Skip to main content Accessibility help

Resource partitioning among African savanna herbivores in North Cameroon: the importance of diet composition, food quality and body mass

  • H. H. de Iongh (a1), C. B. de Jong (a2), J. van Goethem (a1), E. Klop (a1), A. M. H. Brunsting (a2), P. E. Loth (a1) and H. H. T. Prins (a2)...


The relationship between herbivore diet quality, and diet composition (the range of food plants consumed) and body mass on resource partitioning of herbivores remains the subject of an ongoing scientific debate. In this study we investigated the importance of diet composition and diet quality on resource partitioning among eight species of savanna herbivore in north Cameroon, with different body mass. Dung samples of four to seven wild herbivore and one domesticated species were collected in the field during the dry and wet period. Diet composition was based on microhistological examination of herbivore droppings, epidermis fragments were identified to genus or family level. In addition, the quality of the faecal droppings was determined in terms of phosphorus, nitrogen and fibre concentrations. The results showed that there was no significant correlation between body mass and (differences in) diet composition for wet and dry season. When all species are considered, only significant relationships are found by the Spearman rank correlation analyses during the wet season between body mass and phosphorus and nitrogen, but this relationship did not exist during the dry season. When the analyses focuses on ruminants only (thus leaving out hippo), none of the relationships between body mass and diet quality was significant in either season. During the dry season the proportion of graminoids ranged between 10% (small unidentified herbivore species) to 90% (hippopotamus), during the wet season this proportion ranged from 60% (zebu) to 90% (hippopotamus). All species but zebu had more graminoids in their dung during wet season compared with dry season. However all species but hartebeest had more graminoids old stems in their dung during the dry season, compared with the wet season. The niche breadth for food categories consumed by kob (0.300), hippo (0.090), hartebeest (0.350), roan (0.510) and zebu (0.300) was much greater in the dry season than in the wet season for kob (0.120), hippo (0.020), hartebeest (0.190), roan (0.090) and zebu (0.200). When looking at grass taxa consumed, the niche breadth of kob (0.220), hartebeest (0.140), and roan (0.250) was also greater in the dry season when compared with the wet season for kob (0.050), hartebeest (0.120) and roan (0.120). The opposite was found for zebu and hippo. Comparison of the species’ diet compositions with randomized data showed that dietary overlap between different herbivore species was much higher than what would be expected on the basis of chance, demonstrating surprisingly limited niche separation between species. This offers potential for competition, but it is more likely that the high niche overlap indicates absence of competition, due to low herbivore densities and abundant food resources, permitting species to share non-limiting resources. With increasing herbivore densities and subsequent increasing scarcity of resources, the relationship between diet quality and body mass in combination with increased niche separation is expected to become more visible.


