, Volume 165, Issue 1, pp 89–99 | Cite as

Landscape-scale feeding patterns of African elephant inferred from carbon isotope analysis of feces

  • Jacqueline CodronEmail author
  • Daryl Codron
  • Julia A. Lee-Thorp
  • Matt Sponheimer
  • Kevin Kirkman
  • Kevin J. Duffy
  • Judith Sealy
Behavioral ecology - Original Paper


The African elephant (Loxodonta africana) is a large-bodied, generalist herbivore that eats both browse and grass. The proportions of browse and grass consumed are largely expected to reflect the relative availability of these resources. We investigated variations in browse (C3 biomass) and grass (C4) intake of the African elephant across seasons and habitats by stable carbon isotope analysis of elephant feces collected from Kruger National Park, South Africa. The results reflect a shift in diet from higher C4 grass intake during wet season months to more C3 browse-dominated diets in the dry season. Seasonal trends were correlated with changes in rainfall and with nitrogen (%N) content of available grasses, supporting predictions that grass is favored when its availability and nutritional value increase. However, switches to dry season browsing were significantly smaller in woodland and grassland habitats where tree communities are dominated by mopane (Colophospermum mopane), suggesting that grasses were favored here even in the dry season. Regional differences in diet did not reflect differences in grass biomass, tree density, or canopy cover. There was a consistent relationship between %C4 intake and tree species diversity, implying that extensive browsing is avoided in habitats characterized by low tree species diversity and strong dominance patterns, i.e., mopane-dominated habitats. Although mopane is known to be a preferred species, maintaining dietary diversity appears to be a constraint to elephants, which they can overcome by supplementing their diets with less abundant resources (dry season grass). Such variations in feeding behavior likely influence the degree of impact on plant communities and can therefore provide key information for managing elephants over large, spatially diverse, areas.


Browse Grass Kruger Park Megaherbivore Mixed-feeder 



We thank the staff of the Kruger National Park, in particular R. Grant, I. Whyte, H. Biggs, A. de Buys, A. Gaylard, C. Trennery, R. Jansen van Vuuren, D. Nariandas, O. Mathebula, W. Dhinda, and J. Baloyi. We thank J. Sealy, J. Lanham, I. Newton, and T. Duda at the University of Cape Town, and T. Cerling, D. Dearing, and J. Ehleringer at the University of Utah. E. Codron assisted with fieldwork. Three anonymous reviewers are thanked for their useful comments on the manuscript. This project was supported by grants from the University of KwaZulu-Natal, National Research Foundation (NRF, RSA), University of Cape Town, START/PACOM, and the National Science Foundation (NSF, USA). Experiments in this study comply with the current laws of the Republic of South Africa, the country in which they were performed.


