Non-reproductive sexual behavior 

consists of sexual activities animals participate in that 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, social conflict remediation, 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, and
acts of affection.[2] There have also been observations of sex with cub participants,[3] same-
sex sexual interaction,[4][5] as well as sex with dead animals.[6]

Social interaction and bonding

Lions are known to engage in sex to create bonds and interact with each other. Lions live in a
social group known as a pride that consists of 2–18 females and 1–7 males. The females
found in these prides were born into the pride. The males enter the pride from other prides.
The success of reproduction for each individual lion is dependent on the number of male lions
found in their social group. Male lions create coalitions and search for prides to take over.
Successful coalitions have usually created a strong bond with each other and will take over
prides. Once winning in a competition, all current males in the pride will be kicked out and
left to find another pride. While in search for another pride these males will often engage in
sexual behavior with each other; creating a strong bond in this new coalition created.[7][8]

Sex plays a fundamental role in the social lives of bonobos. Female bonobos have been
observed to engage in sexual activities to create bonds with dominant bonobos. Having
created this bond with the male, they will share food with each other and not compete with
each other.[1] All members of a bonobo group are potential sex partners, regardless of age
combination or gender combination. In "Biological Exuberance," Bagemihl writes: "when
new females (usually adolescents) join a troop, they often pair up with an older female with
whom they have most of their sexual and affectionate interactions." In addition, bonobos need
not limit themselves to a single partner: "These bond need not be exclusive - either party may
have sex with other females or males - but such mentorlike pairings can last for a year or
more until the newcomer is fully integrated into the troop." Pairings between younger and
older male bonobos are also common: "typically an adolescent male spreads his legs and
presents his erect penis to an adult male, who takes the shaft in his hand and caresses it with
up-and-down movements."[9]

Social pairings between youth and adult bonobos happen across gender combinations: "Both
adult males and females interact sexually with adolescents and juveniles (three-to-nine-year-
olds). In fact, young females go through a five-to-six-year period sometimes referred to as
adolescent sterility (although no pathology is involved) during which they actively participate
in heterosexual mating (often with adults) but never get pregnant. Sexual behavior between
adults and infants of both sexes is common - about a third of the time it is initiated by the
infant and may involve genital rubbing and full copulatory postures (including penetration of
an adult female by a male infant)."[9]

Social conflict remediation

Several species utilize sexual activity as a way to resolve disagreement. Bonobos are one
species famously known for using sexual behavior as a means of resolution of social conflict.
[4]

In a study concentrated on primate conflict resolution, researchers wanted to observe primates

in conflict. How primates coped and resolved conflicts was a main concern in this study.
Researchers stated that after primates participated in a heated, physical fight; both primates
involved in the fight would hug and have a mouth-to-mouth kiss. This action was considered
as a demonstration of affection and reconciliation.[1]

Proximate causes
Awareness in species is difficult to determine. Learned behaviors that have been
demonstrated in laboratories have provided good evidence that animals have instinct and a
reward system. The behavior of laboratory animals demonstrates a mental experience wherein
the animal's instincts tell it if it carries out a certain action, it will then receive what it needs.
[10] For example, the lab rat will push the lever because it knows food will fall out of the hole
in the wall. It does not need awareness, but it does seem to work on a reward system. The lab
rat learned the action needed to be fed.

Studies of the brain have proven that pleasure and displeasure are an important component in
the lives of animals.[11] It has been established that the limbic neural mechanism that
generates reactions are very similar across all mammals. Many studies have concentrated on
the brain reward system and how similar it is across mammals. Through extensive research,
scientists have been able to conclude that the brain reward system in animals is extremely
similar to that of humans. The mechanism of core pleasure reaction is significantly important
for animals and humans.[11]

Case study
Learn more
This section relies largely or entirely upon a single source. (August
2021)

In a case study, female Japanese macaques were studied to find evidence of possible female

copulatory orgasms. The frequency of orgasms did not correlate with the age or rank of the
Japanese macaques. Researchers observed that the longer and higher number of pelvic thrusts,
the longer copulation lasted. There was an orgasmic response in 80 of the 240 Japanese
macaques studied.[12]

Reward system
Evolutionary principles have predicted that the reward system is part of the proximate
mechanism underlying the behavior. Because animals possess a brain reward system they are
motivated to perform in different ways by desire and reinforced by pleasure.[10] Animals
establish security of food, shelter, social contact, and mating because proximate mechanism,
if they do not seek these necessities they will not survive.[13]

All vertebrates share similarities in body structure; they all have a skeleton, a nervous system,
a circulatory system, a digestive system and excretory system. Similar to humans, non-human
animals also have a sensory system. The sensory system is responsible for the basic five
senses from touch to tasting. Most of the physiological and biochemical responses found in
animals are found in humans. Neurophysiologists have not found any fundamental difference
between the structure and function of neurons and synapse between humans and other
animals.[10]

Case study
Learn more
This section relies largely or entirely upon a single source. (August
2021)

