Do fences constrain predator movements on an evolutionary scale? Home range, food intake and movement patterns of large predators reintroduced to Addo Elephant National Park, South Africa

  • Matt W. Hayward
  • Gina J. Hayward
  • Dave J. Druce
  • Graham I. H. Kerley
Original Paper


Fencing conservation areas is ubiquitous in South Africa, however, the impact of these on predator ecology has not been tested. We used relationships between prey abundance and predator space use to create equations to predict the home range size of lions Panthera leo and leopards Panthera pardus. We then successfully tested these predictions using published data (Phinda, Makalali) and home range estimates from radio collared individuals reintroduced to Addo Elephant National Park. Spotted hyaena Crocuta crocuta ranges also seem food dependent. Lion home ranges in Addo (114 ± 5 km2) required 180 fixes to be accurately estimated, spotted hyaena ranges (91 ± 10 km2) required 200 fixes, and the solitary leopard had 295 fixes for a range of 38 km2. There were no sexual differences in home range sizes of lions or hyaenas. The daily food intake rate of lions, measured during continuous follows, was 9.8 kg per female equivalent unit. Dominant male lions (14.3 km for 8.3 kg) traveled furthest but obtained the least amount of food per day compared to subordinate males (8.9 km for 16.0 kg) and females (5.8 km for 17.9 kg). Subordinate males traveled the fastest and during the day, to avoid competition and harassment from the dominant males. From an evolutionary viewpoint, the use of fences for conservation has not affected the natural behaviour of the predators as they still conform to predictions derived from unfenced reserves; that is, prey abundance is the key factor in determining space use of large predators.


Behavioural ecology Competition avoidance GPS/GSM radio collars Home range Food intake Intra-specific competition Optimal foraging Reintroduction Resource dispersion hypothesis 



This project was supported by Budget Rent-a-Car, Eveready Batteries, CR Kennedy Pty Ltd, Leica, Continental Tyres, Wayne Linklater and the Centre for African Conservation Ecology at the Nelson Mandela Metropolitan University, and was approved by the South African National Parks service (Project No. 2004-03-01GKER). MWH was assisted in the field by Ivan Philipson, Katie Gough, Heidi Grolsh and Michael Somers, and was supported by a Nelson Mandela Metropolitan University and then a National Research Foundation post doctoral fellowship. The write up of this research project has been supported by a Marie Curie Transfer of Knowledge Fellowship BIORESC of European Community’s Sixth Framework Programme under contract number MTKD-CT-2005-029957. Larry Ward and Phil Hayward of Lean & Hayward Pty Ltd assisted by converting latitude and longitude to UTM coordinates.


  1. Bailey TN (1993) The African leopard: ecology and behavior of a solitary field. Columbia University Press, New YorkGoogle Scholar
  2. Bauer H, De Iongh HH (2005) Lion (Panthera leo) home ranges and livestock conflicts in Waza National Park, Cameroon. Afr J Ecol 43:208–214. doi: 10.1111/j.1365-2028.2005.00570.x CrossRefGoogle Scholar
  3. Bauer H, Van der Merwe S (2004) Inventory of free-ranging lions Panthera leo in Africa. Oryx 38:26–31. doi: 10.1017/S0030605304000055 CrossRefGoogle Scholar
