References
Adler, P.B. (2013). Testing the storage effect with long-term observational data. In Temporal dynamics and ecological processes(eds Kelly, C.K., Bowler, M.G., Fox, G.A.). Cambridge University Press, Cambridge, UK, pp. 82-101.
Adler, P.B. & Drake, J.M. (2008). Environmental variation, stochastic extinction, and competitive coexistence. Am. Nat. , 172, E186-E195.
Adler, P.B., Ellner, S.P. & Levine, J.M. (2010). Coexistence of perennial plants: an embarrassment of niches. Ecol. Lett. , 13, 1019-1029.
Adler, P.B., HilleRisLambers, J. & Levine, J.M. (2007). A niche for neutrality. Ecol. Lett. , 10, 95-104.
Ambrose, S.H. (1991). Effects of diet, climate and physiology on nitrogen isotope abundances in terrestrial foodwebs. J. Archaeol. Sci. , 18, 293–317.
Aranibar, J.N., Anderson, I.C., Epstein, H.E., Feral, C.J.W., Swap, R.J., Ramontsho, J. et al. (2008). Nitrogen isotope composition of soils, C3 and C4 plants along land use gradients in southern Africa.J. Arid Environ. , 72, 326-337.
Arsenault, R. & Owen-Smith, N. (2008). Resource partitioning by grass height among grazing ungulates does not follow body size relation.Oikos , 117, 1711-1717.
Arsenault, R. & Owen-Smith, N. (2011). Competition and coexistence among short-grass grazers in the Hluhluwe-iMfolozi Park, South Africa.Can. J. Zool. , 89, 901-908.
Barabás, G., D’Andrea, R. & Stump, S.M. (2018). Chesson’s coexistence theory. Ecol. Monogr. , 88, 277-303.
Bell, R.H.V. (1971). A grazing ecosystem in the Serengeti. Sci. Am. , 225, 86-93.
Broekman, M.J.E., Muller-Landau, H.C., Visser, M.D., Jongejans, E., Wright, S.J. & de Kroon, H. (2019). Signs of stabilisation and stable coexistence. Ecol. Lett. , 22, 1957-1975.
Burnham, K.P. & Anderson, D.R. (2001). Kullback-Leibler information as a basis for strong inference in ecological studies. Wildl. Res. , 28, 111-119.
Carmel, Y., Suprunenko, Y.F., Kunin, W.E., Kent, R., Belmaker, J., Bar-Massada, A. et al. (2017). Using exclusion rate to unify niche and neutral perspectives on coexistence. Oikos , 126, 1451-1458.
Cerling, T.E. & Harris, J.M. (1999). Carbon isotope fractionation between diet and bioapatite in ungulate mammals and implications for ecological and paleoecological studies. Oecologia , 120, 347–363.
Chesson, P. (2000). Mechanisms of maintenance of species diversity.Annu. Rev. Ecol. Syst. , 31, 343-366.
Clauss, M., Kaiser, T.M. & Hummel, J. (2008). The morphophysiological adaptations of browsing and grazing mammals. In The ecology of browsing and grazing (eds Gordon, I.J., Prins, H.H.T.). Springer, Heidelberg, pp. 149-178.
Clauss, M., Müller, D.W.H. & Codron, D. (2020). Within-niche pace of life acceleration as a fundamental evolutionary principle: a mammal pilot test case. Evol. Ecol. Res. , 20, online at http://www.evolutionary-ecology.com/forthcoming.html.
Codron, D., Brink, J.S., Rossouw, L., Clauss, M., Codron, J., Lee-Thorp, J.A. et al. (2008). Functional differentiation of African grazing ruminants: an example of specialized adaptations to very small changes in diet. Biol. J. Linn. Soc. , 94, 755–764.
Codron, D. & Clauss, M. (2010). Rumen physiology constrains diet niche: linking digestive physiology and food selection across wild ruminant species. Can. J. Zool. , 88, 1129-1138.
Codron, D. & Codron, J. (2009). Reliability of δ13C and δ15N in faeces for reconstructing savanna herbivore diet. Mamm Biol , 74, 36–48.
Codron, D., Codron, J., Lee-Thorp, J.A., Sponheimer, M. & de Ruiter, D. (2005). Animal diets in the Waterberg based on stable isotopic composition of faeces. S. Afr. J. Wildl. Res. , 35, 43-52.
Codron, D., Hofmann, R.R. & Clauss, M. (2019). Morphological and physiological adaptations for browsing and grazing. In The ecology of browsing and grazing II (eds Gordon, I.J., Prins, H.H.T.). Springer, Berlin, Germany, pp. 81-125.
Codron, J., Lee-Thorp, J.A., Sponheimer, M. & Codron, D. (2013). Plant stable isotope composition across habitat gradients in a semi-arid savanna: implications for environmental reconstruction. J Quat Sci , 28, 301-310.
Cole, L.C. (1954). The Population Consequences of Life History Phenomena. Quart. Rev. Biol. , 29, 103-137.
