbabwe. MSc thesis, Department of Biological Sciences, University of Zimbabwe, Harare: 1–134.Google Scholar
  64. Sillero-Zubiri C. and Gottelli D. 1992. Population ecology of spotted hyaena in an equatorial mountain forest. African Journal of Ecology 30(4): 292–300.CrossRefGoogle Scholar
  65. Skinner J. D., Funston P. J., Van Aarde R. J., Van Dyk G. and Haupt M. A. 1992. Diet of spotted hyaenas in some mesic and arid Southern African game reserves adjoining farmland. South African Journal of Wildlife Research 22: 119–121.Google Scholar
  66. Smith J. E., Kolowski J. M., Graham K. E., Dawes S.E., and Holekamp K. E. 2008. Social and ecological determinants of fission-fusion dynamics in the spotted hyaena. Animal Behaviour 76: 619–636.CrossRefGoogle Scholar
  67. Somers M. J., Graf J. A., Szykman M., Slotow R. and Gusset M. 2008. Dynamics of a small re-introduced population of wild dogs over 25 years: Allee effects and the implications of sociality for endangered species’ recovery. Oecologia 158: 239–247.PubMedCrossRefGoogle Scholar
  68. Statsoft 2003. STATISTICA data analysis software system, version 6.1. Tulsa, Oklahoma, StatSoft Inc.Google Scholar
  69. Terborgh J., Lopez L., Nuńez V. P., Rao M., Shahabuddin G., Orihuela G., Riveros M., Ascanio R., Adler G. H., Lambert T. D. and Balbas L. 2001. Ecological meltdown in predator-free forest fragments. Science 294: 1923–1926.PubMedCrossRefGoogle Scholar
  70. Whateley A. 1980. Spotted hyaena changes clans. Lammergeyer 28: 45.Google Scholar
  71. Whateley A. 1981. Density and home range of spotted hyaenas in Umfolozi Game Reserve, Natal. Lammergeyer 31: 15–20.Google Scholar
  72. Whateley A. and Brooks P. M. 1978. Numbers and movements of spotted hyaenas in Hluhluwe Game Reserve. Lammergeyer 26: 44–52.Google Scholar
  73. Whateley A. and Brooks P. M. 1985. The carnivores of the Hluhluwe and Umfolozi Game Reserves: 1973–1982. Lammergeyer 35: 1–28.Google Scholar
  74. White P. J. and Garrott R. A. 2005. Yellowstones’s ungulates after wolves — expectations, realizations, and predictions. Biological Conservation 125: 141–152.CrossRefGoogle Scholar

Copyright information

© Mammal Research Institute, Bialowieza, Poland 2009

Authors and Affiliations

  • Jan A. Graf
    • 1
  • Michael J. Somers
    • 2
  • Micaela Szykman Gunther
    • 3
    • 4
  • Rob Slotow
    • 1
  1. 1.School of Biological and Conservation SciencesUniversity of KwaZulu-NatalDurbanSouth Africa
  2. 2.Centre for Wildlife Management, Centre for Invasion BiologyUniversity of PretoriaPretoriaSouth Africa
  3. 3.Center for Species Survival, Conservation and Research Center, National Zoological ParkSmithsonian InstitutionFront RoyalUSA
  4. 4.Department of WildlifeHumboldt State UniversityArcataUSA

Personalised recommendations