Population regulation of African buffalo in the Mara–Serengeti ecosystem
Holly T. Dublin A and Joseph O. Ogutu B C D
+ Author Affiliations
- Author Affiliations
A ℅ IUCN ESARO, Wasaa Conservation Centre, PO Box 68200, Nairobi 00200, Kenya.
B University of Hohenheim, Institute of Crop Science, Biostatistics Unit, Fruwirthstrasse 23, 70599 Stuttgart, Germany.
C International Livestock Research Institute (ILRI), Old Naivasha Road, Box 30709, Nairobi 00100, Kenya.
D Corresponding author. Email: jogutu2007@gmail.com
Wildlife Research 42(5) 382-393 https://doi.org/10.1071/WR14205
Submitted: 5 October 2014 Accepted: 16 July 2015 Published: 31 August 2015
Journal Compilation © CSIRO Publishing 2015 Open Access CC BY-NC-ND
Abstract
Context: The processes regulating ungulate populations have been the focus of numerous studies. For the African buffalo (Syncerus caffer Sparrman) population inhabiting the Mara–Serengeti ecosystem, rinderpest was the primary regulatory factor up to the mid-1960s. Following reduction of rinderpest and buffalo population increase, interspecific competition for food, notably with cattle and wildebeest (Connochaetes taurinus Burchell), was thought to be the primary regulatory factor in the ecosystem.
Aims: We analysed buffalo population trends and the relationship between buffalo population growth and rainfall and density dependence in the Mara–Serengeti ecosystem and discuss the findings in the context of the key ecosystem processes governing buffalo population dynamics in African savannas, namely, food limitation, competition, predation, disease and land use changes.
Methods: We analysed buffalo population dynamics in the Mara–Serengeti ecosystem in relation to rainfall and density dependence feedback between 1984 and 2010.
Key results: Buffalo population growth was both significantly density-dependent and positively correlated with the dry season rainfall after, but not before, a severe drought in 1993. Buffalo numbers crashed by 48.6% in 1984–85 and by 76.1% in 1993–94 during severe droughts when food availability was lowest and competition with the more numerous cattle and wildebeest was highest.
Conclusions: Recovery of buffalo numbers to pre-drought levels took 8–9 years after the 1984–85 drought but was much slower, with buffaloes numbering merely 36% of their 1993 population (12 895 animals) 18 years after the 1993–94 drought despite intermittent periods of high rainfall, probably due to demographic and/or reproductive factors, heightened competition with livestock, land use changes in the adjoining pastoral ranches, lion predation and recurrent severe droughts.
Implications: Our findings demonstrate how food limitation caused by droughts associated with the hemispheric El Niño–Southern Oscillation can cause severe declines in and threaten the persistence of large ungulate populations. The findings also portray how density-dependent food limitation, competition, predation, land use changes and other factors can accentuate the effect of droughts and greatly prolong population recovery.
Additional keywords: climate change, competition, disease, food limitation, land use change, predation.
References
Anderson, E. C., Jago, M., Mlengeya, T., Timms, S., Payne, A., and Hirji, K. (1990). A serological survey of rinderpest antibody in wildlife and sheep and goats in northern Tanzania. Epidemiology and Infection 105, 203–214.| A serological survey of rinderpest antibody in wildlife and sheep and goats in northern Tanzania.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK3czks1SrtA%3D%3D&md5=1c257e3f7c97d21c909852f688922c11CAS | 2384143PubMed |
Arcese, P., Hando, J., and Campbell, K. (1995). Historical and present-day anti-poaching efforts in Serengeti. In ‘Serengeti II: Dynamics, Management, and Conservation of an Ecosystem’. (Eds A. R. E. Sinclair and P. Arcese.) pp. 506–533. (University of Chicago Press: Chicago, IL.)
Begon, M., Harper, J. L., and Townsend, C. R. (1990). ‘Ecology: Individuals, Populations and Communities.’ (Blackwell Science: Oxford, UK.)
