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Reproductive status influences group size and persistence of bonds in male plains zebra (Equus burchelli)

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Abstract

Animal groups arise from individuals’ choices about the number, characteristics, and identity of associates. Individuals make these choices to gain benefits from their associations. As the needs of an individual change with its phenotype, so too we expect the nature of its associations to vary. In this paper, we investigate how the social priorities of male plains zebra (Equus burchelli) depend on reproductive state. An adult male is either a bachelor, and lacking mating access, or a stallion defending a harem. Multiple harems and bachelor males aggregate in larger herds. Herds frequently split and merge, affording males opportunities to change associates. Over a 4-year period, we sampled the herd associations in a population of 500–700 zebras. To isolate the effects of reproductive state on male social behavior, we account for potential confounding factors: changes in population size, grouping tendencies, and sampling intensity. We develop a generally applicable permutation procedure, which allows us to test the null hypothesis that social behavior is independent of male status. Averaging over all individuals in the population, we find that a typical bachelor is found in herds containing significantly more adults, bachelors, and stallions than the herds of a typical stallion. Further, bachelors’ bonds with each other are more persistent over time than those among stallions. These results suggest that bachelors form cohesive cliques, in which we may expect cooperative behaviors to develop. Stallion–stallion associations are more diffuse, and less conducive to long-term cooperation.

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References

  • Beacham JL (2003) Models of dominance hierarchy formation: effects of prior experience and intrinsic traits. Behaviour 140:1275–1303

    Article  Google Scholar 

  • Beauchamp G (1999) The evolution of communal roosting in birds: origin and secondary losses. Behav Ecol 10:675–687

    Article  Google Scholar 

  • Boinski S (1994) Affiliation patterns among male Costa-Rican squirrel-monkeys. Behaviour 130:191–209

    Article  Google Scholar 

  • Burt DB, Peterson AT (1993) Biology of cooperative-breeding scrub jays (Aphelocoma coerulescens) Of Oaxaca, Mexico. Auk 110:207–214

    Google Scholar 

  • Caraco T, Giraldeau LA (1991) Social Foraging - Producing And Scrounging In A Stochastic Environment. J Theor Biol 153:559–583

    Article  Google Scholar 

  • Chapman CA, Wrangham RW, Chapman LJ (1995) Ecological constraints on group-size - an analysis of spider monkey and chimpanzee subgroups. Behav Ecol Sociobiol 36:59–70

    Article  Google Scholar 

  • Chase ID, Tovey C, Spangler-Martin D, Manfredonia M (2002) Individual differences versus social dynamics in the formation of animal dominance hierarchies. Proc Natl Acad Sci USA 99:5744–5749

    Article  PubMed  CAS  Google Scholar 

  • Clutton-Brock T, Price O, Albon S, Jewell P (1991) Persistent Instability and population regulation in Soay sheep. J Anim Ecol 60:593–608

    Article  Google Scholar 

  • Clutton-Brock T, Stevenson I, Marrow P, MacColl A, Houston A, JM M (1996) Population fluctuations, reproductive costs and life-history tactics in female Soay sheep. J Anim Ecol 65:675–689

    Article  Google Scholar 

  • Clutton-Brock TH, Brotherton PNM, O'Riain MJ, Griffin AS, Gaynor D, Kansky R, Sharpe L, McIlrath GM (2001) Contributions to cooperative rearing in meerkats. Anim Behav 61:705–710

    Article  Google Scholar 

  • Couzin ID, Krause J, Franks NR, Levin SA (2005) Effective leadership and decision-making in animal groups on the move. Nature 433:513–516

    Article  PubMed  CAS  Google Scholar 

  • Danchin E, Wagner RH (1997) The evolution of coloniality: the emergence of new perspectives. Trends Ecol Evol 12:342–347

    Article  Google Scholar 

  • Fischhoff IR, Sundaresan SR, Cordingley J, Larkin HM, Sellier MJ, Rubenstein DI (2007a) Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii. Anim Behav 73:825–831

    Article  Google Scholar 

  • Fischhoff IR, Sundaresan SR, Cordingley JE, Rubenstein DI (2007b) Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions. Behav Ecol 18:725–729

    Article  Google Scholar 

  • Gibbs JP, Droege S, Eagle P (1998) Monitoring populations of plants and animals. Bioscience 48:935–940

    Article  Google Scholar 

  • Goessmann C, Hemelrijk C, Huber R (2000) The formation and maintenance of crayfish hierarchies: behavioral and self-structuring properties. Behav Ecol Sociobiol 48:418–428

    Article  Google Scholar 

  • Hamilton WD (1971) Geometry For Selfish Herd. J Theor Biol 31:295

    Article  PubMed  CAS  Google Scholar 

  • Hasselquist D (1998) Polygyny in great reed warblers: a long-term study of factors contributing to male fitness. Ecology 79:2376–2390

    Article  Google Scholar 

  • Hoare DJ, Ruxton GD, Godin JGJ, Krause J (2000) The social organization of free-ranging fish shoals. Oikos 89:546–554

    Article  Google Scholar 

  • Horn HS (1968) Adaptive significance of colonial nesting in Brewers Blackbird (Euphagus cyanocephalus). Ecology 49:682

    Article  Google Scholar 

  • James R, Croft DP, Krause J (2009) Potential banana skins in animal social network analysis. Behav Ecol Sociobiol. doi:10.1007/s00265-009-0742-5

