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Snow
      The room was suddenly rich and the great bay-window was
      Spawning snow and pink roses against it
      Soundlessly collateral and incompatible:
      World is suddener than we fancy it.

      World is crazier and more of it than we think,
      Incorrigibly plural. I peel and portion
      A tangerine and spit the pips and feel
      The drunkenness of things being various.

      And the fire flames with a bubbling sound for world
      Is more spiteful and gay than one supposes —
      On the tongue on the eyes on the ears in the palms of one's hands —
      There is more than glass between the snow and the huge roses.

      — LOUIS MACNEICE

      ... hugest whole creation may be less incalculable than a single kiss
      — E. E. CUMMINGS
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Introduction
      The most beautiful thing we can experience is the mysterious.
It is the source of all true art and science. He to whom this
emotion is a stranger, who can no longer pause to wonder and
stand rapt in awe, is as good as dead: his eyes are closed.
      — ALBERT EINSTEIN¹

Any book on homosexuality and transgender in animals is necessarily unfinished, a work in progress. The subject is so vast, the types of behaviors so varied, and the number of species involved so large, as to defy any attempt at comprehensiveness. And the scientific research in this area is only in its infancy: new developments and discoveries are continually being made, and the extent of uncharted and as yet unknowable terrain is so great as to render any attempt at completeness hopelessly premature.

Notwithstanding such formidable challenges, this book endeavors to present a reasonably extensive and up-to-date account of the subject. To help narrow the field, certain parameters have been chosen: only examples of homosexual behavior or transgender that have been scientifically documented, for example, are covered in this book (such documentation includes published reports in scientific journals and monographs, and/or firsthand observations by zoologists, wildlife biologists, and other trained animal observers, corroborated by multiple sources whenever possible). Not only does this limit the number of species to be included (many more cases undoubtedly occur but have not been so documented), it establishes a uniform and verifiable platform of data on which to base further discussion. In addition, the book focuses primarily on mammals and birds — not because other types of animals are somehow less interesting or "important," but simply because space and time limitations necessitate that not all species can be covered. These two groups are considered to be sufficiently representative and to have a broad enough appeal to warrant their inclusion, however arbitrary the exclusion of others may be.

Even with these parameters in place, however, an enormous amount of ground must still be covered. In addition to discussing an extensive array of species (nearly 300 mammals and birds), the book draws upon more than two centuries of scientific {2} research. Some of the findings reported here in a few sentences represent literally lifetimes of work on the part of biologists, who often devote their entire careers to studying one very specific and complex aspect of one type of behavior, in one particular population of one particular species. With this in mind, the book should be seen not as a final, definitive pronouncement on the subject, but rather as a beginning or overture, an invitation to further research and discussion.

Any account of homosexuality and transgender in animals is also necessarily an account of human interpretations of these phenomena. Because animals cannot speak directly for themselves the way people can, we must rely on human observations of their behavior. This presents both special challenges and unique advantages to the study of the subject. On the one hand, certain behaviors such as sexual acts can be observed directly (and even quantified), which is often extremely difficult, impossible, or unethical to do in studies of sexuality among people (especially stigmatized or alternative forms of sexuality). On the other hand, we are in the dark about the internal experiences of the animal participants: as a result, the biases and limitations of the human observer — in both the gathering and interpretation of data — come to the forefront in this situation. In many ways this is the reverse of what occurs in some studies of homosexuality among people (including well-informed historical or anthropological studies of different cultures or time periods). With people, we can often speak directly to individuals (or read written accounts) about what their sexuality and associated phenomena mean — and so get a sense of their emotional and motivational states — without necessarily being able to verify their actual sexual behaviors. With animals, in contrast, we can often directly observe their sexual (and allied) behaviors, but can only infer or interpret their meanings and motivations. As a result, many contentious assertions, theories, interpretations, and explanations have been put forward (and continue to be made) within the field of zoology about the function(s) and meaning (s) of homosexuality and transgender. This book seeks to address this historical and very human dimension of the subject, while still maintaining a focus on the animals, their behaviors and lives.