Corresponding author

1Corresponding author. Email:,


Hide All
ALIPAYO, D., VALDEZ, R., HOLECHECK, J. L. & CARDENAS, M. 1992. Evaluation of microhistological analysis for determining ruminant diet botanical composition. Journal of Rangeland Management 45:148152.
ARSENAULT, R. & OWEN-SMITH, N. 2002. Facilitation versus competition in grazing herbivore assemblages. Oikos 97:313318.
AUBRÉVILLE, A. 1970. Flore du Cameroun 9: Légumineuses − Césalpinioidées. Museum National d'Histoire Naturelle, Paris. 376 pp.
BELL, R. H. V. 1971. A grazing ecosystem in the Serengeti. Scientific American 225:8693.
BRABANT, P. & HUMBEL, F. X. 1974. Notice explicative de la carte pédologique de Poli, No 51. ORSTOM, Paris. 123 pp.
CID, M. S. & BRIZUELA, M. A. 1990. Grass blade and sheath quantification by microhistological analysis. Journal of Wildlife Management 54:349352.
CLAUSS, M., SCHWARM, A., ORTMANN, S., ALBER, D., FLACH, E. J., KHÜNE, R., HUMMEL, J., STREICH, W. J. & HOFER, H. 2004. Intake, ingesta retention, particle size distribution and digestibility in the Hippopotamidae. Comparative Biochemistry and Physiology, Part A 139:449459.
CLAUSS, M., SCHWARMM, A., ORTMANN, S., STREICH, W. J. & HUMMEL, J. 2007. A case of non-scaling in mammalian physiology? Body size, digestive capacity, food intake, and ingesta passage in mammalian herbivores. Comparative Biochemistry and Physiology Part A 148:249265.
CODRON, D., LEE-THORP, J. A., SPONHEIMER, M., CODRON, J., DE RUITER, D. & BRINK, J. S. 2007. Significance of diet type and diet quality for ecological diversity of African ungulates. Journal of Animal Ecology 76:526537.
CROMSIGT, J. P. & OLFF, H. 2006. Resource partitioning among savanna grazers mediated by local heterogeneity: an experimental approach. Ecology 87:15321541.
DE JONG, C. B., GILL, R. M. A., VAN WIEREN, S. E. & BURLTON, F. W. E. 1995. Diet selection in Kielder Forest by roe deer Capreolus capreolus in relation to plant cover. Forest Ecology and Management 79:9197.
DE JONG, C. B., VAN WIEREN, S. E., GILL, R. M. A. & MUNRO, R. 2004. Relationship between diet and liver carcinomas in roe deer in Kielder Forest and Galloway forest. Veterinary Records 155:197200.
DEMMENT, M. W. 1982. The scaling of ruminoreticulum size with body weight in East African ungulates. African Journal of Ecology 20:4347.
DEMMENT, M. W. & VAN SOEST, P. J. 1985. A nutritional explanation for body size patterns of ruminant and non-ruminant herbivores. American Naturalist 125:641672.
GOERING, H. K. & VAN SOEST, P. J. 1970. Forage fiber analyses (apparatus, reagents, procedures and some applications). Agricultural Handbook No.379, Agricultural Resources Service, Washington.
GORDON, I. J. & ILLIUS, A. W. 1989. Resource partitioning by ungulates on the Isle of Rhum. Oecologia 79:383389.
GORDON, I. J. & ILLIUS, A. W. 1994. The functional significance of the browser–grazer dichotomy in African ruminants. Oecologia 98:167175.
GORDON, I. J. & ILLIUS, A. W. 1996. The nutritional ecology of African ruminants: a reinterpretation. Journal of Animal Ecology 65:1828.
GOTELLI, N. J. & GRAVES, G. R. 1996. Null models in ecology. Smithsonian Institution Press, Washington DC. 368 pp.
HANSEN, R. M., MUGAMBI, M. M. & NAUNI, S. M. 1985. Diets and trophic ranking of ungulates of the northern Serengeti. Journal of Wildlife Management 49:823829.
HOMOLKA, M. & HEROLDOVÁ, M. 1992. Similarity of the results of stomach and fecal contents analyses in studies of the ungulate diet. Folia Zoologica 41:193208.
HURLBERT, S. H. 1978. The measurement of niche overlap and some relatives. Ecology 59:6777.
IASON, G. R. & VAN WIEREN, S. E. 1999. Digestive and ingestive adaptations of mammalian herbivores to low quality forage. Pp. 337369 in Olff, H., Brown, V. K. & Drent, R. H. (eds.). Herbivores: between plants and predators. Blackwell, Oxford.
ILLIUS, A. W. & GORDON, I. J. 1992. Modelling the nutritional ecology of ungulate herbivores: evolution of body size and competitive interactions. Oecologia 89:428434.
JARMAN, P. J. 1974. The social organization of antelopes in relation to their ecology. Behaviour 48:215266.
KINGDON, J. 1997. The Kingdon field guide to African mammals. Academic Press, San Diego. 465 pp.
KLEIBER, M. 1975. The fire of life: an introduction to animal energetics. Krieger, Huntington. 453 pp.
KLOP, E. & VAN GOETHEM, J. 2008. Savanna fires govern community structure of ungulates in Bénoué National Park, Cameroon. Journal of Tropical Ecology 24:3947.
KLOP, E., VAN GOETHEM, J. & DE IONGH, H. H. 2007. Resource selection by grazing herbivores on post-fire regrowth in a West African woodland savanna. Wildlife Research 34:7783.
LEGENDRE, P. & LEGENDRE, L. 1998. Numerical ecology. (Second edition). Elsevier, Amsterdam. 852 pp.
LEVINS, R. 1968. Evolution in changing environments: some theoretical explorations. Princeton University Press, Princeton. 120 pp.
