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{83} Chapter 3
Two Hundred Years of Looking at Homosexual Wildlife
     1764: ... three or four of the young [Bantam] cocks remaining where they could have no communication with hens ... each endeavoured to tread his fellow, though none of them seemed willing to be trodden. Reflection on this odd circumstance hinted to me, why the natural appetites, in some of our own species, are diverted into wrong channels.
     — GEORGE EDWARDS, Gleanings of Natural History

     1964: Another example of an irreversible sexual abnormality concerns an orang-utan. This ape, a young male, was kept with another young male and they spent a great deal of time playing together. This included some sex play and anal intercourse was observed on a number of occasions.
     — DESMOND MORRIS, "The Response of Animals to a Restricted Environment"

     1994: There are several explanations for homosexual behavior in non-human animals. First, it is possible that the pursuers misidentified male 42 as a female because the plumage of after-second-year female Tree Swallows resembles that of males ...
     — MICHAEL LOMBARDO et al., "Homosexual Copulations by Male Tree Swallows"¹
{84}
Animal homosexuality is by no means a "new" discovery by modern science. Some of the earliest statements regarding homosexual behavior in animals date back to ancient Greece, while the first detailed scientific studies of same-sex behavior were made in the 1700s and 1800s. From the very beginning, descriptions of homosexuality in animals were accompanied by attempts to interpret or explain its occurrence, and observers who witnessed the behavior were almost invariably puzzled, astonished, and even upset by the simple fact of its existence. As the quotes above illustrate, many of these same attitudes have continued to this day. With more than 200 years of scientific attention devoted to the subject, how is it that so many people today — many scientists included — are unaware of the full extent and characteristics of animal homosexuality, and/or continue to be puzzled by its occurrence? This chapter seeks to answer this question, first by chronicling the history of the study of homosexuality in animals, and then by documenting the systematic omissions and negative attitudes of many zoologists in dealing with this phenomenon. As we will see, a history of the scientific study of animal homosexuality is necessarily also a history of human attitudes toward homosexuality.

A Brief History of the Study of Animal Homosexuality

The history of animal homosexuality in Western scientific thought begins with the early speculations of Aristotle and the Egyptian scholar Horapollo on "hermaphroditism" in hyenas, homosexuality in partridges, and variant genders and sexualities in several other species.² Although much of their thinking was infused with mythology and anthropomorphism, and there are notable inaccuracies in their observations (the Spotted Hyena, for example, is not hermaphroditic), the discussions of these scholars represent the first recorded thoughts on homosexuality and transgender in animals. The earliest scientific observations of animal homosexuality are those of the noted French naturalist (and count) Georges-Louis Leclerc de Buffon, whose monumental fifteen-volume Histoire naturelle generale et particuliere (1749-67) includes observations of same-sex behavior in birds. Additional observations on homosexuality in birds were made in the eighteenth century by the British biologist George Edwards, and (as indicated above) they also include some of the first pronouncements about the supposed "causes" and "abnormality" of such behavior.³

The beginning of the modern study of animal homosexuality was heralded by a number of early descriptions of same-sex behavior in insects (e.g., by Alexandre Laboulmene in 1859 and Henri Gadeau de Kerville in 1896), small mammals (e.g., by R. Rollinat and E. Trouessart on Bats in 1895), and birds (e.g., by J. Whitaker on Swans in 1885 and Edward Selous on Ruffs in 1906), while the German scientist Ferdinand Karsch offered, in the year 1900, one of the first general surveys of the phenomenon. ⁴ Since then, the scientific study of animal homosexuality has expanded enormously to include a wide variety of investigations, reported in close to 600 scientific articles, monographs, dissertations, technical reports, and other publications {85} in over ten different languages. These range from field observations of animals that only anecdotally mention homosexual behavior, to more extensive descriptions of homosexuality in a wide range of species studied in the wild, to observations of captive animals (including at many zoos and aquariums throughout the world), to experiments on laboratory animals, to more recent studies devoted to examining all aspects of homosexual behavior in a particular species (often in the wild), to more comprehensive general surveys of the phenomenon. Some reports have received wide attention, such as the discovery of female pairing in various Gull and Tern species that initiated a flurry of scientific and media interest in the late seventies and early eighties. On the other hand, many reports of animal homosexuality have gone unnoticed even by other zoologists, languishing in small specialty or regional journals such as The Bombay Journal of Natural History, Ornis Fennica (the journal of the Finnish Ornithological Society), Revista Brasileira de Entomologia (the Brazilian Journal of Entomology), or the Newsletter of the Papua New Guinea Bird Society. In a few cases, well-known scientists have published descriptions of animal homosexuality, including Desmond Morris on Orang-utans, Zebra Finches, and Sticklebacks, Dian Fossey on Gorillas, and Konrad Lorenz on Greylag Geese, Ravens, and Jackdaws. ⁵ Aristocracy has even been involved: in addition to Count Buffon's observations in the eighteenth century, in the 1930s the Marquess of Tavistock in England coauthored a report on bird behavior with scientist G. C. Low that included descriptions of same-sex pairs in captive waterfowl. Like Desmond Morris's account of same-sex activity in Orang-utans quoted above, however, his report was somewhat less than "objective," containing as it did a statement about how "ludicrous" were a pair of male Mute Swans that remained together and built a nest each year.⁶

While most scientific studies of homosexuality in animals have simply involved careful and systematic observation and recording of behavioral patterns (occasionally {86} supplemented by photographic documentation), in some cases more elaborate measures have been employed. The study of animal behavior has now become extremely sophisticated and even "high-tech," and many of these techniques have been applied with great effect to the recording, analysis, and interpretation of same-sex activities and their social context. DNA testing, for example, has been employed to ascertain the parentage of eggs belonging to lesbian pairs of Snow Geese, to determine the genetic relatedness of female Oystercatchers and Bonobos who engage in same-sex activity, to verify the sex of Roseate Terns (some of whom form homosexual pairs), and to investigate the genetic determinants of mating behavior in different categories of male Ruffs. The extent and characteristics of homosexual pair-bonding in Silver Gulls and Bottlenose Dolphins have been revealed by long-term demographic studies that identified and marked large numbers of individuals, who were then monitored over extended periods. Because most sexual activity in Red Foxes takes place at night, investigators only discovered same-sex mounting in this species by setting up infrared, remote-control video cameras that automatically recorded the animals' nocturnal activities (night photography was also required to document similar activity in wild Spotted Hyenas). Radio tracking (biotelemetry) of individual Grizzlies revealed the activities of bonded female pairs, while similar techniques applied to Red Foxes yielded information about their dispersal patterns and overall social organization that relate to the occurrence of same-sex mounting. Videography, including "frame-by-frame" analysis of taped behavioral sequences, has been utilized in the study of courtship interactions in Griffon Vultures and Victoria's Riflebirds, as well as of communicative interactions during Bonobo sexual encounters (both same-sex and opposite-sex). One ornithologist even x-rayed the eggs belonging to a homosexual pair of Black-winged Stilts to see if they were fertile (they weren't).⁷

Unfortunately, in a few cases scientists have subjected animals to more extreme experimental treatments, procedures, or "interventions." During several studies of captive animals, same-sex partners in Rhesus Macaques, Bottlenose Dolphins, Cheetahs, Long-eared Hedgehogs, and Black-headed Gulls (among others) were forcibly separated, either because their activities were considered "unhealthy," or in order to study their reaction and subsequent behavior on being reunited, or to try to coerce the animals to mate heterosexually. A female pair of Orange-fronted Parakeets was forcibly removed from their nest — which they had successfully defended against a heterosexual pair — in order to "allow" the opposite-sex pair to breed in their stead (based in part on the mistaken assumption that female pairs are unable to be parents). Female Stumptail Macaques had electrodes implanted in their uteri in order to monitor their orgasmic responses during homosexual encounters, while female Squirrel Monkeys were deafened to monitor the effect on vocalizations made during homosexual activities.

Although intended ostensibly to reveal important behavioral and developmental effects, the "treatments" applied to animals have in some cases been disturbingly similar to those administered to homosexual people in an attempt to "cure" them (separation or removal of partners, hormone therapy, castration, lobotomy, and electroshock, among others). Numerous primates, rodents, and hoofed mammals, {87} for example, have been subjected to hormone injections to see how this might affect their homosexual behavior or intersexuality. Macaques were castrated as part of behavioral studies that included investigations of homosexual activity, as were White-tailed Deer to determine the "cause" of transgender in this species. Cats have even been lobotomized in order to study the effect on their (homo)sexuality. In some cases, biologists have gone so far as to kill individuals participating in same-sex activities (e.g., Common Garter Snakes, Hooded Warblers, Gentoo Penguins) in order to take samples of their internal reproductive organs.⁸ The reasons for this — usually to verify their sex or to determine the condition of their reproductive systems, including the presence of any "abnormalities" — reveal the incredulity as well as the often distorted preconceptions that many scientists harbor about homosexuality. As we will see in the next sections, these attitudes often carry over into the "interpretation" or "explanation" of homosexuality/transgender as well.


A drawing from 1896 showing two male Scarab Beetles copulating with each other. This is one of the first scientific illustrations of animal homosexuality to be published.     

"A Lowering of Moral Standards Among Butterflies": Homophobia in Zoology
      ... I have talked with several (anonymous at their request) primatologists who have told me that they have observed both male and female homosexual behavior during field studies. They seemed reluctant to publish their data, however, either because they feared homophobic reactions ("my colleagues might think that I am gay") or because they lacked a framework for analysis ("I don't know what it means"). If anthropologists and primatologists are to gain a complete understanding of primate sexuality, they must cease allowing the folk model (with its accompanying homophobia) to guide what they see and report.
     — primatologist LINDA WOLFE, 1991⁹

There is an astounding amount and variety of scientific information on animal homosexuality — yet most of it is inaccessible even to biologists, much less to the general public. What has managed to appear in print is often hidden away in obscure journals and unpublished dissertations, or buried even further under outdated value judgments and cryptic terminology. Most of this information, however, simply remains unpublished, the result of a general climate of ignorance, disinterest, and even fear and hostility surrounding discussion of homosexuality that exists to this day — not only in primatology (as Linda Wolfe describes), but throughout the field of zoology. Equally disconcerting, popular works on animals routinely omit any mention of homosexuality, even when the authors are clearly aware that such information is available in the original scientific material. As a result, most people don't realize the full extent to which homosexuality permeates the natural world.

Scientists are human beings with human flaws, living in a particular culture at a particular time. Although the profession demands standards of "objectivity" and nonjudgmental attitudes, a survey of the history of science shows that this has not always been the case. For example, the sexism of much biological thinking has been {88} exposed by a number of feminist biologists over the past two decades.¹⁰ They have shown that not only are scientists fallible human beings, but most are men — and their scientific theorizing has often been (and in many cases continues to be) detrimentally colored by their own and their culture's (often negative) attitudes toward women. This observation can be taken a step further: scientists (who are often heterosexual) frequently project, consciously or unconsciously, society's negative attitudes toward homosexuality onto their subject matter. As a result, both scientific and popular understanding of the subject have suffered.¹¹

There are notable exceptions, of course. A number of scientists have presented relatively value-neutral descriptions of same-sex activity in various species without feeling a need to overlay their own commentary on the behavior, and several authors have recognized that homosexual activity is a "natural" or routine component of the behavioral repertoire in certain animals. Zoologist Anne Innis Dagg, for example, offered a groundbreaking survey of the phenomenon among mammals in 1984 that was light-years ahead of her contemporaries, while the more recent work of primatologist Paul L. Vasey is beginning to directly address some of the inadequacies and biases of previous studies.¹² Aside from these few examples, though, the history of the scientific study of animal homosexuality has been — and continues to be — a nearly unending stream of preconceived ideas, negative "interpretations" or rationalizations, inadequate representations and omissions, and even overt distaste or revulsion toward homosexuality — in short, homophobia.¹³ Moreover, not until the 1990s did zoologists begin to address such biased attitudes: Paul Vasey and Linda Wolfe are, so far, the only scientists to acknowledge in print that there may be a problem in their profession (and Wolfe the only one to name this specifically as homophobia). The full extent, history, and ramifications of the problem, however, have not been previously discussed or documented.

