Biological Exuberance

Stonewall Inn Editions

Chapter One

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. HALDANE

In dimly lit undergrowth of a Central American rain forest,jewel-like male hummingbirds flit through the vegetation, pausing briefly tomate now with a male, now with a female. A whale glides through the dark and icywaters of the Arctic, then surges toward the surface in a playful frenzy ofchurning water and splashing, her fins and tail caressing another female.Drifting off to sleep, two male monkeys lie gently in each other's arms, cradledby one of the ancient jungles of Asia. A herd of deer picks its way cautiouslythrough a semidesert scrub of Texas, each animal simultaneously male butnot-quite-male, with half-developed, velvety antlers and diminutive, fine-bonedproportions. In a protected New Zealand inlet, a pair of female gulls--mated forlife--tend their chicks together. Tiny midges swarm above a bleak tundra ofnorthern Europe, a whirlwind of mating activity as males couple with each otherin midair. Circling and prancing around her partner, a female antelope courtsanother female in an ageless, elegant ritual staged on the African savanna.

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

On every continent, animals of the same sex seek each other out and haveprobably been doing so for millions of years. They court each other, usingintricate and beautiful mating dances that are the result of eons of evolution.Males caress and kiss each other, showing tenderness and affection toward oneanother rather than just hostility and aggression. Females form long-lastingpair-bonds--or maybe just meet briefly for sex, rolling in passionate embracesor mounting one another. Animals of the same sex build nests and homestogether, and many homosexual pairs raise young without members of the oppositesex. Other animals regularly have partners of both sexes, and some even live incommunal groups where sexual activity is common among all members, male andfemale. Many creatures are "transgendered," crossing or combiningcharacteristics of both males and females in their appearance or behavior. Amidthis incredible variety of different patterns, one thing is certain: the animalkingdom is most definitely not just heterosexual.

Homosexual behavior occurs in more than 450 different kinds of animalsworldwide, and is found in every major geographic region and every major animalgroup. It should come as no surprise, then, that animal homosexuality is not asingle, uniform phenomenon. Whether one is discussing the forms it takes, itsfrequency, or its relationship to heterosexual activity, same-sex behavior inanimals exhibits every conceivable variation. This chapter presents a broadoverview of animal homosexuality and places it in the context of a number ofother phenomena involving alternative genders and sexualities.

The Many Forms of Animal Homosexuality

For most people, "homosexuality" means one thing: sex. While it's true thatanimals of the same gender often interact sexually with each other, this isonly one aspect of same-sex expression. Animal homosexuality represents a vastand diverse range of activities: it is neither a monolithic nor an exclusivelysexual phenomenon. This section offers a survey of the full range of homosexualactivity found in the animal world, organized around five major behavioralcategories: courtship, affection, sex, pair-bonding, and parenting. While thesecategories are not mutually exclusive and often blend imperceptibly into oneanother, they offer a useful introduction to the multiplicity of homosexualexpression in the animal kingdom.

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

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

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

Same-sex courtship assumes a dizzying array of forms, and zoologists oftenuse evocative or colorful names as the technical terms to designate these moststriking of animal behaviors (which are usually part of heterosexualinteractions as well). Many species perform elaborate dances or kineticdisplays, such as the "strutting" of female Sage Grouse, who spread theirfanlike tails; or the spectacular acrobatics and plumage displays of Birds ofParadise and Superb Lyrebirds; or the courtship encounters of Cavies, who"rumba," "rumble," "rump," and "rear" each other in an alliterative panoply ofchoreographed behaviors. In other cases, subtler poses, stylized postures, ormovements are used, such as the foreleg kicking found in the courtship displaysof many hoofed mammals; "rear-end flirtation" in male Nilgiri Langurs andCrested Black Macaques; ritual preening and bowing during courtshipinteractions in Penguins; "tilting" and "begging" postures in Black-billedMagpies; "jerking" by female Koalas; and "courtship feeding"--a ritual exchangeof food gifts seen in same-sex (and opposite-sex) interactions among Antbirds,Black-headed and Laughing Gulls, Pukeko, and Eastern Bluebirds. Sometimes twocourting individuals perform mutual or synchronized displays, such as the"triumph ceremonies" of male Greylag Geese and Black Swans; the "mutualecstatic" 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 ofGalahs. Many birds have breathtaking aerial displays, including tandem flyingin Griffon Vultures, shuttle displays and "dive-bombing" in Anna'sHummingbirds, "hover-flying" in Black-billed Magpies, "song-dancing" inGreenshanks, and the "bumblebee flight" of Red Bishop Birds.

