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Notes to Part I

When no specific references are noted for a particular species in part 1, the information and sources will be found in the profile itself in part 2. When a note is included for a profiled species (e.g., to provide more detailed information), the citation format includes the species name, author, year, and (in most cases) page numbers of the source, referring to the full references in the profile. References for species that are not profiled in part 2 are included directly in notes.

Introduction

1. Einstein, A. (1930) “What I Believe,” Forum and Century 84(4):193-94.

Chapter 1. The Birds and the Bees

Haldane, J. B. S. (1928) Possible Worlds and Other Papers, p. 298 (New York: Harper & Brothers).

Animal names that are capitalized refer to a species or group of closely related species that is profiled in part 2, or whose references are included in the appendix.

Homosexuality among primates, for example, has been traced back to at least the Oligocene epoch, 24-37 million years ago (based on its distribution among contemporary primates; Vasey 1995:195). Some scientists place its original appearance even earlier in the evolutionary line leading to mammals, at around 200 million years ago (Baker and Bellis 1995:5), and it has probably existed for much longer among other animal groups. Vasey, P. L. (1995) “Homosexual Behavior in Primates: A Review of Evidence and Theory,” International Journal of Primatology 16:173-204; Baker, R., and M. A. Bellis (1995) Human Sperm Competition: Copulation, Masturbation, and Infidelity (London: Chapman and Hall).

See note 29, as well as part 2 and the appendix, for more detailed tabulations (including discussion of species not included in this tally).

For further discussion of sexual orientation in animals, as well as comparisons between animal and human homosexuality, see chapter 2. Following Vasey (“Homosexual Behavior in Primates,” p. 175), the term homosexual is used to designate primarily the form of behaviors without necessarily implying anything about their “function or context or the actors’ ages and motivation.” For further consideration of the terminology used to describe same-sex activity in animals, including discussion of alternative definitions of the term homosexual(ity) as it is applied to animals (and some of the controversies that have surrounded its use in the zoological literature), see chapter 3. For more on the “functions” and contexts of homosexual behavior, see chapters 4-5.

Guianan Cock-of-the-Rock (Endler and Théry 1996); Anna’s Hummingbird (Hamilton 1965); Buff-breasted Sandpiper (Myers 1989).

For a general survey of play-fighting, see Aldis, O. (1975) Play Fighting. (New York: Academic Press).

Spinner Dolphin (Norris et al. 1994:250).

Ingestion of semen by both males and females during masturbation in heterosexual contexts also occurs among Golden Monkeys (Clarke 1991:371).

Supernormal clutches have also been reported for pairs of male Emus, probably because more than one female has laid in their nest. What might be termed “subnormal” clutches—i.e., nests containing fewer eggs than are usually found for heterosexual pairs—are reported for female pairs of Blue Tits. And “super-supernormal” clutches occasionally occur in heterosexual pairs of Roseate Terns: as a result of within-species parasitism and possibly also egg transfer (see chapter 5 for more on these phenomena), some nests contain more than double the number of eggs found even in supernormal clutches (as is also true for “dump” nests in many Ducks and Geese).

For discussion, and refutation, of the idea that same-sex pairs form in species such as these solely for the purpose of raising offspring, see chapter 5. In some birds such as grouse (e.g., sharp-tailed grouse, prairie chickens, white-tailed ptarmigan) and ducks (e.g., eiders, buffleheads) broods from more than one female are combined or “amalgamated” but no same-sex coparenting occurs (one female, or a heterosexual pair, look after all the offspring); cf. Bergerud and Gratson 1988:545 (Grouse); Afton 1993 (Ducks); Eadie, J. McA., E P. Kehoe, and T. D. Nudds (1988) “Pre-Hatch and Post-Hatch Brood Amalgamation in North American Anatidae: A Review of Hypotheses,” Canadian Journal of Zoology 66:1709-21.

Ring-billed Gull (Conover 1989:148).

In some bird species in which same-sex pairs are unable to obtain fertile eggs on their own (or in which homosexual parenting has yet to be observed in the wild), parenting skills have been demonstrated by supplying homosexual pairs with “foster” eggs or young in captivity. Same-sex pairs of Flamingos, White Storks, Black-headed Gulls, Steller’s Sea Eagles, Barn Owls, and Gentoo Penguins, for example, have all successfully hatched such eggs and/or raised foster chicks.

Black Swan (Braithwaite 1981:140—42); for more details, see chapter 5 and part 2.

Ring-billed Gull (Conover 1989:148); Western Gull (Hayward and Fry 1993:17—18); see chapter 2 for further discussion of same-sex pairs being limited to nonoptimal territories. Several other studies point to the possibility of more “attentive” parenting by female homosexual pairs. Researchers have found that female Ring-billed Gulls in same-sex pairs, for example, may have higher levels of progesterone—a female hormone associated with nest-building and incubation behavior—than females in heterosexual pairs (Kovacs and Ryder 1985); see chapter 4 for more on the hormonal profiles of animals involved in same-sex activity. In a related set of observations, some investigators have documented more “intense” nesting behavior in female homosexual pairs than heterosexual pairs in some captive studies. Allen and Erickson (1982:346, 350), for instance, found that female pairs of Ring Doves are more persistent incubators than heterosexual pairs, being less likely to abandon their nests and terminate incubation when they have infertile eggs than are heterosexual pairs. Brockway (1967:76) found that female Budgerigars in homosexual pairs begin continuous occupation of their nests significantly sooner than females in heterosexual pairs. However, because female pairs begin noncontinuous occupation of their nests significantly later than heterosexual pairs in this species, the overall amount of their nesting activity and the timing of their egg-laying essentially evens out.

See chapter 5 for further discussion of homosexual activity in communal groups and the often complex relationship between “helpers” and same-sex activity.

In many species, young may also be raised in heterosexual trios, i.e., family units with three parents in which only opposite-sex bonding is present between the adults. See chapter 5 for some examples.

For discussion of single parenting in animals where two (heterosexual) parents usually raise the young, as well as examples of other heterosexual parenting arrangements that deviate from the species-typical pattern, see chapter 5.

For additional discussion of male bias in biological studies, see chapters 3 and 5.

Rhesus Macaque (Altmann 1962:383; Lindburg 1971:69); Hamadryas Baboon (Abegglen 1984:63); Gelada Baboon (Bernstein 1970:94); Tasmanian Native Hen (Ridpath 1972:30); Gray-headed Flying Fox (Nelson 1965:546).

Pukeko (Jamieson and Craig 1987a:1251 ); Bonobo (Thompson-Handler et al. 1984:349; Kano 1992:187; Kitamura 1989:55—57); Stumptail Macaque (Chevalier-Skolnikoff 1974:101, 110); Red Deer (Hall 1983:278); Red-necked Wallaby (LaFollette 1971:96); Northern Quoll (Dempster 1995:29).

Pig-tailed Macaque (Oi 1990a:350—51): Galah (Rogers and McCulloch 1981); Pronghorn (Kitchen 1974:44).

Gorilla (Fischer and Nadler 1978:660—61; Yamagiwa 1987a:12, 1987b:37).

Pukeko (Jamieson and Craig 1987a:1251—52); Flamingo (C. E. King, personal communication). In Lesser Flamingos, however, the reverse appears to be true: males but not females achieve cloacal contact during their homosexual mounts (Alraun and Hewston 1997:176).

Japanese Macaque (Hanby 1974:838-40; Wolfe 1984:149; Fedigan 1982:143).

For further discussion of comparisons between animal and human homosexuality, see chapter 2.

These formulas are also used to estimate the number of bisexual/heterosexual trios in a population; see Conover and Aylor 1985:127 (Ring-billed Gull).

Kob (Buechner and Schloeth 1965); Long-tailed Hermit (Stiles and Wolf 1979). Likewise, up to 30 homosexual pairs of Herring Gulls have been counted in some locations—a relatively high number of same-sex associations to be present at one time—but in colonies that number more than 10,000 pairs, this amounts to less than I percent of the total number of pairs (Shugart 1980:426—27).

Same-sex courtship, sexual, pair-bonding, and /or parenting behaviors have been documented in the scientific literature in at least 167 species of mammals, 132 birds, 32 reptiles and amphibians 15 fishes, and 125 insects and other invertebrates, for a total of 471 species (see part 2 and appendix for a complete list). These figures do not include domesticated animals (at least another 19 species; see the appendix), nor species in which only sexually immature animals/juveniles engage in homosexual activities (for a survey of the latter in mammals, see Dagg 1984). For a number of reasons, this tally is likely to be an underestimate (especially for species other than mammals and birds, which are not as thoroughly covered): see chapter 3 for further discussion. It should also be pointed out that species totals may differ depending on the classificatory schema used; in some taxonomies, for example, animals that in this book are lumped together as subspecies are considered separate species (e.g., the various subspecies of Savanna Baboons, Flamingos, or Wapiti/Red Deer). This roster also excludes a wide variety of other cases in which the evidence for homosexual activities is not definitive, such as:

1. species in which homosexual activity is suspected (and sometimes included in comprehensive surveys of homosexual behavior, such as Dagg 1984) but in which the sex of participants has not yet been coin-firmed (e.g., one-horned rhinoceros [Laurie 1982:323], yellow-bellied marmot [Armitage 1962:327], South African cliff swallow [Earlé 1985:46], band-tailed barbfhroat hummingbird [Harms and Ahumada 1992], calliope hummingbird [Armstrong 1988], ringed Parakeet [Hardy 1964]).

2. bird species in which supernormal clutches have been documented without any direct evidence of same-sex pairs (e.g., numerous gulls and other bird species; see note 70, chapter 4).

3. same-sex trios or joint parenting arrangements with little or no conclusive evidence of courtship, sex, or pair-bonding between the like-sexed coparenis (e.g., bobolink [Bollinger et al. 1986], various ducks. grouse [cf. note 11, this chapter] ).

4. bird species in which males associate in “pairs” or form “partnerships” with other males for joint displays during heterosexual courtships, but in which no overt courtship or sexual behavior occers between such partners or other same-sex individuals (e.g., several manakins of the genera Chiroxiphia, Pipra, Machaeropterus, and Masius—note however that males in these species often court “female-plumaged. birds, the sex of most of which has not been determined, while in two other species, some of these individuals have been determined to be males; wild turkey; king bird of paradise and possibly other birds of paradise. For further references, see McDonald 1989: (007 and Trainer and McDonald 1993:779).

