Biological Exuberance: Animal Homosexuality and Natural Diversity

Male, pp. 638—41 (Philadelphia: W. B. Saunders).

21

Calculations are based on data in the following sources: Bonobo (Idani 1991:90-91 [tables 5—6]); Red Deer (Hall 1983:278 [table 2]); Bonnet Macaque (Sugiyama 1971:259-60 [tables 8-9]); Pig-tailed Macaque (Tokuda et al. 1968:288, 290 [tables 3,5]); Kob (Buechner and Schloeth 1965:219 [table 2]).

22

R. Wrangham, quoted in Bull, C. (1997) “Monkey Love,” The Advocate, June 10, 735:58. This is but one example of the often misleading statements about animal homosexuality that are perpetuated by both scientists and the popular press. See chapter 3 for further discussion.

23

Females had an average of 5.2 different female partners and 4.1 male partners and ranged 4-9 female partners (out of a pool of 10) and 1—9 male partners (out of a pool of 10) (Idani 1991:91). Of course, all of these figures represent only a relatively short “snapshot” of Bonobo behavior (covering three months), but it is likely that longer-term or lifetime patterns exhibit a comparable spectrum of variation. Because female Bonobos are neither exclusively heterosexual nor exclusively homosexual, de Waal (1997:192) advocates use of the term pansexual to descibe their sexual orientation. This characterization is as appropriate as bisexual as long as it is understood that individuals exhibit a range of same-sex versus opposite-sex interactions (i.e., there are many gradations of “pansexuality” or “bisexuality” in this species).

24

Of course, many other factors besides sexual “preference” are involved in the choice of mates, especially with regard to the availability and specific characteristics of partners. The fact that only some animals ever engage in homosexual or heterosexual activity, however, is an important indication that differences in sexual orientation probably also exist at an individual level. For further discussion of the role that partner availability may play in the occurrence of homosexual (and heterosexual) activity, see chapter 4.

25

Silver Gull: data for 131 females tracked in the wild over their entire lives, from Mills 1991:1525 (table 1); Black-headed Gull: data for 27 males in a captive population studied for seven years, based on van Rhijn and Groothuis 1985:161 (table 3); Japanese Macaque: averages for 46-58 females over two consecutive years in a semiwild population, from Wolfe 1979:526; Galah: based on pair-bonding data over six years pooled from two captive populations comprising 27 birds, from Rogers and McCulloch 1981.

26

Vasey, “Homosexual Behavior in Primates,” p 197.

27

For explicit observations of the nonchalant responses of surrounding animals, including the heterosexual mates or parents of individuals involved in same-sex activity, see Common Chimpanzee (Kortlandt 1962:132); Gorilla (Harcourt et al. 1981:276); White-handed Gibbon (Edwards and Todd 1991:232—33); Japanese Macaque (Wolfe 1984; Vasey 1995:190); Killer Whale (Jacobsen 1986:152); Gray Whale (Darling 1978:55); Northern Fur Seal (Bartholomew 1959:168); African Buffalo (Mloszewski 1983:186); Lion (Chavan 1981:364); Rufous Rat Kangaroo (Johnson 1980:356); Dwarf Cavy (Rood 1970:442); Laughing Gull (Noble and Wurm 1943:205); Sage Grouse (Scott 1942:495). In the majority of cases where homosexual activities draw no response from surrounding animals, scientific observers simply make no comment about the behavior of the other animals. In one species, the Blue- bellied Roller, same-sex (and opposite-sex) “display” mounting is only performed when other birds are present to watch (but not intervene).

28

African Buffalo (Mloszewski 1983:186); Musk-ox (Tener 1965:75).

29

Other species in which harassment of heterosexual but not homosexual activity has been reported include Proboscis Monkeys (Yeager 1990a:224), Squirrel Monkeys (DuMond 1968:121-22; Baldwin and Baldwin 1981:304), Lechwe (Nefdt 1995), Wolves (Zimen 1976, 1981; Derix et al.1993), Red-necked Wallabies (Johnson 1989:275), Gray Squirrels (Koprowski 1992a:393; 1993:167—68), Kittiwakes (Chardine 1986), and King Penguins (Stonehouse 1960:32). In Hanuman Langurs, more than 83 percent of heterosexual copulations are harassed while harassment of homosexual ones only occurs occasionally (Sommer 1989a:208; Srivastava et al. 1991:497); for greater interruption of heterosexual mounts in Japanese Macaques, see Hanby 1974:840; in Moor Macaques, see Matsumura and Okamoto 1998:227-28. See also chapter 5 for further discussion of harassment of heterosexual matings.