The unique historical moment we find ourselves in also necessitates the book being geared as much as possible toward specialist and nonspecialist alike, and informs the organization and two-part structure of the book. Because of the current inaccessibility of a large body of scientific information, a primary aim is to present the technical material to a general (nonacademic) readership, without sacrificing accuracy or sensationalizing what is often a controversial and difficult subject matter. However, because no comprehensive survey (and synthesis) of this material is yet available within the scientific literature — indeed, many zoologists are themselves unaware of much of this material — and because a considerable amount of misinformation and misunderstanding surrounds the subject even among trained biologists, the volume will also be of interest to the scientific community. Consequently, every effort has been made to provide full documentation in the form of notes and references, and to include relatively exhaustive and detailed coverage of a wide range of species. However, this more technical material is positioned in such a way that it can easily be skipped by readers who do not wish to delve into such matters.
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In a book such as this which is intended for both an academic and a nonacademic readership, the question of terminology poses special challenges. I have attempted to steer a course between more accessible but overly anthropomorphic or loaded vernacular, on the one hand, and more "neutral" but highly technical jargon or awkward circumlocutions, on the other. In particular, homosexual(ity) and same-sex are utilized as the labels of choice. Since the words gay and lesbian are burdened with human connotations (cultural, psychological, historical, and/or political) and may not be regarded as appropriate designations for animals, I have been careful to avoid using these terms throughout most of the book (as pointed out in chapter 1). When referring specifically to animals and their behaviors, for example, gay is never employed, while lesbian is used only sparingly (it occurs in less than 3 percent of the more than 3,000 instances in the text where animal homosexuality is named). Even then, lesbian is usually reserved only for cases of linguistic expedience, when alternate phrasings such as "female homosexual(ity)" or "same-sex ... among/between females" would become repetitive, cumbersome, or otherwise infelicitous.

Nevertheless, it is important to recognize that a precedent has been established within the zoological discourse for using the less "neutral" (or more culture-bound) designations. The words gay and lesbian are applied by scientists to animals and their behaviors in a number of scholarly publications spanning the past quarter century, including three separate instantiations in the prestigious journal Nature. As in Biological Exuberance, lesbian is more widely used than gay, e.g., "lesbian females" in Fruit Flies (Cook 1975), a "lesbian pair" in Black-billed Magpies (Baeyens 1979), a Common Chimpanzee behaving "in a lesbian fashion" (de Waal 1982),"gay" Snow Geese (Diamond 1989), "gay courtship" in Long-legged Flies (Dyte 1989), "lesbian behavior" in Bonobos (Kano 1992), "lesbian pairs" in Black Stilts (Reed 1993), "lesbian females" in Lesser Flamingos (Alraun and Hewston 1997), "lesbian copulations" in Oystercatchers (Heg and van Treuren 1998); see chapter 3. As for other terms such as transvestism and transsexual(ity), these are also used in the zoological literature with meanings largely divorced from their human connotations (though other labels are employed as well, such as male/female mimicry or sequential hermaphroditism).

It should also be pointed out that the term homosexual — which many people feel is preferable to gay or lesbian when referring to animals — is not devoid of anthropomorphism. It too is a culture-specific, historical construction with very particular human connotations (the same for other putatively "neutral" designations such as hermaphroditism, mimicry, etc.). In fact, a wide variety of terms used routinely in the zoological literature — e.g., courtship, parent(ing), monogamy, adoption, consort(ship) or, for that matter, heterosexual, male, etc. — carry the same baggage of human referents. In addition, the range of variation between (and within) animal species in behaviors that are labeled with the same terms is sometimes as great as — if not greater than — the variation in corresponding behaviors between animals and people. In other words, the differences between "mothers" (or "homosexual copulations") in flies and chimpanzees probably equal if not exceed the differences between "mothers" (or "homosexual copulations") in chimpanzees and humans. Yet such terms are applied to a wide range of animals with the understanding that a {4} given word can have variant meanings in different contexts, and that the human connotations are specifically not implied when such vocabulary is used in a zoological context. This issue is discussed more fully in chapter 3, where I offer a careful rationale for the continued use of such terms — specifically with reference to the supposedly anthropomorphic/centric label homosexual and the historical reluctance of zoologists to utilize even this designation.

