Wikipedia

Homosexual behavior in animals

(Redirected from Homosexual behavior in spotted hyenas)
For homosexuality in humans, see Homosexuality.

Various non-human animal species exhibit behavior that can be interpreted as homosexual or bisexual, often referred to as same-sex sexual behavior (SSSB) by scientists. This may include same-sex sexual activity, courtship, affection, pair bonding, and parenting among same-sex animal pairs.[1][2][3] Various forms of this are found among a variety of vertebrate and arthropod taxonomic classes. 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.

Two male mallards (Anas platyrhynchos)

Scientists observe same-sex sexual behavior in animals in different degrees and forms among different species and clades. A 2019 paper states that it has been observed in over 1,500 species.[4] Although same-sex interactions involving genital contact have been reported in many animal species, they are routinely manifested in only a few, including humans.[5] Other than humans, the only known species to exhibit exclusive homosexual orientation is the domesticated sheep (Ovis aries), involving about 10% of males.[6][7][8] The motivations for and implications of these behaviors are often lensed through anthropocentric thinking; Bruce Bagemihl states that any hypothesis is "necessarily an account of human interpretations of these phenomena".[9]: 2 

Proposed causes for same-sex sexual behavior vary across species. Theories include mistaken identity (especially for arthropods), sexually antagonistic selection, balancing selection, practice of behaviors needed for reproduction, expression of social dominance or submission, and social bonding.[10] Genetic, hormonal, and neurological variations as a basis for individual behavioral differences within species have been proposed, and same-sex sexual behavior has been induced in laboratory animals by these means.

In relation to humans

 
Two male African golden wolves (Canis anthus)
 
A cow "bulling" during oestrus.
Courtship behavior in male fruit flies

Applying the term homosexual to animals

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.[11] 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.[12] Thus homosexual behavior has been given a number of terms over the years. According to 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.[9]: 122–166 

Bailey et al. says:[2]

Homosexual: in animals, this has been used to refer to same-sex behavior that is not sexual in character (e.g. 'homosexual tandem running' in termites), same-sex courtship or copulatory behavior occurring over a short period of time (e.g. 'homosexual mounting' in cockroaches and rams) or long-term pair bonds between same-sex partners that might involve any combination of courting, copulating, parenting and affectional behaviors (e.g. 'homosexual pair bonds' in gulls). In humans, the term is used to describe individual sexual behaviors as well as long-term relationships, but in some usages connotes a gay or lesbian social identity. Scientific writing would benefit from reserving this anthropomorphic term for humans and not using it to describe behavior in other animals, because of its deeply rooted context in human society.

Animal preference and motivation is always inferred from behavior. In wild animals, researchers will as a rule not be able to map the entire life of an individual, and must infer from frequency of single observations of behavior. The correct usage of the term homosexual is that an animal exhibits homosexual behavior or even same-sex sexual behavior; however, this article conforms to the usage by modern research,[17] 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.[15][page needed]

Nature

The observation of homosexual behavior in animals can be seen as both an argument for and against the acceptance of homosexuality in humans, and has been used especially against the claim that it is a peccatum contra naturam ("sin against nature"). For instance, homosexuality in animals was cited by the American Psychological Association and other groups in their amici curiae brief to the United States Supreme Court in Lawrence v. Texas, which ultimately struck down the sodomy laws of 14 states.[18][19]

Research

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.[9]: 164  However, if the definition of "homosexual behavior" is made to include animals that participate in both same-sex and opposite-sex mating activities, hormonal differences have been documented among key sex hormones, such as testosterone and estradiol, when compared to those who participate solely in heterosexual mating.[9]: 164 

Many of the animals used in laboratory-based studies of homosexuality do not appear to spontaneously exhibit these tendencies often in the wild. Such behavior is often elicited and exaggerated by the researcher during experimentation through the destruction of a portion of brain tissue, or by exposing the animal to high levels of steroid hormones prenatally.[20][page needed] Information gathered from these studies is limited when applied to spontaneously occurring same-sex behavior in animals outside of the laboratory.[20]

Homosexual behaviour in animals has been discussed since classical antiquity. The earliest written mention of animal homosexuality appears to date back to 2,300 years ago, when Aristotle (384–322 BC) described copulation between pigeons, partridges and quails of the same sex.[21][22] The Hieroglyphics of Horapollo, written in the 4th century AD by the Egyptian writer Horapollo, mentions "hermaphroditism" in hyenas and homosexuality in partridges.[21] The first review of animal homosexuality was written by the zoologist Ferdinand Karsch-Haack in 1900.[21][23]

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,[13] innocent confusion, lack of interest, distaste, scientists fearing loss of their grants or even from a fear of "being ridiculed by their colleagues".[24][25] 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."[26] They also noted "Not every sexual act has a reproductive function ... that's true of humans and non-humans."[26] Studies have demonstrated homosexual behavior in a number of species,[27] but the true extent of homosexuality in animals is not known.

 
Two male giraffes in Kenya

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[28] 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,[16] but also in non-monogamous species like sheep. One report on sheep found that 8% of rams exhibited homosexual preferences—that is, even when given a choice, they chose male over female partners.[29] In fact, apparent homosexual individuals are known from all of the traditional domestic species, from sheep, cattle and horses to cats, dogs and budgerigars.[9]: 81  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.[30][31]

Basis

Sexual behaviors often require a significant energy investment. When sexual behaviors produce offspring, there is an obvious benefit for the animal. However, the benefit from performing homosexual behaviors (which cannot result in the production of offspring) is less obvious, and some scientists have called it a "Darwinian paradox" because it is non-reproductive. A number of non-exclusive different explanations for the emergence of such traits have been put forward.[32]

Physiological basis

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,[33] in addition to the size of the animal's gonads,[9]: 164  play a direct role in the sexual behavior and preference exhibited by that animal. Others firmly argue no evidence to support these claims exists when comparing animals of a specific species exhibiting homosexual behavior exclusively and those that do not. Ultimately, empirical support from comprehensive endocrinological studies exist for both interpretations.[33][20] Researchers found no evidence of differences in the measurements of the gonads, or the levels of the sex hormones of exclusively homosexual western gulls and ring-billed gulls.[9]: 164 

Additional studies pertaining to hormone involvement in homosexual behavior indicate that when administering treatments of testosterone and estradiol to female heterosexual animals, the elevated hormone levels increase the likelihood of homosexual behavior. Additionally, boosting the levels of sex hormones during an animal's pregnancy appears to increase the likelihood of it birthing a homosexual offspring.[33]

Genetic basis

Researchers found that disabling the fucose mutarotase[34] (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.[35][36] 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.[37]

Neurobiological basis

In March 2011, research showed that serotonin is involved in the mechanism of sexual orientation of mice.[38][39] A study conducted on fruit flies found that inhibiting the dopamine neurotransmitter inhibited lab-induced homosexual behavior.[40]

Other hypotheses

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.[41]

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.[42]

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.[43]

