Biological Exuberance: Animal Homosexuality and Natural Diversity

23

Ocellated Antbird (Willis 1973:31); on heterosexual divorce in Antbirds, see Willis (1983:414).

24

Gorilla (Fischer and Nadler 1978:660-61); Western Gull (Hunt et al. 1984:160); Guianan Cock-of-the-Rock (Trail 1985a:238, 240); Red Fox (Macdonald 1987:101); de Waal 1989a:25 (Bonobo). For descriptions of nonstandard mounting positions (lateral, head-to-tail) in heterosexual contexts, see (for example) Japanese Macaque (Hanby and Brown 1974:156, 164); Boto (Best and da Silva 1989:15); Bottlenose/Spotted Dolphins (Herzing and Johnson 1997:92, 96); Waterbuck (Spinage 1969:41-42); Mountain Sheep (Geist 1971:139— 40); Mountain Goat (Hutchins 1984:268); Takhi (Boyd and Houpt 1994:202); Collared Peccary (Byers and Bekoff 1981:771); Warthog (Cumming 1975:118-19); Koala (Smith 1980c:48); Ruff (Hogan-Warburg 1966:176); Hammerhead (Brown 1982:171; Campbell 1983:11); Flamingo (Shannon 1985:229); Chaffinch (Marler 1956:114); red-winged blackbird (Monnett, C., L. M. Rotterman, C. Worlein, and K. Halupka [1984] “Copulation Patterns of Red-winged Blackbirds [Agelaius phoeniceus],” p.759, American Naturalist 124:757— 64). Of these, subjective or derogatory terms are only used in Monnett et al. 1984 (“abnormal,” “aberrant”) and Hutchins 1984 (“clumsy,” “awkward”). Nonstandard homosexual mounting positions such as sideways or head-to-tail mounts have usually been classified as “mistakes” or “incomplete” mounting attempts by zoologists who insist on viewing homosexual interactions only in terms of how closely they resemble “standard” heterosexual intercourse. In other words, anything that deviates from genital penetration (or cloacal contact in birds) in the front-to-back position used by males with females is an “error.” Because these mounting positions are often used by female animals (when they mount individuals of either sex), a further, sexist, interpretation is also frequently overlaid on these behaviors: it is claimed that they represent an “imperfect” attempt on the part of females to imitate male mounting behavior. An equally valid perspective, however, is that these represent alternative or more “fluid” sexual interchanges—not bound by the “requirement” of genital penetration—rather than flawed imitations of heterosexual postures. A parallel example can be found in the behavior of “sideways presenting” in female Crab-eating Macaques: previously classified as “disoriented” or “inadequate,” this posture was later found to be a systematic behavioral variant (Emory and Harris 1978). For further discussion and a critique of the widespread view that homosexuality is an imperfect approximation of heterosexuality, see chapter 4. For evidence that heterosexual sex is not focused exclusively on vaginal penetration and ejaculation, see chapter 5.

25

Laughing Gull (Hand 1981:138-39); Black-headed Gull (van Rhijn and Groothuis 1985:161); Herring Gull (Shugart et al. 1988:934); inclusion of infertile eggs in hatching rates of female pairs: Kovacs and Ryder 1983:661- 62, Ryder and Somppi 1979:3 (Ring-billed Gull); Burger, J., and M. Gochfield (1996) “Laridae (Gulls),” p. 584, in J. del Hoyo, A. Elliott, and J. Sargatal, eds., Handbook of the Birds of the World, vol. 3, Hoatzin to Auks, pp. 572-623 (Barcelona: Lynx Edicions); shared characteristics of heterosexual and homosexual supernormal clutches: Kovacs and Ryder 1983:660-62, Lagrenade and Mousseau 1983, Ryder and Somppi 1979:3 (Ring-billed Gull and other species) (on the lower productivity of supernormal clutches attended by heterosexual pairs in species other than Gulls, see Sordahl, T. A. [1997] “Breeding Biology of the American Avocet and Black- necked Stilt in Northern Utah,” pp. 350, 352, Southwestern Naturalist 41:348— 54); equivalent parenting abilities of homosexual and heterosexual pairs: Hunt and Hunt 1977:1467, Hayward and Fry 1993:17-18 (Western Gull); Conover 1989:148 (Ring-billed Gull); Nisbet et al. 1998:314 (Roseate Tern); “runaways” from heterosexual parents: Pierotti and Murphy 1987 (Western Gull and other species); Brown et al. 1995 (Ring-billed Gull); Roberts and Hatch 1994 (Kittiwake).

