Editing Desmostylia

{{Short description|Extinct order of mammals}}
{{Automatic taxobox
| fossil_range = [[Oligocene]]-[[Miocene]], {{Fossil range|30.8|7.2}}
| image = Desmostylus ROM.jpg
| image_caption = ''[[Desmostylus]]'', [[Royal Ontario Museum]]
| image2 = Paleoparadoxia BW.jpg
| image2_caption = Restoration of ''[[Paleoparadoxia]]''
| taxon = Desmostylia
| authority = {{Harvnb|Reinhart|1959}}
| subdivision_ranks = Families and genera
| subdivision = * Family †[[Desmostylidae]]<br /><small>{{Harvnb|Osborn|1905}}</small>
** †''[[Behemotops]]''<br /><small>{{Harvnb|Domning|Ray|McKenna|1986}}</small>
** †''[[Desmostylus]]''<br /><small>{{Harvnb|Marsh|1888}}</small>
** †''[[Cornwallius]]''<br /><small>{{Harvnb|Hay|1923}}</small>
** †''[[Kronokotherium]]''<br /><small>{{Harvnb|Pronina|1957}}</small>
** †''[[Ounalashkastylus]]''<br /><small>Chiba, Kentaro 2016</small>
* Family †[[Paleoparadoxiidae]]<br /><small>{{Harvnb|Reinhart|1959}}</small>
** †''[[Ashoroa]]''<br /><small>{{Harvnb|Inuzuka|2000}}</small>
** †''[[Paleoparadoxia]]''<br /><small>{{Harvnb|Reinhart|1959}}</small>
** †''[[Neoparadoxia]]''<br /><small>{{Harvnb|Barnes|2013}}</small>
}}

The '''Desmostylia''' (from [[Ancient Greek|Greek]] δεσμά ''desma'', "bundle", and στῦλος ''stylos'', "pillar")<ref name="Gingerich-2005-intro">{{Harvnb|Gingerich|2005| loc=Introduction}}</ref> are an [[extinct]] [[order (biology)|order]] of [[aquatic mammal]]s native to the North Pacific from the early [[Oligocene]] ([[Rupelian]]) to the late [[Miocene]] ([[Tortonian]]) ({{Mya|30.8|7.25}}). Desmostylians are the only known extinct order of marine mammals.<ref name="Gheerbrant-etal-2005">{{Harvnb|Gheerbrant|Domning|Tassy|2005|pp=95–6}}</ref>

The Desmostylia, together with [[Sirenia]] and [[Proboscidea]] (and possibly [[Embrithopoda]]), have traditionally been assigned to the [[afrotheria]]n [[clade]] [[Tethytheria]], a group named after the paleoocean [[Tethys Ocean|Tethys]] around which they originally evolved. The relationship between the Desmostylia and the other orders within the Tethytheria has been disputed; if the common ancestor of all tethytheres was semiaquatic, the Proboscidea became secondarily terrestrial; alternatively, the Desmostylia and Sirenia could have evolved independently into aquatic mammals.<ref>{{Harvnb|Uhen|2007|p=515}}</ref> The assignment of Desmostylia to Afrotheria has always been problematic from a biogeographic standpoint, given that Africa was the locus of the early evolution of the Afrotheria while the Desmostylia have only been found along the [[Pacific Rim]]. That assignment has been seriously undermined by a [[:Image:Cooper et al. parsimony analyses consensus tree for anthracobunid phylogeny.png|2014 cladistic analysis]] that places [[anthracobunid]]s and desmostylians, two major groups of putative non-African afrotheres, close to each other within the [[laurasiatheria]]n order [[Perissodactyla]].<ref name="Cooper2014">{{Harvnb|Cooper|Seiffert|Clementz|Madar|2014}}</ref> However, a subsequent study shows that, while anthracobunids are definite perissodactyls, desmostylians share the same number of characteristics necessary for either [[Paenungulata]] or Perissodactyla, making their former assessment as afrotheres a possibility.<ref>{{cite journal | last1 = Gheerbrant | first1 = Emmanuel | last2 = Filippo | first2 = Andrea | last3 = Schmitt | first3 = Arnaud | year = 2016 | title = Convergence of Afrotherian and Laurasiatherian Ungulate-Like Mammals: First Morphological Evidence from the Paleocene of Morocco | journal = PLOS ONE | volume = 11 | issue = 7| page = e0157556 | doi = 10.1371/journal.pone.0157556 | pmid=27384169 | pmc=4934866| bibcode = 2016PLoSO..1157556G | doi-access = free }}</ref>