Animals sometimes exploit specific spatial and environmental elements intheir 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 RegentBowerbirds, and the traditional group or communal display areas known as leksfound in animals as diverse as Kob antelopes, Long-tailed Hermit Hummingbirds,and Ruffs. In other species, dramatic chases that may cover great distances arepart of same-sex interactions: aerial pursuits occur in Greenshanks, GoldenPlovers, Bank Swallows, and Chaffinches; ground chases take place duringcourtships in Mule Deer, Cheetahs, Whiptail Wallabies, and Redshanks; aquaticpursuits occur in Australian Shelducks; while Black-billed Magpies combine bothground and aerial pursuit in their courtship behavior known aschase-hopping. Perhaps most amazing of all are the light-related displays of anumber of bird species, which are designed to utilize specific properties ofsunlight or other luminosity in the bird's environment. GuiananCock-of-the-Rock, for example, position their leks and courtship displays inspecial "light environments" that maximize the visibility of the birds througha sophisticated interaction of the ambient light, the reflectance andcoloration of the bird's own (brilliant orange) plumage, and the forestgeometry in which they are located. Anna's Hummingbirds precisely orient thetrajectory of their stunning aerial climbs and dives to face into the sun,thereby showing off their iridescent plumage to its best. As a male swoopstoward the object of his attentions (either male or female), he resembles abrilliant glowing ember that grows in intensity as he gets closer. To advertisetheir presence on the lek, male Buff-breasted Sandpipers perform a wing-raisingdisplay that exploits the midnight sun of their arctic habitat. Seen from adistance, the brilliant white underwings of each bird flash momentarily againstthe dull tundra background, reflecting the weak late-night sunlight and therebycreating a luminous semaphore that attracts other birds, both male and female,to their territories.

In addition to spectacular visual displays, homosexual courtship--like thecorresponding heterosexual behaviors--can involve a veritable cacophony ofdifferent sounds. Female Kob antelopes whistle, male Gorillas pant, femaleRufous Rat Kangaroos growl, male Blackbuck antelopes bark, female Koalasbellow, male Ocellated Antbirds carol, female Squirrel Monkeys purr, and maleLions moan and hum. The "snap-hiss" ceremonial calls of Black-crowned NightHerons, the croaking of male Moose, "geckering" and "snirking" of female RedFoxes, the chirp-squeaks of male West Indian Manatees, "yip-purr" calls ofHammerheads, 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 maleCaribou, and pulsive scream-calls in Bowhead Whales are just some of thevocalizations heard during same-sex courtship and related interactions.Sometimes pairs of birds execute synchronized vocal displays, as in the duetsof rolling calls performed by Greylag gander pairs, or the precisely syncopated"moo" calling of pairs of male Calfbirds. In a few cases, courtship activitiesinvolve nonvocal sounds or sounds produced in unusual ways. Male GuiananCock-of-the-Rock, Ruffed Grouse, Victoria's Riflebirds, and Red Bishop Birds,for example, make distinctive whistling, drumming, or clapping sounds bybeating or fanning their wings (which in some cases have specially modified,sound-producing feathers), while male Anna's Hummingbirds produce a shrillpopping sound as a result of air passing through their tail feathers duringdisplay flights. Some of the most extraordinary sounds during same-sexcourtship 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 repertoireof courtship splashing sounds made by kicking their feet during displaysvariously named the paddle-kick, plonk-kick, and whistle-kick. Finally, someDolphins appear to engage in a sort of sonic "foreplay": male Atlantic SpottedDolphins have been observed stimulating their partner's genitals with pulsedsound waves, using a type of vocalization known as a genital buzz.

In most species the same courtship behaviors are used in both homosexual andheterosexual interactions. Sometimes, however, same-sex courtship involves onlya subset of the movements and behaviors found in opposite-sex displays. Forexample, when Canada Geese court each other homosexually, they perform aneck-dipping ritual also found in heterosexual courtships, but do not adopt thespecial posture that males and females use after mating. In animals like theWestern Gull or Kob antelope, individuals vary as to how many courtshipbehaviors they use in same-sex interactions. Some exhibit only one or two ofthe typical courtship postures and movements, while others go through theentire elaborate courtship sequence. Perhaps most interesting are thosecreatures that have a special courtship pattern found only in homosexualinteractions. Male Ostriches, for example, perform a unique "pirouette dance"only when courting other males, while female Rhesus Macaques engage incourtship 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 overtlysexual (they do not involve direct contact of the genitals) but that donevertheless have clear sexual or erotic overtones. These are referred to asaffectionate activities and are found in nearly a quarter of theanimals in which some form of homosexual activity occurs. Although many ofthese behaviors (grooming, embracing, play-fighting) can occur in othercontexts, their erotic nature in a same-sex context is usually obvious: the twoanimals may be visibly sexually aroused, the behavior may directly precede orfollow homosexual copulation or courtship, or the affectionate activity mayoccur 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 suchas Gray-headed Flying Foxes and Vampire Bats engage in erotic same-sex groomingand licking; male Mountain Sheep rub their horns and faces on other males,sometimes becoming sexually aroused; Whales and Dolphins stroke and rub eachother with their flippers or tail flukes, as well as rub bodies together; whilenumerous primates such as Apes, Macaques, and Baboons frequently caress andgroom each other in both sexual and nonsexual contexts. A few birds such asHumboldt Penguins, Pukeko, Black-billed Magpies, and Parrots also indulge inpreening--the avian equivalent of grooming--in their homosexual interactions orpair-bonds.

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

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

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

Many other types of affectionate and contactual behaviors occur betweenanimals of the same sex. Sometimes animals gently bite, nibble, or chew on eachother's ears (female Hoary Marmots), or wings and chests (Gray-headed FlyingFoxes), or rumps (male Dwarf Cavies), or necks (male Savanna Baboons). MaleAfrican Elephants intertwine their trunks, while female Japanese Macaquessometimes suck each other's nipples, and male Crested Black Macaques andSavanna Baboons affectionately pat or grab other males' rear ends. Pairs ofanimals may sit, huddle, or lie together in close proximity, sometimes touchinghands 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 hispartner's hands resting on his loins. Male Lions and female Long-earedHedgehogs slide the lengths of their bodies along their partner's, while maleBowhead Whales, Killer Whales, and Gray Seals roll their bodies over eachother, and same-sex companions in Gray Whales and Botos swim side by side whilegently touching each other with their fins.