  1. Barnes RFW (1982) Elephant feeding behavior in Ruaha National Park, Tanzania. Afr J Ecol 20:123–136CrossRefGoogle Scholar
  2. Cerling TE, Harris JM (1999) Carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies. Oecologia 120:347–363CrossRefGoogle Scholar
  3. Cerling TE, Harris JM, Leakey MG (1999) Browsing and grazing in elephants: the isotope record of modern and fossil proboscideans. Oecologia 120:364–374CrossRefGoogle Scholar
  4. Clauss M, Streich WJ, Schwarm A, Ortmann S, Hummel J (2007) The relationship of food intake and ingesta passage predicts feeding ecology in two different megaherbivore groups. Oikos 116:209–216CrossRefGoogle Scholar
  5. Clemens ET, Maloiy GMO (1983) Nutrient digestibility and gastrointestinal electrolyte flux in the elephant and rhinoceros. Comp Biochem Physiol A Physiol 75:653–658CrossRefGoogle Scholar
  6. Coates DB, van der Weide APA, Kerr JD (1991) Changes in faecal δ13C in response to changing proportions of legume (C3) and grass (C4) in the diet of sheep and cattle. J Agric Sci 116:287–295CrossRefGoogle Scholar
  7. Codron J (2008) Annals of ivory: perspectives on African elephant Loxodonta africana (Blumenbach 1797) feeding ecology from a multi-decadal record. PhD thesis. University of Cape Town, Cape TownGoogle Scholar
  8. Codron D, Lee-Thorp JA, Sponheimer M et al (2005) Assessing diet in savanna herbivores using stable carbon isotope ratios of faeces. Koedoe 48:115–124Google Scholar
  9. Codron J, Codron D, Lee-Thorp JA et al (2005) Taxonomic, anatomical, and spatio-temporal variations in the stable carbon and nitrogen isotopic compositions of plants from an African savanna. J Archaeol Sci 32:1757–1772Google Scholar
  10. Codron J, Lee-Thorp JA, Sponheimer M, Codron D, Grant RC, De Ruiter DJ (2006) Elephant (Loxodonta africana) diets in Kruger National Park, South Africa: spatial and landscape differences. J Mammal 87:27–34CrossRefGoogle Scholar
  11. Codron D, Lee-Thorp JA, Sponheimer M, Codron J (2007a) Nutritional content of savanna plant foods: implications for browser/grazer models of ungulate diversification. Eur J Wildl Res 53:100–111CrossRefGoogle Scholar
  12. Codron D, Lee-Thorp JA, Sponheimer M, Codron J (2007b) Stable carbon isotope reconstruction of ungulate diet changes through the seasonal cycle. S Afr J Wildl Res 37:117–125CrossRefGoogle Scholar
  13. Cumming DHM Fenton MB, Rautenbach IL et al (1997) Elephants, woodlands and biodiversity in Southern Africa. S Afr J Sci 93:231–236Google Scholar
  14. De Boer WF, Ntumi CP, Correia AU, Mafuca JM (2000) Diet and distribution of elephant in the Maputo Elephant Reserve, Mozambique. Afr J Ecol 38:188–201CrossRefGoogle Scholar
  15. Dublin HT, Sinclair ARE, McGlade J (1990) Elephants and fire as causes of multiple stable states in the Serengeti-Mara woodlands. J Anim Ecol 59:1147–1164CrossRefGoogle Scholar
  16. Grant CC, Davidson T, Funston PJ, Pienaar DJ (2002) Challenges faced in the conservation of rare antelope: a case study on the northern basalt plains of the Kruger National Park. Koedoe 45:45–66Google Scholar
  17. Guy PR (1976) The feeding behavior of elephant (Loxodonta africana) in the Sengwa area, Rhodesia. S Afr J Wildl Res 6:55–63Google Scholar
  18. Jachmann H, Bell RHV (1985) Utilization by elephants of the Brachystegia woodlands of the Kasungu National Park, Malawi. Afr J Ecol 23:245–258CrossRefGoogle Scholar
  19. Kerley GIH, Landman M (2006) The impacts of elephants on biodiversity in the Eastern Cape Subtropical Thickets. S Afr J Sci 102:395–402Google Scholar
  20. Koch PL, Heisinger J, Moss C, Carlson RW, Fogel ML, Behrensmeyer AK (1995) Isotopic tracking of change in diet and habitat use in African elephants. Science 267:1340–1343CrossRefPubMedGoogle Scholar
  21. Laws RM, Parker ISC, Johnstone RCB (1974) Elephants and their habitats: the ecology of elephants in North Bunyoro, Uganda. Clarendum, OxfordGoogle Scholar
  22. Lee-Thorp JA, van der Merwe NJ (1987) Carbon isotope analysis of fossil bone apatite. S Afr J Sci 83:712–715Google Scholar
  23. Lewis DM (1991) Observations of tree growth, woodland structure and elephant damage of Colophospermum mopane in Luangwa Valley, Zambia. Afr J Ecol 29:207–221CrossRefGoogle Scholar
  24. Meissner HH, Zacharias PJK, O’reagain PJ (1999) Forage quality (feed value). In: Tainton NM (ed) Veld management in South Africa. University of Natal Press, Pietermaritzburg, pp 139–168Google Scholar
  25. Norton-Griffiths M (1979) The influence of grazing, browsing, and fire on the vegetation dynamics of the Serengeti. In: Sinclair ARE, Norton-Griffiths M (eds) Serengeti: dynamics of an ecosystem. University of Chicago Press, Chicago, pp 310–352Google Scholar
  26. O’Connor TG, Goodman PS, Clegg B (2007) A functional hypothesis of the threat of local extirpation of woody plant species by elephant in Africa. Biol Conserv 136:329–345CrossRefGoogle Scholar
