Biological Exuberance: Animal Homosexuality and Natural Diversity

ISBN-13: 9780312253776

Media Type: Paperback

Publisher: St. Martin's Publishing Group

Publication Date: 04-10-2000

Pages: 768

Product Dimensions: 6.00(w) x 9.00(h) x 1.70(d)

Series: Stonewall Inn Editions (Paperback)

Bruce Bagemihl, Ph.D., is a biologist and researcher who has served on the faculty of the University of British Columbia, where he taught linguistics and cognitive science. He is the author of Biological Exuberance and has published diverse essays and scientific articles on issues pertaining to language, biology, gender, and sexuality. He lives in Seattle, Washington.

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Biological Exuberance

Part 1

A Polysexual, Polygendered World

Chapter 1

The Birds and the Bees

The universe is not only queerer than we suppose, it is queerer than we can suppose.

—evolutionary biologist J. B. s. HALDANE1

In the dimly lit undergrowth of a Central American rain forest, jewel-like male hummingbirds flit through the vegetation, pausing briefly to mate now with a male, now with a female. A whale glides through the dark and icy waters of the Arctic, then surges toward the surface in a playful frenzy of churning water and splashing, her fins and tail caressing another female. Drifting off to sleep, two male monkeys lie gently in each other's arms, cradled by one of the ancient jungles of Asia. A herd of deer picks its way cautiously through a semidesert scrub of Texas, each animal simultaneously male but not-quite-male, with half-developed, velvety antlers and diminutive, fine-boned proportions. In a protected New Zealand inlet, a pair of female gulls—mated for life—tend their chicks together. Tiny midges swarm above a bleak tundra of northern Europe, a whirlwind of mating activity as males couple with each other in midair. Circling and prancing around her partner, a female antelope courts another female in an ageless, elegant ritual staged on the African savanna.

Although biologist J. B. S. Haldane was not (necessarily) referring to homosexuality when he spoke of the "queerness" of the natural world, little did he know how accurate his statement would turn out to be. The world is, indeed, teeming with homosexual, bisexual, and transgendered creatures of every stripe and feather. From the Southeastern Blueberry Bee of the United States to more than 130 different bird species worldwide, the "birds and the bees," literally, are queer.2

On every continent, animals of the same sex seek each other out and have probably been doing so for millions of years.3 They court each other, using intricate and beautiful mating dances that are the result of eons of evolution. Males caress and kiss each other, showing tenderness and affection toward one another rather than just hostility and aggression. Females form long-lasting pair-bonds—or maybe just meet briefly for sex, rolling in passionate embraces or mounting one another. Animals of the same sex build nests and homes together, and many homosexual pairs raise young without members of the opposite sex. Other animals regularly have partners of both sexes, and some even live in communal groups where sexual activity is common among all members, male and female. Many creatures are "transgendered," crossing or combining characteristics of both males and females in their appearance or behavior. Amid this incredible variety of different patterns, one thing is certain: the animal kingdom is most definitely not just heterosexual.

Homosexual behavior occurs in more than 450 different kinds of animals worldwide, and is found in every major geographic region and every major animal group.4 It should come as no surprise, then, that animal homosexuality is not a single, uniform phenomenon. Whether one is discussing the forms it takes, its frequency, or its relationship to heterosexual activity, same-sex behavior in animals exhibits every conceivable variation. This chapter presents a broad overview of animal homosexuality and places it in the context of a number of other phenomena involving alternative genders and sexualities.

The Many Forms of Animal Homosexuality

For most people, "homosexuality" means one thing: sex. While it's true that animals of the same gender often interact sexually with each other, this is only one aspect of same-sex expression. Animal homosexuality represents a vast and diverse range of activities: it is neither a monolithic nor an exclusively sexual phenomenon. This section offers a survey of the full range of homosexual activity found in the animal world, organized around five major behavioral categories: courtship, affection, sex, pair-bonding, and parenting. While these categories are not mutually exclusive and often blend imperceptibly into one another, they offer a useful introduction to the multiplicity of homosexual expression in the animal kingdom.

