15 - Natural and sexual selection and the evolution of multi-level societies: insights from zebras with comparisons to primates  pp. 266-279

Natural and sexual selection and the evolution of multi-level societies: insights from zebras with comparisons to primates

By Daniel I. Rubenstein and Mace Hack

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Animal societies derive from the social relationships that exist among its members (Hinde, 1983). Behavioural ecologists have traditionally focused on the core relationships defining a mating system as a means toward understanding the role of ecology in the evolution of sociality (e.g. Jarman, 1974; Bradbury & Vehrencamp, 1977). Typically, these core relationships emerge from the operation of both natural and sexual selection and how they differently affect the behaviour of females and males (Rubenstein, 1986; van Schaik, 1989). However, emphasis on mating systems has tended to marginalise the importance to social evolution of interactions and relationships that extend beyond the basic breeding unit. This is even true for the small subset of species with multi-level societies, where breeding units and other social subgroups are themselves organised into more complex social groups within a population. By examining how natural and sexual selection operate within multi-level societies, however, a more complete understanding of the function and evolution of sociality emerges than would by investigating the dynamics of mating systems alone (e.g. Dunbar, 1988).

Although the societies of many primate species are multi-levelled, the relative simplicity of societies of plains zebras (Equus burchelli) where only two tiers exist – the core breeding units and the herds they often comprise – can provide insights into the rules that give form to multi-level societies. In this chapter we begin by highlighting the environmental and sociosexual factors that shape zebra mating systems and herd dynamics.