  4. Berger KM, Gese EM (2007) Does interference competition with wolves limit the distribution and abundance of coyotes? J Anim Ecol 76:1075–1085. doi: 10.1111/j.1365-2656.2007.01287.x PubMedCrossRefGoogle Scholar
  5. Berger J, Swenson JE, Persson I-L (2001) Recolonizing carnivores and naive prey: conservation lessons from Pleistocene extinctions. Science 291:1036–1039. doi: 10.1126/science.1056466 PubMedCrossRefGoogle Scholar
  6. Bertram BCB (1975) Weights and measures of lions. Af Wildl J 13:141–143Google Scholar
  7. Blumenschine RJ, Caro TM (1986) Unit flesh weights of some East African bovids. Afr J Ecol 24:273–286. doi: 10.1111/j.1365-2028.1986.tb00371.x CrossRefGoogle Scholar
  8. Bothma JdP, le Riche EAN (1984) Aspects of the ecology and the behaviour of the leopard Panthera pardus in the Kalahari Desert. Koedoe Suppl 84:259–279Google Scholar
  9. Boydston EE, kapheim KM, Szykman M et al (2003) Individual variation in space use by female spotted hyenas. J Mammal 84:1006–1018. doi: 10.1644/BOS-038 CrossRefGoogle Scholar
  10. Bradfield M (2005) Addo Elephant National Park—official guide. South African National Parks, Port ElizabethGoogle Scholar
  11. Carbone C, Gittleman JL (2002) A common rule for the scaling of carnivore density. Science 295:2273–2276. doi: 10.1126/science.1067994 PubMedCrossRefGoogle Scholar
  12. Carbone C, Mace GM, Roberts SC et al (1999) Energetic constraints on the diet of terrestrial carnivores. Nature 402:286–288. doi: 10.1038/46266 PubMedCrossRefGoogle Scholar
  13. Creel S, Creel NM (1996) Limitation of African wild dogs by competition with larger carnivores. Conserv Biol 10:526–538. doi: 10.1046/j.1523-1739.1996.10020526.x CrossRefGoogle Scholar
  14. de Solla SR, Bonduriansky R, Brooks RJ (1999) Eliminating autocorrelation reduces biological relevance of home range estimates. J Anim Ecol 68:221–234. doi: 10.1046/j.1365-2656.1999.00279.x CrossRefGoogle Scholar
  15. Druce D, Genis H, Greatwood S et al (2004) Population demography and spatial ecology of a reintroduced lion population in the Greater Makalali Conservancy, South Africa. Koedoe 47:103–118Google Scholar
  16. Elliott JP, Cowan IM (1978) Territoriality, density, and prey of the lion in Ngorongoro Crater, Tanzania. Can J Zool 56:1726–1734CrossRefGoogle Scholar
  17. Eloff FC (1973) Lion predation in the Kalahari Gemsbok National Park. J SA Wildl Manage Assoc 3:59–63Google Scholar
  18. Funston PJ (2001) Kalahari Transfrontier Lion Project, ed. South African National Parks, Botswana Department of Wildlife and National Parks, Endangered Wildlife Trust, Carnivore Conservation Group, WWF. The Green Trust, PretoriaGoogle Scholar
  19. Funston PJ, Mills MGL, Biggs HC et al (1998) Hunting by male lions: ecological influences and socioecological implications. Anim Behav 56:1333–1345. doi: 10.1006/anbe.1998.0884 PubMedCrossRefGoogle Scholar
  20. Funston PJ, Mills MGL, Richardson PRK et al (2003) Reduced dispersal and opportunistic territory acquisition in male lions (Panthera leo). J Zool (Lond) 259:131–142. doi: 10.1017/S0952836902003126 CrossRefGoogle Scholar
  21. Gittleman JL, Harvey PH (1982) Carnivore home-range size, metabolic needs and ecology. Behav Ecol Sociobiol 10:57–63. doi: 10.1007/BF00296396 CrossRefGoogle Scholar
  22. Green B, Anderson J, Whateley A (1984) Water and sodium turnover and estimated food consumption in free-living lions (Panthera leo) and spotted hyaenas (Crocuta crocuta). J Mammal 65:593–599. doi: 10.2307/1380842 CrossRefGoogle Scholar