The Perversion of Scientific Discourse
      From a distance this might be mistaken for fighting, but perverted sexuality is the real keynote ... . In fact, the birds seem sometimes hardly to understand themselves, or to know where their feelings are leading them ... . My principal observation during the earlier part of the time ... was a repetition of what I have before noted in regard to the sexual perversion, as one calls it — a term which serves to save one the trouble of thinking ... .
    — from a scientific description of Ruffs in 1906

     Three unnatural tending bonds were observed: ... On July 16 a two-year-old bull closely tended a yearling bull for at least four hours in the Wichita Refuge and attempted mounting with penis unsheathed ... .
    — from a scientific description of American Bison in 1958

     Among aberrant sexual behaviors, anoestrous does were very occasionally seen to mount one another ... .
    — from a scientific description of Waterbuck in 1982¹⁴
{89}
In many ways, the treatment of animal homosexuality in the scientific discourse has closely paralleled the discussion of human homosexuality in society at large. Homosexuality in both animals and people has been considered, at various times, to be a pathological condition; a social aberration; an "immoral," "sinful," or "criminal" perversion; an artificial product of confinement or the unavailability of the opposite sex; a reversal or "inversion" of heterosexual "roles"; a "phase" that younger animals go through on the path to heterosexuality; an imperfect imitation of heterosexuality; an exceptional but unimportant activity; a useless and puzzling curiosity; and a functional behavior that "stimulates" or "contributes to" heterosexuality. In many other respects, however, the outright hostility toward animal homosexuality has transcended all historical trends. One need only look at the litany of derogatory terms, which have remained essentially constant from the late 1800s to the present day, used to describe this behavior: words such as strange, bizarre, perverse, aberrant, deviant, abnormal, anomalous, and unnatural have all been used routinely in "objective" scientific descriptions of the phenomenon and continue to be used (one of the most recent examples is from 1997). In addition, heterosexual behavior is consistently defined in numerous scientific accounts as "normal" in contrast to homosexual activity.¹⁵

The entire history of ideas about, and attitudes toward, homosexuality is encapsulated in the titles of zoological articles (or book chapters) on the subject through the ages: "Sexual Perversion in Male Beetles" (1896), "Sexual Inversion in Animals" (1908), "Disturbances of the Sexual Sense [in Baboons]" (1922), "Pseudomale Behavior in a Female Bengalee [a domesticated finch]" (1957), "Aberrant Sexual Behavior in the South African Ostrich" (1972), "Abnormal Sexual Behavior of Confined Female Hemichienus auritus syriacus [Long-eared Hedgehogs]" (1981), "Pseudocopulation in Nature in a Unisexual Whiptail Lizard" (1991).¹⁶ The prize, though, surely has to go to W. J. Tennent, who in 1987 published an article entitled "A Note on the Apparent Lowering of Moral Standards in the Lepidoptera." In this unintentionally revealing report, the author describes the homosexual mating of Mazarine Blue butterflies in the Atlas Mountains of Morocco. The entomologist's behavioral observations, however, are prefaced with a lament: "It is a sad sign of our times that the National newspapers are all too often packed with the lurid details of declining moral standards and of horrific sexual offences committed by our fellow Homo sapiens; perhaps it is also a sign of the times that the entomological literature appears of late to be heading in a similar direction."¹⁷ Declining moral standards — in butterflies?! Remember, these are descriptions by scientists in respected scholarly publications of phenomena occurring in nature!

In addition to such labels as unnatural, abnormal, and perverse, a variety of other negative (or less than impartial) designations have also been employed in the scientific literature. Once again, these span the decades. Mounting among Domestic Bulls is characterized as a "male homosexual vice" (1983), echoing a description from nearly a century earlier in which same-sex activities between male Elephants are classified as "vices" and "crimes of sexuality" that are "prohibited by the rules of at least one Christian denomination" (1892). Courtship and mounting between male Lions is called an "atypical sexual fixation" (1942); same-sex relations in {90} Buff-breasted Sandpipers are described in an article on "sexual nonsense" in this species (1989); while courtship and mounting between female Domestic Turkeys are referred to as "defects in sexual behavior" (1955). Homosexual activities in Spinner Dolphins (1984), Killer Whales (1992), Caribou (1974), and Adelie Penguins (1998) are characterized as "inappropriate" (or as being directed toward "inappropriate" partners), and same-sex courtship among Black-billed Magpies (1979) and Guianan Cock-of-the-Rock (1985) is called "misdirected." In what is perhaps the most oblique designation, one scientist uses the term heteroclite (meaning "irregular" or "deviant") to refer to Sage Grouse engaging in homosexual courtship or copulations (1942).¹⁸

Besides labeling same-sex behavior with derogatory or biased terms, many scientists have felt the need to embellish their descriptions of homosexuality with other sorts of value judgments. Repeatedly referring to same-sex activity in female Long-eared Hedgehogs as "abnormal," for example, one zoologist matter-of-factly reported that he separated the two females he was studying for fear that they might actually "suffer damage" from continuing to engage in this behavior. Similarly, in describing pairs of female Eastern Gray Kangaroos, another scientist suggested that only in cases where there was no (overt) homosexual behavior between the females could bonding be considered to represent a "positive relationship between the two animals." In the 1930s, homosexual pairing in Black-crowned Night Herons was labeled a "real danger," while one biologist (upon learning the true sex of the birds) referred to his discovery and reporting of same-sex activities in King Penguins as "regrettable disclosures" and "damaging admissions" about "disturbing" activities. More than 50 years later, a scientist suggested that homosexual behavior between male Gorillas in zoos would be "disturbing to the public" were it not for the fact that people would be unable to distinguish it from "normal heterosexual mating behavior." Same-sex pairing in Lorikeets has been described as an "unfortunate" occurrence, while mounting activity between female Red Foxes has been characterized as being part of a "Rabelaisian mood." Finally, in describing the behavior of Greenshanks, an ornithologist used unabashedly florid and sympathetic language to characterize an episode of heterosexual copulation, referring to it as a "lovely act of mating" and concluding, "The grace, movement, and passion of this mating had created a poem of ecstasy and delight." In contrast, homosexual copulations in the same species were given only cursory descriptions, and one episode was even characterized as a "bizarre affair."¹⁹

In a direct carryover from attitudes toward human homosexuality, same-sex activity is routinely described as being "forced" on other animals when there is no evidence that it is, and a whole range of "distressful" emotions are projected onto the individual who experiences such "unwanted advances."²⁰ One scientist surmises that Mountain Sheep rams "deem it an insult to be treated as a female" (including being mounted by another male), while Rhesus Macaques and Laughing Gulls are described as "submitting" to homosexual mounts even when there is clear evidence that they are willing participants (for example, by initiating the activity). Cattle Egrets who are mounted during homosexual copulations are characterized as "suffering males," while female Sage Grouse mounted by other females are their "victims." {91} Orang-utan males who participate in homosexuality are said to be "forced into nonconformist sexual behavior" by their partners even though they display none of the obvious signs of distress (such as screaming and struggling violently) that are characteristic of female Orangs during heterosexual rapes. Scientists describing same-sex courtship in Kob antelopes imply that females try to "avoid" homosexual attentions by circling around the other female or butting her on the shoulder. In fact, these actions are a formally recognized ritual behavior called mating-circling that is a routine part of heterosexual courtships, and not indicative of disinterest or "unwillingness" on the part of the courted female. Females who do not want to be mounted (by male partners) actually drop their hindquarters to the ground (a behavior not observed in homosexual contexts). Same-sex courtship in Ostriches is deemed to be a "nuisance" that goes "on and on" and is perpetrated by "sexually aberrant" males. The calm stance of a courted male (referred to as the "normal" partner) in the face of such homosexual advances is described as "astonishing," while the recipient's occasional acknowledgment of the activity is downplayed in favor of those times when he makes no visible response (interpreted as disinterest). Yearling male Guianan Cock-of-the-Rock are consistently described as "taking advantage of" or "victimizing" adult males that they mount, while their partners are said to "tolerate" such homosexual activity. This is at odds with the descriptions, by the same scientists, of the adult partners as willing participants who actively facilitate genital contact during homosexual mounts and allow the yearlings to remain on their territories (unlike unwanted adult intruders who are chased away or attacked). Finally, male Mallard Ducks that switch from heterosexual to homosexual pairings are described as being "seduced" by other males, while Rhesus Macaques are characterized as reacting with a sort of "homosexual panic" to same-sex advances — both echoing widely held misconceptions about human homosexuality.²¹

In other cases, zoologists have problematized homosexual activity or imputed an inherent inadequacy, instability, or incompetence to same-sex relations, when the supporting evidence for this is scanty or questionable at best and nonexistent at worst. For example, the fact that male homosexual pairs in Greylag Geese engage in higher rates of pair-bonding and courtship behavior is ascribed to an (unsubstantiated) "instability" of same-sex pair-bonds. In fact gander pairs in this species have been documented as lasting for 15 or more years and are described as being, in many cases, more strongly bonded than heterosexual mates.²² Similarly, even though pair-bonds between male Ocellated Antbirds can last for years, one ornithologist insisted on portraying them as "fragile" and liable to dissolve at the mere appearance of a "nubile female." Antbird same-sex pairs do sometimes divorce, but so do heterosexual ones, and any generalizations about the comparative stability of each cannot be made without comprehensive, long-term studies of pair-bonding — which have yet to be undertaken for this species.²³ The fact that sexual activity between female Gorillas generally takes longer than heterosexual copulations is speculatively attributed to "mechanical difficulties" involved in sex between two females — it is apparently inconceivable to the investigator that females might be experiencing closer bonding or greater enjoyment with each other (as reflected by their face-to-face position and other features that also distinguish homosexual {92} from heterosexual activity in this species). In the same vein, accounts of same-sex mounting in Western Gulls, Guianan Cock-of-the-Rock, and Red Foxes refer to the "disoriented," "bumbling," or "fumbling" actions of some individuals — terms that are rarely used to describe nonstandard mounting attempts in heterosexual contexts (even when they are equally "incompetent"). Conversely, one primatologist is willing to concede that affiliative gestures (such as mutual touching, grooming, or preening) between animals of the opposite sex may be "tender" and even "an expression of love and affection," yet similar or identical activities between same-sex participants are never characterized this way.²⁴

This double standard is particularly apparent where descriptions of same-sex pairs in Gulls are concerned. When a male Laughing Gull in a homosexual pair courted and mounted a female, for example, this was taken by one investigator to mean that his pair-bond was unstable and that he was "dissatisfied" with his homosexual partnership (rather than as simply an instance of bisexual behavior). In contrast, homosexual activity by birds in heterosexual pairs is never interpreted as "dissatisfaction" with heterosexuality or as reflecting the tenuousness of opposite-sex bonds. In a study on pair-bonding in Black-headed Gulls, the term "monogamous" (implying stability) was reserved for heterosexual pairs, even though homosexual pairs in this species can also be stable and monogamous, and heterosexual pairs are sometimes nonmonogamous. Likewise, the stability of female pairs of Herring Gulls was claimed to be lower than heterosexual pairs. Yet in making this assessment, researchers were considering females to have broken their pair-bond if they were simply not seen at the nesting colony the following year — when in fact they or their partner could have died, relocated, or been missed by observers. Among those females that were subsequently observed at the colony (a more accurate measure, and the standard way of calculating mate fidelity for heterosexual pairs), the rate of pair stability was in fact nearly identical to that of opposite-sex pairs.

Similarly, the parenting abilities of female pairs in many Gull species are often implied to be substandard because such couples usually hatch fewer chicks than heterosexual pairs. However, calculations of the hatching success of homosexual pairs typically include infertile eggs in the overall count; since many females in same-sex pairs do not mate with males, large numbers of their eggs are infertile and so of course a larger proportion of their clutches do not hatch. In addition, all of the traits taken to indicate poor quality of parenting in some female pairs — e.g., smaller eggs, slower embryonic development, lower hatching rate of fertile eggs, reduced weight and greater mortality of chicks, higher rates of loss or abandonment — are also characteristic of supernormal clutches attended by heterosexual parents (usually polygamous trios). In other words, they are related to the larger-than-average clutch size rather than the sex of the parents per se. In fact, most studies of Gulls have shown that the parenting abilities of homosexual pairs are at least as good as those of heterosexual pairs. Moreover, heterosexual parents in many Gull species can be severely neglectful or overtly violent toward their chicks, causing youngsters to "run away" from their own families and be adopted by others (or even perish). Needless to say, this behavior is never interpreted as being representative {93} of all heterosexual pairs or as impugning heterosexuality in general (even though it is usually far more widespread than homosexual inadequacies).²⁵ Thus, many zoological studies evidence the same inconsistency often found in discussions of human homosexuality: any difficulties or irregularities in same-sex relations are generalized to all homosexual interactions (or else focused on to the exclusion of other examples), whereas comparable problems in opposite-sex relations are seen in the proper perspective, simply for what they are — individual (or idiosyncratic) occurrences that, while noteworthy, do not reflect the entirety of heterosexuality nor warrant disproportionate attention.

