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Nasal concha
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==Animals== [[File:Pferdeschädel.jpg|right|thumb|300px|The horse [[Respiratory system of the horse|breathes]] through [[Nostril|nare]]s (nostrils) which expand during exercise. The nasal passages have two turbinates on either side which increase the surface area to which air is exposed.<br />{{center|1 : Concha nasalis dorsalis }}{{center|2 : Concha nasalis media }}{{center|3 : Concha nasalis ventralis}}]] Generally, in animals, nasal conchae are convoluted structures of thin bone or cartilage located in the [[nasal cavity]]. These are lined with [[mucous membrane]]s that can perform two functions. They can improve the sense of smell by increasing the area available to absorb airborne chemicals, and they can warm and moisten inhaled air, and extract heat and moisture from exhaled air to prevent [[desiccation]] of the lungs. [[Olfaction|Olfactory]] turbinates are found in all living [[tetrapod]]s,{{citation needed|date=January 2015}} and [[respiratory]] turbinates are found in most mammals and birds. Animals with respiratory turbinates can breathe faster without drying out their lungs, and consequently can have a faster metabolism.<ref name=ZC94/> For example, when the [[emu]] exhales, its nasal turbinates condense moisture from the air and absorbs it for reuse.<ref>{{cite journal |author1=Maloney, S. K. |author2=Dawson, T. J. |year=1998 |title=Ventilatory accommodation of oxygen demand and respiratory water loss in a large bird, the Emu (Dromaius novaehollandiae), and a re-examination of ventilatory allometry for birds |journal=Physiological Zoology |volume=71 |issue=6 |pages=712–719 |pmid=9798259|doi=10.1086/515997 |s2cid=39880287 }}</ref> [[Dog]]s and other [[canid]]s possess well-developed nasal turbinates.<ref name=Wang88>Wang (2008) p. 88.</ref> These turbinates allow for heat exchange between small arteries and veins on their [[wiktionary:maxilloturbinal|maxilloturbinate]] (turbinates positioned on [[maxilla]] bone) surfaces in a counter-current heat-exchange system.<ref name=Wang88 /> Dogs are capable of prolonged chases, in contrast to the ambush predation of cats, and these complex turbinates play an important role in enabling this (cats only possess a much smaller and less-developed set of nasal turbinates).<ref name=Wang88 /> This same complex turbinate structure help conserve water in arid environments.<ref name=Wang87>Wang (2008) p. 87.</ref> The water conservation and thermoregulatory capabilities of these well-developed turbinates in dogs may have been crucial adaptations that allowed dogs (including both domestic dogs and their wild prehistoric [[gray wolf]] ancestors) to survive in the harsh [[Arctic]] environment and other cold areas of northern Eurasia and North America, which are both very dry and very cold.<ref name=Wang87 /> [[Reptile]]s and more primitive [[synapsid]]s have olfactory turbinates that are involved in sensing smell rather than preventing desiccation.<ref name=HWJ94>{{cite journal|last=Hillenius|first=W.J.|year=1994|title=Turbinates in therapsids: Evidence for Late Permian origins of mammalian endothermy|journal=Evolution|volume=48|issue=2|pages=207–229|doi=10.2307/2410089|pmid=28568303|jstor=2410089}}</ref> While the maxilloturbinates of mammals are located in the path of airflow to collect moisture, sensory turbinates in both mammals and reptiles are positioned farther back and above the nasal passage, away from the flow of air.<ref name=Ruben2000>{{cite journal|last=Ruben|first=J.A.|author2=Jones, T.D.|year=2000|title=Selective factors associated with the origin of fur and feathers|journal=American Zoologist|volume=40|issue=4|pages=585–596|doi=10.1093/icb/40.4.585|doi-access=free}}</ref> ''[[Glanosuchus]]'' has ridges positioned low in the nasal cavity, indicating that it had maxilloturbinates that were in the direct path of airflow. The maxilloturbinates may not have been preserved because they were either very thin or [[cartilaginous]]. The possibility has also been raised that these ridges are associated with an olfactory [[epithelium]] rather than turbinates.<ref name=KTS06>{{cite journal|last=Kemp|first=T.S.|year=2006|title=The origin of mammalian endothermy: a paradigm for the evolution of complex biological structure|journal=Zoological Journal of the Linnean Society|volume=147|issue=4|pages=473–488|doi=10.1111/j.1096-3642.2006.00226.x|doi-access=free}}</ref> Nonetheless, the possible presence of maxilloturbinates suggests that ''Glanosuchus'' may have been able to rapidly breathe without drying out the nasal passage, and therefore could have been an endotherm.<ref name=ZC94>{{cite journal|last=Zimmer|first=C.