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In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive,
reproductive The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are a ...
, and
urinary tract The urinary system, also known as the urinary tract or renal system, consists of the kidneys, ureters, bladder, and the urethra. The purpose of the urinary system is to eliminate waste from the body, regulate blood volume and blood pressure, c ...
s (if present) of many
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
animals. All
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
s,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
s and birds, and a few mammals (
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals ( Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brai ...
s, tenrecs,
golden mole Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae and as such they are taxonomically distinct from the true moles, family Talpidae, and other mole-like families, all ...
s, and marsupial moles), have this orifice, from which they excrete both
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excreted from the body through the urethra. Cellul ...
and
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a rela ...
; this is in contrast to most
placental Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
mammals, which have two or three separate orifices for evacuation. Excretory openings with analogous purpose in some
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s are also sometimes referred to as cloacae. Mating through the cloaca is known as cloacal
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
, commonly referred to as cloacal kiss. The cloacal region is also often associated with a secretory organ, the cloacal gland, which has been implicated in the scent-marking behavior of some reptiles, marsupials, amphibians, and
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals ( Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brai ...
s.


Etymology

The word is from the
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
verb ''cluo'', "(I) cleanse", thus the noun ''cloaca'', " sewer, drain".


Birds

Birds reproduce using their cloaca; this occurs during a cloacal kiss in most birds. Birds that mate using this method touch their cloacae together, in some species for only a few seconds, sufficient time for
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
to be transferred from the male to the female. For some birds, such as
ostriches Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There a ...
,
cassowaries Cassowaries ( tpi, muruk, id, kasuari) are flightless birds of the genus ''Casuarius'' in the order Casuariiformes. They are classified as ratites (flightless birds without a keel on their sternum bones) and are native to the tropical fo ...
, kiwi,
geese A goose ( : geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera '' Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some other birds, mostly related to the she ...
, and some species of
swan Swans are birds of the family Anatidae within the genus ''Cygnus''. The swans' closest relatives include the geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form the tribe Cygnini. Som ...
s and
duck Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form ...
s, the males do not use the cloaca for reproduction, but have a
phallus A phallus is a penis (especially when erect), an object that resembles a penis, or a mimetic image of an erect penis. In art history a figure with an erect penis is described as ithyphallic. Any object that symbolically—or, more precise ...
. One study has looked into birds that use their cloaca for cooling. The cloaca in birds may also be referred to as the vent. Among falconers the word vent is also a verb meaning "to defecate".


Fish

Among fish, a true cloaca is present only in
elasmobranch Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of g ...
s (sharks and rays) and
lobe-finned fish Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includ ...
es. In
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s and in some
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
es, part of the cloaca remains in the adult to receive the urinary and reproductive ducts, although the anus always opens separately. In
chimaera Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively. At ...
s and most
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tele ...
s, however, all three openings are entirely separated.


Mammals

With a few exceptions noted below, mammals have no cloaca. Even in the marsupials that have one, the cloaca is partially subdivided into separate regions for the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, ...
and
urethra The urethra (from Greek οὐρήθρα – ''ourḗthrā'') is a tube that connects the urinary bladder to the urinary meatus for the removal of urine from the body of both females and males. In human females and other primates, the urethra ...
.


Monotremes

The
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals ( Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brai ...
s (egg-laying mammals) possess a true cloaca.


Marsupials

In
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in ...
s (and a few birds), the genital tract is separate from the anus, but a trace of the original cloaca does remain externally. This is one of the features of marsupials (and monotremes) that suggest their basal nature, as the
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are dis ...
s from which mammals evolved possessed a cloaca, and the earliest animals to diverge into the mammalian class would most likely have had this feature, too. Unlike other marsupials, marsupial moles have a true cloaca, a fact that has been used to argue against a marsupial identity for these mammals.


Placentals

Most adult
placental mammal Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupialia. Placentalia contains the vast majority of extant mammals, which are partly distinguishe ...
s have no remaining trace of the cloaca. In the embryo, the embryonic cloaca divides into a posterior region that becomes part of the anus, and an anterior region that has different fates depending on the sex of the individual: in females, it develops into the
vestibule Vestibule or Vestibulum can have the following meanings, each primarily based upon a common origin, from early 17th century French, derived from Latin ''vestibulum, -i n.'' "entrance court". Anatomy In general, vestibule is a small space or cavity ...
that receives the urethra and vagina, while in males it forms the entirety of the
penile urethra The spongy urethra (cavernous portion of urethra, penile urethra) is the longest part of the male urethra, and is contained in the corpus spongiosum of the penis. It is about 15 cm long, and extends from the termination of the membranous p ...
. However, the tenrecs and
golden mole Golden moles are small insectivorous burrowing mammals endemic to Sub-Saharan Africa. They comprise the family Chrysochloridae and as such they are taxonomically distinct from the true moles, family Talpidae, and other mole-like families, all ...
s, small placental mammals native to Africa, as well as some
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to differ ...
s retain a cloaca as adults. Being placental animals, humans only have an embryonic cloaca, which is split up into separate tracts during the
development of the urinary and reproductive organs The development of the urinary system begins during prenatal development, and relates to the development of the urogenital system – both the organs of the urinary system and the sex organs of the reproductive system. The development continues a ...
. However, a few human
congenital disorder A birth defect, also known as a congenital disorder, is an abnormal condition that is present at birth regardless of its cause. Birth defects may result in disabilities that may be physical, intellectual, or developmental. The disabilities c ...
s result in persons being born with a cloaca, including persistent cloaca and sirenomelia (mermaid syndrome).


Reptiles

In reptiles, the cloaca consists of the ''urodeum'', ''proctodeum'', and ''coprodeum''. Some species have modified cloacae for increased gas exchange (see Reptile respiration and
Reptile reproduction Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
). This is where reproductive activity occurs.


Cloacal respiration in animals

Some
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s, especially those specialized in diving, are highly reliant on cloacal
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellul ...
during dives. They accomplish this by having a pair of accessory air bladders connected to the cloaca which can absorb oxygen from the water. Various
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
, as well as
polychaete Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are made ...
worms and even
crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all th ...
s, are specialized to take advantage of the constant flow of water through the cloacal respiratory tree of
sea cucumbers Sea cucumbers are echinoderms from the class Holothuroidea (). They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. Sea cucumbers are found on the sea floor worldwide. The number of holothurian ...
while simultaneously gaining the protection of living within the sea cucumber itself. At night, many of these species emerge from the anus of the sea cucumber in search of food.Aquarium Invertebrates by Rob Toonen, Ph.D.
/ref>


See also

*
Cloaca (embryology) The cloaca is a structure in the development of the urinary and reproductive organs. The hind-gut is at first prolonged backward into the body-stalk as the tube of the allantois; but, with the growth and flexure of the tail-end of the embryo, ...


References

{{Authority control Animal anatomy Bird anatomy Sex organs Animal reproductive system