Behavioral Ecology and Sociobiology

, Volume 63, Issue 10, pp 1483–1494 | Cite as

Does the risk of encountering lions influence African herbivore behaviour at waterholes?

  • Marion ValeixEmail author
  • Hervé Fritz
  • Andrew J. Loveridge
  • Zeke Davidson
  • Jane E. Hunt
  • Felix Murindagomo
  • David W. Macdonald
Original Pper


A central question in the study of predator–prey relationships is to what extent prey behaviour is determined by avoidance of predators. Here, we test whether the long-term risk of encountering lions and the presence of lions in the vicinity influence the behaviour of large African herbivores at waterholes through avoidance of high-risk areas, increases in group size, changes in temporal niche or changes in the time spent in waterhole areas. In Hwange National Park, Zimbabwe, we monitored waterholes to study the behaviour of nine herbivore species under different risks of encountering lions. We radio-collared 26 lions in the study area which provided the opportunity to monitor whether lions were present during observation sessions and to map longer-term seasonal landscapes of risk of encountering lions. Our results show that the preferred prey species for lions (buffalo, kudu and giraffe) avoided risky waterholes. Group size increased as encounter risk increased for only two species (wildebeest and zebra), but this effect was not strong. Interestingly, buffalo avoided the hours of the day which are dangerous when the long-term and short-term risks of encountering lions were high, and all species showed avoidance of waterhole use at night times when lions were in the vicinity. This illustrates well how prey can make temporal adjustments to avoid dangerous periods coinciding with predator hunting. Additionally, many herbivores spent more time accessing water to drink when the long- and short-term risks of encountering lions were high, and they showed longer potential drinking time when the long-term risk of encountering lions was high, suggesting higher levels of vigilance. This study illustrates the diversity of behavioural adjustments to the risk of encountering a predator and how prey respond differently to temporal variations in this risk.


Buffalo Hwange National Park Landscape of risk Predation risk Predator–prey relationships Temporal niche 



The Director General of the Zimbabwe Parks and Wildlife Management Authority is acknowledged for providing the opportunity to carry out this research and for permission to publish this manuscript. We are indebted to CIRAD as herbivore data were collected in the frame of a CIRAD–CNRS project, the HERD project (Hwange Environmental Research Development), funded by the French “Ministère des Affaires Etrangères”, the “Ambassade de France au Zimbabwe”, the CIRAD, the CNRS and the ANR Biodiversité “BioFun project” ANR-05-BDIV-013-01. This work was made possible with grants from The Darwin Initiative for Biodiversity Grant 162/09/015, The Eppley Foundation, Disney Foundation, Marwell Preservation Trust, Regina B. Frankenburg Foundation, Panthera Foundation and the generosity of Joan and Riv Winant. We thank all the people that participated in the fieldwork, particularly Simon Chamaillé-Jammes and the rangers and ecological staff of HNP. We thank Paul Funston and two anonymous reviewers for their fruitful comments on a previous draft of this manuscript. Relevant animal care protocols were followed and approval received from the appropriate agencies.