Homophobia in the field of zoology is not always this overt or virulent; nevertheless, ignorance or negative attitudes that are not directly expressed usually have identifiable consequences and important ramifications for the way the subject is handled. Discussion of animal homosexuality has in fact been compromised and stifled in the scientific discourse in four principal ways: presumption of heterosexuality, terminological denials of homosexual activity, inadequate or inconsistent coverage, and omission or suppression of information.

Heterosexual Until Proven Guilty
      ... after about twenty minutes I realized that what I was watching was three whales involved in most erotic activities! ... Then one, two, and eventually three penes appeared as the three whales rolled at the same time. Obviously, all three were males! It was almost two hours after the first sighting ... and up to that point I was convinced I was watching mating behavior. A discovery — and a stern reminder that first impressions are deceiving.
     — JAMES DARLING, "The Vancouver Island Gray Whales"²⁶

Many behavioral studies of animals operate under a presumption of heterosexuality: a widespread — if not universal — assumption among field biologists is that all courtship and mating activity is heterosexual unless proven otherwise. This is particularly prevalent in studies of animals in which males and females are not visually distinguishable at a distance. The scientific literature is filled with examples of biologists who were convinced that the sexual, courtship, or pair-bonding activity they had been observing was between a male and female — until confronted with {94} clear evidence of homosexuality, such as a glimpse of two sets of male genitalia, or a nest containing more eggs than just one female could have laid.²⁷

Moreover, many zoologists still routinely determine the sex of animals in the field based on their behavior during sexual activity — with the (often unstated) assumption that there must be both a male (the one doing the mounting) and a female (the one being mounted) in any such interaction. Of course, this automatically eliminates any "chance" of observing homosexual activity in the first place. A field study of Laughing Gulls, for example, utilized the following assumptions in determining the sex of birds: "(1) any bird copulating more than twice on top was presumed a male, and (2) the mate of a male was presumed to be a female." Yet other studies of this species in both the wild and captivity have revealed that male homosexual mounting and pairing do in fact occur in Laughing Gulls. Scientists studying sexual behavior in Common Murres admitted that they probably underestimated the frequency of homosexual mounting because they assumed that sexual activity involved opposite-sex partners unless they had direct evidence to the contrary. Amazingly, this practice is even used in species where homosexual behavior is known to occur from previous studies of either captive or wild animals, such as Kittiwakes and Griffon Vultures.²⁸ True, some biologists have critiqued this method of sex determination — but only on the grounds that it can miss examples of reverse heterosexual mounting (where females mount males).²⁹ And in spite of its obvious shortcomings, behavioral sex determination continues to be employed in recent studies, some of which constitute the first and only documentation of little-known species. One can only guess at how many examples of homosexual activity have been and will continue to be overlooked because of this.³⁰

Even in captivity, the sex of animals is often mistaken, and the consequent "amending" of mating or courtship activity from heterosexual to homosexual sometimes results in elaborate retractions, revisions, and reinterpretations. Renowned German ornithologist Oskar Heinroth, for example, published one of the first descriptions of heterosexual mating in Emus — only to discover that the two birds he had been observing in captivity were in fact both males, prompting him to publish a "correction" to his earlier description three years later. In reviewing the earliest descriptions of courtship behavior in captive Regent Bowerbirds, scientists realized that what had previously been described as heterosexual activity was in fact display behavior performed between two males. This resulted in rather confusing citations of the earlier material such as the following, in which the true sex of the birds is indicated by the later author's bracketed insertions (prefaced by the assertion "I make no apology for revising in brackets his text to make it meaningful"): "'These love-parlours, each one built by a female [immature male] for her [his] sole use ... were of the shape of a horseshoe ... . The female would enter and squat in her [in the immature male's] love-parlour, the tail remaining towards the entrance ... the rejected females [immature males in adult female dress] ... built or partly built three love-parlours in different spots.'" The very first description of "heterosexual" courtship and mating in Dugongs (a marine mammal) was published, ironically, in a scientific article prefaced with lines of romantic verse about the "heaving bosoms" of mermaids and sea nymphs (creatures that the animal has historically been {95} mistaken for). Ironic, because nearly a decade later biologists confirmed that both animals involved in this sexual activity were actually males.³¹

Perhaps the most convoluted — and humorous — mix-up of this sort involves a set of King Penguins that were studied at the Edinburgh Zoo from 1915 to 1930. The various permutations and shufflings of mistaken gender identities (on the part of human observers, not the birds) reached truly Shakespearean complexity. The sex of the penguins was initially determined on the basis of what was thought to be heterosexual behavior, and the birds were given (human) names accordingly. Following this, however, some "puzzling" observations of apparently homosexual activity were made. Subsequent re-pairings and breeding activity eventually revealed — more than seven years later! — that in fact the sex of all but one of the birds had been misidentified by the scientists. At this point a comprehensive "sex change" in the names of the birds was hastily instituted to reflect their true genders: "Andrew" was renamed Ann, "Bertha" turned into Bertrand, "Caroline" became Charles, and "Eric" metamorphosed into Erica ("Dora" had correctly been identified as a female). Ironically, although some previous "homosexual" interactions could be reclassified as heterosexual once the true sex of the birds was known, other less straightforward revisions were also required. Two penguins that had initially been seen engaging in "heterosexual" activity — "Eric" and Dora — later turned out to be same-sexed, while premature observations of lesbian mating between "Bertha" and "Caroline" were confirmed as homosexual — but actually involved the males Bertrand and Charles!³²

Sometimes the presumption of heterosexuality concerns not the sex of animals but the context in which courtship or pairing activity occurs. This can be characterized as a "heterocentric" view of animal behavior, i.e., one that tends to see all forms of social interaction as revolving around heterosexual activity (see chapter 5). For example, female homosexual pairs in a number of birds, such as Snow Geese, Ring-billed Gulls, Red-backed Shrikes, and Blue Tits, were initially thought to represent the female portion of heterosexual trios. The females were erroneously assumed to be bonded not to each other but to a third, male, bird (that had yet to be observed) — to the extent that several researchers felt compelled to provide explicit evidence and argumentation that no male was associated with such female pairs. Likewise, courtship and mounting activity between male Guianan Cock-of-the-Rock was categorized as a form of "disruption" of heterosexual courtships in one study, when in fact the majority of same-sex activity took place outside of heterosexual courtships when females weren't even around. In a similar vein, same-sex behavior in Stumptail Macaques was classified as sexual in one study only if it occurred "during or immediately after or between heterosexual copulations." In summarizing the pairing strategies adopted by widowed Jackdaws, one scientist enumerated only heterosexual mating patterns and failed to include the formation of female homosexual pairs, even though his own data showed that 10 percent of widowed females attracted new female mates. Likewise, one author's discussion of homosexual activity in male Cheetahs focused on a single case where males mounted each other in apparent "frustration" during heterosexual courtship activities, when in fact the majority of same-sex interactions did not occur in this type {96} of context. Finally, sexual activity and bonding between female Bonobos has traditionally been interpreted as a derivative extension of heterosexuality and subsumed under the general patterns of male-female relations. Recent work, however, shows that female bonding and homosexuality in this species are in fact autonomous from heterosexuality, not geared toward attracting opposite-sex partners, and actually much stronger and more primary than male-female bonding.³³

Similar assumptions have frequently guided the treatment of actual sexual behavior, most blatantly when same-sex activity is excluded entirely from the definition of what constitutes sexual activity. One researcher, for example, only considered cases involving "insertion of the penis into the vagina" to be genuine examples of sexual penetration in Savanna (Olive) Baboons, and a study of Right Whales classified behavior as sexual only if it occurred in groups containing both males and females. A recent study of Moose defined sexual mounting behavior solely as "a male mounting a female," while any mounting activity in Cattle Egrets "in which male-female cloacal [genital] contact appeared to be impossible" was classified a priori as "incomplete" or unsuccessful sexual activity.³⁴ Anal and oral intercourse are not the only forms of penetration excluded by these sorts of definitions. In discussing homosexual activity in female Squirrel Monkeys, one scientist bluntly asserted that clitoral penetration — the insertion of one female's clitoris into another's vagina — was anatomically impossible: "Because of the structure of the female genitalia, however, intromission between females is not possible." In fact, the clitoris in Squirrel Monkeys and many other female mammals becomes conspicuously erect during sexual arousal, and actual clitoral penetration has been documented during lesbian sexual activity in Bonobos, and it may also occur in Spotted Hyenas.³⁵ The phallocentric viewpoint expressed in comments such as these is merely the most recent manifestation of attitudes that can be traced back to some of the earliest descriptions of animal homosexuality. In 1922, for example, one scientist wrote of female homosexual interactions in Savanna (Chacma) Baboons, "The physical completion of the act was, of course, impossible and it seemed more like an impulsive action in which there was no real sexual excitement involved."³⁶ This perfectly epitomizes the sort of stereotypes and misinformation that have continued to engulf homosexuality to this day, in both animals and people.

Mock Courtships and Sham Matings
      The attitude that homosexual activity is not "genuine" sexual, courtship, or pair-bonding behavior is also sometimes made explicit in the descriptions and terminology used by researchers. In spite of witnessing two male homosexual mounts during a morning spent observing Ruffs, for example, one ornithologist reported offhandedly that "there were no real copulations" because no heterosexual mounting took place; a similar comment was made by a scientist studying Bonnet Macaques.³⁷ This attitude is also encoded directly in the words used for homosexual behaviors: rarely do animals of the same sex ever simply "copulate" or "court" or "mate" with one another (as do animals of the opposite sex). Instead, male Walruses indulge in "mock courtship" with each other, male African Elephants and {97} Gorillas have "sham matings," while female Sage Grouse and male Hanuman Langurs and Common Chimpanzees engage in "pseudo-matings." Musk-oxen participate in "mock copulations," Mallard Ducks of the same sex form "pseudo-pairs" with each other, and Blue-bellied Rollers have "fake" sexual activity. Male Lions engage in "feigned coitus" with one another, male Orang-utans and Savanna Baboons take part in "pseudo-sexual" mountings and other behaviors, while Mule Deer and Hammerheads exhibit "false mounting." Bonobos, Japanese and Rhesus Macaques, Red Foxes, and Squirrels all perform "pseudo-copulations" with animals of the same sex.³⁸ Amid this abundance of counterfeit sexual activity, one thing is all too real: the level of denial on the part of some zoologists in dealing with this subject.³⁹

Even use of the term homosexual is controversial. Although the majority of scientific sources on same-sex activity classify the behavior explicitly as "homosexual" — and a handful even use the more loaded terms gay or lesbian⁴⁰ — many scientists are nevertheless loath to apply this term to any animal behavior. In fact, a whole "avoidance" vocabulary of alternate, and putatively more "neutral," words has come into use. "Male-male" or "female-female" activity is the most common appellation, although some more oblique designations have also appeared, such as "male-only social interactions" in Killer Whales or "multifemale associations" for same-sex pairs in Roseate Terns and some Gulls. Homosexual activities are also called "unisexual," "isosexual," "intrasexual," or "ambisexual" (meaning single-sex, same-sex, within-sex, and bisexual, respectively) in various species such as Gorillas, Ruffs, Stumptail Macaques, Hooded Warblers, and Rhesus Macaques. The use of "alternate" words such as unisexual is sometimes advocated precisely because of the homophobia evoked by the term homosexual: one scientist reports that an article on animal behavior containing homosexual in its title was widely received with a "lurid snicker" by biologists, many of whom never got beyond the "sensationalistic wording" of the title to actually read its contents.⁴¹

Occasionally there are directly opposing assertions regarding the suitability of the term homosexual for the same behavior and species. In a relatively enlightened treatment of same-sex activities in Giraffes, for example, one zoologist stated, "Such usage [of the term homosexual] is acceptable provided it is used without the usual human connotation of stigma and sexual abnormality ... . In giraffes the erection of the penis, mounting, and even possibly orgasm leaves little doubt as to the sexual motivation behind these actions." In contrast, a decade later another zoologist objected, "Considerable significance has been attached to the fact that necking males sometimes show penis erections and that one may mount the other ... such behavior has been called 'homosexual.' However ... I ... do not feel that the use of the term homosexual, with its usual (human) connotation, is justified in this context."⁴² Ironically, where the first scientist objected only to the stigma associated with the term as applied to people, the second objected to the connotation of genuinely sexual behavior in the term as applied to people.