==Description==
[[File:Desmostylia skeletal diagram (to scale).png|center|thumb|655x655px|Skeletal diagrams: (A) ''[[Ashoroa]]'', (B) ''[[Desmostylus]]'', (C) ''[[Behemotops|Behemetops]]'', (D) ''[[Paleoparadoxia]]'']]
Desmostylians were large, fully aquatic quadrupeds with massive limbs and short tails.<ref name="Gheerbrant-etal-2005" /> The smallest is ''[[Ashoroa laticosta]]'', a relatively large animal at a body length of {{convert|168|cm|ftin|0|disp=or}}, while the largest species reached sizes comparable to [[Steller's sea cow]].<ref name="Nicholas D. Pyenson 2016">[[Nicholas D. Pyenson]], Geerat J. Vermeij, The rise of ocean giants: maximum body size in Cenozoic marine mammals as an indicator for productivity in the Pacific and Atlantic Oceans, Published 5 July 2016.DOI: 10.1098/rsbl.2016.0186</ref>

A desmostylian skull has an elongated and broadened [[Rostrum (anatomy)|rostrum]], with the nasal opening located slightly dorsally. The [[zygomatic arch]]es are prominent (behind the eyes), the [[paroccipital process]]es elongated (downward-pointing processes behind the jaw-joints), and the [[epitympanic sinus]]es open into the [[temporal fossa]]e (cavities above the ear holes).<ref name="Gheerbrant-etal-2005" />

The mandible and maxilla typically have forward-pointing incisors and canine tusks, followed by a long postcanine diastema, partly because of the reduced number of premolars. The cusps of the premolars and molars are composed of densely packed cylinders of thick enamel, giving the order its name ("bundle of columns"). The primitive dental formula is 3.1.4.3, with a trilobate fourth deciduous premolar. The cheek teeth are [[brachydont]] and [[bunodont]] in primitive genera, but [[hypsodont]] in later genera such as ''Desmostylus'', which has many supernumerary cusps.<ref name="Gheerbrant-etal-2005" />

In the [[postcrania]], the [[clavicle]] is absent and the [[sternum]] consists of a series of heavy, paired, plate-like sternebrae. In adults, the joints between the radius and ulna prevent any movements. The metacarpals are longer than metatarsals, and each foot has four digits (digit I is vestigial).<ref name="Gheerbrant-etal-2005" />

==Behaviour==
Their dental and skeletal forms suggest that desmostylians were aquatic [[herbivore]]s dependent on [[littoral]] habitats. Their name refers to their highly distinctive molars, in which each cusp was modified into hollow columns, so a typical molar would have resembled a cluster of pipes, or in the case of worn molars, volcanoes. (This may reflect the close relationship between the [[Paenungulata]], to which this group has been assigned, and the [[Tubulidentata]].)

''Desmostylus'' did not chew or eat like any other known animal.  It clenched its teeth, rooted up plants with the help of tusks and powerful neck, and then sucked them in using strong throat muscles and the shape of the roof of the mouth.<ref>{{cite web
  | url       = https://www.sciencedaily.com/releases/2015/10/151006102733.htm
  | title     = New fossils intensify mystery of short-lived, toothy mammal found in ancient North Pacific, Oddball creature, Desmostylia
  | date      = October 6, 2015
  | website   = www.sciencedaily.com
  | publisher = Southern Methodist University |access-date=October 11, 2015
}}</ref>

Desmostylians are believed to be aquatic because of a combination of characteristics. Their legs seemed to be adapted for terrestrial locomotion, while a number of other parameters confirms their aquatic nature:<ref name="Gingerich-2005-intro" />
* Fossils have been found in marine strata.
* The [[nostril|nares]] are retracted and the [[Orbit (anatomy)|orbits]] are raised like in other aquatic mammals.
* Levels of stable [[isotope]]s in their tooth enamel suggest an aquatic diet and environment (carbon and oxygen) and fresh or brackish water (strontium).
* Their spongy bone structure is similar to that of [[cetaceans]].

