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Ginsberg, J. R., & Rubenstein, D. I. (1990). Sperm competiton and variation in zebra mating behaviour. Behav. Ecol. Sociobiol., 26(6), 427–434.
Abstract: Data are presented on the breeding behavior of two zebra species to test whether intra- and interspecific variation in male reproductive behavior and physiology are correlated with differences in female promiscuity. In one species, plains zebra (Equus burchelli) females live in closed membership single male groups and mate monandrously. In the other species, the Grevy's zebra (E. grevyi) females live in groups whose membership is much more temporary. Typically, associations with individual males are brief and mating is polyandrous. However, some females – those having just given birth – reside with one male for long periods, mating monandrously. These differences in female mating behavior generate variability in the potential for sperm competition. We show that behavioral differences in male investment in reproductive activities correlate with the potential for sperm competition. When mating with promiscuous mares, Grevy's zebra stallions made a greater investment in reproductive behavior (calling, mounting, ejaculations) than did stallions of either species when mating with monandrous females. The evolution of large testes size in the Grevy's zebra, when compared to the congeneric plains zebra, horse, and mountain zebra, allows for this increased investment.
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Rubenstein, D. I.,. (1986). Ecology and sociality in horses and zebras. In D. I. Rubenstein, & R. W. Wrangham (Eds.), Ecological Aspects of Social Evolution (pp. 282–302). Princeton, NJ.: Princeton University Press.
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Rubenstein, D. I. (1989). Life history and social organization in arid adapted ungulates. J. Arid. Environ., 17, 145–156.
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Rubenstein, D. I. (1994). The ecology of female social behaviour in horses, zebras and asses. In P. J. Jarman, & A. R. (Eds.), Animal Societies (pp. 13–28). Kyoto University Press.
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Rubenstein, D. (2012). Wild Equid Movements: Impacts of Habitat Quality, Predation Pressure and Leadership. In K. Krueger (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Animals spend much of their lives on the move searching for food, shelter and mates. As long-legged, large bodied species, equids are well suited for traveling long distances. Understanding where free-ranging equids go, why they choose particular locations, some near and some far, and what impact such movements have on patterns of sociality, demography and human livelihoods are only now coming into focus. This talk will explore how bottom-up factors, such as the abundance and distribution of key resources, and top-down factors, such as the type and location of predators, interact to shape overall movement patterns of equids, how leadership roles develop and what outcomes such movements are likely to have on human activities.
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Rubenstein, D. (2015). Social Networks: Linking Form with Function in Equid Societies. In K. Krueger (Ed.), Proceedings of the 3. International Equine Science Meeting. Proc. 3. Int. Equine. Sci. Mtg, in prep. Wald: Xenophon Publishing.
Abstract: Animal societies develop from interactions and relationships that occur among individuals within populations. The fundamental tenet of behavioral ecology is that ecological factors shape behavior and determine the distribution and associations of individuals on landscapes. As a result, different social systems emerge in different habitats and under different environmental conditions. Since characterizing social systems depends on time and motion studies of individual actions and interactions that are often bilateral, such characterizations are often coarse-grained. If social relationships can be characterized using social networks, however, seemingly similar social organizations often reveal informative differences in terms of deep structure. Thus social network theory should be able to provide insights in to the connections between social form and function. This talk will explore how the network structures of horses, zebras and asses can provide novel insights into the functioning of animal societies with respect to the spread of memes, genes and diseases.
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Sundaresan, S. R., Fischhoff, I. R., Dushoff, J., & Rubenstein, D. I. (2007). Network metrics reveal differences in social organization between two fission-fusion species, Grevy's zebra and onager. Oecologia, 151(1), 140–149.
Abstract: For species in which group membership frequently changes, it has been a challenge to characterize variation in individual interactions and social structure. Quantifying this variation is necessary to test hypotheses about ecological determinants of social patterns and to make predictions about how group dynamics affect the development of cooperative relationships and transmission processes. Network models have recently become popular for analyzing individual contacts within a population context. We use network metrics to compare populations of Grevy's zebra (Equus grevyi) and onagers (Equus hemionus khur). These closely related equids, previously described as having the same social system, inhabit environments differing in the distribution of food, water, and predators. Grevy's zebra and onagers are one example of many sets of coarsely similar fission-fusion species and populations, observed elsewhere in other ungulates, primates, and cetaceans. Our analysis of the population association networks reveals contrasts consistent with their distinctive environments. Grevy's zebra individuals are more selective in their association choices. Grevy's zebra form stable cliques, while onager associations are more fluid. We find evidence that females associate assortatively by reproductive state in Grevy's zebra but not in onagers. The current approach demonstrates the utility of network metrics for identifying fine-grained variation among individuals and populations in association patterns. From our analysis, we can make testable predictions about behavioral mechanisms underlying social structure and its effects on transmission processes.
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