Recent studies using positron emission tomography (PET) and magnetic resonance

imaging (MRI) has provided evidence proving that chemical changes that occur with
emotions are similar between humans and non-human animals. In a study comparing guinea
pigs and humans, it was determined that the distress experienced by offspring separation in a
guinea pig and a human going through depression activates the same region of the brain.
[
citation needed] The opiate receptor was also examined, allowing observation of the pleasure
stimuli. In the procedure both a human and a rat had their receptors blocked with a certain
drug. Once receptors were blocked, both the rat and the human were exposed to pleasurable
food, but both were disinclined to eat the food.[14]

Types of behavior
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 for 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.[4] Male lions are known for head
rubbing, bats engage in licking, and mountain sheep rub horns and faces with each other.
[15] Animals have also engaged in kissing, touching of noses, mouths and muzzles have been
witnessed in African elephants, walruses, and mountain zebras.[5] 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.
[4] 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
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 where all insertion was anal. However, the practice might have been a
consequence of homosexual rearing, as the male orangutan in this dyad had had extensive
same–sex experience.[21]

A case of male homosexual anal insertion with the finger has also reported among orangutans,
[22] and Bruce Bagemihl mentions it as one of the homosexual practices recorded at least
once among male chimpanzees.[9]

Autoeroticism or masturbation
 Vervet monkey
It appears that many animals, both male and female, masturbate, both when partners are
available and otherwise.[23][24] For example, it has been observed in cats,[25] dogs,[26]
[27] male Cape ground squirrels,[28] male deer,[29][30][31] rhinoceroses,[32] boars,[33] and
male monkeys.[34][35]

A review from the University of Pennsylvania School of Veterinary Medicine says:[36]

[The] behavior known within the horse breeding industry as masturbation ...

involves normal periodic erections and penile movements. This behavior,
both from the descriptive field studies cited above and in extensive study of
domestic horses, is now understood as normal, frequent behavior of
male equids.[37] Attempting to inhibit or punish masturbation, for example
by tying a brush to the area of the flank underside where the penis rubs into
contact with the underside, which is still a common practice of horse
managers regionally around the world, often leads to increased masturbation
and disturbances of normal breeding behaviour.[38]
Castration does not prevent masturbation, as it is observed in geldings.[39] Masturbation is
common in both mares and stallions, before and after puberty.

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:

I am informed by a gentleman who is a recognized authority on goats, that

they sometimes take the penis into the mouth and produce actual orgasm,
thus practicing auto-fellatio. As regards ferrets ... "if the bitch, when in heat,
cannot obtain a dog [ie, male ferret] she pines and becomes ill. If a smooth
pebble is introduced into the hutch, she will masturbate upon it, thus
preserving her normal health for one season. But if this artificial substitute is
given to her a second season, she will not, as formerly, be content with it." ...
Blumenbach observed a bear act somewhat similarly on seeing other bears
coupling, and hyenas, according to Ploss and Bartels, have been seen
practicing mutual masturbation by licking each other's genitals.
In his 1999 book, Biological exuberance, Bruce Bagemihl documents that:

Autoeroticism also occurs widely among animals, both male and female. A
variety of creative techniques are used, including genital stimulation using
the hand or front paw (primates, Lions), foot (Vampire Bats, primates),
flipper (Walruses), or tail (Savanna Baboons), sometimes accompanied by
stimulation of the nipples (Rhesus Macaques, Bonobos); auto-fellating or
licking, sucking and/or nuzzling by a male of his own penis (Common
Chimpanzees, Savanna Bonobos, Vervet Monkeys, Squirrel Monkeys,
Thinhorn Sheep, Bharal, Aovdad, Dwarf Cavies); stimulation of the penis by
flipping or rubbing it against the belly or in its own sheath (White-tailed and
Mule Deer, Zebras and Takhi); spontaneous ejaculations (Mountain
Sheep, Warthogs, Spotted Hyenas); and stimulation of the genitals using
inanimate objects (found in several primates and cetaceans).[40]

Many birds masturbate by mounting and copulating with tufts of grass, leaves
or mounds of earth, and some mammals such as primates and dolphins also
rub their genitals against the ground or other surfaces to stimulate
themselves.[40]

Autoeroticism in female mammals, as well as heterosexual and homosexual

intercourse (especially in primates), often involves direct or indirect
 stimulation of the clitoris ... This organ is present in the females of all
mammalian species and several other animal groups.[40]
and that:

Apes and Monkeys use a variety of objects to masturbate with and even

deliberately create implements for sexual stimulation ... often in highly
creative ways.[40]
David Linden, professor of neuroscience at Johns Hopkins University, remarks that:

... perhaps the most creative form of animal masturbation is that of the male
bottlenose dolphin, which has been observed to wrap a live, wriggling eel
around its penis.[41]
Among elephants, female same-sex behaviours have been documented only in captivity
where they are known to masturbate one another with their trunks.[42]

Oral sexEdit
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.