Some select species and groups

See also: List of animals displaying homosexual behavior

Birds

See also: List of birds displaying homosexual behavior and Seabird breeding behavior § Same-sex pairing

Black swans

 
Male black swans (Cygnus atratus)

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.[1][44] 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.[9]: 487–491 [45]

Albatrosses

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%).[46]

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.[47][48]

Blue ducks

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.[49]

Ibises

Main article: American white ibis § Breeding and lifespan

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.[50][51]

Mallards

Main article: Mallard § Breeding
 
Two male mallards (Anas platyrhynchos)

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.[9]: 493  Kees Moeliker of the Natural History Museum Rotterdam has observed one male mallard engage in homosexual necrophilia.[52]

Penguins

Further information: Penguin § Behaviour

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.[53][54]

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.[18] Other penguins in New York zoos have also been reported to have formed same-sex pairs.[55][56]

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.[57][58] Zoos in Japan and Germany have also documented homosexual male penguin couples.[59][60] 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.[61] German gay groups protested at this attempt to break up the male-male pairs[62] 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."[63]

A pair of male Magellanic penguins at the San Francisco Zoo shared a burrow for six years and raised a surrogate chick; the pair split when the male of a pair in the next burrow died and the female sought a new mate.[64]

Buddy and Pedro, a pair of male African penguins, were separated by the Toronto Zoo to mate with female penguins in 2011.[65][66] Buddy has since paired off with a female.[66]

Suki and Chupchikoni are two female African penguins that pair bonded at the Ramat Gan Safari in Israel in 2013. Chupchikoni was assumed to be male until her blood was tested.[67]

Jumbs and Kermit, two male Humboldt penguins at Wingham Wildlife Park, were gven an egg from heterosexual couple Hurricane and Isobel. The egg failed to hatch. However, they were given another abandoned egg from the same couple which successfully hatched on 12 April 2014.[68]

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.[69]

Sphen and Magic, two male gentoo penguins in Sea Life Sydney Aquarium, paired in 2018 and went on to raise two chicks, Lara (formerly Sphengic) and Clancy.[70][71] Sphen died in 2024.[72]

In September 2018 at Odense Zoo in Denmark, a male penguin couple took a chick from a heterosexual couple while they were swimming. Zoo staff speculated that the couple saw the biological parents as neglectful of the chick. The biological parents later confronted the couple and zoo staff returned the chick to them.[73]

Marama and Rocky, two female gentoo penguins at Sea Life London Aquarium, had an "incredibly strong bond" for five years prior to adopting a chick who was born in June 2019.[74] The chick was intentionally not gendered and was given a gender-neutral purple band for identification purposes.[75] They were still a couple in 2021.[76]

Skipper and Ping, two male penguins at Berlin Zoo, adopted an abandoned egg in July 2019.[77]

A female penguin couple named Electra and Viola adopted, incubated and raised an egg from another couple at L'Oceanogràfic in Valencia in 2020.[78]

A second same-sex female couple, Marmalade and Chickpea, formed at Sea Life London during the 2021 mating season.[76]

Vultures

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.[79] 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.[80]

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.[81]

Pigeons

Further information: Pigeon § Courtship and reproduction

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.[82]

Mammals

See also: List of mammals displaying homosexual behavior and Animal sexual behaviour § Mammals

Amazon dolphins

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.[9]: 339–348  In captivity, they have been observed to sometimes perform homosexual and heterosexual penetration of the blowhole, a hole homologous with the nostril of other mammals, making this the only known example of nasal sex in the animal kingdom.[9]: 339–348 [83] The males will sometimes also perform sex with males from the tucuxi species, a type of small porpoise.[9]: 339–348 

American bisons

 
The American bison is a bovine mammal which displays homosexual behavior.
Further information: American bison § Behavior and ecology

Courtship, mounting, and full anal penetration between bulls has been noted to occur among American bison. The Mandan nation Okipa festival concludes with a ceremonial enactment of this behavior, to "ensure the return of the buffalo in the coming season".[84] 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.

Bats

Two male Bonin flying foxes (Pteropus pselaphon) performing fellatio on each other[85]

More than 20 species of bat have been documented to engage in homosexual behavior.[21][85] Bat species that have been observed engaging in homosexual behavior in the wild include:[21]

Bat species that have been observed engaging in homosexual behavior in captivity include the Comoro flying fox (Pteropus livingstonii), the Rodrigues flying fox (Pteropus rodricensis) and the common vampire bat (Desmodus rotundus).[21]

Homosexual behavior in bats has been categorized into 6 groups: mutual homosexual grooming and licking, homosexual masturbation, homosexual play, homosexual mounting, coercive sex, and cross-species homosexual sex.[21][85]

In the wild, the grey-headed flying fox (Pteropus poliocephalus) engages in allogrooming wherein one partner licks and gently bites the chest and wing membrane of the other partner. Both sexes display this form of mutual homosexual grooming and it is more common in males. Males often have erect penises while they are mutually grooming each other. Like opposite-sex grooming partners, same-sex grooming partners continuously utter a "pre-copulation call", which is described as a "pulsed grating call", while engaged in this activity.[21][85]

In wild Bonin flying foxes (Pteropus pselaphon), males perform fellatio or 'male-male genital licking' on other males. Male–male genital licking events occur repeatedly several times in the same pair, and reciprocal genital licking also occurs. The male-male genital licking in these bats is considered a sexual behavior. Allogrooming in Bonin flying foxes has never been observed, hence the male-male genital licking in this species does not seem to be a byproduct of allogrooming, but rather a behavior of directly licking the male genital area, independent of allogrooming.[85] In captivity, same-sex genital licking has been observed among males of the Comoro flying fox (Pteropus livingstonii) as well as among males of the common vampire bat (Desmodus rotundus).[21][85]

In wild Indian flying foxes (Pteropus giganteus), males often mount one another, with erections and thrusting, while play-wrestling.[21] Males of the long-fingered bat (Myotis capaccinii) have been observed in the same position of male-female mounting, with one gripping the back of the other's fur. A similar behavior was also observed in the common bent-wing bat (Miniopterus schreibersii).[21]

In wild little brown bats (Myotis lucifugus), males often mount other males (and females) during late autumn and winter, when many of the mounted individuals are torpid.[21] 35% of matings during this period are homosexual.[86] These coercive copulations usually include ejaculation and the mounted bat often makes a typical copulation call consisting of a long squawk.[21] Similarly, in hibernacula of the common noctule (Nyctalus noctula), active males were observed to wake up from lethargy on a warm day and engage in mating with lethargic males and (active or lethargic) females. The lethargic males, like females, called out loudly and presented their buccal glands with opened mouth during copulation.[21]

Vesey-Fitzgerald (1949) observed homosexual behaviours in all 12 British bat species known at the time: "Homosexuality is common in the spring in all species, and, since the males are in full possession of their powers, I suspect throughout the summer...I have even seen homosexuality between Natterer's and Daubenton's bats (Myotis nattereri and M. daubentonii)."[21]

Bottlenose dolphins

See also: Animal sexual behavior § Bottlenose dolphins, and Bottlenose dolphin § Life history

