Biodiversity & Conservation

, Volume 8, Issue 12, pp 1643–1661

Functional significance of ungulate diversity in African savannas and the ecological implications of the spread of pastoralism

  • J.T. Du Toit
  • D.H.M. Cumming


The African savanna biome supports a higher diversity of ungulate species than is found in any other biome or continent. This exceptional faunal diversity and herbivore biomass density is directly linked to the high spatial heterogeneity of African savanna ecosystems. The dependence of herbivore dietary tolerance on body size translates into important size-related differences between savanna ungulate species in terms of habitat specificity, geographical range, and the share of community resources exploited. Intact savanna ungulate communities, with species distributed across body size classes and feeding guilds (grazer/browser), have strong regulatory influences on savanna ecosystem structure and function. Replacement with livestock systems of low diversity and high biomass density within a narrow body size range has occurred through the removal of competitors, pathogens, and predators, and the widespread provisioning of water. Overgrazing by livestock, coupled with episodic droughts, has caused widespread rangeland degradation and loss of floristic and faunal diversity which, by current models, is unlikely to recover to 'climax’ conditions even with destocking. In selected regions where potential still exists, African savanna biodiversity and human economic development will both be best served by the integration of sustainable wildlife utilization into multispecies animal production systems.

African savannas biodiversity conservation multispecies animal production 


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  1. Abel NOJ and Blaikie PM (1989) Land degradation, stocking rates and conservation policies in the communal rangelands of Botswana Zimbabwe. Land Degradation and Rehabilitation 1: 101–123Google Scholar
  2. Allen-Diaz B (1996) Rangelands in a changing climate: impacts, adaptations, and mitigation. In: Watson RT, Zinyowera MC, Moss RH and Dokken DJ (eds) Climate Change 1995. Impacts, Adaptations and Mitigation of Climate Change: Scientific-Technical Analysis, pp 131–158. Cambridge University Press, CambridgeGoogle Scholar
  3. Andrew MH (1988) Grazing impact in relation to livestock watering points. Trends in Ecology and Evolution 3: 336–339Google Scholar
  4. Bell RHV (1971) A grazing system in the Serengeti. Scientific American 224: 86–93Google Scholar
  5. Bell RHV (1982) The effect of soil nutrient availability on community structure in Africa savannas. In: Huntley BJ and Walker BH (eds) Ecology of Tropical Savannas, pp 193–216. Springer-Verlag, BerlinGoogle Scholar
  6. Bell RHV (1986) Soil-plant-herbivore interactions. In: Bell RHV and McShane-Caluzi E (eds) Conservation and Wildlife Management in Africa, pp 109–130. US Peace Corps, Washington, DCGoogle Scholar
  7. Belsky AJ (1984) Role of small browsing mammals in preventing woodland regeneration in the Serengeti National Park, Tanzania. African Journal of Ecology 22: 271–279Google Scholar
  8. Blackburn TM and Gaston KJ (1994) Abundance-body size relationships: the area you census tells you more. Oikos 75: 303–309Google Scholar
  9. Botkin DB, Melillo JM and Wu LSY (1981) How ecosystem processes are linked to large mammal population dynamics. In: Fowler CW and Smith D (eds) Dynamics of LargeMammal Populations, pp 373–387. Wiley, New YorkGoogle Scholar
  10. Bourliere F and Hadley M (1970) The ecology of tropical savannas. Annual Review of Ecology and Systematics 1: 125–152Google Scholar
  11. Brown JH and Nicoletto PF (1991) Spatial scaling of species composition: body masses of North American land mammals. American Naturalist 138: 1478–1512Google Scholar
  12. Bryant JP, Provenza FD, Pastor J, Reichardt PB, Clausen TP and du Toit JT (1991) Interactions between woody plants and browsing mammals mediated by secondary metabolites. Annual Review of Ecology and Systematics 22: 431–446Google Scholar
  13. Caughley G (1987) The distribution of eutherian body weights. Oecologia 74: 319–320Google Scholar