A word on terminology is in order. In this book, heterosexuality is defined as courtship, affectionate, sexual, pair-bonding, and/or parenting behaviors between animals of the opposite sex, while homosexuality is defined as these same activities when they occur between animals of the same sex. When applied to people, the terms homosexual, gay, or lesbian can refer either to a particular behavior when it occurs between two men or two women, or to an individual whose primary "identity" involves any or all of these activities. Since the notion of identity is inappropriate to ascribe to animals, these terms will be reserved for the behaviors that animals engage in and, where relevant, to describe individuals whose primary "orientation" is toward animals of the same sex where courtship, sexual, and/or pair-bonding activities are concerned. In addition, because the terms gay and lesbian have particularly human connotations, these will generally be avoided in favor of homosexual(ity) or same-sex (although it must be remembered that each of these words can have specific meanings independent of their human connotations when used in relation to animals, and they are employed as cover terms for widely divergent activities even among humans). When a particular individual engages in both homosexual and heterosexual activity, these words are limited to describing the animal'sspecific behaviors (depending on the gender of the animal's partner), while the animal itself is described as bisexual.5

Pirouette Dances, Ecstatic Displays, and Triumph Ceremonies: Courtship Patterns

To attract the attention and interest of a potential partner, animals often perform a series of stylized movements and behaviors prior to mating, sometimes in the form of a complex visual or vocal display. This is known as courtship behavior, and it usually indicates that one animal is advertising his or her presence to prospective mates or is sexually interested in another individual. If the interest is mutual, this may lead to mating or other sexual activity and possibly pair-bonding (for example, in birds). Some animals also use special courtship behaviors to conclude, as well as initiate, sexual activity, or to reinforce their pair-bonds. Courtship behavior is a common feature of homosexual interactions, occurring in nearly 40 percent of the mammals and birds in which same-sex activity has been observed.

Same-sex courtship assumes a dizzying array of forms, and zoologists often use evocative or colorful names as the technical terms to designate these most striking of animal behaviors (which are usually part of heterosexual interactions as well). Many species perform elaborate dances or kinetic displays, such as the "strutting" of female Sage Grouse, who spread their fanlike tails; or the spectacular acrobatics and plumage displays of Birds of Paradise and Superb Lyrebirds; or the courtship encounters of Cavies, who "rumba," "rumble," "rump," and "rear" each other in an alliterative panoply of choreographed behaviors. In other cases, subtler poses, stylized postures, or movements are used, such as the foreleg kicking found in the courtship displays of many hoofed mammals; "rear-end flirtation" in male Nilgiri Langurs and Crested Black Macaques; ritual preening and bowing during courtship interactions in Penguins; "tilting" and "begging" postures in Black-billed Magpies; "jerking" by female Koalas; and "courtship feeding"—a ritual exchange of food gifts seen in same-sex (and opposite-sex) interactions among Antbirds, Black-headed and Laughing Gulls, Pukeko, and Eastern Bluebirds. Sometimes two courting individuals perform mutual or synchronized displays, such as the "triumph ceremonies" of male Greylag Geese and Black Swans; the "mutual ecstatic" and "dabbling" displays of Humboldt and King Penguins, respectively; synchronous aquatic spiraling in male Harbor Seals and Orcas; the elaborate "leapfrogging" and "Catherine wheel" courtship displays by groups of Manakins; and synchronized wing-stretching and head-bobbing in homosexual pairs of Galahs. Many birds have breathtaking aerial displays, including tandem flying in Griffon Vultures, shuttle displays and "dive-bombing" in Anna's Hummingbirds, "hover-flying" in Black-billed Magpies, "song-dancing" in Greenshanks, and the "bumblebee flight" of Red Bishop Birds.

