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Cloaca

From Wikipedia, the free encyclopedia
Posterior opening in zoology
For other uses, seeCloaca (disambiguation).
Cloaca of ared-tailed hawk

Acloaca (/klˈkə/ kloh-AY-kə),pl.:cloacae (/klˈsi/kloh-AY-see or/klˈki/kloh-AY-kee), orvent, is the rearorifice that serves as the only opening for thedigestive (rectum),reproductive, andurinary tracts (if present) of manyvertebrate animals. Allamphibians,reptiles,birds,cartilaginous fish[1] and a few mammals (monotremes,afrosoricids, andmarsupial moles) have this orifice, from which they excrete bothurine andfeces; this is in contrast to mostplacental mammals, which have separate orifices for evacuation and reproduction.Excretory openings with analogous purpose in someinvertebrates are also sometimes called cloacae. Mating through the cloaca is called cloacalcopulation 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,[2] marsupials,[3] amphibians, andmonotremes.[4]

Etymology

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The word is from theLatin verbcluo, "(I) cleanse", thus the nouncloaca, "sewer, drain".[5][6][7]

Birds

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See also:Bird anatomy § Reproductive and urogenital systems
Cloaca of a female bird
Cloaca of a male bird
Aroseate spoonbill excreting urine in flight

Birds reproduce using their cloaca; this occurs during a cloacal kiss in most birds.[8] Birds that mate using this method touch their cloacae together, in some species for only a few seconds, sufficient time forsperm to be transferred from the male to the female.[9] Forpalaeognaths andwaterfowl, the males do not use the cloaca for reproduction, but have aphallus.[10]

One study[11] has looked into birds that use their cloaca for cooling.[12]

Amongfalconers, the word vent is also a verb meaning "to defecate".

Fish

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Among fish, a true cloaca is present only inelasmobranchs (sharks and rays) andlobe-finned fishes. Inlampreys and in someray-finned fishes, part of the cloaca remains in the adult to receive the urinary and reproductive ducts, although the anus always opens separately. Inchimaeras and mostteleosts, however, all three openings are entirely separated.[13]

Mammals

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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 theanus andurethra.

Diagrams to illustrate the changes in the cloaca in mammals during development.A, early embryonic stage, showing the cloaca receiving the urinary bladder, the rectum, and the Wolffian duct, as in non-therian vertebrates.B, later stage, showing the beginning of the fold which divides the cloaca into a ventralurogenital sinus which receives theurinary bladder, Wolffian ducts, andureters, and into a dorsal part which receives therectum.C, further progress of the fold, dividing the cloaca into urogenital sinus and rectum; the ureter has separated from the Wolffian duct and is shifting anteriorly.D, completion of the fold, showing complete separation of the cloaca into ventral urogenital sinus and dorsal rectum.[14]

Monotremes

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Themonotremes (egg-laying mammals) possess a true cloaca.[15]

Marsupials

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Further information:Marsupial § Reproductive system
Cloacal opening in anAustralian brushtail possum

Inmarsupials, the genital tract is separate from the anus, but a trace of the original cloaca does remain externally.[13] This is one of the features of marsupials (and monotremes) that suggest their basal nature, as theamniotes from which mammals evolved had a cloaca, and probably so did the earliestmammals.

Unlike other marsupials,marsupial moles have a true cloaca.[16] This fact has been used to argue that they are not marsupials.[17][18][unreliable source?]

Placental mammals

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Most adultplacental mammals have no cloaca. In the embryo, theembryonic cloaca divides into a posterior region that becomes part of theanus, and an anterior region that develops depending on sex: in males, it forms thepenile urethra, while in females, it develops into thevestibule orurogenital sinus that receives the urethra and vagina.[13][19] However, some placental mammals retain a cloaca as adults: those are members of the orderAfrosoricida (small mammals native to Africa) as well aspikas,beavers, and someshrews.[20][21][22][23]

Being placental mammals, humans have an embryonic cloaca which divides into separate tracts during thedevelopment of the urinary and reproductive organs. However, a few humancongenital disorders result in persons being born with a cloaca, includingpersistent cloaca andsirenomelia (mermaid syndrome).

Reptiles

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In reptiles, the cloaca consists of theurodeum,proctodeum, andcoprodeum.[24][25] Some species have modified cloacae for increased gas exchange (seereptile respiration andreptile reproduction). This is where reproductive activity occurs.[26]

Cloacal respiration in animals

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Further information:Cloacal respiration

Someturtles, especially those specialized in diving, are highly reliant on cloacalrespiration during dives.[27] They accomplish this by having a pair of accessory air bladders connected to the cloaca, which can absorb oxygen from the water.[28]

Sea cucumbers use cloacal respiration. The constant flow of water through it has allowed variousfish,polychaete worms and evencrabs 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.[29]

