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Animal non-reproductive sexual behavior encompasses sexual activities that non-human animals participate in which do not lead to the reproduction of the species. Although procreation continues to be the primary explanation for sexual behavior in animals, recent observations on animal behavior have given alternative reasons for the engagement in sexual activities by animals.[1] Animals have been observed to engage in sex for social interaction bonding, exchange for significant materials, affection, mentorship pairings, sexual enjoyment, or as demonstration of social rank. Observed non-procreative sexual activities include non-copulatory mounting (without insertion, or by a female, or by a younger male who does not yet produce semen), oral sex, genital stimulation, anal stimulation, interspecies mating, same-sex sexual interaction,[2][3] and acts of affection, although it is doubted that they have done this since the beginning of their existence.[4] There have also been observations of sex with cub participants,[5] as well as sex with dead animals.[6]
A cow "bulling" during oestrus
Engagements of sexual activities during non-breeding seasons have been observed in the animal kingdom. Dolphins and Japanese macaques are two of the many species that engage in sexual activities that do not lead to fertilization. Great varieties of non-copulatory mounting are expressed in several species. Male lions engage in mounting with other male lions, especially when in search of another pride.[7] The varieties of mounting include mounting without erections, mounting with erection but no insertion, and mounting from the side.
Expressions of affection are displayed in the animal kingdom as well. Affectionate behaviors do not include insertion or genital rubbing, but are still seen as a manner of sexual behavior. An affectionate activity can be as simple as licking.[2] Felids are known for head rubbing, bats engage in licking, and mountain sheep rub horns and faces with each other.[15] Kissing, touching of noses, mouths and muzzles have been witnessed in African elephants, walruses, and mountain zebras.[3] Primates also engage in kissing that is incredibly similar to human display of kissing. Chimpanzees have full mouth-to-mouth contact, and bonobos kiss with their mouth open and mutual tongue stimulation.[2] There are a variety of acts to show affection such as African elephants intertwining their trunks, giraffes engaging in "necking", and Hanuman langurs cuddling with each other in a front to back sitting position.
Non-penetrative genital stimulation is very common throughout the animal kingdom. Different forms of self and partner genital stimulation have been observed in the animal kingdom. Oral sex has been observed throughout the animal kingdom, from dolphins to primates. Bonobos have been observed to transition from a simple demonstration of affection to non-penetrative genital stimulation.[1][15] Animals perform oral sex by licking, sucking or nuzzling the genitals of their partner.[9][15] Another form of genital stimulation is masturbation.
Masturbation is widespread throughout mammals for both males and females. It is less common in birds. There are several techniques, in which animals engage in masturbation from using paws, feet, flippers, tails, and sometimes using objects like sticks, pebbles, and leaves.[9] Masturbation occurs more often in primate species with large testes relative to their body size.[16]
Anal insertion with the penis (both in heterosexual and male homosexual dyads, i.e. pairs of animals) has been observed among some primate species. Male homosexual anal insertion has been recorded in Old World primate species, including gorillas, orangutans, and some members of the Macaca genus (namely, stumptail, rhesus, and Japanese macaques).[17][18][19] It has also been recorded in at least two New World primate species, the squirrel monkey and the spider monkey.[18][20] Morris (1970) also described one heterosexual orangutan dyad for whom all penetration was performed anally. However, the practice might have been a consequence of homosexual rearing, as the male orangutan in this dyad had had extensive sameโsex experience.[21] Anal insertion has also been observed among bonobo, with the observation described as 'anal intromission'. [22]
A case of male homosexual anal insertion with the finger has also been reported among orangutans,[23] and Bruce Bagemihl mentions it as one of the homosexual practices recorded at least once among male chimpanzees.[9]
Male vervet monkey masturbating
Many animals, both male and female, masturbate, both when partners are available and otherwise.[24][25] For example, it has been observed in cats,[26] dogs,[27][28] male Cape ground squirrels,[29] male deer,[30][31][32] rhinoceroses,[33] boars,[34] male monkeys,[35][36] cetaceans,[37] and otters.[38]
A review from the University of Pennsylvania School of Veterinary Medicine says:[39]
Sexologist Havelock Ellis in his 1927 Studies in the Psychology of Sex identified bulls, goats, sheep, camels and elephants as species known to practice autoeroticism, adding of some other species:
Animals of several species are documented as engaging in both autofellatio and oral sex. Although easily confused by laypeople, autofellatio and oral sex are separate, sexually oriented behaviors, distinct from non-sexual grooming or the investigation of scents.