Furthermore, within this book such terminology is not used in a vacuum: it is accompanied by explicit discussion of the meanings of all such labels when applied to animals — including overt disavowal of their human connotations and extensive consideration of the inappropriateness of making unwarranted human-animal comparisons (see chapter 2). In order to contextualize the discussion, I also address a number of related issues such as the precedent for employing these words within the zoological literature; the problems inherent in any choice of terminology; and the widespread use within scientific discourse of anthropomorphic labels and descriptions for heterosexual animals and behaviors. Finally and perhaps most importantly, I point out in Biological Exuberance that terminological debates themselves are not ahistorical — they reflect and embody very specific cultural and historical streams both within the scientific community and in society at large; they recapitulate (and lag behind) debates regarding "appropriate" terminology for homosexuality in humans; and the effect of such debates within the scientific discourse has often been to distract from the phenomena designated by such terms rather than to clarify them.

Virtually no terminology for animal behavior — particularly sexual behavior — is entirely free of human (cultural, historical, etc.) associations. When confronted with this situation, we have two options: construct an alternative vocabulary of relatively opaque labels and unwieldy circumlocutions that attempts to avoid such bias (but inevitably falls short of this ideal); or use the already available terms with careful qualification of their meanings and an understanding of their historical context, such that they become uncoupled from their anthropomorphic connotations. In Biological Exuberance, I opt for the latter.

 
The book is organized into two complementary sections. Part 1, A Polysexual, Polygendered World, offers a wide-ranging exploration of all aspects of animal homosexuality and transgender: their diversity, history, and meanings. Part 2, A Wondrous Bestiary, presents a series of profiles of individual homosexual, bisexual, and transgendered animals. Where the first part of the book follows a linear, narrative progression, part 2 is organized in a nonlinear, reference format. The two halves of the book are linked via the animals themselves: throughout part 1, the reader is referred to specific animals that are profiled in part 2 and may at any point consult those profiles to supplement the narrative (names of profiled species or groups of related species are capitalized to indicate their inclusion in part 2 and the appendix). Alternatively, those readers more interested in a general cross-species survey or the interpretive/historical aspect can focus almost exclusively on part 1, while those who wish to gain a more in-depth understanding of particular animals can focus primarily on part 2. This dual structure allows the reader to access information {5} on animal homosexuality/transgender in a variety of ways, suited to his or her own reading style.

Chapter 1, "The Birds and the Bees," presents a broad overview of animal homosexuality and transgender, exploring the full range of behaviors and phenomena covered by these terms. Comparisons between animal and human homosexuality are the focus of chapter 2, "Humanistic Animals, Animalistic Humans," including a discussion of the advisability and implications of making such comparisons in the first place. This chapter also exposes the false dichotomy of the "nature versus nurture" debate, by examining the sociocultural dimensions of homosexuality within animal communities. Next, the history of the scientific study of animal homosexuality is chronicled in chapter 3, "Two Hundred Years of Looking at Homosexual Wildlife." This includes documentation of systematic prejudices within the field of zoology in dealing with this subject, which have often hampered our understanding of the phenomenon. Chapter 4, "Explaining (Away) Animal Homosexuality," continues the historical perspective by examining the many attempts to interpret and determine the "function" or "cause" of animal homosexuality and transgender. Most such efforts to find an "explanation" have failed outright or are fundamentally misguided — particularly when they try to show how homosexuality might contribute to heterosexual reproduction. In the next chapter, "Not for Breeding Only," animal life and sexuality are shown not to be organized exclusively around reproduction. A wide range of nonprocreative heterosexual activities are described and exemplified, as are the diverse ways that homosexual, bisexual, heterosexual, and transgendered animals structure their relationship to breeding.