Some animals have developed unique forms of touching that combine severaldifferent types of affectionate activities along with courtship and sexualbehaviors. Male Giraffes engage in "necking", a multifaceted activity thatincorporates elements of play-fighting, courtship, and sexuality, in which theyrub their necks along each other's body while also licking, sniffing, andbecoming sexually aroused by one another. In Giraffes and other species, thesetypes of activities sometimes involve multiple animals interactingsimultaneously in near "orgies" of bodily contact. Spinner Dolphins, forexample, participate in "wuzzles"--group sessions of mutual caressing andsexual activity (both same-sex and opposite-sex)--while West Indian Manateeshave a similar sort of "free-for-all" group activity known as cavorting, whichcan involve rubbing, chasing, and sexual interactions, among many otheractivities. Among birds, Hammerheads, Acorn Woodpeckers, and Blue-belliedRollers have ritualized bouts of courtship and mounting activity that mayinvolve groups of individuals and both same-sex and opposite-sex partners. Thedistinctive and, in many cases, unabashedly sensual and playful aspects of someof these activities are aptly reflected in the descriptive names given to themby zoologists. In fact, the term wuzzle--though used as a technicaldesignation for this behavior in the scientific literature--is actually anonsense word coined by a marine biologist, whose whimsical "etymology" for thename could be right out of Lewis Carroll: "The term comes from W. E. Schevillof Woods Hole Oceanographic Institution, who, when asked what the behavior was,replied without hesitation, `Why, it looks like a wuzzle to me.'"

Mounting, Diddling, and Bump-Rumping: Sexual Techniques

Affectionate activity often leads to, or is inseparable from, overtly sexualbehavior--defined here as any contact between two or more animals involvinggenital stimulation. Stumptail and Crab-eating Macaques, for example, kisstheir same-sex partners during sexual mounting. In fact, mounting is the mostcommon type of sexual behavior found in homosexual contexts: one animal climbson top of the other in a position similar to heterosexual intercourse, usuallyfrom behind in a front-to-back position (that is, one animal mounted on theback of the other). More than 95 percent of mammal and bird species use thisposition, for both male and female homosexual interactions. On the other hand,some animals--particularly primates such as Gorillas, Bonobos, and White-handedGibbons--use a face-to-face position (in addition to, or instead of), and insome cases this is more common in homosexual encounters than in heterosexualones. Belly-to-belly copulation is also the norm for both homosexual andheterosexual 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 sometimesinteract in a supine or semirecumbent position, one individual behind the otherwith her partner between her legs or sitting "in her lap" (which may also bedone in a face-to-face position). Occasionally female Warthogs, Rhesus andJapanese Macaques, Koalas, and Takhi mount their female partner from the siderather than from behind; lateral mounts also sometimes occur duringheterosexual 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 onlytwo individuals at a time, but group sexual (and courtship) activity--involvinganywhere 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 maleanal intercourse occurs in some species (for example, Orang-utans, RhesusMacaques, Bison, and Bighorn rams), while female penetration of various typesoccurs during lesbian interactions in Orang-utans (insertion of the finger intothe vagina), Bonobos (insertion of the erect clitoris into the vulva), andBottlenose and Spinner Dolphins (insertion of a fin or tail fluke into thefemale's genital slit). Simple pelvic thrusting and rubbing of the genitals onthe rump of the other animal is widespread in both male and female homosexualmounts (occurring in the Northern Fur Seal, Lion, and Proboscis Monkey, amongothers), and simple genital-to-genital touching is the form of homosexual (andheterosexual) contact in species where males do not have a penis (as in mostbirds, such as the Pukeko and Tree Swallow). A more unusual type of malehomosexual contact involves various forms of non-anal penetration. InWhales and Dolphins, both males and females have a genital slit or opening; whennot aroused, the male's penis is contained in the cavity leading to this slit.Homosexual activity in Bowhead Whales, Bottlenose Dolphins, and Botos sometimesinvolves 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 Botosoccasionally insert the penis into a male partner's blowhole (on the top of hishead!), while male Orang-utans have even been observed retracting their penis toform a sort of "hollow" or concavity that another male can penetrate. Clitoralrubbing 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 theirpenises together or on each other's body. In male Bonobos, mutual genitalrubbing sometimes takes the form of an activity with the colorful name of "penisfencing," in which the males hang suspended by their arms and rub their erectorgans against each other.

Oral sex of various kinds also occurs in a number of species. This mayinvolve 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 Thin-horn Sheep and Vampire Bats; genital licking in femaleSpotted Hyenas and male Cheetahs); mouthing, nuzzling, or "kissing" of genitals(female Gorillas, male Savanna Baboons, Crab-eating Macaques, and West IndianManatees); and genital sniffing in female Pronghorns and Marmots as well asscrotal sniffing in Whiptail and Red-necked Wallabies. Male Stumptail Macaqueseven perform mutual fellatio in a sixty-nine position, while males of a numberof primate species (including Gibbons, Bonnet and Crested Black Macaques, andNilgiri Langurs) sometimes actually eat or swallow their partner's (or theirown) semen--though usually after mutual genital rubbing or manual stimulationrather than oral sex. Dwarf Cavies and Rufous Bettongs occasionally indulge inanal licking, nuzzling, and sniffing with same-sex (and opposite-sex) partners.Another sort of "oral" sexual activity is called beak-genital propulsion andoccurs among both male and female Bottlenose and Spinner Dolphins: one animalinserts its snout or "beak" into the genital slit of another, simultaneouslystimulating and propelling its partner forward while swimming (a similarbehavior in Orcas, involving simple nuzzling or touching of the genitals withthe snout, is known as beak-genital orientation).