  27. Owen-Smith RN (1988) Megaherbivores—The influence of very large body size on ecology. Cambridge University Press, CambridgeGoogle Scholar
  28. Post DM (2002) Using stable isotopes to estimate trophic position: Models, methods, and assumptions. Ecology 83:703–718CrossRefGoogle Scholar
  29. Rogosic J, Estell R, Skobic D, Martinovic A, Maric S (2006) Role of species diversity and secondary compound complementarity on diet selection of Mediterranean shrubs by goats. J Chem Ecol 32:1279–1287CrossRefPubMedGoogle Scholar
  30. Scholes RJ, Bond WJ, Eckhardt HC (2003) Vegetation dynamics in the Kruger ecosystem. In: du Toit JT, Rogers KH, Biggs HC (eds) The Kruger experience. Island Press, Washington D.C., pp 242–262Google Scholar
  31. Skarpe C et al (2004) The return of the giants: ecological effects of an increasing elephant population. Ambio 33:276–282PubMedGoogle Scholar
  32. Smallie JJ, O’Connor TG (2000) Elephant utilization of Colophospermum mopane: possible benefits of hedging. Afr J Ecol 38:352–359CrossRefGoogle Scholar
  33. Smit IPJ, Grant CC, Whyte IJ (2007) Landscape-scale sexual segregation in the dry season distribution and resource utilization of elephants in Kruger National Park, South Africa. Divers Distrib 13:225–236CrossRefGoogle Scholar
  34. Sorensen J, Heward E, Dearing M (2005) Plant secondary metabolites alter the feeding patterns of a mammalian herbivore (Neotoma lepida). Oecologia 146:415–422CrossRefPubMedGoogle Scholar
  35. Sponheimer M et al (2003) An experimental study of carbon-isotope fractionation between diet, hair, and feces of mammalian herbivores. Can J Zool 81:871–876CrossRefGoogle Scholar
  36. Stokke S, du Toit JT (2000) Sex and size related differences in the dry season feeding patterns of elephants in Chobe National Park, Botswana. Ecography 23:70–80CrossRefGoogle Scholar
  37. Styles CV, Skinner JD (2000) The influence of large mammalian herbivores on growth form and utilization of mopane trees, Colophospermum mopane, in Botswana’s Northern Tuli Game Reserve. Afr J Ecol 38:95–101CrossRefGoogle Scholar
  38. Tieszen LL, Boutton TW, Ottichilo WK, Nelson DE, Brandt DH (1989) An assessment of long-term food habits of Tsavo elephants based on stable carbon and nitrogen isotope ratios of bone collagen. Afr J Ecol 27:219–226CrossRefGoogle Scholar
  39. Trollope WSW, Potgieter ALF, Zambatis N (1989) Assessing veld condition in the Kruger National Park using key grass species. Koedoe 32:68–75Google Scholar
  40. van der Merwe NJ, Lee Thorp JA, Bell RHV (1988) Carbon isotopes as indicators of elephant diets in African environments. Afr J Ecol 26:163–172CrossRefGoogle Scholar
  41. Van Soest PJ (1994) Nutritional ecology of the ruminant, 2nd edn. Comstock, IthacaGoogle Scholar
  42. Venter FJ, Scholes RJ, Eckhardt HC (2003) The abiotic template and its associated vegetation pattern. In: du Toit JT, Rogers KH, Biggs HC (eds) The Kruger experience. Island Press, Washington D.C., pp 83–129Google Scholar
  43. Viljoen PJ (1989) Habitat selection and preferred food plants of a desert-dwelling elephant population in the Northern Namib Desert, South West Africa/Namibia. Afr J Ecol 27:227–240CrossRefGoogle Scholar
  44. Vogel JC (1978) Isotopic assessment of the dietary habits of ungulates. S Afr J Sci 74:298–301Google Scholar
  45. Vogel JC, Talma AS, Hall-Martin AJ, Viljoen PJ (1990) Carbon and nitrogen isotopes in elephants. S Afr J Sci 86:147–150Google Scholar
  46. Wiggins N, McArthur C, Davies N (2006) Diet switching in a generalist mammalian folivore: fundamental to maximising intake. Oecologia 147:650–657CrossRefPubMedGoogle Scholar
  47. Williamson BR (1975) The condition and nutrition of elephant in Wankie National Park. Arnoldia 7:1–20Google Scholar
  48. Wing LD, Buss IO (1970) Elephants and forests. Wildl Monogr 19:3–92Google Scholar
  49. Wrench JM, Meissner HH, Grant CC, Casey NH (1996) Environmental factors that affect the concentration of P and N in faecal samples collected for the determination of nutritional status. Koedoe 39:1–6Google Scholar

Copyright information

© Springer-Verlag 2010

Authors and Affiliations

  • Jacqueline Codron
    • 1
    • 2
    • 7
    Email author
  • Daryl Codron
    • 1
    • 3
  • Julia A. Lee-Thorp
    • 4
  • Matt Sponheimer
    • 5
  • Kevin Kirkman
    • 1
  • Kevin J. Duffy
    • 6
  • Judith Sealy
    • 2
  1. 1.School of Biological and Conservation SciencesUniversity of KwaZulu-NatalScottsvilleRSA
  2. 2.Department of ArchaeologyUniversity of Cape TownRondeboschRSA
  3. 3.Clinic for Zoo Animals, Exotic Pets and Wildlife, Vetsuisse FacultyUniversity of ZurichZurichSwitzerland
  4. 4.Division of Archaeological, Environmental and Geographical SciencesBradford UniversityBradfordUK
  5. 5.Department of AnthropologyUniversity of Colorado at BoulderBoulderUSA
  6. 6.Centre for Systems ResearchDurban Institute of TechnologyDurbanRSA
  7. 7.ZurichSwitzerland

Personalised recommendations