The final chapter of part 1, "A New Paradigm: Biological Exuberance," calls for a radical rethinking of the way we view the natural world. This revisioning begins with an exploration of another, alternative set of human interpretations: traditional beliefs about animal homosexuality/transgender in indigenous cultures. Particular attention is paid to the ways in which these ideas are relevant to contemporary scientific inquiry. As it turns out, Western science has a lot to learn from aboriginal cultures about systems of gender and sexuality. In the remainder of the chapter, a synthesis of a number of "new" sciences is suggested, including chaos theory, post-Darwinian evolutionary theorizing, biodiversity studies, and the theory of General Economy. The approach taken throughout this chapter is exploratory rather than explanatory. Ultimately, this synthesis leads to a worldview in which animal homosexuality and other nonreproductive behaviors suddenly "make sense," while still remaining, paradoxically, "inexplicable" — a worldview that is also remarkably consistent with indigenous perspectives on gender and sexuality.

In the second half of the book, A Wondrous Bestiary, the reader is treated to a series of individual profiles of homosexual, bisexual, and transgendered animals, from Antbirds to Zebras. Each profile is a verbal and visual "snapshot" of one (or several closely related) species, allowing the reader to "meet" the animal and "get to know" it in detail. Part 2 is divided into two major sections, one for mammals and one for birds, each of which in turn is organized around the formal subgroupings of animals in that category. The section on mammals, for example, includes separate groupings for primates, marine mammals, hoofed mammals, and {6} so on. Each profile within these groupings contains a wealth of information — everything from detailed descriptions of courtship displays to statistics on frequency of homosexual behaviors, to background information on the animal's social organization.

Although its focus is primarily on animal homosexuality and transgender, the book actually moves far beyond these subjects to consider much broader patterns in nature and human society. Sexual and gender variance in animals offer a key to a new way of looking at the world, symbolic of the larger paradigm shifts currently underway in a number of natural and social sciences. The discussion is rooted in the basic facts about animal homosexuality and nonreproductive heterosexuality, information that is presented most fully in the individual animal profiles. Using these to expose the hidden assumptions behind the way biology looks at natural systems, a fresh perspective is developed, based on the melding of contemporary scientific insights with traditional knowledge from indigenous cultures. Taking a broad interdisciplinary perspective, the narrative builds upon a solid foundation of scientific and cultural research to arrive at some conclusions that have the potential to fundamentally alter the way we think about the world and our position in it. Biological Exuberance is, ultimately, a meditation on the nature of life itself, and a celebration of its paradoxes and pluralities.

As such, the book seeks not only to convey "the facts" about animal behavior but, perhaps as importantly, to capture something of their "poetry" as well. The beauty and mystery of nature can be found in many forms. And one particular form of natural beauty is the diversity of sexuality and gender expression throughout the animal world. In addition to being interesting from a purely scientific standpoint, these phenomena are also capable of inspiring our deepest feelings of wonder, and our most profound sense of awe.

{xi} Acknowledgments

This project has been a labor of love, one that would not have been possible without the participation and contributions of numerous individuals and organizations. I owe a tremendous debt of gratitude to the many people who helped bring this book to life.