Auto-fellatio or oral sex in animals is documented in spiders (Darwin's

bark[43] and widow[44]), brown bears,[45] stump-tailed macaques,[46] Tibetan macaques,
[47] wolves,[48] goats, primates, hyenas,[49][self-published source] bats,[50] cape ground
squirrels[28] 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.[51]

Homosexual behaviour
Main articles: Homosexual behavior in animals and Bisexuality § 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.
[52]
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.[53] Organisers of the Against Nature? exhibit stated that
"homosexuality has been observed among 1,500 species, and that in 500 of those it is well
documented."[54]

To turn the approach on its head: No species has been found in which
homosexual behaviour has not been shown to exist, with the exception of
species that never have sex at all, such as sea urchins and aphis. Moreover, a
part of the animal kingdom is hermaphroditic, truly bisexual. For them,
homosexuality is not an issue.[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.[4][13]
[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.

"Humans have created the myth that sexuality can be justified only by reproduction, which by
definition limits it to hetero sex," says Michael Bronski, author of The Pleasure Principle:
Culture, Backlash, and the Struggle for Gay Freedom. "But here is an animal society that uses
homosexuality to improve its social life."

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 — bonobos just love sex play," de Waal
said in an interview. "They have so much sex, it gets boring."

Homosexual behaviour is found in 6–10% of rams (sheep) and associated with variations in
cerebral mass distribution and chemical activity.[56]

Approximately eight percent of [male] rams exhibit sexual preferences [that

is, even when given a choice] for male partners (male-oriented rams) in
contrast to most rams, which prefer female partners (female-oriented rams).
We identified a cell group within the medial preoptic area/anterior
hypothalamus of age-matched adult sheep that was significantly larger in
adult rams than in ewes ...
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]

Homosexual pair-bonding and parentingEdit

Homosexual pair-bonding can be established several ways; two of the main ways are pair
bonding as partners or as companions.[9] As partners, both animals will engage in sexual
activities with each other. In companion bonding, sexual engagement is not necessary in the
relationship. This form of homosexuality is more of a partnership and friendship; they spend
all their time together. More than 70 species of birds engage in one of these two bonding.[9]

Homosexual parenting (sometimes referred to as cooperative breeding) occurs in a wide

variety of species in the animal kingdom.[9] Homosexual parenting can occur in different
ways, one of the most common being two females (typically related) coming together to help
one another raise their offspring. An example of this is in meadow vole populations. Summer
is peak breeding season for meadow voles; however, going into winter and spring there is a
division between the male and female meadow vole populations. They prefer communal
nesting (because of the thermoregulatory benefits), and therefore, in the winter and spring
female meadow voles will commonly not only nest with another female, but nurse their
offspring together as well. This kind of communal nursing, and same-sex social bonds, among
meadow voles is actually thought to benefit the young — increasing growth and survival
rates.[61]

Homosexual parenting is especially present among certain species of birds,[9] one of the most
famous examples being Laysan albatross. It is fairly uncommon among different species for
unrelated individuals of the same sex to raise offspring together, but female-female pairings
in Laysan albatross populations are one of the exceptions. This same-sex pairing and mutual
cooperation in chick-rearing often occurs in the Laysan albatross populations which have
uneven sex ratios (and an overall greater surplus of females). Also, Laysan albatross are
known for being monogamous, and this tendency actually allows same-sex parenting to
persist.[62]

Genital-genital rubbing
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".[63][64] 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.[65]

Genital rubbing has been observed once among male orangutans[22] 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.[66] It has been observed among bull manatees,
in conjunction with "kissing",[52] and is also common among homosexually active mammals.
[52]

Inter-species sex
See also: Animal hybrid, Sexual imprinting, and Zoophilia

A dog mates with a coyote to produce a dog-coyote hybrid.

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.
[67] Nevertheless, animals in the wild have been observed to attempt sexual activity with
other species.[68] It is mostly documented among species that belong to the same genus, but
sometimes occurs between species of distant taxa.[69] 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.[70]

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.[69] It may result in fitness loss
because of the waste of time, energy, and nutrients.[69]

Male sea otters have been observed forcibly copulating with seals,[71][72] and male seals

have been observed forcibly copulating with penguins.[73] Inter-species sexual behavior has
also been observed in sea lions.[74] 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.[69] 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.[69]

The Japanese macaque has been observed attempting to mate with the sika deer.[75]

Sex involving juveniles

Male stoats (Mustela erminea) will sometimes mate with infant females of their species.
[76] This is a natural part of their reproductive biology – they have a delayed gestation period,
so these females give birth the following year when they are fully grown.
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 ten-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.[77][78]

It appears to be common in the Adélie penguin.[79]

Among insects, there have been reports of immature females being forcibly copulated with.
[80]

Juvenile male chimpanzees have been recorded mounting and copulating with immature

chimps. Infants in bonobo societies are often involved in sexual behaviour.[81] 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.[82]

Necrophilia

A male black and white tegu mounts a female that has been dead for two days and attempts
to mate. Photo by Ivan Sazima.[83]
Main article: Necrophilia § 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 Adélie penguin.
[79] 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 feedback—well, didn't get any feedback."[84] The case study earned
Moeliker an Ig Nobel Prize in biology, awarded for research that cannot or should not be
reproduced.[85]