  23. Hayward MW (2006) Prey preferences of the spotted hyaena Crocuta crocuta and evidence of dietary competition with lion Panthera leo. J Zool 270:606–614. doi: 610.1111/j.1469-7998.2006.00184.x CrossRefGoogle Scholar
  24. Hayward MW, Hayward GJ (2007) Activity patterns of reintroduced lion Panthera leo and spotted hyaena Crocuta crocuta in Addo Elephant National Park, South Africa. Afr J Ecol 45:135–141. doi: 10.1111/j.1365-2028.2006.00686.x CrossRefGoogle Scholar
  25. Hayward MW, Kerley GIH (2005) Prey preferences of the lion (Panthera leo). J Zool (Lond) 267:309–322. doi: 10.1017/S0952836905007508 CrossRefGoogle Scholar
  26. Hayward MW, de Tores PJ, Augee ML et al (2004) Home range and movements of the quokka Setonix brachyurus (Macropodidae: Marsupialia), and its impact on the viability of the metapopulation on the Australian mainland. J Zool (Lond) 263:219–228. doi: 10.1017/S0952836904005060 CrossRefGoogle Scholar
  27. Hayward MW, Henschel P, O’Brien J et al (2006) Prey preferences of the leopard (Panthera pardus). J Zool (Lond) 270:298–313Google Scholar
  28. Hayward MW, Adendorff J, Moolman LC et al (2007a) The successful reintroduction of leopard Panthera pardus to the Addo Elephant National Park. Afr J Ecol 45:103–104. doi: 10.1111/j.1365-2028.2006.00673.x CrossRefGoogle Scholar
  29. Hayward MW, Adendorff J, O’Brien J et al (2007b) Practical considerations for the reintroduction of large, terrestrial, mammalian predators based on reintroductions to South Africa’s Eastern Cape Province. Open Conserv Biol J 1:1–11. doi  10.2174/1874-8392/2107
  30. Hayward MW, Adendorff J, O’Brien J et al (2007c) The reintroduction of large carnivores to the Eastern Cape Province, South Africa: an assessment. Oryx 41:205–214. doi: 10.1017/S0030605307001767 CrossRefGoogle Scholar
  31. Hayward MW, Hofmeyr M, O’Brien J et al (2007d) Testing predictions of the prey of the lion (Panthera leo) derived from modelled prey preferences. J Wildl Manage 71:1567–1575. doi: 10.2193/2006-264 CrossRefGoogle Scholar
  32. Hayward MW, O’Brien J, Kerley GIH (2007e) Carrying capacity of large African predators: predictions and tests. Biol Conserv 139:219–229. doi: 210.1016/j.biocon.2007.1006.1018 CrossRefGoogle Scholar
  33. Hemson G (2003) The ecology and conservation of lions: human-wildlife conflict in semi-arid Botswana. PhD Dissertation, University of OxfordGoogle Scholar
  34. Hofer H, East ML (1993) The commuting system of Serengeti spotted hyaenas: how a predator copes with migratory prey. III. Attendance and maternal care. Anim Behav 46:575–589. doi: 10.1006/anbe.1993.1224 CrossRefGoogle Scholar
  35. Hunter LTB (1998) The behavioural ecology of reintroduced lions and cheetahs in the Phinda Resource Reserve, kwaZulu-Natal, South Africa. PhD Thesis, University of PretoriaGoogle Scholar
  36. Hunter LTB, Skinner JD (1998) Vigilance behaviour in African ungulates: the role of predation pressure. Behaviour 135:195–211Google Scholar
  37. Jetz W, Carbone C, Fulford J et al (2004) The scaling of animal space use. Science 306:266–268. doi: 10.1126/science.1102138 PubMedCrossRefGoogle Scholar
  38. Kenward RE, Hodder KH (1992) Ranges V—an analysis system for biological location data. Institute of Terrestrial Ecology, Furzebrook Research Station, WarehamGoogle Scholar
  39. le Roux PG, Skinner JD (1989) A note on the ecology of the leopard (Panthera pardus Linnaeus) in the Londolozi Game Reserve, South Africa. Afr J Ecol 27:167–171. doi: 10.1111/j.1365-2028.1989.tb00940.x CrossRefGoogle Scholar
