Unexplained sexual segregation in polygamous ungulates: A defense of an ontogenetic approach

Behavioural Processes

Volumn 38, Issue 2, 1996, Pages 131-154

  Bon, Richard ^a   Campan, Raymond ^b  

^a FEDERAL UNIVERSITY OF PARÁ   (Brazil)

^b UNIVERSITÉ DE TOULOUSE   (France)

Author keywords

Causal and ontogenetic approach; Sexually dimorphic ungulate; Social segregation; Spatial segregation

Indexed keywords

ANIMAL EXPERIMENT; CATTLE; FEMALE; MALE; NONHUMAN; ONTOGENY; REVIEW; SEX DIFFERENCE; SEXUAL BEHAVIOR; SEXUAL MATURATION; SOCIAL BEHAVIOR; SOCIAL INTERACTION; UNGULATE;

ANIMALIA; BOS TAURUS; UNGULATA;

ACTIVITY BUDGET; OPPORTUNISTIC FEEDING; POLYGAMY; PREDATION RISK; SEXUAL SEGREGATION; UNGULATE;

EID: 0031457711     PISSN: 03766357     EISSN: None     Source Type: Journal    
DOI: 10.1016/S0376-6357(96)00029-0     Document Type: Article

References (155)

1. Alexander, C., Lynch, J.J. and Mottershead, B.E., 1979. Use of shelter and selection of lambing sites by shorn and unshorn ewes in paddocks with closely or widely spaced shelters. Appl. Anim. Ethol., 5: 51-69.
2. Appleby, M., 1983. The probability of linearity in hierarchies. Anim. Behav., 31: 600-608.
3. Armstrong, D.P., 1991. Levels of cause and effect as organizing principles for research in animal behaviour. Can. J. Zool., 69: 823-829.
4. Arnold, G.W., Boundy, C.A.P., Morgan, P.D. and Bartle, G., 1975. The roles of sight and hearing in the lamb in the location and discrimination between ewes. Appl. Anim. Ethol., 5: 43-50.
5. Arnold, G.W., Wallace, S.R. and Maller, R.A., 1979. Some factors involved in natural weaning processes in sheep. Appl. Anim. Ethol., 5: 43-50.
6. Arnold, G.W., Wallace, S.R. and Rea, W.A., 1981. Associations between individuals and home-range behaviour in natural flocks of three breeds of domestic sheep. Appl. Anim. Ethol., 7: 239-257.
7. Arnold, G.W. and Grassia, A., 1985. Spatial relationships between ewes and lambs. Appl. Anim. Behav. Sci., 14: 253-261.
8. Beier, P., 1987. Sex differences in quality of white-tailed deer diets. J. Mamm., 68: 323-329.
9. Bekoff, M., 1977. Mammalian dispersal and the ontogeny of individual behavioral phenotypes. Am. Nat., 111: 715-732.
10. Bekoff, M., 1984. Social play behaviour. BioScience. 34: 228-233.
11. Bekoff, M., 1989. Behavioral development of terrestrial carnivores. In: J.L. Gittleman (Editor). Carnivore Behavior, Ecology, and Evolution, Cornell University Press, New York, NY. pp. 89-124.
12. Berger, J., 1979a. Social ontogeny and behavioural diversity: consequences for bighorn sheep Ovis canadensis inhabiting desert and mountain environments. J. Zool., Lond., 188: 251-266.
13. Berger, J., 1979b. Weaning, social environments and the ontogeny of spatial associations in bighorn sheep. Biol. Behav., 4: 363-372.
14. Berger, J., 1980. The ecology, structure and functions of social play in bighorn sheep (Ovis canadensis). J. Zool., Lond., 192: 531-542.
15. Bernstein, I.S. and Williams, L.E. (Editors). 1986. In: The study of social organization: Comparative Primate Biology, Vol. 2A: Behavior, Conservation, and Ecology, Alan Liss, New York, NY. pp. 195-213.
16. Berry, M. and Signoret, J.P., 1984. Sex play and behavioural sexualization in the pig. Reprod. Nutr. Dév., 24: 507-513.
17. Biben, M., 1989. Effects of social environment on play in squirrel monkeys: resolving Harlequin's dilemna. Ethology, 81: 72-82.
18. Bon, R., 1985. Approche dynamique de l'éco-éthologie d'une population de mouflons de Corse (Ovis ammon musimon) dans la Réserve Nationale de Chasse du Caroux-Espinouse, D.E.A., Université Paul Sabatier, Toulouse.