Based on a comparison of trunk and limb proportions, {{Harvnb|Gingerich|2005}} concluded<ref>{{Harvnb|Gingerich|2005| loc=Discussion}}</ref> that desmostylians were more terrestrial than aquatic and clearly fore limb-dominated swimmers, hence they were more similar to "sea bears" than "[[Thalassocnus|sea sloths]]" (as proposed by other researchers.) However, a more recent and detailed analysis of desmostylian bone structure has revealed them to be fully aquatic, like [[sirenians]] and [[cetaceans]],<ref name="Hayashi2013">{{Harvnb|Hayashi|Houssaye|Nakajima|Kentaro|2013}}</ref> with their limbs being incapable of supporting their own weight on land. More recent studies vindicate this assessment, as desmostylians had a thoracic morphology more similar to sirenians and modern cetaceans than to that of semiaquatic mammals.<ref>{{cite journal | doi = 10.1111/joa.12518 | volume=229 | title=Farewell to life on land - thoracic strength as a new indicator to determine paleoecology in secondary aquatic mammals | year=2016 | journal=Journal of Anatomy | pages=768–777 | last1 = Ando | first1 = Konami | issue=6 | pmid=27396988 | pmc=5108153}}</ref> Its less dense bone structure suggests that ''Desmostylus'' had a lifestyle of active swimming and possibly feeding at the surface, while other desmostylians were primarily slow swimmers and/or bottom walkers and sea grass feeders.<ref name="Hayashi2013" />

==Habitat==
A 2017 study on ''[[Desmostylus]]'' and ''[[Paleoparadoxia]]'' shows that the former preferred areas shallower than 30 m, while the latter occurred in deep, offshore waters.<ref>{{cite journal | last1 = Matsui | first1 = Kumiko | last2 = Sashida | first2 = Katsuo | last3 = Agematsu | first3 = Sachiko | last4 = Kohno | first4 = Naoki | year = 2017 | title = Habitat preferences of the enigmatic Miocene tethythere Desmostylus and Paleoparadoxia (Desmostylia; Mammalia) inferred from the depositional depth of fossil occurrences in the Northwestern Pacific realm | journal = Palaeogeography, Palaeoclimatology, Palaeoecology | volume = 471 | pages = 254–265 | doi = 10.1016/j.palaeo.2017.02.005 | bibcode = 2017PPP...471..254M }}</ref>

==Distribution==
Desmostylian [[fossil]]s are known from the northern [[Pacific Rim]],<ref>{{Harvnb|Gingerich|2005| loc=Abstract}}</ref> from southern [[Japan]] through [[Russia]], the [[Aleutian Islands]], and the Pacific coast of [[North America]] to the southern tip of [[Baja California]]. They range from the Early [[Oligocene]] to the late [[Miocene]].

==Extinction==
Desmostylians, being fully marine herbivores, are thought to have been outcompeted ecologically by [[dugongid]] sirenians. In particular, later species like ''[[Neoparadoxia]]'' are more specialised than previous forms, suggesting increased divergence to compete with sirenians, and sirenian diversity appears to increase with desmostylian decline.<ref>Barnes, L.. (2013, September 11) .A new genus and species of late Miocene Paleoparadoxiid (Mammalia, Desmostylia) from California. Natural History Museum of Los Angeles County Contributions in Science, (521), 51–114</ref> Both desmostylians and [[North Pacific]] dugongids were apparently [[kelp]] specialists, as opposed to marine herbivorous mammals from other regions, with diets primarily composed of [[seagrass]].<ref name="Nicholas D. Pyenson 2016"/>

==Classification==
[[File:Desmostylia.jpg|thumb|Restoration of ''Desmostylus'' and ''Paleoparadoxia'']]
[[File:Paleoparadoxia Natural History Museum of Los Angeles County 20110330.jpg|thumb|''Neoparadoxia cecilialina'' on display at the Natural History Museum of Los Angeles County]]
[[File:Paleoparadoxia Skeleton.jpg|thumb|''Paleoparadoxia'' skeleton]]
[[File:Desmostylus Skull.jpg|thumb|Skull of ''Desmostylus'']]
The [[type species]] ''Desmostylus hesperus'' was originally classified from a few teeth and [[vertebrae]] as a sirenian by {{Harvnb|Marsh|1888}}, but doubts arose a decade later when more complete fossils were discovered in Japan. {{Harvnb|Osborn|1905}} also proposed that they belonged to the Sirenia.<ref>{{Paleodb|reference|25349|H. F. Osborn 1905}}. Retrieved 17 March 2013.</ref> One of the most comprehensive collections of desmostylian teeth was amassed by paleontologist [[John C. Merriam]], who concluded on the basis of the [[molar (tooth)|molar]] structure and repeated occurrence in marine beds that the animals had been aquatic, and were probably sirenian.