  40. Macdonald DW (1983) The ecology of carnivore social behaviour. Nature 301:379–384. doi: 10.1038/301379a0 CrossRefGoogle Scholar
  41. Makacha S, Schaller GB (1969) Observations on lions in the Lake Manyara National Park, Tanzania. Af Wildl J 7:99–103Google Scholar
  42. Marker L, Dickman AJ (2005) Factors affecting leopard (Panthera pardus) spatial ecology, with particular reference to Namibian farmlands. S Afr J Wildl Res 35:105–115Google Scholar
  43. McDougal C (1988) Leopard and tiger interactions at Royal Chitwan National Park, Nepal. J Bombay Nat Hist Soc 85:609–610Google Scholar
  44. Mills MGL (1992) A comparison of methods used to study food habits of large African carnivores. In: McCulloch C, Barret RH (eds) Wildlife 2001: Populations. Elsevier, LondonGoogle Scholar
  45. Mills MGL, Gorman ML (1997) Factors affecting the density and distribution of wild dogs in the Kruger National Park. Conserv Biol 11:1397–1406. doi: 10.1046/j.1523-1739.1997.96252.x CrossRefGoogle Scholar
  46. Mizutani F, Jewell PA (1998) Home-range and movements of leopards (Panthera pardus) on a livestock ranch in Kenya. J Zool (Lond) 244:269–286. doi: 10.1111/j.1469-7998.1998.tb00031.x CrossRefGoogle Scholar
  47. Nilsen EB, Linnell JDC (2006) Intra-specific variation and taxa-sampling affects the home range–body mass relationship. Acta Theriol (Warsz) 51:225–232Google Scholar
  48. Ogutu JO, Dublin HT (2004) Spatial dynamics of lions and their prey along an environmental gradient. Afr J Ecol 42:8–42. doi: 10.1111/j.0141-6707.2004.00440.x CrossRefGoogle Scholar
  49. Packer C, Pusey AE (1982) Cooperation and competition within coalitions of males lions: kin selection or game theory? Nature 296:740–742. doi: 10.1038/296740a0 CrossRefGoogle Scholar
  50. Packer C, Scheel D, Pusey AE (1990) Why lions form groups: food is not enough. Am Nat 136:1–19. doi: 10.1086/285079 CrossRefGoogle Scholar
  51. Packer C, Hilborn R, Mosser A et al (2005) Ecological change, group territoriality, and population dynamics in Serengeti lions. Science 307:390–392. doi: 10.1126/science.1105122 PubMedCrossRefGoogle Scholar
  52. Radloff FGT, du Toit JT (2004) Large predators and their prey in a southern African savanna: a predator’s size determines its prey size range. J Anim Ecol 73:410–423. doi: 10.1111/j.0021-8790.2004.00817.x CrossRefGoogle Scholar
  53. Rahimi S, Owen-Smith N (2007) Movement patterns of sable antelope in the Kruger National Park from GPS/GSM collars: a preliminary assessment. S Afr J Wildl Res 37:143–151CrossRefGoogle Scholar
  54. Rapson JA, Bernard RTF (2007) Interpreting the diet of lions (Panthera leo); a comparison of various methods of analysis. S Afr J Wildl Res 37:179–187CrossRefGoogle Scholar
  55. Rudnai J (1973) Reproductive biology of lions (Panthera leo massaica Neumann) in Nairobi National Park. Af Wildl J 11:241–253Google Scholar
  56. Rudnai J (1974) The pattern of lion predation in Nairobi Park. Af Wildl J 12:213–225Google Scholar
  57. Schaller GB (1972) The Serengeti lion. University of Chicago Press, ChicagoGoogle Scholar
  58. Smuts GL (1976) Population characteristics and recent history of lions in two parts of the Kruger National Park. Koedoe 19:153–164Google Scholar
  59. Smuts GL (1979) Diet of lions and spotted hyaenas assessed from stomach contents. S Afr J Wildl Res 9:19–25Google Scholar
