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Journal of Ethology

, Volume 26, Issue 1, pp 137–144 | Cite as

Female–female competition or male mate choice? Patterns of courtship and breeding behavior among feral horses (Equus caballus) on Assateague Island

  • David M. Powell
Article

Abstract

Previous research on the Assateague horses found that high-ranking females had more surviving offspring than low-ranking females. Variance in reproductive success may be the result of a variety of proximate processes that affect sexual behavior such as mate choice and mate competition. A study was done to determine whether patterns of courtship, social, and sexual behavior could be identified that would suggest mate choice and/or mate competition. Behavioral data were collected from approximately 40 sexually mature mares living in harem bands. Stallions showed more interest in the eliminations of dominant mares than subordinate mares. Males also engaged in significantly more high-intensity (e.g., mounts and copulations) sexual behavior with dominant mares than subordinate mares, and there was a trend for males to engage in more low-intensity (e.g., flehmen and ano-genital sniffing) sexual behavior with dominant mares than subordinate mares. There was no effect of mare rank on spatial relationships with the stallion; however, dominant mares did attempt to restrict reproductive access to the stallion by harassing and disrupting copulations. Higher foaling rates among dominant mares on Assateague Island could therefore be the result of rank-related mate choice by stallions and direct female competition for mating opportunities.

Keywords

Mate choice Female competition Dominance Horses Copulation harassment 

Notes

Acknowledgements

I must thank my field staff: Paul Varga, Tim Hadlock, and 63 Earthwatch volunteers for their assistance with this research. I also thank Allison Turner, Jack Kumer, and Carl Zimmerman at the Assateague Island National Seashore for permission to work at the site and for access to records on the horses. This work was supported by a Pre-doctoral Fellowship from the Smithsonian Institution, the Department of Zoological Research of the National Zoological Park, the Biology of Small Populations NSF training grant at the University of Maryland (Grant #BIR 9602266), Earthwatch, Sigma Xi, and the Abbott and Nelson Funds of the Smithsonian Institution. I would like to thank Devra Kleiman, James Dietz, Gerald Wilkinson, Sue Carter-Porges, Estelle Russek-Cohen and two anonymous reviewers for their comments on this research.