In 1926, the Austrian palaeontologist [[Othenio Abel]] suggested origins with [[monotreme]]s, like the [[platypus|duck-billed platypus]], and in 1933, he even created the order "Desmostyloidea", which he placed within the [[Multituberculata]]. Abel died shortly after World War II, and his classification won few supporters and has been ignored since.<ref>{{Harvnb|Domning|Ray|McKenna|1986|p=34}}</ref>

Because desmostylians were originally known only from skull fragments, teeth, and bits of other bones, general agreement was that they had had flippers and a fin-like tail. The discovery of a complete skeleton from [[Sakhalin Island]] in 1941, however, showed that they possessed four legs, with bones as stout as a [[hippopotamus]]', and justified the creation of a new order for the desmostylians, described by {{Harvnb|Reinhart|1959}}.

A major find was announced in October 2015 after scientists examined an extensive group of giant, tusked, quadruped, marine mammal fossils. This northernmost to date species discovery had been unearthed during excavation for the construction of a school in [[Unalaska]], in the [[Aleutian Islands]].<ref>[http://www.adn.com/article/20151007/giant-extinct-mammal-identified-unalaska-fossils Giant extinct mammal identified from Unalaska fossils], [[Alaska Dispatch News]], Mike Dunham, October 7, 2015. Retrieved 8 October 2015.</ref> A rendition of a group was drawn by Alaskan artist [[Ray Troll]].<ref>[http://www.adn.com/slideshow/photos-new-species-extinct-marine-mammal-identified-unalaska-fossils Photos: New species of extinct marine mammal identified from Unalaska fossils], [[Alaska Dispatch News]], Mike Dunham, October 7, 2015. Retrieved 8 October 2015.</ref>

Despite their similarities to manatees and elephants, desmostylians were entirely unlike any living creatures. [[Douglas Emlong]]'s 1971 discovery of the new genus ''[[Behemotops]]'' from [[Oregon]] showed that early desmostylians had more proboscidean-like teeth and jaws than later ones. Despite this discovery, their relationships to manatees and proboscids remain unresolved. The analysis of Cooper ''et al.'' (2014) indicates the similarities with manatees and elephants may be a result of [[convergent evolution|convergence]] and that they may instead be basal [[perissodactyls]].<ref name="Cooper2014"/>

{{Harvnb|Barnes|2013}} proposed a new classification of Paleoparadoxiidae:<ref>{{Harvnb|Barnes|2013|pp=103, 108–109}}</ref>
* Order '''Desmostylia''' Reinhart, 1953
** Family Paleoparadoxiidae Reinhart, 1959
*** Subfamily Behemotopsinae (Inuzuka, 1987)
**** ''[[Behemotops]]'' Domning, Ray, and McKenna, 1986
***** ''Behemotops proteus'' Domning, Ray, and McKenna, 1986 (including ''Behemotops emlongi'' Domning, Ray, and McKenna, 1986)
***** ''Behemotops katsuiei'' Inuzuka, 2000b
*** Subfamily Paleoparadoxiinae (Reinhart, 1959)
**** ''[[Archaeoparadoxia]]''
***** ''Archaeoparadoxia weltoni'' (Clark, 1991)
**** ''[[Paleoparadoxia]]'' Reinhart, 1959
***** ''Paleoparadoxia tabatai'' (Tokunaga, 1939), (= ''Paleoparadoxia media'' Inuzuka, 2005)
**** ''[[Neoparadoxia]]'' Barnes 2013
***** ''Neoparadoxia repenningi'' (Domning and Barnes, 2007)
***** ''Neoparadoxia cecilialina'' Barnes 2013