Dolphins of several species engage in homosexual acts, though it is best studied in the bottlenose dolphins.[9][page needed] Sexual encounters between females take the shape of "beak-genital propulsion", where one female inserts her beak in the genital opening of the other while swimming gently forward.[87] 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.[88]

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.[89] 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.[90]

Elephants

Further information: Elephant § Mating

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.[9]: 427–430 

Giraffes

Further information: Giraffe § Necking

Male giraffes have been observed to engage in remarkably high frequencies of homosexual behavior. After aggressive "necking", it is common for two male giraffes to caress and court each other, leading up to mounting and climax. Such interactions between males have been found to be more frequent than heterosexual coupling.[91] 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.[9]: 391–393 

Humpback whales

In 2022, photographers captured the first-ever images of humpback whales mating, both of which were males, off the coast of Maui. The whales were observed for 30 minutes, with one whale using its penis to penetrate the genital slit of the other.[92][93]

Marmots

 
 
Olympic marmot (left) and hoary marmot (right)

Homosexual behavior is quite common in wild marmots.[9]: 525–527  In Olympic marmots (Marmota olympus) and hoary marmots (Marmota caligata), females often mount other females as well as engage in other affectionate and sexual behaviors with females of the same species.[9]: 525–527  They display a high frequency of these behaviors especially when they are in heat.[9]: 525–527 [94] 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.[9]: 525–527 [95]

Lions

Further information: Sexual behavior of lions
Male lions mating

Both male and female lions have been seen to interact homosexually.[98] Male lions pair-bond for a number of days and initiate homosexual activity with affectionate nuzzling and caressing, leading to mounting and thrusting. About 8% of mountings have been observed to occur with other males. Pairings between females are held to be fairly common in captivity but have not been observed in the wild.

Polecats

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.[99][page needed]

Primates

Same-sex sexual behaviors have been observed in New World monkeys, Old World monkeys, and apes. 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 behavior. 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;[100] these behaviors are a conserved trait among Old World monkeys and apes.[101] Among Old World monkeys, same-sex genital interactions are much more common in multi-male systems as opposed to uni-male systems.[102]

Bonobos
See also: Bonobo § Sociosexual behaviour
 
Two male bonobos

Bonobos form a matriarchal society, unusual among apes. They are fully bisexual: both males and females engage in hetero- and homosexual behavior, being noted for female–female sex in particular, including between juveniles and adults.[103] Roughly 60% of all bonobo sexual activity occurs between two or more females. While the homosexual bonding system in bonobos represents the highest frequency of homosexuality known in any primate species, homosexuality has been reported for all great apes, as well as a number of other primate species.[109]

Dutch primatologist Frans de Waal, who extensively observed and filmed bonobos, believed that sexual activity is the bonobo's way of avoiding conflict. Anything that arouses the interest of more than one bonobo at a time, not just food, tends to result in sexual contact. If two bonobos approach a cardboard box thrown into their enclosure, they will briefly mount each other before playing with the box. Such situations lead to squabbles in most other species. But bonobos are quite tolerant, perhaps because they use sex to divert attention and to defuse tension.[103][110]

Bonobo sex often occurs in aggressive contexts totally unrelated to food. A jealous male might chase another away from a female, after which the two males reunite and engage in scrotal rubbing. Or after a female hits a juvenile, the latter's mother may lunge at the aggressor, an action that is immediately followed by genital rubbing between the two adults.[103]

Chimpanzees

Compared to bonobos, less research has been conducted on same-sex sexual behaviors in chimpanzees (Pan troglodytes). In contrast to bonobos, same-sex sexual behaviors are less common in Kibale chimpanzees (Pan troglodytes schweinfurthii) and primarily occur between adult males.[111] While less frequent than in female bonobos, same-sex sexual behaviors between male chimpanzees occur regularly and in a variety of contexts. Female chimpanzees are much less likely to engage in same-sex sexual behaviors than males.

Gibbons

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.[112]

Gorillas
See also: Gorilla § Social structure

Both male and female mountain gorillas (Gorilla gorilla beringei) have been observed engaging in same-sex sexual behaviors.[113] Homosexual behavior among male gorillas has been studied.[114] 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.[115] Among all-male "bachelor" groups, same-sex sexual interactions are extremely common.[116] 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.[117] Same-sex sexual behaviors are natural and common among mountain gorillas.

Orangutans

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.[118]

Sociosexual behaviors among orangutans (Pongo spp.) are extremely common and invoke a variety of evolutionary strategies.[119][120] However, only one study of same-sex sexual behavior has been published.[121] 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.

Monkeys

Among monkeys[clarification needed], Lionel Tiger and Robin Fox conducted a study on how Depo-Provera contraceptives lead to decreased male attraction to females.[122]

Japanese macaques
Further information: Japanese macaque § Behavior

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.[9]: 302–305 

Sheep

Main article: Homosexual behavior in sheep

Ovis aries, the common domesticated sheep, has attracted much attention due to the fact that around 8–10% of rams have an exclusive homosexual orientation.[127] Such rams prefer to court and mount other rams only, even in the presence of estrous ewes.[6] Moreover, around 18–22% of rams are bisexual.[124]

Several observations indicate that male–male sexual preference in rams is sexually motivated. Rams routinely perform the same courtship behaviors (including foreleg kicks, nudges, vocalizations, anogenital sniffs and flehmen) prior to mounting other males as observed when other rams court and mount estrous females. Furthermore, pelvic thrusting and ejaculation often accompany same-sex mounts by rams.[126]

Spotted hyenas

See also: Animal sexual behaviour § Spotted hyena, and Spotted hyena § Reproduction and development
 
The spotted hyena is a moderately large, terrestrial carnivore native to Africa.

The family structure of the spotted hyena is matriarchal, and dominance relationships with strong sexual elements are routinely observed between related females. Due largely to the female spotted hyena's unique urogenital system, which looks more like a penis rather than a vagina, early naturalists thought hyenas were hermaphroditic males who commonly practiced homosexuality.[128][failed verification] 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,"[129] and they have "been masculinized without being defeminized".[128][failed verification]

Study of this unique genitalia and aggressive behavior in the female hyena has led to the understanding that more aggressive females are better able to compete for resources, including food and mating partners.[128][130] Research has shown that "elevated levels of testosterone in utero"[131] contribute to extra aggressiveness; both males and females mount members of both the same and opposite sex,[131][132] who in turn are possibly acting more submissive because of lower levels of testosterone in utero.[129]

Reptiles

Lizards

Further information: Animal sexual behaviour § Lizards

Several species of whiptail lizard (especially in the genus Aspidoscelis) consist only of females that have the ability to reproduce through parthenogenesis.[133] 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.[134]

Some species of sexually reproducing geckos have also been found to display homosexual behavior, e.g. the day geckos Phelsuma laticauda and Phelsuma cepediana.[135]

Tortoises

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.[136]

Insects and arachnids

There is evidence of same-sex sexual behavior in at least 110 species of insects and arachnids.[137] 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".[137]

Scharf et al. continues: "SSS behavior has been reported in most insect orders, and Bagemihl (1999) provides a list of ~100 species of insects demonstrating such behavior. Yet, this list lacks detailed descriptions, and a more comprehensive summary of its prevalence in invertebrates, as well as ethology, causes, implications, and evolution of this behavior, remains lacking".[137]

Dragonflies

 
The head of darner dragonfly (Basiaeschna janata)

Male homosexuality has been inferred in several species of dragonflies (the order Odonata). The cloacal pinchers of male damselflies and dragonflies inflict characteristic head damage to females during sex. A survey of 11 species of damsel and dragonflies[138][139] has revealed such mating damages in 20 to 80% of the males too, indicating a fairly high occurrence of sexual coupling between males.