5. species in which the only form of documented “same-sex” activity involves individuals mounting het-crosexual copulating pairs, such that the mounting activity is not necessarily limited to like-sexed individuals or the same-sex motivation/orientation is not clear (e.g., camel and Dagg 1981:92], Buller’s albatross [Warham 1967:129]).

6. species in which the only same-sex activity is mounting that appears to be exclusively aggressive in character with no sexual component (e.g., collared lemming, degu, ground squirrel; of. Dagg 1984 and sources cited therein; see also chapter 3 for further discussion of aggressive or “dominance” mounting and the difficulty of distinguishing this from sexual mounting); and species in which the only same-sex activities are “affectionate” behaviors or “platonic” companionships unaccompanied by either signs of sexual arousal or overt courtship or sexual behaviors.

7. other inconclusive cases, such as species reported in secondary sources as exhibiting homosexuality but whose original sources do not definitively document same-sex activity (e.g., avocets, reported in Terres [1980:813], with no mention of source, as engaging in homosexual mounting; Makkink [1936] and Hamilton [1975]—the most comprehensive primary field studies of this species and the most likely sources for this information—describe ritual mountings and masturbation “eruptive copulations”] but no homosexual mounting).

Armitage, K. B. (1962) “Social Behavior of a Colony of the Yellow-bellied Marmut (Marmota flaviventris).” Animal Behavior 10:319-31; Armstrong, D. P. (1988) “Persistent Attempts by a Male Calliope Hummingbird, Stellula calliope, to Copulate with Newly Fledged Conspecifics,” Canadian Field-Naturalist 102:259—60; Bollinger, E. K., T. A. Gavin, C. J. Hibbard, and J. T. Wootton (1986) “Two Male Bobolinks Feed Young at the Same Nest,” Wilson Bulletin 98:154—56; Dagg, A. I. (1984) “Homosexual Behavior and Female-Wale Mounting in Mammals—a First Survey,” Mammal Review 14:155—85; Earlé, R. A. (1985) “A Description of the Social, Aggressive, and Maintenance Behavior of the South African Cliff Swallow Hirundo spilodera (Aves: Hirundinidae),” Navorsinge van die nasionale Museum, Bloemfontein 5:37—50; Gauthier-Pilters, H., and A. I. Dagg (1981) The Camel: Its Evolution, Ecology, Behavior, and Relationship to Man (Chicago: University of Chicago Press); Hardy, J.W. (1964) “Ringed Parakeets Nesting in Los Angeles, California,” Condor 65:445—47; Harms, K. E., and J.A. Ahumada (1992) “Observations of an Adult Hummingbird Provisioning an Incubating Adult,” Wilson Bulletin 104:369-70; Laurie, A. ( 1982) “Behavioral Ecology of the Greater One-horned Rhinoceros (Rhinoceros unicornis),” Journal of Zoology, London 196:307—41; Makkink, G. F. (1936) “An Attempt at an Ethogram of the European Avocet (Recurvirostra avosetta L.), With Ethological and Psychological Remarks,” Ardea 25:1-63; McDonald, D. B. (1989) “Correlates of Male Mating Success in a Lekking Bird with Male-Male Cooperation,” Animal Behavior 37:1007—22; Terres, J. K. (1980) The Audubon Society Encyclopedia of North American Birds (New York: Alfred A. Knopf); Trainer, J. M., and D. B. McDonald (1993) “Vocal Repertoire of the Long-tailed Manakin and Its Relation to Male-Male Cooperation,” Condor 95:769—81; Warham, J. (1967) “Snares Island Birds,” Notornis 14:122—39.

According to Wilson (1992), approximately 1,032,000 species of animals are currently known to science, although the number of actually occurring species is undoubtedly much higher—on the order of 10-100 million—and there are many complexities in estimating the total number of species. For further discussion, see Wilson, E. O. (1992) The Diversity of Life, pp. 131ff (Cambridge, Mass.: Belknap Press); Wilson, E. O. (1988) “The Current State of Biological Diversity,” in E. O. Wilson, ed., BioDiversity, pp. 3-18. (Washington, D.C.: National Academy Press); May, R. M. (1988) “How Many Species Are There on Earth?” Science 241:1441—49.

Le Boeuf and Mesnick 1991:155 (Elephant Seal); see also Wilson, E. O. (1975) Sociobiology: The New Synthesis (Cambridge and London: Belknap Press). This figure is borne out by the data on homosexuality: the average number of observation hours for scientific studies in which homosexual behavior has been observed is approximately 1,050 hours (based on data from 47 species in which the number of observation hours has been recorded).

Marten, M., J. May, and R. Taylor (1982) Weird and Wonderful Wildlife, p. 7. (San Francisco: Chronicle Books). A somewhat more precise estimate of the number of species that have been adequately studied can be obtained for a subset of animals by using the Zoological Record (a comprehensive electronic database that indexes more than a million zoological source documents, including articles from over 6,000 journals worldwide, over the past 20 years). The Zoological Record for the period 1978-97 lists 825 mammal species in which at least some aspects of courtship, sexual, pair-bonding, mating-system, and/or parenting behaviors have been studied (the behavior categories in which homosexuality, if present, is likely to be found). Homosexual behavior has been documented in 133 of these species, or approximately 16 percent—comparable to the lower range obtained using the estimate of Marten et al. (The following subject headings/behavior categories indexed by Zoological Record were used in compiling this estimate: Courtship, Lek, Sexual Display, Precopulatory Behavior, Copulation, Mating, Pair Formation, Monogamy, Polygamy, Cooperative Breeding, Breeding Habits, Parental Care, Care of Young, Homosexuality).

See chapter 4 for further discussion of these factors.

For species that do not engage in “heterosexual” mating at all, e.g., parthenogenetic or hermaphrodite animals, see the next section.

Clapham, P. J. (1996) “The Social and Reproductive Biology of Humpback Whales: An Ecological Perspective,” p. 37, Mammal Review 26:27-49.

Scott, P. E. (1994) “Lucifer Hummingbird (Calothorax lucifer),” in A. Poole and F. Gill, eds., The Birds of North America: Life Histories for the 21st Century, no. 134, p. 9. (Philadelphia: Academy of Natural Sciences; Washington, D.C.: American Ornithologists’ Union); Dejong, M. J. (1996) “Northern Rough-winged Swallow (Stelgidopteryx serripennis),” in Poole and Gill, The Birds of North America, no. 234, p. 9; Kricher, J. C. (1995) “Black-and-white Warbler (Pheucticus melanocephalus),” in Poole and Gill, The Birds of North America, no. 158, p. 9; O‘Brien, R. M. (1990) “Red-tailed Tropicbird (Phaethon rubricauda)”, in S. Marchant and P. J. Higgins, eds., Handbook of Australian, New Zealand, and Antarctic Birds, vol. 1, part B, p. 940 (Melbourne: Oxford University Press); Johnsgard, P. A. (1983) Cranes of the World (Bloomington: Indiana University Press); Powers, D. R. (1996) “Magnificent Hummingbird (Eugenes fulgens),” in Poole and Gill, The Birds of North America, no. 221, p. 10; Hill, G. E. (1994) “Black-headed Grosbeak (Pheucticus melanocephalus),” in Poole and Gill, The Birds of North America, no. 143, p. 8; Victoria’s Riflebird (Frith and Cooper 1996:103; Gilliard 1969:13); Cheetah (Caro 1994:42); Lepson, J. K., and L. A. Freed (1995) “Variation in Male Plumage and Behavior of the Hawaii Akepa,” Auk 112:402-14; Spotted Hyena (Frank 1996:117); Agile Wallaby (Stirrat and Fuller 1997:75); Birds of Paradise (Davis and Beehler 1994:522); Nelson, S. K., and S. G. Sealy (1995) “Biology of the Marbled Murrelet: Inland and at Sea (Symposium Introduction),” Northwestern Naturalist 76:1—3; Orang-utan (Schürmann 1982; Schürmann and van Hoof 1986; Maple 1980); Rowe, S., and R. Empson (1996) “Observations on the Breeding Behavior of the Tanga’eo or Mangaia Kingfisher (Halcyon tuta rufi-collaris),” Notornis 43:43-48; Common Chimpanzee (Gagneux et al. 1997; Wrangham 1997); Harbor Seal (Perry and Amos 1998).

Emu (Coddington and Cockburn 1995; Heinroth 1924); Black-rumped Flameback (Neelakantan 1962); Nilgiri Langur (Poirier 1970a,b); Harbor Seal (Johnson 1976:45); Northern Quoll (Dempster 1995); Gray-capped Social Weaver (Collias and Collias 1980); Walrus (Miller 1975; Fay et al. 1984); Acorn Woodpecker (Koenig and Stacey 1990:427); Australian Shelduck (Riggert 1977:20); Killer Whale (Jacobsen 1990:78; Rose 1992:1—2). See also Lutz and Voight 1994 for the first documentation of copulatory behavior—between two males—in two previously unknown species of deep-sea octopuses (a group in which heterosexual mating has yet to be observed in any species). Other animals in which same-sex activity has been documented and in which heterosexual activity has rarely been observed include Musk-oxen (Smith 1976:62), Red-necked Wallabies (Johnson 1989a:275), Vicunas (Koford 1957:182-84), Musk Ducks (Lowe 1966:285), and Ruffed Grouse (Johnsgard 1983:295). See chapter 4 for further discussion of the often insurmountable difficulties in attempting to observe and study sexual activity under field conditions.

Ring-billed Gull (Conover and Aylor 1985). See chapter 4 for discussion of some of the pitfalls of equating homosexual pairings with supernormal clutches.

Dragonflies (Dunkle 1991).

Pukeko (Craig 1980); Pronghorn (Kitchen 1974). The behavior was probably classified as rare in Pronghorn because the amount of same-sex activity was not being compared to the amount of opposite-sex activity, but rather to the total amount of “dominance” behavior (which it was classified as), consisting primarily of nonsexual activities. For further discussion and critique of homosexuality interpreted as a “dominance” activity, see chapter 3.

Killer Whale (Rose 1992:116); Regent Bowerbird (Lenz 1994:266 [table 2]); White-handed Gibbon (Edwards and Todd 1991:233 [table 1]); Crab-eating Macaque (Thompson 1969:465).

Giraffe (Pratt and Anderson 1985, 1982, 1979). In a study of another population of Giraffes, only three mounts between males were recorded, but only 400 hours of observation were involved (Dagg and Foster 1976:124).

Mountain Sheep (Geist 1968:210-11 [tables 4, 6]); 1971:152 [table 30]).

Gray Heron (Ramo 1993:116-17). In some species (e.g., Little Egrets, Little Blue Herons) quantitative information is only available for the proportion of promiscuous copulations that are homosexual (the higher figure). Where both proportions are available (e.g., Cattle Egrets, Gray Herons), an average of the two is taken when calculating cross-species comparisons of frequency (see below). Different frequencies can also be obtained depending on whether a distinction between copulatory and noncopulatory (or “ritualized”) mounting is taken into account. (See chapter 3 for discussion of the—sometimes arbitrary—distinction between these two types of mounting.) In some cases, such as Pig-tailed and Crested Black Macaques, a sizable difference obtains. In Pigtails, 82 percent of mounting is same-sex if only noncopulatory mounts are considered (Oi 1990a:350-51 [table 4]), whereas 7-23 percent of all mounts are same-sex if “full” heterosexual copulations are included in the calculation (Bernstein 1967:226-7; Oi 1996:345). In Crested Blacks, roughly a third of noncopulatory mounts are between males, but overall this constitutes only about 8 percent of mounting activity when combined with copulatory (heterosexual) mounts (C. Reed, personal communication). In cases such as these, the latter (smaller) percentage is taken to be the overall rate of same-sex activity. For other species, however, the two rates are comparable. In Common Chimpanzees, for example, Nishida and Hosaka (1996:122, 129 [tables 9.7, 9.17a]) recorded 61 ritualized mounts between males compared to 123 male-female copulations (nonritualized), yielding a rate of 33 percent same-sex mountings. This is comparable to the exclusively noncopulatory figures of Bygott (1974; cited in Hanby 1974:845 [Japanese Macaque]), who found that 4 of 14 ritualized mounts (29 percent) occurred between males.

Of course, these cross-species comparisons refer only to those animals in which homosexual behavior has been observed and in which the appropriate quantitative information is available. In many species same-sex behavior is more or less common than the maximum and minimum figures obtained from these measures, but it has not been quantified and therefore cannot be compared to these examples. For these calculations, if multiple frequency proportions were available for the same species—either because of population, seasonal, or behavioral differences (as discussed above)—these were averaged prior to being combined with the figures for other species. Proportions for courtship behavior are based on quantitative information from 21 species (avg = 23 percent same-sex activity), for sexual behavior from 77 species (avg = 26 percent), for pairing behavior from 45 species (avg = 14 percent), and for population percentages from 56 species (avg = 27 percent). For the purposes of comparison, tallies of observed homosexual and heterosexual behaviors in each species are assumed to be representative of actually occurring frequencies. The statistics for pairing and sexual behavior do not include the many species in which the only form of pair-bonding, coparenting, or observed sexual behavior is between same-sex individuals, i.e., in which 100 percent of pairs, coparents, or observed sexual interactions are homosexual (were these to be included, the proportions would be considerably, and perhaps unrepresentatively, higher). In five of these cases, however—Northern Elephant Seals, Cheetahs, Grizzly Bears, Lesser Scaup Ducks, and Greater Rheas—the proportion of all families or nests that are tended by same-sex pairs or trios (as opposed to single individuals) is substituted for the proportion of same-sex pairs. For population calculations, figures represent only the sex or activity that has been quantified (e.g., for species in which only females form same-sex pairs, or in which only female pairs have been tallied, only the proportion of females in such pairs is included). Moreover, for many bird species-especially those in which only a small fraction of individuals participate in same-sex activity—population percentages are not available. However, in species with same-sex pairing, the proportion of homosexual pairs is roughly comparable to the proportion of individuals engaging in same-sex activity (the two differ, of course, if there are sizable numbers of nonbreeding birds that do not form pairs with either sex). In order not to bias the sample toward species with relatively high population percentages, therefore, pairing proportions for wild-bird populations have been substituted. In species where no other population data are available, these figures are taken as a rough estimate of the proportion of individuals involved in same-sex activity.

The term transgender, when it is applied to people, has two uses: as a cover term that refers to a wide variety of gender-crossing or gender-mixing phenomena, including transsexuality (with various degrees of hormonal and/or surgical transformation), transvestism (including cross-dressers, drag queens and kings, and female and male impersonators), intersexuality (including various forms of hermaphroditism), and even extremes of butch-femme presentation. It is also used as a designation for a specific form of gender mixing in which an individual lives full-time in the gender opposite to his or her anatomy (e.g., a man who passes for a woman without undergoing the full physical transitioning of a transsexual). For more discussion and exemplification see Feinberg, L. (1996) Transgender Warriors: Making History from Joan of Arc to RuPaul (Boston: Beacon Press).

Foltz, D. W, H. Ochman, J. S. Jones, S. M. Evangelisti, and R. K. Selander (1982) “Genetic Population Structure and Breeding Systems in Arionid Slugs (Mollusca: Pulmonata),” Biological Journal of the Linnean Society 17:225-41.

Of course, the term transvestism, when applied to people, refers primarily the wearing clothing of the opposite sex (and all of the attendant social and cultural repercussions). In its zoological usage, however, it simply refers to physical or behavioral attributes that are typical of the opposite sex in that species. For scientific use of this term, see the references in the notes for this section, as well as Weinrich, J. D. (1980) “Homosexual Behavior in Animals: (A New Review of Observations from the Wild, and Their Relationship to Human Sexuality,” in R. Forleo and W. Pasini, eds., Medical Sexology: The Third International Congress, pp. 288-95. (Littleton, Mass.: PSG Publishing).

Owen, D. F. (1988) “Mimicry and Transvestism in Papilio phorcas (Lepidoptera: Papilionidae),” Journal of the Entomological Society of Southern Africa 51:294-96; Weldon, P. J., and G. M. Burghardt (1984) “Deception Divergence and Sexual Selection,” Zeitschrift für Tierpsychologie 65:89-102.

Rohwer, S., S. D. Fretwell, and D. M. Niles (1980) “Delayed Maturation in Passerine Plumages and the Deceptive Acquistion of Resources,” American Naturalist 115:400-437.

Estes, R. D. (1991) “The Significance of Horns and Other Male Secondary Sexual Characters in Female Bovids,” Applied Animal Behavior Science 29:403-51; Guthrie, R. D., and R. G. Petocz (1970) “Weapon Au-tomimicry Among Mammals,” American Naturalist 104:585-88.

Kirwan, G. M. (1996) “Rostratulidae (Painted-Snipes),” p. 297, in J. del Hoyo, A. Elliott, and J. Sargatal, eds., Handbook of the Birds of the World, vol. 3, Hoatzin to Auks, pp. 292-301 (Barcelona: Lynx Edicións). Examples such as these are often termed sex-role reversal by biologists.

See the discussion of homosexual gender roles and interpretations of homosexuality as “pseudoheterosexuality” in chapter 4.

Bighorn Sheep (Berger 1985). See also chapter 2 for further discussion of human and animal comparisons.

Policansky, D. (1982) “Sex Change in Plants and Animals,” Annual Review of Ecology and Systematics 13:471—95; Forsyth, A. (1986) A Natural History of Sex: The Ecology and Evolution of Mating Behavior, chapter 13. (New York: Scribner’s).

For surveys of transsexuality in fishes, see Potts, G. W., and R. J. Wootton, eds., (1984) Fish Reproduction: Strategies and Tactics (London: Academic Press); Warner, R. R. (1978) “The Evolution of Hermaphroditism and Unisexuality in Aquatic and Terrestrial Vertebrates,” in E. S. Reese and F. J. Lighter, eds., Contrasts in Behavior: Adaptations in the Aquatic and Terrestrial Environments, pp. 77-101 (New York: Wiley); Warner, R. R. (1975) “The Adaptive Significance of Sequential Hermaphroditism in Animals,” American Naturalist 109:61-82; Warner, R. R. (1984) “Mating Behavior and Hermaphroditism in Coral Reef Fishes,” American Scientist 72:128—36; Policansky, “Sex Change”; Armstrong, C. N. (1964) Intersexuality in Vertebrates Including Man (London: Academic); Smith, C. L. (1975) “The Evolution of Hermaphroditism in Fishes,” in R. Reinboth, ed., Intersexuality in the Animal Kingdom, pp. 295-310 (New York: Springer-Verlag); Smith, C. L. (1967) “Contribution to a Theory of Hermaphroditism,” Journal of Theoretical Biology 17:76-90.

Robertson, D. R., and R. R. Warner (1978) “Sexual Patterns in the Labroid Fishes of the Western Caribbean, II: The Parrotfishes (Scaridae),” Smithsonian Contributions to Zoology 255:1—26; Warner, R.R., and I. F. Downs (1977) “Comparative Life Histories: Growth versus Reproduction in Normal Males and Sex-changing Hermaphrodites in the Striped Parrotfish, Scarus croicensis,” Proceedings of the Third International Symposium on Coral Reefs 1(Biology):275-82; Thresher, R. E. (1984) Reproduction in Reef Fishes (Neptune City, N.J.: T.F.H. Publications).

Paketi: Jones, G. P. (1980) “Growth and Reproduction in the Protogynous Hermaphrodite Pseudolabrus celidotus (Pisces: Labridae) in New Zealand,” Copeia 1980:660-75; Ayling, T. (1982) Sea Fishes of New Zealand, p. 255 (Auckland: Collins). Humbug damselfish: Coates, D. (1982) “Some Observations on the Sexuality of Humbug Damselfish, Dascyllus aruanus (Pisces, Pomacentridae) in the Field,” Zeitschrift für Tierpsychologie 59:7-18. Red Sea anemonefish: Fricke, H. W. (1979) “Mating System, Resource Defence, and Sex Change in the Anemonefish Amphiprion akallopisos,” Zeitschrift für Tierpsychologie 50:313-26. Lantern bass and others: Petersen, C. W., and E. A. Fischer (1986) “Mating System of the Hermaphroditic Coral-reef Fish, Serranus baldwini,” Behavioral Ecology and Sociobiology 19:171—78; Nakashima, Y, K. Karino, T. Kuwamura, and Y. Sakai (1997) “A Protogynous Wrasse May Have a Functionally Simultaneous Hermaphrodite Phase,” in M. Taborsky and B. Taborsky, eds., Contributions to the XXV International Ethological Conference, p. 214, Advances in Ethology no. 32 (Berlin: Blackwell Wissenschafrs-Verlag). Coral goby: Kuwamura, T., Y. Nakshima, and Y. Yogo (1994) “Sex Change in Either Direction by Growth-Rate Advantage in the Monogamous Coral Goby, Paragobiodon echinocephalus,” Behavioral Ecology 5:434-38; Nakashima, Y., T. Kuwamura, and Y. Yogo (1995) “Why Be a Both-ways Sex Changer?” Ethology 101:301-7.

Chapter 2. Humanistic Animals, Animalistic Humans

Names for individual animals in each species, and the activities they engaged in, are from the following sources: Gorillas (Yamagiwa 1987a, Stewart 1977); Bottlenose Dolphins (Tavolga 1966); West Indian Manatees (Hartman 1971); Siamangs (Fox 1977); Bonobos (Idani 1991); Crested Black Macaques (Poirier 1964); Rhesus Macaques (Reinhardt et al. 1986); Japanese Macaques (Sugiyama 1960); Crab-eating Macaques (Hamilton 1914); Asiatic Mouflons (Pfeffer 1967); Grizzly Bears (Craighead 1979); Long-eared Hedgehogs (Poduschka 1981); Greylag Geese (Lorenz 1991); White-handed Gibbons (Edwards and Todd 1991); Orangutans (Rijksen 1978).

For cross-cultural and other surveys of the wide variety of human homosexualities, see Ford, C. S., and F. A. Beach (1951) Patterns of Sexual Behavior (New York: Harper and Row); Bell, A. P., and M. S. Weinberg (1978) Homosexualities: A Study of Diversity Among Men and Women (New York: Simon and Schuster); Blackwood, E., ed., (1986) The Many Faces of Homosexuality: Anthropological Approaches to Homosexual Behavior (New York: Harrington Park Press); Greenberg, D. F. (1988) The Construction of Homosexuality (Chicago and London: University of Chicago Press); Murray, S.O.,ed., (1992) Oceanic Homosexualitites (New York: Garland); Plummer, K., ed., (1992) Modern Homosexualities: Fragments of Lesbian and Gay Experience (London: Routledge); Murray, S. O. (1995) Latin American Homosexualities (Albuquerque: University of New Mexico Press); Murray, S., and W. Roscoe (1997) Islamic Homosexualities: Culture, History, and Literature (New York: New York University Press).

Kangaroos: Dagg, A. I. (1984) “Homosexual Behavior and Female-Male Mounting in Mammals—a First Survey,” p. 179, Mammal Review 14:155—85. Bighorn Sheep: Weinrich, J. D. (1987) Sexual Landscapes, p. 294 (New York: Charles Scribner’s Sons). Bottlenose Dolphins (Caldwell and Caldwell 1977:804).

For some discussion of the parameters, complexities, and variations found in prison homosexuality (including male “pairing” as opposed to “rape”), see Donaldson, S. (1993) “A Million lockers, Punks, and Queens: Sex Among American Male Prisoners and Its Implications for Concepts of Sexual Orientation,” lecture delivered at the Columbia University Seminar on Homosexualities; Wooden, W. S., and J. Parker (1982) Men Behind Bars: Sexual Exploitation in Prison (New York: Plenum). For discussion of similar factors in other types of “situational” homosexuality (i.e., evidence for the nonmonolithic nature of sexual activity in all-male groups), see Williams, W. L. (1986) “Seafarers, Cowboys, and Indians: Male Marriage in Fringe Societies on the Anglo-American Frontier,” chapter 8 in The Spirit and the Flesh: Sexual Diversity in American Indian Culture, pp. 152—74 (Boston: Beacon Press).

Donaldson, S., and W. R Dynes (1990) “Typology of Homosexuality,” in W. R. Dynes, ed., Encyclopedia of Homosexuality, vol. 2, pp. 1332-37 (New York and London: Garland). Donaldson and Dynes’s typology uses three main axes, representing sexual orientation, gender expression, and temporal or chronological patterning. This triaxial schema has been expanded here to include a number of other axes.

For discussion of “axes” not specifically considered here, such as gendered homosexual interactions and the complex manifestation of gender roles in same-sex contexts, see chapter 4.

Weinrich, J. D. (1982) “Is Homosexuality Biologically Natural?” in W. Paul, J. D. Weinrich, J. C. Gonsiorek, and M. E. Hotveldt, eds., Homosexuality: Social, Psychological, and Biological Issues, pp. 197-211 (Beverly Hills, Calif.: SAGE Publications).

Gadpaille, W. J. (1980) “Cross-Species and Cross-Cultural Contributions to Understanding Homosexual Activity,” Archives of General Psychiatry 37:349-56; Dagg, “Homosexual Behavior and Female-Male Mounting in Mammals”; Vasey, P. L. (1995) “Homosexual Behavior in Primates: A Review of Evidence and Theory,” International Journal of Primatology 16:173-204.

This does not include domesticated species, in which the evidence for exclusive homosexuality is sometimes even more conclusive, as in the recent behavioral and physiological studies of domesticated sheep; see Adler, T. (1996) “Animals’ Fancies (Why Members of Some Species Prefer Their Own Sex,” Science News 151:8-9; Resko et al. 1996; Perkins et al. 1992, 1995. The question of homosexual orientation or “preference” also ties in to the common misconception that animal homosexuality is largely a matter of “necessity” or “last resort,” i.e., a response to the absence or unavailability of the opposite sex. This issue will be addressed more fully in chapter 4.

Silver Gull (Mills 1991); Greylag Goose (Huber and Martys 1993); Humboldt Penguin (Scholten 1992 and personal communication). In addition, a pair-bond between (captive) male Yellow-backed Lories has been documented as lasting more than 14 years (Low 1977:134), although (unlike the other species) this occurred in the absence of birds of the opposite sex.

Galah (Rogers and McCulloch 1981); Common Gull (Riddiford 1995); Black-headed Gull (van Rhijn and Groothuis 1985, 1987); Great Cormorant (Fukuda 1992); Bicolored Antbird (Willis 1967, 1972); Black Swan (Braithwaite 1981); Ring-billed Gull (Kovacs and Ryder 1981); Western Gull (Hunt and Hunt 1977); Hooded Warbler (Niven 1993). See also Clarke (1982:71) for documentation of a pair-bond between captive male White-fronted Amazon Parrots that lasted for at least two years.

Bottlenose Dolphin (Wells 1991, 1995; Wells et al. 1987). Another possible example is male Cheetahs, who form life-long pair-bonds or “coalitions” with each other. Although many such individuals mate with females, and sexual activity specifically between male pair-members has only recently been verified in captivity (Ruiz-Miranda et al. 1998), it is likely that at least some paired males have few if any heterosexual contacts for significant portions of their lives (especially in view of the fact that opposite-sex mating opportunities for such males are often reduced [Caro 1994:252, 304]).

Similarities and differences between homosexual and heterosexual patterns are discussed in more detail in the following section “Sexual Virtuosos.”

Kleiman, D. G. (1977) “Monogamy in Mammals,” Quarterly Review of Biology 52:39-69; Clutton-Brock, T. G. (1989) “Mammalian Mating Systems,” Proceedings of the Royal Society of London, Series B 235:339-72. In most of these pair-forming mammals homosexuality has not been reported; however, same-sex activity does occur among Gibbons, Rufous-naped Tamarins, and Wolves, but not between animals that are bonded to each other was mates. In Gibbons, homosexual interactions are incestuous—between father and son(s). In an oblique sense, then, this pattern of homosexuality involves a sort of “monogamy,” in that sexual activity is not sought outside of the primary relationship or family unit. Interestingly, Gibbons do sometimes seek promiscuous copulations with partners other than their mate—but all such cases reported so far involve heterosexual, rather than homosexual, infidelities (see, for example, Palombit 1994a,b).

Gorilla (Robbins 1995:29, 30, 38); Hanuman Langur (Rajpurohit et al. 1995:292).

Of course, it has often been claimed that homosexuality in such contexts is strictly “situational” or “by default,” i.e., due to the absence of females. This is a great oversimplification, assuming as it does that males are necessarily “forced” into living in same-sex groups and engaging in homosexual activities; for further discussion and evidence against this interpretation, see chapter 5. Moreover, homosexual behavior “by default” is still homosexual behavior. If the “motivation” or desire of participants is to be factored in so as to distinguish “genuine” homosexuality, then the same must be done for opposite-sex interactions. The fact is that heterosexual behavior in many contexts is also “situational,” not actively sought out by one or both partners, or even overtly resisted, yet it still falls under the umbrella of “heterosexuality” (see chapters 4 and 5 for further discussion and exemplification).

Mountain Zebra (Rasa and Lloyd 1994:172); American Bison (Komers et al. 1992:197, 201).

Nilgiri Langur (Hohmann 1989:445-47); Ruff (van Rhijn 1991; Hogan-Warburg 1966); White-handed Gibbon (Edwards and Todd 1991); Red Fox (Macdonald 1980:137; Schantz 1984:200; Storm and Montgomery 1975:239).

For further examples of these various forms of sequential bisexuality, consult the index. Most, if not all, of these patterns are also attested in human bisexualities; for an example of seasonal bisexuality (one of the less well known patterns), see the description of a medieval Persian practice in which men had female partners in winter and male ones in the summer (Murray and Roscoe, Islamic Homosexualities, p. 139).

Kinsey, A. C., W. B. Pomeroy, and C. E. Martin (1948) Sexual Behavior in the Human Male, pp. 638—41 (Philadelphia: W. B. Saunders).

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]).

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.

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).

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.

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.

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

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).

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

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.

Bonobo (de Waal 1995:48, 1997:117, 120; Hashimoto 1997:12); Jackdaw (Röell 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]).

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

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

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

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.

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 (Röell 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).

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).

Manakadan, R. (1991) “A Flock of One-Legged Greenshanks Tringa nebularia,” Journal of the Bombay Natural History Society 88:452.

Ring-billed Gull (Kovacs and Ryder 1983; Fetterolf et al. 1984); Japanese Macaque (Wolfe 1986:272; Gouzoules and Goy 1983:41); Greylag Goose (Huber and Martys 1993:161-62); Mallard Duck (Schutz 1965). Heg and van Treuren (1998:689) also found that bisexual trios of Oystercatchers are just as common on optimal as suboptimal territories.

Weinrich, Sexual Landscapes, p. 308.

Bonobo (de Waal 1997:107); Bottlenose Dolphin (Wells et al. 1987:294); Orang-utan (Galdikas 1981:285, 297, 1995:172; Kaplan and Rogers 1994:82).

Undoubtedly other species will be discovered that also exhibit this full range of characteristics. Many of these features are already known to characterize Stumptail Macaque sexuality, for example, including hidden estrous cycles (cf. de Waal 1989:150), anal and oral intercourse, and pairlike “sexual friendships” or “preferred partners” (and much remains to be learned about this species in the wild). Similarly, Japanese Macaques have pair-bonded consortships, face-to-face sexual encounters, and “social class” differences in sexual/pairing activity (cf. Corradino 1990:360), while Gorillas have face-to-face copulation, bonding or “preferred partners,” hidden estrous cycles (cf. Wolfe 1991:125), and oral sexual activities. Certain of these characteristics also occur individually in animal groups other than primates and cetaceans: a face-to-face mating position, for instance, is occasionally used by snow leopards, while Ruffs have a highly structured “class” system among males involving (among other features) differing sexual behaviors (Freeman, H. [1983] “Behavior in Adult Pairs of Captive Snow Leopards [Panthera uncia],” Zoo Biology 2:1-22). Another erroneous claim about human uniqueness is that no animal exhibits a type of homosexuality sometimes known as “(mutual) androphilia,” an interaction involving two adult males neither of whom adopts a stereotypically “feminine” gender presentation or sexual behavior (on the supposed absence of this in animals, see Houser, W. [1990] “Animal Homosexuality,” in W. R. Dynes, ed., Encyclopedia of Homosexuality, vol. 1, pp. 60-63 [New York and London: Garland]). In fact, exactly this sort of homosexuality occurs in Greylag Goose and Mallard Duck male pairs, as well as in a number of other species; see chapter 4 for further discussion of gender roles (or their absence) in homosexual interactions.

Weinrich, Sexual Landscapes, p. 305 (where this idea is formulated as the “technique puzzle” and characterized as “a disturbing generalization”); Masters, W.H., and V.E. Johnson (1979) Homosexuality in Perspective (Boston: Little, Brown).

Likewise, the durations of homosexual as opposed to heterosexual acts (such as mounting) are usually comparable. In some species, however, homosexual interactions generally last longer (e.g., Gorillas, White-handed Gibbons, American Bison, West Indian Manatees), while in others heterosexual encounters typically last longer (e.g., Harbor Seals, Red Foxes, Humboldt Penguins, Long-tailed Hermit Hummingbirds). In many species homosexual interactions do exhibit greater variability or flexibility in terms of the role differentiation of partners (see chapter 4 for further discussion).

Bonobo (Kitamura 1989:53—57, 61; Kano 1992:187); Gorilla (Fischer and Nadler 1978:660—61; Yamagiwa 1987a:12—14, 1987b:37; Harcourt and Stewart 1978:611—12); Hanuman Langur (Weber and Vogel 1970:76; Srivastava et al. 1991:496-97).

Japanese Macaque (Hanby and Brown 1974:164; Hanby 1974:838-40).

Flamingo (C. E. King, personal communication).

The head-to-tail position does occur in interspecies homosexual interactions with Tucuxi Dolphins. Same-species versus cross-species differences in mounting position (independent of the sex of the partner) are also found in other cetaceans. Among Bottlenose Dolphins, for example, a belly-to-belly mating position is typical of same-species contacts, both homosexual and heterosexual (cf. McBride and Hebb 1948:115, among others), while a sideways, dorsoventral position occurs in interspecies encounters with Atlantic Spotted Dolphins of both sexes (Herzing and Johnson 1997:92, 96).

Anderson, S. (1993) “Stitchbirds Copulate Front to Front,” Notornis 40:14; Tyrrell, E. Q. (1990) Hummingbirds of the Caribbean, pp. 114, 155 (New York: Crown Publishers); “Red-capped Plover, Charadrius ruficapillus ,” in S. Marchant and P. J. Higgins, eds. (1993) Handbook of Australian, New Zealand, and Antarctic Birds, vol. 2, pp. 836-47 (Melbourne: Oxford University Press); Wilkinson, R., and T. R. Birkhead (1995) “Copulation Behavior in the Vasa Parrots Coracopsis vasa and C. nigra,” Ibis 137:117—19; Kilham, L. (1983) Life History Studies of Woodpeckers of Eastern North America, pp. 49—50, 143, 160 (Cambridge, Mass.: Nuttall Ornithological Club); Southern, W. E. (1960) “Copulatory Behavior of the Red-headed Woodpecker,” Auk 77:218-19.

Vasey, “Homosexual Behavior in Primates,” p. 195.

For further details see the primate profiles in part 2, as well as the discussion of nonreproductive heterosexualities in chapter 5.

For discussion of cultural traditions in animals, including references to many specific cases, see Bonner, J. T. (1980) The Evolution of Culture in Animals (Princeton, N.J.: Princeton University Press); Galef, B. G., Jr. (1995) “Why Behavior Patterns That Animals Learn Socially Are Locally Adaptive,” Animal Behavior 49:1325—34; Lefebvre, L. (1995) “The Opening of Milk Bottles by Birds: Evidence for Accelerating Learning Rates, but Against the Wave-of-Advance Model of Cultural Transmission,” Behavioral Processes 34:43-54; Menzel, E.W., Jr., ed. (1973) Precultural Primate Behavior (Symposia of the Fourth International Congress of Primatology, vol. 1) (Basel: S. Karger); Gould, J. L., and C. G. Gould (1994) The Animal Mind (New York: Scientific American Library). For an excellent survey of animal cultural traditions, see Mundinger, P. C. (1980) “Animal Cultures and a General Theory of Cultural Evolution,” Ethology and Sociobiology 1:183-223.

Japanese Macaque (Itani 1959; Gouzoules and Goy 1983:47; Eaton 1978; Wolfe 1984:152); Stumptail Macaque (Chevalier-Skolnikoff 1976:512; Bertrand 1969:193-94); Savanna Baboon (Ransom 1981:139). In Hanuman Langurs, mounting between females may also have a cultural component, since it exhibits wide variability not only between individuals but also between geographic areas. It occurs frequently in some regions (e.g., Jodhpur, India), less frequently in others (e.g., Abu and Sariska in India), rarely in others (e.g., Sri Lanka), and not at all in still others (e.g., some parts of Nepal) (Srivastava et al.1991:504—5 [table V]). Heterosexual courtship patterns in Common Chimpanzees also exhibit cultural variations (cf. Nishida 1997:394, among others).

Bonobo (Savage-Rumbaugh et al. 1977; Savage-Rumbaugh and Wilkerson 1978; Savage and Bakeman 1978; Roth 1995; S. Savage-Rumbaugh, personal communication). Drawings, verbal descriptions, and “glosses” of the hand signals and their meanings in the accompanying illustration are based on these sources. For alternate descriptions of some of these gestures, as well as gestures used in nonsexual situations, see de Waal 1988:214-21.

Bonobo (Savage-Rumbaugh and Wilkerson 1978:334; Roth 1995:75, 88).

Bonobo (Savage-Rumbaugh et al. 1977:108).

Linguists studying the structure of American Sign Language, for example, have identified a continuum of iconicity in signs, ranging from transparent signs (quasi-mimetic gestures whose meaning is readily identifiable from their form, even to nonsigners) to translucent signs (gestures in which a connection between meaning and form can be discerned but not automatically identified without knowing the meaning of the sign) to opaque signs (gestures in which all form-meaning correspondences have been lost). According to these criteria, the Bonobo gestures would fall primarily in the transparent-translucent range. For further discussion see Klima, E.S., and U. Bellugi (1979) The Signs of Language, especially chapter 1, “Iconicity in Signs and Signing” (Cambridge: Harvard University Press).

This gestural system has only been observed in captivity, albeit in “untrained” Bonobos. Studies of wild Bonobos have so far revealed a less elaborate communicative repertoire associated with sexual interactions, although researchers have identified similar types of communicative exchanges prior to some episodes of sexual activity (e.g., Kitamura 1989:54-55; Enomoto 1990:473-75). It must also be remembered that many behaviors are easily missed in the field (especially given the particular difficulties of observing wild Bonobos; cf. de Waal 1997:12, 63—64, 70, 76—77); hence it is possible that more elaborate gestural repertoires do occur in wild Bonobos but have yet to be observed. For more on the issue of behaviors that are only observed in captivity as opposed to the wild, see chapter 4.

Hewes, G. W. (1973) “Primate Communication and the Gestural Origin of Language,” Current Anthropology 14:5-24; Hewes, G. W. (1976) “The Current Status of the Gestural Theory of Language Origin,” in S. Harnad, H. Steklis, and J. Lancaster, eds., Origins and Evolution of Language and Speech. Annals of the New York Academy of Science, vol. 280, pp. 482-504 (New York: New York Academy of Sciences).

Bonobo (Roth 1995:4-45).

Beck, B. B. (1980) Animal Tool Behavior: The Use and Manufacture of Tools by Animals (New York: Garland); Goodall 1986:545—48, 559 (Common Chimpanzee); van Lawick-Goodall, J., H. van Lawick, and C. Packer (1973) “Tool-Use in Free-living Baboons in the Gombe National Park, Tanzania,” Nature 241:212-13; McGrew, W.C. (1992) Chimpanzee Material Culture: Implications for Human Evolution (Cambridge: Cambridge University Press); Berthelet, A., and J. Chavaillon, eds., (1993) The Use of Tools by Human and Nonhuman Primates (Oxford: Clarendon Press); Weinberg, S.M., and D.K. Candland (1981) “‘Stone-Grooming’ in Macaca fuscata,” American Journal of Primatology 1:465—68.

Orang-utan (Rijksen 1978:262-63; Nadler 1982:241; Harrison 1961:61).

Common Chimpanzee (Bingham 1928:148-50; Kollar et al. 1968:456-57; Goodall 1986:559-60; McGrew, Chimpanzee Material Culture, p. 183); Bonobo (Takeshita and Walraven 1996:428; Walraven et al. 1993:28, 30; Becker, C. [1984] Orang-Utans und Bonobos im Spiel: Untersuchungen zum Spielverhalten von Menschenaffen [Orang-utans and Bonobos at Play: Investigations on the Play Behavior of Apes], pp. 149, 152, 193-94 [Munich: Profil-Verlag]). Female Japanese Macaques are also reported to use inanimate objects for masturbation (Rendall and Taylor 1991:321), although it is not clear whether this involves use of “tools” or simply rubbing of genitals against a surface (as is found in many other species). Masturbatory tool use is also occasionally reported for animals other than primates; see, for example, Shadle’s description of male and female Porcupines holding sticks in their forepaws while straddling the object in order to stimulate their genitals (Shadle, A. R. [1946] “Copulation in the Porcupine,” Journal of Wildlife Management 10:159—62; Shadle, A. R., M. Smelzer, and M. Metz [1946] “The Sex Reactions of Porcupines (Erethizon d. dorsaturn) Before and After Copulation,” Journal of Mammalogy 27:116—21). Objects or “tools” are also sometimes employed by Common Chimpanzees and Bonobos during heterosexual courtship and solicitations (cf. McGrew, Chimpanzee Material Culture, pp. 82, 188; Nishida 1997:385, 394 [Common Chimpanzee]; de Waal 1997:120 [Bonobo]).

Bonnet Macaque (Sinha 1997). Sinha (1997:23) believes that this female was using the tools to “scratch” her vagina, possibly because of “some irritation,” whose presence, however, was never confirmed. Sexual stimulation is also compatible with the observed behaviors (instead of, or along with, “scratching”), especially considering that masturbation without the use of tools occurs regularly in Bonnet Macaques of both sexes (cf. Makwana 1980:11; Kaufman and Rosenblum 1966:221; Rahaman and Parthasarathy 1969:155).

See, for example, (Rawson, P. (1973) Primitive Erotic Art, especially pp. 20, 71 (New York: G. P. Putnam’s Sons); Kinsey, A. C., W. B. Pomeroy, C. E. Martin, and P. H. Gebhard (1953) Sexual Behavior in the Human Female, p. 136 (Philadelphia: W. B. Saunders). Examples of tools utilized for sexual stimulation of partners (rather than self-stimulation) have yet to be reported for any nonhuman species. For a recent discussion of the role of sexual pleasure in the evolution of tool use among both nonhuman primates and early humans, see Vasey, P. L. (1998) “Intimate Sexual Relations in Prehistory: Lessons from Japanese Macaques,” World Archaeology 29:407—25.

For further discussion of these (and other) examples as well as cultural variation in the occurrence of incest and its taboos, see Leavitt, G. C. (1990) “Sociobiological Explanations of Incest Avoidance: A Critical Review of Evidential Claims,” American Anthropologist 92:971-93; Arens, W. (1986) The Original Sin: Incest and Its Meaning (New York and Oxford: Oxford University Press); Livingstone, F. B. (1980) “Cultural Causes of Genetic Change,” in G. W. Barlow and J. Silverberg, eds., Sociobiology: Beyond Nature/Nurture? pp. 307—29, AAAS Selected Symposium, no. 35 (Boulder: Westview Press); Schneider, D. M. (1976) “The Meaning of Incest,” Journal of the Polynesian Society 85:149—69.

For an overview of a variety of kinship restrictions on Melanesian homosexual relations, see Schwimmer, E. (1984) “Male Couples in New Guinea,” pp. 276-77, in G. H. Herdt, ed., Ritualized Homosexuality in Melanesia, pp. 248—91 (Berkeley: University of California Press); Murray, S. O. (1992) “Age-Stratified Homosexuality: Introduction,” pp. 10-12, in Murray, Oceanic Homosexualities, pp. 293-327. For more on New Guinean homosexualities, see chapter 6.

Leavitt, “Sociobiological Explanations,” pp. 974-75; Livingstone, “Cultural Causes,” p. 318. For arguments that in animals some forms of inbreeding (such as between cousins) may actually have beneficial genetic and social effects and are preferred in some species (e.g., Vervet monkeys, Japanese quail), see Moore and Ali 1984 (Bonnet Macaque); Bateson, P. (1982) “Preferences for Cousins in Japanese Quail,” Nature 295:236—37; Shields, W. M. (1982) Philopatry, Inbreeding, and the Evolution of Sex (Albany: State University of New York Press); Cheney, D. M., and R. M. Seyfarth (1982) “Recognition of Individuals Within and Between Groups of Free-Ranging Vervet Monkeys,” American Zoologist 22:519-30.

Japanese Macaque (Wolfe 1979; Chapais and Mignault 1991; Vasey 1996:543; Chapais et al. 1997); Hanuman Langur (Srivastava et al. 1991:509 [table II]; Sommer and Rajpurohit 1989:304, 310); Bonobo (Hashimoto et al. 1996:315—16). Bonobo mother-daughter homosexual relations occasionally occur (Thompson-Handler et al. 1984:355).

Savanna Baboon (Smuts and Watanabe 1990:167-70).

Berndt, R., and C. Berndt (1945) “A Preliminary Report of Field Work in the Ooldea Region, Western South Australia,” pp. 245, 260-66, Oceania 15:239-75; Meggitt, M. J. (1962) Desert People: A Study of the Walbiri Aborigines of Central Australia, pp. 262-63. (Sydney: Angus and Robertson); Eibl-Eibesfeldt, I. (1977) “Patterns of Greeting in New Guinea,” pp. 221,226, in S.A. Wurm, ed., New Guinea Area Languages and Language Study, Vol. 3: Language, Culture, Society, and the Modern World, pp. 209—47, Pacific Linguistics Series C, no. 40 (Canberra: Australian National University Press). For more on ritualized homosexuality among the Bedamini and other New Guinean peoples, see chapter 6.

Interestingly, homosexual activity in another group of highly intelligent creatures—whales and dolphins—also has many of the hallmarks of cultural activity identified here. Same-sex activity varies considerably between individuals, populations, and time periods in a number of cetaceans. For example, sexual interactions between male Killer Whales appear to differ in frequency and occurrence depending on the geographic area (Rose 1992:7), while male pairs of Bottlenose Dolphins exhibit different characteristics in various populations (see chapter 5). In addition, “incest taboos” appear to be operative in most male Killer Whale homosexual interactions (Rose 1992:112), sexual activity in Bottlenose Dolphins sometimes has a ritualistic or “greeting” component (Ostman 1991:313), while Bottlenose Dolphins of both sexes have also been observed “masturbating” or stimulating their genitals using inanimate objects (Caldwell and Caldwell 1972:430).

Hamer, D., and P. Copeland (1994) The Science of Desire: The Search for the Gay Gene and the Biology of Behavior, p. 213 (New York: Simon and Schuster).

Ward, J. (1987) “The Nature of Heterosexuality,” in G. E. Hanscombe and M. Humphries, eds., Heterosexuality, pp. 145-69. (London: GMP Publishers).

Weinrich, J. D. (1982) “Is Homosexuality Biologically Natural?” in W. Paul, J. D. Weinrich, J. C. Gonsiorek, and M. E. Hotvedt, eds., Homosexuality: Social, Psychological, and Biological Issues, pp. 197-208 (Beverly Hills, Calif: SAGE Publications). For an early discussion of animal homosexuality in relation to the question of “naturalness,” see Gide, A. (1911/1950) Corydon (New York: Farrar, Straus, and Co.).

Weinrich, ibid.; Plant, R. (1986) The Pink Triangle: The Nazi War Against Homosexuals, pp. 27, 185 (New York: Henry Holt); Grau, G., ed., (1995) Hidden Holocaust? Gay and Lesbian Persecution in Germany 1933—45, p. 284 (London: Cassell); Mann, M. (1797/1866) The Female Review: Life of Deborah Sampson, the Female Soldier in the War of the Revolution, p. 225 (Boston: J. K. Wiggin & W. P. Lunt) [excerpts reprinted in Katz, J. (1976) Gay American History, pp. 212—214 (New York: Thomas Y. Crowell)]. Boswell, J. (1980) Christianity, Social Tolerance, and Homosexuality: Gay People in Western Europe from the Beginning of the Christian Era to the Fourteenth Century, p. 309 (Chicago: University of Chicago Press).

For a summary and overview of such experimental studies (e.g., involving hormones), see Mondimore, F. M. (1996) A Natural History of Homosexuaity, pp. 111—13, 129-30 (Baltimore: Johns Hopkins University Press). These studies, typically involving laboratory rats, also invariably overlook the fact that the homosexual behaviors “induced” by hormones and other experimental treatments occur spontaneously in the wild ancestors of the laboratory animals involved, e.g., (European) Brown Rats (cf. Barnett 1958). Concerning further pitfalls in extrapolating from laboratory animals, as well as a general discussion of the “nature versus nurture” controversy, see Byne, W. (1994) “The Biological Evidence Challenged,” Scientific American 270(5):50— 55; LeVay, S., and D. H. Hamer (1994) “Evidence for a Biological Influence in Male Homosexuality,” Scientific American 270(5):44—49.

Weinrich, “Is Homosexuality Biologically Natural?” p. 207.

See chapter 5, as well as the animal profiles in part 2, for specific examples.

For explicit statements by gay-bashers to the effect that homosexuality is “not natural,” see Comstock, G. D. (1991) Violence Against Lesbians and Gay Men, p. 74 (New York: Columbia University Press).

Middleton, S., and D. Liittschwager (1996) “Parting Shots?” Sierra 81(1):40—45.

For documentation of these activities, see the following sources: Ligon, J. D. (1970) “Behavior and Breeding Biology of the Red-cockaded Woodpecker,” Auk 87:255-78; Lennartz, M. R., R.G. Hooper, and R. F. Harlow (1987) “Sociality and Cooperative Breeding of Red-cockaded Woodpeckers, Picoides borealis,” Behavioral Ecology and Sociobiology 20:77-88; Walters, J. R., P. D. Doerr, and J. H. Carter III (1988) “The Cooperative Breeding System of the Red-cockaded Woodpecker,” Ethology 78:275-305; Walters, J. R. (1990) “Red-cockaded Woodpeckers: A ‘Primitive’ Cooperative Breeder,” in P. B. Stacey and W. D. Koenig, eds., Cooperative Breeding in Birds: Long-Term Studies of Ecology and Behavior, pp. 69-101 (Cambridge: Cambridge University Press); Haig, S. M., J. R. Walters, and J. H. Plissner (1994) “Genetic Evidence for Monogamy in the Cooperatively Breeding Red-cockaded Woodpecker,” Behavioral Ecology and Sociobiology 34:295-303; Rossell, C. R., Jr., and J. J. Britcher (1994) “Evidence of Plural Breeding by Red-cockaded Woodpeckers,” Wilson Bulletin 106:557-59.

For a complete list of references, see the appendix. For an example of an anecdotal, nonscientific account, see O‘Donoghue, B. P. (1996) My Lead Dog Was a Lesbian: Mushing Across Alaska in the Iditarod—the World’s Most Grueling Race, p. 42 (New York: Vintage). Interestingly, a female dog that showed interest in both females and males is described in this book as “sexually confused” and willing to mount “any dog within reach”—some of the same subjective characterizations that appear in the scientific descriptions of bisexuality and homosexuality among wild animals (see chapters 3 and 4).

Ford and Beach, Patterns of Sexual Behavior, p. 142; Denniston, R. H. (1980) “Ambisexuality in Animals,” p. 34, in J. Marmor, ed., Homosexual Behavior: A Modern Reappraisal, pp. 25—40 (New York: Basic Books).

Kelley, K. (1978) Playboy interview: Anita Bryant, Playboy, May 1978, p. 82. Quoted in Weinrich, “Is Homosexuality Biologically Natural?” p. 198.

Lillian Faderman, interviewed in the Seattle Gay News, October 21, 1994, p. 26. See also Faderman’s mention of same-sex activities in her pet terriers in Faderman, L. (1998) “Setting Love Straight,” The Advocate, February 17, 753:72.

Chapter 3. Two Hundred Years of Looking at Homosexual Wildlife

Edwards, G. (1758-64) Gleanings of Natural History. Exhibiting figures of quadrupeds, birds, insects, plants, etc., many of which have not, till now, been either figured or described, vol. 3, p. xxi (London: Royal College of Physicians); Orang-utan (Morris 1964:502); Tree Swallow (Lombardo et al. 1994:555).

For discussion of observations of animal homosexuality in nonwestern scientific traditions, particularly those of indigenous cultures, see chapter 6.

Horapollo (1835) Hieroglyphica, Greek text edited by Conradus Leemans (Amsterdam: J. Muller; English translation by George Boas [New York: Pantheon, 1950]); Cory, A. T., ed. and trans., (1840) The Hieroglyphics of Horapollo Nilous (London: Pickering); Buffon, G. L. L. Count de (1749-67) Histoire naturelle générale et particulière (Natural History, General and Particular), 15 vols. (Paris: De l’Imprimerie royale); Edwards, Gleanings of Natural History. For an annotated bibliography of these and other early references on animal homosexuality, see Dynes, W. R. (1987) “Animal Homosexuality,” in Homosexuality: A Research Guide, pp. 743—49 (New York and London: Garland Publishing) For further discussions of Aristotle, Horapollo, and others, see Boswell, J. (1980) Christianity, Social Tolerance, and Homosexuality: Gay People in Western Europe from the Beginning of the Christian Era to the Fourteenth Century, especially chapters 6 and 11 (Chicago and London: University of Chicago Press).

Laboulmène 1859, Gadeau de Kerville 1896 (insects); Rollinat and Trouessart 1895, 1896 (Bats); Whitaker 1885 (Mute Swan); Selous 1906-7 (Ruff); Karsch, F. (1900) “Päderastie und Tribadie bei den Tieren auf Grund der Literatur” (Pederasty and Tribadism Among Animals on the Basis of the Literature), Jahrbuch für sexuelle Zwischenstufen 2:126—60.

Morris 1964 (Orang-utan), Morris 1954 (Zebra Finches), Morris 1952 (Ten-spined Stickleback); Fossey 1983, 1990, Harcourt, Stewart, and Fossey 1981, Harcourt, Fossey, Stewart, and Watts 1980 (Gorilla); Lorenz 1979, 1991 (Greylag Goose), Lorenz 1935, 1972 (Jackdaw, Raven).

Mute Swan (Low and M. of Tavistock 1935:147).

Snow Goose (Quinn et al. 1989); Oystercatcher (Heg and van Treueren 1998); Bonobo (Hashimoto et al. 1996; Roth 1995; Savage-Rumbaugh et al. 1977); Roseate Tern (Sabo et al. 1994); Ruff (Lank et al. 1995); Silver Gull (Mills 1989, 1991); Bottlenose Dolphin (Wells 1991, 1995; Wells et al. 1987); Red Fox (Macdonald 1980; Storm and Montgomery 1975); Spotted Hyena (Mills 1990); Grizzly Bear (Craighead and Craighead 1972; Craighead et al. 1995); Griffon Vulture (Mouze and Bagnolini 1995); Victoria’s Riflebird (Frith and Cooper 1996); Black-winged Stilt (Kitagawa 1988a).

Separation—Rhesus Macaque (Erwin and Maple 1976); Bottlenose Dolphin (McBride and Hebb 1948); Cheetah (Ruiz-Miranda et al. 1998); Long-eared Hedgehog (Poduschka 1981); Black-headed Gull (van Rhijn 1985; van Rhijn and Groothuis 1987); see also Clarke 1982:71 (White-fronted Amazon Parrot); removal—Orange-fronted Parakeet (Hardy 1963:187); electrodes—Stumptail Macaque (Goldfoot et al. 1980); deafening—Squirrel Monkey (Talmage-Riggs and Anschel 1973); castration—Crab-eating, Rhesus Macaques (Hamilton 1914); White-tailed Deer (Taylor et al. 1964); lobotomy—Domestic Cats (Green et al. 1957); killing, tissue collection—Common Garter Snake (Noble 1937); Hooded Warbler (Niven 1993); Gentoo Penguin (Roberts 1934). For primate hormonal treatment studies relating to homosexuality, see the literature survey in Vasey, P. L. (1995) “Homosexual Behavior in Primates: A Review of Evidence and Theory,” International Journal of Primatology 16:173-204. For examples of hormonal treatments administered to transgendered animals, see Savanna Baboon (Bielert 1984b, 1985); White-tailed Deer (Thomas et al. 1970).

Wolfe, L. D. (1991) “Human Evolution and the Sexual Behavior of Female Primates,” p. 130, in J. D. Loy and C. B. Peters, eds., Understanding Behavior: What Primate Studies Tell Us About Human Behavior, pp. 121-51 (New York: Oxford University Press). For another example of the extent to which scientific information about animal homosexuality remains unpublished (thereby perpetuating inaccuracies), see Weinrich’s account of how he had to obtain much of his information from personal conversations and letters with zoologists—a procedure that was still necessary, a decade later, in the preparation of this book (Weinrich, J. D. [1987] Sexual Landscapes, p. 308 [New York: Charles Scribner’s Sons]).

See, for example, Hubbard, R., M. Henifin, and B. Fried, eds., (1979) Women Look at Biology Looking at Women: A Collection of Feminist Critiques (Cambridge: Schenkman); Hrdy, S. B., and G. C. Williams (1983) “Behavioral Biology and the Double Standard,” in S. K. Wasser, ed., Social Behavior of Female Vertebrates, pp. 3-17 (New York: Academic Press); Shaw, E., and J. Darling (1985) Female Strategies (New York: Walker and Company); Kevles, B. (1986) Females of the Species: Sex and Survival in the Animal Kingdom (Cambridge, Mass.: Harvard University Press); Haraway, D. (1989) Primate Visions: Gender, Race, and Nature in the World of Modern Science (New York: Routledge); Gowaty, P. A., ed. (1996) Feminism and Evolutionary Biology: Boundaries, Intersections, and Frontiers (New York: Chapman Hall); Cunningham, E., and T. Birkhead (1997) “Female Roles in Perspective,” Trends in Ecology and Evolution 12:337-38. On the general male-centeredness of most biological theorizing, see Eberhard, W. G. (1996) Female Control: Sexual Selection by Cryptic Female Choice, pp. 34-36. (Princeton: Princeton University Press); Batten, M. (1992) Sexual Strategies (New York: Putnam’s); Gowaty, P. A. (1997) “Principles of Females’ Perspectives in Avian Behavioral Ecology,” Journal of Avian Biology 28:95-102.

This is not to suggest, of course, that only scientists who are themselves homosexual can deal with the subject in an unbiased way. Certainly many contemporary heterosexual biologists do not harbor negative views about homosexuality, while some gay and lesbian zoologists have undoubtedly perpetuated the silences and prejudices of their field. (There are also those who believe that being homosexual actually invalidates a gay or lesbian scientist’s objectivity on the subject. However, if sexual orientation resulted in such bias, then heterosexual zoologists should confine themselves only to research topics that have nothing to do with breeding or male-female relations.) Nevertheless, sexism and male bias in biology have been exposed most directly through the work of women and feminist scientists, and it is likely that similar insights regarding heterosexism and homophobia will be forthcoming from openly gay, lesbian, or bisexual zoologists—that is, once such people no longer have to fear losing tenure, research grants, or jobs because of their outspokenness. Regardless of their own sexual orientation, however, many zoologists have avoided studying homosexuality or speaking widely about their results because the topic is still far from being considered a “legitimate” area of inquiry (see, for example, Wolfe’s commentary above; also, Anne Perkins’s decision not to discuss her findings on homosexuality in domestic sheep until after she had secured tenure, reported in “Counting Sheep,” Advocate, July 8, 1997, 737:21). A parallel situation exists in the fields of anthropology and history, where denial, omission, suppression, and condemnation of information about human homosexuality have long been carried out by researchers studying other cultures or historical periods. For a particularly good discussion of this phenomenon, see Read, K. E. (1984) “The Nama Cult Recalled,” in G. H. Herdt, ed., Ritualized Homosexuality in Melanesia, pp. 211-47 (Berkeley: University of California Press). On the myth of observer “objectivity” where discussion of homosexuality by anthropologists is concerned, see Lewin, E., and W. L. Leap, eds. (1996) Out in the Field: Reflections of Lesbian and Gay Anthropologists (Urbana and Chicago: University of Illinois Press). For further discussion of indigenous human homosexualities, see chapter 6.

Dagg, A. I. (1984) “Homosexual Behavior and Female-Male Mounting in Mammals—a First Survey,” Mammal Review 14:155-85; Vasey, “Homosexual Behavior in Primates”; Vasey 1996, 1998 (Japanese Macaque); Vasey, P. L. (in press) “Homosexual Behavior in Male Birds,” “Homosexual Behavior in Male Primates,” in W. R. Dynes, ed., Encyclopedia of Homosexuality, 2nd ed., vol. 1: Male Homosexuality (New York: Garland Press). See also the recent bibliography: Williams, J. B. (1992) Homosexuality in Nonhuman Primates: A Bibliography: 1940—1992 (Seattle: Primate Information Center). For descriptions of animal homosexuality that are relatively value neutral (i.e., that do not view homosexual behavior as inherently problematic), or for accounts that are not overly concerned with finding a “cause” or “explanation” for the behavior, see, for example, Yeager 1990a (Proboscis Monkey); Marlow 1975 (Australian, New Zealand Sea Lions); Sowls 1974, 1984 (Collared Peccary); Schaller 1967 (Blackbuck, Barasingha); Braithwaite 1981 (Black Swan); King 1994 (Flamingo); Riddiford 1995 (Common Gull); Smith 1988 (Lyrebird); Neelakantan 1962 (Black-rumped Flameback); and Rogers and McCulloch 1981, Rowley 1990 (Galah). For descriptions of homosexual activity that recognize it as a routine or “normal” behavioral phenomenon, see Porton and White 1996 (Gorilla); Akers and Conaway 1979 (Rhesus Macaque); Eaton 1978, Fedigan 1982, Wolfe 1984, 1986, Chapais and Mignault 1991, Vasey 1996 (Japanese Macaque); Chevalier-Skolnikoff 1976 (Stumptail Macaque); Wells et al. 1987, Wells 1991, Wells et al. 1998 (Bottlenose Dolphin); Rose 1992 (Killer Whale); Hartman 1971, 1979 (West Indian Manatee); Lott 1983 (American Bison); and Coe 1967 (Giraffe). In addition, a number of zoologists in their personal communications with me have been refreshingly free of the negative judgments or interpretations that unfortunately characterize most of the field; among them, B. J. Ens (Oystercatchers), C. B. Frith (Birds of Paradise), M. Fujioka (Egrets), M. Fukuda (Great Cormorants), D. Heg (Oystercatchers), D. L. Herzing (Dolphins), C. E. King (Flamingos), W. D. Koenig (Acorn Woodpeckers), D. F. Lott (American Bison), M. Martys (Greylag Geese), M. G. L. Mills (Spotted Hyenas), C. Reed (Crested Black Macaques), S. Savage-Rumbaugh (Bonobos), C. J. Scholten (Humboldt Penguins), L. H. Smith (Superb Lyrebirds), Y. Sugiyama (primates), and P. L. Vasey (Japanese Macaques, other species).

While the word homophobia means, literally, an irrational fear of homosexuality, the term is also applied to instances of disgust, revulsion, hatred, or open hostility, as well as more subtle prejudicial feelings of discomfort, distaste, or dislike toward homosexuality or homosexual individuals (not necessarily accompanied by fear). For more discussion and further references on the nature and consequences of homophobia, see Blumenfeld, W. J., ed. (1992) Homophobia: How We All Pay the Price (Boston: Beacon Press).

Ruff (Selous 1906—7:420, 423); American Bison (McHugh 1958:25); Waterbuck (Spinage 1982:118).

The appellations abnormal, aberrant, unnatural, or perverted, for example, have been applied by scientists to homosexual behavior or transgender in at least 30 different species of mammals and birds (often in multiple sources for each species), and as recently as the mid-1980s in some published accounts (Kittiwake [Coulson and Thomas 1985:20]; Bighorn Sheep [Berger 1985]). Even more recently (Finley et al. 1997:914—15, 917), same-sex courtship and sexual activity in Fruit Flies (as well as refusal of heterosexual advances) have been characterized as “abnormal,” “aberrant,” and a “defect,” and similar terms have also been used by some zoologists in their personal communications with me. Somewhat less derogatory designations such as odd (including odd couples), peculiar, irregular, or bizarre have been used to describe homosexuality or transgender in at least 15 other species of mammals and birds. Many other examples can of course be found in descriptions of reptiles, amphibians, fishes, insects, and other creatures. Heterosexual behaviors or individuals are characterized as “normal” in opposition to homosexual behaviors or individuals in the following published scientific sources, among others: Common Chimpanzee (Adang et al. 1987:242); Gorilla (Harcourt 1988:59); Kob (Buechner and Schloeth 1965:2219); Canada Goose (Collias and Jahn 1959:484); Black-winged Stilt (Kitagawa 1988a:64); Black-headed Gull (van Rhijn and Groothuis 1985:161); Lovebirds (Dilger 1960:667); Hooded Warbler (Niven 1994:192); Ostrich (Sauer 1972:729).

Gadeau de Kerville (1896); Grollet and L. Lepinay (1908) “L’inversion sexuelle chez les animaux” (Sexual Inversion in Animals), Revue de l’hypnotisme 23:34-37; Savanna Baboon (Marais 1922/1969); Bengalese Finch (Masatomi 1957); Ostrich (Sauer 1972); Long-eared Hedgehog (Poduschka 1981); Whiptail Lizard (Crews and Young 1991).

Mazarine Blue (Tennent 1987:81-82).

Domestic Cattle (Klemm et al. 1983:187); Elephants (Rosse 1892:799); Lion (Cooper 1942:26-28); Buff-breasted Sandpiper (Myers 1989); Domestic Turkey (Hale 1955:1059); Spinner Dolphin (Wells 1984:470); Killer Whale (Rose 1992:112); Caribou (Bergerud 1974:420); Adélie Penguin (Davis et al. 1998:137); Black-billed Magpie (Baeyens 1979:39-40); Guianan Cock-of the-Rock (Trail 1985a:238-39); Sage Grouse (Scott 1942:494). Other terms, while not necessarily derogatory, reflect scientists’ particular interpretations of such behavior as substitute or counterproductive activities: same-sex mounting in Gorillas is called “vicarious” sexual activity (Fossey 1983:74, 188—89), and homosexual mounting in African Buffalo is categorized as “barren sexual behavior” (Mloszewski 1983:186). See also the subsequent section “Mock Courtships and Sham Matings” for discussion of the widespread use of terms such as false or mock sexual behavior to characterize homosexual activity, and chapters 4 and 5 for other interpretations of homosexuality.

Long-eared Hedgehog (Poduschka 1981:84, 87); Eastern Gray Kangaroo (Grant 1974:74); Black-crowned Night Heron (Noble et al. 1938:29); King Penguin (Gillespie 1932:95, 98); Gorilla (Harcourt 1988:59); Lorikeets (Low 1977:24); Red Fox (Macdonald 1987:101); Greenshank (Nethersole-Thompson and Nethersole-Thompson 1979:112—13; Nethersole-Thompson 1951:109).

This is not to say, of course, that homosexual “advances” are never unwanted. Various forms of nonconsensual courtship or sexual approaches between animals of the same sex have been reported in about a quarter of the mammal and bird species exhibiting homosexuality. However, in many cases they co-occur with “consensual” homosexual interactions in the same species, from which they are clearly distinguished by behavioral indications of unwillingness on the part of one partner. As in nonconsensual heterosexual interactions (which are reported in more than a third of the species in which homosexual behavior has been documented and in general are equally, if not more, prevalent in animals—see chapter 5), there is actually a continuum of disinterest and “refusal” behavior. An animal may signal its unwillingness by not permitting any sexual approaches or contact at all, by permitting sexual contact but not facilitating the interaction, or by actively interrupting contact (either by trying to get away or by attacking the other animal). Assertions by scientists of “unwanted” homosexual attentions are usually anthropomorphic projections made regardless of whether such behavioral evidence is present (or what degree of nonconsensuality is involved).

Mountain Sheep (Geist 1975:100); Rhesus Macaque (Carpenter 1942:137, 151-52); Laughing Gull (Noble and Wurm 1943:205—6); Cattle Egret (Fujioka and Yamagishi 1981:139); Sage Grouse (Gibson and Bradbury 1986:396); Orang-utan (Rijksen 1978:264-65); Kob (Buechner and Schloeth 1965:211-12, 217, 219); Ostrich (Sauer 1972:729, 733); Guianan Cock-of-the-Rock (Trail and Koutnik 1986:210-11, 215); Mallard Duck (Schutz 1965:458); Rhesus Macaque (Kempf 1917:136). One zoologist also reveals something of his own misconceptions concerning both homosexual and heterosexual intercourse when he expresses surprise that a female Bonobo “on the bottom” during a lesbian interaction does not appear to mind—in fact, visibly enjoys—being in that position: “If we were on the bottom being held down, we would probably feel submissive and inferior, but female pygmy chimpanzees seem not to take it that way … the female on the bottom … looks proud and affectionate” (Kano 1992:193).

Greylag Goose (Huber and Martys 1993:161); see Lorenz (1991:241—42) on gander pairs being more closely bonded than heterosexual pairs.

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

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.

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 Edicións); 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).

Gray Whale (Darling 1977:10—11).

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.

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).

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

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.

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).

King Penguin (Gillespie 1932:96-120).