Alexander, R. D. 1974. The evolution of social behavior. Annual Review of Ecological Systems, 5, 325–83
Altmann, J. 1990. Primate males go where females are. Animal Behaviour, 39, 193–4
Andleman, S. J. 1986. Ecological and social determinants of Cercopithecine mating patterns. In Ecological Aspects of Social Evolution: Birds and Mammals, ed. D. I. Rubenstein & R. W. Wrangham. Princeton, NJ: Princeton University Press, pp. 201–16
Barton, R. A. 2000. Socioecology of baboons: the interaction of male and female strategies. In Primate Males: Causes and Consequences of Variation in Group Composition, ed. P. M. Kappeler. Cambridge: Cambridge University Press, pp. 97–107
Berger, J. 1986. Wild Horses of the Great Basin: Social Competition and Population Size. Chicago, IL: University of Chicago Press
Berger, J. 1987. Reproductive fates of dispersers in a harem-dwelling ungulate: the wild horse. In Mammalian Dispersal Patterns: The Effects of Social Structure on Population Genetics, ed. B. D. Chepko-Sade & Z. T. Halpin. Chicago, IL: University of Chicago, pp. 41–54
Bradbury, J. W. & Vehrencamp, S. L. 1977. Social organization and foraging in emballonurid bats. III. Mating systems. Behavioral Ecology and Sociobiology, 2, 1–17
Brown, J. L. 1987. Helping and Communal Breeding in Birds. Princeton, NJ: Princeton University
Byrne, R., Whiten, A. & Henzi, S. 1990. Social relationships in mountain baboons: leadership and affiliation in a non-female-bonded monkey. American Journal of Primatology, 20, 313–329
Clutton-Brock, T. H. 1989. Mammalian mating systems. Proceedings of the Royal Society of London, Series B, 236, 339–72
Dunbar, R. I. M. 1986. The social ecology of gelada baboons. In Ecological Aspects of Social Evolution, ed. D. I. Rubenstein & R. W. Wrangham. Princeton, NJ: Princeton University, pp. 332–51
Dunbar, R. I. M. 1988. Primate Social Systems. Ithaca: Cornell University
Emlen, S. T. & Oring, L. W. 1977. Ecology, sexual selection, and the evolution of mating systems. Science, 197, 215–23
Feh, C. 1999. Alliances and reproductive success in Camargue horses. Animal Behaviour, 57, 705–13
Feist, J. D. & McCullough, D. R. 1976. Behavior patterns and communication in feral horses. Zeitschrift für Tierpsychologie, 41, 337–71
Ginsberg, J. R. 1989. The ecology of female behaviour and male mating success in the Grevy's zebra. Symposia of the Zoological Society of London, 61, 89–110
Groves, C. P. 1974. Horses, Asses, and Zebras in the Wild. Hollywood, FL: Curtis Books
Hinde, R. A. 1983. A conceptual framework. In Primate Social Relationships, ed. R. A. Hinde. Oxford: Blackwell, pp. 1–7
Jarman, P. J. 1974. The social organisation of antelope in relation to their ecology. Behaviour, 48, 215–67
Joubert, E. 1972. The social organisation and associated behaviour in the Hartmann zebra Equus zebra hartmannae. Madoqua, 1, 17–56
Klingel, H. 1969a. Reproduction in the plains zebra, Equus burchelli boehmi: behaviour and ecological factors. Journal of Reproduction and Fertility, Supplement, 6, 339–45
Klingel, H. 1969b. The social organisation and population ecology of the plains zebra (Equus quagga). Zoologica Africana, 4, 249–63
Klingel, H. 1974. A comparison of the social behaviour of the Equidae. In The Behaviour of Ungulates and Its Relation to Management, ed. V. Geist & F. Walther. Morges, Switzerland: IUCN Publications, pp. 124–32
Klingel, H. 1977. Observations on social organization and behaviour of African and Asiatic wild asses. Zeitschrift für Tierpsychologie, 44, 323–31
Kummer, H. 1968. Social Organization of Hamadrayas Baboons. A Field Study. Chicago, IL: University of Chicago
Linklater, W. L. 2000. Adaptive explanation in socio-ecology: lessons from the Equidae. Biological Reviews of the Cambridge Philosophical Society, 75, 1–20
Linklater, W. L., Cameron, E. Z., Minot, E. O. & Stafford, K. J. 1999. Stallion harassment and the mating system of horses. Animal Behaviour, 58, 295–306
Lloyd, P. H. & Rasa, O. A. E. 1989. Status, reproductive success and fitness in Cape mountain zebra (Equus zebra zebra). Behavioral Ecology and Sociobiology, 25, 411–20
McCracken, G. F. & Bradbury, J. W. 1981. Social organization and kinship in the polygynous bat Phyllostomus hastatus. Behavioral Ecology and Sociobiology, 8, 11–34
Miller, R. 1979. Band organization and stability in Red Desert feral horses. In Proceedings of a Conference on the Ecology and Behavior of Feral Equids, ed. R. H. Denniston. Laramie, WY: University of Wyoming, pp. 113–23
Moehlman, P. D. 1979. Behavior and ecology of feral asses (Equus asinus). National Geographic Society Research Reports, 405–11
Penzhorn, B. L. 1984. A long-term study of social organisation and behaviour of Cape mountain zebras Equus zebra zebra. Zeitschrift für Tierpsychologie, 64, 97–146
Pereira, M. E. & Fairbanks, L. A. 1993. Juvenile Primates. New York, NY: Oxford University Press
Rubenstein, D. I. 1981. Behavioural ecology of island feral horses. Equine Veterinary Journal, 13, 27–34
Rubenstein, D. I. 1986. Ecology and sociality in horses and zebras. In Ecological Aspects of Social Evolution, ed. D. I. Rubenstein & R. W. Wrangham. Princeton, NJ: Princeton University, pp. 282–302
Rubenstein, D. I. 1989. Life history and social organization in arid adapted ungulates. Journal of Arid Environments, 17, 145–56
Rubenstein, D. I. 1994. The ecology of female social behaviour in horses, zebras and asses. In Animal Societies: Individuals, Interactions and Organisation, ed. P. J. Jarman & A. Rossiter. Kyoto: Kyoto University, pp. 13–28
Rutberg, A. T. 1990. Inter-group transfer in Assateague pony mares. Animal Behaviour, 40, 945–52
Scheel, D. 1993. Profitability, encounter rates, and prey choice of African lions. Behavioral Ecology, 4, 90–7
Sigg, H., Stolba, A., Abegglen, J.-J. & Dasser, V. 1982. Life history of hamadryas baboons: physical development, infant mortality, reproductive parameters and family relationships. Primates, 23, 473–87
Stammbach, E. 1987. Desert, forest, and montane baboons: multilevel societies. In Primate Societies, ed. B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham & T. T. Struhsaker. Chicago, IL: University of Chicago, pp. 112–20
Stroeh, O. 2001. The Effects of Management Strategies on the Behavioural Ecology of the Shackleford Banks Male Horses. B. Sc. thesis
Turner, J. W. J. 1992. Seasonal mountain lion predation on a feral horse population. Canadian Journal of Zoology, 70, 929–34
van Schaik, C. P. 1989. The ecology of social relationships among female primates. In Comparative Socioecology, ed. V. Standen & R. A. Foley. Oxford: Blackwell, pp. 195–218
Woodward, S. L. 1979. The social system of feral asses (Equus asinus). Zeitschrift für Tierpsychologie, 49, 304–16