Fruit flies

Male Drosophila melanogaster flies bearing two copies of a mutant allele in the fruitless gene court and attempt to mate exclusively with other males.[140] The genetic basis of animal homosexuality has been studied in the fly D. melanogaster.[141] Here, multiple genes have been identified that can cause homosexual courtship and mating.[142] These genes are thought to control behavior through pheromones as well as altering the structure of the animal's brains.[143][144] These studies have also investigated the influence of environment on the likelihood of flies displaying homosexual behavior.[145][146]

Bed bugs

Main article: Bed bug § Reproduction

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.

See also

References

  1. ^ a b Braithwaite, L. W. (1981). "Ecological studies of the Black Swan III – Behaviour and social organization". Wildlife Research. 8. Canberra, Australia: CSIRO: 134–146. doi:10.1071/WR9810135.
  2. ^ a b Bailey, N. W.; Zuk, M. (August 2009). "Same-sex sexual behavior and evolution". Trends in Ecology and Evolution. 24 (8): 439–46. Bibcode:2009TEcoE..24..439B. doi:10.1016/j.tree.2009.03.014. PMID 19539396.
  3. ^ "Same-sex Behavior Seen In Nearly All Animals, Review Finds". ScienceDaily (Press release). 17 June 2009.
  4. ^ Monk, Julia D.; Giglio, Erin; Kamath, Ambika; Lambert, Max R.; McDonough, Caitlin E. (December 2019). "An alternative hypothesis for the evolution of same-sex sexual behaviour in animals". Nature Ecology and Evolution. 3 (12): 1622–1631. Bibcode:2019NatEE...3.1622M. doi:10.1038/s41559-019-1019-7. ISSN 2397-334X. PMID 31740842. S2CID 256708244.
  5. ^ Bailey JM, Vasey PL, Diamond LM, Breedlove SM, Vilain E, Epprecht M (September 2016). "Sexual Orientation, Controversy, and Science". Psychological Science in the Public Interest. 17 (2): 45–101. doi:10.1177/1529100616637616. PMID 27113562. S2CID 42281410.
  6. ^ a b c Poiani A, Dixson AF (2010). Animal Homosexuality: A Biosocial Perspective. Cambridge University Press. p. 179. ISBN 9781139490382. This makes O. aries (ram) only the second mammal known, apart from humans, capable of displaying exclusive homosexuality.
  7. ^ Levay S (2017). Gay, Straight, and The Reason Why: The Science of Sexual Orientation (Second ed.). Cambridge, Massachusetts: Oxford University Press. pp. 38, 119. ISBN 978-0-19-029737-4 – via Google Books.
  8. ^ Raymond, Michel; Crochet, Pierre-André (October 2023). "Carving Non-Proximal Explanations for Same-Sex Sexual Orientation". Archives of Sexual Behavior. 52 (7): 3007–3012. doi:10.1007/s10508-022-02497-z. PMID 36469147. There are numerous reports of homosexual behavior in many animal species in the wild, but there are no reports of exclusive same-sex sexual orientation (Bagemihl, 2000). It is, however, well-established that some domestic rams exhibit exclusive same-sex sexual orientation.
  9. ^ a b c d e f g h i j k l m n o p q r s t u v Bagemihl B (1999). Biological Exuberance: Animal Homosexuality and Natural Diversity (Stone Wall Inn ed.). New York City: St. Martin's Press. ISBN 9780312253776 – via Internet Archive.
  10. ^ Balfour, Vicki L.; Shuker, David M. (November 2020). "Same-sex sexual behaviour". Current Biology. 30 (22): R1345–R1346. Bibcode:2020CBio...30R1345B. doi:10.1016/j.cub.2020.09.005. PMID 33202225. Retrieved 25 June 2024.
  11. ^ The first known use of the word Homoseksuäl is found in Benkert Kertbeny, K. M. (1869): Paragraph 143 des Preussichen Strafgesetzebuches vom 14/4-1851 und seine Aufrechterhaltung als Paragraph 152 im Entwurf eines Strafgesetzbuches fur den Norddeutschen Bundes, Leipzig, 1869. Reprinted in Jahrbuch fur sexuelle Zwischenstufen 7 (1905), pp. 1–66
  12. ^ Dorit R (September–October 2004). "Rethinking Sex". American Scientist. Archived from the original on 25 June 2010. Retrieved 11 September 2007.
  13. ^ a b Roughgarden, Joan (2004). Evolutions rainbow: Diversity, gender and sexuality in nature and people. Berkeley, California: University of California Press. pp. 13–183. ISBN 9780520957978 – via Google Books.
  14. ^ Vasey PL (1995). "Homosexual behaviour in primates: A review of evidence and theory". International Journal of Primatology. 16 (2): 173–204. doi:10.1007/bf02735477. S2CID 26021360.
  15. ^ a b Sommer V, Vasey PL (2006). Homosexual Behaviour in Animals, An Evolutionary Perspective. Cambridge: Cambridge University Press. ISBN 978-0-521-86446-6.
  16. ^ a b Douglas K (7 December 2009). "Homosexual selection: The power of same-sex liaisons". New Scientist. Retrieved 21 December 2009.
  17. ^ [9]: 122–166 [13][14][15][page needed][16]
  18. ^ a b Smith D (7 February 2004). "Love That Dare Not Squeak Its Name". The New York Times. Retrieved 10 September 2007.
  19. ^ "Brief for Amici Curiae in Support of Petitioners, Lawrence v. Texas" (PDF). American Psychological Association. January 2003.
  20. ^ a b c Somer V, Vasey PL, eds. (2010). Homosexual Behavior In Animals: An Evolutionary Perspective. Cambridge: Cambridge Press. ISBN 978-0-521-18230-0. ASIN 0521182301.
  21. ^ a b c d e f g h i j k l m n o Riccucci M (2011). "Same-sex sexual behaviour in bats". Hystrix It. J. Mammal. 22 (1): 139–47. doi:10.4404/hystrix-22.1-4478.
  22. ^ Aristotle; Pliny (transl. Rackham, 1947, Vol. 3: 399; Aristotle, transl. Peck, 1970, Vol. 2: 233; see also: Aelian, transl. Wilson, 1997: 37). "Oxford Academic Zoologic Journal".{{cite web}}: CS1 maint: multiple names: authors list (link) CS1 maint: numeric names: authors list (link)
  23. ^ Karsch, F. (1900). "Päderastie und Tribadie bei den Tieren auf Grund Literatur". Jahrbuch für sexuelle Zwischenstufen. 2. hdl:2027/mdp.39015016472477.