When it comes to heterosexual activities, however, scientists are not at all adverse to making analogies with human behaviors. Opposite-sex courtship-feeding in birds is described as "romantic" and reminiscent of human lovers kissing, male canaries whose vocalizations attract female partners are said to sing "sexy" songs, while avian {98} heterosexual monogamy and foster-parenting are compared to similar activities in people (in spite of the acknowledged differences in the behaviors involved). Even more flagrant anthropomorphizing sometimes occurs: male-female interactions in Savanna Baboons, for example, are likened to "May-December romances," "flirting," and other human courtship rituals in a "singles bar"; polyandry in Tasmanian Native Hens is termed "wife-sharing"; opposite-sex bonds between cranes who readily pair with one another are characterized as "magic marriages"; and heterosexually precocious male Bonobos are dubbed "little Don Juans." Female fireflies that lure males of other species by courting and then eating them are labeled "femmes fatales," and one scientist even uses the term gang-bang to describe group courtship and forced heterosexual activity in Domestic Goats. Regardless of whether these characterizations are appropriate, among zoologists it is still more acceptable (in practice if not in theory) to draw human analogies where heterosexuality is concerned.⁴³

Many scientists' denial that same-sex courtship, sexual, pair-bonding, and/or parenting activities should be put in the category of "homosexuality" are based on spurious or overly restrictive interpretations of the phenomenon (or the word). For example, Konrad Lorenz claims that gander pairs in Greylag Geese are not actually "homosexual" because sexual behavior is not necessarily an important component of such associations (not all members of gander pairs engage in sexual activity), and because not all such birds pair exclusively with other males over their entire lifetime. By the same criteria, however, opposite-sex pairs would fail to qualify as "heterosexual": sexual activity is not an important component of male-female pairings in this species (as Lorenz himself acknowledges), and not all such birds pair exclusively with opposite-sex partners during their lives. Yet Lorenz has no qualms about labeling such pairs "heterosexual."⁴⁴ In fact, what we have here is simply an attempt to equate homosexuality with only one characteristic or type of same-sex activity (sexual versus pair-bonding, or sequential bisexuality versus exclusive homosexuality).

In a parallel discussion of female pairs in Western Gulls, one researcher suggests that previous descriptions of such pairs as "homosexual" or "lesbian" or "gay" is inappropriate because they do not resemble homosexual pairings in humans.⁴⁵ But which homosexual pairings, in which humans? As discussed in chapter 2, there is no single type of same-sex pair-bonding in people: homosexual couples differ vastly in a wide range of factors such as their sexual behavior, social status, formation process, sexual orientation of members, participation in parenting, duration, and so on, and they vary enormously between different cultures, historical periods, and individuals. Assuming, however, that this author is referring to Euro-American lesbian couples, it is difficult to see what specific similarities are required before the label of homosexual would be considered acceptable. Same-sex pairs in both Gulls and humans engage in a variety of courtship, pair-bonding, sexual, and parenting activities and exhibit parallel variability in their formation, social status, and the sexual orientation of their partners. In fact, it is fallacious to suggest that a same-sex activity should resemble some human behavior before we can label it homosexual. A more reasonable approach (the one used in this book as well as in many scientific sources) is to take comparable behaviors in the same or closely related species as the {99} point of reference: any activity between two animals of the same sex that involves behaviors independently recognized (usually in heterosexual contexts) as courtship, sexual, pair-bonding, or parenting activities is classified as "homosexual." By this criterion, same-sex pairs of Gulls are "homosexual" because all of the characteristics they exhibit are well-established components of pair-bonding in heterosexual pairs of the same species — to the extent that same-sex couples have often been mistaken for heterosexual ones and unhesitatingly labeled a "mated pair" before their true sex was discovered.

More generally, a number of scientists have suggested that the term homosexual should be reserved for overt sexual behavior, and that it is inappropriate to apply this word to other behavior categories such as same-sex courtship, pair-bonding, or parenting arrangements. We might characterize this as a "narrow" definition of homosexuality (such as that assumed by Lorenz). On the other hand, homosexuality, as the term is used in this book, refers not only to overt sexual behavior between animals of the same sex, but also to related activities that are more typically associated with a heterosexual or breeding context. This usage is consistent with a number of studies in the zoological literature, in which the word is employed as a cover term for both sexual and related behaviors (e.g., courtship, pairing, parenting).⁴⁶ We might characterize this as a "broad" definition of homosexuality. Although overt sexual behavior is by far the single most common type of same-sex activity found in various species — hence the original terminology — the other behavior categories also occur in a sizable proportion of cases in which same-sex activities have been documented. In many (but not all) species, behaviors of various categories co-occur (e.g., sexual and courtship activity with pair-bonding, courtship or bonding with parenting, and so on). There are also numerous cases where only one behavior type is instantiated, or where several behavior categories co-occur in the same species but are not necessarily observed in the same individuals (e.g., sexual behavior may be seen between some animals, courtship behavior between others, etc.). In some cases this represents actual discontinuities of behaviors; in others, it represents observational gaps. When the term homosexuality is employed in the broad sense for these cases, it is always with the understanding that only selected behavior categories or co-occurrences may be involved (as in observations of heterosexual behavior).⁴⁷

The difference between these two usages of the term homosexual can be illustrated with an example involving two different forms of same-sex activity (each widely attested in birds, sometimes both in the same species). On one hand, consider two female birds that are pair-bonded to each other for life, regularly engage in courtship activity with one another, build a nest each year in which they jointly lay eggs, and on one occasion raise chicks together (fathered via a single heterosexual copulation that season by one of the partners), yet never mount each other. On the other hand, consider a male bird who is mated to a female partner for life — with whom he regularly copulates and raises offspring — but who participates in a single copulation with another male (and never again engages in such behavior for the remainder of his life). A narrow definition of homosexuality would require us to consider the first case to be somehow less "homosexual" than the second simply {100} because no overt sexual behavior takes place between the two females. A broad view of homosexuality, on the other hand, recognizes that both cases involve homosexual behavior — but of two distinct types that need to be carefully distinguished in terms of their social context as well as the other sexual and pairing activities of the participants (since both scenarios actually exemplify contrasting forms of bisexuality). Unlike the narrow definition, this usage acknowledges the complexities and variability of same-sex interactions in the animal world, while providing a useful framework for cross-species comparisons and generalizations; it also offers the possibility of more precise and nuanced characterizations of sexual orientation.

Most scientists are understandably wary of anthropomorphizing animals with terms that have wide applicability in a human context — as well they should be — and obviously not all zoologists who avoid the word homosexual are motivated by homophobia. Nevertheless, the lengths that are taken to circumvent terminology that can easily be clarified with a simple explanatory statement often border on the absurd.⁴⁸

"Not Included in the Tabulated Statistics"
      Even when homosexual behavior is recognized as such, detailed study of it is often omitted or passed over, or the phenomenon is marginalized and trivialized. For instance, numerous published reports on the courtship and copulation behavior of animals provide excruciatingly detailed descriptions and statistics on frequency of mounts, number of ejaculations, duration of penile erections, number of thrusts, timing of estrous cycles, total number of sexual partners, and so on and so forth — but all for heterosexual interactions. In contrast, homosexual activity is often mentioned only in passing, not deemed worthy of the exhaustive coverage that is afforded "real" sexual behavior.⁴⁹ In a detailed study of Spinner Dolphin sexual activity, for example, only heterosexual behavior is quantified and given a thorough statistical treatment, even though the author recognizes the prominence of homosexual activity in this species and actually states directly that its frequency exceeds that of heterosexual behavior. Another study of the same species mentions homosexual copulations without providing the total number observed, unlike heterosexual matings. In a tabulation of homosexual and heterosexual activity in Kob antelopes, the number of male partners of each female is cataloged while the number of female partners is not. Likewise, articles on Crested Black Macaque and Brown Capuchin sexual behavior acknowledge the occurrence of female homosexual activity yet offer no statistics on this behavior, even though it is said to be more common (in Crested Blacks) than male homosexual activity (which, along with heterosexual behavior, is quantified). Finally, graphs of the frequency of various Giraffe activities in one study fail to provide adequate information on homosexual mounts: all same-sex interactions are lumped into the category of "sparring" (a form of fighting) without distinguishing actual sparring from necking (a ritualized, nonviolent form of play-fighting and affection) or mounting activity.⁵⁰

Sometimes certain aspects of homosexual activity are excluded or arbitrarily eliminated from an overall analysis or tabulation — often resulting in a distorted picture of same-sex interactions (regardless of whether the omission is deliberate {101} or well-motivated). For instance, a female Western Gull who exhibited the most overt sexual activity with her female partner was "not included in the tabulated statistics" of a study comparing heterosexual and homosexual behaviors. By failing to incorporate data from this individual (intentionally or not), researchers undoubtedly helped foster the (now widely cited) impression that sexual activity is a uniformly negligible aspect of female pairing in this species. Along the same lines, scientists surveying pair formation in Black-crowned Night Herons only tabulated homosexual couples that they considered to be "caused" by the "crowded" conditions of captivity. They ignored a male pair whose formation could not be attributed to such conditions and also overlooked the fact that such "crowded" conditions regularly occur in wild colonies of the same species. And all data concerning same-sex pairs or coparents in Laughing Gulls, Canary-winged Parakeets, Greater Rheas, and Zebra Finches were excluded from general studies of pair-bonding, nesting, or other behaviors in these species.⁵¹

The significance of homosexual activity is sometimes also downplayed in discussions of its prevalence or frequency. Certainly many variables must be considered when trying to quantify same-sex activity, and the task is rarely straightforward (as we saw in chapter 1). Nevertheless, in some instances homosexual frequency is interpreted or calculated so as to give the impression that same-sex activity is less common than it really is or else is de-emphasized in terms of its importance relative to other species. In Gorillas, for example, homosexual activity in females is classified as "rare" because investigators observed it "only" 10 times on eight separate days. However, these figures are incomplete unless compared with the frequency of heterosexual interactions during the same period. In fact, 98 episodes of heterosexual mating were recorded during the same period, which means that 9 percent of all sexual activity was homosexual — a significant percentage when compared to other species.⁵² Similarly, investigators studying lesbian pairs in Western Gulls state, "We have estimated female-female pairs make up only 10-15 percent of the population" (emphasis added), when in fact this is one of the higher rates recorded for homosexual pairs in any bird species (and certainly the highest rate reported at that time). Homosexual mounting in female Spotted Hyenas is claimed to be much less frequent than in other female mammals, yet no specific figures are offered; the one species that is mentioned in comparison is the Guinea Pig, a domesticated rodent that is not necessarily the best model for a wild carnivore.⁵³

It is also important to consider the behavioral type and context when evaluating frequency. Homosexual copulations in Tree Swallows, for example, have been characterized as "exceedingly rare" because they have been observed only infrequently and are much less common than heterosexual matings between pair-bonded birds. However, homosexual copulations are nonmonogamous matings (i.e., they typically involve birds that are not paired to one another and may even have heterosexual mates); it is insufficient in this case to compare the frequency rates of two different kinds of copulation (within-pair and extra-pair). In fact, the more comparable heterosexual behavior — nonmonogamous copulations involving males and females — are also "rarely" seen. Early observers considered them to be exceedingly uncommon (or nonexistent), while a later study documented only two {102} such matings during four years of observation, and subsequent research has yielded consistently low levels of observed promiscuous (heterosexual) copulations. Yet scientists now know that such matings must be common because of the high rates of offspring resulting from them — in some populations, more than three-quarters of all nestlings (as verified by DNA testing). Thus, it is likely that the frequency of homosexual nonmonogamous matings has been similarly underestimated.⁵⁴

Many scientists, on first observing an episode of homosexual activity, are also quick to classify the behavior as an exceptional or isolated occurrence for that species. In contrast, a single observed instance of heterosexuality is routinely interpreted as representative of a recurrent behavior pattern, even though it may occur (or be observed) extremely rarely or exhibit wide variation in form or context. This sets up a double standard in assessing and interpreting the prevalence of each behavior type, especially since opposite-sex mating can be a less than ubiquitous or uniform feature of an animal's social life (see chapter 5). It also conflicts with the patterns established for other species. In repeated instances, homosexual activity was initially recorded in only one episode, dyad, or population (and usually interpreted — or dismissed — as an isolated example), but was then confirmed by subsequent research as a regular feature of the behavioral repertoire of the species — often spanning many decades, geographic areas, and behavioral contexts. ⁵⁵ It is no longer possible to claim that homosexuality is an anomalous occurrence in a certain species simply because it has only been observed a handful of times.