Another type of activity found during homosexual interactions ismasturbation, in which one animal stimulates its own or its partner's genitalswith a finger, hand, foot, flipper, or some other appendage. For example, maleSavanna Baboons often touch, grab, or fondle the genitals of another male--thisbehavior is known aptly as diddling--while male Bottlenose Dolphins and WestIndian Manatees sometimes rub another male's penis with their flippers. MaleRhesus and Crested Black Macaques, female Gorillas, male Vampire Bats, femaleProboscis Monkeys, and male Walruses sometimes masturbate themselves whenmounting, courting, or interacting sexually with another animal of the samesex. Mutual masturbation in a side-by-side sixty-nine position occurs in femaleCrested Black Macaques, while male Bonnet and Stumptail Macaques masturbateeach other and even fondle one another's scrotums. Another form of mutualmasturbation in these species involves two males backing up toward each otherand fondling each other's genitals between their legs. In Bonobos and CommonChimpanzees, individuals often rub their anal and genital regions togetherwhile in this rump-to-rump position, prompting zoologists to give thesebehaviors names like "rump-rubbing" and "bump-rump." Other more unusual formsof "manual" stimulation include mutual genital stimulation using trunks infemale Elephants, and anal stimulation and penetration with fingers by maleCommon 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 same sex.Homosexual pair-bonding takes many different forms, but two broad categoriescan be recognized: "partners," who engage in sexual or courtship activities witheach other, and "companions," who are bonded to each other but do not necessarilyengage in overt sexual activity with one another. More than a third of themammals and birds in which homosexual activity occurs have at least one ofthese types of same-sex bonding. The archetypal example of a "partnership" isthe mated pair: two individuals who are strongly bonded to one another in a waythat is equivalent to heterosexually paired animals of the same species.Partners engage directly in courtship, sexual, and/or parenting behaviors; theyusually spend a significant amount of time with each other; and they do similaractivities together. This is found primarily in birds (more than 70 differentspecies)--not surprisingly, since heterosexual pairing is typical of featheredcreatures (but generally rare in other animal groups). Examples of homosexualmates are found in male Black Swans and Black-headed Gulls, and femaleBlack-winged Stilts and Silver Gulls (among many others). In mammals,partnerships take many different forms, including "consortships" in femaleRhesus and Japanese Macaques, "sexual friendships" in Stumptail and Crab-eatingMacaques, "tending bonds" between male Bison, and "coalitions" between maleBonnet Macaques, Savanna Baboons, and Cheetahs. Some animals, while notnecessarily forming same-sex bonds, do have "preferred" or "favorite" sexualand affectionate partners with whom they tend to interact more often than withothers: this is true for Bonobos, Gorillas, Killer Whales, and Dwarf Cavies,among others.

Many forms of same-sex partnership are exclusive or monogamous, and partnersmay even actively defend their pair-bond against the intrusion of outsideindividuals (for instance in male Gorillas, female Japanese Macaques, and maleLions). Animals of the same sex sometimes also compete with each other for theattentions of homosexual partners, as in male Gorillas and Blue-winged Teals;female Orangutans, Japanese Macaques, and Orange-fronted Parakeets may evencompete with males for "preferred" female partners. Some partnerships,however, are "open" or nonmonogamous: female Bonobos and Rhesus Macaques, forinstance, may have sexual relations with several different "favorite" partnersor consorts (of both sexes). Males in homosexual pairs of Greylag Geese,Laughing Gulls, Humboldt Penguins, and Flamingos sometimes engage in"promiscuous" copulations with birds (male or female) other than their mate(heterosexual pairs in these species are also sometimes nonmonogamous). Anotherform of nonmonogamy occurs among lesbian pairs in a number of Gulls and otherbirds: one or both females sometimes mate with a male (while still maintainingtheir same-sex bond) and are thereby able to fertilize their eggs and becomeparents.

The second main type of homosexual pairing is the "companionship." Twoanimals of the same sex may bond with each other, often spending most of theirtime together exclusive of the opposite sex, but they do not necessarily engagein recognizable courtship or sexual activities with each other. For example,older African Elephant bulls sometimes form long-lasting associations with ayounger "attendant" male: these animals are loners, spending all their timewith each other rather than with other Elephants, helping each other, and neverengaging in heterosexual activity. Male Calfbird companions display and traveltogether and also sometimes share a "home" with one another (a special perchknown as a retreat where they spend time away from the display court). Similarsame-sex associations are found in many other species, including Orang-utans,Gray Whales, Grizzly Bears, Vampire Bats, and Superb Lyrebirds. Youngersame-sex attendants are known as satellites in male Moose and shadows in maleWalruses, while companions are called duos in male Hanuman Langurs andspinsters in female Warthogs--the latter is something of a misnomer, though,since Warthog companions do occasionally participate in sexual activity withmales or females, but not necessarily with their companions.