Cromsigt, J.P.G.M. & Olff, H. (2006). Resource partitioning among savanna grazers mediated by local heterogeneity: an experimental approach. Ecology , 87, 1532-1541.
de Iongh, H.H., de Jong, C.B., van Goethem, J., Klop, E., Brunsting, A.M.H., Loth, P.E. et al. (2011). Resource partitioning among African savanna herbivores in North Cameroon: the importance of diet composition, food quality and body mass. J. Trop. Ecol. , 27, 503-513.
Demment, M.W. & Van Soest, P.J. (1985). A nutritional explanation for body-size patterns of ruminant and nonruminant herbivores. Am. Nat. , 125, 641-672.
du Toit, J.T. (1990). Feeding-height stratification among African browsing ruminants. Afr. J. Ecol. , 28, 55-61.
Duncan, R.P., Forsyth, D.M. & Hone, J. (2007). Testing the metabolic theory of ecology: allometric scaling exponents in mammals.Ecology , 88, 324-333.
Ellner, S.P., Snyder, R.E. & Adler, P.B. (2016). How to quantify the temporal storage effect using simulations instead of math. Ecol. Lett. , 19, 1333-1342.
Ellner, S.P., Snyder, R.E., Adler, P.B. & Hooker, G. (2019). An expanded modern coexistence theory for empirical applications.Ecol. Lett. , 22, 3-18.
Foose, T.J. (1982). Trophic strategies of ruminant versus nonruminant ungulates. University of Chicago Chicago.
Fraser, D. & Rybczynski, N. (2014). Complexity of ruminant masticatory evolution. J. Morphol. , 275, 1093-1102.
Fraser, D. & Theodor, J.M. (2011). Comparing ungulate dietary proxies using discriminant function analysis. J. Morphol. , 272, 1513-1526.
Fryxell, J.M., Sinclair, A.R.E. & Caughley, G. (2014). Wildlife Ecology, Conservation, and Management, third edition . Wiley-Blackwell, Malden.
Gordon, I.J. & Illius, A.W. (1989). Resource partitioning by ungulates on the Isle of Rhum. Oecologia , 79, 383-389.
Gordon, I.J. & Prins, H.H.T. (2019). The ecology of browsing and grazing II . Springer, Berlin, Germany.
Grainger, T.N., Levine, J.M. & Gilbert, B. (2019). The invasion criterion: a common currency for ecological research. Trends Ecol. Evol. , 34, 925-935.
HilleRisLambers, J., Adler, P.B., Harpole, W.S., Levine, J.M. & Mayfield, M.M. (2012). Rethinking community assembly through the lens of coexistence theory. Ann. Rev. Ecol. Evol. Syst. , 43, 227-248.
Hopcraft, J.G.C., Anderson, T.M., Pérez-Vila, S., Mayemba, E. & Olff, H. (2012). Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers. J. Anim. Ecol. , 81, 201-213.
Hubbell, S.P. (2001). The unified neutral theory of biodiversity and biogeography . Princeton University Press, Princeton, MA.
Hutchinson, G.E. (1961). The paradox of the plankton. Am. Nat. , 95, 137-145.
Jackson, A.L., Inger, R., Parnell, A.C. & Bearhop, S. (2011). Comparing isotopic niche widths among and within communities: SIBER – Stable Isotope Bayesian Ellipses in R. J. Anim. Ecol. , 80, 595-602.
Janis, C.M. & Ehrhardt, D. (1988). Correlation of relative muzzle width and relative incisor width with dietary preference in ungulates.Zool. J. Linn. Soc. , 92, 267-284.
Kartzinel, T.R., Chen, P.A., Coverdale, T.C., Erickson, D.L., Kress, W.J., Kuzmina, M.L. et al. (2015). DNA metabarcoding illuminates dietary niche partitioning by African large herbivores. Proc. Natl. Acad. Sci. U.S.A. , 112, 8019-8024.
Kleynhans, E.J., Jolles, A.E., Bos, M.R.E. & Olff, H. (2011). Resource partitioning along multiple niche dimensions in differently sized African savanna grazers. Oikos , 120, 591-600.
Kuznetsova, A., Brockhoff, P.B. & Christensen, R.H.B. (2017). lmerTest Package: tests in linear mixed effects models. J. Stat. Soft. , 82, 1-26.
Lande, R. (1993). Risks of population extinction from demographic and environmental stochasticity and random catastrophes. Am. Nat. , 142, 911-927.
Letten, A.D., Ke, P.-J. & Fukami, T. (2017). Linking modern coexistence theory and contemporary niche theory. Ecol. Monogr. , 87, 161-177.
Macandza, V.A., Owen-Smith, N. & Cain, I.I.I.J.W. (2012a). Dynamic spatial partitioning and coexistence among tall grass grazers in an African savanna. Oikos , 121, 891-898.
Macandza, V.A., Owen-Smith, N. & Cain, J.W. (2012b). Habitat and resource partitioning between abundant and relatively rare grazing ungulates. J. Zool. , 287, 175-185.
