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—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

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 générale et particulière (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 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

The earliest photographic record of animal homosexuality: a pair of male Mute Swans photographed in 1923 on the nest they built together in Scotland. A female pair in the same species was first observed in 1885.

While most scientific studies of homosexuality in animals have simply involved careful and systematic observation and recording of behavioral patterns (occasionallysupplemented 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