See also

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References

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  1. ^"Fish Groups".Florida Museum. Retrieved2 February 2025.
  2. ^Carl Gans; David Crews (June 1992).Hormones, Brain, and Behavior. University of Chicago Press.ISBN 978-0-226-28124-7.
  3. ^R. F. Ewer (11 December 2013).Ethology of Mammals. Springer.ISBN 978-1-4899-4656-0.
  4. ^Harris, R. L.,Cameron, E. Z., Davies, N. W., & Nicol, S. C. (2016).Chemical cues, hibernation and reproduction in female short-beaked echidnas (Tachyglossus aculeatus setosus): implications for sexual conflict. In Chemical Signals in Vertebrates 13 (pp. 145–166). Springer, Cham.
  5. ^Cassell's Latin Dictionary, Marchant, J.R.V, & Charles, Joseph F., (Eds.), Revised Edition, 1928, p.103
  6. ^Harper, Douglas."cloaca".Online Etymology Dictionary.
  7. ^cloaca. Charlton T. Lewis and Charles Short.A Latin Dictionary onPerseus Project.
  8. ^Michael L. Morrison; Amanda D. Rodewald; Gary Voelker; Melanie R. Colón; Jonathan F. Prather (3 September 2018).Ornithology: Foundation, Analysis, and Application. JHU Press.ISBN 978-1-4214-2471-2.
  9. ^Lynch, Wayne (2007)."The Cloacal Kiss".Owls of the United States and Canada. JHU Press. p. 151.ISBN 978-0-8018-8687-4.
  10. ^Julian Lombardi (1998).Comparative Vertebrate Reproduction. Springer.ISBN 978-0-7923-8336-9.Archived from the original on 26 March 2014. Retrieved5 December 2012.
  11. ^Hoffman, Ty C. M.; Walsberg, Glenn E.; DeNardo, Dale F. (2007)."Cloacal evaporation: an important and previously undescribed mechanism for avian thermoregulation".The Journal of Experimental Biology.210 (5):741–9.Bibcode:2007JExpB.210..741H.doi:10.1242/jeb.02705.PMID 17297135.
  12. ^Hager, Yfke (2007)."Cloacal Cooling".The Journal of Experimental Biology.210 (5): i.Bibcode:2007JExpB.210S...1H.doi:10.1242/jeb.02737.
  13. ^abcRomer, Alfred Sherwood; Parsons, Thomas S. (1977).The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 396–399.ISBN 978-0-03-910284-5.
  14. ^Libbie Henrietta Hyman,A laboratory manual for comparative vertebrate anatomy. 1922 (1920s)
  15. ^Mervyn Griffiths (2 December 2012).The Biology of the Monotremes. Elsevier Science.ISBN 978-0-323-15331-7.
  16. ^Gadow, Hans (20 August 2009)."On the Systematic Position of Notoryctes typhlops".Proceedings of the Zoological Society of London.60 (3):361–433.doi:10.1111/j.1469-7998.1892.tb06835.x.
  17. ^Riedelsheimer, B.; Unterberger, Pia; Künzle, H.; Welsch, U. (November 2007). "Histological study of the cloacal region and associated structures in the hedgehog tenrec Echinops telfairi".Mammalian Biology.72 (6):330–341.Bibcode:2007MamBi..72..330R.doi:10.1016/j.mambio.2006.10.012.
  18. ^Chimento, Nicolás; Agnolin, Federico (22 December 2014),Morphological evidence supports Dryolestoid affinities for the living Australian marsupial mole Notoryctes, PeerJ PrePrints,doi:10.7287/peerj.preprints.755
  19. ^Linzey, Donald W. (2020).Vertebrate Biology: Systematics, Taxonomy, Natural History, and Conservation. Johns Hopkins University Press. p. 306.ISBN 978-1-42143-733-0.
  20. ^Symonds, Matthew R. E. (February 2005). "Phylogeny and life histories of the 'Insectivora': controversies and consequences".Biological Reviews.80 (1):93–128.doi:10.1017/S1464793104006566.PMID 15727040.S2CID 21132866.
  21. ^Jonathan Kingdon; David Happold; Thomas Butynski; Michael Hoffman; Meredith Happold; Jan Kalina (2020).Mammals of Africa: Volumes I-VI. Bloomsbury Publishing. p. 216.ISBN 978-1-40818-996-2. Retrieved13 November 2024.
  22. ^Lukefahr, Steven D.; McNitt, James. I; Cheeke, Peter Robert; Patton, Nephi M. (2022).Rabbit Production. CABI. p. 3.ISBN 978-1-78924-978-1. Retrieved27 December 2024.
  23. ^Rosell, Frank; Campbell-Palmer, Róisín (2022).Beavers: Ecology, Behaviour, Conservation, and Management. OUP Oxford. p. 74.ISBN 978-0-19257-199-1. Retrieved27 December 2024.
  24. ^Stephen J. Divers; Douglas R. Mader (13 December 2005).Reptile Medicine and Surgery - E-Book. Elsevier Health Sciences.ISBN 978-1-4160-6477-0.
  25. ^C. Edward Stevens; Ian D. Hume (25 November 2004).Comparative Physiology of the Vertebrate Digestive System. Cambridge University Press. pp. 23–.ISBN 978-0-521-61714-7.
  26. ^Orenstein, Ronald (2001).Turtles, Tortoises & Terrapins: Survivors in Armor. Firefly Books.ISBN 978-1-55209-605-5.
  27. ^Dunson, William A. (1960). "Aquatic Respiration in Trionyx spinifer asper".Herpetologica.16 (4):277–83.JSTOR 3889486.
  28. ^The Straight Dope - Is it true turtles breathe through their butts?
  29. ^Aquarium Invertebrates by Rob Toonen, Ph.D.
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