  24. ^ "1,500 Animal Species Practice Homosexuality". News-medical.net. 23 October 2006. Archived from the original on 28 May 2007. Retrieved 10 September 2007.
  25. ^ "Homosexuality in the Animal kingdom". nhm.uio.no. Natural History Museum, University of Oslo. February 2009. Retrieved 6 October 2017.
  26. ^ a b Moskowitz C (19 May 2008). "Homosexuality Common in the Wild, Scientists Say". Fox News. Archived from the original on 28 May 2008. Retrieved 2 July 2008.
  27. ^ Dagg, Anne Innis (1984). "Homosexual behaviour and female-male mounting in mammals—a first survey". Mammal Review. 14 (4): 155–185. doi:10.1111/j.1365-2907.1984.tb00344.x.
  28. ^ Thierry Lodé La guerre des sexes chez les animaux Eds O Jacb, Paris, 2006, ISBN 2-7381-1901-8
  29. ^ a b Roselli CE, Larkin K, Resko JA, Stellflug JN, Stormshak F (February 2004). "The volume of a sexually dimorphic nucleus in the ovine medial preoptic area/anterior hypothalamus varies with sexual partner preference". Endocrinology. 145 (2): 478–83. doi:10.1210/en.2003-1098. PMID 14525915.
  30. ^ Zimmer, Carl (3 October 2023). "Same-Sex Behavior Evolved in Many Mammals to Reduce Conflict, Study Suggests". The New York Times. Archived from the original on 20 October 2023 – via Internet Archive.
  31. ^ Gómez, José M.; et al. (3 October 2023). "The evolution of same-sex sexual behaviour in mammals". Nature. 14 (5719): 5719. Bibcode:2023NatCo..14.5719G. doi:10.1038/s41467-023-41290-x. PMC 10547684. PMID 37788987.
  32. ^ Savolainen, Vincent (2016). "Evolution of Homosexuality". Encyclopedia of Evolutionary Psychological Science. Springer Publishing. pp. 1–8. doi:10.1007/978-3-319-16999-6_3403-1. ISBN 978-3-319-16999-6. S2CID 220448278.
  33. ^ a b c Adler T (4 January 1997). "Animal's Fancies". Society for Science and the Public. 151 (1): 8–9. doi:10.2307/3980720. JSTOR 3980720.
  34. ^ "WikiGenes - Collaborative Publishing". Archived from the original on 10 June 2016.
  35. ^ Moore M (8 July 2010). "Female mice 'can be turned lesbian by deleting gene'". The Telegraph. London. Archived from the original on 10 July 2010. Retrieved 17 November 2010.
  36. ^ Park D, Choi D, Lee J, Lim DS, Park C (July 2010). "Male-like sexual behavior of female mouse lacking fucose mutarotase". BMC Genetics. 11. BioMed Central: 62. doi:10.1186/1471-2156-11-62. PMC 2912782. PMID 20609214.
  37. ^ Terry J (2000). "'Unnatural Acts' in Nature: The Scientific Fascination with Queer Animals". GLQ: A Journal of Lesbian and Gay Studies. 6 (2): 151–193. doi:10.1215/10642684-6-2-151. S2CID 145504677.
  38. ^ "Sexual preference chemical found in mice". BBC News. 23 March 2011. Retrieved 24 March 2011.
  39. ^ Liu Y, Jiang Y, Si Y, Kim JY, Chen ZF, Rao Y (April 2011). "Molecular regulation of sexual preference revealed by genetic studies of 5-HT in the brains of male mice". Nature. 472 (7341): 95–9. Bibcode:2011Natur.472...95L. doi:10.1038/nature09822. PMC 4094133. PMID 21441904.
  40. ^ Zuk M (22 November 2011). "Same-sex insects: what do bees-or at least flies-have to tell us about homosexuality?". Natural History. 119 (10): 22.
  41. ^ Braithwaite, L.Wayne. "Ecological studies of the black swan". Australian Wildlife Research. hdl:102.100.100/292286. Retrieved 3 December 2021.
  42. ^ MacFarlane, Geoff (2010). "Homosexual behaviour in birds: frequency of expression is related to parental care disparity between the sexes". Animal Behaviour. 80 (3): 375–390. doi:10.1016/j.anbehav.2010.05.009. S2CID 53148085. Retrieved 3 December 2021.
  43. ^ Monk, Julia (2019). "An alternative hypothesis for the evolution of same-sex sexual behaviour in animals". Nature Ecology & Evolution. 3 (12): 1622–1631. Bibcode:2019NatEE...3.1622M. doi:10.1038/s41559-019-1019-7. PMID 31740842. S2CID 208144026.
  44. ^ Braithwaite, L. W. (1970). "The Black Swan". Australian Natural History. 16: 375–379.
  45. ^ "Oslo gay animal show draws crowds". BBC News. 19 October 2006. Retrieved 15 June 2009.
  46. ^ Young LC, Zaun BJ, Vanderwerf EA (August 2008). "Successful same-sex pairing in Laysan albatross". Biology Letters. 4 (4): 323–5. doi:10.1098/rsbl.2008.0191. PMC 2610150. PMID 18505710.
  47. ^ Frost, Natasha (29 November 2021). "Climate Change Is Driving Some Albatrosses to 'Divorce,' Study Finds". The New York Times. ISSN 0362-4331. Retrieved 29 November 2021.
  48. ^ "Climate crisis pushes albatross 'divorce' rates higher – study". The Guardian. 24 November 2021. Retrieved 29 November 2021.
  49. ^ Lite, Jordan. "Gay ducks derail repopulation plan". Scientific American Blog Network. Retrieved 4 December 2021.
  50. ^ Milton J (1 December 2010). "Mercury causes homosexuality in male ibises". Nature. doi:10.1038/news.2010.641.
  51. ^ Frederick P, Jayasena N (June 2011). "Altered pairing behaviour and reproductive success in white ibises exposed to environmentally relevant concentrations of methylmercury". Proceedings. Biological Sciences. 278 (1713): 1851–7. doi:10.1098/rspb.2010.2189. PMC 3097836. PMID 21123262.
  52. ^ Moeliker CW (9 November 2001). "The First Case of Homosexual Necrophilia in the Mallard Anas platyrhynchos (Aves: Anatidae)" (PDF). Deinsea. 8 (2001): 243–247. Retrieved 6 October 2014.
  53. ^ "Unearthed study on 'sexual depravity' in penguins". abc.net.au. 10 June 2012. Retrieved 25 March 2018.
  54. ^ Russell, Douglas G. D.; Sladen, William J. L.; Ainley, David G. (2012). "Dr. George Murray Levick (1876–1956): unpublished notes on the sexual habits of the Adélie penguin". Polar Record. 48 (4): 387–393. Bibcode:2012PoRec..48..387R. doi:10.1017/S0032247412000216. S2CID 146584734.