Autofellatio and oral sex in animals is documented in spiders (Darwin's bark[46] and widow[47]), brown bears,[48] stump-tailed macaques,[49] Tibetan macaques,[50] wolves,[51] goats, primates, bats,[52] cape ground squirrels[29] and sheep (see section Masturbation for details).
In the greater short-nosed fruit bat, copulation by males is dorsoventral and the females lick the shaft or the base of the male's penis, but not the glans, which has already penetrated the vagina. While the females do this, the penis is not withdrawn and research has shown a positive relationship between length of the time that the penis is licked and the duration of copulation. Post copulation genital grooming has also been observed.[53]
Main articles: Homosexual behavior in animals and Bisexuality ยง Among other animals
Two male mallards, Anas platyrhynchos. 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.[54]
The presence of same-sex sexual behaviour was not scientifically reported on a large scale until recent times. Homosexual behaviour does occur in the animal kingdom outside humans, especially in social species, particularly in marine birds and mammals, monkeys, and the great apes. As of 1999, the scientific literature contained reports of homosexual behavior in at least 471 wild species.[55]
Homosexual behavior exists on a spectrum, and may or may not involve insertion. Apart from sexual activity, it can refer to homosexual pair-bonding, homosexual parenting and homosexual acts of affection. Engaging in homosexual behavior may allow species to obtain benefits such as gaining practice, relieving tension, and experiencing pleasure.[2][12][15] Georgetown University professor Janet Mann has specifically theorised that homosexual behaviour, at least in dolphins, is an evolutionary advantage that minimizes intraspecies aggression, especially among males.
After studying bonobos for his book Bonobo: The Forgotten Ape, primatologist Frans de Waal, a professor of psychology at Emory University in Atlanta, says that such expressions of intimacy are consistent with the homosexual behaviour of what he terms "the erotic champions of the world". "Same-sex, opposite-sex โ bonobos just love sex play," de Waal said in an interview. "They have so much sex, it gets boring."
Homosexual behaviour is found in 6โ10% of rams (sheep) and associated with variations in cerebral mass distribution and chemical activity.[56]
Male-male copulation has been observed in captive penguins[59] and homosexual behaviour has been observed among bats, in particular, the fruit bat.[60]
See also: Tribadism ยง Among female bonobos, and Frot ยง Among other animals
Genital-genital rubbing, or GG rubbing, among non-human animals is sexual activity in which one animal rubs his or her genitals against the genitals of another animal. The term GG rubbing is frequently used by primatologists to describe this type of sexual intimacy among female bonobos, and is stated to be the "bonobo's most typical sexual pattern, undocumented in any other primate".[61][62] The term is sometimes used in reference to GG rubbing among male bonobos, under the term "penis fencing", which is the non-human form of frot that human males engage in. Such rubbing between males is thought, according to varying evolutionary theorists, to have existed before the development of hominids into humans and bonobos, and may or may not have occurred in the homosexual activity of both of these genetically related species.[63]
Genital rubbing has been observed once among male orangutans[23] and several times in a small group of lar gibbons, where two males thrust their genitals together, sometimes resulting in ejaculation in one of the partners.[64] It has been observed among bull manatees, in conjunction with "kissing",[54] and is also common among homosexually active mammals.[54]
See also: Animal hybrid, Sexual imprinting, and Zoophilia
Some animals opportunistically mate with individuals of another species. This is more commonly observed in domesticated species and animals in captivity, possibly because captivity is associated with a decrease in aggression and an increase in sexual receptivity.[65] Nevertheless, animals in the wild have been observed to attempt sexual activity with other species.[66] It is mostly documented among species that belong to the same genus, but sometimes occurs between species of distant taxa.[67] Alfred Kinsey cites reports of sexual activity involving a female eland with an ostrich, a male dog with a chicken, a male monkey with a snake, and a female chimpanzee with a cat.[68]
A 2008 review of the literature found 44 species pairs that had been observed attempting interspecies mating, and 46 species pairs that had completed interspecies matings, not counting cases that had resulted in hybridization. Most were known from laboratory experiments, but field observations had also been made.[67] It may result in fitness loss because of the waste of time, energy, and nutrients.[67]