Bell, R. (1970). The use of the herb layer by grazing ungulates in the Serengeti. In ‘Animal Populations in Relation to Their Food Resources’. (Ed. A. Watson.) pp. 111–124. (Blackwell: Oxford, UK.)
Bender, E. A., Case, T. J., and Gilpin, M. E. (1984). Perturbation experiments in community ecology: theory and practice. Ecology 65, 1–13.
| Perturbation experiments in community ecology: theory and practice.Crossref | GoogleScholarGoogle Scholar |
Bhola, N., Ogutu, J. O., Piepho, H.-P., Said, M. Y., Reid, R. S., Hobbs, N. T., and Olff, H. (2012a). Comparative changes in density and demography of large herbivores in the Maasai Mara Reserve and its surrounding human-dominated pastoral ranches in Kenya. Biodiversity and Conservation 21, 1509–1530.
| Comparative changes in density and demography of large herbivores in the Maasai Mara Reserve and its surrounding human-dominated pastoral ranches in Kenya.Crossref | GoogleScholarGoogle Scholar |
Bhola, N., Ogutu, J. O., Said, M. Y., Piepho, H.-P., and Olff, H. (2012b). The distribution of large herbivore hotspots in relation to environmental and anthropogenic correlates in the Mara region of Kenya. Journal of Animal Ecology 81, 1268–1287.
| The distribution of large herbivore hotspots in relation to environmental and anthropogenic correlates in the Mara region of Kenya.Crossref | GoogleScholarGoogle Scholar | 22640527PubMed |
Boutton, T. W., Tieszen, L. L., and Imbamba, S. K. (1988). Seasonal changes in the nutrient of east African grassland vegetation. African Journal of Ecology 26, 103–115.
| Seasonal changes in the nutrient of east African grassland vegetation.Crossref | GoogleScholarGoogle Scholar |
Braun, H. M. H. (1973). Primary production in the Serengeti: purpose, methods and some results of research. Annals of the University of Abidjan 6, 171–188.
Bronner, G. (1990). ‘Vegetation and Land Use in the Matthews Range Area, Samburu District.’ (Gebrüder Borntraeger Verlag: Berlin.)
Burnham, K. P., and Anderson, D. R. (2002). ‘Model Selection and Multi-Model Inference.’ 2nd edn. (Springer Verlag: New York, NY.)
Butt, B., Shortridge, A., and WinklerPrins, A. M. G. A. (2009). Pastoral herd management, drought coping strategies and cattle mobility in southern Kenya. Annals of the Association of American Geographers 99, 309–334.
| Pastoral herd management, drought coping strategies and cattle mobility in southern Kenya.Crossref | GoogleScholarGoogle Scholar |
Caron, A., Cross, P. C., and du Toi, J. T. (2003). Ecological implications of bovine tuberculosis in African buffalo herds. Ecological Applications 13, 1338–1345.
| Ecological implications of bovine tuberculosis in African buffalo herds.Crossref | GoogleScholarGoogle Scholar |
Coe, M. J., Cumming, D. H., and Phillipson, J. (1976). Biomass and production of large African herbivores in relation to rainfall and primary production. Oecologia 22, 341–354.
Connor, E., and Simberloff, D. (1979). The assembly of species: chance or competition. Ecology 60, 1132–1140.
| The assembly of species: chance or competition.Crossref | GoogleScholarGoogle Scholar |
Cross, P. C., Heisey, D. M., Bowers, J. A., Hay, C. T., Wolhuter, J., Buss, P., Hofmeyr, M., Miche, A. L., Bengis, R. G., Bird, T. L. F., du Toit, J. T., and Getz, M. W. (2009). Disease, predation and demography: assessing the impacts of bovine tuberculosis on African buffalo by monitoring at individual and population levels. Journal of Applied Ecology 46, 467–475.
| Disease, predation and demography: assessing the impacts of bovine tuberculosis on African buffalo by monitoring at individual and population levels.Crossref | GoogleScholarGoogle Scholar |
Darling, F. F. (1960). ‘An Ecological Reconnaissance of the Mara Plains in Kenya Colony.’ Wildlife Monograph no. 5, 41 pp. (The Wildlife Society: Washington, DC.)