19. Bon, R., 1991. Trajectoires sociales chez le Mouflon de Corse: rôle des processus d'attachement dans la sociogenèse. PhD thesis, Paul Sabatier University, Toulouse.
20. Bon, R., 1992. Social and spatial sexual segregation of polygamous ungulates: proximate factors. In: F. Spitz, G. Janeau, G. Gonzalez and S. Aulagnier (Editors). Ongulés/Ungulates 91. S.F.E.P.M., Paris. pp. 195-198.
21. Bon, R. and Campan, R., 1989. Social tendencies of the Corsican mouflon Ovis ammon musimon in the Caroux-Espinouse (South of France). Behav. Process., 19: 57-78.
22. Bon, R., Dubois, M. and Maublanc, M.E., 1993a. Does age influence between-rams companionship in mouflon (Ovis gmelini.)? Rev. Ecol. (Terre vie). 47: 57-64.
23. Bon, R., Badia, J., Maublanc, M.E., and Recarte, J.M., 1993b. Population structure and social grouping of mouflons during the rut. Z. Säugetierkunde. 58: 294-301.
24. Bon, R., Joachim, J. and Maublanc. M.E., 1995. Do lambs affect feeding habitat use of lactating female mouflons in spring within an area free of predators? J. Zool. Lond., 235: 43-51.
25. Bouissou, M.F and Andrieu, S., 1977. Etablissement des relations de dominance-soumission chez les ovins domestiques. IV. Etablissement des relations chez les jeunes. Biol. Behav., 2: 97-107.
26. Bouissou, M.F. and Andrieu, S., 1978. Etablissement des relations préférentielles chez les bovins domestiques. Behaviour, 64: 148-157.
27. Bouissou, M.F. and Hövels, J., 1976. Effet des conditions d'élevage sur le comportement des génisses dans une situation de compétition alimentaire. Ann. Zootech., 25: 213-219.
28. Bouissou, M.F. and Hövels, J., 1976b. Effet d'un contact précoce sur quelques aspects du comportement social des bovins domestiques. Biol. Behav., 1: 17-36.
29. Bowyer, R.T., 1984. Sexual segregation in southern mule deer. J. Mamm., 65: 410-417.
30. Brown, S.G., 1988. Play behaviour in Lowland gorillas: age differences, sex differences, and possible functions. Primates. 29: 219-228.
31. Byers, J.A., 1984. Play in ungulates In: P.K. Smith (Editor). Play in animals and humans. Basil Blackwell, Oxford, pp. 43-65.
32. Byers, J.A., 1986. Natural variation in early experience in Pronghorn fawns: sources and consequences. In: L. Passera and J.P. Lachaud (Editors). Readings from the 19th I.E.C. The individual and society. Privat, Toulouse. pp. 81-92.
33. Carranza, J., 1984. Organización y estructura. diferenciación de comportamientos y configuración espacial. en el sistema social del ciervo (Cervus elaphus L.). PhD thesis. Universidad de Extremadura.
34. Cederlund, B.-M., 1987. Parturition and early development of moose (Alces alces L.) calves. Swedish Wildl. Res. Suppl.. 1: 399-422.
35. Cederlund, G., Sandegren, F. and Larsson, K., 1987. Summer movements of female moose and dispersal of their offspring. J. Wildl. Manag., 51: 342-352.
36. Chalmers, N.R., 1987. Developmental pathways in behaviour. Anim. Behav., 35: 659-674.
37. Cheney, D.L., 1978. The play partners of immature baboons. Anim. Behav., 26: 1038-1050.
38. Clutton-Brock, T.H., Albon, S.D., Gibson, R.M. and Guinness, F.E., 1979. The logical stag: adaptive aspects of fighting in red deer (Cervus elaphus L.). Anim. Behav., 27: 211-225.
39. Clutton-Brock, T.H., F.E. Guinness and S.D. Albon. 1982a. Red Deer. Behavior and ecology of two sexes. Edinburgh University Press, Edinburgh.
40. Clutton-Brock, T.H., Iason, G.R., Albon, S.D. and Guinness, F.E., 1982b. Effects of lactation on feeding behaviour and habitat use in wild red deer hinds. J. Zool., Lond., 198: 227-236.
