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Sexual behavior among non human species that is interpreted as homosexual
homosexuality in humans

Various non human animal species exhibit behavior that can be interpreted as homosexual or bisexual. This may include same sex sexual activity, courtship, affection, pair bonding, and parenting among same sex animal pairs. Various forms of this are found in every major geographic region and every major animal group. The sexual behavior of non human animals takes many different forms, even within the same species, though homosexual behavior is best known from social species. Scientists perceive homosexual behavior in animals to different degrees. A 2019 paper states that same sex sexual behavior has been observed in over 1,500 species. A previous 1999 book by Canadian biologist Bruce Bagemihl states same sex behavior (comprising courtship, sexual, pair bonding, and parental activities) has been documented in over 450 species of animals worldwide. Although same sex interactions involving genital contact have been reported in hundreds of animal species, they are routinely manifested in only a few, including humans. Simon LeVay stated that "[a]lthough homosexual behavior is very common in the animal world, it seems to be very uncommon that individual animals have a long lasting predisposition to engage in such behavior to the exclusion of heterosexual activities. Thus, a homosexual orientation, if one can speak of such thing in animals, seems to be a rarity." The motivations for and implications of these behaviors are lensed through anthropocentric thinking; Bagemihl notes that any hypothesis is "necessarily an account of human interpretations of these phenomena". One species in which exclusive homosexual orientation occurs is the domesticated sheep (Ovis aries). In October 2023, biologists reported studies of animals (over 1,500 different species) that found same sex behavior (not necessarily related to human orientation) may help improve social stability by reducing conflict within the groups studied.

The term homosexual was coined by the Hungarian writer and campaigner Karl Maria Kertbeny in 1868 to describe same sex sexual attraction and sexual behavior in humans. Its use in animal studies has been controversial for two main reasons: animal sexuality and motivating factors have been and remain poorly understood, and the term has strong cultural implications in western society that are irrelevant for species other than humans. Thus homosexual behavior has been given a number of terms over the years. According to Bruce Bagemihl, when describing animals, the term homosexual is preferred over gay, lesbian, and other terms currently in use, as these are seen as even more bound to human homosexuality. applying the term homosexuality to all sexual behavior (copulation, genital stimulation, mating games and sexual display behavior) between animals of the same sex. In most instances, it is presumed that the homosexual behavior is but part of the animal's overall sexual behavioral repertoire, making the animal "bisexual" rather than "homosexual" as the terms are commonly understood in humans.

A majority of the research available concerning homosexual behavior in animals lacks specification between animals that exclusively exhibit same sex tendencies and those that participate in heterosexual and homosexual mating activities interchangeably. This lack of distinction has led to differing opinions and conflicting interpretations of collected data amongst scientists and researchers. For instance, Bruce Bagemihl, author of the book Biological Exuberence: Animal Homosexuality and Natural Diversity, emphasizes that there are no anatomical or endocrinological differences between exclusively homosexual and exclusively heterosexual animal pairs. Information gathered from these studies is limited when applied to spontaneously occurring same sex behavior in animals outside of the laboratory. The Hieroglyphics of Horapollo, written in the 4th century AD by the Egyptian writer Horapollo, mentions "hermaphroditism" in hyenas and homosexuality in partridges. Academic research into the ubiquity of same sex sexual behavior was not carried out on a large scale, possibly due to observer bias caused by social attitudes to same sex sexual behavior, Georgetown University biologist Janet Mann states "Scientists who study the topic are often accused of trying to forward an agenda, and their work can come under greater scrutiny than that of their colleagues who study other topics." They also noted "Not every sexual act has a reproductive function that's true of humans and non humans." but the true extent of homosexuality in animals is not known. Some researchers believe this behavior to have its origin in male social organization and social dominance, similar to the dominance traits shown in prison sexuality. Others, particularly Bagemihl, Joan Roughgarden, Thierry Lodé and Paul Vasey suggest the social function of sex (both homosexual and heterosexual) is not necessarily connected to dominance, but serves to strengthen alliances and social ties within a flock. While reports on many such mating scenarios are still only anecdotal, a growing body of scientific work confirms that permanent homosexuality occurs not only in species with permanent pair bonds, In fact, apparent homosexual individuals are known from all of the traditional domestic species, from sheep, cattle and horses to cats, dogs and budgerigars.