In some cases, conflicting verbal assessments of the prevalence of homosexual activity are offered by the same investigators, when the actual quantitative data show a relatively high occurrence. Homosexual courtship/copulation in Pukeko, for example, is described as being both "common" and "relatively rare" — the actual rate of 7 percent of all sexual activity is in fact fairly high compared to other species (and the same-sex courtship rates are even higher). Likewise, a report on Black-headed Gulls states, "Homosexual pairs were also rare," then a few pages later counterasserts that "male-male bonds occurred rather commonly" — and at approximately 16 percent of all pairs observed, the actual rates support the latter interpretation more than the former.⁵⁶ Not only are these assessments inconsistent and unfair with regard to the observed rates of homosexuality, they also run counter to a standard cross-species measure of heterosexual frequency. Although there is no absolute or universal criterion for what is "rare" or "common," biologists do recognize a "threshold" of 5 percent as being significant where at least one heterosexual behavior is concerned — polygamy. When this mating system is exhibited by only a minority of the population (as is true in many birds, for example), it is nevertheless considered to be a "regular" feature of the species' behavioral repertoire when its incidence reaches 5 percent. This is certainly far less than the rate of homosexuality in many species where same-sex behavior is regarded as "uncommon" or "exceptional."⁵⁷

In a vivid example of the marginalization that often surrounds discussion of animal homosexuality, scientists sometimes find their own descriptions of same-sex activity published with "amendments," "asides," or "explanations" inserted by {103} journal or reprint editors who are uncomfortable with the content or appellation. For example, one ornithologist's description of homosexual activity in House Sparrows and Brown-headed Cowbirds was embellished with a note from the editor of the journal where it appeared, offering several implausible "reinterpretations" of the behavior that eliminated any sexual motivation. Likewise, when descriptions of homosexual activity in Baboons from the 1920s were reprinted nearly half a century later, a scientist who penned the introduction to the new edition felt compelled to annotate the offending passages with the "modern" viewpoint that such activity is not really homosexual. And editors of the journal British Birds scrambled to try to "explain" a case of homosexual pairing in male Kestrels as actually involving a "male-plumaged female" (i.e., a female bird that looked exactly like a male). They added in their published postscript to the article that this putative plumage variation was, in their opinion, "of much more interest than the copulation or attempted one between the two males" that was the primary focus of the author's report.⁵⁸

In a similar vein, one scientist who observed a pair of female Chaffinches hedged his bets by saying only that "female-plumaged" birds were involved, leaving open the possibility that one might still have been a male (and consequently part of a heterosexual pair) — even though there was absolutely no evidence that either bird could have been a male. He finally had to concede that the birds "were surely females." Sometimes this strategy backfires, as in the case of an early description of courtship display in Regent Bowerbirds (mentioned previously), in which the presumed "female-plumaged" birds both turned out to be males — and therefore still participants in homosexual activities.⁵⁹ These cases show that scientists are sometimes reluctant even to commit to the sex of the animals they are observing if it seems that homosexuality might be involved — in stark contrast to the haste with which they usually judge (or assume) participants to be opposite-sexed on the scantiest of evidence.

The Love That Dare Not Bark Its Name
      Although the first reports of homosexual behavior among primates were published >75 years ago, virtually every major introductory text in primatology fails to even mention its existence.
     — primatologist PAUL L. VASEY, 1995⁶⁰

     In the 1890s, Oscar Wilde's lover, Lord Alfred Douglas, characterized homosexuality as "the Love that dare not speak its name," referring to the silence and stigma surrounding disclosure of homosexual interests and discussion of same-sex activities. ⁶¹ An analogue to this silencing and stigmatization exists in the pages of zoology journals, monographs, and textbooks, and in the wider scientific discourse. Discussion of homosexual activity in animals has frequently been stifled or eliminated, and a number of examples can only be considered active suppression of information on the subject. When several comprehensive reference works devoted to every conceivable aspect of an animal's biology and behavior are published, including {104} chapters by scientists who originally observed homosexuality in the species, and yet consistently no mention is made of that homosexual behavior, one has to wonder about the "objectivity" of these scientific endeavors.

At one extreme, there are cases of apparently deliberate removal of information. In 1979, a report on Killer Whale behavior was issued by the Moclips Cetological Society, a nonprofit scientific organization devoted to whale study. Sexual activity between males — classified explicitly as "homosexuality" in the report — was discussed at some length, concluding with the statement, "Homosexual behavior has been observed in many animals including cetaceans, canids, and primates, and, in some cases, it has significance for social order." A year later, when this report was published as a government document for the U.S. Marine Mammal Commission, all mention of homosexuality was eliminated even though the remainder of the report was intact.⁶² At the other extreme are cases where homosexuality is discussed but is buried in unpublished dissertations, obscure technical reports, foreign-language journals, or in articles whose titles give no clue as to their content. For example, the earliest reports of same-sex courtship and mounting in wild Musk-oxen appeared in an unpublished master's thesis at the University of Alaska and a (published) report for the Canadian Wildlife Service. Consequently, a study on homosexual activity in captive Musk-oxen conducted more than 20 years after the initial discovery fails to mention any occurrence of this behavior in the wild. Similarly, the first reports of Walrus homosexual activity, complete with photographs, were published in an article with the rather opaque title of "Walrus Ethology I: The Social Role of Tusks and Applications of Multidimensional Scaling," while all records of homosexual behavior in Harbor Seals are contained in unpublished reports and conference proceedings that are only available at a handful of libraries in the world. This perhaps explains why virtually every subsequent discussion of homosexuality in animals omits any mention of these two species.⁶³

Between these extremes are numerous examples where homosexuality is "overlooked" or fails to gain mention. Describing itself as "the culmination of years of intensive research and writing by more than 70 authors" — all experts on the species — the massive book White-tailed Deer: Ecology and Management (1984) presents in minute detail every imaginable aspect of this animal's biology and behavior, no matter how obscure or rare. There's even room in the book's nearly 900 pages for lengthy discussion of "abnormal" and pathological phenomena (a category in which homosexual activity is often placed). Although the chapter on behavior was coauthored by the scientist who originally described homosexual mounting in White-tailed Deer, there is no mention anywhere in the book of this particular behavior. Nor is there discussion of the transgendered deer found in Texas, even though a whole chapter is devoted to this regional population. A decade later, the same scenario was repeated when another volume of the same scope and on the same species was put out by the same publishers. Similarly, a standard scientific source book, The Gray Whale, Eschrichtius robustus (1984), omits any reference to homosexuality in this species even though it includes a chapter by the first biologist to record same-sex activity in Gray Whales.⁶⁴ Several comprehensive reference volumes on woodpeckers fail to mention homosexual copulations in {105} Black-rumped Flamebacks, even though no other (hetero)sexual behavior has ever been observed in this species. This omission cannot be due to the putative rarity or "insignificance" of such behavior, since one book does mention another behavior that has only ever been observed once in wild woodpeckers — bathing.⁶⁵ Other in-depth surveys of individual species follow suit, eliminating any mention of homosexuality even when they make direct use of other information from the very sources that describe same-sex activity.⁶⁶

Because of the omission and inaccessibility of information on animal homosexuality in the scientific literature, many zoologists are themselves unaware of the full extent of the phenomenon. One of the most unfortunate consequences of this is that misinformation (and absence of information) about the subject is widely disseminated and perpetuated from one source to the next. On discovering homosexual activity in a particular species they are studying firsthand — and being unable to find more than a handful of comparable examples in a cursory literature search — many zoologists acquire the mistaken impression that their observations of this behavior are somehow unique or unusual. At that point they may issue blanket statements to the effect that homosexual activity is rare or previously unreported in the form or species they are observing. Such statements are then often repeated by other biologists and become definitive pronouncements on the subject. As recently as 1993, for example, a scientist reporting on Hooded Warblers could claim that male homosexual pairs had not previously been seen in wild birds — when, in fact, such pairs were documented more than a quarter century earlier in Antbirds, Orange-fronted Parakeets, Golden Plovers, and Mallard Ducks, and thereafter in Black Swans, Scottish Crossbills, Black-billed Magpies, and Pied Kingfishers, among others.⁶⁷ Scientists studying same-sex pairs of Black-headed Gulls in captivity asserted in 1985 that this behavior had yet to be seen in this species in the wild — apparently unaware of a description of a male homosexual pair in wild Black-headed Gulls published in a Russian zoology journal just a year earlier. And researchers who discovered same-sex matings in Adelie and Humboldt Penguins and in Kestrels stated that they did not know of any comparable phenomena in other species of penguins or birds of prey, when in fact homosexual activity in King Penguins, Gentoo Penguins, and Griffon Vultures had previously been reported in the literature.⁶⁸

Sadly, omission and misinformation on the subject of animal homosexuality have ramifications far beyond the individual scientific articles in which they occur. Reference works such as those mentioned above are frequently consulted by researchers in other fields, and they are also the source of much of the information on animal behavior that is presented to the general public. As the quote at the beginning of this subsection indicates, the cycle is also perpetuated through each new generation of scientists as the textbooks they use (or the professors who instruct them) continue to offer inaccurate or incomplete information on the subject (when they aren't completely silent on the topic). It is no surprise, then, that many scientists — and, by extension, most nonscientists — continue to harbor the erroneous impression that homosexuality does not exist in animals or is at best an isolated and anomalous phenomenon. When erasure and silence surround the subject {106} among zoologists, misinformation and prejudice readily fill in the gaps — both in the scientific community and beyond.

To conclude this examination of homophobic attitudes in the scientific establishment, one simple observation can be made: given the considerable obstacles encountered in the recording, analysis, and discussion of the subject, it is remarkable that any descriptions of animal homosexuality make it to the pages of scientific journals and monographs (or to a wider audience). A great deal of progress is being made, and the situation today is certainly improved over that of even a decade ago. Moreover, none of this discourse would even be possible without the invaluable work of zoologists and wildlife biologists who study animals firsthand and report their findings — however flawed that study and reporting may be at times. Nevertheless, the examples of animal homosexuality currently contained in the zoological literature represent only the tip of the iceberg. Many more remain to be discovered, recorded, and granted the scientific attention that has so repeatedly been denied them in the past.

Anything but Sex

As we have seen, one way that zoologists have tried to avoid classifying same-sex activity as "homosexuality" is by using terminology and behavioral categories that deny it is sexual activity at all. This approach also extends to the interpretations, explanations, and "functions" attributed to same-sex behavior, even when it involves the most overt and explicit of activities. Astounding as it sounds, a number of scientists have actually argued that when a female Bonobo wraps her legs around another female, rubbing her own clitoris against her partner's while emitting screams of enjoyment, this is actually "greeting" behavior, or "appeasement" behavior, or "reassurance" behavior, or "reconciliation" behavior, or "tension-regulation" behavior, or "social bonding" behavior, or "food exchange" behavior — almost anything, it seems, besides pleasurable sexual behavior. ⁶⁹ Similar "interpretations" have been proposed for many other species (involving both males and females), allowing scientists to claim that these animals do not really engage in "genuine" (i.e., purely sexual) homosexual activity. But what heterosexual activity is ever "purely" sexual?
{107}
Most biologists are not as candid as Valerius Geist, who, in Mountain Sheep and Man in the Northern Wilds, readily admits to his discomfort and homophobia in trying to "explain" homosexuality in Bighorn Rams as "aggressive" or "dominance" behavior:

      «I still cringe at the memory of seeing old D-ram mount S-ram repeatedly ... . True to form, and incapable of absorbing this realization at once, I called these actions of the rams aggressosexual behavior, for to state that the males had evolved a homosexual society was emotionally beyond me. To conceive of those magnificent beasts as "queers" — Oh God! I argued for two years that, in [wild mountain] sheep, aggressive and sexual behavior could not be separated ... . I never published that drivel and am glad of it ... . Eventually I called the spade a spade and admitted that rams lived in essentially a homosexual society.»⁷⁰

This section will examine a number of nonsexual interpretations, including attempts to classify homosexuality as dominance or aggressive behavior, as a form of play, as a social interaction that relieves group tension, and as a greeting activity. In many cases, these "explanations" are not so much genuine attempts to understand the phenomenon as they are ways of denying its existence in the first place. Often these interpretations are simply incompatible with the facts, especially where "dominance" is involved. Furthermore, while in many instances animal homosexuality does have components of all these (nonsexual) activity types, this does not cancel its sexual aspects. As Paul L. Vasey observes, "Just because a behavior which is sexual in form serves some social role or function doesn't mean it cannot be simultaneously sexual."⁷¹ Indeed, both animal and human heterosexualities also share aspects of these nonsexual functions without losing their classification as "sexual" activities.