  14. Coe MJ, Cumming DHM and Phillipson J (1976) Biomass and production of large African herbivores in relation to rainfall and primary production. Oecologia 22: 341–354Google Scholar
  15. Coe MJ and Coe C (1987) Large herbivores, acacia trees and bruchid beetles. South African Journal of Science 83: 624–635Google Scholar
  16. Cumming DHM (1982) The influence of large herbivores on savanna structure in Africa. In: Huntley BJ and Walker BH (eds) Ecology of Tropical Savannas, pp 217–245. Springer-Verlag, BerlinGoogle Scholar
  17. Cumming DHM (1993) Multispecies systems: progress, prospects and challenges in sustaining range animal production and biodiversity in East and Southern Africa. Proceedings of the 7th World Conference on Animal Production, Edmonton, Canada 1: 145–159Google Scholar
  18. Cumming DHM(1994) Are multispecies systems a viable landuse option for southern African rangelands? In: Hofmann RR and Schwartz HJ (eds) Proc. Int. Symp. onWild and Domestic Ruminants in Extensive Land Use Systems, pp 203–234. Humboldt University, BerlinGoogle Scholar
  19. Damuth J (1981) Population density and body size in mammals. Nature 290: 699–700Google Scholar
  20. Damuth J (1987) Interspecific allometry of population density in mammals and other animals: the dependence of body mass and population energy use. Biological Journal of the Linnaean Society 31: 193–246Google Scholar
  21. Dean WRJ and Macdonald IAW (1994) Historical changes in stocking rates of domestic livestock as a measure of semi-arid and arid rangeland degradation in the Cape Province, South Africa. J. Arid Environ. 26: 281–298Google Scholar
  22. Demment MW and van Soest PJ (1985) A nutritional explanation for body-size patterns of ruminant and non-ruminant herbivores. American Naturalist 125: 641–672Google Scholar
  23. Denbow JR and Wilmsen EN (1986) Advent and cause of pastoralism in the Kalahari. Science 234: 1509–1515Google Scholar
  24. du Toit JT (1990a) The feeding ecology of a very small ruminant, the steenbok (Raphicerus campestris). African Journal of Ecology 31: 35–48Google Scholar
  25. du Toit JT (1990b) Giraffe feeding on Acacia flowers: predation or pollination? African Journal of Ecology 28: 63–68Google Scholar
  26. du Toit JT (1995) Determinants of the composition and distribution of wildlife communities in southern Africa. Ambio 24: 2–6Google Scholar
  27. du Toit JT and Owen Smith N (1989) Body size, population metabolism, and habitat specialization among large African herbivores. American Naturalist 133: 736–740Google Scholar
  28. du Toit JT, Bryant JP and Frisby K (1990) Regrowth and palatability of Acacia shoots following pruning by African savanna browsers. Ecology 71: 149–154Google Scholar
  29. Dorst J and Dandelot P (1970) A Field Guide to the Larger Mammals of Africa. Houghton Mifflin, BostonGoogle Scholar
  30. Dublin HT (1995)Vegetation dynamics in the Serengeti-Mara ecosystem: the role of elephants, fire, and other factors. In: Sinclair ARE and Arcese P (eds) Serengeti II: Dynamics, Management, and Conservation of an Ecosystem, pp 71–90. University of Chicago Press, Chicago, ILGoogle Scholar
  31. Dublin HT, Sinclair ARE and McGlade J (1990) Elephants and fire as causes of multiple stable states in the Serengeti-Mara woodlands. Journal of Animal Ecology 59: 1147–1164Google Scholar
  32. Dyson-Hudson N (1985) Pastoral production systems and livestock development projects: an East African perspective. In: Cernea MM (ed) Putting People First: Sociological Variables in Rural Development, pp 157–186. Oxford University Press, OxfordGoogle Scholar
  33. East R (1984) Rainfall, soil nutrient status and biomass of large African savanna mammals. African Journal of Ecology 22: 245–270Google Scholar
  34. East R (1988) Antelopes: Global Survey and Regional Action Plans. Part 1. East and Northeast Africa. IUCN, GlandGoogle Scholar
  35. East R (1989) Antelopes: Global Survey and Regional Action Plans. Part 2. Southern and South-Central Africa. IUCN, GlandGoogle Scholar
  36. East R (1990) Antelopes: Global Survey and Regional Action Plans. Part 3. West and Central Africa. IUCN, GlandGoogle Scholar