A definite physiological explanation or reason for homosexual activity in animal species has not been agreed upon by researchers in the field. Numerous scholars are of the opinion that varying levels (either higher or lower) of the sex hormones in the animal, in addition to the size of the animal's gonads, (FucM) gene in laboratory mice – which influences the levels of estrogen to which the brain is exposed – caused the female mice to behave as if they were male as they grew up. "The mutant female mouse underwent a slightly altered developmental programme in the brain to resemble the male brain in terms of sexual preference" said professor Chankyu Park of the Korea Advanced Institute of Science and Technology in Daejon, South Korea, who led the research. His findings were published in the BMC Genetics journal on July 7, 2010. Another study found that by manipulating a gene in fruit flies (Drosophila), homosexual behavior appeared to have been induced. However, in addition to homosexual behavior, several abnormal behaviors were also exhibited apparently due to this mutation.

In March 2011, research showed that serotonin is involved in the mechanism of sexual orientation of mice. A study conducted on fruit flies found that inhibiting the dopamine neurotransmitter inhibited lab induced homosexual behavior.

One proposal for the adaptive function of homosexual behavior is the formation of alliances and mutual social benefit to the animals. Studies support this in specific species, such as black swans, where a quarter of mate pairs consist of two males, who mate with a female and chase her away once she lays the egg, then raise it themselves. These M M pairs have great success in defending their territory and resources, and keep their young alive until fledgling 80% of the time, compared to 30% for M F pairs. Studies done on homosexual behavior in birds showed a negative correlation between relative parental investment and F F homosexual behaviors, i. e. females that invested more time and care into their young relative to males had less homosexual encounters. Similarly, there was a negative correlation between relative parental investment and M M homosexual behaviors. This meant that species exhibiting a high degree of polygamy (where females often are the exclusive caretakers of the young) F F sexual behaviors were very rare, whereas in a socially monogamous species (in which a M F pair works together to care for young) they were much more common. The trend was opposite for males, in polygamous species M M sexual behaviors were quite common and in socially monogamous species they were rare. The study argues that release from parental care, a very energy intensive investment, allows the opportunities for homosexual behaviors to be exhibited, and higher parental care prevents homosexual behaviors from occurring because of the energy cost of the behaviors. A 2019 paper hypothesized that when sex first began to evolve, there was no distinction between homosexuality and heterosexuality, and animals mated with other members of their species indiscriminately. This is a contrast to most perspectives, which try to find explanations for the evolution of homosexual behaviors and separate it completely from the evolution of heterosexual behaviors. The study states that it is unlikely that sexual behaviors evolved simultaneously to the evolution of traits necessary to recognize a compatible sexual mate, such as size, shape, odor, and color. As those secondary sex characteristics evolved, sexuality would have become more discriminatory, leading to less homosexuality, but homosexual behaviors would rarely have had enough cost to be selected against and removed entirely from a population. Additionally, the cost of homosexual behavior would be offset by the cost of mate recognition, which requires psychological adaptations, and excessive discrimination in mate choice can lead to missing out of mating opportunities. With indiscriminate mating, these factors are irrelevant. The paper notes that in some species, especially where survival is very difficult and each energy related decision could mean the animal's death, homosexual behavior would be strongly selected against, leading strictly heterosexual species.

An estimated one quarter of all black swan pairings are of males. They steal nests, or form temporary threesomes with females to obtain eggs, driving away the female after she lays the eggs. The males spent time in each other's society, guarded the common territory, performed greeting ceremonies before each other, and (in the reproductive period) pre marital rituals, and if one of the birds tried to sit on the other, an intense fight began. More of their cygnets survive to adulthood than those of different sex pairs, possibly due to their superior ability to defend large portions of land. The same reasoning has been applied to male flamingo pairs raising chicks.

Female Laysan albatrosses, on the north western tip of the island of Oahu, Hawaii, form pairs for co growing offspring. On the observed island, the number of females considerably exceeds the number of males (59% N=102/172), so 31% of females, after mating with males, create partnerships for hatching and feeding chicks. Compared to male female couples, female partnerships have a lower hatching rate (41% vs 87%) and lower overall reproductive success (31% vs. 67%). Warming waters from climate change have led to increased foraging times and thus increased mortality among female black browed albatrosses on the Antipodes Islands in New Zealand. The skewed gender imbalance has led to many male albatrosses forming homosexual relationships. Male male pairs now comprise between 2–5% of the albatross population on the islands.

In 2009, a UK based captive breeding program for blue ducks (involving two males and one female) was derailed when the two males paired with each other instead of with the female that they were assigned to mate with.