MacArthur, R.H. & Levins, R. (1967). The limiting simlarity, convergence, and divergence of coexisting species. Am. Nat. , 101, 377-385.
Malindie, S., Buschke, F. & Codron, D. (2020). Carbon isotope differences between grassland and savanna herbivores reveal environmentally driven rather than phylogenetically conserved niches.J. Zool. , doi:10.1111/jzo.12762.
May, R.M. (1973). Stability and complexity in model ecosystems . Princeton University Press, Princeton.
Murray, M.G. & Illius, A.W. (2000). Vegetation modification and resource competition in grazing ungulates. Oikos , 89, 501-508.
Newsome, S.D., Martinez del Rio, C., Bearhop, S. & Phillips, D.L. (2007). A niche for isotopic ecology. Front. Ecol. Environ. , 5, 429-436.
Owen-Smith, N. (2006). Demographic determination of the shape of density dependence for three African ungulate populations. Ecol. Monogr. , 76, 93-109.
Pande, J., Fung, T., Chisholm, R. & Shnerb, N.M. Mean growth rate when rare is not a reliable metric for persistence of species. Ecol. Lett. , n/a.
Pansu, J., Guyton, J.A., Potter, A.B., Atkins, J.L., Daskin, J.H., Wursten, B. et al. (2019). Trophic ecology of large herbivores in a reassembling African ecosystem. J. Ecol. , 107, 1355-1376.
Pianka, E.R. (1973). The structure of lizard communities. Annu. Rev. Ecol. Syst. , 4, 53-74.
Prins, H.H.T., De Boer, W.F., Van Owveren, H., Correia, A.U., Mafuca, J. & Olff, H. (2006). Co-existence and niche segregation of three small bovid species in southern Mozambique. Afr. J. Ecol. , 44.
Prins, H.H.T. & Olff, H. (1998). Species-richness of African grazer assemblages: towards a functional explanation. In Dynamics of Tropical Communities (eds Newbery, D.M., Prins, H.H.T., Brown, N.). Blackwell Science, Oxford, pp. 449-490.
R_Core_Team. (2015). R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL http://www.R-project.org/.
Robbins, C., Felicetti, L. & Sponheimer, M. (2005). The effect of dietary protein quality on nitrogen isotope discrimination in mammals and birds. Oecologia , 144, 534-540.
Rosenzweig, M.L. & McCord, R.D. (1991). Incumbent replacement: evidence for long-term evolutionary progress. Ecology and Evolutionary Biology , 17, 202-213.
Saavedra, S., Rohr, R.P., Bascompte, J., Godoy, O., Kraft, N.J.B. & Levine, J.M. (2017). A structural approach for understanding multispecies coexistence. Ecol. Monogr. , 87, 470-486.
Scheffer, M. & van Nes, E.H. (2006). Self-organized similarity, the evolutionary emergence of groups of similar species. Proc. Natl. Acad. Sci. U.S.A. , 103, 6230-6235.
Schluter, D. (2000). The ecology of adaptive radiation . Oxford University Press, Oxford.
Schulze, E.-D., Williams, R.J., Farquhar, G.D., Schulze, W., Langridge, J., Miller, J.M. et al. (1998). Carbon and nitrogen isotope discrimination and nitrogen nutrition of trees along a rainfall gradient in northern Australia. Aust. J. Plant Physiol. , 25, 413-425.
Sinclair, A.R.E. (2003). Mammal population regulation, keystone processes and ecosystem dynamics. Philos. Trans. R. Soc. Lond. B , 358, 1729-1740.
Sinclair, A.R.E., Mduma, S. & Brashares, J.S. (2003). Patterns of predation in a diverse predator–prey system. Nature , 425, 288-290.
Tieszen, L., Hein, D., Qvortrup, D., Troughton, J. & Imbamba, S. (1979). Use of δ13C values to determine vegetation selectivity in east African herbivores. Oecologia , 37, 351-359.
Van Soest, P.J. (1994). Nutritional ecology of the ruminant . second edn. Comstock, Ithaca, New York.
Vergnon, R., van Nes, E.H. & Scheffer, M. (2012). Emergent neutrality leads to multimodal species abundance distributions. Nature Comm. , 3, 663.
Wainwright, C.E., HilleRisLambers, J., Lai, H.R., Loy, X. & Mayfield, M.M. (2019). Distinct responses of niche and fitness differences to water availability underlie variable coexistence outcomes in semi-arid annual plant communities. J. Ecol. , 107, 293-306.
Zerbe, P., Clauss, M., Codron, D., Lackey, L.B., Rensch, E., Streich, W.J. et al. (2012). Reproductive seasonality in captive wild ruminants: implications for biogeographical adaptation, photoperiodic control, and life history. Biol Rev (Camb) , 87, 965-990.
Table 1. Descriptive statistics summarizing mean (with standard deviations in parentheses) isotopic compositions, isotopic niche widths (standard ellipse areas, SEAc ), and average population sizes (N ) of grassland herbivores.