Dennis, B., Ponciano, J. M., Subhash, R. L., Taper, M. L., and Staples, D. F. (2006). Estimating density dependence, process noise, and observation error. Ecological Monographs 76, 323–341.
| Estimating density dependence, process noise, and observation error.Crossref | GoogleScholarGoogle Scholar |
Diamond, J. M., and Gilpin, M. E. (1982). Examination of the null model of Connor and Simberloff for species co-occurrences on islands. Oecologia 52, 64–74.
| Examination of the null model of Connor and Simberloff for species co-occurrences on islands.Crossref | GoogleScholarGoogle Scholar |
Dobson, A. (1995). The ecology and epidemiology of rinderpest virus in the Serengeti and Ngorongoro conservation area. In ‘Serengeti II: Dynamics, Management, and Conservation of an Ecosystem’. (Eds A. R. E. Sinclair and P. Arcese.) pp. 485–505. (University of Chicago Press: Chicago, IL.)
Dublin, H. T. (1991). Dynamics of the Mara–Serengeti woodlands: an historical perspective. Forest Conservation History 35, 169–178.
Dublin, H. T. (1995). Vegetation dynamics in the Mara–Serengeti ecosystem: the role of elephants, fire, and other factors. In ‘Serengeti II: Dynamics, Management, and Conservation of an Ecosystem’. (Eds A. R. E. Sinclair and P. Arcese.) pp. 71–90. (University of Chicago Press, Chicago, IL.)
Dublin, H. T., and Douglas-Hamilton, I. (1987). Status and trends of elephants in the Serengeti-Mara ecosystem. African Journal of Ecology 25, 19–33.
| Status and trends of elephants in the Serengeti-Mara ecosystem.Crossref | GoogleScholarGoogle Scholar |
Dublin, H. T., Sinclair, A. R. E., Boutin, S., Anderson, E., Jago, M., and Arcese, P. (1990a). Does competition regulate ungulate populations: further evidence from Serengeti, Tanzania. Oecologia 82, 283–288.
| Does competition regulate ungulate populations: further evidence from Serengeti, Tanzania.Crossref | GoogleScholarGoogle Scholar |
Dublin, H. T., Sinclair, A. R. E., and McGlade, J. (1990b). Elephants and fire as causes of multiple stable states in the Mara–Serengeti woodlands. Journal of Animal Ecology 59, 1147–1164.
| Elephants and fire as causes of multiple stable states in the Mara–Serengeti woodlands.Crossref | GoogleScholarGoogle Scholar |
East, R. (1984). Rainfall, soil nutrient status and biomass of large African savanna mammals. African journal of Ecology 22, 245–270.
Estes, R. D., Atwood, J. L., and Estes, A. B. (2006). Downward trends in Ngorongoro Crater ungulate populations 1986–2005: conservation concerns and the need for ecological research. Biological Conservation 131, 106–120.
| Downward trends in Ngorongoro Crater ungulate populations 1986–2005: conservation concerns and the need for ecological research.Crossref | GoogleScholarGoogle Scholar |
Ferrar, A. A., and Walker, B. H. (1974). An analysis of herbivore/habitat relationships in Kyle National Park, Rhodesia. Journal of South African Wildlife Management Association 4, 137–147.
Field, C. R. (1968). The food habits of some wild ungulates in Uganda. PhD Thesis, University of Cambridge, UK.