LESLIE, D. M., BROWN, R. T. & JENKS, J. A. 2007. Facts from feces: nitrogen still measures up as a nutritional index for mammalian herbivores. Journal of Wildlife Management 72:14201433.
MACARTHUR, R. H. & LEVINS, R. 1967. The limiting similarity, convergence and divergence of co-exiting species. American Naturalist 101:377385.
MAYAKA, T. B. 2002. Value wildlife! An ecological and economic assessment of wildlife use in Northern Cameroon. Ph.D. thesis, Leiden University, Leiden.
METCALFE, C. R. 1960. Anatomy of the monocotyledons, Vol. 1: Gramineae. Clarendon Press, Oxford.
MYSTERUD, A. 2000. Diet overlap among ruminants in Fennoscandia. Oecologia 124:130137.
NAMGAIL, T. 2009. Biogeography of mammalian herbivores in Ladakh: distribution in relation to environmental and geographical barriers. Ph.D. thesis, Wageningen University and Research Centre.
NAMGAIL, T., FOX, J. L. & BHATANAGAR, Y. V. 2004. Habitat segregation between sympatric Tibetan argali Ovis ammon hodgsoni and blue sheep Pseudois nayaur in the Indian Trans-Himalaya. Journal of Zoology (London) 262:5763.
PÉREZ-BARBERIA, F. J., ELSTON, D. A., GORDON, I. J. & ILLIUS, A. W. 2004. The evolution of phylogenetic differences in the efficiency of digestion in ruminants. Proceedings of the Royal Society of London, Biological Sciences 271:10811090.
PIANKA, E. R. 1973. The structure of lizard communities. Annual Review of Ecology and Systematics 4:5374.
PRINS, H.H.T. & OLFF, H. 1998. Species-richness of African grazer assemblages: towards a functional explanation. Pp. 449490 in Newbery, D. M., Prins, H. H. T. & Brown, N. D. (eds.). Dynamics of tropical communities. Blackwell Science, Oxford.
PRINS, H. H. T., DE BOER, W. F., VAN OEVEREN, H., CORREIA, A., MAFUCA, J. & OLFF, H. 2006. Co-existence and niche segregation of three small bovid species in southern Mozambique. African Journal of Ecology 44:186198.
PUTMAN, R. J. 1984. Facts from faeces. Mammalian Review 14:7997.
PUTMAN, R. J. 1996. Competition and resource partitioning in temperate ungulate assemblies. Chapman and Hall, London. 135 pp.
RICHTER, H. V. & CUMMING, G. S. 2006. Food availability and annual migration of the straw-colored fruit bat (Eidolon helvum). Journal of Zoology 268:3544.
ROBBINS, C. T., SPALINGER, D. E. & VAN HOVEN, W. 1995. Adaptation of ruminants to browse and grass diets: are anatomical-based browser–grazer interpretations valid? Oecologia 103:208213.
SCHWARM, A., CLAUSS, M., FLACH, E. J. & TACK, C. 2003. Passage rate and digestibility coefficients in captive Hippopotamidae – a pilot study. Verhaltnisse Zaugetiere 41:413418.
SCHWARM, A., ORTMANN, S., HOFER, H., STREICH, W. J., FLACH, E. J., KHÜNE, R., HUMMEL, J., CASTELL, J. C., SCHWARM, C. & CLAUSS, M. 2006. Digestion studies in captive Hippopotamidae: a group of large ungulates with an unusually low metabolic rate. Journal of Animal Physiology and Animal Nutrition 90:300308.
SPARKS, D. R. & MALECHEK, C. 1968. Estimating percentage dry weight in diets using a microscopic technique. Journal of Rangeland Management 21:264265.
STARK, M. A. & HUDSON, R. J. 1985. Plant communities’ structure in Bénoué National Park, Cameroon: a cluster association analysis. African Journal of Ecology 23:2127.
STEWART, D. R. M. 1967. Analysis of plant epidermis in faeces: a technique for studying the food preferences of grazing herbivores. Journal of Applied Ecology 4:83111.
STUART, C. & STUART, T. 2000. A field guide to the tracks and signs of Southern and East African Wildlife. Struik Publishers, Cape Town. 310 pp.
VAN DE VIJVER, C. A. D. M., POOT, P. & PRINS, H. H. T. 1999. Causes of increased nutrient concentrations in post-fire regrowth in an East African savanna. Plant and Soil 214:173185.
VAN SOEST, P. J. 1994. Nutritional ecology of the ruminant. Cornell University Press, New York. 479 pp.
VERWEIJ, R. J. T., VERRELST, J., LOTH, P., HEITKÖNIG, I. M. A. & BRUNSTING, A. M. H. 2006. Grazing lawns contribute to the subsistence of mesoherbivores on dystrophic savannas. Oikos 114:108116.
VOETEN, M. M. & PRINS, H. H. T. 1999. Resource partitioning between sympatric wild and domestic herbivores in the Tarangire region of Tanzania. Oecologia 120:287294.
WEGGE, P., SHRESTHA, A. K. & MOE, S. R. 2006. Dry season diets of sympatric ungulates on lowland Nepal: competition and facilitation in alluvial tall grasslands. Ecological Research 21:698706.


Related content

Powered by UNSILO

Resource partitioning among African savanna herbivores in North Cameroon: the importance of diet composition, food quality and body mass

  • H. H. de Iongh (a1), C. B. de Jong (a2), J. van Goethem (a1), E. Klop (a1), A. M. H. Brunsting (a2), P. E. Loth (a1) and H. H. T. Prins (a2)...


Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed.