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      {683} Chapter 3. Two Hundred Years of Looking at Homosexual Wildlife

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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).

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

      3
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 generale et particuliere (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).

      4
Laboulmene 1859, Gadeau de Kerville 1896 (insects); Rollinat and Trouessart 1895, 1896 (Bats); Whitaker 1885 (Mute Swan); Selous 1906-7 (Ruff); Karsch, F. (1900) "Paderastie und Tribadie bei den Tieren auf Grund der Literatur" (Pederasty and Tribadism Among Animals on the Basis of the Literature), Jahrbuch fur sexuelle Zwischenstufen 2:126-60.

      5
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).

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

      7
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).

      8
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).

      9
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]).

      10
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.
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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.

      12
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).

      13
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).

      14
Ruff (Selous 1906-7:420, 423); American Bison (McHugh 1958:25); Waterbuck (Spinage 1982:118).

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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).
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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).

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Mazarine Blue (Tennent 1987:81-82).

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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); Adelie 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.

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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).

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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).

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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).

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

      23
Ocellated Antbird (Willis 1973:31); on heterosexual divorce in Antbirds, see Willis (1983:414).

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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, {686} 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.

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

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Gray Whale (Darling 1977:10-11).

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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.

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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).

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

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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.

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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).

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King Penguin (Gillespie 1932:96-120).

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

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

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Squirrel Monkey (Talmage-Riggs and Anschel 1973:70-71); Bonobo (Savage-Rumbaugh and Wilkerson 1978:338; Savage and Bakeman 1978:614); Spotted Hyena (Burr 1996:118-19). For conflicting information on the occurrence of clitoral penetration in Spotted Hyenas, see Glickman (1995). See also Morris (1956:261), who defines courtship as "the heterosexual reproductive communication system leading up to the consummatory sexual act" (Morris, D. [1956] "The Function and Causation of Courtship Ceremonies," in M. Autuori and Fondation Singer-Polignac, L'instinct dans le comportement des animaux et de l'homme [Paris: Masson et Cie.])
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Savanna (Chacma) Baboon (Marais 1922/1969:215).

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Ruff (van Rhijn 1991:21); Bonnet Macaque (Nolte 1955:179).

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Walrus (Miller and Boness 1983:305); African Elephant (Shelton 1965:163-64); Gorilla (Maple, T. [1977] "Unusual Sexual Behavior of Nonhuman Primates," in J. Money and H. Musaph, eds., Handbook of Sexology, pp. 1169-70 [Amsterdam: Excerpta Medica]); Sage Grouse (Scott 1942:495); Hanuman Langur (Mohnot 1984:349); Common Chimpanzee (Kortlandt 1962:132); Musk-ox (Reinhardt 1985:297-98); Mallard Duck (Lebret 1961:111-12); Blue-bellied Roller (Moynihan 1990:17); Lion (Cooper 1942:26-28); Orang-utan (Rijksen 1978:257); Savanna Baboon (Noe 1992:295, 311); Mule Deer (Halford et al. 1987:107); Hammerhead (Brown 1982:171; Campbell 1983:11); Bonobo (Thompson-Handler et al. 1984:358; de Waal 1987:319, 1997:102); Japanese Macaque (Green 1975:14); Rhesus Macaque (Reinhardt et al. 1986:56); Red Fox (Macdonald 1980:137); Squirrels (Ferron 1980; Horwich 1972; Reilly 1972). A few of these terms are also applied to nonreproductive heterosexual activities, in which case the attribution of "falseness" refers to the fact that the behavior does not result in procreation rather than to a same-sex context per se. See chapter 5 for further discussion of the parallel treatment of nonreproductive heterosexual behaviors as "abnormal" in the history of zoology.

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The categorization of homosexual activity as less than "genuine" sexual activity is an important issue, and the various ways that same-sex activity is desexualized will be examined in greater detail in the next section.

      40
Bonobo (Kano 1992); Common Chimpanzee (de Waal 1982); Snow Goose (Diamond 1989); Lesser Flamingo (Alraun and Hewston 1997); Oystercatcher (Heg and van Treuren 1998); Black-billed Magpie (Baeyens 1979); Black Stilt (Reed 1993); Fruit Flies (Cook 1975); Long-legged Fly sp. (Dyte 1989).

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Gowaty, P. A. (1982) "Sexual Terms in Sociobiology: Emotionally Evocative and Paradoxically, Jargon," Animal Behavior 30:630-31. The title of the article in question (Abele and Gilchrist 1977, on Acanthocephalan Worms) also contained the word rape, so it is possible that scientists were "snickering" at this as well. Gowaty suggests replacing, along with unisexual for homosexual, all "loaded" terminology with more "neutral" words, e.g., forced copulation for rape, kleptogamy for cuckoldry, one-male social unit for harem. Many of her arguments for such alternate terminology are valid, e.g., that the "loaded" terms are often scientifically inaccurate. Notably, however, her principal argument against the word homosexual is not that it is inaccurate, but that use of this term is "sensationalistic" and triggers the prejudices of other scientists, thereby preventing them from seeing past the word to what it describes. It should also be pointed out that many formerly controversial terms for heterosexual behaviors are now acceptable in scientific circles. The word divorce, for example, was first greeted with an "uproar" when used to describe the break-up of pair-bonds in birds, and numerous scientists suggested replacing it with more "neutral" words; yet the term is now widely used in the ornithological literature (Milius, S. [1998] "When Birds Divorce: Who Splits, Who Benefits, and Who Gets the Nest," p. 153, Science News 153:153-55).

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Giraffe (Coe 1967:320; Leuthold, W. [1977] African Ungulates: A Comparative Review of Their Ethology and Behavioral Ecology, p. 130 [Berlin: Springer-Verlag]).

      43
Connor, J. (1997) "Courtship Testing," Living Bird 16(3)31-32; Depraz, V, G. Leboucher, L. Nagle, and M. Kreutzer (1997) "'Sexy' Songs of Male Canaries: Are They Necessary for Female Nest-Building?" in M. Taborsky and B. Taborsky, eds., Contributions to the XXV International Ethological Conference, p. 122, Advances in Ethology no. 32 (Berlin: Blackwell Wissenschafrs-Verlag); Emlen, S. T., and N. J. Demong (1996) "All in the Family," Living Bird 15(3):30-34; Savanna Baboon (Smuts 1985:223, 1987:39, 43); Tasmanian Native Hen (Goldizen et al. 1998); Mirande, C. M., and G. Archibald (1990) "Sexual Maturity and Pair Formation in Captive Cranes at the International Crane Foundation," in AAZPA Annual Conference Proceedings, pp. 216-25 (Wheeling, W.Va.: American Association of Zoological Parks and Aquariums); Bonobo (de Waal 1997:117); Eisner, T., M.A. Goetz, D. E. Hill, S. R. Smedley and J. Meinwald (1997) "Firefiy 'Femmes Fatales' Acquire Defensive Steroids (Lucibufagins) from Their Firefly Prey," Proceedings of the National Academy of Sciences 94:9723-28; Domestic Goat (Shank 1972:500). See also the discussion of red-cockaded woodpecker "family values" in chapter 2.

      44
Greylag Goose (Lorenz 1991:241-43) (see also Lorenz's own assertion, in this same passage, that such male pairs are not simply platonic "friendships" between males but are equivalent to male-female mated pairs). Analogously, Kortlandt (1949) (Great Cormorant) labels same-sex pairs "pseudohomosexual" rather than "homosexual" if their members later form heterosexual bonds, once again equating "true" homosexuality with lifetime, exclusive same-sex pairing. See chapter 2 for more on the dubious notion of "true" homosexuality and its relation to more sophisticated characterizations of sexual orientation. Lorenz's unwillingness to apply the term homosexual to gander pairs and thereby invite human-animal comparisons (or imply full heterosexual-homosexual equivalence) is especially problematic in light of his activities during the Third Reich. As a member of the Nazi party in Austria and an official lecturer for its Office of Race Policy, Lorenz did not hesitate to draw analogies between animals and people to support and develop the doctrines of "biological degeneracy," "racial purity," and the "elimination" of "inferior" or "asocial" elements (Deichmann, U. [1996] Biologists under Hitler, especially "Konrad Lorenz, Ethology, and National Socialist Racial Doctrine," pp. 179-205. Cambridge, Mass.: Harvard University Press). Among his most blatant assertions in this regard are statements (in 1943) that physical and moral "decay" in people is "identical" to the effects of domestication on animals and (in 1940) that the "defective type" among humans is like "the domesticated animal that {688} can be bred in the dirtiest stable and with any sexual partner" (ibid., pp. 186, 188; cf. Lorenz's [1935/1970:203] surmise that same-sex pairing in Jackdaws only occurs in captive animals and is not a feature of "natural" populations). He also asserted (in 1941) that "Precisely in the large field of instinctive behavior, humans and animals can be directly compared ... . We confidently venture to predict that these studies will be fruitful for both theoretical as well as practical concerns of race policy" (ibid., p. 186). The subject of how the antihomosexual climate of Nazi Germany and the Nazi sympathies of some biologists helped shape the scientific discourse on animal homosexuality deserves further investigation. Many zoological studies of this phenomenon, after all, were written in Germany and Austria during this period or were heavily influenced by work that had its genesis during this time. Moreover, one of the earliest scientific surveys of animal homosexuality (Karsh, "Paderastie und Tribadie bei den Tieren" [1990]), appeared in the periodical Jahrbuch fur sexuelle Zwischenstufen (Yearbook for Sexual Intermediate Types), published by the noted Jewish homosexual Magnus Hirschfield, whose mammoth archives and library of sexology were later destroyed by the Nazis.

      45
Western Gull (Hayward and Fry 1993). See also Diamond and Burns, who suggest that same-sex pairing in Gulls should be referred to as "joint brooding" or "coparenting" rather than as homosexuality, thereby emphasizing its supposed reproductive functions (Diamond, M., and J. A. Burns [1995] "Human-Nonhuman Comparisons in Sex: Valid and Invalid," paper presented at the 24^th International Ethological Conference, Honolulu, Hawaii). For arguments that same-sex pairing is not primarily a reproductive behavior, see chapters 4 and 5.

      46
For examples of scientists who use the term homosexual (or lesbian or gay) even when no overt sexual activity is involved (i.e., to refer to related behaviors such as courtship, pairing, or parenting), see Sauer 1972 (Ostrich), Nethersole-Thompson 1975 (Scottish Crossbill), Wingfield et al. 1980 (Western Gull), Braithwaite 1981 (Black Swan), Smith 1988 (Lyrebird), Diamond 1989 (Snow Goose), Reed 1993 (Black Stilt).

      47
And in fact just such a "broad" definition of heterosexuality is required in many cases. "Heterosexual pairs" in which little or no sexual activity occurs between partners have been reported for Greylag Geese (as mentioned above) and Lesser Scaup Ducks (Afton 1985:150), among others; see also Loy (1971:26) for "sexual" bonds between male and female Rhesus Macaques that do not involve mounting or copulation, and Smuts (1985:18, 163-66, 199, 213) on the platonic "pair-bonds" or "friendships" between male and female Savanna Baboons. In addition, in some "heterosexual pairs" of splendid fairy-wrens all offspring are fathered by males other than the female's pair-bonded mate (i.e., she does not copulate — or at least is not fertilized by — her partner); see Russell, E., and L. Rowley (1996) "Partnerships in Promiscuous Splendid Fairy-wrens," in J. M. Black, ed., Partnerships in Birds: The Study of Monogamy, pp. 162-73 (Oxford: Oxford University Press). For an example of a "broad" definition of (hetero)sexuality that encompasses courtship activities in addition to overt copulatory behavior, see Tinbergen, N. (1965) "Some Recent Studies of the Evolution of Sexual Behavior," in F. A. Beach, ed., Sex and Behavior, pp. 1-33 (New York: John Wiley and Sons).

      48
In discussing the possible dangers of anthropomorphism in terminology, the comments of biologist John Bonner are instructive: "An anthropologist might find the use of words such as slaves or castes for ant colonies most undesirable ... . For instance, it implies that the most repugnant human morals are ascribed to the members of some species of ant ... . Much worse, it could imply that if ants have slavery, it is a natural thing to do and therefore quite justified in a human society. These arguments are not quite rational and can only be advanced under extreme fervor of one sort or another. A more reasoned objection would be that the motivations of ants and men might differ radically, but by using the same words this distinction is lost. A biologist, on the other hand, feels that the points made above are too obvious to interfere with the dual use of the words. He does not see any problem: in both ant and human slavery individuals forcibly capture members of their own species or related species and cause their captives to do work for the benefit of the captors. It is unnecessary to drag in all the possible political, psychological, or strictly human nuances; a very simple definition of the word is sufficient. There is no need to be tyrannized by words. If a biologist may not use the common words, he will be forced to invent a whole new set of jargon terms for nonhuman societies, an unfortunate direction since there are too many jargon words in any science as it is. I hope it will be sufficient if I make it clear in the beginning that words either invented or frequently used for human societies will also be used for animal societies with the understanding that I am not implying anything human in their meaning; they are to be considered simple descriptions of conditions." (Bonner, J. T. [1980] The Evolution of Culture in Animals, pp. 9-10. [Princeton, N.J.: Princeton University Press].) Unfortunately, this eminently reasonable position has not been adopted by most biologists where homosexuality is concerned; for a counterview, sec Gowaty, "Sexual Terms in Sociobiology."

      49
Examples of species in which homosexual activity is given only cursory treatment compared to heterosexual activity are too numerous to list, but include White-tailed Deer (Hirth 1977), Wapiti (Harper et al. 1967), Fat-tailed Dunnart (Ewer 1968), Matschie's Tree Kangaroo (Hutchins et al. 1991), Wattled Starling (Sontag 1991), Sage Grouse (Wiley 1973, Gibson and Bradbury 1986), and Canary-winged Parakeet (Arrowood 1988). In a few studies, however, detailed quantitative and descriptive information is provided on homosexual behavior; see, for example, Kitamura 1989, Kano 1992, de Waal 1987, 1995, 1997 (Bonobo); Edwards and Todd 1991 (White-handed Gibbon); Hanby 1974, Eaton 1978, Chapais and Mignault 1991, Vasey 1996 (Japanese Macaque); Pratt and Anderson 1985 (Giraffe); Jamieson and Craig 1987a (Pukeko); van Rhijn and Groothuis 1985, 1987 (Black-headed Gull); Rogers and McCulloch 1981 (Galah). For further discussion of how same-sex activity has frequently not been considered "genuine" sexual behavior, see the next section.
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      50
Spinner Dolphin (Wells 1984:468; Bateson 1974); Kob (Buechner and Schloeth 1965:219 [table 21]); Crested Black Macaque (Dixson 1977); Brown Capuchin (Linn et al. 1995); Giraffe (Dagg and Foster 1976:75-77).

      51
Western Gull (Hunt et al. 1984:160) (see Hayward and Fry [1993:16, 18] for a recent reiteration of the findings of this study, in which sexual activity is once again downplayed); Black-crowned Night Heron (Noble et al. 1938:28-29); on comparable levels of crowding in wild colonies, see Gross 1923:13-15; Davis 1993:6; Kazantzidis et al. 1997:512); Laughing Gull (Hand 1985:128); Canary-winged Parakeet (Arrowood 1988. 1991); Greater Rhea (Fernandez and Reboreda 1998:341); Zebra Finch (Burley 1981:722).

      52
Gorilla (Harcourt 1979a:255). Harcourt et al. (1981:266) also characterize heterosexual copulation as "rare." In addition, they report directly observing only 69 episodes of heterosexual mating (other copulations were heard but not seen) compared to 10 episodes between females, which yields an even higher proportion of nearly 13 percent homosexual activity.

      53
Western Gull (Hunt et al 1980:474); Spotted Hyena (Glickman 1993; Burr 1996:118-19); for further discussion of comparisons between wild and captive animals, see the next chapter

      54
Tree Swallow (M. P. Lombardo, personal communication: Venier et al. 1993:413; Lombardo 1986; Leffelaar and Robertson 1984:78). Similarly, homosexual mounting is claimed to be very rare in Northern Fur Seals, yet most heterosexual matings in this species are missed by observers because they occur at night (Gentry 1998:75-77, 107, 145).

      55
For specific examples, see Nilgiri Langur (Poirier 1970; Hohmann 1989). White tailed Deer (Hirth 1977: Rue 1989), Mule Deer (Geist 1981; Halford et al. 1987; Wong and Parker 1988), Red Deer (Lincoln et al. 1970; Guiness et al. 1971; Hall 1983), American Bison (McHugh 1958: Lott 1983 and personal communication), Red Squirrel (Layne 1954; Smith 1968; Ferron 1980), Mallard Duck Ramsey 1956; Lebret 1961; Schutz 1965; Bossema and Roemers 1985; Geh 1987), Ruff (Selous 1906-7; Bogan-Warburg 1966; Scheufler and Stiefel 1985; van Rhijn 1991), Oystercatcher (Makkink 1942; Heg and van Treuren 1998), Hooded Warbler (Niven 1994 and personal communication), Cliff Swallow (Emlen 1954, Barlow et al. 1963; Brown and Brown 1996), Red-backed Shrike (Owen 1946; Ashby 1958; Pounds 1972), Victoria's Riflebird (Bourke and Austin 1947; Frith and Cooper 1996; C. B. Frith, personal communication), Sage Grouse (Scott 1942; Patterson 1952; Wiley 1973; Gibson and Bradbury 1986), Acorn Woodpecker (MacRoberts and MacRoberts 1976; Troetschler 1976; W. D. Koenig, personal communication), Gentoo Penguin (Robert 1934; Wheater 1976; Stevenson 1983), and the examples of wild versus captive observations in notes 99-100, chapter 4.

      56
Pukeko (Craig 1980:594; Jamieson and Craig 1987a;1252); Blak-headed Gull (van Rhijn and Groothuis 1985:161, 165).

      57
For example, Vasey ("Homosexual Behavior in Primates," p. 181) sets up a general frequency scale in which homosexual behavior is classified as "rare" if it occurs "5 percent or less frequently as heterosexual behavior" and "occasional" if it occurs "6-24 percent as frequently as heterosexual behavior"; it is regarded as "frequent" only if it occurs "25 percent or more frequently."Certainly this scale is to be commended for its standardization and multipoint assessment criteria (which also include nonquantitative measures); yet (like most scales) it is not without arbitrarines, and it is at odds with the heterosexual "5 percent" criterion. The "Polygyny threshold" model, which recognizes a frequency of [?]5 percent as significant for "minority" heterosexual mating systems (i.e., polygamy in otherweise monogamous species) was originally proposed by Verner, J., and M. P. Willson (1966) "The Influence of Habitats on Mating Systems of North American Passerine Birds," Ecology 47:143-47. The 5 percent threshold continues to be widely used as a criterion for "regular" polygyny — for more recent examples, see Quinney 1983 (Tree Swallow); Moller, A. P. (1986) "Mating Systems Among European Passerines. A Review," Ibis 128:234-50; Petit, L. J. (1991) "Experimentally Induced Polygyny in a Monogamous Bird Species: Prothonotary Warblers and the Polygyny Threshold," Behavioral Ecology and Sociobiology 29:177-87.

      58
House Sparrow/Cowbird (Griffin 1959); Savanna Baboon (Marais 1922/1969:214-18); Kestrel (Olsen 1985). Regarding the House Sparrow/Cowbird case, a number of subsequent researchers (e.g., Selander and LaRue 1961; Rothstein 1980) have also interpreted this behavior as "aggression" or "appeasement." Aside from the fact that the activity involving homosexual mounting is not identical to strictly "aggressive" or "preening invitation" displays in Cowbirds (cf. Laskey 1950), a "nonsexual" interpretation cannot explain why Cowbirds "tolerate" homosexual mountings from Sparrows and even actively solicit them. Moreover, the function(s) of these "head-down" displays remain controversial and speculative independent of any homosexual activity (cf. Scott and Grumstrup-Scott 1983). Specific arguments against an "aggressive" or "appeasement" interpretation of these types of behaviors (regardless of whether any same-sex mounting is involved) are presented in Verbeek, N. A. M., R. W. Butler, and H. Richardson (1981) "Interspecific Allopreening Solicitation in Female Brewer's Blackbirds," Condor 83:179-80. A parallel example involves Stonor (1937:88), who "reinterprets" Selous's (1906-7) early descriptions of homosexual mountings by female Ruffs as involving heterosexual activity by "female-plumaged" males. More recent observers (e.g., Hogan-Warburg 1966, van Rhijn 1991) have corroborated Selous's original observations, confirming not only the existence of both female and male homosexual activity, but also "female plumaged" males (i,e., the so-called naked-nape males) that participate in homosexuality.
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      59
Chaffinch (Marjakangas 1981); Regent Bowerbird (Phillipps 1905; Marshall 1954). Similarly, early reports of courtship activity between male Swallow-tailed Manakins by Sick (1959, 1967) were discounted by Foster (1981:174), who tried to claim that the younger male birds being courted by adult males were actually females that had malelike plumage or were male observers or participants in nonsexual aggressive displays. However, Sick (1959:286) verified the male sex of these birds by dissecting them, and he stated explicitly (Sick 1967:17) that no aggression was involved in the displays. Moreover, it is clear from his descriptions (Sick 1959:286) that the display type that Foster (1981) claimed was aggressive occurs in the absence of younger males, not in their presence. Foster's categorization of such displays as aggressive also appears to be based primarily on the fact that they occur between males, rather than on any inherent differences in the behaviors: as Foster (1981:172; 1984:58) admits, such displays are "extremely similar to" and "strongly reminiscent" of courtship behaviors. That Foster was unable to directly observe courtship displays of the type that Sick reported between males may also be due to geographic or subspecies differences in behaviors: Sick studied a population in Brazil while Foster observed birds in Paraguay. Other elements of the courtship displays between the two populations do appear to differ significantly, such as the vocalizations used and the direction in which males fly during the display (in Brazil, the male farthest from the courted bird begins the courtship "wheel," while in Paraguay the bird closest to the courted bird begins). It should also be pointed out that Snow (1963) independently observed courtship between males in the closely related Blue-backed Manakin.

      60
Vasey "Homosexual Behavior in Primates," p. 197; for a similar observation, see Wolfe, "Human Evolution and the Sexual Behavior of Female Primates," p. 130.

      61
Hyde, H. M. (1970) The Love That Dared Not Speak Its Name: A Candid History of Homosexuality in Britain, p. 1 (Boston: Little, Brown and Company).

      62
Killer Whale (Balcomb et al. 1979:23); published version: Balcomb, K. C., III, J. R. Boran, R. W. Osborne, and N. J. Haenel (1980) "Observations of Killer Whales (Orcinus orca) in Greater Puget Sound, State of Washington," report no. MMC-78/13 to U.S. Marine Mammal Commission, NTIS# PB80-224728. (Washington, D.C.: U.S. Department of Commerce).

      63
Musk-ox (Smith 1976; Tener 1965; Reinhardt 1985); Walrus (Miller 1976); Harbor Seal (Johnson 1974, 1976; Johnson and Johnson 1977).

      64
Halls, L. K., ed., (1984) White-tailed Deer: Ecology and Management (Harrisburg, Pa.: Stackpole Books); Gerlach, D., S. Atwater, and J. Schnell, eds., (1994) Deer (Mechanicsburg, Pa.: Stackpole Books); Jones, M. L., S. L. Swartz, and S. Leatherwood, eds., (1984) The Gray Whale, Eschrictius robustus (Orlando: Academic Press). In contrast, a similarly comprehensive book on Mule Deer does mention homosexual activity (Geist 1981), as does another volume on White-tailed Deer (Rue 1989).

      65
Woodpeckers (Short 1982; Winkler et al. 1995); Skutch, A. F. (1985) Life of the Woodpecker, p. 44 (Santa Monica: Ibis Publishing). For a similar omission of all information on homosexuality in Parrots by the standard "comprehensive" guide to this bird family, see Forshaw (1989).

      66
See, for example, Fay (1982) on Walruses, Birkhead (1991) on Magpies, Lowther and Cink (1992) on House Sparrows, Davis (1993) on Black-crowned Night Herons, Lowther (1993) on Brown-headed Cowbirds, Telfair (1994) on Cattle Egrets, Burger (1996) on Laughing Gulls, Russell (1996) on Anna's Hummingbirds, and Ciaranca et al. (1997) on Mute Swans.

      67
Hooded Warbler (Niven 1993:190); Antbirds (Willis 1967, 1972, 1973); Orange-fronted Parakeet (Buchanan 1966); Golden Plover (Nethersole-Thompson and Nethersole-Thompson 1961, 1986); Mallard Duck (Lebret 1961); Black Swan (Braithwaite 1970, 1981); Scottish Crossbill (Nethersole-Thompson 1975); Black-billed Magpie (Baeyens 1981a); Pied Kingfisher (Moynihan 1990). Similar statements have been made by Konrad Lorenz (1991:241 [Greylag Goose]), who claimed that long-term pair-bonding between males only occurs in Geese and Ducks; and Hunt and Hunt (1977:1467 [Western Gull]), who were unaware of any previous reports of homosexual pairing in wild birds.

      68
Black-headed Gull (van Rhijn and Groothuis 1985:165; Kharitonov and Zubakin 1984); Adelie Penguin (Davis et al. 1998:136); Humboldt Penguin (Scholten 1992:8); Kestrel (Olsen 452). Similar statements have been issued by scientists studying other species — Sylvestre (1985:64), for example, reported not being aware of any previous records of homosexual activity in Botos, even though fairly extensive descriptions were available in Layne and Caldwell (1964), Caldwell et al. (1966), Spotte (1967), and Pilleri et al. (1980). Walther (1990:308) claimed that courtship betweeen male hoofed mammals had not been observed in the wild, when in fact such behavior had been reported in numerous prior studies, including in Pronghorn, Blackbuck, Mountain Sheep (Bighorn, Thinhorn, Asiatic Mouflon), Mountain Goats, Musk-oxen, Bharal, and Markhor (Walther, F. R. [1990] "Bovids: Introduction," in Grzimek's Encyclopedia of Mammals, vol. 5, pp. 290-324 [New York: McGraw-Hill]).

      69
See, for example, Takahata et al. (1996:149), who ask, "Is GG-rubbing a sexual behavior?" and conclude that its "nonsexual" aspects are more prominent, because of its association with tension reduction, feeding, reassurance, participation by nonestrous females, and the fact that Bonobos (unlike Japanese Macaques) do not form "exclusive homosexual female-female pairs." None of these characteristics, in fact, negate a fully "sexual" interpretation. In particular, the fact that Bonobos do not form same-sex pairs or consortships hardly argues against the sexual nature of their genital rubbing — it simply indicates that homosexual interactions in this species do not involve extensive pair-bonding. By these criteria, Bonobo heterosexual interactions would have to be considered nonsexual as well, since they are often associated with the same "social" or "nonsexual" situations, nor do individuals form "exclusive heterosexual male-female pairs." See also Kuroda (1980:190), who considers genital rubbing between females to be "uninterpretable" when it occurs in contexts that are not related to tension reduction or food exchange; and Kano (1980:253-54, 1992:139,1990:66-67, 69), who classifies same-sex activities in Bonobos as primarily "social" rather than "sexual" and ascribes to them the primary "functions" of greeting, reassurance, reconciliation, and food-sharing (while nevertheless recognizing that sexual aspects may be secondarily involved in some cases). As recently as 1997, researchers were still speculating about, and emphasizing, the nonsexual "functions" of Bonobo homosexual activity (Hohmann and Fruth 1997).
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Mountain Sheep (Geist 1975:97-98).

      71
Vasey, P. L. (1997, August 19) "Summary: Homosexual or Dominance Behavior? (Discussion)," message posted to Primate Talk (on-line discussion list).

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Rhesus Macaque (Hamilton 1914). A standard and widely cited exposition of the dominance interpretation is Wickler, W. (1967) "Socio-sexual Signals and Their Intra-specific Imitation Among Primates," in D. Morris, ed., Primate Ethology, pp. 69-147 (London: Weidenfield and Nicolson).

      73
On the occurrence of dominance hierarchies in various mammals and birds without homosexuality, and further references, see Wilson, E. O. (1975) Sociobiology: The New Synthesis, p. 283 (Cambridge and London: Belknap Press); Welty, J. C., and L. Baptista (1988) The Life of Birds, 4th ed., pp. 206-210 (New York: W. B. Saunders).

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For explicit statements on the absence, unimportance, or irrelevance of dominance hierarchies in these species or populations, see Gorilla (Yamagiwa 1987a:25; Robbins 1996:957); Savanna (Olive) Baboon (Rowell 1967b:507-8); Bottlenose Dolphin (Shane et al. 1986:42); Zebras (Penzhorn 1984:113; Schilder 1988:300); Musk-ox (Smith 1976:92-93); Koala (Smith 1980:187); Buff-breasted Sandpiper (Lanctot and Laredo 1992:7); Tree Swallow (Lombardo et al. 1994:556). In Gorilla all-male groups, dominance is not a central organizing feature of social interactions (including homosexual interactions) even though some semblance of a dominance "hierarchy" may exist and males clearly have different ranks. The same may also be true for Hanuman Langur all-male groups (Weber and Vogel 1970:75) and Collared Peccary mixed-sex groups (Sowls 1997:151-53) in which same-sex interactions occur. In Buff-breasted Sandpipers, although mounting between males may be accompanied by aggression and therefore superficially appears related to "dominance," there is in fact no evidence that a dominance hierarchy exists in this species or constitutes an important aspect of its social organization. In some of these species (e.g., Zebras, Musk-oxen, Bottlenose Dolphins) dominance hierarchies are more prominent in captivity, although homosexual activity occurs in both wild and captive contexts. Finally, J. Steenberg (personal communication) suggests that mounting between female Matschie's Tree Kangaroos is a dominance display, yet Hutchins et al. (1991:154-56, 161) found no clear-cut dominance hierarchy in the study population where this behavior was observed.

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Tasmanian Native Hen (Ridpath 1972:81) (in this species, hierarchies can be induced in wild birds by provisioning them with food, but dominance plays no role in their unprovisioned activities — including homosexual mounting, which is not associated in any way with induced dominance); Little Blue Heron (Werschkul 1982:383-84); white-browed sparrow weaver and other weavers (Collias, N. E., and E. C. Collias [1978] "Cooperative Breeding Behavior in the White-browed Sparrow Weaver," Auk 95:472-84; Collias, N. E., and E. C. Collias [1978] "Group Territory, Dominance Hierarchy, Co-operative Breeding in Birds, and a New Factor," Animal Behavior 26:308-9). Likewise, dominance systems occur in most Macaques, yet homosexual behavior is apparently absent in some species, e.g., the Barbary Macaque (Macaca sylvanus) — see Vasey, "Homosexual Behavior in Primates," pp. 178-79; for an extensive summary of research on this species with no mention of same-sex mounting, see Fa, J. E., ed. (1984) The Barbary Macaque: A Case Study in Conservation (New York: Plenum Press). However, recent work seems to suggest that same-sex mounting may in fact occur: Di Trani, C. M. P. (1998) "Conflict Causes and Resolution in Semi-Free-Ranging Barbary Macaques (Macaca sylvanus )," Folia Primatologica 69:47-48. Therefore, this example must be interpreted with caution, like many other instances involving an apparent "absence" of homosexual behavior (see chapter 4 for further discussion).

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Wolf (Zimen 1976, 1981); Spotted Hyena (Frank 1986); Squirrel Monkey (Baldwin and Baldwin 1981:294-95; Castell and Heinrich 1971:187-88); Bottlenose Dolphin (Samuels and Gifford 1997:82, 88-90). In Red Squirrels, both sexes have dominance systems yet same-sex mounting is much more prominent among males (Ferron 1980:135-36); in Bonobos, a dominance system is much more developed or important among males (de Waal 1997:72-74), yet homosexual activities occur in both sexes. A related observation is that in Bighorn Sheep, both sexes have well-defined dominance systems and exhibit same-sex mounting, yet only among males does it have some correlation with homosexual activity.

      77
For examples of animals that participate in interspecies homosexual mounting, see the profiles for Crabeating Macaque, Bottlenose Dolphin, Walrus, Greenshank, Orange Bishop Bird, and House Sparrow. On the occurrence of interspecies dominance hierarchies, see, for example, Fisler, G. F. (1977) "Interspecific Hierarchy at an Artificial Food Source," Animal Behavior 25:240-44; Morse, D. H. (1974) "Niche Breadth as a Function of Social Dominance," American Naturalist 108:818-30.

      78
Rhesus Macaque (Reinhardt et al. 1986:56); Japanese Macaque (Chapais and Mignault 1991:175-76; Vasey et al. 1998); Common Chimpanzee (Nishida and Hosaka 1996:122 [table 9.7]). See also Bygott 1974 — cited in Hanby 1974:845 [Japanese Macaque] — who found that 59 percent of mounts between male Chimps were by subordinates on dominants or by equally ranked participants.

      79
Musk-ox (Reinhardt 1985:298). In Cattle Egrets, Fujioka and Yamagishi (1981:139) stated that males attempting homosexual copulations always rank higher than or equal to the males they mount. Yet two males in their study population who mounted other males were apparently not part of the dominance hierarchy (cf. their table 3), while the highest-ranking male did not participate in any same-sex mounts. M. Fujioka (personal communication) concedes that the rank of the males may not actually be an important factor in their homosexual mounting.
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Crested Black Macaque (Dixson 1977:77; Poirier 1964:96); American Bison (Reinhardt 1985:218, 222, 1987:8); Pig-tailed Macaque (Oi 1990a:350); Red Deer (Hall 1983:278); Pukeko (Jamieson and Craig 1987b:319-22); Japanese Macaque (Chapais and Mignault 1991:175-76); Bighorn Sheep (Shackleton 1991:179-80).

      81
A further argument is provided by "pile-up mounts," i.e., when three individuals are all mounted (stacked) on each other. In this case, the mounter-mountee relations rarely if ever follow dominance lines: either they occur in species without dominance hierarchies (e.g., Sage Grouse, Common Murre), or else it is not the case that the middle animal is both higher-ranking than the animal it is mounting but lower-ranking than the animal who is mounting it (e.g., Wolf, Bonobo). For more on pile-up mounts, see chapter 4.

      82
Common Chimpanzee (Nishida and Hosaka 1996:122; Bygott 1974 [cited in Hanby 1974:845 (Japanese Macaque)]); White-faced Capuchin (Manson et al. 1997:771, 780); Blackbuck (Dubost and Feer 1981:89 — 90); Cavies (Rood 1972:36); Gray-capped Social Weaver (Collias and Collias 1980:218, 220). Although mounting between male Musk-oxen in captivity seems to follow dominance lines (Reinhardt 1985), in wild herds Smith (1976) found no dominance hierarchy within (as opposed to between) sex/age classes. Same-sex mounting in the wild occurs among age-mates (who are therefore essentially equal in rank, e.g., two-year-old males mount each other).

      83
See, for example, Bertrand 1969:191 (Stumptail Macaque); Simonds 1965:183, Sugiyama 1971:259 (Bonnet Macaque); Bernstein 1972:406 (Pig-tailed Macaque); Dixson et al. 1975:195-96 (Talapoin Monkey); Kaufmann 1974:309 (Whiptail Wallaby).

      84
A distinction between consensual and nonconsensual homosexual mounts is found in more than 30 different species of mammals and birds. Direct evidence of sexual arousal and stimulation on the part of animals being mounted is also available in many species, including orgasmic (and other) responses in female Japanese, Rhesus, Stumptail, and Pig-tailed Macaques being mounted; erection and masturbation by male mountee Rhesus, Pig-tailed, and Crested Black Macaques; thrusting by male Bonobos being mounted; and stimulation of the mountee's clitoris by her partner's thrusting in Hanuman Langurs and Japanese Macaques. In addition to direct and indirect genital stimulation during mounting, it is quite likely that male animals being penetrated during anal intercourse also experience stimulation of the prostate gland (which presses against the wall of the rectum). In human males, direct stimulation of the prostate — for instance, during anal intercourse — can be highly arousing and may precipitate or enhance orgasm. A similar capacity is probably present in all male mammals. Although direct evidence (in the form of firsthand accounts) of the pleasurable or arousing nature of this activity is, of course, lacking in nonhuman animals, there is some indirect evidence. A standard technique of inducing erection and ejaculation (for purposes of artificial insemination) in male mammals is through anal and/or prostate stimulation. Known as electroejaculation, this technique involves insertion of an anal probe and stimulation of the rectum — especially in the area of the prostate gland — with a mild electrical current as well as back-and-forth (thrusting) movements of the probe. This technique has proven effective in numerous species of mammals, including virtually all of those in which male homosexual mounting and /or anal penetration occur. For further information on electroejaculation, see Watson, P. F., ed. (1978) Artificial Breeding of Non-Domestic Animals, Symposia of the Zoological Society of London no. 43, especially pp. 109, 129, 208-10, 221, 295 (London: Academic Press).

      85
Hanuman Langur (Srivastava et al. 1991:506-7); for a similar assessment with regard to homosexual activity between males in this species, see Weber and Vogel (1970:77-78). See also Rowell (1967a:23), who states that "sexual" and "dominance" mounts in Savanna (Yellow) Baboons are virtually indistinguishable, and Enomoto (1990:473), who remarks on the difficulty of discriminating between sexual and ritualized dominance mounting in Bonobos because of the gradation between the two. Weinrich (Sexual Landscapes, p. 294), in discussing mounting between male Mountain Sheep, also points out how sexuality and dominance can both be part of the same behavior and suggests an analogy with human sexuality. Indeed, elements of consensual "dominance" or power-play, although rarely acknowledged, are often a part of human lovemaking and sexual pleasure, ranging along a continuum from gentle "love bites" to full sadomasochism (and nonconsensual dominance also figures prominently in many human sexual interactions, especially heterosexual ones).

      86
Japanese Macaque (Wolfe 1986:268); Rhesus Macaque (Akers and Conaway 1979:78); Greylag Goose (Lorenz 1991:206); Black-winged Stilt (Kitagawa 1989:65, 69) (see also the distinction between same-sex courtship and aggressive/appeasing kantling in Ostriches [Sauer 1972:731; Bertram 1992:15, 50-51]). For species such as these that have a clear distinction between mounts in sexual and nonsexual contexts, only the former are considered (in this book and in most sources) to be homosexual behavior. As noted in chapter 1, some species classified by Dagg (1984) as exhibiting homosexuality (e.g., bush squirrels and degus) are excluded from our roster on the basis of this criterion, because all same-sex mounting in these species appears to fall into this genuinely nonsexual category; see Viljoen, S. (1977) "Behavior of the Bush Squirrel, Paraxerus cepapi cepapi," Mammalia 41:119-66; Fulk, G. W. (1976) "Notes on the Activity, Reproduction, and Social Behavior of Octodon degus," Journal of Mammology 57:495-505.

      87
Walrus (Miller 1975:607); Gray Seal (Anderson and Fedak 1985); Oystercatcher (Ens, B. J., and J. D. Goss-Custard [1986] "Piping as a Display of Dominance by Wintering Oystercatchers Haematopus ostralegus," Ibis 128:382-91). Early observers of this species (e.g., Makkink 1942) misinterpreted the piping display as a courtship activity because it often occurs between males and females.
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      88
For details of the way that dominance is expressed in these species, see Savanna (Yellow) Baboon (Maxim and Buettner-Janusch 1963:169); Hamadryas Baboon (Stammbach, E. [1978] "On Social Differentiation in Groups of Captive Female Hamadryas Baboons," Behavior 67:322-38); Bottlenose Dolphin (Samuels and Gifford 1997); Killer Whale (Rose 1992:108-9); Caribou (Espmark, J. [1964] "Studies in Dominance-Subordination Relationship in a Group of Semi-Domestic Reindeer (Rangifer tarandus L.)," Animal Behavior 12:420-26); Blackbuck (Dubost and Feer 1981:97-100); Wolf (Zimen 1976, 1981); Bush Dog (Macdonald 1996); Spotted Hyena (Frank 1986:1511); Grizzly Bear (Craighead et al. 1995:109ff); Black Bear (Stonorov and Stokes 1972:235, 242); Red-necked Wallaby (Johnson 1989:267); Canada Goose (Collias and Jahn 1959:500-501); Scottish Crossbill (Nethersole-Thompson 1975:53); Black-billed Magpie (Birkhead 1991); Jackdaw (Roell 1978); Acorn Woodpecker (Stanback 1994); Galah (Rowley 1990:57). In Pronghorns, mounting between males was originally claimed to represent a dominance activity (Kitchen 1974), yet more recent studies of dominance in this species have not included same-sex mounting (Bromley 1991).

      89
In some cases, sexual behaviors other than mounting can be correlated with dominance. For example, grooming between males in Nilgiri Langurs and Crested Black Macaques is often performed by a subordinate animal on a more dominant one. Nevertheless, it is apparent that this activity has a clearly sexual component as well: one or both males may become intensely aroused, developing an erection and even ejaculating during the grooming (see Poirier 1970a:334 for Nilgiri Langurs and Poirier 1964:146-47 for Crested Black Macaques). Similarly, adult (dominant) Bonobos often masturbate or massage the genitals of adolescent (subordinate) males, but again, the activity involves clear sexual stimulation (cf. de Waal 1987, 1995, 1997). Also, Squirrel Monkey genital displays are sometimes correlated with dominance, but there are also cases where the association is less than definitive, or where they occur in clearly sexual contexts between animals of the same sex (cf. Talmage-Riggs and Anschel 1973:70; Travis and Holmes 1974:55; Baldwin and Baldwin 1981:295-97; Castell and Heinrich 1971:187-88).

      90
One cannot help but surmise that it is the heterosexism of many biologists that has led them to focus on mounting behavior to the exclusion of other activities in their appeal to dominance factors — for only in mounting can the positions of the participants be clearly analogized to those of a male and female in a heterosexual interaction. As Fedigan (1982:101 [Japanese Macaque]) points out, underlying the entire discussion of dominance in same-sex interactions is the assumption that homosexual mounting is essentially a transposition from heterosexual copulation — and that males "dominate" females in such interactions. For further evidence against this view, see the discussion of homosexuality as a form of "pseudoheterosexuality" in chapter 4.

      91
Possible exceptions are same-sex courtship interactions in Mountain Sheep (Geist 1968, 1971), Musk-oxen (Reinhardt 1985), and Cavies (Rood 1972), which have been interpreted as reflecting dominance. Additionally, mounting or other sexual behaviors within a same-sex pair-bond — common in many bird species — does not fit easily into a dominance interpretation, since this usually involves ongoing interaction with only one other animal (rather than the establishment of hierarchical positions within a network of individuals).

      92
Giraffe (Pratt and Anderson 1985:774-75, 780-81); Crested Black Macaque (Dixson 1977:77-78; Reed et al. 1997:255); Stumptail Macaque (Bernstein 1980:40); Pig-tailed Macaque (Giacoma and Messeri 1992:187); Savanna (Olive) Baboon (Owens 1976:250-51); Squirrel Monkey (Baldwin and Baldwin 1981:295-97; Baldwin 1968:296, 311); Red Squirrel (Ferron 1980:136); Spinifex Hopping Mouse (Happold 1976:147); American Bison (Reinhardt 1985:222-23); Pukeko (Lambert et al. 1994); Sociable Weaver (Collias and Collias 1980:246, 248; in the latter instance, the inconsistency in dominance status was not one of the cases of temporary reversals of dominance that were occasionally seen in this species). In female Squirrel Monkeys, dominance hierarchies are not considered to be a salient feature of social organization in the wild (Baldwin and Baldwin 1981:294-95). However, even when dominance systems appear to develop (e.g., in some captive situations), investigators have found that the rank of females based on their homosexual activities does not agree with other measures of rank (Anschel and Talmage-Riggs 1978:602 [table 1]).

      93
For some reevaluation and/or critiques of the concept of dominance, see Gartlan, J. S. (1968) "Structure and Function in Primate Society," Folia Primatologica 8:89-120; Bernstein 1970 (Crab-eating Macaque); Richards, S. M. (1974) "The Concept of Dominance and Methods of Assessment," Animal Behavior 22:914 — 30; Ralls, K. (1976) "Mammals in Which Females Are Larger Than Males," Quarterly Review of Biology 51:245-76; Lockwood, R. (1979) "Dominance in Wolves: Useful Construct or Bad Habit?" in E. Klingham-mer, ed., Behavior and Ecology of Wolves, pp. 225-44 (New York: Garland); Baldwin and Baldwin 1981 (Squirrel Monkey); Bernstein, I. S. (1981) "Dominance: The Baby and the Bathwater," Behavioral and Brain Sciences 4:419-57; Hand, J. L. (1986) "Resolution of Social Conflicts: Dominance, Egalitarianism, Spheres of Dominance, and Game Theory," Quarterly Review of Biology 61:201-20; Walters, J. R., and R. M. Seyfarth (1987) "Conflict and Cooperation," in B. B. Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wrangham, and T. T. Struhsaker, eds., Primate Societies, pp. 306-17 (Chicago and London: University of Chicago Press); Drews, C. (1993) "The Concept and Definition of Dominance in Animal Behavior," Behavior 125:283-313; Lambert et al. 1994 (Pukeko).

      94
Fedigan 1982:92-93 (Japanese Macaque).

      95
Bonobo (Kano 1992:253-54; Kitamura 1989:57, 63); Gorilla (Harcourt et al. 1981:276; Yamagiwa 1987a:25; Harcourt 1988:59); Hanuman Langur (J. J. Moore, in Weinrich 1980:292); Japanese Macaque (Vasey 1996:549; Chapais and Mignault 1991:175-76; Tartabini 1978:433, 435; Hanby 1974:841); Rhesus Macaque (Akers and Conaway 1979:78; Reinhardt et al. 1986:55; Gordon and Bernstein 1973:224); Pig-tailed Macaque (Tokuda et al. 1968:293); Crested Black Macaque (Dixson 1977:77-78; Poirier 1964:20, 49; Reed et al. 1997:255); Savanna Baboon (Owens 1976:256); Gelada Baboon (Mori 1979:134-35; R.Wrangham, in Weinrich 1980:291); Squirrel Monkey (Talmage-Riggs and Anschel 1973:70); Bottlenose Dolphin (Caldwell and Caldwell 1972:427); Blackbuck (Dubost and Feer 1981:89-90); Giraffe (Pratt and Anderson 1985:774 — 75, 780); American Bison (Reinhardt 1985:222, 1987:8); Red Squirrel (Ferron 1980:136); Little Blue Heron (Werschkul 1982:383-84); Tree Swallow (Lombardo et al. 1994:556).
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      96
For examples of earlier claims of a dominance connection being refuted by later studies, see Common Chimpanzee (Yerkes 1939:126-27; Nishida 1970:57 — Bygott 1974 [cited in Hanby 1974:845 (Japanese Macaque)]; Nishida and Hosaka 1996:122 [table 9.7]); Hanuman Langur (Weber 1973:484 — Srivastava et al. 1991:506 — 7; J. J. Moore, in Weinrich 1980:292); Rhesus Macaque (Carpenter 1942 — Akers and Conaway 1979:78; Reinhardt et al. 1986; Gordon and Bernstein 1973:224); Japanese Macaque (Sugiyama 1960:136 — Hanby 1974:841; Chapais and Mignault 1991:175-76); Bonnet Macaque (Rahaman and Parthasarathy 1968:68, 263 — Makwana 1980:10); Pig-tailed Macaque (Tokuda et al. 1968 — Oi 1990a:353-54); Killer Whale (Balcomb et al. 1979:23 — Rose 1992:108-9); Giraffe (Dagg and Foster 1976, Leuthold 1979:27-29 — Pratt and Anderson 1985:774-75); Blackbuck (Schaller 1967 — Dubost and Feer 1981:89-90); American Bison (Lott 1974:391 — Reinhardt 1986:222-23); Wolf (Schenkel 1947 — van Hooff and Wensing 1987:232). In addition, a parallel example in Laughing Gulls involves an indirect refutation of the relevance of dominance. Noble and Wurm (1943:205-6) linked homosexual mounting in Laughing Gulls to the supposedly lower rank of the male being mounted, citing as evidence of his lower status the fact that the mounted male did not "dominate" his female mate. In a more recent detailed study of interactions between partners in heterosexual pairs, however, Hand (1985) concluded that males do not in general dominate their female mates in this species — thus invalidating the earlier claim that being mounted homosexually was correlated with "lower status." Studies that attribute homosexual activity to dominance with little or no supporting evidence include Orang-utan (Rijksen 1978:257); Squirrel Monkey (DuMond 1968:124); West Indian Manatee (Rathbun et al. 1995:150); Pied Kingfisher (Moynihan 1990:19).

      97
Whiptail Wallaby (Kaufmann 1974:307, 309); Rhesus Macaque (Gordon and Bernstein 1973:224). Kaufmann concluded that Whiptail Wallaby homosexual mountings are themselves probably not dominance-related, however, because dominant animals generally invite subordinate ones to mount (the opposite of the "usual" dominance pattern).

      98
Bighorn Sheep (Hogg 1987:120; Hass and Jenni 1991:471); Crested Black Macaque (Poirier 1964:54).

      99
Vasey, "Homosexual Behavior in Primates," p. 191.

      100
Orang-utan (Maple 1980:118).

      101
West Indian Manatee (Rathbun et al. 1995:150). See also the suggestion in Buss (1990:19-21) that sexual arousal in male African Elephants during same-sex play-fighting serves to dull pain. While this is possible, it is rather far-fetched, considering that such ritual fights (described by Buss as "erotic") are rarely violent.

      102
Vasey, P. L. (in press) "Homosexual Behavior in Male Birds," in W. R. Dynes, ed., Encyclopedia of Homosexuality, 2nd ed., vol. 1: Male Homosexuality (New York: Garland Press).

      103
American Bison (Reinhardt 1985:222) (cf. also Kaufmann [1974:107] on Whiptail Wallabies, who asserts, "Though tail-lashing seems clearly a sign of sexual arousal, it was occasionally performed by males when they were approached by subordinate males in nonsexual situations"); Asiatic Mouflon (McClelland 1991:80); Stumptail Macaque (O'Keefe and Lifshitz 1985:149); Dugong (Nair et al. 1975:14); Laysan Albatross (Frings and Frings 1961:311); Dwarf Mongoose (Rasa 1979a:365); Bonnet Macaque (Nolte 1955:179). Similarly, Frank et al. (1990:308) state that genital erections in Spotted Hyenas have no "sexual significance" unless displayed by a male toward a female during courtship. The "desexing" of this behavior stems, in large part, from the fact that erections are frequently displayed between animals of the same sex (especially females) and in situations that do not involve (heterosexual) mounting (e.g., during the "meeting ceremony"). While erections undoubtedly have "nonsexual" connotations outside of a mounting context (see, for example, East et al. 1993), it seems overly restrictive to eliminate all "sexual significance" from situations that do not fall into the category of heterosexual courtship and mating.

      104
Redshank (Hale and Ashcroft 1983:21). For a summary of the historical interpretation of this behavior, see also Cramp and Simmons 1983:533.

      105
Crested Black Macaque (Dixson 1977:71, 76; Poirier 1964:147). Dixson (1977:77) does concede that the distinction between sexual and nonsexual mounts and solicitations is a subjective one, but only in heterosexual contexts — homosexual interactions are assumed to be self-evidently nonsexual.

      106
Vicuna (Koford 1957:183, 184); Musk-ox (Smith 1976:51); Giraffe (Dagg and Foster 1976:127; Pratt and Anderson 1985:777-78; Leuthold 1979:27, 29); Bank Swallow (Beecher and Beecher 1979:1284); Savanna Baboon (Smuts 1985:18, 148-49, 163-66, 199, 213); Rhesus Macaque (Loy 1971:26); Oystercatcher (Makkink 1942; Ens and Goss-Custard, "Piping as a Display of Dominance").

      107
Crested Black Macaque (Dixson 1977:70-71); Bottlenose Dolphin (Ostman 1991:313; Dudok van Heel and Mettivier 1974:12; Saayman and Tayler 1973); Spinner Dolphin (Norris and Dohl 1980a:845; Norris et al. 1994:199); Common Murre (Birkhead 1978a:326); Blue-bellied Roller (Moynihan 1990).

      108
Rhesus Macaque (see, for example, Sade 1968:32-33); Japanese Macaque (Hanby 1974:843, 845; Wolfe, "Human Evolution and the Sexual Behavior of Female Primates," p. 129).

      109
For further discussion see chapter 5. On a related point, aggressive behaviors may accompany homosexual interactions in some species and are therefore used to argue that such behavior is not "really" sexual. However, aggression is also characteristic of heterosexual relations in many species, where such male-female interactions are still classified as "sexual."
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      110
Kob (Buechner and Schloeth 1965:218); Giraffe (Pratt and Anderson 1985:774-75); northern jacana (del Hoyo, J., A. Elliott, and J. Sargatal, eds. [1996] Handbook of the Birds of the World, vol. 3: Hoatzin to Auks, p. 282. [Barcelona: Lynx Edicions]); Orang-utan (Galdikas 1981:286).

      111
Walrus (Dittrich 1987:168); Musk-ox (Smith 1976:62); Bighorn Sheep (Hogg 1984:527; Geist 1971:139); Asiatic Mouflon (McClelland 1991:81); Grizzly Bear (Craighead et al. 1995:161); Olympic Marmot (Barash 1973:212); White-tailed Deer (Hirth 1977:43); Orang-utan (Galdikas 1981:286); White-faced Capuchin (Manson et al. 1997:775); Northern Fur Seal (Gentry 1998:172); Ruff (Hogan-Warburg 1966:167-68). Additionally, in one study of Matschie's Tree Kangaroos — a species in which researchers deny that mounting between females is (homo)sexual (J. Steenberg, personal communication) — all mounts observed between animals of the opposite sex were "incomplete" in that they did not involve penetration or thrusting (Hutchins et al. 1991:158). Another study of the same population found both that "full" copulations between males and females were infrequent, and that in heterosexual contexts females showed few overt signs of sexual interest, since the behavioral cues for female sexual arousal are extremely subtle (Dabek 1994:84, 93-94, 116).

      112
Morrill and Robertson 1990 (Tree Swallow); Scott, M. P., and T. N. Tan (1985) "A Radiotracer Technique for the Determination of Male Mating Success in Natural Populations," Behavioral Ecology and Sociobiology 17:29-33. More recently, a copulation-verification technique using fluorescent powder has been tested for rodents. Dusted on males, the powder is transferred to females during mating and can be checked using ultraviolet light. Ironically, during the testing of this procedure, pairs of females were used as "controls" since it was assumed that they would not engage in mounting behavior with one another. Nevertheless, 12 percent of female pairs showed transfer of powder — but of course this was interpreted by researchers as evidence of nonsexual contact between such females (Ebensperger, L. A., and R. H. Tamarin [1997] "Use of Fluorescent Powder to Infer Mating Activity of Male Rodents," Journal of Mammalogy 78:888-93).

      113
Rhesus Macaque (Erwin and Maple 1976); field report of penetration and ejaculation (Sade 1968:27); see also Kempf (1917:134) for an even earlier documentation of anal penetration between (captive) male Rhesus Macaques. Walther (1990:308) makes a parallel claim that mounting activity between male hoofed mammals does not constitute (homo)sexual behavior because erection and anal penetration are not always observed (Walther, "Bovids: Introduction"). On a related point, Tuttle (1986:289) takes great pains to point out that rump-rubbing and mounting between male Bonobos do not "qualify" as genital contact because, "pace certain sodomites, the anus is not a genital organ (International Anatomical Nomenclature Commitee, 1977, p. A49)." Tuttle does, however, accept that sexual activity between females — which he calls "bizarre homosexual hunching" (ibid., p. 282) — qualifies as genital contact (Tuttle, R. H. [1986] Apes of the World: Their Social Behavior, Communication, Mentality, and Ecology [Park Ridge, N.J.: Noyes Publications]). For a recent survey of homosexual behavior in primates that (wisely) drops the occurrence of penetration, arousal, and/or orgasm as a defining criterion of the behavior, see Vasey, "Homosexual Behavior in Primates," p. 175.

      114
On a similar gradation of mounting behavior in male birds, see Moynihan 1955:105 (Black-headed Gull).

      115
For specific arguments against homosexual activity as a form of tension reduction in various species, see Yamagiwa 1987a:23, 1987b:37 (Gorilla); Edwards and Todd 1991:234-35 (White-handed Gibbon); Vasey 1996:549-50 (Japanese Macaque); R. Wrangham, in Weinrich 1980:291 (Gelada Baboon). Against homosexuality as a form of play, see Talmage-Riggs and Anschel 1973:71 (Squirrel Monkey); Lombardo et al. 1994:556 (Tree Swallow). Against homosexuality as reconciliation or reassurance behavior, see Vasey 1996:550 (Japanese Macaque); Akers and Conaway 1979:78 (Rhesus Macaque); Lombardo et al. 1994:556 (Tree Swallow). Against homosexual activities as a means of forging coalitions or alliances, see Silk 1994:285-87 (Bonnet Macaque) (and also Silk 1993:187 for arguments that coalition-bonding between males in this species is not "functional" in terms of enhancing the males' status, access to resources, or inclusive fitness). Against homosexuality as a gesture of appeasement or placation, see Manson et al. 1997:783 (White-faced Capuchin); Ferron 1980:136 (Red Squirrel); Lombardo et al. 1994:556 (Tree Swallow). Against homosexual relations as "kinship alliances" between individuals who associate with each other primarily because they are related (so-called kin selection), see Fernandez and Reboreda 1995:323 (Greater Rhea); Heg and van Treuren 1998:688-89, Ens 1998:635 (Oystercatcher); Afton 1993:232 (Lesser Scaup Duck); Rose 1992:104, 112 (Killer Whale); Hashimoto et al. 1996:316 (Bonobo). See also Vasey, "Homosexual Behavior in Primates," for a summary and review of the evidence against many of these nonsexual "explanations."

      116
Japanese Macaque (Vasey 1996).

      117
Bonobo (de Waal 1987, 1995 [among others]; Savage-Rumbaugh and Lewin 1994:110); Gorilla (Yamagiwa 1987a:23, 1987b:37).

      118
See Silk 1994:285-87 (Bonnet Macaque) for more detailed discussion.

      119
Signs of sexual arousal such as these have been documented for homosexual interactions in more than 90 species of mammals and birds. In addition, a number of scientists have themselves asserted the clearly sexual character of same-sex interactions (in addition to, or instead of, nonsexual aspects); see, for example, de Waal 1995:45-46 (Bonobo); Yamagiwa 1987a, Harcourt 1988:59, Porton and White 1996:724 (Gorilla); Edwards and Todd 1991 (White-handed Gibbon); Weber and Vogel 1970:76-77 (Hanuman Langur); Vasey 1996:550, Rendall and Taylor 1991:324, Wolfe 1984:147 (Japanese Macaque); Akers and Conaway 1979:78-79 (Rhesus Macaque); Chevalier-Skolnikoff 1976:525 (Stumptail Macaque); Srivastava et al. 1991 (Hanuman Langur); R. Wrangham, in Weinrich 1980:291 (Gelada Baboon); Manson et al. 1997:775-76 (White-faced Capuchin); Herzing and Johnson 1997:85, 90 (Bottlenose/Atlantic Spotted Dolphins); Saulitis 1993:58 (Killer Whale); Darling 1978:60, 1977:10-11 (Gray Whale); Coe 1967:320 (Giraffe); Rue 1989:313 (White-tailed Deer); Buss 1990:20 (African Elephant); Heg and van Treuren 1998:688 (Oystercatcher); Davis et al. 1998 (Adelie Penguin); Stiles 1982:216 (Anna's Hummingbird). For use of the word erotic to characterize same-sex interactions, see, for example, de Waal 1987:323, 1997:103-4, Kano 1992:192, 1990:66 (Bonobo); Darling 1977:10-11 (Gray Whale); Mathews 1983:72 (Walrus); Buss 1990:19 (African Elephant).

      120
Occasionally, however, multiple "functions" are granted to heterosexual behavior; see, for example, Lindburg 1971 (Rhesus Macaque); de Waal 1987, 1995, 1997, Kano 1990:67 (Bonobo); Manson et al. 1997 (White-faced Capuchin); Hanby, J. (1976) "Sociosexual Development in Primates," in P. P. G. Bateson and P. H. Klopfer, eds., Perspectives in Ethology, vol. 2, pp. 1-67 (New York: Plenum Press).
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