Biological Exuberance: Animal Homosexuality and Natural Diversity

Useful Construct or Bad Habit?” in E. Klingham-mer, ed., Behavior and Ecology of Wolves, pp. 225-44 (New York: Garland); Baldwin and Baldwin 1981 (Squirrel Monkey); Bernstein, I. S. (1981) “Dominance: The Baby and the Bathwater,” Behavioral and Brain Sciences 4:419-57; Hand, J. L. (1986) “Resolution of Social Conflicts: Dominance, Egalitarianism, Spheres of Dominance, and Game Theory,” Quarterly Review of Biology 61:201-20; Walters, J. R., and R. M. Seyfarth (1987) “Conflict and Cooperation,” in B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham, and T. T. Struhsaker, eds., Primate Societies, pp. 306-17 (Chicago and London: University of Chicago Press); Drews, C. (1993) “The Concept and Definition of Dominance in Animal Behavior,” Behavior 125:283-313; Lambert et al. 1994 (Pukeko).

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Fedigan 1982:92-93 (Japanese Macaque).

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Bonobo (Kano 1992:253-54; Kitamura 1989:57, 63); Gorilla (Harcourt et al. 1981:276; Yamagiwa 1987a:25; Harcourt 1988:59); Hanuman Langur (J. J. Moore, in Weinrich 1980:292); Japanese Macaque (Vasey 1996:549; Chapais and Mignault 1991:175-76; Tartabini 1978:433, 435; Hanby 1974:841); Rhesus Macaque (Akers and Conaway 1979:78; Reinhardt et al. 1986:55; Gordon and Bernstein 1973:224); Pig-tailed Macaque (Tokuda et al. 1968:293); Crested Black Macaque (Dixson 1977:77-78; Poirier 1964:20, 49; Reed et al. 1997:255); Savanna Baboon (Owens 1976:256); Gelada Baboon (Mori 1979:134-35; R.Wrangham, in Weinrich 1980:291); Squirrel Monkey (Talmage-Riggs and Anschel 1973:70); Bottlenose Dolphin (Caldwell and Caldwell 1972:427); Blackbuck (Dubost and Feer 1981:89-90); Giraffe (Pratt and Anderson 1985:774— 75, 780); American Bison (Reinhardt 1985:222, 1987:8); Red Squirrel (Ferron 1980:136); Little Blue Heron (Werschkul 1982:383-84); Tree Swallow (Lombardo et al. 1994:556).

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For examples of earlier claims of a dominance connection being refuted by later studies, see Common Chimpanzee (Yerkes 1939:126-27; Nishida 1970:57—Bygott 1974 [cited in Hanby 1974:845 (Japanese Macaque)]; Nishida and Hosaka 1996:122 [table 9.7]); Hanuman Langur (Weber 1973:484—Srivastava et al. 1991:506— 7; J. J. Moore, in Weinrich 1980:292); Rhesus Macaque (Carpenter 1942—Akers and Conaway 1979:78; Reinhardt et al. 1986; Gordon and Bernstein 1973:224); Japanese Macaque (Sugiyama 1960:136—Hanby 1974:841; Chapais and Mignault 1991:175-76); Bonnet Macaque (Rahaman and Parthasarathy 1968:68, 263—Makwana 1980:10); Pig- tailed Macaque (Tokuda et al. 1968—Oi 1990a:353-54); Killer Whale (Balcomb et al. 1979:23—Rose 1992:108-9); Giraffe (Dagg and Foster 1976, Leuthold 1979:27—29—Pratt and Anderson 1985:774-75); Blackbuck (Schaller 1967—Dubost and Feer 1981:89—90); American Bison (Lott 1974:391—Reinhardt 1986:222-23); Wolf (Schenkel 1947—van Hooff and Wensing 1987:232). In addition, a parallel example in Laughing Gulls involves an indirect refutation of the relevance of dominance. Noble and Wurm (1943:205-6) linked homosexual mounting in Laughing Gulls to the supposedly lower rank of the male being mounted, citing as evidence of his lower status the fact that the mounted male did not “dominate” his female mate. In a more recent detailed study of interactions between partners in heterosexual pairs, however, Hand (1985) concluded that males do not in general dominate their female mates in this species—thus invalidating the earlier claim that being mounted homosexually was correlated with “lower status.” Studies that attribute homosexual activity to dominance with little or no supporting evidence include Orang-utan (Rijksen 1978:257); Squirrel Monkey (DuMond 1968:124); West Indian Manatee (Rathbun et al. 1995:150); Pied Kingfisher (Moynihan 1990:19).