Research has shown that the environmental pollutant methylmercury can increase the prevalence of homosexual behavior in male American white ibis. The study involved exposing chicks in varying dosages to the chemical and measuring the degree of homosexual behavior in adulthood. The results discovered was that as the dosage was increased the likelihood of homosexual behavior also increased. The endocrine blocking feature of mercury has been suggested as a possible cause of sexual disruption in other bird species.

Mallards form male female pairs only until the female lays eggs, at which time the male leaves the female. Mallards have rates of male male sexual activity that are unusually high for birds, in some cases, as high as 19% of all pairs in a population.

Penguins have been observed to engage in homosexual behaviour since at least as early as 1911. George Murray Levick, who documented this behaviour in Adélie penguins at Cape Adare, described it as "depraved". The report was considered too shocking for public release at the time, and was suppressed. The only copies that were made available privately to researchers had the English text partly written in Greek letters, to prevent this knowledge becoming more widely known. The report was unearthed only a century later, and published in Polar Record in June 2012. In early February 2004, The New York Times reported that Roy and Silo, a male pair of chinstrap penguins in the Central Park Zoo in New York City, had successfully hatched and fostered a female chick from a fertile egg they had been given to incubate. Other penguins in New York zoos have also been reported to have formed same sex pairs. In Odense Zoo in Denmark, a pair of male king penguins adopted an egg that had been abandoned by a female, proceeding to incubate it and raise the chick. Zoos in Japan and Germany have also documented homosexual male penguin couples. The couples have been shown to build nests together and use a stone as a substitute for an egg. Researchers at Rikkyo University in Tokyo found 20 homosexual pairs at 16 major aquariums and zoos in Japan. The Bremerhaven Zoo in Germany attempted to encourage reproduction of endangered Humboldt penguins by importing females from Sweden and separating three male pairs, but this was unsuccessful. The zoo's director said that the relationships were "too strong" between the homosexual pairs. German gay groups protested at this attempt to break up the male male pairs but the zoo's director was reported as saying "We don't know whether the three male pairs are really homosexual or whether they have just bonded because of a shortage of females nobody here wants to forcibly separate homosexual couples." A pair of male Magellanic penguins who had shared a burrow for six years at the San Francisco Zoo and raised a surrogate chick, split when the male of a pair in the next burrow died and the female sought a new mate. Buddy and Pedro, a pair of male African penguins, were separated by the Toronto Zoo to mate with female penguins. Buddy has since paired off with a female. In 2014 Jumbs and Hurricane, two Humboldt penguins at Wingham Wildlife Park became the center of international media attention as two male penguins who had pair bonded a number of years earlier and then successfully hatched and reared an egg given to them as surrogate parents after the mother abandoned it halfway through incubation. As of 2018, two female King penguins at Kelly Tarltons in Auckland, New Zealand, called Thelma and Louise (named after the 1991 film) have been in a relationship for eight years, when most of the other eligible penguins switch partners each mating season, regardless of their orientation. The two penguins were both taking care of an egg that Thelma hatched, but is unknown whether it was fertilized.

In 1998, two male griffon vultures named Dashik and Yehuda, at the Jerusalem Biblical Zoo, engaged in "open and energetic sex" and built a nest. The keepers provided the couple with an artificial egg, which the two parents took turns incubating, and 45 days later, the zoo replaced the egg with a baby vulture. The two male vultures raised the chick together. A few years later, however, Yehuda became interested in a female vulture that was brought into the aviary. Dashik became depressed, and was eventually moved to the zoological research garden at Tel Aviv University where he too set up a nest with a female vulture. Two male vultures at the Allwetter Zoo in Muenster built a nest together, although they were picked on and their nest materials were often stolen by other vultures. They were eventually separated to try to promote breeding by placing one of them with female vultures, despite the protests of German homosexual groups.

Both male and female pigeons sometimes exhibit homosexual behavior. In addition to sexual behavior, same sex pigeon pairs will build nests, and hens will lay (infertile) eggs and attempt to incubate them.

The Amazon river dolphin or boto has been reported to form up in bands of 3–5 individuals engaging in sexual activity. The groups usually comprise young males and sometimes one or two females. Sex is often performed in non reproductive ways, using snout, flippers and genital rubbing, without regard to gender. The males will sometimes also perform sex with males from the tucuxi species, a type of small porpoise. Also, mounting of one female by another (known as "bulling") is extremely common among cattle. The behaviour is hormone driven and synchronizes with the emergence of estrus (heat), particularly in the presence of a bull.