|year=1994|title=The Importance of Noses|journal=Discover|volume=15|issue=8|url=http://discovermagazine.com/1994/aug/theimportanceofn416}}</ref><ref name=HWJ94/><ref name=KTS06/> The bones of nasal turbinates are very fragile and seldom survive as fossils. In particular none have been found in fossil birds.<ref name=Witmer2001>{{cite journal |author=Witmer, L.M. |title=Nostril Position in Dinosaurs and Other Vertebrates and Its Significance for Nasal Function |journal=Science |date=August 2001|issue=5531 |pages=850–853 |doi=10.1126/science.1062681 |volume=293 |pmid=11486085 |citeseerx=10.1.1.629.1744 |s2cid=7328047 }}</ref> But there is indirect evidence for their presence in some fossils. Rudimentary ridges like those that support respiratory turbinates have been found in advanced Triassic [[cynodont]]s, such as ''[[Thrinaxodon]]'' and ''[[Eucynodontia|Diademodon]]''. This suggests that they may have had fairly high metabolic rates.<ref name=Brink>{{Cite journal| last=Brink | first=A.S. | title=A study on the skeleton of ''Diademodon'' | journal=Palaeontologia Africana | volume=3 | pages=3–39 |year=1955 }}</ref><ref name=Kemp>{{Cite book| last=Kemp | first=T.S. | title=Mammal-like reptiles and the origin of mammals | publisher=Academic Press | year=1982 | location=London | pages=363| isbn=978-0-12-404120-2}}</ref><ref>{{Cite journal| last=Hillenius | first=W.H. | title=The evolution of nasal turbinates and mammalian endothermy | journal=Paleobiology | volume=18 | issue=1 | pages=17–29 |year=1992 | jstor = 2400978 | doi=10.1017/S0094837300012197 | bibcode=1992Pbio...18...17H | s2cid=89393753 }}</ref><ref>{{Cite journal| last=Ruben | first=J. | title=The evolution of endothermy in mammals and birds: from physiology to fossils | journal=Annual Review of Physiology | volume=57 | pages=69–95 |year=1995 | doi=10.1146/annurev.ph.57.030195.000441 | pmid=7778882 }}</ref> The paleontologist [[John Ruben]] and others have argued that no evidence of nasal turbinates has been found in dinosaurs. All the dinosaurs they examined had nasal passages that they claimed were too narrow and too short to accommodate nasal turbinates, so dinosaurs could not have sustained the breathing rate required for a mammal-like or bird-like metabolic rate while at rest, because their lungs would have dried out.<ref name=Ruben2000/><ref name=Ruben1997>{{cite journal |author=Ruben, J.A., Jones, T.D., Geist, N.R. and Hillenius, W. J. |title=Lung structure and ventilation in theropod dinosaurs and early birds |journal=Science |date=November 1997 |volume=278 |issue=5341 |pages=1267–1270 |doi=10.1126/science.278.5341.1267 |bibcode = 1997Sci...278.1267R }}</ref><ref name=Ruben1996>{{cite journal |author=Ruben, J.A., Hillenius, W.J., Geist, N.R., Leitch, A., Jones, T.D., Currie, P.J., Horner, J.R., and Espe, G. |title=The metabolic status of some Late Cretaceous dinosaurs |journal=Science |date=August 1996 |volume=273 |issue=5279 |pages=1204–1207 |doi=10.1126/science.273.5279.1204 |bibcode = 1996Sci...273.1204R |s2cid=84693210 |url=http://doc.rero.ch/record/14694/files/PAL_E1414.pdf }}</ref> However, objections have been raised against this argument. Nasal turbinates are absent or very small in some birds, such as [[ratite]]s, [[Procellariiformes]] and [[Falconiformes]]. They are also absent or very small in some mammals, such as anteaters, bats, elephants, whales and most primates, although these animals are fully endothermic and in some cases very active.<ref name="Bang1966">{{cite journal |author=Bang, B.G. | year=1966 | title=The olfactory apparatus of Procellariiformes |journal=Acta Anatomica | volume=65 | pages=391–415 |doi=10.1159/000142884 |pmid=5965973 |issue=1 }}</ref><ref name="Bang1977">{{cite journal |author=Bang, B.G. | year=1971 | title= Functional anatomy of the olfactory system in 23 orders of birds |series=79 |journal=Acta Anatomica | volume= 79| pages=1–76 | pmid=5133493 | doi=10.1159/isbn.978-3-318-01866-0| isbn=978-3-8055-1193-3 }}</ref><ref name="Scott1954">{{cite journal |author=Scott, J.H. | year=1954 | title=Heat regulating function of the nasal mucous membrane |journal=Journal of Larynology and Otology | volume=68 |issue=5 |pages=308–317 | pmid = 13163588 | doi = 10.1017/S0022215100049707 | s2cid=32082759 }}</ref><ref name="Coulombeetal1965">{{cite journal|author=Coulombe, H.N., Sam H. Ridgway, S.H., and Evans, W.E. | year=1965 | title=Respiratory water exchange in two species of porpoise | journal=Science | volume=149 | issue=3679 | pages=86–88 | doi=10.1126/science.149.3679.86|pmid=17737801|bibcode = 1965Sci...149...86C | s2cid=38947951 }}</ref> Furthermore, ossified turbinate bones have been identified in the [[Ankylosauridae|ankylosaurid]] dinosaur ''[[Saichania]]''.<ref name="Maryańska1977">{{cite journal |last=Maryańska |first=T. |author-link =Teresa Maryańska| year=1977 |title=Ankylosauridae (Dinosauria) from Mongolia |journal=Palaeontologia Polonica |volume=37 |pages=85–151 }}</ref>
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