==References==
{{Reflist}}

==Further reading==
{{Refbegin|30em}}
* {{Cite journal
 |last         = Barnes
 |first        = Lawrence G.
 |title        = A New Genus of Late Miocene Paleoparadoxiid (Mammalia, Desmostylia) from California
 |year         = 2013
 |journal      = Contributions in Science
 |volume       = 521
 |pages        = 51–114
 |url          = http://www.nhm.org/site/sites/default/files/pdf/contrib_science/CS521.51-114.pdf
 |access-date  = 14 January 2014
 |archive-date = 7 November 2015
 |archive-url  = https://web.archive.org/web/20151107060556/http://www.nhm.org/site/sites/default/files/pdf/contrib_science/CS521.51-114.pdf
 |url-status   = dead
}}<!-- {{Harvnb|Barnes|2013}} -->
* {{Cite journal
  | doi     = 10.1371/journal.pone.0109232
  | pmid    = 25295875
  | title   = Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls
  | journal = PLOS ONE
  | volume  = 9
  | issue   = 10
  | pages   = e109232
  | date    = 2014-10-08
  | last1   = Cooper
  | first1  = L. N.
  | last2   = Seiffert
  | first2  = E. R.
  | last3   = Clementz
  | first3  = M.
  | last4   = Madar
  | first4  = S. I.
  | last5   = Bajpai
  | first5  = S.
  | last6   = Hussain
  | first6  = S. T.
  | last7   = Thewissen
  | first7  = J. G. M.
  | pmc=4189980
| bibcode = 2014PLoSO...9j9232C
 | doi-access = free
 }}<!-- {{Harvnb|Cooper|Seiffert|Clementz|Madar|2014}} -->
* {{Cite journal
  | last1   = Domning
  | first1  = Daryl P.
  | last2   = Barnes
  | first2  = Lawrence G.
  | title   = A new name for the 'Stanford skeleton' of ''Paleoparadoxia'' (Mammlia, Desmostylia)
  | year    = 2007
  | journal = Journal of Vertebrate Paleontology
  | volume  = 27
  | issue   = 3
  | pages   = 748–751
  | doi     = 10.1671/0272-4634(2007)27[748:annfts]2.0.co;2
}}<!-- {{Harvnb|Domning|Barnes|2007}} -->
* {{Cite journal
  | last1      = Domning
  | first1     = Daryl P.
  | last2      = Ray
  | first2     = Clayton Edward
  | last3      = McKenna
  | first3     = Malcolm C.
  | title      = Two new Oligocene desmostylians and a discussion of Tethytherian systematics
  | year       = 1986
  | journal    = Smithsonian Contributions to Paleobiology
  | volume     = 59
  | issue     = 59
  | pages      = 1–56
  | url        = http://www.sil.si.edu/SmithsonianContributions/Paleobiology/pdf_lo/SCtP-0059.pdf
  | access-date = 15 December 2013
  | oclc       = 12974759
  | doi        = 10.5479/si.00810266.59.1
}}<!-- {{Harvnb|Domning|Ray|McKenna|1986}} -->
* {{Cite journal
  | last       = Gingerich
  | first      = Philip D.
  | author-link = Philip D. Gingerich
  | title      = Aquatic Adaptation and Swimming Mode Inferred from Skeletal Proportions in the Miocene Desmostylian ''Desmostylus''
  | year       = 2005
  | journal    = Journal of Mammalian Evolution
  | volume     = 12
  | issue      = 1/2
  | url        = http://deepblue.lib.umich.edu/bitstream/handle/2027.42/44971/10914_2005_Article_5719.pdf
  | access-date = 17 March 2013
  | doi        = 10.1007/s10914-005-5719-1
  | pages      = 183–194
| hdl= 2027.42/44971
  | s2cid = 7812089
 | hdl-access= free
  }}<!-- {{Harvnb|Gingerich|2005}} -->
* {{Cite journal
  | last       = Hay
  | first      = O. P.
  | author-link = Oliver Perry Hay
  | title      = Characteristics of sundry fossil vertebrates
  | year       = 1923
  | journal    = Pan-American Geologist
  | volume     = 39
  | pages      = 101–20
  | oclc       = 38855496
}}<!-- {{Harvnb|Hay|1923}} -->
* {{Cite journal
  | last1   = Hayashi
  | first1  = S.
  | last2   = Houssaye
  | first2  = A.
  | last3   = Nakajima
  | first3  = Y.
  | last4   = Kentaro
  | first4  = C.
  | last5   = Ando
  | first5  = T.
  | last6   = Sawamura
  | first6  = H.
  | last7   = Inuzuka
  | first7  = N.
  | last8   = Kaneko
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  | last9   = Osaki
  | first9  = T.
  | title   = Bone Inner Structure Suggests Increasing Aquatic Adaptations in Desmostylia (Mammalia, Afrotheria)
  | year    = 2013
  | journal = PLOS ONE
  | volume  = 8
  | issue   = 4
  | page    = e59146
