Large carnivores inhabiting ecosystems with heterogeneously distributed environmental resources with strong seasonal variations frequently employ opportunistic foraging strategies, often typified by seasonal switches in diet. In semi-arid ecosystems, herbivore distribution is generally more homogeneous in the wet season, when surface water is abundant, than in the dry season when only permanent sources remain. Here, we investigate the seasonal contribution of the different herbivore species, prey preference and distribution of kills (i.e. feeding locations) of African lions in Hwange National Park, Zimbabwe, a semi-arid African savanna structured by artificial waterholes. We used data from 245 kills and 74 faecal samples. Buffalo consistently emerged as the most frequently utilised prey in all seasons by both male (56%) and female (33%) lions, contributing the most to lion dietary biomass. Jacobs’ index also revealed that buffalo was the most intensively selected species throughout the year. For female lions, kudu and to a lesser extent the group “medium Bovidae” are the most important secondary prey. This study revealed seasonal patterns in secondary prey consumption by female lions partly based on prey ecology with browsers, such as giraffe and kudu, mainly consumed in the early dry season, and grazers, such as zebra and suids, contributing more to female diet in the late dry season. Further, it revealed the opportunistic hunting behaviour of lions for prey as diverse as elephants and mice, with elephants taken mostly as juveniles at the end of the dry season during droughts. Jacobs’ index finally revealed a very strong preference for kills within 2 km from a waterhole for all prey species, except small antelopes, in all seasons. This suggested that surface-water resources form passive traps and contribute to the structuring of lion foraging behaviour.
References
[1]
Radloff FGT, du Toit JT (2004) Large predators and their prey in a southern African savanna: a predator’s size determines its prey range size. J Anim Ecol 73: 410–423.
[2]
Owen Smith N (2008) Changing vulnerability to predation related to season and sex in an African ungulate assemblage. Oikos 117: 602–610.
[3]
Bacon MM, Becic GM, Epp MT, Boyce MS (2011) Do GPS clusters really work? Carnivore diet from scat analysis and GPS telemetry methods. Wildl Soc Bull 35: 409–415.
[4]
Tambling CJ, Laurence SD, Bellan SE, Cameron EZ, du Toit JT, et al. (2012) Estimating carnivoran diets using a combination of carcass observations and scats from GPS clusters. J Zool 286: 102–109.
[5]
De Boer WF, Vis MJP, de Knegt HJ, Rowles C, Kohi EM, et al. (2010) Spatial distribution of lion kills determined by the water dependency of prey species. J Mamm 91: 1280–1286.
[6]
Hayward MW, Kerley GIH (2005) Prey preferences of the lion (Panthera leo). J Zool 267: 309–322.
[7]
Hayward MW (2006) Prey preferences of the spotted hyaena (Crocuta crocuta) and degree of dietary overlap with the lion (Panthera leo). J Zool 270: 606–614.
[8]
Hayward MW, Henschel P, O’Brien J, Hofmeyr M, Balme G, et al. (2006) Prey preferences of the leopard (Panthera pardus). J Zool 270: 298–313.
[9]
Western D (1975) Water availability and its influence on the structure and dynamics of a savannah large mammal community. E Afr Wildl J 13: 265–286.
[10]
De Leeuw J, Maweru MN, Okello OO, Maloba M, Nguru P, et al. (2001) Distribution and diversity of wildlife in northern Kenya in relation to livestock and permanent water points. Biol Conserv 100: 297–306.
[11]
Redfern JV, Grant R, Biggs H, Getz WM (2003) Surface-water constraints on herbivore foraging in the Kruger National Park, South Africa. Ecology 84: 2092–2107.
[12]
Smit IPJ, Grant CG, Devereux BJ (2007) Do artificial waterholes influence the way herbivores use the landscape? Herbivore distribution patterns around rivers and artificial surface water sources in a large African savanna park. Biol Conserv 136: 85–99.
[13]
Estes RD (1991) The behavior guide to African mammals. University of California Press, London, UK.
[14]
Thrash I, Theron GK, Bothma JdP (1995) Dry season herbivore densities around drinking troughs in the Kruger National Park. J Arid Environ 29: 213–219.
[15]
Valeix M (2011) Temporal dynamics of dry-season water-hole use by large African herbivores in two years of contrasting rainfall in Hwange National Park, Zimbabwe. J Trop Ecol 27: 163–170.
[16]
Owen-Smith N (1982) Factors influencing the consumption of plant products by large herbivores. In: Huntley BJ, Walker BH, editors. Ecology of tropical savanna. Berlin: Springer-Verlag. 359–404.
[17]
Walker BH, Emslie RH, Owen-Smith RN & Scholes RJ (1987) To cull or not to cull: lessons from a southern African drought. J Appl Ecol 24: 381–401.
[18]
Mills MGL (1995) Notes on wild dog Lycaon pictus and lion Panthera leo population trends during a drought in the Kruger National Park. Koedoe 38: 95–99.
[19]
Loveridge AJ, Hunt JE, Murindagomo F, Macdonald DW (2006) The influence of drought on predation of elephant calves by lions in an African wooded savannah. J Zool 270: 253–530.
[20]
Fritz H, Loison A (2006). Large herbivores across biomes. In: Danell K, Duncan P, Bergstr？m R, Pastor J (eds) Large herbivore ecology, ecosystem dynamics and conservation. Cambridge University Press, Cambridge, 19–49.
[21]
Owen-Smith N, Mills MGL (2008) Shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator prey web. Ecology 84: 1120–1133.