References

  1. Ahnesjo I, Vincent A, Alatalo R, Halliday T, Sutherland WJ (1993) The role of females in influencing mating patterns. Behav Ecol 4:187–189CrossRefGoogle Scholar
  2. Altmann J (1974) Observational study of behavior: sampling methods. Behaviour 49:227–267PubMedGoogle Scholar
  3. Asa CS, Goldfoot DA, Ginther OJ (1979) Sociosexual behavior and the ovulatory cycle of ponies (Equus caballus) observed in harem groups. Horm Behav 13:49–65PubMedCrossRefGoogle Scholar
  4. Barnard P, Markus MB (1989) Male copulation frequency and female competition for fertilizations in a promiscuous brood parasite, the pin-tailed whydah (Vidua macroura). Ibis 131:421–425Google Scholar
  5. Bebié N, McElligott AG (2006) Female aggression in red deer: does it indicate competition for mates? Mamm Biol 71:347–355CrossRefGoogle Scholar
  6. Bennett NC, Faulkes CG, Jarvis JUM (1999) Socially induced infertility, incest avoidance, and the monopoly of reproduction in cooperatively breeding African mole rats, family Bathyergidae. Adv Study Behav 28:75–114CrossRefGoogle Scholar
  7. Berenstain L, Wade TD (1983) Intrasexual selection and male mating strategies in baboons and macaques. Int J Primatol 4:201–235CrossRefGoogle Scholar
  8. Berger J (1986) Wild horses of the Great Basin. University of Chicago Press, ChicagoGoogle Scholar
  9. Berger J (1989) Female reproductive potential and its apparent evaluation by male mammals. J Mammal 70:347–358CrossRefGoogle Scholar
  10. Berglund A, Rosenqvist G (1993) Selective males and ardent females in pipefishes. Behav Ecol Sociobiol 32:331–336CrossRefGoogle Scholar
  11. Berglund A, Magnhagen C, Bisazza A, Konig B, Huntingford F (1993) Female-female competition over reproduction. Behav Ecol 4:184–187CrossRefGoogle Scholar
  12. Brandtmann G, Scandura M, Trillmich F (1999) Female-female conflict in the harem of a snail cichlid (Lamprologus ocellatus). Behaviour 136:1123–1144CrossRefGoogle Scholar
  13. Bro-Jørgensen J (2002) Overt female mate competition and preference for central males in a lekking antelope. Proc Nat Acad Sci USA 99:9290–9293PubMedCrossRefGoogle Scholar
  14. Brown JL, Brown ER, Sedransk J, Ritter S (1997) Dominance, age and reproductive success in a complex society: a long term study of the Mexican jay. Auk 114:279–286Google Scholar
  15. Burley N, Moran N (1979) The significance of age and reproductive experience in the mate preferences of feral pigeons, Columba livia. Anim Behav 27:686–698CrossRefGoogle Scholar
  16. Bush SL, Bell DJ (1997) Courtship and female competition in the Majorcan midwife toad (Alytes muletensis). Ethology 103:292–303CrossRefGoogle Scholar
  17. Clutton-Brock TH, Guinness FE, Albon SD (1982) Red deer: behavior and ecology of two sexes. Univ of Chicago Press, ChicagoGoogle Scholar
  18. Cunningham EJA, Birkhead TR (1998) Sex roles and sexual selection. Anim Behav 56:1311–1321PubMedCrossRefGoogle Scholar
  19. Darwin C (1871) The descent of man, and selection in relation to sex. Murray Press, LondonGoogle Scholar
  20. Derix RRWM, van Hoof JARAM (1995) Male and female partner preferences in a captive wolf pack (Canis lupus): specificity versus spread of sexual attention. Behaviour 132:128–148Google Scholar
  21. Dewsbury DA (1982) Ejaculate cost and male choice. Am Nat 119:601–610CrossRefGoogle Scholar
  22. Estes RD (1972) The role of the vomeronasal organ in mammalian reproduction. Mammalia 36:315–341CrossRefGoogle Scholar
  23. Estes RD (1991) The behavior guide to African mammals. University of California Press, BerkeleyGoogle Scholar
  24. Frame LH, Malcolm JR, Frame GW, van Lawick H (1979) Social organization of African wild dogs (Lycaon pictus) on the Serengeti plains, Tanzania (1967–1978). Z Tierpsychol 50:225–249Google Scholar
  25. Gwynne DT (1981) Sexual difference theory: Mormon crickets show role reversal in mate choice. Science 213:779–780PubMedCrossRefGoogle Scholar
  26. Hart BL (1983) Flehmen behavior and vomeronasal organ function. In: Müller-Schwarze D, Silverstein RM (eds) Chemical signals in vertebrates. Plenum Press, New York, pp 87–103Google Scholar
  27. Houpt KA, Keiper R (1982) The position of the stallion in the equine dominance hierarchy of feral and domestic ponies. J Anim Sci 54:945–950Google Scholar
  28. Keiper RR (1985) The Assateague ponies. Tidewater, Centreville, p 185Google Scholar
  29. Keiper R, Houpt K (1984) Reproduction in feral horses: an eight year study. Am J Vet Res 45:991–995PubMedGoogle Scholar
  30. Keiper RR, Sambraus HH (1986) The stability of equine dominance hierarchies and the effects of kinship, proximity, and foaling status on hierarchy rank. Appl Anim Behav Sci 16:121–130CrossRefGoogle Scholar
  31. Kirkpatrick JF, Liu IKM, Turner JW Jr, Naugle R, Keiper R (1992) Long-term effects of porcine zona pellucida immunocontraception on ovarian function in feral horses (Equus caballus). J Reprod Fertil 94:437–444PubMedCrossRefGoogle Scholar
  32. Krebs JR, Davies NB (1987) An introduction to behavioral ecology, 2nd edn. Sinauer, Sutherland, p 257Google Scholar