  | doi     = 10.1371/journal.pone.0059146
  | oclc    = 837402105
  | pmid    = 23565143
  | pmc     = 3615000
| bibcode = 2013PLoSO...859146H
 | doi-access = free
 }}<!-- {{Harvnb|Hayashi|Houssaye|Nakajima|Kentaro|2013}} -->
* {{Cite journal
  | last    = Inuzuka
  | first   = N.
  | title   = Primitive late Oligocene desmostylians from Japan and Phylogeny of the Desmostylia
  | year    = 2000
  | journal = Bulletin of the Ashoro Museum of Paleontology
  | volume  = 1
  | pages   = 91–123
}}<!-- {{Harvnb|Inuzuka|2000}} -->
**{{cite web |title=N. Inuzuka 2000 |website=Fossilworks |url=https://paleobiodb.org/classic/displayRefResults?reference_no=16701 }}
* {{Cite journal
  | last       = Marsh
  | first      = O. C.
  | author-link = Othniel Charles Marsh
  | title      = Notice of a new fossil sirenian, from California
  | year       = 1888
  | journal    = American Journal of Science
  | volume     = 25
  | issue      = 8
  | pages      = 94–96
  | doi = 10.2475/ajs.s3-35.205.94
 | bibcode = 1888AmJS...35...94M
 | s2cid = 129254943
 | url        = http://marsh.dinodb.com/marsh/Marsh%201888%20-%20Notice%20of%20a%20new%20fossil%20sirenian,%20from%20California.pdf
  | access-date = 15 December 2013
  | oclc       = 79838746
}}<!-- {{Harvnb|Marsh|1888}} -->
* {{Cite journal
  | last       = Osborn
  | first      = H. F.
  | author-link = Henry Fairfield Osborn
  | title      = Ten years progress in the mammalian Palaeontology of North America
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}}<!-- {{Harvnb|Osborn|1905}} -->
* {{Cite journal
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  | title   = A new desmostylid, ''Kronokotherium brevimaxillare'' gen. nov., sp. nov., from the Miocene deposits of Kamchatka
  | year    = 1957
  | journal = [[Proceedings of the USSR Academy of Sciences|C. R. Acad. Sci. URSS]]
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}}<!-- {{Harvnb|Pronina|1957}} -->
* {{Cite journal
  | last       = Reinhart
  | first      = Roy Herbert
  | author-link = Roy Herbert Reinhart
  | title      = A review of the Sirenia and Desmostylia
  | year       = 1959
  | journal    = University of California Publications in Geological Sciences
  | volume     = 36
  | issue      = 1
  | pages      = 1–146
  | oclc       = 3474601
}}<!-- {{Harvnb|Reinhart|1959}} -->
* {{Cite book
  | last1      = Gheerbrant
  | first1     = Emmanuel
  | last2      = Domning
  | first2     = Daryl P.
  | last3      = Tassy
  | first3     = Pascal
  | chapter    = Paenungulata
  | title      = The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades | editor1-last = Rose | editor1-first = Kenneth D. | editor2-last = Archibald | editor2-first = J. David
  | year       = 2005
  | publisher  = JHU Press
  | chapter-url        = https://books.google.com/books?id=DhchVG_rbQ8C&pg=PA95
  | access-date = 15 June 2013
  | isbn       = 9780801880223
  | oclc       = 55801049
}}<!-- {{Harvnb|Gheerbrant|Domning|Tassy|2005}} -->
* {{Cite journal
  | last    = Uhen
  | first   = Mark D.
  | title   = Evolution of Marine Mammals: Back to the Sea After 300 Million Years
  | year    = 2007
  | journal = Anatomical Record
  | volume  = 290
  | issue   = 6
  | pages   = 514–22
  | oclc    = 137264359
  | doi     = 10.1002/ar.20545
  | pmid    = 17516441
| doi-access= free
  }}<!-- {{Harvnb|Uhen|2007}} -->
{{Refend}}

==External links==
{{Wikispecies}}
{{Wiktionary}}
{{Commons category}}
* {{Cite web
  | title      = Introduction to Desmostylia
  | publisher  = University of California Museum of Paleontology
  | url        = http://www.ucmp.berkeley.edu/mammal/mesaxonia/desmostylia.html
  | date       = January 2009
  | access-date = 16 June 2013
}}

{{Paenungulata}}
{{Taxonbar|from=Q1139385}}

[[Category:Desmostylians| ]]
[[Category:Tortonian extinctions]]
[[Category:Prehistoric placental mammals]]
[[Category:Rupelian first appearances]]
[[Category:Taxa described in 1959]]
[[Category:Taxa named by Roy Herbert Reinhart]]
[[Category:Mammal orders]]
[[Category:Prehistoric tetrapod orders]]