The Dominant Paradigm
      In many animal societies, individuals can be ranked with respect to each other on the basis of a number of factors — aggression, access to food or heterosexual mating opportunities, age and/or size, and so on. The resulting hierarchy of individuals and their interaction within this system is often subsumed under the term dominance. Many scientists have suggested that mounting and other sexual behaviors between animals of the same sex are not in fact sexual behavior at all, but rather express dominance relations between the two individuals. The usual interpretation is that the "dominant" partner mounts the "subordinate" one and thereby asserts or solidifies his or her ranking relative to that individual. This "explanation" of homosexual behavior is firmly entrenched within the scientific establishment: one of the earliest statements of this position is a 1914 description of same-sex mounting in Rhesus Macaques, and since then dominance factors have regularly been invoked in discussions of animal homosexuality.⁷² Most scientists have appealed to dominance as an explanation for animal homosexuality only in relation to the particular {108} species (or at most, animal subgrouping) that they are studying — and sometimes only for one sex within that species — without regard for a broader range of considerations. Once the full panoply of animal types, behaviors, and forms of social organization is taken into account, however, it becomes quite clear that dominance has little, if any, explanatory power. While dominance may be relevant in a few specific cases, it cannot account for the full range of homosexual interactions found throughout the natural world. Moreover, even in particular instances where dominance seems to be important, mitigating factors usually render its influence suspect, if not irrelevant.

At the most basic level, dominance is neither a sufficient nor a necessary condition for the occurrence of homosexual behavior in a species. Just because an animal has a dominance-based or ranked form of social organization does not mean that it exhibits homosexuality, and just because homosexual behavior occurs in a species does not mean that it has a dominance hierarchy. For example, many animals with dominance hierarchies have never been reported to engage in homosexual mounting. Dominance systems are found in "the vast majority of mammal species forming groups with any degree of social complexity" — most primates, seals, hoofed mammals, kangaroos, and rodents, for instance — yet only a fraction of these participate in same-sex mounting. Specific examples of birds with dominance hierarchies but no reported homosexuality include curlews, silvereyes, Harris's sparrows, European jays, black-capped chickadees, marabou storks, white-crowned sparrows, and Steller's jays.⁷³ Conversely, homosexuality is found in many animals that do not have a dominance hierarchy or in which the relative ranking of individuals plays only a minor role in their social system: for example, some populations of Gorillas, Savanna (Olive) Baboons, Bottlenose Dolphins, Mountain and Plains Zebras, Musk-oxen, Koalas, Buff-breasted Sandpipers, and Tree Swallows.⁷⁴

Often, the relevance of dominance to homosexuality contrasts sharply in two closely related species: Pukeko have a well-defined dominance hierarchy that some scientists believe impacts on the birds' homosexual behavior, yet in the related Tasmanian Native Hen, same-sex mounting occurs in the absence of a dominance hierarchy. Male homosexual mounting has been claimed to correlate with dominance in Cattle Egrets, yet in Little Blue Herons this connection is expressly denied. And the white-browed sparrow weaver (and several other species of weaver birds) has an almost identical social organization and dominance system as the Gray-capped Social Weaver, yet mounting between males is only found in the latter species.⁷⁵ Not only cross-species but also cross-gender comparisons are relevant here. A particularly good example of the problematic relationship between dominance and same-sex activity becomes apparent when one looks at males and females within the same species. In many animals both sexes have their own dominance hierarchies, yet homosexuality occurs in only one sex — male but not female Wolves, for example, and female but not male Spotted Hyenas. A corollary to this is that in some species, only one sex exhibits a stable dominance hierarchy, yet homosexuality occurs among both males and females. In Squirrel Monkeys, for example, female interactions are not consistently organized around a dominance or rank system, yet same-sex mounting and genital displays are not limited to males. In Bottlenose {109} Dolphins, stable dominance hierarchies (if they exist at all) are more prominent among females, yet homosexual activity occurs in both sexes.⁷⁶ Finally, homosexual mounting sometimes occurs between animals of different species. Although cross-species dominance relations have been documented (e.g., in birds), in the majority of the cases involving homosexual activity there is no well-established hierarchical relationship between the participating animals of different species.⁷⁷ Clearly, then, dominance cannot be the only factor involved in the occurrence of homosexuality in a given species.

Even in animals where there is a clear dominance hierarchy, same-sex mounting is often not correlated with an individual's rank, and it rarely follows the idealized scenario of "dominant mounts subordinate, always and without exception." In many species, there is simply no correlation between rank and mounting behavior, since subordinate animals frequently mount dominant ones. In Rhesus Macaques, for example, 36 percent of mounts between males are by subordinates on dominants, while 42 percent of all female Japanese Macaque homosexual mounts go "against" the hierarchy, as do 43 percent of mounts between male Common Chimpanzees. ⁷⁸ Both dominant-subordinate and subordinate-dominant mounting occur in Bonobos, Lion-tailed Macaques, Squirrel Monkeys, Gelada Baboons, and Ruffs, among others, while mounting of older, larger, and/or higher-ranking animals by younger, smaller, and/or subordinate individuals has also been reported for numerous species: Common Marmosets, Australian and New Zealand Sea Lions, Walruses, Bottlenose Dolphins, White-tailed Deer, Mule Deer, Pere David's Deer, Wapiti, Moose, Mountain Goats, Red Foxes, Spotted Hyenas, Whiptail Wallabies, Rufous Rat Kangaroos, Moco, Prea, Guianan Cock-of-the-Rock, Emus, and Acorn Woodpeckers. Oftentimes, while a large proportion of mounts may seem to follow the dominance hierarchy in a particular species, mounting by subordinates on dominants also takes place in the same species. This is true for Hanuman Langurs, Bonnet Macaques, Musk-oxen, Bighorn and Thinhorn Sheep, Cattle Egrets, and Sociable Weavers.⁷⁹

The precise opposite of the "standard" dominance-based system of mounting is often found as well: mountings by subordinates on dominants occur more frequently than the reverse in many species. In Crested Black Macaques, for example, 60-95 percent of mounts are subordinate on dominant, while nearly two-thirds of Bison male homosexual mounts are by subordinates on dominants. To complicate things further, this is often combined with a gender difference in the relationship between mounting and dominance, with female mounts "following" the hierarchy and male mounts going "against" it. For instance, in Pig-tailed Macaques mounting between females is usually by a dominant individual on a subordinate one, but more than three-quarters of mounts between males are just the opposite. Similarly, in both Red Deer and Pukeko, females tend to mount lower-ranking animals while males tend to mount higher-ranking ones. There are often individual or geographic differences as well: in some consortships between female Japanese Macaques, all mounting is done by the lower-ranking individual on the higher-ranking partner, while in some populations of Bighorn Sheep, mounting of dominant rams by subordinates is much more prevalent than in other populations.⁸⁰ Furthermore, {110} homosexual mounting in many species is reciprocal, which means that partners exchange positions — mounter becomes mountee, and vice versa — either in the same mounting session or in alternation over longer periods of time. This behavior, which is found in at least 30 different species, is potent evidence of the irrelevance of dominance for homosexual interactions, since mounting should only occur unidirectionally if it strictly followed the rank of the participating individuals.⁸¹ Finally, in some species mounting can also occur between individuals of the same or close ranks — for example, in Common Chimpanzees, White-faced Capuchins, Musk-oxen, Blackbucks, Cavies, and Gray-capped Social Weavers.⁸²

In a dominance-based view of homosexual mounting, it is often assumed that the animal being mounted is somehow a less willing participant in the interaction, "submitting" to the will of the more dominant individual, who thereby asserts his or her "superiority." In fact, in more than 30 species the mounted animal actually initiates the interaction, "presenting" its hindquarters to the other individual as an invitation to mount, sometimes even actively facilitating anal penetration (among males) or other aspects of the interaction. Where the presenting animal is subordinate, this could be interpreted as simply a reinforcement of the dominance system, but in a number of species it is actually the more dominant individual who presents and actively encourages the lower-ranking animal to mount.⁸³ In addition, dominance "explanations" often ignore the clear differences between consensual and nonconsensual mounts (or rapes), as well as evidence for sexual arousal and even enjoyment on the part of mounted animals.⁸⁴

The relationship between sexuality and dominance is complex and multifaceted, differing greatly from the frequent simplistic equating of homosexual mounting with nonsexual rank-based or aggressive behavior. In many species a gradation or continuum exists between sexual mounts and dominance mounts, with one type "blending" into the other so that any distinction between the two is essentially arbitrary. Thus, same-sex mounting can have an unmistakable sexual component even when it still follows a dominance pattern. Among Hanuman Langurs, for example, usually only dominant females mount subordinate ones, yet so inextricably linked are signs of sexual excitement with this behavior that scientists have concluded, "It seems virtually impossible to separate 'sexual mounting' from 'dominance mounting.' ... Sexual arousal and dominance are obviously not mutually exclusive in langur females, since mounting between females is related to both dominance and sexuality."⁸⁵ At the other end of the spectrum, in some species a sharp distinction does in fact exist between two types of mounting, both of which occur between same-sex partners: a nonsexual form associated with dominance and/or aggression, and a clearly sexual form that occurs in other contexts (often within a homosexual pair-bond or consortship). This is true for female Japanese Macaques, Rhesus Macaques, and Black-winged Stilts, and male Greylag Geese, among others.⁸⁶

Finally, in some animals dominance and mounting are entirely separate, with social rank being expressed through obviously nonsexual activities. For example, male Walrus dominance interactions involve fighting and tusk displays that usually occur during the breeding season and often involve younger animals. Male {111} homosexual mounting is not associated with either of these activities and usually takes place in the nonbreeding season among males of all age groups (a similar pattern is also seen in Gray Seals). Oystercatchers use a special ritualized "piping display" (neck arched, bill pointed downward, accompanied by shrill piping notes) to negotiate their dominance interactions, while same-sex mounting and courtship occur in other contexts.⁸⁷ Dominance in many other animals is expressed through fighting and aggressive encounters, access to food or feeding frequency, body size or age, physical displacement (causing another individual to move off through posture, threats, staring, or other activities), access to heterosexual mating opportunities, or a combination of these or other factors, and specifically does not involve mounting or the other homosexual interactions that occur in these species. Savanna (Yellow) Baboons, (female) Hamadryas Baboons, Bottlenose Dolphins, Killer Whales, Caribou, Blackbucks, Wolves, Bush Dogs, Spotted Hyenas, Grizzly Bears, Black Bears, Red-necked Wallabies, Canada Geese, Scottish Crossbills, Black-billed Magpies, Jackdaws, Acorn Woodpeckers, and Galahs are all species in which this is the case.⁸⁸

Another limitation in looking at homosexual interactions from the perspective of dominance is that only mounting behavior lends itself to such an interpretation. A whole host of other homosexual activities do not fit neatly into the dominance paradigm — either because, by their very nature, they are reciprocal activities, or because neither participant can be assigned a clearly "dominant" or "subordinate" status on the basis of what "position" it assumes during the activity. For example, mutual genital rubbing — in which two animals rub their genitals on each other without any penetration — often occurs with neither participant "mounting" the other. Gibbon and Bonobo males frequently engage in this activity when hanging suspended from a branch, facing each other in a more "egalitarian" position. In aquatic animals such as Gray Whales, West Indian Manatees, Bottlenose Dolphins, and Botos, males rub their penises together or stimulate each other while rolling and clasping one another in constantly shifting, fluid body positions that defy any categorization as "mounter" or "mountee." Reciprocal rump rubbing and genital stimulation — found in Chimpanzees and some Macaques — also renders meaningless a dominance-based view of homosexual interactions. When two males or two females back toward each other and rub their anal and genital regions together, sometimes also manually stimulating each other's genitals — which one is "dominating" the other? Or when a male Vampire Bat grooms his partner, licking his genitals while simultaneously masturbating himself, which one is behaving "submissively"? By the same token, Crested Black Macaque females have a unique form of mutual masturbation in which they stand side by side facing in opposite directions and stimulate each other's clitoris — again, because of the pure reciprocity, it makes little sense to interpret this behavior as expressing some sort of hierarchical relationship between the partners.

Genital rubbing, masturbation of one's partner, oral sex, anal stimulation other than mounting, and sexual grooming occur among same-sexed individuals in more than 70 species — yet virtually all of these forms of sexual expression fall outside the realm of clear-cut dominance relationships.⁸⁹ These more mutual, reciprocal, or dominance-ambiguous sexual activities are commonly found alongside {112} homosexual mounting behavior in the same species — but the former are typically ignored when a dominance analysis is advocated.⁹⁰ Ironically, another entire sphere of homosexual activity eludes a dominance interpretation — any same-sex interaction that is not overtly sexual. Courtship, affectionate, pair-bonding, and parenting behaviors that do not involve genital contact or direct sexual arousal — yet still occur between same-sex partners — are routinely omitted from any discussion of the relevance of dominance to the expression of homosexuality.⁹¹ The exclusion of nonsexual behaviors such as these from dominance considerations contrasts, paradoxically, with the way that mounting behavior itself is ultimately rendered nonsexual by its inclusion under the category of dominance.