Sometimes more than two animals bond together, forming a "trio" (in eitherpartnership or companionship form). This arrangement can consist of threeanimals all of the same sex who are bonded with each other, as occasionallyhappens among female Ring-billed Gulls and male African Elephants, White-tailedDeer, and Black-headed Gulls. Trios can also be bisexual, consisting of twofemales and one male (e.g., Canada Geese, Common Gulls, and Jackdaws) or twomales and one female (Greylag Geese, Black Swans, Sociable Weavers); inOystercatchers, both types occur. In either form of a bisexual trio, there issignificant bonding, courtship, and/or sexual behavior between the two animalsof the same sex. This distinguishes such associations from heterosexual trios,in which two animals of the same sex are bonded with an opposite-sexedindividual but not to each other. Same-sex trios of closely bonded male GreylagGeese or female Grizzly Bears are also sometimes known as triumvirates, whilebisexual (and heterosexual) trios in Flamingos are called triads. In a fewspecies, "quartets" involving simultaneous homosexual and heterosexual bondsbetween four individuals sometimes occur: in Greylag Geese and Black-headedGulls, for instance, three males and a female sometimes bond with each other,while in Galahs, two males and two females may associate in a quartet withvarious bonding arrangements between them.

Homosexual pair-bonds vary not only in their type, but also in theirduration. Same-sex bonding often follows the species-typical pattern forheterosexual pairing in terms of how long it lasts. In species such as theGreylag Goose, for example, which remain mated for life (or else for manyconsecutive years), male pairs are also generally long-lasting or lifelong,while in Bison, tending bonds usually last only a few days or hours in bothheterosexual and homosexual situations. In some cases, long-term pair bondinginvolves continuous association throughout the year, as among male OcellatedAntbirds. This contrasts with seasonal association, for example among severalspecies of Gulls, in which females re-pair with the same female only during themating season. Homosexual pairs may also be of shorter duration thanheterosexual ones in some species: Black-headed Gull male couples, for example,appear to be more prone to divorce than heterosexual ones. However, in manycases homosexual pairings, particularly companionships, actually exceedheterosexual ones in their stability and duration. Among Lions and Elephants,for example, the bond between male companions is closer and longer-lasting thanany heterosexual bonds (which, in these and many other species, are virtuallynonexistent beyond mating), while mated gander pairs in Greylag Geese are oftenmore strongly bonded than heterosexual pairs. Consortships between JapaneseMacaque females sometimes develop into yearlong friendships, unlike themajority of heterosexual associations in this species. In fact, in a number ofanimals the only pair-bonds that occur are homosexual, not heterosexual. MaleBottlenose Dolphins, for example, form lifelong partnerships with each other,while males and females in this species do not generally pair-bond with oneanother at all. Other animals with same-sex but not opposite-sex pairings(often in the form of companionships) include Musk-oxen, Wapiti, White-tailedDeer, Warthogs, Cheetahs, Eastern Gray Kangaroos, Red Squirrels, and Calfbirds.

Formidable Fathers and Supernormal Mothers: Homosexual Parenting

Same-sex pairs in many species (especially birds) raise young together. Notonly are they competent parents, homosexual pairs sometimes actually exceedheterosexual ones in the number of eggs they lay, the size of their nests, orthe skill and extent of their parenting. How are such animals able to haveoffspring in the first place if they are in homosexual associations? Manydifferent strategies are used, including several in which one or both partnersare the biological parent(s) of the young they raise together. The most commonparenting arrangement of this type is found in lesbian pairs of several Gull,Tern, and Goose species: one or both female partners copulate with a male tofertilize her eggs. No bonding or long-term association develops between thefemale and the male (who is essentially a "sperm donor" to the homosexualpair), and the youngsters are then jointly raised by both females without anyassistance from a male parent. Because female birds can lay eggs regardless ofwhether they are fertilized, however, each partner in a lesbian pair usuallycontributes a full clutch of eggs to their nest even if she hasn't mated with amale. As a result, female homosexual pairs often lay what are calledsupernormal clutches, that is, double the number of eggs usually found in nestsof heterosexual pairs.

Sometimes two female animals who already have offspring join forces, bondingtogether and raising their young as a same-sex family unit (among mammals,female coparents may even suckle each other's young), this occurs in GrizzlyBears, Red Foxes, Warthogs, Dwarf Cavies, Lesser Scaup Ducks, and Sage Grouse.Notably, heterosexual pairs do not occur in these species, and most offspringare otherwise raised by single females. In some species, a nonbreeding animalbonds with a (single) breeding animal and helps parent its young: this occursin Squirrel Monkeys, Northern Elephant Seals, Jackdaws (where a widowed femalewith young may pair with a single female), and Greater Rheas (where one malemay help another incubate his eggs and then raise the young together). In mostsuch joint parenting arrangements (as opposed to homosexual mated pairs), thereis not necessarily any overt courtship or sexual activity between the bondedcoparents, although in some species (e.g., Squirrel Monkeys, Northern ElephantSeals, Emus, Sage Grouse), homosexual activity does occur in contextsother than between coparents. Still other birds (e.g., Greylag Geese,Common Gulls, Oystercatchers) may form bisexual parenting trios, mating withthe opposite-sexed partner(s) in their association while maintaining homosexualand heterosexual bonds simultaneously, with all three birds then raising theresulting offspring together. A variation on this arrangement in Black Swansinvolves a sort of "surrogate motherhood": established male homosexual pairssometimes associate temporarily with a female, mating with her to father theirown offspring. Once the eggs are laid, however, they chase her away and raisethe cygnets on their own as a homosexual couple.