41. Clutton-Brock, T.H., Iason, G.R. and Guinness, F.E., 1987. Sexual segregation and density-related changes in habitat use in male and female red deer (Cervus elaphus). J. Zool. Lond., 211: 275-289.
42. Clutton-Brock, T.H., Major, M. and Guinness, F.E., 1985. Population regulation in male and female red deer. J. Anim. Ecol., 54: 831-846.
43. Colmenares, F., 1992. Clans and harems in a colony of hamadryas and hybrid baboons: male kinship, familiarity and the formation of brother-teams. Behaviour, 121: 61-94.
44. Crook, J.H., 1989. Introduction: socioecological paradigms. evolution and history: perspectives for the 1990s. In: V. Standen and R.A. Foley (Editors). Comparative Socioecology. The behavioural ecology of humans and other mammals, Blackwell Scientific Publications. Oxford.
45. Crowell-Davis, S.L., 1987. Play development in Welsh pony (Equus caballus) foals. Appl. Anim. Behav. Sci., 18: 119-131.
46. Curio, E., 1994. Causal and functional questions: how are they linked? Anim. Behav., 47: 99-1021.
47. Dubois, M., Quenette, P.Y., Bideau, E. and Magnac, M.P., 1993. Seasonal range use by European mouflon rams in medium altitude mountains. Acta Theriol., 38: 185-198.
48. Dunbar, R.I.M., 1988. The evolutionary implications of social behaviour. In: H.C. Plotkin (Editor). The role of behavior in evolution. MIT, Cambridge. MA, pp. 105-188.
49. Duncan, P., Harvey, P.H. and Wells, S.M., 1984. On lactation and associated behaviour in a natural herd of horses. Anim. Behav., 32: 255-263.
50. Eaton, W.O. and Enns, L.R., 1986. Sex differences in human motor activity level. Psychol. Bull., 100: 19-28.
51. Eisenberg, J.F., 1981. The Mammalian Radiations. An Analysis of Trends in Evolution. Adaptation, and Behavior. University Chicago Press. Chicago, IL.
52. Espmark, Y., 1971. Mother-young relationship and ontogeny of behaviour in reindeer (Rangifer tarandus L.). Z. Tierpsychol., 29: 42-81.
53. Estep, D.Q., Crowell-Davis, S.L., Earl-Costello, S.-A and Beatey, S.A., 1993. Changes in the social behaviour of drafthorse (Equus caballus) mares coincident with foaling. Appl. Anim. Behav. Sci., 35: 199-213.
54. Estes, R.D. and Estes, R.K., 1979. The birth and survival of wildebeest calves. Z. Tierpsychol., 50: 45-95.
55. Fagen, R., 1981. Animal Play Behavior. Oxford University Press. Oxford.
56. Festa-Bianchet, M., 1986. Site fidelity and seasonal range use by bighorn rams. Can. J. Zool., 64: 2126-2132.
57. Festa-Bianchet, M., 1988. Seasonal range selection in bighorn sheep: conflicts between forage quality, forage quantity, and predator avoidance. Oecologia. 75: 580-586.
58. Festa-Bianchet, M., 1991. The social system of bighorn sheep: grouping patterns, kinship and female dominance rank. Anim. Behav., 42: 71-82.
59. Francisci, F., Focardi, S. and Boitani, L., 1985. Male and female Alpine ibex: phenology of space use and herd size. In: S. Lovari (Editor). The Biology and Management of Mountain Ungulates. Croom Helm. London, pp. 124-133.
60. Franklin, W.L. and Lieb, J.W., 1979. The social organization of a sedentary population of North American elk: a model for understanding other populations. In: M.S. Boyce and L. D. Hayden-Wing (Editors), North American Elk: Ecology, Behaviour and Management. Laramie, New York, NY. pp. 185-198.
61. Freeman, L.C., Freeman, S.C. and Romney, A.K., 1992. The implications of social structure for dominance hierarchies in red deer. Cervus elaphus L. Anim. Behav., 44: 239-245.