Field, C. R. (1972). Food habits of some wild ungulates in Uganda by analysis of stomach contents. East African Wildlife Journal 10, 17.
| Food habits of some wild ungulates in Uganda by analysis of stomach contents.Crossref | GoogleScholarGoogle Scholar |
Field, C. R. (1976). Palatability factors and nutritive values of the food of buffaloes (Syncerus caffer) in Uganda. East African Wildlife Journal 14, 181–201.
| Palatability factors and nutritive values of the food of buffaloes (Syncerus caffer) in Uganda.Crossref | GoogleScholarGoogle Scholar |
Fritz, H., and Duncan, P. (1994). On the carrying capacity for large ungulates of African savanna ecosystems. Proceedings of the Royal Society of London B: Biological Sciences 256, 77–82.
| 1:STN:280:DyaK2c3nsl2qtQ%3D%3D&md5=e59927ebacf550eb0a7c5865a7ac99daCAS |
Georgiadis, N. J., Olwero, J. N., and Romañach, S. S. (2007). Savanna herbivore dynamics in a livestock-dominated landscape: I. Dependence on land use, rainfall, density, and time. Biological Conservation 137, 461–472.
Gilpin, M. E., and Diamond, J. M. (1982). Factors contributing to non randomness in species co-occurrences on islands. Oecologia 52, 75–84.
| Factors contributing to non randomness in species co-occurrences on islands.Crossref | GoogleScholarGoogle Scholar |
Gwynne, M. D., and Bell, R. H. V. (1968). Selection of vegetation components by grazing ungulates in the Serengeti National Park. Nature 220, 390–393.
| Selection of vegetation components by grazing ungulates in the Serengeti National Park.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaF1M%2FhsVKqtA%3D%3D&md5=bdf8ac4e29e7c2c243900d5980b783faCAS | 5684885PubMed |
Haas, L., Hofer, H., East, M., Wohlsein, P., Liess, B., and Barrett, T. (1996). Canine distemper virus infection in Serengeti spotted hyaenas. Veterinary Microbiology 49, 147–152.
| Canine distemper virus infection in Serengeti spotted hyaenas.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK2s%2FhsVyiug%3D%3D&md5=2f7bbfadc1c23b203bdb3a276e0d5e19CAS | 8861651PubMed |
Hilborn, R., Arcese, P., Borner, M., Hando, J., Hopcraft, G., Loibooki, M., Mduma, S., and Sinclair, A. R. E. (2006). Effective enforcement in a conservation area. Science 314, 1266.
| Effective enforcement in a conservation area.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28Xht1agt7vF&md5=9b6a35a4107d05a4552c716476fe119fCAS | 17124316PubMed |
Jarman, P. J. (1971). Diets of large mammals in the woodlands around Lake Kariba, Rhodesia. Oecologia 8, 157–178.
| Diets of large mammals in the woodlands around Lake Kariba, Rhodesia.Crossref | GoogleScholarGoogle Scholar |
Jarman, P. J. (1972). Seasonal distribution of large mammal populations in the unflooded middle Zambezi valley. Journal of Applied Ecology 9, 283–299.
| Seasonal distribution of large mammal populations in the unflooded middle Zambezi valley.Crossref | GoogleScholarGoogle Scholar |
Kanga, E. M., Ogutu, J. O., Olff, H., and Santema, P. (2011). Population trend and distribution of the vulnerable common hippopotamus Hippopotamus amphibius in the Mara Region of Kenya. Oryx 45, 20–27.
| Population trend and distribution of the vulnerable common hippopotamus Hippopotamus amphibius in the Mara Region of Kenya.Crossref | GoogleScholarGoogle Scholar |
Keddy, P. A. (1989). ‘Competition.’ (Chapman: London.)
Kenward, M. G., and Roger, J. H. (1997). Small sample inference for fixed effects from restricted maximum likelihood. Biometrics 53, 983–997.
| Small sample inference for fixed effects from restricted maximum likelihood.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK2svntVGitw%3D%3D&md5=bfe664687c7909a6b15d97dee7b2847aCAS | 9333350PubMed |
Kock, R., Chalmers, W. S. K., Mwanzia, J., Chillingworth, C., Wambua, J., Coleman, P. G., and Baxendale, W. (1998). Canine distemper antibodies in lions of the Masai Mara. The Veterinary Record 142, 662–665.
| Canine distemper antibodies in lions of the Masai Mara.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK1czjsFWjsw%3D%3D&md5=3111a0d73f6393faf517198cc85382ceCAS | 9670445PubMed |
Kock, R. A., Wambua, J. M., Mwanzia, J., Wamwayi, H., Ndungu, E. K., Barrett, T., Kock, N. D., and Rossiter, P. B. (1999). Rinderpest epidemic in wild ruminants in Kenya 1993–1997. The Veterinary Record 145, 275–283.