In a number of cases, homosexual pairs raise young without being thebiological parents of the offspring they care for. Some same-sex pairs adoptyoung: two female Northern Elephant Seals occasionally adopt and coparent anorphaned pup, while male Hooded Warblers and Black-headed Gulls may adopt eggsor entire nests that have been abandoned by females, and pairs of male Cheetahsoccasionally look after lost cubs. Sometimes female birds "donate" eggs tohomosexual couples through a process known as parasitism: in many birds,females lay eggs in nests other than their own, leaving the parenting duties tothe "host" couple. This occurs both within the same species, and (morecommonly) across species, and usually involves heterosexual hosts. Male pairsof Hooded Warblers, however, sometimes receive eggs from Brown-headed Cowbirds(and possibly also from females of their own species) in this way;within-species parasitism may also provide eggs for male pairs of Black-headedGulls and female pairs in Roseate and Caspian Terns. The opposite situation isthought to occur in Ring-billed Gulls: researchers believe that somehomosexually paired females actually lay eggs in nests belonging toheterosexual pairs. Finally, some birds in same-sex pairs take over or "kidnap"nests from heterosexual pairs (e.g., in Black Swans, Flamingos) or occasionally"steal" individual eggs (e.g., in Caspian and Roseate Terns, Black-headedGulls); homosexual pairs in captivity also raise foster young provided to them.

In a detailed study of parental behavior by female pairs of Ring-billedGulls, scientists found no significant differences in quality of care providedby homosexual as opposed to heterosexual parents. They concluded that there wasnot anything that male Ring-billed Gull parents provided that two females couldnot offer equally well. This case is not exceptional: homosexual parents aregenerally as good at parenting as heterosexual ones. Examples of same-sex pairssuccessfully raising young have been documented in at least 20 species, and ina few cases, homosexual couples actually appear to have an advantage overheterosexual ones. Pairs of male Black Swans, for example, are often able toacquire the largest and best-quality territories for raising young because oftheir combined strength. Such fathers--dubbed "formidable" adversaries by onescientist--consequently tend to be more successful at raising offspring thanmost heterosexual pairs. And in many species in which single parenting is therule (because there is no heterosexual pair-bonding), same-sex pairs provide aunique opportunity for young to be raised by two parents (e.g., SquirrelMonkeys, Grizzly Bears, Lesser Scaup Ducks). Moreover, in some Gulls, femalepairs are consigned (for a variety of reasons) to less than optimalterritories, yet they still successfully raise young: in many cases theycompensate by investing more parental effort--and are more dutiful in caringfor their chicks--than male-female pairs. There are exceptions, of course: somefemale pairs of Gulls, for instance, tend to lay smaller eggs and raise fewerchicks (although this is also true of heterosexual trios attending supernormalclutches), while same-sex parents in Jackdaws, Canada Geese, and Oystercatchersmay experience parenting difficulties such as egg breakage ornonsynchronization of incubation duties. By and large, though, same-sex couplesare competent and occasionally even superior parents.

Birds in homosexual pairs often build a nest together. Usually theyconstruct a single nest the way most heterosexual pairs do, but othervariations also occur: female Common Gulls and Jackdaws sometimes make "twin"or "joint" nests containing two cups in the same bowl, while male Greater Rheasand female Canada Geese may use "double" nests consisting of two adjacent ortouching nests. Female Mute Swans occasionally construct two separate nests inwhich both birds lay eggs. Nests belonging to male couples in some species(e.g., Flamingos and Great Cormorants) are often impressive structures,exceeding the size of heterosexual nests because both males contribute equallyto their construction (in heterosexual pairs of these species, usually only onesex builds the nest, or males and females make unequal contributions). Manysame-sex pairs construct nests regardless of whether they lay fertile eggs.Male pairs of Mute Swans, Flamingos, Black-crowned Night Herons, and GreatCormorants, for example, usually build nests even though they never acquireeggs, and the male "parents" may even sit on the nests as if they containedeggs, while female pairs frequently build nests in which they lay supernormalclutches that are entirely infertile. Same-sex parents often share incubationduties, either taking turns sitting on their nest (the most commonarrangement), or else incubating simultaneously on a single nest (femaleRed-backed Shrikes, male Emus) or side by side on a twin or double nest (femaleJackdaws, male Greater Rheas).

In addition to parenting by homosexual couples, some animals raise young inalternative family arrangements, usually a group of several males or femalesliving together. Gorilla babies, for example, grow up in mixed-sex, polygamousgroups where their mothers may have lesbian interactions with each other, whilePukeko and Acorn Woodpeckers live and raise their young in communal breedinggroups where many, if not all, group members engage in courtship and sexualactivities with one another (both same-sex and opposite-sex). In suchsituations, individuals that engage in homosexual courtship or copulationactivities may either reproduce directly because they also mate heterosexually(Pukeko), or they may assist members of their group in raising young withoutreproducing themselves (Acorn Woodpeckers). Other alternative familyconstellations include bisexual trios (mentioned above), homosexual trios (asin Grizzly Bears, Dwarf Cavies, Lesser Scaup Ducks, and Ring-billed Gulls)where three mothers jointly parent their offspring, and even quartets, in whichfour animals of the same (Grizzlies) or both sexes (Greylag Geese) are bondedto each other and all raise their young together.