26

Gray Whale (Darling 1977:10—11).

27

In fact, it can safely be said that no scientific study of wild animals has yet been undertaken with the expectation that homosexual activity would be observed—same-sex behavior is invariably a “surprise.” In contrast, many a field study has been initiated for the express purpose of studying heterosexual mating—and has quite often been treated to the unexpected occurrence of same-sex activity and/or the absence (or rarity) of opposite-sex interactions.

28

Laughing Gull (Burger and Beer 1975:312); Common Murre (Hatchwell 1988:167); Kittiwake (Chardine 1986:1416, 1987:516); Griffon Vulture (Blanco and Martinez 1996:247).

29

Grebes (Nuechterlein and Storer 1989:344—45).

30

For a recent example concerning a little-known species, see Dyrcz, A. (1994) “Breeding Biology and Behavior of the Willie Wagtail Rhipidura leucophrys in the Mdang Region, Papua New Guinea,” Emu 94:17—26.

31

Emu (Heinroth 1924, 1927); Regent Bowerbird (Gilliard 1969:341); Dugong (Jones 1967; Nair et al. 1975:14). The visual resemblance between younger male and adult female Superb Lyrebirds has also resulted in some misidentifications and revised interpretations of this species’ behavior in the wild. Although Smith (1968:88—89, 1988:30—32, 75—78) and Lill (1979a:496) state clearly (and offer photographic documentation) that adult males court (and even mount) younger males, the identification of some individuals has not been so straightforward. One bird photographed as it was being courted by an adult male (including full courtship displays) was first identified as “possibly” a male (Smith 1968:60), then as a female (Smith 1988:30). However, after a careful review of the plumage characteristics of adult females and younger males, L. H. Smith has confirmed (personal communication) that the younger bird in this case was indeed a male and in fact was most likely the adult male’s own son. Unfortunately, the earlier published reports in which the sex of the younger bird was unclear may have led Reilly (1988:32) to state erroneously that males never perform full courtship displays toward other males. For additional photographs of males performing full displays to other males, see Smith (1988:77) and p. 13 (this book).

32

King Penguin (Gillespie 1932:96-120).

33

Snow Goose (Quinn et al. 1989); Ring-billed Gull (Kovacs and Ryder 1981); Red-backed Shrike (Pounds 1972); Blue Tit (Blakey 1996); Guianan Cock-of-the-Rock (Trail and Koutnik 1986); Stumptail Macaque (Chevalier- Skolnikoff 1976:522 [table III]); Jackdaw (Roell 1979:126-27); Cheetah (Eaton and Craig 1973:248, 250); Bonobo (Parish 1996:65, 86; de Waal 1997:112—15). Similarly, in citing Hartman’s (1971) original descriptions of homosexuality in West Indian Manatees, Ronald et al. (1978:37) focus on examples of same-sex activity that occur in conjunction with heterosexual behaviors and downplay those that are independent of opposite-sex encounters (even though such independent encounters are equally, if not more, prevalent). On a related point, genes that are thought to control homosexual activity in Fruit Flies have been given names by scientists that refer solely to their (negative) effect on heterosexuality and breeding. One gene has been labeled dissatisfaction (alluding to the fact that carriers of this gene, in addition to being interested in homosexual activity, typically refuse or are “dissatisfied” with heterosexual advances), while another has been called fruitless (alluding to the fact that carriers, in addition to courting individuals of both sexes, are infertile) (Finley et al. 1997:917).

34

Savanna (Olive) Baboon (Owens 1976:254); Right Whale (Clark 1983:169); Moose (Van Ballenberghe and Miquelle 1993:1688); Cattle Egret (Fujioka and Yamagishi 1981:136).

35

Squirrel Monkey (Talmage-Riggs and Anschel 1973:70-71); Bonobo (Savage-Rumbaugh and Wilkerson 1978:338; Savage and Bakeman 1978:614); Spotted Hyena (Burr 1996:118-19). For conflicting information on the occurrence of clitoral penetration in Spotted Hyenas, see Glickman (1995). See also Morris (1956:261), who defines courtship as “the heterosexual reproductive communication system leading up to the consummatory sexual act”