Male sea otters have been observed forcibly copulating with seals,[69][70] and male seals have been observed forcibly copulating with penguins.[71] Inter-species sexual behavior has also been observed in sea lions.[72] Male grasshoppers of the species Tetrix ceperoi often mount other species of either sex and even flies, but are normally repelled by the larger females.[67] Males of the spider mite species Panonychus citri copulate with female Panonychus mori mites almost as often as with their own species, even though it does not result in reproduction.[67]
The Japanese macaque has been observed attempting to mate with the sika deer.[73]
"Mock mating" of desert fox pups
In one reported observation, a male spotted hyena attempted to mate with a female hyena, but she succeeded in driving him off. He eventually turned to her 10-month-old cub, repeatedly mounting and ejaculating on it. The cub sometimes ignored this and sometimes struggled "slightly as if in play". The mother did not intervene.[74][75]
It appears to be common in the Adรฉlie penguin.[76]
Among insects, there have been reports of immature females being forcibly copulated with.[77]
Juvenile male chimpanzees have been recorded mounting and copulating with immature chimps. Infants in bonobo societies are often involved in sexual behaviour.[78] Immature male bonobos have been recorded initiating genital play with both adolescent and mature female bonobos. Copulation-like contact between immature bonobo males and mature female bonobos increases with age and continues until the male bonobo has reached juvenile age. In contrast, adult gorillas do not show any sexual interest in juvenile or infant members of their species. Primates regularly have sex in full view of infants, juveniles and younger members of their species.[79]
A male black and white tegu mounts a female that has been dead for two days and attempts to mate[80]
Main article: Necrophilia ยง Other animals
Necrophilia describes when an animal engages in a sexual act with a dead animal. It has been observed in mammals, birds, reptiles and frogs.[6] It sometimes occurs in the Adรฉlie penguin.[76] Homosexual necrophilia has been reported between two male mallard ducks. One duck was believed to be pursuing another duck with the goal of rape (a common aspect of duck sexual behaviour) when the second duck collided with a window and died immediately. The observer, Kees Moeliker, suggested that "when one died, the other one just went for it and didn't get any negative feedbackโwell, didn't get any feedback."[81] The case study earned Moeliker an Ig Nobel Prize in biology.[82]
Types of behavior
A male Eastern grey kangaroo attempting to copulate with a bush
A cow "bulling" during oestrusEngagements of sexual activities during non-breeding seasons have been observed in the animal kingdom. Dolphins and Japanese macaques are two of the many species that engage in sexual activities that do not lead to fertilization. Great varieties of non-copulatory mounting are expressed in several species. Male lions engage in mounting with other male lions, especially when in search of another pride.[7] The varieties of mounting include mounting without erections, mounting with erection but no insertion, and mounting from the side.
Expressions of affection are displayed in the animal kingdom as well. Affectionate behaviors do not include insertion or genital rubbing, but are still seen as a manner of sexual behavior. An affectionate activity can be as simple as licking.[2] Felids are known for head rubbing, bats engage in licking, and mountain sheep rub horns and faces with each other.[15] Kissing, touching of noses, mouths and muzzles have been witnessed in African elephants, walruses, and mountain zebras.[3] Primates also engage in kissing that is incredibly similar to human display of kissing. Chimpanzees have full mouth-to-mouth contact, and bonobos kiss with their mouth open and mutual tongue stimulation.[2] There are a variety of acts to show affection such as African elephants intertwining their trunks, giraffes engaging in "necking", and Hanuman langurs cuddling with each other in a front to back sitting position.
Non-penetrative genital stimulation is very common throughout the animal kingdom. Different forms of self and partner genital stimulation have been observed in the animal kingdom. Oral sex has been observed throughout the animal kingdom, from dolphins to primates. Bonobos have been observed to transition from a simple demonstration of affection to non-penetrative genital stimulation.[1][15] Animals perform oral sex by licking, sucking or nuzzling the genitals of their partner.[9][15] Another form of genital stimulation is masturbation.