  1. Barreto GR, Macdonald DW (1999) The response of water voles, Arvicola terrestris, to the odours of predators. Anim Behav 57:1107–1112PubMedCrossRefGoogle Scholar
  2. Bednekoff PA, Lima SL (2004) Risk allocation and competition in foraging groups: reversed effects of competition if group size varies under risk of predation. Proc Roy Soc Lond B 271:1491–1496CrossRefGoogle Scholar
  3. Berger J (1991) Pregnancy incentive and predation constraints in habitat shifts: experimental and field evidence for wild bighorn sheep. Anim Behav 41:61–77CrossRefGoogle Scholar
  4. Brashares JS, Arcese P (2002) Role of forage, habitat and predation in the behavioural plasticity of a small African antelope. J Anim Ecol 71:626–638CrossRefGoogle Scholar
  5. Brown JS, Laundré JW, Gurung M (1999) The ecology of fear: optimal foraging, game theory, and trophic interactions. J Mamm 80:385–399CrossRefGoogle Scholar
  6. Burger J, Gochfield M (1994) Vigilance in African mammals: differences among mothers, other females and males. Behaviour 131:154–274CrossRefGoogle Scholar
  7. Creel S, Winnie JA (2005) Responses of elk herd size to fine-scale spatial and temporal variation in the risk of predation by wolves. Anim Behav 69:1181–1189CrossRefGoogle Scholar
  8. Creel S, Winnie JA, Maxwell B, Hamlin K, Creel M (2005) Elk alter habitat selection as an antipredator response to wolves. Ecology 86:3387–3397CrossRefGoogle Scholar
  9. Dehn MM (1990) Vigilance for predators: detection and dilution effects. Behav Ecol Sociobiol 26:337–342Google Scholar
  10. Doncaster CP (1994) Factors regulating local variations in abundance: field tests on hedgehogs, Erinaceus europaeus. Oikos 69:182–192CrossRefGoogle Scholar
  11. Fenn MGP, Macdonald DW (1995) Use of middens by red foxes: risk reverses rhythms of rats. J Mamm 76:130–136CrossRefGoogle Scholar
  12. Firth D (1991) Generalized linear models. In: Hinkley DV, Reid N, Snell EJ (eds) Statistical theory and modelling. Chapman and Hall, LondonGoogle Scholar
  13. Fryxell JM (1991) Forage quality and aggregation by large herbivores. Am Nat 138:478–498CrossRefGoogle Scholar
  14. Hamilton WD (1971) Geometry for the selfish herd. J Theor Biol 31:295–311PubMedCrossRefGoogle Scholar
  15. Hebblewhite M, Merrill EH, McDonald TL (2005) Spatial decomposition of predation risk using resource selection functions: an example in a wolf–elk predator–prey system. Oikos 111:101–111CrossRefGoogle Scholar
  16. Hemson G, Johnson P, South A, Kenward R, Ripley R, Macdonald DW (2005) Are kernel the mustard? Data from global positioning system (GPS) collars suggests problems for kernel home-range analyses with least-squares cross-validation. J Anim Ecol 74:455–463Google Scholar
  17. Hopcraft GJC, Sinclair ARE, Packer C (2005) Planning for success: Serengeti lions seek prey accessibility rather than abundance. J Anim Ecol 75:559–566Google Scholar
  18. Hunter LTB, Skinner JD (1998) Vigilance behaviour in African ungulates: the role of predation pressure. Behaviour 135:195–211Google Scholar
  19. Jarman PJ (1974) The social organisation of antelope in relation to their ecology. Behaviour 48:215–267CrossRefGoogle Scholar
  20. Kotler BP, Brown JS, Hasson O (1991) Factors affecting gerbil foraging behaviour and rates of owl predation. Ecology 72:2249–2260CrossRefGoogle Scholar
  21. Kotler BP, Brown JS, Bouskila A (2004) Apprehension and time allocation in gerbils: the effects of predatory risk and energetic state. Ecology 85:917–922CrossRefGoogle Scholar
  22. Krause J (1994) Differential fitness returns in relation to spatial position in groups. Biol Rev 69:187–206PubMedCrossRefGoogle Scholar
  23. Liley S, Creel S (2008) What best explains vigilance in elk: characteristics of prey, predators, or the environment? Behav Ecol 19:245–254CrossRefGoogle Scholar
  24. Lima SL (1995) Back to the basics of anti-predatory vigilance: the group size effect. Anim Behav 49:11–20CrossRefGoogle Scholar