[22]
Holdo R (2003) Woody plant damage by African elephants in relation to leaf nutrients in western Zimbabwe. J Trop Ecol 19: 189–196.
[23]
Loveridge AJ, Searle AW, Murindagomo F, Macdonald DW (2007) The impact of sport-hunting on the lion population in a protected area. Biol Conserv 134: 548–558.
[24]
Tambling C, Cameron EZ, du Toit JT, Getz WM (2010) Methods for locating African lion kills using global positioning system movement data. J Wildl Manage 74: 549–556.
[25]
Funston PJ, Mills MGL, Biggs HC, Richardson PRK (1998) Hunting by male lions: ecological influences and socioecological implications. Anim Behav 56: 1333–1345.
[26]
Lehmann MB, Funston PJ, Owen CR, Slotow R (2008) Feeding behaviour of lions (Panthera leo) on a small reserve. S Afr J Wildl Res 38: 66–78.
[27]
Marucco F, Pletscher DH, Boitani L (2008) Accuracy of scat samples for carnivore diet analysis: wolves in the Alps as a case study. J Mamm 89: 665–673.
[28]
Van Kesteren F (2006) Analysis of the diet of African Lions (Panthera leo) in Hwange National Park, Zimbabwe. Masters thesis, University of Oxford, Oxford.
[29]
Marker LL, Muntifering JR, Dickman AJ, Mills MGL, Macdonald DW (2003) Quantifying prey preferences of free ranging Namibian Cheetahs. South Afr J Wildl Res 33: 45–53.
[30]
Perrin MR, Campbell BS (1979) Key to the mammals of the Andries Vosloo Kudu Reserve (Eastern Cape), based on their hair morphology, for use in predator scat analysis. South Afr J Wildl Res 10: 1–14.
[31]
Keogh HJ (1983) A photographic reference system of the microstructure of the hair of southern African bovids. South Afr J Wildl Res 13: 89–132.
[32]
Keogh HJ (1985) A photographic reference system based on the cuticular scale patterns and groove of the hair of 44 species of southern African Cricetidae and Muridae. South Afr J Wildl Res 15: 109–159.
[33]
Buys D, Keogh HJ (1984) Notes on the microstructure of hair of the Orycteropodidae, Elephantidae, Equidae, Suidae and Giraffidae. South Afr J Wildl Res 14: 111–119.
[34]
Cumming DHM, Cumming GS (2003) Ungulate community structure and ecological processes: body size, hoof area and trampling in African savannas. Oecologia 134: 560–568.
Chamaillé-Jammes S, Valeix M, Bourgarel M, Murindagomo F, Fritz H (2009) Seasonal density estimates of common large herbivores in Hwange National Park, Zimbabwe. Afr J Ecol 47: 804–808.
[37]
Schmidt-Nielsen K (1984) Scaling: why is animal size so important? Cambridge Press University, Cambridge, UK.
[38]
Karanth KU, Sunquist ME (1995) Prey selection by tiger, leopard and dhole in tropical forests. J Anim Ecol 64: 439–450.
[39]
Funston PJ, Mills MGL (2006) The influence of lion predation on the population dynamics of common large ungulates in the Kruger National Park. South Afr J Wildl Res 36: 9–22.
[40]
Loveridge AJ, Valeix M, Davidson Z, Murindagomo F, Fritz H, et al. (2009) Changes in home range size of African lions in relation to pride size and prey biomass in a semi-arid savanna. Ecography 32: 953–962.
[41]
Campbell K, Borner M (1995) Population trends and distribution of Serengeti herbivores: Implications for management. In: Sinclair ARE, Arcese P, editors. Serengeti II. Dynamics, management and conservation of an ecosystem. Chicago: University of Chicago Press. 117–145.
[42]
Fritz H, Loreau M, Chamaillé-Jammes S, Valeix M, Clobert J (2011) A food web perspective on large herbivore community limitation. Ecography 34: 196–202.
[43]
Hopcraft GC, Sinclair ARE, Packer C (2005) Planning for success: Serengeti lions seek prey accessibility rather than abundance. J Anim Ecol 74: 559–566.
[44]
Davidson Z, Valeix M, Loveridge AJ, Hunt JE, Johnson PJ, et al. (2012) Environmental determinants of habitat and kill site selection in a large carnivore: scale matters. J Mamm 93: 677–685.
[45]
Valeix M, Loveridge AJ, Davidson Z, Madzikanda H, Fritz H, et al. (2010) How key habitat features influence large terrestrial carnivore movements: waterholes and African lions in a semi-arid savanna of north-western Zimbabwe. Landsc Ecol 25: 337–351.
[46]
Valeix M, Chamaillé-Jammes S, Loveridge AJ, Davidson Z, Hunt JE, et al. (2011) Understanding patch departure rules for large carnivores: lion movements support a patch-disturbance hypothesis. Am Nat 179: 269–275.
[47]
Mosser A, Fryxell JM, Eberly L, Packer C (2009) Serengeti real estate: density vs. fitness-based indicators of lion habitat quality. Ecol Lett 12: 1050–1060.
[48]
Harrington R, Owen-Smith N, Viljoen PC, Biggs HC, Mason DR, et al. (1999) Establishing the causes of the roan antelope decline in the Kruger National Park, South Africa. Biol Conserv 90: 69–78.
[49]
Rogers CML (1993) A woody vegetation survey of Hwange National Park. Department of National Parks and Wildlife Management, Zimbabwe. 176p.
[50]
Hayward MW (2011) Scarcity in the prey community yields anti-predator benefits. Acta Oecol 37: 314–320.