Masturbation is widespread throughout mammals for both males and females. It is less common in birds. There are several techniques, in which animals engage in masturbation from using paws, feet, flippers, tails, and sometimes using objects like sticks, pebbles, and leaves.[9] Masturbation occurs more often in primate species with large testes relative to their body size.[16]
Anal insertion
editAnal insertion with the penis (both in heterosexual and male homosexual dyads, i.e. pairs of animals) has been observed among some primate species. Male homosexual anal insertion has been recorded in Old World primate species, including gorillas, orangutans, and some members of the Macaca genus (namely, stumptail, rhesus, and Japanese macaques).[17][18][19] It has also been recorded in at least two New World primate species, the squirrel monkey and the spider monkey.[18][20] Morris (1970) also described one heterosexual orangutan dyad for whom all penetration was performed anally. However, the practice might have been a consequence of homosexual rearing, as the male orangutan in this dyad had had extensive sameโsex experience.[21] Anal insertion has also been observed among bonobo, with the observation described as 'anal intromission'. [22]
A case of male homosexual anal insertion with the finger has also been reported among orangutans,[23] and Bruce Bagemihl mentions it as one of the homosexual practices recorded at least once among male chimpanzees.[9]
Autoeroticism or masturbation
edit_(6011903243).jpg)
Male vervet monkey masturbatingMany animals, both male and female, masturbate, both when partners are available and otherwise.[24][25] For example, it has been observed in cats,[26] dogs,[27][28] male Cape ground squirrels,[29] male deer,[30][31][32] rhinoceroses,[33] boars,[34] male monkeys,[35][36] cetaceans,[37] and otters.[38]
A review from the University of Pennsylvania School of Veterinary Medicine says:[39]
Castration does not prevent masturbation, as it is observed in geldings.[42] Masturbation is common in both mares and stallions, before and after puberty.[citation needed]
Sexologist Havelock Ellis in his 1927 Studies in the Psychology of Sex identified bulls, goats, sheep, camels and elephants as species known to practice autoeroticism, adding of some other species:
In his 1999 book, Biological Exuberance, Bruce Bagemihl documents that:
and that:
David Linden, professor of neuroscience at Johns Hopkins University, remarks that:
Among elephants, female same-sex behaviours have been documented only in captivity where they are known to masturbate one another with their trunks.[45]
Oral sex
editAnimals of several species are documented as engaging in both autofellatio and oral sex. Although easily confused by laypeople, autofellatio and oral sex are separate, sexually oriented behaviors, distinct from non-sexual grooming or the investigation of scents.
Autofellatio and oral sex in animals is documented in spiders (Darwin's bark[46] and widow[47]), brown bears,[48] stump-tailed macaques,[49] Tibetan macaques,[50] wolves,[51] goats, primates, bats,[52] cape ground squirrels[29] and sheep (see section Masturbation for details).
In the greater short-nosed fruit bat, copulation by males is dorsoventral and the females lick the shaft or the base of the male's penis, but not the glans, which has already penetrated the vagina. While the females do this, the penis is not withdrawn and research has shown a positive relationship between length of the time that the penis is licked and the duration of copulation. Post copulation genital grooming has also been observed.[53]
Homosexual behavior
editMain articles: Homosexual behavior in animals and Bisexuality ยง Among other animals
Two male mallards, Anas platyrhynchos. 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.[54]The presence of same-sex sexual behaviour was not scientifically reported on a large scale until recent times. Homosexual behaviour does occur in the animal kingdom outside humans, especially in social species, particularly in marine birds and mammals, monkeys, and the great apes. As of 1999, the scientific literature contained reports of homosexual behavior in at least 471 wild species.[55]
Homosexual behavior exists on a spectrum, and may or may not involve insertion. Apart from sexual activity, it can refer to homosexual pair-bonding, homosexual parenting and homosexual acts of affection. Engaging in homosexual behavior may allow species to obtain benefits such as gaining practice, relieving tension, and experiencing pleasure.[2][12][15] Georgetown University professor Janet Mann has specifically theorised that homosexual behaviour, at least in dolphins, is an evolutionary advantage that minimizes intraspecies aggression, especially among males.
After studying bonobos for his book Bonobo: The Forgotten Ape, primatologist Frans de Waal, a professor of psychology at Emory University in Atlanta, says that such expressions of intimacy are consistent with the homosexual behaviour of what he terms "the erotic champions of the world". "Same-sex, opposite-sex โ bonobos just love sex play," de Waal said in an interview. "They have so much sex, it gets boring."