  37. Frost P, Medina E, Menaut J-C, Solbrig O, Swift M and Walker B (1986) Responses of Savannas to Stress and Disturbance: A Proposal for a Collaborative Programme of Research. Biology International, Special Issue No. 10, IUBS, ParisGoogle Scholar
  38. Gandar MV (1982) Trophic ecology and plant/herbivore energetics. In: Huntley BJ and Walker BH (eds) Ecology of Tropical Savannas, pp 514–534 Springer-Verlag, BerlinGoogle Scholar
  39. Huntley BJ (1982) Southern African savannas. In: Huntley BJ and Walker BH (eds) Ecology of Tropical Savannas, pp 101–119. Springer-Verlag, BerlinGoogle Scholar
  40. Huxley J (1961) 1. The Wild Riches of Africa; 2. The Wild Protein; 3. Wildlife as a World Asset. The Observer, LondonGoogle Scholar
  41. Janzen DH (1992) A south-north perspective on science in the management, use, and economic development of biodiversity. In: Sandlund OT, Hindar K and Brown AHD (eds) Conservation of Biodiversity for Sustainable Development, pp 27–52. Scandinavian University Press, OsloGoogle Scholar
  42. Jarman PJ (1974) The social organization of antelope in relation to their ecology. Behaviour 48: 215–266Google Scholar
  43. Justice C, Scholes R and Frost P (1994) African Savannas and the Global Atmosphere: Research Agenda. IGBP (Report No. 31), StockholmGoogle Scholar
  44. Laws RH, Parker ISC and Johnstone RCB (1975) Elephants and Their Habitats. Clarendon Press, OxfordGoogle Scholar
  45. Macdonald DW (1984) The Encyclopaedia of Mammals: 2. George Allen & Unwin, LondonGoogle Scholar
  46. Marquet PA, Navarette SA and Castilla JC (1995) Body size, population density, and the Energetic Equivalence Rule. Journal of Animal Ecology 64: 325–332Google Scholar
  47. Matzke G (1983) A reassessment of the expected development consequences of tsetse control efforts in Africa. Social Science in Medicine 17: 531–537Google Scholar
  48. McNaughton SJ (1984) Grazing lawns: animals in herds, plant form and coevolution. Am. Nat. 124: 863–886Google Scholar
  49. McNaughton SJ (1985) Ecology of a grazing ecosystem: the Serengeti. Ecological Monographs 55: 259–294Google Scholar
  50. McNaughton SJ and Chapin FS III (1985) Effects of phosphorus nutrition and defoliation on C4 graminoids from the Serengeti Plains. Ecology 66: 1617–1629Google Scholar
  51. McNaughton SJ and Georgiadis NJ (1986) Ecology of African grazing and browsing mammals. Annual Review of Ecology and Systematics 17: 39–65Google Scholar
  52. McNaughton SJ, Ruess RW and Seagle SW (1988) Large mammals and process dynamics in African ecosystems. BioScience 38: 794–800Google Scholar
  53. McNeely JA (1993) Economic incentives for conserving biodiversity: lessors from Africa, Ambio 22: 144–150Google Scholar
  54. Menaut J-C (1983) The vegetation of African savannas. In: Bourliere F (ed) Tropical Savannas, pp 109–149. Elsevier, A msterdamGoogle Scholar
  55. Miller MF (1994) The costs and benefits of Acacia seed consumption by ungulates. Oikos 71: 181–187Google Scholar
  56. Milton SJ, Dean WRJ, du Plessis MA and Siegfried WR (1994) A conceptual model of arid rangeland degradation. BioScience 44: 70–76Google Scholar
  57. O'Connor TG (1985) A synthesis of field experiments concerning the grass layer in the savanna regions of southern Africa. South African National Scientific Programmes Report, No. 114.Google Scholar
  58. O'Connor TG (1991) Local extinction in perennial grasslands: a life-history approach. American Naturalist 137: 735–773Google Scholar
  59. O'Connor TG and Campbell BM (1986) Hippopotamus habitat relationships on the Lundi River, Gonarezhou National Park, Zimbabwe. African Journal of Ecology 24: 7–26Google Scholar
  60. Oesterheld M, Sala OE and McNaughton SJ (1992) Effect of animal husbandry on herbivore-carrying capacity at a regional scale. Nature 356: 234–236Google Scholar
  61. Owen-Smith RN (1988) Megaherbivores: The Influence of Very Large Body Size on Ecology. Cambridge University Press, CambridgeGoogle Scholar
  62. Owen-Smith N and Cumming DHM (1993) Comparative foraging strategies of grazing ungulates in African savanna grasslands. Proceedings of the XVII International Grasslands Congress New Zealand 691–698Google Scholar