Finally, some animals that have homosexual interactions are "single parents."Many female mammals, for example, that court or mate with other females alsomate heterosexually and raise the resulting young on their own or in female-onlygroups (as is typical for exclusively heterosexual females in the same speciesas well). This is especially prevalent among mammals with polygamous orpromiscuous heterosexual mating systems, such as Kob and Pronghorn antelopes andNorthern Fur Seals (where males, and sometimes females, usually mate with morethan one partner). Males in many polygamous species are often bisexual as well,fathering offspring in addition to courting or mating with other males;typically, however, they do not actively parent their offspring regardless ofwhether they are bisexual or exclusively heterosexual.

What's Good for the Goose...:
Comparisons of Male and Female Homosexuality

Is homosexuality more characteristic of male animals or female animals? And doesit assume different forms in the two sexes--or, to paraphrase a popular saying,is the behavior of the "goose" essentially similar to that of the "gander"? Asit so happens, homosexuality in three species of Geese--Canada, Snow, andGreylag--exemplifies some of the major patterns of male and femalehomosexuality and the range of variation found throughout the rest of theanimal world. In Canada Geese, both males and females participate in the samebasic type of homosexual activity, forming same-sex pairs and engaging in somecourtship activities. Within these same-sex bonds, however, there are genderdifferences in some less common behaviors: sexual activity is morecharacteristic of females (especially if they are part of a bisexual trio), asis nest-building and parenting activity. There are also differences in thefrequency of participation of the two sexes: although same-sex pairs arerelatively common, accounting for more than 10 percent of pairs in somepopulations, a greater proportion of the male population participates insame-sex pairing. In contrast, homosexual activity in Snow Geese is vastlydifferent in males than in females, although it is relatively infrequent inboth sexes. Females form long-lasting pair-bonds with other females in whichsexual activity is not necessarily very prominent, although parenting activityis: both partners lay eggs in a joint nest and raise their young together (theyfertilize their eggs by mating with males). Ganders, on the other hand, limittheir homosexual activity to same-sex mounting of other males duringheterosexual group rape attempts and do not form same-sex pairs (althoughinterspecies gander pairs with Canada Geese sometimes do occur). Finally, inGreylag Geese homosexual activity is found exclusively in males, who formgander pairs that engage in a variety of courtship, sexual, pair-bonding, andparenting activities.

When we look at the full range of species and behaviors, we find that malehomosexuality is slightly more prevalent, overall, than female homosexuality,although the two are fairly close. Same-sex activity (of all forms) occurs inmale mammals and birds in about 80 percent of the species in whichhomosexuality has been observed, and between females in just over 55 percent ofthese (the figures add up to more than 100 percent because both maleand female homosexuality are found in some species). It must also be kept inmind that the prevalence of female homosexuality may actually be greater thanthese figures indicate, but has simply not been documented as systematicallyowing to the general male bias of many biological studies. There is alsovariation between different animal subgroupings: in carnivores, marsupials,waterfowl, and shorebirds, for example, male and female homosexuality arealmost equally common (in terms of the number of species in which each isfound), while in marine mammals and perching birds male homosexuality is moreprevalent. And in many species same-sex activity occurs only among males (e.g.,Boto, a freshwater dolphin) or only among females (e.g., Puku, an Africanantelope).

The frequency of same-sex behavior in males versus females can also beassessed within a given species, and once again, many different patterns arefound: in Rhesus Macaques, Hamadryas and Gelada Baboons, and Tasmanian NativeHens, for example, 80-90 percent of all same-sex mounting is between males,while homosexual activity is also more prevalent among male Gray-headed FlyingFoxes. In other species, female homosexual activity assumes prominence:more than 70 percent of same-sex copulations in Pukeko are between females,and 70-80 percent of homosexual activity in Bonobos is lesbian. Females accountfor almost two-thirds of same-sex behaviors in Stumptail Macaques and RedDeer, while homosexual activity is also more typical of females in Red-neckedWallabies and Northern Quolls. In some species, however, male homosexualityis so predominant that same-sex activity in females is often missed byscientific observers or rarely mentioned (e.g., Giraffes, Blackbuck, BighornSheep), while the reverse is true in other species (e.g., Hanuman Langurs,Herring Gulls, Silver Gulls). In contrast, Pig-tailed Macaque same-sexmounting, Galah pair-bonds, and Pronghorn homosexual interactions are fairlyequally distributed between the two sexes (although actual same-sex mounting ismore common in male Pronghorns).

As with the species of Geese mentioned above, gender differences are alsoapparent in various behavior types. Of those mammal and bird species in whichsome form of homosexual behavior occurs, each of the activities of courtship,affectionate, sexual, or pair-bonding are generally more prevalent in maleanimals. They occur among males in 75-95 percent of the species in which theyare found, while among females these activities occur in 50-70 percent of thespecies (again, however, the possible gender bias of the studies these figuresare based on must be kept in mind). The one exception is same-sex parenting,which is performed by females in more than 80 percent of the species where thisbehavior occurs, but by males in just over half of the species that have someform of such parenting. Of course, not all these forms of same-sex interactionalways co-occur in the same species, and animals sometimes differ as to whichactivities males as opposed to females of the same species tend to participatein (as in the Geese). In Silver and Herring Gulls, for example, females formsame-sex pairs that undertake parenting duties while males engage in homosexualmounting; in Cheetahs and Lions, both sexes engage in sexual activity, butmales in each species also develop same-sex pair-bonds while female Cheetahsparticipate in same-sex courtship activities. In Ruffs, males engage in sexual,courtship, and (occasional) pairing activity with each other, while Reeves (thename for females of this sandpiper species) participate primarily in sexualactivity with one another.