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Whiptail Wallaby (Kaufmann 1974:307, 309); Rhesus Macaque (Gordon and Bernstein 1973:224). Kaufmann concluded that Whiptail Wallaby homosexual mountings are themselves probably not dominance-related, however, because dominant animals generally invite subordinate ones to mount (the opposite of the “usual” dominance pattern).

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Bighorn Sheep (Hogg 1987:120; Hass and Jenni 1991:471); Crested Black Macaque (Poirier 1964:54).

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Vasey, “Homosexual Behavior in Primates,” p. 191.

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Orang-utan (Maple 1980:118).

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West Indian Manatee (Rathbun et al. 1995:150). See also the suggestion in Buss (1990:19-21) that sexual arousal in male African Elephants during same-sex play-fighting serves to dull pain. While this is possible, it is rather far-fetched, considering that such ritual fights (described by Buss as “erotic”) are rarely violent.

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Vasey, P. L. (in press) “Homosexual Behavior in Male Birds,” in W. R. Dynes, ed., Encyclopedia of Homosexuality, 2nd ed., vol. 1: Male Homosexuality (New York: Garland Press).

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American Bison (Reinhardt 1985:222) (cf. also Kaufmann [1974:107] on Whiptail Wallabies, who asserts, “Though tail-lashing seems clearly a sign of sexual arousal, it was occasionally performed by males when they were approached by subordinate males in nonsexual situations”); Asiatic Mouflon (McClelland 1991:80); Stumptail Macaque (O’Keefe and Lifshitz 1985:149); Dugong (Nair et al. 1975:14); Laysan Albatross (Frings and Frings 1961:311); Dwarf Mongoose (Rasa 1979a:365); Bonnet Macaque (Nolte 1955:179). Similarly, Frank et al. (1990:308) state that genital erections in Spotted Hyenas have no “sexual significance” unless displayed by a male toward a female during courtship. The “desexing” of this behavior stems, in large part, from the fact that erections are frequently displayed between animals of the same sex (especially females) and in situations that do not involve (heterosexual) mounting (e.g., during the “meeting ceremony”). While erections undoubtedly have “nonsexual” connotations outside of a mounting context (see, for example, East et al. 1993), it seems overly restrictive to eliminate all “sexual significance” from situations that do not fall into the category of heterosexual courtship and mating.

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Redshank (Hale and Ashcroft 1983:21). For a summary of the historical interpretation of this behavior, see also Cramp and Simmons 1983:533.

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Crested Black Macaque (Dixson 1977:71, 76; Poirier 1964:147). Dixson (1977:77) does concede that the distinction between sexual and nonsexual mounts and solicitations is a subjective one, but only in heterosexual contexts—homosexual interactions are assumed to be self-evidently nonsexual.

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Vicuna (Koford 1957:183, 184); Musk-ox (Smith 1976:51); Giraffe (Dagg and Foster 1976:127; Pratt and Anderson 1985:777-78; Leuthold 1979:27, 29); Bank Swallow (Beecher and Beecher 1979:1284); Savanna Baboon (Smuts 1985:18, 148—49, 163-66, 199, 213); Rhesus Macaque (Loy 1971:26); Oystercatcher (Makkink 1942; Ens and Goss-Custard, “Piping as a Display of Dominance”).

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Crested Black Macaque (Dixson 1977:70—71); Bottlenose Dolphin (Ostman 1991:313; Dudok van Heel and Mettivier 1974:12; Saayman and Tayler 1973); Spinner Dolphin (Norris and Dohl 1980a:845; Norris et al. 1994:199); Common Murre (Birkhead 1978a:326); Blue-bellied Roller (Moynihan 1990).

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Rhesus Macaque (see, for example, Sade 1968:32-33); Japanese Macaque (Hanby 1974:843, 845; Wolfe, “Human Evolution and the Sexual Behavior of Female Primates,” p. 129).

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For further discussion see chapter 5. On a related point, aggressive behaviors may accompany homosexual interactions in some species and are therefore used to argue that such behavior is not “really” sexual. However, aggression is also characteristic of heterosexual relations in many species, where such male-female interactions are still classified as “sexual.”

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Kob (Buechner and Schloeth 1965:218); Giraffe (Pratt and Anderson 1985:774-75); northern jacana (del