Animals sometimes exploit specific spatial and environmental elements in their courtship activities as well. Special display courts are used in same-sex (and opposite-sex) interactions in many species, including the "drumming logs" of male Ruffed Grouse, the elaborate architectural creations of Regent Bowerbirds, and the traditional group or communal display areas known as leks found in animals as diverse as Kob antelopes, Long-tailed Hermit Hummingbirds, and Ruffs. In other species, dramatic chases that may cover great distances are part of same-sex interactions: aerial pursuits occur in Greenshanks, Golden Plovers, Bank Swallows, and Chaffinches; ground chases take place during courtships in Mule Deer, Cheetahs, Whiptail Wallabies, and Redshanks; aquatic pursuits occur in Australian Shelducks; while Black-billed Magpies combine both ground and aerial pursuit in their courtship behavior known as chase-hopping. Perhaps most amazing of all are the light-related displays of a number of bird species, which are designed to utilize specific properties of sunlight or other luminosity in the bird's environment. Guianan Cock-of-the-Rock, for example, position their leks and courtship displays in special "light environments" that maximize the visibility of the birds through a sophisticated interaction of the ambient light, the reflectance and coloration of the bird's own (brilliant orange) plumage, and the forest geometry in which they are located. Anna's Hummingbirds precisely orient the trajectory of their stunning aerial climbs and dives to face into the sun, thereby showing off their iridescent plumage to its best. As a male swoops toward the object of his attentions (either male or female), he resembles a brilliant glowing ember that grows in intensity as he gets closer. To advertise their presence on the lek, male Buff-breasted Sandpipers perform a wing-raising display that exploits the midnight sun of their arctic habitat. Seen from a distance, the brilliant white underwings of each bird flash momentarily against the dull tundra background, reflecting the weak late-night sunlight and thereby creating a luminous semaphore that attracts other birds, both male and female, to their territories.6

In addition to spectacular visual displays, homosexual courtship—like the corresponding heterosexual behaviors—can involve a veritable cacophony of different sounds. Female Kob antelopes whistle, male Gorillas pant, female Rufous Rat Kangaroosgrowl, male Blackbuck antelopes bark, female Koalas bellow, male Ocellated Antbirds carol, female Squirrel Monkeys purr, and male Lions moan and hum. The "snap-hiss" ceremonial calls of Black-crowned Night Herons, the croaking of male Moose, "geckering" and "snirking" of female Red Foxes, the chirp-squeaks of male West Indian Manatees, "yip-purr" calls of Hammerheads, the yelping and babble-singing of Black-billed Magpies, "lip-smacking" in several Macaque species, the humming call of Pukeko, "stutters" and "chirps" of male Cheetahs, the "vacuum-slurping" of male Caribou, and pulsive scream-calls in Bowhead Whales are just some of the vocalizations heard during same-sex courtship and related interactions. Sometimes pairs of birds execute synchronized vocal displays, as in the duets of rolling calls performed by Greylag gander pairs, or the precisely syncopated "moo" calling of pairs of male Calfbirds. In a few cases, courtship activities involve nonvocal sounds or sounds produced in unusual ways. Male Guianan Cock-of-the-Rock, Ruffed Grouse, Victoria's Riflebirds, and Red Bishop Birds, for example, make distinctive whistling, drumming, or clapping sounds by beating or fanning their wings (which in some cases have specially modified, sound-producing feathers), while male Anna's Hummingbirds produce a shrill popping sound as a result of air passing through their tail feathers during display flights. Some of the most extraordinary sounds during same-sex courtship are made by aquatic animals: Walruses generate eerily metallic "bell" sounds by striking special throat pouches with their flippers and castanet-like "knocks" by chattering their teeth, while Musk Ducks have an entire repertoire of courtship splashing sounds made by kicking their feet during displays variously named the paddle-kick, plonk-kick, and whistle-kick. Finally, some Dolphins appear to engage in a sort of sonic "foreplay": male Atlantic Spotted Dolphins have been observed stimulating their partner's genitals with pulsed sound waves, using a type of vocalization known as a genital buzz.

In most species the same courtship behaviors are used in both homosexual and heterosexual interactions. Sometimes, however, same-sex courtship involves only a subset of the movements and behaviors found in opposite-sex displays. For example, when Canada Geese court each other homosexually, they perform a neckdipping ritual also found in heterosexual courtships, but do not adopt the special posture that males and females use after mating. In animals like the Western Gull or Kob antelope, individuals vary as to how many courtship behaviors they use in same-sex interactions. Some exhibit only one or two of the typical courtship postures and movements, while others go through the entire elaborate courtship sequence. Perhaps most interesting are those creatures that have a special courtship pattern found only in homosexual interactions. Male Ostriches, for example, perform a unique "pirouette dance" only when courting other males, while female Rhesus Macaques engage in courtship games such as "hide-and-seek" that are unique to lesbian interactions.