[[File:Homosexual Fellatio Erect Penis Licking between Male Bonin Flying Foxes Pteropus pselaphon pone.0166024. s002. ogv|thumb|Two male Bonin flying foxes (Pteropus pselaphon) performing fellatio on each other Bat species that have been observed engaging in homosexual behavior in the wild include: These coercive copulations usually include ejaculation and the mounted bat often makes a typical copulation call consisting of a long squawk. Between males, homosexual behaviour includes rubbing of genitals against each other, which sometimes leads to the males swimming belly to belly, inserting the penis in the other's genital slit and sometimes anus. Janet Mann, Georgetown University professor of biology and psychology, argues that the strong personal behavior among male dolphin calves is about bond formation and benefits the species in an evolutionary context. She cites studies showing that these dolphins later in life as adults are in a sense bisexual, and the male bonds forged earlier in life work together for protection as well as locating females to reproduce with. Confrontations between flocks of bottlenose dolphins and the related species Atlantic spotted dolphin will sometimes lead to cross species homosexual behaviour between the males rather than combat.

African and Asian male elephants will engage in same sex bonding and mounting. Such encounters are often associated with affectionate interactions, such as kissing, trunk intertwining, and placing trunks in each other's mouths. Male elephants, who often live apart from the general herd, often form "companionships", consisting of an older individual and one or sometimes two younger males with sexual behavior being an important part of the social dynamic. Unlike heterosexual relations, which are always of a fleeting nature, the relationships between males may last for years. The encounters are analogous to heterosexual bouts, one male often extending his trunk along the other's back and pushing forward with his tusks to signify his intention to mount. Same sex relations are common and frequent in both sexes, with Asiatic elephants in captivity devoting roughly 45% of sexual encounters to same sex activity. In one study, up to 94% of observed mounting incidents took place between two males. The proportion of same sex activities varied between 30 and 75%, and at any given time one in twenty males were engaged in non combative necking behavior with another male. Only 1% of same sex mounting incidents occurred between females.

Homosexual behavior is quite common in wild marmots. A homosexual encounter often begins with a greeting interaction in which one female nuzzles her nose on the other female's cheek or mouth, or both females touch noses or mouths. Additionally, a female may gently chew on the ear or neck of her partner, who responds by raising her tail. The first female may sniff the other's genital region or nuzzle that region with her mouth. She may then proceed to mount the other female, during which the mounting female gently grasps the mounted female's dorsal neck fur in her jaws while thrusting. The mounted female arches her back and holds her tail to one side to facilitate their sexual interaction.

European polecats (Mustela putorius) were found to engage homosexually with non sibling animals. Exclusive homosexuality with mounting and anal penetration in this solitary species serves no apparent adaptive function.

Same sex sexual behaviors have been observed in New World monkeys, Old World monkeys, and apes. Although, these behaviors are much more common in Old World monkeys than New World monkeys, and all ape genera have been observed engaging in same sex sexual behaviors. Prosimians have not been observed engaging in same sex sexual behavior. Same sex mounting, for instance, has been observed in 16% of New World primate genera. In contrast, 76% of Old World monkey genera contain at least one species that has been observed in male male mounts, and 70% of genera for female female mounts. Same sex mounting has been observed in all ape genera. Same sex sexual behaviors likely evolved following the divergence of New World monkeys; these behaviors are a conserved trait among Old World monkeys and apes. Among Old World monkeys, same sex genital interactions are much more common in multi male systems as opposed to uni male systems.

There is one published study of same sex sexual behavior in white handed gibbons (Hylobates lar). The authors report on observations of an adult male mounting an adolescent male. In gibbons, same sex sexual behavior is likely rare.

Both male and female mountain gorillas (Gorilla gorilla beringei) have been observed engaging in same sex sexual behaviors. Homosexual behavior among male gorillas has been studied. This behavior occurs more often in all male bachelor packs in the wild and it is believed to play a role in social bonding. Homosexual behavior among female mountain gorillas has also been documented. Among all male "bachelor" groups, same sex sexual interactions are extremely common. Female mountain gorillas engage in same sex sexual behaviors frequently as well, and coincided in prevalence in accordance with intersexual copulations as well. As opposed to conferring social status, same sex sexual behaviors are likely reflective of elevated arousal in female mountain gorillas. Same sex sexual behaviors are natural and common among mountain gorillas.

