Cloacal
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A cloaca ( ), : cloacae ( or ), or vent, is the rear orifice that serves as the only opening for the digestive (
rectum The rectum (: rectums or recta) is the final straight portion of the large intestine in humans and some other mammals, and the gut in others. Before expulsion through the anus or cloaca, the rectum stores the feces temporarily. The adult ...
),
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 al ...
, and urinary tracts (if present) of many
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
animals. All
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s,
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,
birds 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 ...
,
cartilaginous fish Chondrichthyes (; ) is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...
and a few mammals (
monotreme Monotremes () are mammals of the order Monotremata. They are the only group of living mammals that lay eggs, rather than bearing live young. The extant monotreme species are the platypus and the four species of echidnas. Monotremes are typified ...
s, afrosoricids, and marsupial moles, etc.) have this orifice, from which they excrete both
urine Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...
and
feces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) 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 ...
; this is in contrast to most placental mammals, which have separate orifices for evacuation and reproduction. Excretory openings with analogous purpose in some
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s are also sometimes called cloacae. Mating through the cloaca is called cloacal copulation and cloacal kissing. 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 mammals of the order Monotremata. They are the only group of living mammals that lay eggs, rather than bearing live young. The extant monotreme species are the platypus and the four species of echidnas. Monotremes are typified ...
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 spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
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 (: sperm or sperms) is the male reproductive Cell (biology), 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 ...
to be transferred from the male to the female. For palaeognaths and
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which i ...
, the males do not use the cloaca for reproduction, but have a phallus. One study has looked into birds that use their cloaca for cooling. 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 modern sharks ( division Selachii), and batomorphs (division Batomorphi, including rays, skates, and sawfish). Members of this subclass are characterised by h ...
s (sharks and rays) and
lobe-finned fish Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
es. In
lamprey Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
s and in some
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
es, part of the cloaca remains in the adult to receive the urinary and reproductive ducts, although the anus always opens separately. In chimaeras and most
teleost Teleostei (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...
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 In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...
and
urethra The urethra (: urethras or urethrae) is the tube that connects the urinary bladder to the urinary meatus, through which Placentalia, placental mammals Urination, urinate and Ejaculation, ejaculate. The external urethral sphincter is a striated ...
.


Monotremes

The
monotreme Monotremes () are mammals of the order Monotremata. They are the only group of living mammals that lay eggs, rather than bearing live young. The extant monotreme species are the platypus and the four species of echidnas. Monotremes are typified ...
s (egg-laying mammals) possess a true cloaca.


Marsupials

In
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
s, 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 tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s from which mammals evolved had a cloaca, and probably so did the earliest
mammals A mammal () is a vertebrate animal of the 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 middle e ...
. Unlike other marsupials, marsupial moles have a true cloaca. This fact has been used to argue that they are not marsupials.


Placentals

Most adult
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 distinguished ...
s have no cloaca. In the embryo, the embryonic cloaca divides into a posterior region that becomes part of the
anus In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...
, and an anterior region that develops depending on sex: in males, it forms the penile urethra, while in females, it develops into the vestibule or
urogenital sinus The urogenital sinus is a body part of a human or other Placentalia, placental only present in the development of the urinary system, development of the urinary and development of the reproductive organs, reproductive organs. It is the ventral p ...
that receives the urethra and vagina. However, some placentals retain a cloaca as adults: those are members of the order Afrosoricida (small mammals native to Africa) as well as
pika A pika ( , or ) is a small, mountain-dwelling mammal native to Asia and North America. With short limbs, a very round body, an even coat of fur, and no external tail, they resemble their close relative the rabbit, but with short, rounded ears. ...
s,
beaver Beavers (genus ''Castor'') are large, semiaquatic rodents of the Northern Hemisphere. There are two existing species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers are the second-large ...
s, and 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 dif ...
s. Being placental animals, humans have an embryonic cloaca which divides 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 as ...
. However, a few human
congenital disorder A birth defect is an abnormal condition that is present at childbirth, birth, regardless of its cause. Birth defects may result in disability, disabilities that may be physical disability, physical, intellectual disability, intellectual, or dev ...
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). This is where reproductive activity occurs.


Cloacal respiration in animals

Some
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s, especially those specialized in diving, are highly reliant on cloacal respiration during dives. They accomplish this by having a pair of accessory air bladders connected to the cloaca, which can absorb oxygen from the water. Sea cucumbers use cloacal respiration. The constant flow of water through it has allowed various
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
,
polychaete Polychaeta () is a paraphyletic class of generally marine Annelid, annelid worms, common name, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called c ...
worms and even
crab Crabs are decapod crustaceans of the infraorder Brachyura (meaning "short tailed" in Greek language, Greek), which typically have a very short projecting tail-like abdomen#Arthropoda, abdomen, usually hidden entirely under the Thorax (arthropo ...
s to specialize to take advantage of it while living protected inside the cucumber. At night, many of these species emerge through the anus of the sea cucumber in search of food.Aquarium Invertebrates by Rob Toonen, Ph.D.
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See also

* Cloaca (embryology)


References

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