Within each of the categories of courtship, sexual, pairing, and parentingbehaviors, further gender distinctions can be drawn. Consider various types ofsexual behavior. Mounting as a same-sex activity is ubiquitous and occursfairly regularly in both males and females (although there are exceptions--inAfrican Elephants, for example, sexual activity between males assumes the formof mounting while female same-sex interactions consist of mutual masturbation).Oral sex (which includes activities as diverse as fellatio, cunnilingus,genital nuzzling and sniffing, and beak-genital propulsion) is about equallyprevalent in both sexes. Group sexual activity is more common in males (onlyoccurring among females in 6 species, including Bonobos and Sage Grouse), asare interactions between adults and adolescents (only occurring among femalesin 9 species, including Hanuman Langurs, Japanese Macaques, Ring-billed Gulls,and Jackdaws, but among males in more than 70 species). Although penetration isalso more typical of male homosexual interactions, there are notable exceptions(e.g., Bonobos, Orang-utans, and Dolphins, as mentioned previously). Genderdifferences sometimes also manifest themselves in the minutiae of varioussexual acts. Same-sex mounting in Gorillas, for instance, is performed in bothface-to-face and front-to-back positions, but the two sexes differ in thefrequency with which these two positions are used: females prefer theface-to-face position, adopting it in the majority of their sexualinteractions, while males use it less often, in only about 17 percent of theirhomosexual mounting episodes. In contrast, the frequency of full genitalcontact during homosexual copulations is nearly identical for both sexes ofPukeko: females achieve cloacal contact in about 23 percent of their same-sexmounts while males do so in about 25 percent of theirs (in comparison, genitalcontact occurs in a third to half of all heterosexual mounts). Among Flamingos,though, genital contact is more characteristic of copulations between femalesthan between males.

Or consider pair-bonding and parenting. Stable, long-lasting pair-bonds aregenerally not more characteristic of females (contrary to what one mightinitially expect); mated pairs or partnerships are almost equally common inboth sexes (in terms of number of species in which homosexual pair-bondingoccurs), while same-sex companionships are more prevalent between males.Likewise, long-term pair-bonds are just as likely to be found between males asfemales, while nonmonogamy and divorce occur in male and female couples inroughly equal numbers of species. Nor are male couples less successful parents:male coparents or partners are not overrepresented among the few species inwhich same-sex parents occasionally experience parenting difficulties. One areawhere a gender difference in same-sex parenting does manifest itself is in theway that homosexual pairs "acquire" offspring. Particularly among birds, femalecouples can raise their own offspring by simply having one or both partnersmate with males without interrupting their homosexual pair-bond. This option isnot as widely available for male couples, who usually father their ownoffspring by forming a longer-lasting (prior or simultaneous) association witha female (as in Black Swans, Greylag Geese, and Greater Rheas).

Japanese Macaques offer a particularly compelling example of the multipleways that homosexual activity can differ between males and females. Althoughhomosexual mounting occurs in both sexes in this species, males and femalesdiffer in the specific details of their sexual interactions. Homosexual mountsare usually initiated by the mounter between males but by the mountee betweenfemales, make use of a wider variety of mounting positions between females, areaccompanied by a unique vocalization only between males, and involve pelvicthrusting and multiplemounts more often between females than between males. Thetwo sexes also differ in their partner selection and pair-bonding activities:females generally form strongly bonded consortships and have fewer partners thanmales, while the latter tend to interact sexually with more individuals anddevelop less intense bonds (although some do have "preferred" male partners).Finally, there is a seasonal difference in male as opposed to female same-sexinteractions: homosexual mounting is more common outside the breeding season inmales but during the breeding season in females, while same-sex bonds infemales, but not males, may extend into yearlong associations that transcend thebreeding season.

Whether we're talking about ganders and geese or Ruffs and Reeves, whetherit's Botos and Bonobos or Pukus and Pukeko, male and female homosexuality can beeither surprisingly similar to each other or decidedly distinctive from oneanother. In any case, a complex intersection of factors is involved in theexpression of homosexuality in each gender. As with other aspects of animalhomosexuality, preconceived ideas about how males and females act must bereassessed and refined when considering the full range of animal behaviors. Insome species such as Silver Gulls, male and female homosexualities conform tostereotypes commonly held about similar human behaviors: females form stable,long-lasting lesbian pairbonds and raise families while males participate inpromiscuous homosexual activity. In other species these gender stereotypes areturned completely on their heads, as in Black Swans, where only males formlong-term same-sex couples and raise offspring, and Sage Grouse, where onlyfemales engage in group "orgies" of homosexual activity. And in the majority ofcases, male and female homosexualities present their own unique blends ofbehaviors and characteristics that defy any simplistic categorization--such asBonobos, where sexual penetration occurs in female rather than male same-sexactivity, where sexual interactions between adults and adolescents are aprominent feature of female interactions, and where males do not form stronglybonded relationships with each other the way females do, but engage in lesshomosexual activity overall and more affectionate activity such as openmouthedkissing. Once again, the diversity of animal homosexuality reveals itself downto the very last detail of expression.

Continues...
Excerpted from Biological Exuberanceby Bruce Bagemihl Copyright © 2000 by Bruce Bagemihl. Excerpted by permission.
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