62. Geist, V., 1968. On delayed social and physical maturation in mountain sheep. Can. J. Zool., 46: 899-904.
63. Geist, V., 1971. Mountain sheep: a Study in Behavior and Evolution. University Chicago Press, Chicago, IL.
64. Geist, V., 1974. On the relationship of social evolution and ecology in ungulates. Am. Zool., 14: 205-220.
65. Geist, V., 1982. Adaptative behavioural strategies. In: J.W. Thomas and D.E. Toweill (Editors), Elk of North America. Stackpole Books. Harrisburg. PA. pp. 219-277.
66. Gerard, J.F. and Richard-Hansen, C., Maublanc, M.L. and Bideau, E., 1993. Probable exaptations within the 'distributed' herd. Rev. Ecol. (Terre vie). 48: 239-248.
67. Goldfoot, D.A., Wallen, K., Neff, K., McBrair, D.A. and Goy, M.C., 1984. Social influences on the display of sexually dimorphic behavior in rhesus monkeys: isosexual rearing. Arch. Sex. Behav., 13: 395-412.
68. Gonzalez, G., 1984. Ecoéthologie du Mouflon et de l'Isard dans le massif du Carlit (Pyrénées Orientales), PhD thesis, Université Paul Sabatier. Toulouse.
69. Gosling, L.M., 1969. Parturition and related behaviour in Coke's Hartebeest. Alcelaphus buselaphus cokie Günther, J. Reprod. Fert., Suppl., 6: 265-286.
70. Gould, S.J., 1977. Ontogeny and phylogeny. Harvard University Press, Cambridge, MA.
71. Green, W.C.H., 1992. The development of independence in bison: pre-weaning spatial relations between mothers and calves. Anim. Behav., 43: 759-773.
72. Grubb, P., 1974a. The rut and behaviour of Soay rams. In: P.A. Jewell, C. Milner and J.M. Boyde (Editors). Island Survivors: the Ecology of the Soay Sheep of St. Kilda, Athlone Press, London. pp. 195-223.
73. Grubb, P., 1974b. Social organization of Soay sheep and the behaviour of ewes and lambs. In: P.A. Jewell, C. Milner and J.M. Boyde (Editors), Island Survivors: the Ecology of the Soay Sheep of St. Kilda, Athlone Press, London, pp. 131-159.
74. Grubb, P., 1974c. Mating activity and the social significance of rams in a feral sheep community. In: V. Geist and F. Walther (Editors), The Behaviour of Ungulates and its Relation to Management, IUCN No. 50, Morges, Switzerland, pp. 457-476.
75. Grubb, P. and Jewell, P.A., 1966. Social grouping and home range in feral soay sheep. Symp. Zool. Soc. Lond., 18: 179-210.
76. Guinness, F.E., Hall, M.J. and Cockerill, R.A., 1979. Mother-offspring association in red deer (Cervus elaphus L.) on Rhum. Anim. Behav., 27: 536-544.
77. Hall, S.J.G. 1989. Chillingham cattle: social and maintenance behaviour in an ungulate that breeds all year round. Anim. Behav., 38: 215-225.
78. Hass, C.C. and Jenni, D.A., 1991. Structure and ontogeny of dominance relationships among bighorn rams. Can. J. Zool., 69: 471-476.
79. Hass, C.C. and Jenni, D.A., 1993. Social play among juvenile bighorn sheep: structure, development, and relationship to adult behavior. Ethology. 93: 105-116.
80. Hinch, G.N., Lécrivain, E., Lynch, J.J. and Elvin, R.L., 1987. Changes in maternal-young associations with increasing age of lambs. Appl. Anim. Behav. Sci., 17: 305-318.
81. Hinde, R.A., 1974. Biological bases of human social behaviour, McGraw-Hill, New York.
82. Hinde, R.A., 1976. Interactions, relationships and social structure. Man, II: 1-17.
83. Hirth, D.H., 1977. Social behavior of white-tailed deer in relation to habitat. Wildl. Monogr., 53: 1-55.
84. Hirth, D.H., 1985. Mother-young behaviour in white-tailed deer, Odocaileus virginianus. South. Nat., 30: 297-302.
85. Hoffmann, R., 1985. Social behaviour in immature males of a feral horse population. Z. Säugetierkunde, 50: 304-314.