  55. ^ "They're in love. They're gay. They're penguins... And they're not alone". Columbia University. Columbia News Service. 10 June 2002. Archived from the original on 23 January 2009.
  56. ^ "Central Park Zoo's gay penguins ignite debate", 2002-02-07, San Francisco Chronicle
  57. ^ "Denmark's Gay Penguins Become Fathers". Huffington Post. 11 November 2012.
  58. ^ "Gay Penguins Become Dads". BuzzFeed. 9 November 2012.
  59. ^ "Cold Shoulder for Swedish Seductresses | Germany". Deutsche Welle. 10 February 2005. Retrieved 17 November 2010.
  60. ^ "Gay penguin couple adopts abandoned egg in German zoo". CBC News. 5 June 2009.
  61. ^ "Ananova Article on "Tempting Gay Penguins Straight"". Ananova.com. Retrieved 17 November 2010.
  62. ^ "Followup Ananova Article on German authorities stopping trying to change the penguins' sexual orientation, after GLBTQI organizations protest". Ananova.com. Archived from the original on 8 February 2009. Retrieved 17 November 2010.
  63. ^ "Gay outrage over penguin sex test". BBC News. 14 February 2005. Retrieved 4 May 2010.
  64. ^ May M (14 July 2009). "Widow a wedge between zoo's male penguin pair". San Francisco Chronicle. Retrieved 17 November 2010.
  65. ^ McCormack S (14 November 2011), "Gay Penguins Reunion: Buddy And Pedro Will Be Back Together By Spring", Gay Voices, The Huffington Post, retrieved 16 November 2011
  66. ^ a b "Same-sex penguin pair pursue female partners". CBC News. 12 December 2011.
  67. ^ Schuster R (19 December 2013). "Lesbian Penguins Shack Up at Israeli Zoo". The Jewish Daily Forward.
  68. ^ Wingham Wildlife Park
  69. ^ "Auckland's gay penguin couple prepare for special Christmas bundle of joy". Newshub. Archived from the original on 24 December 2018. Retrieved 24 December 2018.
  70. ^ Bowles, Nellie (15 January 2019). "The Gay Penguins of Australia". The New York Times. ISSN 0362-4331. Retrieved 22 August 2024.
  71. ^ Press (21 August 2024). "When globally famous gay penguin Sphen died in Sydney, his partner began to sing". The Guardian. ISSN 0261-3077. Retrieved 22 August 2024.
  72. ^ "Gay penguin Sphen dies in Australian aquarium". Deutsche Welle. 22 August 2024. Retrieved 22 August 2024.
  73. ^ Kacala, Alexander (27 September 2018). "Gay penguins 'kidnap' chick from straight parents at Denmark zoo". NBC News. Retrieved 22 August 2024.
  74. ^ Konstantinides, Anneta. "A same-sex penguin couple at a London aquarium become mothers for the first time after adopting a chick". Business Insider. Retrieved 22 August 2024.
  75. ^ Gawronski, Quinn (10 September 2019). "Gay penguins at London aquarium are raising 'genderless' chick". NBC News. Retrieved 22 August 2024.
  76. ^ a b "Penguins: Same-sex couples formed at London aquarium". BBC News. Retrieved 22 August 2024.
  77. ^ "Berlin gay penguins adopt abandoned egg". BBC News. 12 August 2019. Retrieved 22 August 2024.
  78. ^ Silva, Cynthia (26 August 2020). "Lesbian penguin 'super moms' welcome chick at Spanish aquarium". NBC News. Retrieved 22 August 2024.
  79. ^ Silver E (2 August 1999). "Gay vulture couple raise surrogate chicks". The Independent. London. Retrieved 21 September 2009.
  80. ^ Lis J (21 September 2009). "'Gay' vulture couple split up at Jerusalem zoo, then become fathers". Haaretz. Archived from the original on 16 June 2013. Retrieved 21 September 2009.
  81. ^ "Gay vultures split up to go straight | Showbiz: Latest News | STV Entertainment". Entertainment.stv.tv. Archived from the original on 21 November 2010. Retrieved 17 November 2010.
  82. ^ Jankowiak, Łukasz; Tryjanowski, Piotr; Hetmański, Tomasz; Skórka, Piotr (26 January 2018). "Experimentally evoked same-sex sexual behaviour in pigeons: better to be in a female-female pair than alone". Scientific Reports. 8 (1): 1654. Bibcode:2018NatSR...8.1654J. doi:10.1038/s41598-018-20128-3. ISSN 2045-2322. PMC 5785962. PMID 29374281.
  83. ^ Sylvestre, J.-P. (Some Observations on Behavior of Two Orinoco Dolphins (Inia geoffrensis humboldtiaba [Pilleri and Gihr 1977]), in Captivity, at Duisburg Zoo. Aquatic mammals no 11, pp. 58–65 article Archived 22 October 2016 at the Wayback Machine
  84. ^ Bagemihl B (May 2000). "Left-Handed Bears & Androgynous Cassowaries: Homosexual/transgendered animals and indigenous knowledge". Whole Earth Magazine. Archived from the original on 15 August 2000. Retrieved 13 December 2013 – via Internet Archive.
  85. ^ a b c d e f g Sugita N (November 2016). "Homosexual Fellatio: Erect Penis Licking between Male Bonin Flying Foxes Pteropus pselaphon". PLOS ONE. 11 (11): e0166024. Bibcode:2016PLoSO..1166024S. doi:10.1371/journal.pone.0166024. PMC 5100941. PMID 27824953.
  86. ^ Havens A (2006). Dewey T, Shefferly N (eds.). "Myotis lucifugus". Animal Diversity Web.
  87. ^ Norris KS, Dohl TP (1980). "Behaviour of the Hawaiian spinner dolphin, Stenella longirostris". Fishery Bulletin. 77: 821–849.
  88. ^ Wells RS (1995). "Community structure of Bottlenose Dolphins near Sarasota, Florida". Paper presented at the 24th International Ethological Conference, Honoluly, Hawaii.
  89. ^ "Same Sex Couples Common in the Wild". livescience.com. 16 May 2008. Retrieved 25 March 2018.
  90. ^ Herzing DL, Johnson CM (1997). "Interspecific Interaction between Spotted Dolphins (Stenella frontalis) and Bottlenose Dolphins (Tursiops truncatus) in the Bahamas, 1985–1995". Aquatic Mammals. 23: 85–99.
  91. ^ Coe MJ (1967). ""Necking" behavior in the giraffe". Journal of Zoology. 151 (3): 313–321. doi:10.1111/j.1469-7998.1967.tb02117.x.
  92. ^ Kuta, Sarah. "These Photos Are the First to Show Humpback Whales Mating—and Both Are Males". Smithsonian Magazine. Retrieved 11 March 2024.