Kisses, Wuzzles, and Necking: Affectionate Behaviors

Many animals of the same sex touch each other in ways that are not overtly sexual (they do not involve direct contact of the genitals) but that do nevertheless haveclear sexual or erotic overtones. These are referred to as affectionate activities and are found in nearly a quarter of the animals in which some form of homosexual activity occurs. Although many of these behaviors (grooming, embracing, play-fighting) can occur in other contexts, their erotic nature in a same-sex context is usually obvious: the two animals may be visibly sexually aroused, the behavior may directly precede or follow homosexual copulation or courtship, or the affectionate activity may occur in a same-sex pair-bond.

One type of affectionate activity is simple grooming or rubbing. Male Lions "head-rub" and roll around with each other before having sex together; Bats such as Gray-headed Flying Foxes and Vampire Bats engage in erotic same-sex grooming and licking; male Mountain Sheep rub their horns and faces on other males, sometimes becoming sexually aroused; Whales and Dolphins stroke and rub each other with their flippers or tail flukes, as well as rub bodies together; while numerous primates such as Apes, Macaques, and Baboons frequently caress and groom each other in both sexual and nonsexual contexts. A few birds such as Humboldt Penguins, Pukeko, Black-billed Magpies, and Parrots also indulge in preening—the avian equivalent of grooming—in their homosexual interactions or pair-bonds.

Some animals also "kiss" each other: male African Elephants, female Rhesus Macaques, male West Indian Manatees and Walruses, female Hoary Marmots, and male Mountain Zebras (among others) all touch mouths, noses, or muzzles during their homosexual encounters. Even some birds, such as Black-billed Magpies, engage in mutual beak-nibbling or "billing" as part of same-sex courtship. In primates, kissing (in both homosexual and heterosexual contexts) can bear a startling resemblance to the corresponding human activity: a number of species such as Squirrel Monkeys and Common Chimpanzees engage in full mouth-to-mouth contact, while male Bonobos kiss each other with "passionate" openmouthed kisses with considerable mutual tongue stimulation.

Numerous species of Monkeys and Apes also "hug" or embrace same-sex partners in homosexual contexts (usually face-to-face, although male Bonobos and Vervets also embrace while standing in a front-to-back position). Among non-primates, female Bottlenose Dolphins clasp each other during homosexual activity, male West Indian Manatees embrace one another underwater, while Gray-headed Flying Foxes wrap their wing-membranes around same-sex partners while stimulating each other. A striking form of same-sex embracing is the "sleeping huddle" found in Stumptail and Bonnet Macaques: a pair of males often sleep together in a front-to-back position, one male wrapping his arms around the other and sometimes even holding on to his partner's penis. A similar sleeping arrangement occurs, surprisingly, among male Walruses, who often sleep in same-sex pairs or extended "chains" of males, all clasping each other in a front-to-back position as they float at the water's surface.

A number of mammals also engage in mock battles or "play-fights" that have erotic overtones. Although they superficially resemble aggressive behavior, these "battles" or "contests" do not involve any physical violence and are clearly distinguished from actual cases of aggressive or territorial behavior in these species. Male African Elephants, for example, frequently become sexually aroused and develop erections when they perform ritualized erotic jousting matches, while numerous hoofed mammals such as male Giraffes, Bison, Blackbuck antelopes, and Mule Deer mount each other during play-fights or ritualistic jousting. Among primates such as Orang-utans, Gibbons, and Proboscis Monkeys, males sometimes engage in playful wrestling matches that can develop into sexual encounters, while male Australian and New Zealand Sea Lions also indulge in play-fighting combined with same-sex mounting. Although play-fighting is most common among male mammals, female Cheetahs sometimes engage in "mock fighting" with each other as part of same-sex courtship sequences, while female (and male) Galahs and Orange-fronted Parakeets in same-sex pairs have playful "fencing bouts" with their bills.7

Many other types of affectionate and contactual behaviors occur between animals of the same sex. Sometimes animals gently bite, nibble, or chew on each other's ears (female Hoary Marmots), or wings and chests (Gray-headed Flying Foxes), or rumps (male Dwarf Cavies), or necks (male Savanna Baboons). Male African Elephants intertwine their trunks, while female Japanese Macaques sometimes suck each other's nipples, and male Crested Black Macaques and Savanna Baboons affectionately pat or grab other males' rear ends. Pairs of animals may sit, huddle, or lie together in close proximity, sometimes touching hands or putting an arm around the shoulder (female Gorillas, Squirrel Monkeys, and Japanese Macaques, male Siamangs), while male Hanuman Langurs "cuddle" together by sitting back-to-front, one male between the other's legs with his partner's hands resting on his loins. Male Lions and female Long-eared Hedgehogs slide the lengths of their bodies along their partner's, while male Bowhead Whales, Killer Whales, and Gray Seals roll their bodies over each other, and same-sex companions in Gray Whales and Botos swim side by side while gently touching each other with their fins.

