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Zhang, Y., Cao, Q. S., Rubenstein, D. I., Zang, S., Songer, M., Leimgruber, P., et al. (2015). Water Use Patterns of Sympatric Przewalski's Horse and Khulan: Interspecific Comparison Reveals Niche Differences. Plos One, 10(7), e0132094.
Abstract: Acquiring water is essential for all animals, but doing so is most challenging for desert-living animals. Recently Przewalski's horse has been reintroduced to the desert area in China where the last wild surviving member of the species was seen before it vanished from China in the1960s. Its reintroduction placed it within the range of a close evolutionary relative, the con-generic Khulan. Determining whether or not these two species experience competition and whether or not such competition was responsible for the extinction of Przewalski's horses in the wild over 50 years ago, requires identifying the fundamental and realized niches of both species. We remotely monitored the presence of both species at a variety of water points during the dry season in Kalamaili Nature Reserve, Xinjiang, China. Przewalski's horses drank twice per day mostly during daylight hours at low salinity water sources while Khulans drank mostly at night usually at high salinity water points or those far from human residences. Spatial and temporal differences in water use enables coexistence, but suggest that Przewalski's horses also restrict the actions of Khulan. Such differences in both the fundamental and realized niches were associated with differences in physiological tolerances for saline water and human activity as well as differences in aggression and dominance.
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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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Rubenstein, D. I., & Hohmann, M. E. (1989). Parasites and Social Behavior of Island Feral Horses. Oikos, 3, 312–320.
Abstract: The horses of Shackelford Banks, North Carolina, USA, are heavily parasitized by intestinal worms and harassed by dipterans, and although both types influence behavior only internal parasites affect bodily condition and the structuring of horse society. Thirteen species of internal parasites were identified, but only 4 of 13 groups contain them all and even within groups differences among individuals are large. Among individuals ova emissions vary ranging from 50 to 76,875 eggs per gram. The most important environmental factors influencing egg production are season and a group's location on the island, presumably because of salinity and soil differences and their effects on ova survival. Of the social and life history factors, age, and group size, but neither reproductive state nor dominance status are important. The fitness consequences of internal parasitism may be large since the number emitted is negatively correlated with next year's bodily condition. Biting fly burdens are also affected by a variety of environmental factors. In general, horses are covered with more flies on sunny days, when winds are moderately brisk, when occupying dunes, and around mid-day. In contrast to endoparasites, fly burden is affected by reproductive condition and dominance status and tends to decrease as groups increase in size. Since groups do not grow very large, nor do females attempt to bring groups together, the negative effects of endoparasites appear to overide those associated with ectoparasites. Consequently, endoparasites appear to exert a stronger influence on social structure, even though ectoparasites seem to play a stronger role in shaping details of behavior.
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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., & Hack, M. A. (1992). Horse signals: The sounds and scents of fury. Evol. Ecol., 6(3), 254–260.
Abstract: During contests animals typically exchange information about fighting ability. Among feral horses these signals involve olfactory or acoustical elements and each type can effectively terminate contests before physical contact becomes necessary. Dung transplant experiments show that for stallions, irrespective of rank, olfactory signals such as dung sniffing encode information about familiarity suggesting that such signals can be used as signatures. As such they can provide indirect information about fighting ability as long as opponents associate identity with past performance. Play-back experiments, however, show that vocalizations, such as squeals, directly provide information about status regardless of stallion familiarity. Sonographs reveal that squeals of dominants are longer than those of subordinates and that only those of dominants have at their onset high-frequency components.
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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. (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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Parker, G. A., & Rubenstein, D. I. (1981). Role assessment, reserve strategy, and acquisition of information in asymmetric animal conflicts. Anim. Behav., 29(1), 221–240.
Abstract: It was formerly argued that alternative evolutionarily stable strategies (ESSs) are possible for animal contests characterized by some asymmetry that can be perceived with perfect accuracy. Where roles A and B refer to the asymmetry between opponents, ESSs are: [`]fight when A, retreat when B', and vice versa. Either can be an ESS, but only if the [`]reserve strategy' (=what an animal does when it fights) is sufficiently damaging. We examine the [`]war of attrition' (winner = opponent that persists longer). In a population at either ESS, reserve strategy is never normally shown; it is therefore subject to drift unless the selective action of rare individuals which break the convention is considered. These could arise either by mutation or by mistakes in role assessment. When mutations and mistakes simply specify that occasionally an animal fights when it [`]should' retreat, selection adjusts reserve strategy to a level where only one ESS (the [`]commonsense' ESS) is possible, if the asymmetry is relevant to payoff. Thus for asymmetries in fighting ability or resource value, the individual with the lower score will retreat. However, we are particularly concerned with cases where both payoff-relevant aspects (fighting ability and resource value) are asymmetric. If opponents sustain contest costs at rates KA and KB, and their resource values are VA and VB, an [`]optimal assessor' strategy defined by the interaction between the two asymmetries, is a unique ESS. It obeys the rule [`]fight on estimating role A, where VA/KA>VB/KB; retreat in B'. If mistakes can occur in both roles, but are very rate, the ESS is not fundamentally altered though there will be infinitesimal tendencies for persisting in role B. Selection to improve assessment abilities intensifies as abilities improve, but is weak if roles A and B are rather similar. Over a range of similarity between roles, an [`]owner wins' convention may be adopted if ownership correlates positively with role A and an individual cannot tell when it would otherwise pay him to break the convention. We also examine a contest in which information about roles can be acquired only during a contest itself, and at a cost. Much depends on the rate at which information is acquired relative to the rate at which costs are expended, and on whether contests normally escalate in intensity, remain at the same level, or de-escalate. Selection favours short contests when costs are high relative to resource value, where the outcome of a round contains much information about fighting ability, and where the actual disparity in fighting ability is large.
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