| Rinderpest epidemic in wild ruminants in Kenya 1993–1997.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BD3c%2FksVWgug%3D%3D&md5=038921094d38f23579bc7ee71880c93bCAS | 10579537PubMed |
Lamprey, R. H., and Reid, R. S. (2004). Expansion of human settlement in Kenya’s Maasai Mara: what future for pastoralism and wildlife? Journal of Biogeography 31, 997–1032.
Lamprey, H. K. (1963). Ecological separation of large mammal species in Tarangire Game Reserve, Tanzania. East African Wildlife Journal 1, 63–92.
| Ecological separation of large mammal species in Tarangire Game Reserve, Tanzania.Crossref | GoogleScholarGoogle Scholar |
Lande, R. (1987). Extinction thresholds in demographic models of territorial populations. American Naturalist 130, 624–635.
| Extinction thresholds in demographic models of territorial populations.Crossref | GoogleScholarGoogle Scholar |
Lande, R. (1993). Risks of population extinction from demographic and environmental stochasticity and random catastrophes. American Naturalist 142, 911–927.
| Risks of population extinction from demographic and environmental stochasticity and random catastrophes.Crossref | GoogleScholarGoogle Scholar |
Law, R., and Watkinson, A. R. (1989). Competition. In ‘Ecological Concepts’. (Eds J. M. Cherret and R. Law.) (Blackwell Scientific Publications: Oxford, UK.)
Marshal, J. P., Owen-Smith, N., Whyte, I. J., and Stenseth, N. C. (2011). The role of El Niño–Southern Oscillation in the dynamics of a savanna large herbivore population. Oikos 120, 1175–1182.
| The role of El Niño–Southern Oscillation in the dynamics of a savanna large herbivore population.Crossref | GoogleScholarGoogle Scholar |
McNaughton, S. J., and Tarrants, J. L. (1983). Grass leaf silicification: natural selection for and inducible defense against herbivores. Proceedings of the National Academy of Sciences of the United States of America 80, 790–791.
| Grass leaf silicification: natural selection for and inducible defense against herbivores.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC3cnivVehsQ%3D%3D&md5=d61e87358ecf98f6902fb71d6be93313CAS | 16578767PubMed |
Metzger, K. L., Sinclair, A. R. E., Hilborn, R., Hopcraft, J. G. C., and Mduma, S. A. R. (2010). Evaluating the protection of wildlife in parks: the case of African buffalo in Serengeti. Biodiversity and Conservation 19, 3431–3444.
| Evaluating the protection of wildlife in parks: the case of African buffalo in Serengeti.Crossref | GoogleScholarGoogle Scholar |
Mills, M. G. L., Biggs, H. C., and Whyte, I. J. (1995). The relationship between rainfall, lion predation and population trends in African herbivores. Wildlife Research 22, 75–88.
| The relationship between rainfall, lion predation and population trends in African herbivores.Crossref | GoogleScholarGoogle Scholar |
Mduma, S. A., Sinclair, A. R. E., and Hilborn, R. (1999). Food regulates the Serengeti wildebeest: a 40-year record. Journal of Animal Ecology 68, 1101–1122.
Norton-Griffiths, M. (1978). ‘Counting Animals.’ (African Wildlife Foundation: Nairobi, Kenya.)
Norton-Griffiths, M., Herlocker, D., and Pennycuick, L. (1975). Patterns of rainfall in the Serengeti ecosystem. East African Wildlife Journal 13, 347–374.
| Patterns of rainfall in the Serengeti ecosystem.Crossref | GoogleScholarGoogle Scholar |
Obara, G. A. O. (1999). The changes in the woody vegetation cover of Masai Mara National Reserve, Kenya. MSc Thesis, University of Kent, UK.