Some animals have developed unique forms of touching that combine several different types of affectionate activities along with courtship and sexual behaviors. Male Giraffes engage in "necking", a multifaceted activity that incorporates elements of play-fighting, courtship, and sexuality, in which they rub their necks along each other's body while also licking, sniffing, and becoming sexually aroused by one another. In Giraffes and other species, these types of activities sometimes involvemultiple animals interacting simultaneously in near "orgies" of bodily contact. Spinner Dolphins, for example, participate in "wuzzles"—group sessions of mutual caressing and sexual activity (both same-sex and opposite-sex)—while West Indian Manatees have a similar sort of "free-for-all" group activity known as cavorting, which can involve rubbing, chasing, and sexual interactions, among many other activities. Among birds, Hammerheads, Acorn Woodpeckers, and Blue-bellied Rollers have ritualized bouts of courtship and mounting activity that may involve groups of individuals and both same-sex and opposite-sex partners. The distinctive and, in many cases, unabashedly sensual and playful aspects of some of these activities are aptly reflected in the descriptive names given to them by zoologists. In fact, the term wuzzle—though used as a technical designation for this behavior in the scientific literature—is actually a nonsense word coined by a marine biologist, whose whimsical "etymology" for the name could be right out of Lewis Carroll: "The term comes from W. E. Schevill of Woods Hole Oceanographic Institution, who, when asked what the behavior was, replied without hesitation, 'Why, it looks like a wuzzle to me.'"8

Mounting, Diddling, and Bump-Rumping: Sexual Techniques

Affectionate activity often leads to, or is inseparable from, overtly sexual behavior—defined here as any contact between two or more animals involving genital stimulation. Stumptail and Crab-eating Macaques, for example, kiss their same-sex partners during sexual mounting. In fact, mounting is the most common type of sexual behavior found in homosexual contexts: one animal climbs on top of the other in a position similar to heterosexual intercourse, usually from behind in a front-to-back position (that is, one animal mounted on the back of the other). More than 95 percent of mammal and bird species use this position, for both male and female homosexual interactions. On the other hand, some animals—particularly primates such as Gorillas, Bonobos, and White-handed Gibbons—use a face-to-face position (in addition to, or instead of), and in some cases this is more common in homosexual encounters than in heterosexual ones. Belly-to-belly copulation is also the norm for both homosexual and heterosexual interactions in Dolphins. Occasionally more unusual or "creative" mounting positions are used, particularly by female animals. In Bonobos, Stumptail Macaques, and Japanese Macaques, for instance, females sometimes interact in a supine or semirecumbent position, one individual behind the other with her partner between her legs or sitting "in her lap" (which may also be done in a face-to-face position). Occasionally female Warthogs, Rhesus and Japanese Macaques, Koalas, and Takhi mount their female partner from the side rather than from behind; lateral mounts also sometimes occur during heterosexual interactions in these (and other) species. And in some animals a "backward," head-to-tail mounting position is occasionally used, e.g., in Botos, Hammerheads, Ruffs, and Western Gulls. Most same-sex interactions involve only two individuals at a time, but group sexual (and courtship) activity—involving anywhere from three or four (Giraffes, Lions) to six or more (Bowhead Whales, Mountain Sheep) partners—occurs in over 25 different species.

