Biological Exuberance: Animal Homosexuality and Natural Diversity

Crab-eating Macaque (Poirier and Smith 1974); Pukeko (Craig 1980:594); Rhesus Macaque (Lindburg 1971:14, 69); Tree Swallow (Stutchbury and Robertson 1985, 1987b); Galah (Rogers and McCulloch 1981:90); Scarlet Ibis (Elbin and Lyles 1994:90–91); Flamingo (King 1994:104–5); Nilgiri Langur (Hohmann 1989:449); Little Egret (M. Fujioka, personal communication); Little Blue Heron (Werschkul 1982:382).

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Black Stilt (Reed 1993:772); Humboldt Penguin (Scholten 1992:6 and personal communication); Savanna (Yellow) Baboon (Rowell 1967a:16, 22—23 [tables 2, 3]); Mallard Duck (Lebret 1961:108 [table I]).

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Pig-tailed Macaque (Oi 1990a:340); Bottlenose Dolphin (Wells 1991:222); Cheetah (Eaton and Craig 1973:252); Koala (Smith 1980:184); Canada Goose (Collias and Jahn 1959:484); Flamingo (C. E. King, personal communication); Lesser Flamingo (Alraun and Hewston 1997:175–76).

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Japanese Macaque (Chapais and Mignault 1991:172; Wolfe 1984:155); Giraffe (Dagg and Foster 1976:28, 124, 144); Greylag Goose (Huber and Martys 1993:160). Likewise, in Northern Fur Seal populations with up to 40 or more females for every male, a number of behavioral and other factors insure that nearly every female is still able to mate heterosexually (Gentry 1998:167, 192–93). For Macaques, some researchers have suggested that females resort to homosexuality when deprived of “novel” male partners rather than of males per se (i.e., when they “run out” of new partners or become overly familiar with them) (Wolfe 1984:155, 1986:274 [Japanese Macaque]; Huynen 1997 [Rhesus Macaque]). As Vasey (1996:550) points out, however, this explanation is flawed because the females they turn to are no more “novel” than the males are (and probably even less so, owing to the high levels of female bonding and familiarity in these species). In addition, some females continue to choose other females as partners even in populations that have novel males.

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Gorilla (Robbins 1996; Fossey 1983, 1984; Harcourt et al. 1981); Hanuman Langur (Weber and Vogel 1970) Crested Black Macaque (Reed et al. 1997; Dixson 1977); Squirrel Monkey (DuMond 1968; Travis and Holmes 1974; Akers and Conaway 1979; Denniston 1980; Mendoza and Mason 1991); Walrus (Miller and Boness 1983; Sjare and Stirling 1996); Lion (Schaller 1972; Chavan 1981); Mallard Duck (Bossema and Roemers 1985; Schutz 1965:457–59); Black-headed Gull (Kharitonov and Zubakin 1984); West Indian Manatee (Hartman 1971, 1979); Cheetah (Eaton and Craig 1973; Eaton 1974a). In some of these cases (e.g., Gorillas, Hanuman Langurs) homosexual activity among males is much more common in same-sex groups although it still occurs sporadically or “residually” in mixed groups; in other cases (e.g., Squirrel Monkeys, Crested Black Macaques) homosexual activity is equally if not more common in at least some mixed-sex groups.

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Squirrel Monkey (Talmage-Riggs and Anschel 1973:68, 71); Long-eared Hedgehog (Poduschka 1981:81).

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Silver Gull (Mills 1991:1523, 1526); Mallard Duck (Schutz 1965:442); Canada Goose (Collias and Jahn 1959:500); Jackdaw (Röell 1979:126, table 1); Lesser Scaup Duck (Bellrose 1976:344); Caribou (Bergerud 1974:432).

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Flamingo (Wilkinson 1989:53-54; King 1994:105; C. E. King, personal communication); Laughing Gull (Hand 1981:138-39); Humboldt Penguin (Scholten 1992:5); Gentoo Penguin (Stevenson 1983:192); Pied Kingfisher (Moynihan 1990:19; Reyer 80:220); Peach-faced Lovebird (Fischdick et al 1984:314); Galah (Rogers and McCulloch 1981:90); Bicolored Antbird (Willis 1967:112).

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Cattle Egret (Fujioka 1986b:421-22); emperor and other penguins (Williams, T. D. [1995] The Penguins: Spheniscidae, pp. 80, 160 [Oxford: Oxford University Press]); dipper (Wilson, J. D. [1996] “The Breeding Biology and Population History of the Dipper Cinclus cinclus on a Scottish River System,” Bird Study 43:108—18); Oystercatcher (Heg and van Treuren 1998); Australian noisy miner (Dow, D. D., and M. J. Whitmore [1990] “Noisy Miners: Variations on the Theme of Communality,” in P. B. Stacey and W. D. Koenig, eds., Cooperative Breeding in Birds: Long-Term Studies in Behavior, pp. 559--92 [Cambridge: Cambridge University Press]); spotted sandpiper (Oring, L. W., J. M. Reed, and S. J. Maxson [1994] “Copulation Patterns and Mate Guarding in the Sex-Role Reversed, Polyandrous Spotted Sandpiper, Actitis macularia,” Animal Behavior 47:1065-72).

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