  60. Spong G (2002) Space use in lions, Panthera leo, in the Selous Game Reserve: social and ecological factors. Behav Ecol Sociobiol 52:303–307. doi: 10.1007/s00265-002-0515-x CrossRefGoogle Scholar
  61. Stander PE (1991) Demography of lions in the Etosha National Park, Namibia. Madoqua 18:1–9Google Scholar
  62. Stander PE, Haden PJ, Kaqece et al (1997) The ecology of asociality in Namibian leopards. J Zool (Lond) 242:343–364Google Scholar
  63. Stuart CT, Stuart T (2000) Field guide to the larger mammals of Africa. Struik Publishers, Cape TownGoogle Scholar
  64. Swihart RK, Slade NA (1985a) Influence of sampling intervals on estimates of home-range size. J Wildl Manage 49:1019–1025. doi: 10.2307/3801388 CrossRefGoogle Scholar
  65. Swihart RK, Slade NA (1985b) Testing for independence of observations in animal movements. Ecology 66:1176–1184. doi: 10.2307/1939170 CrossRefGoogle Scholar
  66. Taru M, Kanda T, Sunobe T (2002) Alternative mating tactics of the gobiid fish, Bathygobius fuscus. J Ethol 20:9–12. doi: 10.1007/s10164-002-0047-x CrossRefGoogle Scholar
  67. Tilson RL, Henschel JR (1986) Spatial arrangement of spotted hyaena groups in a desert environment, Namibia. Afr J Ecol 24:173–180. doi: 10.1111/j.1365-2028.1986.tb00358.x CrossRefGoogle Scholar
  68. van Dyk G, Slotow R (2003) The effects of fences and lions on the ecology of African wild dogs reintroduced to Pilanesberg National Park, South Africa. Afr Zool 38:79–94Google Scholar
  69. van Orsdol KG (1982) Ranges and food habits of lions in Rwenzori National Park, Uganda. Symp Zool Soc Lond 49:325–340Google Scholar
  70. van Orsdol KG (1984) Foraging behaviour and hunting success of lions in Queen Elizabeth National Park, Uganda. Afr J Ecol 22:79–99. doi: 10.1111/j.1365-2028.1984.tb00682.x CrossRefGoogle Scholar
  71. van Orsdol KG, Hanby JP, Bygott JD (1985) Ecological correlates of lion social organisation (Panthera leo). J Zool 206:97–112Google Scholar
  72. Viljoen PC (1993) The effects of changes in prey availability on lion predation in a large natural ecosystem in northern Botswana. Symp Zool Soc Lond 65:193–213Google Scholar
  73. Vlok JHJ, Euston-Brown DIW, Cowling RM (2003) Acocks’ Valley Bushveld 50 years on: new perspectives on the delimitation, characterisation and origin of subtropical thicket vegetation. S Afr J Bot 69:27–51Google Scholar
  74. West PM, Packer C (2002) Sexual selection, temperature, and the lion’s mane. Science 297:1339–1343. doi: 10.1126/science.1073257 PubMedCrossRefGoogle Scholar
  75. Woodroffe R, Ginsberg JR (2000) Ranging behaviour and vulnerability to extinction in carnivores. In: Gosling M, Sutherland WJ (eds) Behaviour and conservation. Cambridge University Press, CambridgeGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2008

Authors and Affiliations

  • Matt W. Hayward
    • 1
    • 2
    • 3
  • Gina J. Hayward
    • 2
  • Dave J. Druce
    • 2
    • 4
  • Graham I. H. Kerley
    • 2
  1. 1.Marie Curie Research Fellow, Mammal Research Institute (EU Centre of Excellence)Polish Academy of ScienceBialowiezaPoland
  2. 2.Centre for African Conservation EcologyNelson Mandela Metropolitan UniversityPort ElizabethSouth Africa
  3. 3.School of Biological, Earth and Environmental ScienceUniversity of New South WalesKensingtonAustralia
  4. 4.Regional EcologistEzemvelo KZN Wildlife, HluhluwekwaZulu NatalSouth Africa

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