Ogutu, J. O., and Owen-Smith, N. (2003). ENSO, rainfall and temperature influences on extreme population declines among African savanna ungulates. Ecology Letters 6, 412–419.
| ENSO, rainfall and temperature influences on extreme population declines among African savanna ungulates.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J. O., Bhola, N., and Reid, R. S. (2005). The effects of pastoralism and protection on the density and distribution of carnivores and their prey in the Mara ecosystem of Kenya. Journal of Zoology 265, 281–293.
| The effects of pastoralism and protection on the density and distribution of carnivores and their prey in the Mara ecosystem of Kenya.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J. O., Piepho, H.-P., Dublin, H. T., Bhola, N., and Reid, R. S. (2008). El Niño–Southern Oscillation, rainfall, temperature and normalized difference vegetation index fluctuations in the Mara–Serengeti ecosystem. African Journal of Ecology 46, 132–143.
| El Niño–Southern Oscillation, rainfall, temperature and normalized difference vegetation index fluctuations in the Mara–Serengeti ecosystem.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J. O., Piepho, H.-P., Dublin, H. T., Bhola, N., and Reid, R. S. (2009). Dynamics of Mara–Serengeti ungulates in relation to land use changes. Journal of Zoology 278, 1–14.
| Dynamics of Mara–Serengeti ungulates in relation to land use changes.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J. O., Owen‐Smith, N., Piepho, H.-P., and Said, M. Y. (2011). Continuing wildlife population declines and range contraction in the Mara region of Kenya during 1977–2009. Journal of Zoology 285, 99–109.
| Continuing wildlife population declines and range contraction in the Mara region of Kenya during 1977–2009.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J. O., Owen-Smith, N., Piepho, H.-P., Kuloba, B., and Endebe, J. (2012). Dynamics of ungulates in relation to climatic and land use changes in an insularized African savanna ecosystem. Biodiversity and Conservation 21, 1033–1053.
| Dynamics of ungulates in relation to climatic and land use changes in an insularized African savanna ecosystem.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J. O., Owen-Smith, N., Piepho, H.-P., Said, M. Y., Kifugo, S., and Reid, R. S. (2013). Changing wildlife populations in Nairobi National Park and adjoining Athi-Kaputiei Plains: collapse of the migratory wildebeest. The Open Conservation Biology Journal 7, 11–26.
| Changing wildlife populations in Nairobi National Park and adjoining Athi-Kaputiei Plains: collapse of the migratory wildebeest.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J. O., Piepho, H.-P., Said, M. Y., and Kifugo, S. (2014). Herbivore dynamics and range contraction in Kajiado: climate and land use changes, population pressures, governance, policy and human–wildlife conflicts. Open Journal of Ecology 7, 9–31.
| Herbivore dynamics and range contraction in Kajiado: climate and land use changes, population pressures, governance, policy and human–wildlife conflicts.Crossref | GoogleScholarGoogle Scholar |
Ogutu, J., Piepho, H.-P., Said, M., Ojwang, G., Kifugo, S. C., Wargute, P. W., and Njino, L. W. (2015). Extreme wildlife declines and concurrent increase in livestock numbers in Kenya rangelands. PLoS One, , .
Ottichilo, W. K., de Leeuw, J., Skidmore, A. K., Prins, H. H. T., and Said, M. Y. (2000). Population trends of large non-migratory wild herbivores and livestock in Masai Mara ecosystem, Kenya, between 1977 and 1997. African Journal of Ecology 38, 202–216.
| Population trends of large non-migratory wild herbivores and livestock in Masai Mara ecosystem, Kenya, between 1977 and 1997.Crossref | GoogleScholarGoogle Scholar |
Owen-Smith, N., and Mills, M. G. L. (2006). Manifold interactive influences on the population dynamics of a multispecies ungulate assemblage. Ecological Monographs 76, 73–92.
| Manifold interactive influences on the population dynamics of a multispecies ungulate assemblage.Crossref | GoogleScholarGoogle Scholar |
Owen-Smith, N., and Mills, M. G. L. (2008). Shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator–prey web. Ecology 89, 1120–1133.
| Shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator–prey web.Crossref | GoogleScholarGoogle Scholar | 18481536PubMed |
Pienaar, U. V. (1969). Observations on developmental biology, growth and some aspects of the population biology of African buffalo (Syncerus caffer Sparrman) in the Kruger National Park. Koedoe 12, 29–52.
| Observations on developmental biology, growth and some aspects of the population biology of African buffalo (Syncerus caffer Sparrman) in the Kruger National Park.Crossref | GoogleScholarGoogle Scholar |
Plowright, W. (1982). The effect of rinderpest and rinderpest control on wildlife in Africa. Symposium of the Zoological Society of London 50, 1–28.
Roelke-Parker, M. E., Munson, L., Packer, C., Kock, R., Cleaveland, S., Carpenter, M., Obrien, S. J., Pospischil, A., Hofmann-Lehmann, R., Lutz, H., Mwamengele, G. L. M., Mgasa, M. N., Machange, G. A., Summers, B. A., and Appel, M. J. G. (1996). A canine distemper virus epidemic in Serengeti lions (Panthera leo). Nature 379, 441–445.
| A canine distemper virus epidemic in Serengeti lions (Panthera leo).Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK28XovVagsg%3D%3D&md5=d896645f378d61ccf74932db581fde43CAS | 8559247PubMed |
Ruppert, D., Wand, M. P., and Carroll, R. J. (2003). ‘Semiparametric Regression.’ (Cambridge University Press: Cambridge, UK.)
Ryan, S. J., Knechtel, C. V., and Getz, W. M. (2007). Ecological cues, gestation length, and birth timing in African buffalo (Syncerus caffer). Behavioural Ecology 18, 635–644.
| Ecological cues, gestation length, and birth timing in African buffalo (Syncerus caffer).Crossref | GoogleScholarGoogle Scholar |
Saltz, D., and Rubenstein, D. I. (1995). Population dynamics of a reintroduced Asiatic wild ass (Equus hemionus) herd. Ecological Applications 5, 327–335.
| Population dynamics of a reintroduced Asiatic wild ass (Equus hemionus) herd.Crossref | GoogleScholarGoogle Scholar |
SAS Institute Inc. (2014). ‘SAS System for Windows.’ Version 9.4. (SAS Institute Inc.: Carey, NC.)
Schaller, G. B. (1972). ‘The Serengeti Lion: a Study of Predator-Prey Relations.’ (Chicago University Press: Chicago, IL.)
Serneels, S., Said, M. Y., and Lambin, E. F. (2001). Land cover changes around a major east African wildlife reserve: the Mara ecosystem (Kenya). International Journal of Remote Sensing 22, 3397–3420.
| Land cover changes around a major east African wildlife reserve: the Mara ecosystem (Kenya).Crossref | GoogleScholarGoogle Scholar |
Simberloff, D. (1983). Competition theory, hypothesis testing, and other community ecological buzzwords. American Naturalist 122, 626–635.
| Competition theory, hypothesis testing, and other community ecological buzzwords.Crossref | GoogleScholarGoogle Scholar |
Simberloff, D. (1984). Properties of coexisting bird species in two archipelagos. In ‘Ecological Communities: Conceptual Issues and the Evidence’. (Eds D. R. Strong, D. Simberloff, L. G. Abele and A. B. Thistle.) pp. 234–253. (Princeton University Press: Princeton, NJ.)
Sinclair, A. R. E. (1973a). Population increases of buffalo and wildebeest in the Serengeti. East African Wildlife Journal 11, 93–107.
| Population increases of buffalo and wildebeest in the Serengeti.Crossref | GoogleScholarGoogle Scholar |
Sinclair, A. R. E. (1973b). Regulation and population models for a tropical ruminant. East African Wildlife Journal 11, 307–316.
| Regulation and population models for a tropical ruminant.Crossref | GoogleScholarGoogle Scholar |
Sinclair, A. R. E. (1974a). The natural regulation of buffalo populations in East Africa. III. Population trends and mortality. East African Wildlife Journal 12, 185–200.