  63. Parker ISC (1989) The Raw Ivory Trade: 1979–1989. Report to CITES, Lausanne, Switzerland.Google Scholar
  64. Pastor J, Naiman RJ, Dewey B and McInnes P (1988) Moose, microbes, and the boreal forest, BioScience 38: 770–777Google Scholar
  65. Pellew R (1983) The impacts of elephant, giraffe, and fire upon the Acacia tortilis woodlands of the Serengeti. African Journal of Ecology 21: 41–74Google Scholar
  66. Phillipson J (1973) The biological efficiency of protein production by grazing and other land-based systems. In: Jones JGW (ed) The Biological Efficiency of Protein Production, pp 217–223. Cambridge University Press, CambridgeGoogle Scholar
  67. Prins HHT and van der Jeugd HP (1993) Herbivore population crashes and woodland structure in East Africa. Journal of Ecology 81: 305–314Google Scholar
  68. Silva M and Downing JA (1995) The allometric scaling of density and body mass: a nonlinear relationship for terrestrial mammals. American Naturalist 145: 704–727Google Scholar
  69. Sinclair ARE (1975) The resource limitation of trophic levels in tropical grassland ecosystems. Journal of Animal Ecology 44: 497–520Google Scholar
  70. Sinclair ARE (1995) Equilibria in plant-herbivore interactions. In: Sinclair ARE and Arcese P (eds) Serengeti II: Dynamics, Management, and Conservation of an Ecosystem, pp 91–113. University of Chicago Press, Chicago, ILGoogle Scholar
  71. Skarpe C (1991) Impact on grazing in savanna ecosystems. Ambio 20: 351–356Google Scholar
  72. Smithers RHN (1983) The mammals of the Southern African Subregion. University of Pretoria Press, PretoriaGoogle Scholar
  73. Stuart SN and Adams RJ (1990) Biodiversity in Sub-Saharan Africa and its Islands. IUCN-SSC (Occasional Paper No. 6), GlandGoogle Scholar
  74. Tilman D and El Haddi A (1992) Drought and biodiversity in grasslands. Oecologia 89: 257–264Google Scholar
  75. Tshuma J, Logan JWM and Pearce MJ (1988) Termites attacking field crops, pasture and forest trees in Zimbabwe. Zimbabwe Journal of Agricultural Research 26: 87–97Google Scholar
  76. Turpie JK and Crowe TM (1994) Patterns of distribution, diversity and endemism of larger African mammals. South African Journal of Zoology 29: 19–32Google Scholar
  77. Vesey-Fitzgerald DF (1960) Grazing succession among East African game animals. Journal of Mammalogy 41: 161–172Google Scholar
  78. Vrba Es (1992) Mammals as a key to evolutionary theory. Journal of Mammalogy 73: 1–28Google Scholar
  79. Walker BH and Noy Meir I (1982) Aspects of the stability and resilience of savanna ecosystems. In: Huntley BJ and Walker BH (eds) Ecology of Tropical Savannas, pp 556–590. Springer-Verlag, BerlinGoogle Scholar
  80. Walker BH, Emslie RH, Owen-Smith RN and Scholes RJ (1987) To cull or not to cull: lessons from a southern African drought. Journal of Applied Ecology 24: 381–401Google Scholar
  81. Walker BH (1989) Diversity and stability in ecosystem conservation. In: Western D and Pearl M (eds) Conservation for the 21st Century, pp 121–130. Oxford University Press, OxfordGoogle Scholar
  82. Walker BH (1993) Rangeland ecology: understanding and managing change. Ambio 22: 80–87Google Scholar
  83. Walter H (1971) Ecology of Tropical and Subtropical Vegetation. Oliver Boyd, EdinburghGoogle Scholar
  84. Westoby M, Walker BH and Noy-Meir I (1989) Opportunistic management for rangelands not at equilibrium. Journal of Range Management 42: 266–274Google Scholar
  85. World Conservation Monitoring Centre (1992) Global Biodiversity: Status of the Earth's Living Resources. Chapman & Hall, LondonGoogle Scholar
  86. Young MD and Solbrig OT (1993) Providing an environmentally sustainable, economically profitable and socially equitable future for the world's savannas. In: Young MD and Solbrig OT (eds) The World's Savannas, pp 321–344. UNESCO Man and the Biosphere Series, ParisGoogle Scholar

Copyright information

© Kluwer Academic Publishers 1999

Authors and Affiliations

  • J.T. Du Toit
    • 1
  • D.H.M. Cumming
    • 2
  1. 1.Mammal Research Institute, Department of Zoology and EntomologyUniversity of PretoriaPretoriaSouth Africa
  2. 2.WWF Southern Africa Regional Programme OfficeCauseway, HarareZimbabwe

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