Homosexual behavior forms part of the natural repertoire of sexual or sociosexual behavior of orangutans. Male homosexual behavior occurs both in the wild and in captivity, and it occurs in both adolescent and mature individuals. Homosexual behavior in orangutans is not an artifact of captivity or contact with humans. Sociosexual behaviors among orangutans (Pongo spp.) are extremely common and invoke a variety of evolutionary strategies. However, only one study of same sex sexual behavior has been published. In Sumatran orangutans (Pongo abelii), same sex sexual behavior has been documented among two dyads of unflanged adolescent males. Oral contact with genitals and behavior similar to male female forced copulation between sexually mature sub adult males were observed.

Among monkeys, Lionel Tiger and Robin Fox conducted a study on how Depo Provera contraceptives lead to decreased male attraction to females.

With the Japanese macaque, also known as the "snow monkey", same sex relations are frequent, though rates vary between troops. Females will form "consortships" characterized by affectionate social and sexual activities. In some troops up to one quarter of the females form such bonds, which vary in duration from a few days to a few weeks. Often, strong and lasting friendships result from such pairings. Males also have same sex relations, typically with multiple partners of the same age. Affectionate and playful activities are associated with such relations. Such rams prefer to court and mount other rams only, even in the presence of estrous ewes. Early writings such as Ovid's Metamorphoses and the Physiologus suggested that the hyena continually changed its sex and nature from male to female and back again. In Paedagogus, Clement of Alexandria noted that the hyena (along with the hare) was "quite obsessed with sexual intercourse". Many Europeans associated the hyena with sexual deformity, prostitution, deviant sexual behavior, and even witchcraft. The reality behind the confusing reports is the sexually aggressive behavior between the females, including mounting between females. Research has shown that "in contrast to most other female mammals, female Crocuta are male like in appearance, larger than males, and substantially more aggressive," and they have "been masculinized without being defeminized". Research has shown that "elevated levels of testosterone in utero" who in turn are possibly acting more submissive because of lower levels of testosterone in utero. Females engage in sexual behavior to stimulate ovulation, with their behavior following their hormonal cycles; during low levels of estrogen, these (female) lizards engage in "masculine" sexual roles. Those animals with currently high estrogen levels assume "feminine" sexual roles. Some parthenogenetic lizards that perform the courtship ritual have greater fertility than those kept in isolation due to an increase in hormones triggered by the sexual behaviors. So, even though asexual whiptail lizards populations lack males, sexual stimuli still increase reproductive success. From an evolutionary standpoint, these females are passing their full genetic code to all of their offspring (rather than the 50% of genes that would be passed in sexual reproduction). Certain species of gecko also reproduce by parthenogenesis. Some species of sexually reproducing geckos have also been found to display homosexual behavior, e. g. the day geckos Phelsuma laticauda and Phelsuma cepediana.

Jonathan, the world's oldest tortoise (an Aldabra giant tortoise), had been mating with another tortoise named Frederica since 1991. In 2017, it was discovered that Frederica was actually probably male all along, and was renamed Frederic.

There is evidence of same sex sexual behavior in at least 110 species of insects and arachnids. Scharf et al. says: "Males are more frequently involved in same sex sexual (SSS) behavior in the laboratory than in the field, and isolation, high density, and exposure to female pheromones increase its prevalence. SSS behavior is often shorter than the equivalent heterosexual behavior. Most cases can be explained via mistaken identification by the active (courting/mounting) male. Passive males often resist courting/mating attempts". has revealed such mating damages in 20 to 80% of the males too, indicating a fairly high occurrence of sexual coupling between males.

Male Drosophila melanogaster flies bearing two copies of a mutant allele in the fruitless gene court and attempt to mate exclusively with other males. The genetic basis of animal homosexuality has been studied in the fly D. melanogaster. Here, multiple genes have been identified that can cause homosexual courtship and mating. These genes are thought to control behavior through pheromones as well as altering the structure of the animal's brains. These studies have also investigated the influence of environment on the likelihood of flies displaying homosexual behavior.

Male bed bugs (Cimex lectularius) are sexually attracted to any newly fed individual and this results in homosexual mounting. This occurs in heterosexual mounting by the traumatic insemination in which the male pierces the female abdomen with his needle like penis. In homosexual mating this risks abdominal injuries as males lack the female counteradaptive spermalege structure. Males produce alarm pheromones to reduce such homosexual mating.