30

Bonobo (de Waal 1995:48, 1997:117, 120; Hashimoto 1997:12); Jackdaw (Roell 1978:29); Guianan Cock-of- the-Rock (Trail and Koutnik 1986:210—11). In a number of species (e.g., Rhesus and Crab-eating Macaques, Spotted Hyenas) a phenomeon known as scapegoating sometimes occurs, in which several individuals combine forces to attack another individual not directly involved in a dispute. Notably, individuals engaging in same-sex activity are not specifically targeted as scapegoats, and this behavior is not in fact generally related to sexual activity at all (Harcourt, A. H., and F. B. M. de Waal, eds., (1992) Coalitions and Alliances in Humans and Other Animals, pp. 87, 91, 129, 240 [Oxford: Oxford University Press]).

31

Savanna Baboon (Marais 1922/1969); Red Deer (Darling 1937); Common Garter Snake (Mason and Crews 1985).

32

Greylag Goose (Lorenz 1979, 1991; Huber and Martys 1993; Schonfeld 1985); Black Swan (Braithwaite 1981).

33

For example, Japanese Macaques, Savanna Baboons, Kob, Mute Swans, Black-winged Stilts, Caspian Terns, Black-billed Magpies.

34

Greylag Goose (Huber and Martys 1993); Black Swan (Braithwaite 1981); Flamingo (King 1994, 1993a,b; E. Stevens, personal communication); Orange-fronted Parakeet (Hardy 1963:187, 1965:150); Laughing Gull (Noble and Wurm 1943:205; Hand 1981:138-39); Rose-ringed Parakeet (Goodwin 1983:87); Nilgiri Langur (Hohmann 1989:452); Lion (Cooper 1942:27-28); Rhesus Macaque (Fairbanks et al. 1977:247); Japanese Macaque (Vasey 1998); Common Chimpanzee (de Waal 1982:64-66); Livingstone’s Fruit Bat (Courts 1996:27); Savanna Baboon (Marais 1922/1969:205-6). Homosexual pairs in the domesticated Bengalese Finch also attack other birds (Masatomi 1959). Additionally, in a number of mammals (e.g., Common Chimpanzees, Bonnet Macaques, Savanna Baboons, Bottlenose and Atlantic Spotted Dolphins, Cheetahs), paired “coalitions” or “alliances” of males that “solidify” their partnership through overt or ritualized sexual, affectionate, and bonding behaviors often cooperate in challenging and attacking other individuals; a similar phenomenon ocurs in homosexually bonded female Oystercatchers that are part of bisexual trios.

35

Brown Capuchin (Linn et al. 1995:50); Rufous Rat Kangaroo (Ganslosser and Fuchs 1988:311); Sage Grouse (Patterson 1952:155-56); Gorilla (Harcourt et al. 1981:276; Fisher and Nadler 1978:660-61); Bonobo (de Waal 1997:114, 130); Canada Goose (Allen 1934:197—98); Wapiti (Franklin and Lieb 1979:188-89); Japanese Macaque (Vasey 1998); Rhesus Macaque (Akers and Conaway 1979:76); Jackdaw (Roell 1979:124-25). In Greenshanks, the female partner of a male involved in homosexual copulation made a threatening call during a same-sex interaction but did not interfere (Nethersole-Thompson 1951:109—10).

36

White-tailed Deer (Thomas et al. 1965). Another possible example of transgendered animals being harassed is found in the paketi (a New Zealand fish), in which Ayling (1982:255) claims that transvestite fish are attacked (“beat up”) when their true sex is discovered; however, Jones (1980), the original source on which this account is based, does not in fact mention such behavior (Ayling, T. [1982] Sea Fishes of New Zealand [Auckland: Collins]; Jones, G. P. [1980] “Growth and Reproduction in the Protogynous Hermaphrodite Pseudolabrus celidotus [Pisces: Labridae] in New Zealand,” Copeia 1980:660—75).