I am especially grateful to all the zoologists, wildlife biologists, natural history photographers, and zoo biologists who generously provided (often previously unpublished) information, data, and/or original photographs in response to my inquiries regarding various species (any errors in fact or interpretation, however, remain solely my responsibility): Arthur A. Allen/David G. Allen (Bird Photographs, New i Geese; John J. Craighead/John W. Craighead (Craighead Wildlife-Wildlands Institute, Montana) — Grizzly Bears; James D. Darling (West Coast Whale Research Foundation, Canada) — Gray Whales; Bruno J. Ens (Institute of Forestry and Nature Research [IBN-DLO], Netherlands) — Oystercatchers; Ron Entius (Artis Zoo, Netherlands) — Flamingos; J. Bristol Foster (Sierra Club of British Columbia) — Giraffes; Clifford B. Frith (Frith & Frith Books, Australia) — Birds of Paradise; Masahiro Fujioka (Applied Ornithology Laboratory, National Agricultural Research Center, Japan) — Egrets; Michio Fukuda (Tokyo Sea Life Park) — Great Cormorants; Valerius Geist (University of Calgary) — Bighorn Sheep; Jeremy Hatch (University of Massachusetts) — Roseate Terns; Dik Heg (University of Groningen, Netherlands) — Oystercatchers; Denise L. Herzing (Wild Dolphin Project/Florida Atlantic University) — Bottlenose/Atlantic Spotted Dolphins; Katherine A. Houpt (New York State College of Veterinary Medicine, Cornell University) — Przewalski's Horses; George L. Hunt Jr. (University of California — Irvine) — Western Gulls; Alan R. Johnson (Station Biologique de la Tour du Valat, France) — Flamingos; Catherine E. King (Rotterdam Zoo, Netherlands) — Flamingos; Tamaki Kitagawa (Ichikawa High School, Japan) — {xii} Black-winged Stilts; Walter D. Koenig (Hastings Natural History Reservation/UC-Berkeley) — Acorn Woodpeckers; Adriaan Kortlandt (United Kingdom) — Great Cormorants; James N. Layne (Archbold Biological Station, Florida) — Botos; Michael P. Lombardo (Grand Valley State University, Michigan) — Tree Swallows; Dale F. Lott (University of California — Davis) — American Bison; Stephen G. Maka (Wildlife/ Environmental Photography, Massachusetts) — Giraffes; Michael Martys (Alpenzoo Innsbruck) — Greylag Geese; Donald B. Miles (Ohio University/University of Washington) — Whiptail Lizard identification; Gus Mills (Hyena Specialist Group, Kruger National Park, South Africa) — Spotted Hyenas; Daniel K. Niven (Smithsonian Environmental Research Unit/Illinois Natural History Survey) — Hooded Warblers; Jenny Norman (Macquarie University, Australia) — Eastern Gray Kangaroos; Yoshiaki Obara (Tokyo University of Agriculture and Technology) — Cabbage White Butterflies, UV perception; David Powell (University of Maryland) — Flamingos; Mitch Reardon (Photo Researchers, New York/Okapia Bild-Archiv, Germany) — African Elephants; Juan C. Reboreda (Universidad de Buenos Aires) — Greater Rheas; Caitlin Reed (University of North Carolina/Cambridge University) — Crested Black Macaques; H. D. Rijksen (Institute of Forestry and Nature Research [IBN-DLO]/Golden Ark Foundation, Netherlands) — Orang-utans; Leonard Lee Rue III/Len Rue Jr. (Leonard Rue Enterprises, New Jersey) — White-tailed Deer, Bighorn Sheep; Susan Savage-Rumbaugh (Language Research Center, Georgia State University) — Bonobos; Carolien J. Scholten (Emmen Zoo, Netherlands) — Humboldt Penguins; John W. Scott/John P. Scott (Bowling Green State University, Ohio) — Sage Grouse; Paul E. Simonds (University of Oregon) — Bonnet Macaques; L. H. Smith (Australia) — Superb Lyrebirds; Judie Steenberg (Woodland Park Zoological Gardens, Washington) — Tree Kangaroos; Elizabeth Stevens (Zoo Atlanta/Disney World Animal Programs) — Flamingos; Yukimaru Sugiyama (Primate Research Institute, Kyoto University) — Bonnet Macaques; Angelika Tipler-Schlager (Austria) — Greylag Geese; Pepper W. Trail (Oregon) — Guianan Cock-of-the-Rock; Paul L. Vasey (Universite de Montreal) — Japanese Macaques, other species; Frans B. M. de Waal (Emory University/Yerkes Regional Primate Research Center, Georgia) — Bonobos; Juichi Yamagiwa (Kyoto University) — Gorillas. Thanks also to the photo archives of Yellowstone National Park and the American Museum of Natural History for supplying images, and to the many publishers, journal editors, and scientists for permission to reprint previously published photographs (see photo credits on pp. 733-735).

