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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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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. (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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