Homosexual behaviour is found in 6โ10% of rams (sheep) and associated with variations in cerebral mass distribution and chemical activity.[56]
Male bighorn sheep are divisible into two kinds: the typical males among whom homosexual behaviour, including intercourse, is common and "effeminate sheep", or "behavioural transvestites", which are not known to engage in homosexual behaviour.[57][58]
Male-male copulation has been observed in captive penguins[59] and homosexual behaviour has been observed among bats, in particular, the fruit bat.[60]
Genital-genital rubbing
editSee also: Tribadism ยง Among female bonobos, and Frot ยง Among other animals
Genital-genital rubbing, or GG rubbing, among non-human animals is sexual activity in which one animal rubs his or her genitals against the genitals of another animal. The term GG rubbing is frequently used by primatologists to describe this type of sexual intimacy among female bonobos, and is stated to be the "bonobo's most typical sexual pattern, undocumented in any other primate".[61][62] The term is sometimes used in reference to GG rubbing among male bonobos, under the term "penis fencing", which is the non-human form of frot that human males engage in. Such rubbing between males is thought, according to varying evolutionary theorists, to have existed before the development of hominids into humans and bonobos, and may or may not have occurred in the homosexual activity of both of these genetically related species.[63]
Genital rubbing has been observed once among male orangutans[23] and several times in a small group of lar gibbons, where two males thrust their genitals together, sometimes resulting in ejaculation in one of the partners.[64] It has been observed among bull manatees, in conjunction with "kissing",[54] and is also common among homosexually active mammals.[54]
Inter-species sex
editSee also: Animal hybrid, Sexual imprinting, and Zoophilia
Some animals opportunistically mate with individuals of another species. This is more commonly observed in domesticated species and animals in captivity, possibly because captivity is associated with a decrease in aggression and an increase in sexual receptivity.[65] Nevertheless, animals in the wild have been observed to attempt sexual activity with other species.[66] It is mostly documented among species that belong to the same genus, but sometimes occurs between species of distant taxa.[67] Alfred Kinsey cites reports of sexual activity involving a female eland with an ostrich, a male dog with a chicken, a male monkey with a snake, and a female chimpanzee with a cat.[68]
A 2008 review of the literature found 44 species pairs that had been observed attempting interspecies mating, and 46 species pairs that had completed interspecies matings, not counting cases that had resulted in hybridization. Most were known from laboratory experiments, but field observations had also been made.[67] It may result in fitness loss because of the waste of time, energy, and nutrients.[67]
Male sea otters have been observed forcibly copulating with seals,[69][70] and male seals have been observed forcibly copulating with penguins.[71] Inter-species sexual behavior has also been observed in sea lions.[72] Male grasshoppers of the species Tetrix ceperoi often mount other species of either sex and even flies, but are normally repelled by the larger females.[67] Males of the spider mite species Panonychus citri copulate with female Panonychus mori mites almost as often as with their own species, even though it does not result in reproduction.[67]
The Japanese macaque has been observed attempting to mate with the sika deer.[73]
Sex involving juveniles
edit
"Mock mating" of desert fox pupsIn one reported observation, a male spotted hyena attempted to mate with a female hyena, but she succeeded in driving him off. He eventually turned to her 10-month-old cub, repeatedly mounting and ejaculating on it. The cub sometimes ignored this and sometimes struggled "slightly as if in play". The mother did not intervene.[74][75]
It appears to be common in the Adรฉlie penguin.[76]
Among insects, there have been reports of immature females being forcibly copulated with.[77]
Juvenile male chimpanzees have been recorded mounting and copulating with immature chimps. Infants in bonobo societies are often involved in sexual behaviour.[78] Immature male bonobos have been recorded initiating genital play with both adolescent and mature female bonobos. Copulation-like contact between immature bonobo males and mature female bonobos increases with age and continues until the male bonobo has reached juvenile age. In contrast, adult gorillas do not show any sexual interest in juvenile or infant members of their species. Primates regularly have sex in full view of infants, juveniles and younger members of their species.[79]
Necrophilia
edit
A male black and white tegu mounts a female that has been dead for two days and attempts to mate[80]Main article: Necrophilia ยง Other animals
Necrophilia describes when an animal engages in a sexual act with a dead animal. It has been observed in mammals, birds, reptiles and frogs.[6] It sometimes occurs in the Adรฉlie penguin.[76] Homosexual necrophilia has been reported between two male mallard ducks. One duck was believed to be pursuing another duck with the goal of rape (a common aspect of duck sexual behaviour) when the second duck collided with a window and died immediately. The observer, Kees Moeliker, suggested that "when one died, the other one just went for it and didn't get any negative feedbackโwell, didn't get any feedback."[81] The case study earned Moeliker an Ig Nobel Prize in biology.[82]
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