86. Hogg, J.T., 1987. Intrasexual competition and mate choice in Rocky Mountain bighorn sheep. Ethology, 75: 119-144.
87. Holekamp, K.E. and Sherman, P.W., 1989. Why male ground squirrels disperse? Am. Sci., 77: 232-239.
88. Horwich, R.H., van Dyke, R. and Cogswell, S.J.H., 1977. Regressive growth periods as a mechanism for herd formation in Siberian ibex (Capra ibex). Zool. Garten, 47: 59-68.
89. Illius, A.W. and Gordon, I.J., 1987. The allometry of food intake in grazing ruminants. J. Anim. Ecol., 56: 989-999.
90. Illius, A.W. and Gordon, I.J., 1990. Variation in foraging behaviour in red deer and the consequences for population demography. J. Anim. Ecol., 59: 89-101.
91. Jacklin, C.N. and Maccoby, E.E., 1978. Social behaviour at thirty-three months in same-sex and mid-sex dyads. Child Dev., 49: 557-569.
92. Jakimchuk, R.D., Fergusson, S.H. and Sopuck, L.G., 1987. Differential habitat use and sexual segregation in the Central Arctic caribou herd. Can. J. Zool., 65: 534-541.
93. Jarman, P.J., 1974. The social organisation of antelop in relation to their ecology. Behaviour, 48: 215-266.
94. Jarman, P.J., 1983. Mating system and sexual dimorphism in large, terrestrial, mammalian herbivores. Biol. Rev., 58: 485-520.
95. Jewell, P., 1986. Survival in a feral population of primitive sheep on St. Kilda, Outer Hebrides, Scotland. Natl. Geogr. Res., 2: 402-406.
96. Krausman, P.R., Leopold, B.D., Seegmiller, R.F. and Torres, S.G., 1989. Relationships between desert bighorn sheep and habitat in western Arizona. Wildl. Monogr., 102: 1-66.
97. Langman, V.A., 1977. Cow-calf relationships in giraffe (Giraffa camelopardalis giraffa). Z. Tierpsychol., 43: 264-286.
98. Lawrence, A.L., 1990. Mother-daughter and peer relationships of Scottish hill sheep. Anim. Behav., 39: 481-486.
99. Leberg, P.L. and Smith, M.H., 1993. Influence of density on growth of white-tailed deer. J. Mamm., 74: 723-731.
100. Leclerc, B. and Lécrivain, E., 1979. Etude de comportements d'ovins domestiques en élevage extensif sur le Causse du Larzac. PhD thesis, Université Rennes, Rennes.
101. Lee, P.C., 1986. Early social development among African elephant calves. Natl. Geogr. Res., 2: 388-401.
102. Legault, F. and Strayer, F.F., 1991. Genèse de la ségrégation sexuelle et différences comportementales chez des enfants d'âge préscolaire. Behaviour, 119: 285-301.
103. Lent, P.C., 1974. Mother-infant relationships in ungulates. In: V. Geist and F. Walther (Editors), The Behaviour of Ungulates and its Relation to Management, Vol. I , IUCN No. 50, Morges, Switzerland, pp. 14-55.
104. Lenarz, M.S., 1979. Social structure and reproductive strategy in desert bighorn sheep (Ovis canadensis mexicana). J. Mamm., 60: 671-678.
105. Leslie, D.M. and Douglas, C.L., 1979. Desert Bighorn sheep on the River Mountains, Nevada. Widl. Monogr., 66: 1-56.
106. Leuthold, W., 1977. Spatial organization and strategy of habitat utilization of elephants in Tsavo National Park, Kenya. Z. Säugertierkunde, 42: 358-379.
107. Lidfors, L. and Jensen, P., 1988. Behaviour of free-ranging beef cows and calves. Appl. Anim. Behav. Sci., 20: 237-247.
108. Lott, D.F. and Minta, S.C., 1983. Random individual association and social group instability in American bison (Bison bison). Z. Tierpsychol., 61: 153-172.
109. Lowe,. V.P.M., 1966. Observations on the dispersal of red deer on Rhum. In: P.A. Jewell and C. Loizos (Editors), Play, Exploration and Territory in Mammals, Symp. Zool. Soc., Lond., 18: pp. 179-210.
110. Main, M.B. and Coblentz, B.E., 1990. Sexual segregation among ungulates: a critique. Wildl. Soc. Bull., 18: 204-210.