The actual type of genital contact varies widely. Full penetration in male anal intercourse occurs in some species (for example, Orang-utans, Rhesus Macaques, Bison, and Bighorn rams), while female penetration of various types occurs during lesbian interactions in Orang-utans (insertion of the finger into the vagina), Bonobos (insertion of the erect clitoris into the vulva), and Bottlenose and Spinner Dolphins (insertion of a fin or tail fluke into the female's genital slit). Simple pelvic thrusting and rubbing of the genitals on the rump of the other animal is widespread in both male and female homosexual mounts (occurring in the Northern Fur Seal, Lion, and Proboscis Monkey, among others), and simple genital-to-genital touching is the form of homosexual (and heterosexual) contact in species where males do not have a penis (as in most birds, such as the Pukeko and Tree Swallow). A more unusual type of male homosexual contact involves various forms of non-anal penetration. In Whales and Dolphins, both males and females have a genital slit or opening; when not aroused, the male's penis is contained in the cavity leading to this slit. Homosexual activity in Bowhead Whales, Bottlenose Dolphins, and Botos sometimes involves insertion of the penis of one male into the genital slit of the other. Other more unusual forms of penetration have also been documented: male Botos occasionally insert the penis into a male partner's blowhole (on the top of his head!), while male Orang-utans have even been observed retracting their penis to form a sort of "hollow" or concavity that another male can penetrate. Clitoral rubbing or other types of genital tribadism are found in female Bonobos, Gorillas, and Rhesus Macaques (among others), while males in several species (e.g., White-handed Gibbons, West Indian Manatees, and Gray Whales) rub their penises together or on each other's body. In male Bonobos, mutual genital rubbing sometimes takes the form of an activity with the colorful name of "penis fencing," in which the males hang suspended by their arms and rub their erect organs against each other.

Oral sex of various kinds also occurs in a number of species. This may involve actual sucking of genitals (fellatio between males in Bonobos, Orang-utans, Siamangs, and Stumptail Macaques); licking of genitals (cunnilingus in Common Chimpanzees, Long-eared Hedgehogs, and Kob antelopes; penis-licking in Thinhorn Sheep and Vampire Bats; genital licking in female Spotted Hyenas and male Cheetahs); mouthing, nuzzling, or "kissing" of genitals (female Gorillas, male Savanna Baboons, Crab-eating Macaques, and West Indian Manatees); and genital sniffing in female Pronghorns and Marmots as well as scrotal sniffing in Whiptail and Red-necked Wallabies. Male Stumptail Macaques even perform mutual fellatio in a sixty-nine position, while males of a number of primate species (including Gibbons, Bonnet and Crested Black Macaques, and Nilgiri Langurs) sometimes actually eat or swallow their partner's (or their own) semen—though usually after mutual genital rubbing or manual stimulation rather than oral sex.9 Dwarf Cavies and Rufous Bettongs occasionally indulge in anal licking, nuzzling, and sniffing with same-sex (and opposite-sex) partners. Another sort of "oral" sexual activity is called beak-genital propulsion and occurs among both male and female Bottlenose and Spinner Dolphins: one animal inserts its snout or "beak" into the genital slit of another, simultaneously stimulating and propelling its partner forward while swimming (a similar behavior in Orcas, involving simple nuzzling or touching of the genitals with the snout, is known as beak-genital orientation).

Another type of activity found during homosexual interactions is masturbation, in which one animal stimulates its own or its partner's genitals with a finger, hand, foot, flipper, or some other appendage. For example, male Savanna Baboons often touch, grab, or fondle the genitals of another male—this behavior is known aptly as diddling—while male Bottlenose Dolphins and West Indian Manatees sometimes rub another male's penis with their flippers. Male Rhesus and Crested Black Macaques, female Gorillas, male Vampire Bats, female Proboscis Monkeys, and male Walruses sometimes masturbate themselves when mounting, courting, or interacting sexually with another animal of the same sex. Mutual masturbation in a side-by-side sixty-nine position occurs in female Crested Black Macaques, while male Bonnet and Stumptail Macaques masturbate each other and even fondle one another's scrotums. Another form of mutual masturbation in these species involves two males backing up toward each other and fondling each other's genitals between their legs. In Bonobos and Common Chimpanzees, individuals often rub their anal and genital regions together while in this rump-to-rump position, prompting zoologists to give these behaviors names like "rump-rubbing" and "bump-rump." Other more unusual forms of "manual" stimulation include mutual genital stimulation using trunks in female Elephants, and anal stimulation and penetration with fingers by male Common Chimpanzees, Siamangs, and Crab-eating Macaques.

Consorts, Satellites, and Triumvirates: Same-Sex Mates and Pair-Bonding

Wild animals often form significant pair-bonds with animals of the