96
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).
97
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).
98
Bighorn Sheep (Hogg 1987:120; Hass and Jenni 1991:471); Crested Black Macaque (Poirier 1964:54).
99
Vasey, “Homosexual Behavior in Primates,” p. 191.
100
Orang-utan (Maple 1980:118).
101
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.
102
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).
103
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.
104
Redshank (Hale and Ashcroft 1983:21). For a summary of the historical interpretation of this behavior, see also Cramp and Simmons 1983:533.
105
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.
106
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”).
107
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).
108
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).
109
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 Hoyo, J., A. Elliott, and J. Sargatal, eds. [1996] Handbook of the Birds of the World , vol. 3: Hoatzin to Auks, p. 282. [Barcelona: Lynx Edicións]); Orang-utan (Galdikas 1981:286).
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Walrus (Dittrich 1987:168); Musk-ox (Smith 1976:62); Bighorn Sheep (Hogg 1984:527; Geist 1971:139); Asiatic Mouflon (McClelland 1991:81); Grizzly Bear (Craighead et al. 1995:161); Olympic Marmot (Barash 1973:212); White-tailed Deer (Hirth 1977:43); Orang-utan (Galdikas 1981:286); White-faced Capuchin (Manson et al. 1997:775); Northern Fur Seal (Gentry 1998:172); Ruff (Hogan-Warburg 1966:167-68). Additionally, in one study of Matschie’s Tree Kangaroos—a species in which researchers deny that mounting between females is (homo)sexual (J. Steenberg, personal communication)—all mounts observed between animals of the opposite sex were “incomplete” in that they did not involve penetration or thrusting (Hutchins et al. 1991:158). Another study of the same population found both that “full” copulations between males and females were infrequent, and that in heterosexual contexts females showed few overt signs of sexual interest, since the behavioral cues for female sexual arousal are extremely subtle (Dabek 1994:84, 93—94, 116).
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Morrill and Robertson 1990 (Tree Swallow); Scott, M. P., and T. N. Tan (1985) “A Radiotracer Technique for the Determination of Male Mating Success in Natural Populations,” Behavioral Ecology and Sociobiology 17:29-33. More recently, a copulation-verification technique using fluorescent powder has been tested for rodents. Dusted on males, the powder is transferred to females during mating and can be checked using ultraviolet light. Ironically, during the testing of this procedure, pairs of females were used as “controls” since it was assumed that they would not engage in mounting behavior with one another. Nevertheless, 12 percent of female pairs showed transfer of powder—but of course this was interpreted by researchers as evidence of nonsexual contact between such females (Ebensperger, L. A., and R. H. Tamarin [1997] “Use of Fluorescent Powder to Infer Mating Activity of Male Rodents,” Journal of Mammalogy 78:888-93).
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