  • Janson CH, Goldsmith ML (1995) Predicting group-size in primates - foraging costs and predation risks. Behav Ecol 6:326–336

    Article  Google Scholar 

  • Klingel H (1975) Social-organization and reproduction in equids. Journal of Reproduction and Fertility 23(Suppl):7–11

    PubMed  Google Scholar 

  • Klingel H (1998) Observations on social organization and behaviour of African and Asiatic Wild Asses (Equus africanus and Equus hemionus) (Reprinted from Z Tierpsychol, vol 44, pg 323–331, 1977). Appl Anim Behav Sci 60:103–113

    Article  Google Scholar 

  • Krause J, Lusseau D, James R (2009) Animal social networks: an introduction. Behav Ecol Sociobiol. doi:10.1007/s00265-009-0747-0

  • Neems RM, Lazarus J, McLachlan AJ (1992) Swarming behavior in male chironomid midges—a cost-benefit-analysis. Behav Ecol 3:285–290

    Article  Google Scholar 

  • Rubenstein DI (1986) Ecology and sociality in horses and zebras. In: Rubenstein DI, Wrangham RW (eds) Ecological aspects of social evolution: birds and mammals. Princeton University Press, Princeton, pp 282–302

    Google Scholar 

  • Rubenstein DI (1994) Ecology of female social behavior in horses, zebras and asses. In: Jarman P, Rossiter A (eds) Animal societies: individuals, interaction and organisation. Kyoto University Press, pp 13–28

  • Rubenstein DI, Hack M (2004) Natural and sexual selection and the evolution of multi-level societies: insights from zebras with comparisons to primates. In: Kappeler PM, van Schaik C (eds) Sexual selection in primates: new and comparative perspectives. Cambridge University Press, Cambridge, pp 266–279

    Google Scholar 

  • Van Schaik CP, Van Noordwijk MA (1989) The special role of male Cebus monkeys in predation avoidance and its effect on group composition. Behav Ecol Sociobiol 24:265–276

    Article  Google Scholar 

  • Vickery WL, Giraldeau LA, Templeton JJ, Kramer DL, Chapman CA (1991) Producers, scroungers, and group foraging. Am Nat 137:847–863

    Article  Google Scholar 

  • Zuberbuhler K, Noe R, Seyfarth RM (1997) Diana monkey long-distance calls: messages for conspecifics and predators. Anim Behav 53:589–604

    Article  Google Scholar 

Download references

Acknowledgments

We thank the Ministry of Education, Government of Kenya for permission to work in Kenya. All work described here complies with the laws of Kenya. We are grateful to Ol Pejeta Conservancy for allowing us to work there and providing field support. For hosting and supporting us during this work, we thank Princeton University, McMaster University, Mpala Research Center, Denver Zoological Foundation, and Ol Pejeta Conservancy. We thank organizers Jens Krause and David Lusseau, and fellow participants at the International Ethological Conference symposium on social networks, for which we initiated this analysis. We acknowledge funding from the US National Science Foundation (IBN-0309233 [SRS, DIR], CNS-025214 [IRF, JEC, DIR], IOB-9874523 [IRF, JEC, DIR], IIS-0705822 [IRF, DIR]), Pew Charitable Trusts award 2000-0002558 “Program in Biocomplexity” (IRF, SRS, DIR), Teresa Heinz Environmental Scholars program (IRF), Smithsonian Institution (IRF), and National Sciences and Engineering Research Council of Canada (JD). Two anonymous reviewers, Jens Krause and Tatiana Czeschlik provided useful comments on earlier drafts.

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Authors and Affiliations

  1. Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, 08544, USA

    Ilya R. Fischhoff, Siva R. Sundaresan, Justine E. Cordingley & Daniel I. Rubenstein

  2. Department of Biology, McMaster University, 1280 Main St W, Hamilton, ON, L8S 4 K1, Canada

    Ilya R. Fischhoff & Jonathan Dushoff

  3. Mpala Research Center, PO Box 155, Nanyuki, Kenya

    Ilya R. Fischhoff, Siva R. Sundaresan, Justine E. Cordingley & Daniel I. Rubenstein

  4. Department of Conservation Biology, Denver Zoological Foundation, 2300 Steele Street, Denver, CO, 80205, USA

    Siva R. Sundaresan

  5. Ecological Monitoring Department, Ol Pejeta Conservancy, Private Bag, Nanyuki, 10400, Kenya

    Justine E. Cordingley

  6. Percy FitzPatrick Institute of African Ornithology, University of Cape Town, Rondebosch 7701, Cape Town, South Africa

    Justine E. Cordingley

Authors
  1. Ilya R. Fischhoff
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  2. Jonathan Dushoff
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  3. Siva R. Sundaresan
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  4. Justine E. Cordingley
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  5. Daniel I. Rubenstein
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Corresponding author

Correspondence to Ilya R. Fischhoff.

Additional information

Communicated by J. Krause

This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).

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Fischhoff, I.R., Dushoff, J., Sundaresan, S.R. et al. Reproductive status influences group size and persistence of bonds in male plains zebra (Equus burchelli). Behav Ecol Sociobiol 63, 1035–1043 (2009). https://doi.org/10.1007/s00265-009-0723-8

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  • DOI: https://doi.org/10.1007/s00265-009-0723-8

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