  33. Lloyd PH, Rasa OAE (1989) Status, reproductive success, and fitness in Cape Mountain zebra (Equus zebra zebra). Behav Ecol Sociobiol 25:411–420CrossRefGoogle Scholar
  34. Mackler SF, Dolan JM Jr (1980) Social structure and herd behavior of Equus przewalskii Poliakov, 1881 at the San Diego Wild Animal Park. Equus 2:55–69Google Scholar
  35. Milner-Gulland EJ, Bukreeva OM, Coulson T, Lushchekina AA, Kholodova MV, Bekenov AB, Grachev IA (2003) Reproductive collapse in saigo antelope harems. Nature 422:135PubMedCrossRefGoogle Scholar
  36. Nakamura M (1998) Multiple mating and cooperative breeding in polygynandrous alpine accentors. I. Competition among females. Anim Behav 55:259–275PubMedCrossRefGoogle Scholar
  37. Niemeyer CL, Anderson JR (1983) Primate harassment of matings. Ethol Sociobiol 4:205–220CrossRefGoogle Scholar
  38. Pluháček J, Bartoš L, Čulík L (2006) High-ranking mares of captive plains zebra Equus burchelli have greater reproductive success than low-ranking mares. Appl Anim Behav Sci 99:315–329CrossRefGoogle Scholar
  39. Powell DM (1999) Preliminary evaluation of porcine zona pellucida (PZP) immunocontraception for behavioral effects in feral horses (Equus caballus). J Appl Anim Welf Sci 2:321–335PubMedCrossRefGoogle Scholar
  40. Powell DM (2000) Evaluation of effects of contraceptive population control on behavior and the role of social dominance in female feral horses, Equus caballus. PhD Thesis, University of Maryland, College ParkGoogle Scholar
  41. Powell DM, Monfort SL (2001) Assessment: effect of porcine zona pellucida (PZP) immunocontraception on estrous cyclicity in feral horses. J Appl Anim Welf Sci 4:271–284CrossRefGoogle Scholar
  42. Preston BT, Stevenson IR, Pemberton JM, Wilson K (2001) Dominant rams lose out by sperm depletion. Nature 409:681–682PubMedCrossRefGoogle Scholar
  43. Rabb GL, Woolpy JH, Ginsburg BE (1967) Social relations in a group of captive wolves. Am Zool 7:305–311Google Scholar
  44. Rasa OAE (1977) The ethology and sociology of the dwarf mongoose (Helogale undulata rufula). Z Tierpsychol 43:337–406Google Scholar
  45. Rendall D, Taylor LL (1991) Female sexual behavior in the absence of male male competition in captive Japanese macaques (Macaca fuscata). Zoo Biol 10:319–328CrossRefGoogle Scholar
  46. Robinson JG (1982) Intrasexual competition and mate choice in primates. Am J Primatol Suppl 1:131–144CrossRefGoogle Scholar
  47. Rood JP (1980) Mating relationships and breeding suppression in the dwarf mongoose. Anim Behav 28:143–150CrossRefGoogle Scholar
  48. Rutberg AT, Greenberg SA (1990) Dominance, aggression frequencies, and modes of aggressive competition in feral pony mares. Anim Behav 40:322–331CrossRefGoogle Scholar
  49. Schein MW, Fohrman MH (1955) Social dominance relationships in a herd of dairy cattle. Br J Anim Behav 3:45–55CrossRefGoogle Scholar
  50. Schilder MBH (1990) Intervention in a herd of semi-captive plains zebras. Behaviour 112:53–83Google Scholar
  51. Schilder MBH, Boer LP (1987) Ethological investigation on a herd of plains zebra in a safari park: time budgets, reproduction, and food competition. Appl Anim Behav Sci 18:45–56CrossRefGoogle Scholar
  52. Seligsohn E (1987) Dominance relationships and reproductive success within bands of feral ponies. PhD Thesis, University of Connecticut, StorrsGoogle Scholar
  53. Slagsvold T, Lifjeld JT (1994) Polygyny in birds: the role of competition between females for male parental care. Am Nat 143:59–94CrossRefGoogle Scholar
  54. Small MF, Smith DG (1982) The relationship between maternal and paternal rank in rhesus macaques (Macaca mulatta). Anim Behav 30:626–627CrossRefGoogle Scholar
  55. Stebbins MC (1974) Social organization in free-ranging Appaloosa horses. PhD Thesis, Idaho State University, PocatelloGoogle Scholar
  56. Szykman M, Engh AL, Van Horn RC, Funk SM, Scribner KT, Holekamp KE (2001) Association patterns among male and female spotted hyenas (Crocuta crocuta) reflect male mate choice. Behav Ecol Sociobiol 50:231–238CrossRefGoogle Scholar
  57. Trivers RL (1972) Parental investment and sexual selection. In: Campbell B (ed) Sexual selection and the descent of man. Aldine, Chicago, pp 136–179Google Scholar
  58. Tyler S (1972) The behavior and social organization of the New Forest ponies. Anim Behav Monogr 5:87–196CrossRefGoogle Scholar
  59. Verrell PA (1995) Males choose larger females as mates in the salamander, Desmognathus santeetlah. Ethology 99:162–171CrossRefGoogle Scholar
  60. Waights V (1996) Female sexual interference in the smooth newt, Triturus vulgaris vulgaris. Ethology 102:736–747CrossRefGoogle Scholar
  61. Waring GH (2003) Horse behavior, 2nd edn. William Andrew, NorwichGoogle Scholar
  62. Zimen E (1975) Social dynamics in the wolf pack. In: Fox MW (ed) The wild canids. Van Nostrand, New York, pp 336–362Google Scholar

Copyright information

© Japan Ethological Society and Springer 2007

Authors and Affiliations

  1. 1.Department of MammalogyWildlife Conservation Society/Bronx ZooBronxUSA

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