  25. Lima SL, Dill LM (1990) Behavioural decisions made under the risk of predation: a review and synthesis. Can J Zool 68:619–640CrossRefGoogle Scholar
  26. Loveridge AJ, Davidson Z, Hunt JE, Valeix M, Elliot N, Stapelkamp B (2007a) Hwange Lion Project Annual Report 2007. Zimbabwe Parks and Wildlife Management Authority, ZimbabweGoogle Scholar
  27. Loveridge AJ, Searle AW, Murindagomo F, Macdonald DW (2007b) The impact of sport-hunting on the lion population in a protected area. Biol Conserv 134:548–558CrossRefGoogle Scholar
  28. McNamara JM, Houston AI (1987) Starvation and predation as factors limiting population size. Ecology 68:1515–1519CrossRefGoogle Scholar
  29. Owen-Smith N, Mills MGL (2008) Predator–prey size relationships in an African large-mammal food web. J Anim Ecol 77:173–183PubMedCrossRefGoogle Scholar
  30. Prins HHT, Iason GR (1989) Dangerous lions and nonchalant buffalo. Behaviour 108:262–293CrossRefGoogle Scholar
  31. Pulliam HR (1973) On the advantages of flocking. J Theor Biol 38:419–422PubMedCrossRefGoogle Scholar
  32. Ripple WJ, Beschta RL (2004) Wolves and the ecology of fear: can predation risk structure ecosystems? BioScience 54:755–766CrossRefGoogle Scholar
  33. Roberts G (1996) Why individual vigilance declines as group size increases. Anim Behav 51:1077–1086CrossRefGoogle Scholar
  34. Rogers CML (1993) A woody vegetation survey of Hwange National Park. Department of National Parks and Wildlife Management, ZimbabweGoogle Scholar
  35. Roth TC, Lima SL (2007) The predatory behavior of wintering Accipiter hawks: temporal patterns in activity of predators and prey. Oecologia 152:169–178PubMedCrossRefGoogle Scholar
  36. Ryan SJ, Knechtel CU, Getz WM (2006) Range and habitat selection of African buffalo in South Africa. J Wildl Manage 70:764–776CrossRefGoogle Scholar
  37. Schaller GB (1972) The Serengeti lion: a study of predator prey relations. Chicago University Press, ChicagoGoogle Scholar
  38. Scheel D (1993) Profitability, encounter rates, and prey choice of African lions. Behav Ecol 4:90–97CrossRefGoogle Scholar
  39. South AB, Kenward RE (2006) Ranges7 v1.0: for the analysis of tracking and location data. Anatrack, WarehamGoogle Scholar
  40. Thrash I, Theron GK, Bothma JP (1995) Dry season herbivore densities around drinking troughs in the Kruger National Park. J Arid Environ 29:213–219CrossRefGoogle Scholar
  41. Underwood R (1982) Vigilance behaviour in grazing African antelopes. Behaviour 79:81–107CrossRefGoogle Scholar
  42. Valeix M, Fritz H, Matsika R, Matsvimbo F, Madzikanda H (2007) The role of water abundance, thermoregulation, perceived predation risk and interference competition in water access by African herbivores. Afr J Ecol 46:402–410CrossRefGoogle Scholar
  43. Valeix M, Loveridge AJ, Chamaillé-Jammes S, Davidson Z, Murindagomo F, Fritz H, Macdonald DW (2009) Behavioral adjustments of African herbivores to predation risk by lions: spatiotemporal variations influence habitat use. Ecology 90:23–30PubMedCrossRefGoogle Scholar
  44. Van Balaan M, Sabelis MV (1993) Coevolution of patch strategies of predator and prey and the consequences for ecological stability. Am Nat 142:646–670CrossRefGoogle Scholar
  45. Ward JF, Austin RM, Macdonald DW (2000) A simulation model of foraging behaviour and the effect of predation risk. J Anim Ecol 69:16–30CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2009

Authors and Affiliations

  • Marion Valeix
    • 1
    • 3
    Email author
  • Hervé Fritz
    • 3
  • Andrew J. Loveridge
    • 1
  • Zeke Davidson
    • 1
  • Jane E. Hunt
    • 1
  • Felix Murindagomo
    • 2
  • David W. Macdonald
    • 1
  1. 1.Wildlife Conservation Research Unit, Zoology DepartmentOxford UniversityAbingdonUK
  2. 2.Zimbabwe Parks and Wildlife Management AuthorityHarareZimbabwe
  3. 3.Université de Lyon, CNRS Université Claude Bernard Lyon 1 UMR 5558, Laboratoire Biométrie et Biologie EvolutiveVilleurbanne cedexFrance

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