| The natural regulation of buffalo populations in East Africa. III. Population trends and mortality.Crossref | GoogleScholarGoogle Scholar |
Sinclair, A. R. E. (1974b). The natural regulation of buffalo populations in East Africa. IV. The food supply as a regulating factor, and competition. East African Wildlife Journal 12, 291–311.
| The natural regulation of buffalo populations in East Africa. IV. The food supply as a regulating factor, and competition.Crossref | GoogleScholarGoogle Scholar |
Sinclair, A. R. E. (1977). ‘The African Buffalo.’ (University of Chicago Press: Chicago, IL.)
Sinclair, A. R. E. (1979). Dynamics of the Serengeti ecosystem: process and pattern. In ‘Serengeti: Dynamics of an Ecosystem’. (Eds A. R. E. Sinclair and M. Norton-Griffiths.) pp. 1–30. (University of Chicago Press: Chicago, IL.)
Sinclair, A. R. E. (1983). The adaptations of African ungulates and their effects on community function. In ‘Ecosystems of the World 13: Tropical Savannas.’ (Ed. F. Bourliere.) pp. 401–426. (Elsevier: Amsterdam.)
Sinclair, A. R. E. (1985). Does interspecific competition or predation shape the African ungulate community? Journal of Animal Ecology 54, 899–918.
| Does interspecific competition or predation shape the African ungulate community?Crossref | GoogleScholarGoogle Scholar |
Sinclair, A. R. E., and Arcese, P. (1995). ‘Serengeti II: Dynamics, Management, and Conservation of an Ecosystem.’ (The University of Chicago Press: Chicago, IL.)
Sinclair, A. R. E., and Norton-Griffiths, M. (1979). ‘Serengeti: Dynamics of an Ecosystem.’ (University of Chicago Press: Chicago, IL.)
Sinclair, A. R. E., and Norton-Griffiths, M. (1982). Does competition or facilitation regulate migrant ungulate populations in the Serengeti? A test of hypotheses. Oecologia 53, 364–369.
| Does competition or facilitation regulate migrant ungulate populations in the Serengeti? A test of hypotheses.Crossref | GoogleScholarGoogle Scholar |
Sinclair, A. R. E., Dublin, H., and Borner, M. (1985). Population regulation of Serengeti wildebeest – a test of the food hypothesis. Oecologia 65, 266–268.
| Population regulation of Serengeti wildebeest – a test of the food hypothesis.Crossref | GoogleScholarGoogle Scholar |
Sitati, N. W. (2003). Human–elephant conflicts in the Maasai Mara dispersal areas of Transmara district. PhD Thesis, University of Kent, UK.
Stewart, D. R. M., and Talbot, L. M. (1962). Census of wildlife on the Serengeti, Mara and Loita plains. East African Agriculture and Forestry Journal 28, 58–60.
Talbot, L. M., and Stewart, D. R. M. (1964). First wildlife census of the entire Mara–Serengeti region, East Africa. The Journal of Wildlife Management 28, 815–827.
| First wildlife census of the entire Mara–Serengeti region, East Africa.Crossref | GoogleScholarGoogle Scholar |
Thouless, C. R. (1992). ‘Samburu and Laikipia Elephant Count 1992.’ (Kenya Wildlife Service Laikipia Elephant Project: Nairobi, Kenya.)
TWCM (1992). ‘Census of Buffalo and Elephant in the Serengeti Ecosystem.’ (Tanzania Wildlife Conservation Monitoring: Arusha, Tanzania.)
TWCM (1999). ‘Total Count of Buffalo and Elephant in the Serengeti Ecosystem, February/March 1998.’ (TWCM/FZS/EU: Arusha, Tanzania.)
Walker, B. H., Emslie, R. H., Owen-Smith, R. N., and Scholes, R. J. (1987). To cull or not to cull: lessons from a southern African drought. Journal of Applied Ecology 24, 381–401.
| To cull or not to cull: lessons from a southern African drought.Crossref | GoogleScholarGoogle Scholar |