111. Mason, W.A., 1978. Ontogeny of social systems. In: D.J. Chivers and J. Herbert (Editors), Recent Advances in Primatology, Vol. I. Behaviour, Academic Press. London, pp. 5-14.
112. Meaney, M.J., 1988. The sexual differentiation of social play. Trends Neurosci., 11: 54-58.
113. Meaney, M.J., Stewart, J. and Beatty, W.W., 1985. Sex differences in social play: the socialization of sex roles. Adv. Study Behav., 15: 1-58.
114. Miquelle, D.G., Peek, J.M. and Van Ballenberghe, V., 1992. Sexual segregation in Alaskan moose. Wildl. Monogr., 122: 1-57.
115. Moore, C.L., 1985. Development of mammalian sexual behavior. In: E.S. Gollin (Editor), The Comparative Development of Adaptative Skills: Evolutionary implications, Lawrence Erlbaum Associate, Hillsdale, NJ, pp. 19-56.
116. Mooring, M.S., 1989. Ontogeny of allogrooming in mule deer (Odocoileus hemionus). J. Mamm., 70: 434-437.
117. Morris, D.W., 1984. Sexual differences in habitat use by small mammals: evolutionary strategy or reproductive constraint? Oecologia, 65: 51-57.
118. Morton, E.S. 1990. Habitat segregation by sex in the hooded warbler: experiments on proximate causation and discussion of its evolution. Am. Nat., 135: 319-333.
119. Moss, C.J. and Poole, J.H., 1983. Relationships and social structure of African elephants. In: R.A. Hinde (Editor), Primate Social Relationships, Blackwell, Oxford. pp. 315-325.
120. Muller-Schwarze, D. and Muller-Schwarze, C., 1983. Play behaviour in free-ranging caribou, Rangifer tarandus. Acta Zool. Fennica, 175: 121-124.
121. Newberry, R.C. and Wood-Gush, D.G.M., 1986. Social relationships of piglets in a semi-natural environment. Anim. Behav., 34: 1311-1318.
122. Nievergelt, B., 1967. Die Zusammensetzung der Gruppen beim Alpensteinbock. Z. Säugetierkunde, 32: 129-144.
123. Nievergelt, B., 1974. A comparison of rutting behaviour and grouping in the Ethiopian and Alpine ibex. In: V. Geist and F. Walther (Editors). The Behaviour of Ungulates and its Relation to Management, IUCN No. 50, Morges, pp. 324-340.
124. Ordway, L.L. and Krausman, P.R., 1986. Habitat use by mule deer. J. Wildl. Manage., 50: 677-683.
125. Orgeur, P. and Signoret, J.P., 1984. Sexual play and its functional significance in the domestic sheep (Ovis aries L.). Physiol. Behav., 33: 111-118.
126. Ozoga, J.J. and Verme, L.J., 1985. Comparative breeding behavior and performance of yearling vs. prime-age white-tailed bucks. J. Wildl. Manage., 49: 364-372.
127. Pfeifer, S., 1985. Sex differences in social play of scimitar-horned oryx calves (Oryx dammah). Z. Tierpsychol., 69: 281-292.
128. Poindron, P., Lévy, F. and Krehbiel, D., 1988. Genital, olfactory, and endocrine interactions in the development of maternal behaviour in the parturient ewe. Psychoneuroendocrinology, 13: 99-125.
129. Polis, G.A., 1984. Age structure component of niche width and intra-specific resource partitioning: can age groups function as ecological species? Am. Nat., 123: 541-564.
130. Pratt, D.M. and Anderson, V.H., 1979. Giraffe cow-calf relationships and social development of the calf in the Serengeti. Z. Tierpsychol., 51: 233-251.
131. Pulliam, H.R. and Caraco. T., 1984. Living in groups: is there an optimal group size? In: J.R. Krebs and N.B. Davies (Editors), Behavioural Ecology: An Evolutionary Approach, Blackwell, Oxford, pp. 122-147.
132. Reinhardt, V. and Reinhardt,. A., 1981. Cohesive relationships in a cattle herd (Bos indicus). Behaviour, 77: 121-151.
