In
zoology
Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...
, copulation is
animal sexual behavior in which a male introduces
sperm into the female's body, especially directly into her
reproductive tract. This is an aspect of
mating. Many
aquatic animal
An aquatic animal is any animal, whether vertebrate or invertebrate, that lives in a body of water for all or most of its lifetime. Aquatic animals generally conduct gas exchange in water by extracting dissolved oxygen via specialised respirato ...
s use
external fertilization, whereas
internal fertilization may have developed from a need to maintain
gamete
A gamete ( ) is a Ploidy#Haploid and monoploid, haploid cell that fuses with another haploid cell during fertilization in organisms that Sexual reproduction, reproduce sexually. Gametes are an organism's reproductive cells, also referred to as s ...
s in a liquid medium in the
Late Ordovician epoch. Internal fertilization with many
vertebrates (such as all
reptile
Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s, some
fish, and most
bird
Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s) occurs via
cloacal copulation, known as cloacal kiss (see also
hemipenis), while most
mammal
A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s copulate
vagina
In mammals and other animals, the vagina (: vaginas or vaginae) is the elastic, muscular sex organ, reproductive organ of the female genital tract. In humans, it extends from the vulval vestibule to the cervix (neck of the uterus). The #Vag ...
lly, and many
basal vertebrates reproduce sexually with external fertilization.
In spiders and insects
Spider
Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
s are often confused with
insect
Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s, but they are not insects; instead, they are
arachnid
Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s. Spiders have separate male and female sexes. Before mating and copulation, the male spider spins a small web and ejaculates on to it. He then stores the sperm in reservoirs on his large
pedipalps, from which he transfers sperm to the female's genitals. The females can
store sperm indefinitely.

For primitive insects, the male deposits
spermatozoa on the substrate, sometimes stored within a special structure;
courtship involves inducing the female to take up the sperm package into her genital opening, but there is no actual copulation. In groups that have reproduction similar to spiders, such as
dragonflies, males extrude sperm into secondary copulatory structures removed from their genital opening, which are then used to inseminate the female. In dragonflies, it is a set of modified
sternites on the second abdominal segment. In advanced groups of insects, the male uses its
aedeagus, a structure formed from the terminal segments of the abdomen, to deposit sperm directly (though sometimes in a capsule called a ''
spermatophore'') into the female's reproductive tract.
In mammals
Sexual behavior can be classified into behavioral states associated with
reward motivation ("
wanting"), reward consummation also known as pleasure ("liking"), and
satiety ("inhibition");
these behavioral states are regulated in mammals by reward-based sexual learning, fluctuations in various neurochemicals (i.e.,
dopamine −
sexual desire also known as "wanting";
norepinephrine −
sexual arousal;
oxytocin
Oxytocin is a peptide hormone and neuropeptide normally produced in the hypothalamus and released by the posterior pituitary. Present in animals since early stages of evolution, in humans it plays roles in behavior that include Human bonding, ...
and
melanocortins −
sexual attraction), and gonadal hormone cycles and further influenced by
sex pheromones and
motor reflexes (i.e.,
lordosis behaviour) in some mammals.
These behavioral states correlate with the phases of the
human sexual response cycle: motivation − excitement; consummation − plateau and orgasm; satiety − refraction.
Sexual learning (a form of
associative learning) occurs when an animal starts to associate bodily features, personality, contextual cues, and other stimuli with genitally-induced sexual pleasure.
Once formed, these associations in turn impinge upon both sexual wanting and sexual liking.
In most female mammals, the act of copulation is controlled by several innate neurobiological processes, including the motor sexual reflex of lordosis. In males, the act of copulation is more complex, because some learning is necessary, but the innate processes (retrocontrol of
penis
A penis (; : penises or penes) is a sex organ through which male and hermaphrodite animals expel semen during copulation (zoology), copulation, and through which male placental mammals and marsupials also Urination, urinate.
The term ''pen ...
intromission in the
vagina
In mammals and other animals, the vagina (: vaginas or vaginae) is the elastic, muscular sex organ, reproductive organ of the female genital tract. In humans, it extends from the vulval vestibule to the cervix (neck of the uterus). The #Vag ...
, rhythmic movement of the pelvis, detection of female pheromones) are specific to copulation. These innate processes direct heterosexual copulation. Female lordosis behaviour became secondary in
Hominidae and is non-functional in humans. Mammals usually copulate in a dorso-ventral posture, although some primate species copulate in a ventro-vental posture.
Most mammals possess a
vomeronasal organ that is involved in pheromone detection, including
sex pheromones.
Despite the fact that humans do not possess this organ, adult humans appear to be sensitive to certain mammalian pheromones that putative
pheromone receptor proteins in the
olfactory epithelium are capable of detecting.
While sex pheromones clearly play a role in modifying sexual behavior in some mammals, the capacity for general pheromone detection and the involvement of
pheromones in human sexual behavior has not yet been determined.
The duration of copulation varies significantly between mammal species, and may be correlated with body mass, lasting longer in large mammals than in small mammals. The duration of copulation may also be correlated with the length of the
baculum in mammals.
Male mammals
ejaculate semen through the penis into the female reproductive tract during copulation. Ejaculation usually occurs after only one intromission in humans,
canids, and
ungulates, but occurs after multiple intromissions in most mammal species.
Copulation can
induce ovulation in mammal species that do not
ovulate spontaneously.
File:Dubim2.webm, Copulating brown bears
Lions (Panthera leo) couple mating ....webm, Copulating lion
The lion (''Panthera leo'') is a large Felidae, cat of the genus ''Panthera'', native to Sub-Saharan Africa and India. It has a muscular, broad-chested body (biology), body; a short, rounded head; round ears; and a dark, hairy tuft at the ...
s
Australian Wildlife- Mating Kangaroos.webm, Copulating grey kangaroos
20150821 Aldabra giant tortoises mating in Tierpark Hellabrunn.webm, Copulating Aldabra giant tortoises
Copulation of Magellanic Penguins.ogv, Copulating Magellanic penguins
See also
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Pelvic thrust
Notes
References
Bibliography
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{{Animal sexual behavior
Animal physiology
Animal sexuality
Mammalian sexuality
Mating
Ethology