  93. ^ Stack, Stephanie H.; Krannichfeld, Lyle; Romano, Brandi (27 February 2024). "An observation of sexual behavior between two male humpback whales". Marine Mammal Science. 40 (3). Bibcode:2024MMamS..40E3119S. doi:10.1111/mms.13119. hdl:10072/430033. ISSN 0824-0469.
  94. ^ Barash DP (1973). "The Social Biology of the Olympic Marmot". Animal Behaviour Monographs. 6 (3): 171–245. doi:10.1016/0003-3472(73)90002-X.
  95. ^ Barash DP (February 1974). "The social behaviour of the hoary marmot (Marmota caligata)". Anim. Behav. 22 (1): 256–61. doi:10.1016/S0003-3472(74)80077-1.
  96. ^ In his discussion of lion same-sex relations, Bagemihl is making use of published work by: J.B. Cooper, "An Exploratory Study on African Lions" in Comparative Psychology Monographs 17:1–48; R.L. Eaton, "The Biology and Social Behavior of Reproduction in the Lion" in Eaton, ed. The World's Cats, vol. II; pp.3–58; Seattle, 1974; G.B. Schaller, The Serengeti Lion; University of Chicago Press, 1972
  97. ^ Srivastav S (15–31 December 2001). "Lion, Without Lioness". TerraGreen: News to Save the Earth. Terragreen. Archived from the original on 14 September 2007. Retrieved 2 September 2007.
  98. ^ [9]: 302–305 [96][97]
  99. ^ Thierry Lodé La guerre des sexes chez les animaux Eds O Jacob, Paris, 2006.ISBN 2-7381-1901-8
  100. ^ Pfau, Daniel; Jordan, Cynthia L.; Breedlove, S. Marc (1 August 2021). "The De-Scent of Sexuality: Did Loss of a Pheromone Signaling Protein Permit the Evolution of Same-Sex Sexual Behavior in Primates?". Archives of Sexual Behavior. 50 (6): 2267–2276. doi:10.1007/s10508-018-1377-2. ISSN 1573-2800. PMID 31016493. S2CID 254251448.
  101. ^ Dixson, Alan (19 August 2010), "Homosexual behaviour in primates", Animal Homosexuality, Cambridge University Press, pp. 381–400, doi:10.1017/cbo9780511762192.010, ISBN 978-0-521-19675-8, retrieved 22 October 2023
  102. ^ MacFarlane, Geoff R.; Vasey, Paul L. (1 May 2016). "Promiscuous primates engage in same-sex genital interactions". Behavioural Processes. 126: 21–26. doi:10.1016/j.beproc.2016.02.016. ISSN 0376-6357. PMID 26930251. S2CID 39603917.
  103. ^ a b c de Waal FB (March 1995). "Bonobo Sex and Society: The behavior of a close relative challenges assumptions about male supremacy in human evolution" (PDF). Scientific American. 272 (3): 82–88. Bibcode:1995SciAm.272c..82W. doi:10.1038/scientificamerican0395-82. PMID 7871411. Archived from the original (PDF) on 1 August 2016. Retrieved 10 March 2019.
  104. ^ "Bonobo". Columbus Zoo and Aquarium. Archived from the original on 17 August 2007.
  105. ^ Dawkins R (2004). "Rendezvous I: Chimpanzees". The Ancestor's Tale: A Pilgrimage to the Dawn of Life. Houghton Mifflin (US). pp. 92–93. ISBN 978-1-155-16265-2.
  106. ^ Laird C. "Social Organization". bio.davidson.edu. Archived from the original on 19 May 2011.
  107. ^ Stanford CB (1998). "The social behavior of chimpanzees and bonobos". Current Anthropology. 39 (4): 399–407. doi:10.1086/204757. S2CID 8452514.
  108. ^ Kano T (1992). The Last Ape: Pygmy Chimpanzee Behavior and Ecology. Stanford University Press. ISBN 9780804716123.
  109. ^ [104][105][106][107][108]
  110. ^ de Waal FB (2001). "Bonobos and Fig Leaves". The Ape and the Sushi Master: Cultural Reflections by a Primatologist. Basic Books. ISBN 978-84-493-1325-7.
  111. ^ Sandel, Aaron A.; Reddy, Rachna B. (2 February 2021). "Sociosexual behaviour in wild chimpanzees occurs in variable contexts and is frequent between same-sex partners". Behaviour. 158 (3–4): 249–276. doi:10.1163/1568539X-bja10062. ISSN 0005-7959. S2CID 234056789.
  112. ^ Edwards, Anna-Marie A. R.; Todd, John D. (1 April 1991). "Homosexual behaviour in wild white-handed gibbons (Hylobates lar)". Primates. 32 (2): 231–236. doi:10.1007/BF02381180. ISSN 1610-7365. S2CID 26662075.
  113. ^ "Does homosexual behaviour exist among gorillas?". Berggorilla & Regenwald Direkthilfe e.V. Retrieved 22 October 2023.
  114. ^ Yamagiwa J (1 January 1987). "Intra- and inter-group interactions of an all-male group of virunga mountain gorillas (Gorilla gorilla beringei)". Primates. 28 (1): 1–30. doi:10.1007/BF02382180. ISSN 1610-7365. S2CID 24667667.
  115. ^ Grueter CC, Stoinski TS (11 May 2016). "Homosexual Behavior in Female Mountain Gorillas: Reflection of Dominance, Affiliation, Reconciliation or Arousal?". PLOS ONE. 11 (5): e0154185. Bibcode:2016PLoSO..1154185G. doi:10.1371/journal.pone.0154185. PMC 4864209. PMID 27167861.
  116. ^ Yamagiwa, Juichi (1 January 1987). "Intra- and inter-group interactions of an all-male group of virunga mountain gorillas (Gorilla gorilla beringei)". Primates. 28 (1): 1–30. doi:10.1007/BF02382180. ISSN 1610-7365. S2CID 24667667.
  117. ^ Australia, University of Western. "Homosexual activity documented in female gorillas for the first time". phys.org. Retrieved 22 October 2023.
  118. ^ Fox EA (November 2001). "Homosexual behavior in wild Sumatran orangutans (Pongo pygmaeus abelii)". American Journal of Primatology. 55 (3): 177–81. doi:10.1002/ajp.1051. PMID 11746281. S2CID 21561581.
  119. ^ Kunz, Julia A.; Duvot, Guilhem J.; Willems, Erik P.; Stickelberger, Julia; Spillmann, Brigitte; Utami Atmoko, Sri Suci; van Noordwijk, Maria A.; van Schaik, Carel P. (December 2021). "The context of sexual coercion in orang-utans: when do male and female mating interests collide?". Animal Behaviour. 182: 67–90. doi:10.1016/j.anbehav.2021.09.012. ISSN 0003-3472.
  120. ^ O'Connell, Caitlin A.; Susanto, Tri Wahyu; Knott, Cheryl D. (November 2020). "Sociosexual behavioral patterns involving nulliparous female orangutans ( Pongo sp.) reflect unique challenges during the adolescent period". American Journal of Primatology. 82 (11): e23058. doi:10.1002/ajp.23058. ISSN 0275-2565. PMID 31583721. S2CID 203659435.