I am profoundly grateful to Michael Denneny, not only for his editorial acumen and invaluable insights, but also for championing this project with unflagging enthusiasm and personal devotion. This book would simply not have come into being without his guiding hand at its helm. I would also like to acknowledge the many other people at St. Martin's Press who worked on this project, including: Robert Cloud, Helene Berinsky, Steven Boldt, and Sarah Rutigliano. A heartfelt thanks as well to Natasha Kern and Oriana Green, who believed in this book from the very, very beginning and helped steer it through the (sometimes treacherous) waters of the publishing industry. I am also grateful to Robert Jones and Eric Steel for their early support of this project.
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A special note of appreciation goes to John Megahan (Museum of Zoology, University of Michigan), whose superb drawings turned my vision of this book into a reality. John brought nearly two hundred animals to life with accuracy, aplomb, and an exuberant visual style, all the while weathering the numerous pressures of this project with grace and good cheer. John would like to thank his wife, Anne, for her invaluable support and feedback during this project. Many thanks also to the other individuals and organizations who contributed illustrations or assisted in the preparation of visual materials: Stuart Kenter and Tom McCarthy (Stuart Kenter Associates) — photo research and permissions; Gary Antonetti and Jon Daugherity (Ortelius Design) — cartography; Phyllis Wood (Phyllis Wood Associates Scientific Illustration) — icon design; Turid Owren — legal services; and the Guild of Natural Science Illustrators.

I would also like to thank the librarians, information specialists, and other staff at the following libraries, who guided me through numerous arcane bibliographic and electronic searches and helped me navigate their collections: University of Washington libraries, University of British Columbia libraries, Simon Fraser University Library, University of California — Los Angeles libraries (including the Rare Books collection of the biomedical library), University of Hawaii libraries, National Marine Mammal Laboratory Library, Woodland Park Zoological Gardens Library, Vancouver Public Aquarium Library, Seattle Public Library, Vancouver Public Library. Many rare journals, books, monographs, dissertations, and technical reports were not available at these institutions and were obtained primarily through the interlibrary loan division of the Seattle Public Library. Thanks to the hardworking staff in this division, and to the following institutions and organizations for loaning or otherwise making available from their library collections rare or hard-to-find items: California Academy of Sciences, Evergreen State College, Humboldt State University, Idaho State University, Montana State University, Oregon Regional Primate Research Center, Portland State University, The Royal Australasian Ornithologists' Union, University of Alaska, University of Alberta, University of California — Davis, University of California — Irvine, University of California — Santa Cruz, University of Kansas, University of Minnesota, University of Montana, University of Oregon, University of Texas — Austin, University of Utah, University of Victoria, University of Wisconsin — Milwaukee, University of Wyoming, U.S. Fish and Wildlife Service Northern Prairie Wildlife Research Center, Vancouver Public Library, Washington State University. For help with translations of scientific articles, thanks to Sergei V. Mihailov (Russian), John R. Van Son (Dutch), and Courtney Searles-Ridge (German).

Finally, I am truly indebted to the many other people who provided personal support, encouragement, feedback, commiseration, and inspiration over the many years that this book was in gestation (and labor!), including Dawn Bates, Nicola Bessell, Thom Feild, Neal Graves, Ed Kaplan, Clara Ma, Nathan Ohren, Jackal Plumb, Michael Rochemont, and Liza White. Thank you most deeply, Nicola, for holding my hand as I leapt into the void ... .
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