133. Richard-Hansen, C., 1992. Association between individually marked isards (Rupicapra pyrenaica): seasonal and inter-annual variations. In: F. Spitz, G. Janeau, G. Gonzalez and S. Aulagnier (Editors), Ongulés/Ungulates 91, SFEPM-IRGM, Paris-Toulouse: pp. 299-304.
134. Richard-Hansen, C., 1993. Social interactions in Isard kids Rupicapra pyrenaica pyrenaica (Artiodactyla: Bovidae): possible influence on development of social bonding. Ethology, 94: 291-305.
135. Rothstein, A. and Griswold, J.G., 1991. Age and sex preferences for social partners by juvenile bison bulls, Bison bison. Anim. Behav., 41: 227-237.
136. Rutberg, A.T., 1986. Dominance and its fitness consequences in American bison cows. Behaviour, 96: 62-91.
137. Rutberg, A.T. and Keiper, R.K., 1993. Proximate causes of natal dispersal in feral ponies: some sex differences. Anim. Behav., 46: 969-975.
138. Sachs, B.D. and Harris, V.S., 1978. Sex differences and developmental changes in selected juvenile activities (play) of domestic lambs. Anim. Behav., 26: 678-684.
139. Scarbrough, D.L. and Krausman, P.R., 1988. Sexual segregation by desert mule deer. Southwestern Nat., 33: 157-165.
140. Seip, D.R. and Bunnell, F.L., 1985. Foraging behaviour and food habits of Stone's sheep. Can. J. Zool., 63: 1638-1646.
141. Shackleton, D.M., 1991. Social maturation and productivity in bighorn sheep: are young males incompetent. Appl. Anim. Behav. Sci., 29: 173-184.
142. Shackleton, D.M. and Shank, C.C., 1984. A review of the social behavior of feral and wild sheep and goats. J. Anim. Sci., 58: 500-509.
143. Shank, C.C., 1982. Age-sex differences in the diets of wintering Rocky Mountain bighorn sheep. Ecology, 63: 627-633.
144. Shank, C.C., 1985. Inter-and intra-sexual segregation of chamois (Rupicapra rupicapra) by altitude and habitat during summer. Z. Säugetierkunde, 50: 117-125.
145. Shillito Walser, E., Hague, P. and Yeomans, M., 1983. Preferences for sibling or mother in Dalesbred and Jacob twin lambs. Appl. Anim. Ethol., 9: 289-297.
146. Sinclair, A. R. E., 1977. The African Buffalo. A Study of Resource Limitation of Populations. University Chicago Press, Chicago, IL.
147. Staines, B.W., Crisp, J.M. and Parish, T., 1982. Differences in the quality of food eaten by red deer (Cervus elaphus) stags and hinds in winter. J. Appl. Ecol., 19: 65-77.
148. Teillaud, P., 1986. Stratégies alimentaires et statut social chez le sanglier en captivité. Behav. Process., 12: 327-347.
149. Tulloch, D.G., 1979. The water buffalo, Bubalus babalis, in Australia: reproductive and parent-offspring behaviour. Aust. Wildl. Res., 6: 265-287.
150. Veissier, I., Le Neindre, P. and Garel, J.P., 1990. Decrease in cow-calf attachment after weaning. Behav. Process., 21: 95-105.
151. Villaret, J.C. and Bon, R., in press. Social and spatial segregation in Alpine ibex (Capra ibex) in Bargy. French Alps. Ethology.
152. Watson, A. and Staines, B.W., 1978. Differences in the quality of wintering areas used by male and female red deer (Cervus elaphus) in Aberdeenshire. J. Zool., Lond., 286: 544-550.
153. Weckerly, F.W., 1993. Intersexual resource partitioning in black-tailed deer: a test of the body size hypothesis. J. Wildl. Manage., 57: 475-494.
154. Wiley, R.H., 1981. Social structure and individual ontogenies: problems of description, mechanism, and evolution. In: P.P.G. Bateson and P.H. Klopfer (Editors), Perspectives in Ethology (Vol. 4), Plenum. New York, NY, pp. 105-133.
155. Wood-Gush, D.G.M., Hunt, K., Carson, K. and Dennison, S.G.C., 1984. The early behaviour of suckler calves in the field. Biol. Behav., 9: 295-306.