  121. ^ Fox, Elizabeth A. (2001). "Homosexual behavior in wild Sumatran orangutans (Pongo pygmaeus abelii)". American Journal of Primatology. 55 (3): 177–181. doi:10.1002/ajp.1051. ISSN 0275-2565. PMID 11746281. S2CID 21561581.
  122. ^ Tiger L (1992). The Pursuit of Pleasure. Transaction Publishers. pp. 66ff. ISBN 978-0-7658-0696-3.
  123. ^ Roselli CE, Stormshak F (May 2009). "The neurobiology of sexual partner preferences in rams". Hormones and Behavior. 55 (5): 611–20. doi:10.1016/j.yhbeh.2009.03.013. PMC 2684522. PMID 19446078.
  124. ^ a b Roselli CE, Stormshak F (March 2009). "Prenatal programming of sexual partner preference: the ram model". Journal of Neuroendocrinology. 21 (4): 359–64. doi:10.1111/j.1365-2826.2009.01828.x. PMC 2668810. PMID 19207819.
  125. ^ Perkins A, Fitzgerald JA, Price EO (July 1992). "Luteinizing hormone and testosterone response of sexually active and inactive rams". Journal of Animal Science. 70 (7): 2086–93. doi:10.2527/1992.7072086x. PMID 1644682. Archived from the original on 30 June 2017. Retrieved 7 July 2015.
  126. ^ a b Roselli CE, Larkin K, Schrunk JM, Stormshak F (November 2004). "Sexual partner preference, hypothalamic morphology and aromatase in rams". Physiology & Behavior. 83 (2): 233–45. doi:10.1016/j.physbeh.2004.08.017. PMID 15488542. S2CID 156571.
  127. ^ [6][29][123][124][125][126]
  128. ^ a b c "Like mother, like cubs: Hyena alpha moms jumpstart cubs with hormonal jolt". Michigan State University. 2006. Archived from the original on 29 June 2007. Retrieved 11 September 2007.
  129. ^ a b Holekamp KE (2003). "Research: Spotted Hyena - Introduction and Overview". Michigan State University, Department of Zoology. Archived from the original on 29 June 2007. Retrieved 11 September 2007.
  130. ^ Selim J (28 April 2006). "Top Dogs: Steroid hormones give hyenas a head start". Discover Magazine. Retrieved 11 September 2007.
  131. ^ a b Forger NG, Frank LG, Breedlove SM, Glickman SE (November 1996). "Sexual dimorphism of perineal muscles and motoneurons in spotted hyenas". The Journal of Comparative Neurology. 375 (2): 333–43. doi:10.1002/(SICI)1096-9861(19961111)375:2<333::AID-CNE11>3.0.CO;2-W. PMID 8915834.
  132. ^ Anna Wilson (Spring 2003). "Sexing the Hyena: Intraspecies Readings of the Female Phallus". Signs. 28 (3). University of Chicago Press: 755–790. doi:10.1086/345320. S2CID 146640802.
  133. ^ Cole CJ, Townsend CR (9 May 2005). "Parthenogenetic lizards as vertebrate systems". Journal of Experimental Zoology. 256 (S4): 174–176. doi:10.1002/jez.1402560436. PMID 1974787.
  134. ^ Anon (9 June 2005). "Geckos: It's Not Always About Sex". Science Daily. Lewis & Clark College. Retrieved 14 December 2008.
  135. ^ Budzinski RM (1997). "Homosexuelles Verhalten bei Geckos der Gattung Phelsuma". Sauria. 19 (3): 33–34. Archived from the original on 7 June 2020. Retrieved 25 July 2020.
  136. ^ Binyon M (19 October 2017). "St Helena tortoise has a gay old time". The Times. Retrieved 20 October 2017.
  137. ^ a b c Scharf I, Martin OY (2013). "Same-sex sexual behavior in insects and arachnids: prevalence, causes, and consequences". Behavioral Ecology and Sociobiology. 67 (11): 1719–1730. Bibcode:2013BEcoS..67.1719S. doi:10.1007/s00265-013-1610-x. S2CID 10899013.
  138. ^ Dunkle SW (1991). "Head damage from mating attempts in dragonflies (Odonata:Anisoptera)". Entomological News. 102 (1): 37–41.
  139. ^ Utzeri C, Belfiore C (1990). "Anomalous tandems in Odonata". Fragmenta Entomologica. 22 (2): 271–288.
  140. ^ Gailey DA, Hall JC (April 1989). "Behavior and cytogenetics of fruitless in Drosophila melanogaster: different courtship defects caused by separate, closely linked lesions". Genetics. 121 (4): 773–85. doi:10.1093/genetics/121.4.773. PMC 1203660. PMID 2542123.
  141. ^ Yamamoto D, Nakano Y (November 1999). "Sexual behavior mutants revisited: molecular and cellular basis of Drosophila mating". Cellular and Molecular Life Sciences. 56 (7–8): 634–46. doi:10.1007/s000180050458. PMC 11146922. PMID 11212311. S2CID 22055533.
  142. ^ Yamamoto D, Ito H, Fujitani K (October 1996). "Genetic dissection of sexual orientation: behavioral, cellular, and molecular approaches in Drosophila melanogaster". Neuroscience Research. 26 (2): 95–107. doi:10.1016/S0168-0102(96)01087-5. PMID 8953572. S2CID 36582275.
  143. ^ Ferveur JF, Savarit F, O'Kane CJ, Sureau G, Greenspan RJ, Jallon JM (June 1997). "Genetic feminization of pheromones and its behavioral consequences in Drosophila males". Science. 276 (5318): 1555–8. doi:10.1126/science.276.5318.1555. PMID 9171057.
  144. ^ Yamamoto D, Fujitani K, Usui K, Ito H, Nakano Y (May 1998). "From behavior to development: genes for sexual behavior define the neuronal sexual switch in Drosophila". Mechanisms of Development. 73 (2): 135–46. doi:10.1016/S0925-4773(98)00042-2. PMID 9622612. S2CID 14531382.
  145. ^ Zhang SD, Odenwald WF (June 1995). "Misexpression of the white (w) gene triggers male-male courtship in Drosophila". Proceedings of the National Academy of Sciences of the United States of America. 92 (12): 5525–9. Bibcode:1995PNAS...92.5525Z. doi:10.1073/pnas.92.12.5525. PMC 41728. PMID 7777542.
  146. ^ Svetec N, Ferveur JF (March 2005). "Social experience and pheromonal perception can change male-male interactions in Drosophila melanogaster". The Journal of Experimental Biology. 208 (Pt 5): 891–8. doi:10.1242/jeb.01454. PMID 15755887.

Further reading

  • Roughgarden, Joan (2013). Evolution's Rainbow: Diversity, Gender, and Sexuality in Nature and People. University of California Press. ISBN 978-0520280458.
 
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