A final indictment of a dominance analysis is that the purported ranking of individuals based on their mounting or other sexual behavior often fails to correspond with other measures of dominance in the species. Male Giraffes, for example, have a well-defined dominance hierarchy in which the rank of an individual is determined by his age, size, and ability to displace other males with specific postures and stares. Homosexual mounting and "necking" behavior is usually claimed to be associated with dominance, yet a detailed study of the relationship between these activities and an individual's social standing according to other measures revealed no connection whatsoever. Mounting position also fails to reflect an individual's rank as measured by aggressive encounters (e.g., threat and attack behavior) and other criteria in male Crested Black Macaques, male Stumptail Macaques, and female Pig-tailed Macaques. In only about half of all male homosexual mounts among Savanna (Olive) Baboons is there a correlation between dominance status, as determined in aggressive or playful interactions, and the role of an animal as mounter or mountee. In male Squirrel Monkeys, dominance status affects an individual's access to food, heterosexual mating opportunities, and the nature of his interactions with other males, yet the rank of males as evidenced by their participation in homosexual genital displays does not correspond in any {113} straightforward way to these other criteria. Among male Red Squirrels, there is no simple relationship between aggressiveness and same-sex mounting: the most aggressive individual in one study population indeed mounted other males the most frequently, yet he was also the recipient of mounts by other males the most often, while the least aggressive male was hardly ever mounted by any other males. This is also true for Spinifex Hopping Mice, in which males typically mount males who are more aggressive than themselves. Similarly, although male Bison fairly consistently express dominance through displays such as chin-raising and head-to-head pushing, these behaviors do not offer a reliable predictor of which will mount the other. Although some mounts between male Pukeko appear to be correlated with the dominance status of the participants (as determined by their feeding behavior, age, size, and other factors), there is no consistent relationship between these measures of dominance and another important indicator of rank — a male's access to heterosexual copulations (or the number of offspring he fathers). Finally, dominance relations in Sociable Weavers are not always uniform across different measures either: one male, for example, was "dominant" to another according to their mounting behavior, yet "subordinate" to him according to their pecking and threat interactions.⁹²

In fact, multiple nonsexual measures of dominance often fail to correspond even among themselves, and this has led some scientists to suggest that the entire concept of dominance needs to be seriously reexamined, if not abandoned altogether. While it may have some relevance for some behaviors in some species, dominance (or rank) is not a fixed or monolithic determinant of animal behavior. Its interaction with other factors is complex and context-dependent, and it should not be accorded the status of a preeminent form of social organization that it has traditionally been granted.⁹³ Primatologist Linda Fedigan advocates a more sophisticated approach to the role of dominance in animal behavior, eloquently summarized in the following statement. Although her comments are specifically about primates, they are relevant for other species as well:

      «We often oversimplify the phenomena categorized together as dominance, as well as overestimating the importance of physical coercion in day-to-day primate life ... . An additional focus on alliances based on kinship, friendship, consortship, and roles, and on social power revealed in phenomena such as leadership, attention-structure, social facilitation, and inhibition, may help us to better understand the dynamics of primate social interaction. Also it may help us to place competition and cooperation among social primates in proper perspective as intertwined rather than opposing forces, and female as well as male primates in their proper perspective as playing major roles in primate "politics" through their participation in alliance systems.»⁹⁴

Considering the wide range of evidence against a dominance analysis of animal homosexuality — as well as a number of explicit statements by zoologists questioning or entirely discounting dominance as a factor in same-sex activities⁹⁵ — it is surprising that this "explanation" keeps reappearing in the scientific literature {114} whenever homosexual behavior is discussed. Yet reappear it does, even in several studies published in the 1990s. As recently as 1995, in fact, dominance was invoked in a discussion of mounting between male Zebras, and this explanation still has enough currency that scientists felt compelled to refute it in a 1994 account of homosexual copulation in Tree Swallows. In looking through the many examples of the way that this "explanation" has been used, it becomes apparent that the relevance of dominance is often asserted without any supporting evidence, then cited and re-cited in subsequent studies to create a chain of misconstrual, as it were, extending across many decades of scientific investigation. Again and again, early characterizations of homosexual activity as dominance behavior — often hastily proposed on the initial (and unexpected) discovery of this behavior in a species — have been refuted by later, more careful investigations of the phenomenon.⁹⁶ Yet frequently only the earlier studies are cited by researchers, perpetuating the myth that this is a valid characterization of the behavior. For example, in a 1974 report that described same-sex mounting in Whiptail Wallabies, a zoologist referred to dominance interpretations of Rhesus Macaque homosexuality even though more recent studies had invalidated — or at the very least, called into question — such an analysis for this species.⁹⁷

At times, the very word dominance itself becomes simply code for "homosexual mounting," repeated mantralike until it finally loses what little meaning it had to begin with. "Dominance" interpretations have in fact been applied to same-sex mounting regardless of how overtly sexual it is. The relatively "perfunctory" mounts between female Tree Kangaroos or male Bonnet Macaques, as well as interactions involving direct clitoral stimulation to orgasm between female Rhesus Macaques, and full anal penetration and ejaculation in Giraffes, have all been categorized as nonsexual "dominance" activities at one time or another. Even though many scientists have gone on record against a dominance interpretation — thereby challenging the stronghold of this analytic framework — information that contradicts a dominance analysis is sometimes troublesomely discounted or omitted from studies. For example, in several reports on dominance in Bighorn rams, same-sex mounting and courtship activities (as well as certain aggressive interactions) were deliberately excluded from statistical calculations because they frequently involved "subordinates" acting as "dominants," i.e., they did not conform to the dominance hierarchy. One scientist even classified some instances of same-sex mounting in Crested Black Macaques as "dysfunctional" because they failed to reflect the dominance system or exhibit any other "useful" properties.⁹⁸

Nor is this merely a question of relevance to scientists, or simply a matter of esoteric academic interpretation. The assertions made by zoologists about the "functions" of homosexual behavior are often picked up and repeated, unsubstantiated, in popular works on animals, becoming part of our "common knowledge" of these creatures. In a detailed survey of primate homosexuality published in 1995, zoologist and anthropologist Paul L. Vasey finally and definitively put the dominance interpretation of homosexuality in its proper perspective, stating that "while dominance is probably an important component of some primate homosexual behavior, it can only partially account for these complex interactions."⁹⁹ We can only {115} hope that his colleagues — and ultimately, those who convey the wonders of animal behavior to all of us — will take these words to heart once and for all.

The Desexing of Homosexual Behavior
      ... two males (Dinding and Durian) regularly mouthed the penis of the other on a reciprocal basis. This behavior, however, may be nutritively rather than sexually motivated.
      — T. L. MAPLE, Orang-utan Behavior¹⁰⁰

      In nearly a quarter of all animals in which homosexuality has been observed and analyzed, the behavior has been classified as some other form of nonsexual activity besides (or in addition to) dominance. Reluctant to ascribe sexual motivations to activities that occur between animals of the same gender, scientists in many cases have been forced to come up with alternative "functions." These include some rather far-fetched suggestions, such as the idea (quoted above) that fellatio between male Orang-utans is a "nutritive" behavior, or that episodes of cavorting and genital stimulation between male West Indian Manatees are "contests of stamina."¹⁰¹ At various times, homosexuality has also been classified as a form of aggression (not necessarily related to dominance), appeasement or placation, play, tension reduction, greeting or social bonding, reassurance or reconciliation, coalition or alliance formation, and "barter" for food or other "favors." It is striking that virtually all of these functions are in fact reasonable and possible components of sexuality — as any reflection on the nature of sexual interactions in humans will reveal — and indeed in some species homosexual interactions do bear characteristics of some or all of these activities. However, in the vast majority of cases these functions are ascribed to a behavior instead of, rather than along with, a sexual component — and only when the behavior occurs between two males or two females. According to Paul L. Vasey, "While homosexual behavior may serve some social roles, these are often interpreted by zoologists as the primary reason for such interactions and usually seen as negating any sexual component to this behavior. By contrast, heterosexual interactions are invariably seen as being primarily sexual with some possible secondary social functions."¹⁰²

Thus, a widespread double standard exists when it comes to classifying behavior as "sexual." Desexing is selectively applied to homosexual but not heterosexual activities, according to a number of different strategies. The first and most obvious is when scientists explicitly classify the same behavior as sexual when it takes place between members of the opposite sex and nonsexual when it involves members of the same sex. This is readily apparent in the following statement: "Mounting [in Bison] can be referred to as 'mock copulation.' It seems appropriate to classify this action as sexual behavior only when it is directed towards females. The gesture, however, was also directed to males which suggests that it also has a social function." Likewise, because a behavior often associated with courtship in Asiatic Mouflons and other Mountain Sheep (the foreleg kick) was observed more frequently between individuals of the same sex than of the opposite sex, one zoologist {116} concluded that this activity must therefore be aggression rather than courtship. Primatologists reassigned what they had initially classified as sexual behavior in Stumptail Macaques to the category of aggressive or dominance behavior when it took place in homosexual pairs, while marine biologists reclassified courtship and mating activity in Dugongs as nonsexual play behavior once they learned both participants were actually male. Ornithologists studying the courtship display of Laysan Albatrosses also questioned whether this behavior was "truly" related to pair-bonding or mating after they discovered that some courting birds were of the same sex. Finally, because (male) Dwarf Mongooses and Bonnet Macaques are as likely to mount same-sex as opposite-sex partners, scientists decided this behavior must be nonsexual.¹⁰³ This is not to say that behaviors cannot have different meanings or "functions" in same-sex versus opposite-sex contexts, only that the erasure by zoologists of sexual interpretations from same-sex contexts has been categorical and nearly ubiquitous.

Not only do zoologists apply nonsexual interpretations to behaviors when they know that the participants are of the same sex, they also do the reverse, assuming that a superficially nonsexual behavior — especially if it involves aggression — must involve animals of the same sex. A particularly interesting example of these assumptions concerns the flip-flop in interpretation of sexual chases in Redshanks, part of the courtship repertoire of this sandpiper. Because of their somewhat aggressive nature, these chases were originally interpreted as a nonsexual, territorial interaction and assumed to involve two males — in spite of the fact that some scientists reported seeing chases between birds of the opposite sex. Subsequently, more detailed study involving banded birds (enabling individual identification) revealed that most chases did in fact involve a male and a female and occurred early in the breeding season — at which point the behavior was reclassified as a form of courtship. However, it was also discovered that in a few instances two males were actually chasing each other — and of course scientists then tried to claim that, only in these cases, the chasing was once again nonsexual (in spite of the fact that the two males often copulated with each other as well).¹⁰⁴

Sometimes the arbitrary categorization of behaviors reaches absurd levels. In a few instances components of one and the same activity are given separate classifications, or the undeniably sexual character of a homosexual interaction is taken to mean only that the activity is "usually" heterosexual. For example, in one report on female Crested Black Macaques, a behavior labeled the "mutual lateral display" is classified as a "sociosexual" activity, i.e., not fully or exclusively sexual. It is described as a "distance-reducing display" or a form of "greeting" that "precedes grooming or terminates aggression between two animals." Yet the fact that females masturbate each other's clitoris during this "display" — about as definitively sexual as a behavior can get — is inexplicably omitted from the description of this activity. Instead, this detail is included separately in the "sexual behavior" section of the report under the heading "masturbation" — a contradictory recognition that, apparently, part of this behavior is not truly sexual yet part of it is! In the same species, males often become sexually aroused while grooming one another, developing erections and sometimes even masturbating themselves to ejaculation. {117} Amazingly, this is interpreted by another investigator not as evidence of the sexual nature of grooming between males, but rather that grooming is probably an activity "typically" performed by females to males prior to copulation. Apparently such overt sexuality could only be a case of misplaced heterosexuality, not "genuine" homosexuality.¹⁰⁵

Often a behavior is automatically assumed to involve courtship or sexuality when its participants are known to be of the opposite sex — and the criteria for a "sexual" interpretation are generally far less stringent than those applied to the corresponding interactions between like-sexed individuals. In other words, heterosexual interactions are given the benefit of the doubt as to their sexual content or motivation, even when there is little or no direct evidence for this or even overt evidence to the contrary. For example, simple genital nuzzling of a female Vicuna by a male — taking place outside of the breeding season, and without any mounting or copulation to accompany it — is classified as sexual behavior, while actual same-sex mounting in the same species is considered nonsexual or "play" behavior. In Musk-oxen, foreleg-kicking in heterosexual contexts is often much more aggressive than in homosexual contexts. The male's blow to a female's spine or pelvis is sometimes so forceful that it can be heard up to 150 feet away, yet this behavior is still classified as essentially courtship-oriented. If this level of aggression were exhibited in foreleg-kicking between males, the behavior would never be considered homosexual courtship (as it is, this classification is granted only grudgingly, accompanied by the obligatory reference to its "dominance" function between males).

When a male Giraffe sniffs a female's rear end — without any mounting, erection, penetration, or ejaculation — he is described as being sexually interested in her and his behavior is classified as primarily, if not exclusively, sexual. Yet when a male Giraffe sniffs another male's genitals, mounts him with an erect penis, and ejaculates — then he is engaging in "aggressive" or "dominance" behavior, and his actions are considered to be, at most, only secondarily or superficially sexual. In one study of Bank Swallows, all chases between males and females were assumed to be sexual even though they were rarely seen to result in copulation. Indeed, the majority of bird studies label dyads composed of a male and female as "[heterosexual] pairs" in spite of the fact that overt sexual (mounting) activity is rarely verified for all such couples. In contrast, most investigators will not even consider classifying same-sex interactions in birds to be courtship, sexual, or pair-bonding activity — even when they involve the same behavior patterns used in heterosexual contexts — unless mounting is observed. Certain associations between male and female Savanna Baboons and Rhesus Macaques are described as "sexual" relationships or "pair-bonds" even though they often do not include sexual activity. In contrast, bonds between same-sex individuals in these species are characterized as nonsexual "coalitions" or "alliances" even though they may involve sexual activities (as well as the same intensity and longevity found in heterosexual bonds). Finally, the "piping display" of the Oystercatcher described earlier was initially assumed to be a courtship behavior, largely because it is a common activity between males and females. Subsequent studies have shown that this is in fact a primarily nonsexual (territorial or dominance) interaction.¹⁰⁶
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Another strategy adopted by scientists when confronted with an apparently sexual behavior occurring between two males or two females is to deny its sexual content in both same-sex and opposite-sex contexts. For example, because female Crested Black Macaques show behavioral signs of orgasm during homosexual as well as heterosexual mounts, one scientist concluded that this behavior is not reliable evidence of female orgasm in either situation. The fact that intercourse and other sexual interactions occur between like-sexed individuals in Bottlenose and Spinner Dolphins is often taken to be "proof" that such behaviors have become largely divorced from their sexual content and are now forms of "greeting" or "social communication," even in heterosexual contexts. Similarly, copulation in Common Murres has many nonreproductive features: in addition to occurring between males, in heterosexual pairs it frequently takes place before the female becomes fertile. Drawing an explicit analogy with "nonsexual" mounting in primates, one ornithologist suggested that in this bird heterosexual mounting must therefore serve an "appeasement" function rather than being principally a sexual behavior, i.e., females invite their male partners to mate in order to deflect aggression from them. Likewise, nonprocreative copulations in both heterosexual and homosexual contexts in Blue-bellied Rollers are categorized as a form of ritualized aggression or appeasement.¹⁰⁷

A difference in form between homosexual and heterosexual behaviors is often interpreted as a difference in their sexual content. The reasoning is that if same-sex activity does not resemble opposite-sex activity, and only opposite-sex activity is by definition sexual, then same-sex activity cannot be sexual. For example, in Rhesus Macaques most heterosexual copulations involve a series of mounts by the male, only the last of which typically involves ejaculation. Because mounts between males are often single rather than series mounts, they are frequently classified as nonsexual, even when they include clear signs of sexual arousal such as erection, pelvic thrusting, penetration, and even ejaculation. A similar interpretation has also been suggested for mounting between male Japanese Macaques. In contrast, significant differences in form also exist between heterosexual copulation in Macaques (with series mounting) and male masturbatory patterns, yet both activities are clearly sexual and are typically classified as such.¹⁰⁸

In other animals the very characteristics that are used to claim that same-sex activities are nonsexual — their briefness, "incompleteness," or absence of signs of sexual arousal, for example — are as typical, if not more typical, of opposite-sex interactions that are classified as sexual behavior. Nearly a third of all mammals in which same-sex mounting occurs also have "symbolic" or "incomplete" heterosexual mounts in which erection, thrusting, penetration, and/or ejaculation do not occur; "ritual" heterosexual mountings are also typical of many bird species.¹⁰⁹ In Kob antelopes, 52 percent of heterosexual copulations involve at least one mount by the male without an erection; in contrast, 56 percent of homosexual mountings between male Giraffes — sometimes classified as nonsexual — do involve erections. Likewise, only one in four to five heterosexual mounts among northern jacanas results in cloacal (genital) contact, and ejaculation probably occurs in less than three-quarters of Orang-utan heterosexual mounts.¹¹⁰ Evidence for sexual arousal {119} or "completed" copulations is often entirely lacking in heterosexual contexts, yet such male-female mounts are still considered "sexual" behavior. In Walruses, Musk-oxen, Bighorn Sheep, Asiatic Mouflons, Grizzly Bears, and Olympic Marmots, for example, penetration and ejaculation are rarely, if ever, directly observable during heterosexual mounts, while male erections are routinely not visible during White-tailed Deer copulations, ejaculation can only be "assumed" to occur in observations of Orang-utan, White-faced Capuchin, and Northern Fur Seal heterosexual mating, and genital contact is difficult to verify during Ruff male-female mounts (among many other species).¹¹¹

In fact, actual sperm transfer during heterosexual copulations in many species is so difficult to observe that biologists have had to develop a variety of special "ejaculation-verification" techniques. In birds such as Tree Swallows, for example, tiny glass beads or "microspheres" of various colors are inserted into males' genital tracts. If the birds ejaculate during a heterosexual mating, these beads are transferred to the female's genital tract, where they can be retrieved by scientists and checked for their color coding to determine which males have actually transferred sperm. For rodents and small marsupials, biologists actually inject several different radioactive substances into males' prostate glands. During ejaculation, these are carried via semen into females, who are then monitored with a sort of "sperm Geiger counter" to determine which males, if any, have inseminated them.¹¹² If such elaborate lengths are required to verify a fundamental and purportedly self-evident aspect of heterosexual mating, is it any wonder that homosexual matings should sometimes appear to be "incomplete"?

Because of such difficulties in observation and interpretation, scientists have often employed similarly extreme measures in an attempt to "verify" homosexual intercourse. In the early 1970s, for example, a controversy arose concerning to what extent, if at all, mounting activity between male animals was truly "sexual." As proof of its "nonsexual" character, some scientists claimed that full anal penetration never occurred in such contexts (thus equating penetration with "genuine" sexuality). Researchers actually went to the trouble of filming captive male Rhesus Macaques mounting each other in order to record examples of anal penetration; they even anesthetized the monkeys afterward to search for the presence of semen in their rectums. Needless to say, the cinematographic proof of anal penetration they obtained did little to quell any subsequent debate about whether such mounts were "sexual" — all it did was institute a revised definition of "sexual" activity. The fact that they were able to document penetration but not ejaculation simply meant that a new "standard" of sexuality could now be applied: only mounts that culminated in ejaculation were to be considered "genuine" sexual behavior. Ironically, none of these researchers were apparently aware of an earlier field report of homosexual activity in Rhesus Macaques in which both anal penetration and ejaculation were observed.¹¹³

This near-obsessive focus on penetration and ejaculation — indeed, on "measuring" various aspects of sexual activity to begin with — reveals a profoundly phallocentric and "goal-oriented" view of sexuality on the part of most biologists. Not just homosexual activity, but noninsertive sexual acts, female sexuality and {120} orgasmic response, oral sex and masturbation, copulation in species (such as birds) where males do not have a penis — any form of sex whatsoever that does not involve penis-vagina penetration falls off the map of such a narrow definition. The fact is that both heterosexual and homosexual activities exist along a continuum with regard to their degree of "sexuality" or "completeness." Male mammalian mounting behavior, for example, can involve partial mounting, full mounting but no thrusting, thrusting but no erection, erection but no penetration, penetration but no ejaculation, ejaculation but no penetration, penetration and ejaculation without series mounting, and so on and so forth.¹¹⁴ Each stage along this continuum has at one time or another been considered a defining threshold of "true" sexual behavior — often so as to exclude same-sex interactions — rather than as one possible manifestation of a broader sexual capacity that is sometimes, but not always, orgasmically (or genitally) focused.

A nonsexual component of homosexual behavior does appear to be valid in a number of species; in equally many species, there are clear arguments against various nonsexual interpretations, and some zoologists have themselves explicitly refuted nonsexual analyses.¹¹⁵ Overall, though, three important points must be considered in relation to nonsexual interpretations of behaviors between animals of the same sex. First, the question of causality — or the primacy of the nonsexual aspect — must be addressed. Just because an apparently sexual behavior is associated with a nonsexual result or circumstance does not mean that the sole function or context of the behavior is nonsexual. For example, female Japanese Macaques often gain powerful allies by forming homosexual associations, since their consorts typically support them in challenging (or defending themselves against) other individuals. However, a detailed study of partner choices showed that such nonsexual benefits are of secondary importance: females choose their consorts primarily on the basis of sexual attraction rather than on whether they will make the best or most strategic allies.¹¹⁶ Likewise, mounting (or other sexual activity) between animals of the same sex is described in many species (e.g., Bonobos) as a behavior that serves to reduce aggression or tension between the participants. Indeed, individuals who mount each other may be less aggressive to one another or may experience less tension in their mutual interaction, and homosexuality probably does serve a tension-reducing function for at least some animals in some contexts (as does heterosexuality). However, the situation is considerably more involved than this. Tension reduction is as likely to be a consequence of an affiliative or friendly relationship between individuals — a relationship that is also expressed through sexual contact — as it is to be a direct result of their sexual activity. Moreover, as some researchers have pointed out for Bonobos, the causal relationship may also be the reverse of what is usually supposed. That sexual behavior and situations involving tension often co-occur in this species can give the impression that sexuality is functioning only to reduce tension, when in fact it may also create or generate its own tension. Indeed, homosexual activity in male Gorillas often results in increased rather than decreased social tension.¹¹⁷

Second, even if behaviors are classified as nonsexual or having a nonsexual component, the behavioral categories to which they are assigned (aggression, greetings, {121} alliance formation, etc.) are not monolithic. Many important questions remain concerning the forms and contexts of such behaviors — questions that are often overlooked once they receive their "classification." Just because we "know" that a given behavior is "nonsexual" does not mean that we then know everything about that behavior. Apparently sexual behaviors between males in both Bonnet Macaques and Savanna Baboons, for example, are classified as social "greetings" interactions. Yet there are fundamental differences between these two species, not only in the types of activities involved, but in the frequency of participation, the types of participants, the social framework and outcome of participation, and so on.¹¹⁸ Ultimately, classifying such behavior as "nonsexual" is as meaningless, misleading, and unilluminating as many investigators claim a sexual categorization is, if it obscures these differences or fails to address their origin.

Finally, the relationship between sexual and nonsexual aspects of behavior is complex and multilayered and does not fit the simple equations that are usually applied, namely same-sex participants = nonsexual, opposite-sex participants = sexual. In many species, there is clear evidence of the genuinely sexual aspect of behaviors between animals of the same sex, using the same criteria that are applied to heterosexual interactions — for example, penile or clitoral erection, pelvic thrusting, penetration (or cloacal contact), and/or orgasm.¹¹⁹ In still other species there is a gradation or cline between sexual and nonsexual behaviors that defies any rigid categorization — or else there is a sharp distinction between the two, with both occurring among animals of the same sex. Most importantly, the sexual and nonsexual aspects of a behavior are not mutually exclusive. An interaction involving genital stimulation between two males or two females can be a form of greeting, or a way of reducing tension or aggression, or a type of play, or a form of reassurance, or any number of different things — and still be a sexual interaction at the same time. Ironically, by denying the sexual component of many same-sex activities and seeking alternative "functions," scientists have inadvertently ascribed a much richer and varied palette of behavioral nuances to homosexual interactions than is often granted to heterosexual ones.¹²⁰ Because heterosexuality is linked so inextricably to reproduction, its nonsexual "functions" are often overlooked, whereas because homosexuality is typically disassociated from reproduction, its sexual aspects are often denied. By bringing these two views together — by recognizing that both same-sex and opposite-sex behaviors can be all these things and sexual, too — we will have come very close indeed to embracing a fully integrated or whole view of animal life and sexuality.
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