View source for Paenungulata

{{Short description|Clade of mammals including elephants, hyraxes, and sea cows}}
{{automatic taxobox
| fossil_range = [[Paleocene]] - Recent, {{fossilrange|61|0}}
| image = Paenungulata crown.png
| image_caption = Top: [[rock hyrax]] ([[Hyracoidea]]), ''[[Arsinoitherium|Arsinoitherium zitteli]]'' (†[[Embrithopoda]]); bottom: [[Asian elephant]] ([[Proboscidea]]), [[West Indian manatee]] ([[Sirenia]]).
| taxon = Paenungulata
| authority = [[George Gaylord Simpson|Simpson]], 1945
| display_parents = 2
| subdivision_ranks = Subgroups
| subdivision = * [[Hyracoidea]]
* †[[South American native ungulates|Sudamericungulata]]?<ref>{{Cite journal|last1=Avilla|first1=Leonardo S.|last2=Mothé|first2=Dimila|date=2021|title=Out of Africa: A New Afrotheria Lineage Rises From Extinct South American Mammals|journal=Frontiers in Ecology and Evolution|language=English|volume=9|doi=10.3389/fevo.2021.654302|issn=2296-701X|doi-access=free}}</ref>
* [[Tethytheria]]
| range_map = Paenungulata map cropped.jpg
}}
'''Paenungulata''' (from [[Latin]] {{lang|la|paene}} 'almost' and {{lang|la|ungulātus}} 'having [[hoof]]s'), also known as '''Uranotheria''', is a [[clade]] of "sub-[[Ungulate|ungulates]]", which groups three [[Extant taxon|extant]] [[mammal]] [[Order (biology)|orders]]: [[Proboscidea]] (including [[elephant]]s), [[Sirenia]] ([[sirenia|sea cow]]s, including [[dugong]]s and [[manatee]]s), and [[Hyracoidea]] ([[hyrax]]es). At least two more possible orders are known only as fossils, namely [[Embrithopoda]] and [[Desmostylia]].<ref group=lower-alpha name=possible_exclusions/>

Molecular evidence indicates that Paenungulata (or at least its extant members) is part of the cohort [[Afrotheria]], an ancient assemblage of mainly [[Africa]]n mammals of great diversity. The other members of this cohort are the orders [[Afrosoricida]] ([[tenrec]]s and [[golden mole]]s), Macroscelidea ([[elephant shrew]]s) and Tubulidentata ([[aardvark]]s).<ref name="comparison"/>

Of the five orders, hyraxes are the most basal, followed by embrithopods; the remaining orders (sirenians and elephants) are more closely related. These latter three are grouped as the '''[[Tethytheria]]''', because it is believed that their common ancestors lived on the shores of the prehistoric [[Tethys Sea]]; however, recent [[myoglobin]] studies indicate that even [[Hyracoidea]] had an aquatic ancestor.<ref>{{cite news |title=One Protein Shows Elephants and Moles Had Aquatic Ancestors |date=2013-06-13 |df=dmy-all |publisher=[[National Geographic Society]] |url=http://phenomena.nationalgeographic.com/2013/06/13/one-protein-shows-elephants-and-moles-had-aquatic-ancestors/|archive-url=https://web.archive.org/web/20130617051932/http://phenomena.nationalgeographic.com/2013/06/13/one-protein-shows-elephants-and-moles-had-aquatic-ancestors/|archive-date=June 17, 2013}}</ref>

== History ==
In 1945, [[George Gaylord Simpson]] used traditional [[Taxonomy (biology)|taxonomic techniques]] to group these spectacularly diverse mammals in the superorder he named Paenungulata ("almost [[ungulate]]s"), but there were many loose threads in unravelling their genealogy.<ref name="Seiffert">{{cite journal |last1=Seiffert |first1=Erik |last2=Guillon |first2=J.M. |year=2007 |title=A new estimate of Afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence |journal=BMC Evolutionary Biology |volume=7 |page=13 |doi=10.1186/1471-2148-7-224 |pmid=17999766 |pmc=2248600 |url= |doi-access=free |bibcode=2007BMCEE...7..224S }}</ref> For example, hyraxes in his Paenungulata had some characteristics suggesting they might be connected to the [[Perissodactyla]] (odd-toed ungulates, such as horses and rhinos). Indeed, early taxonomists placed the Hyracoidea closest to the rhinoceroses because of their dentition.

When genetic techniques were developed for inspecting [[amino acid]] differences among [[haemoglobin]] sequences the most [[Maximum parsimony (phylogenetics)|parsimonious]] [[cladogram]]s depicted Simpson's Paenungulata as an authentic [[clade]] and as one of the first groups to diversify from the basal placental mammals ([[Eutheria]]). The amino acid sequences reject a connection between extant paenungulates and [[perissodactyls]] (odd-toed ungulates).<ref name="Seiffert"/>

However, a [[:Image:Cooper et al. parsimony analyses consensus tree for anthracobunid phylogeny.png|2014 cladistic analysis]] placed [[anthracobunid]]s and desmostylians, two major extinct groups that have been considered to be non-African afrotheres, close to each other within [[Perissodactyla]].<ref name="Cooper2014"/>

==Phylogeny==
{{cladogram|style=font-size:100%;margin-top:25px;margin-bottom:25px|align=left|caption=A cladogram of [[Afrotheria]] based on molecular evidence<ref name=tabuce>{{cite journal|url=http://phylodiversity.net/azanne/csfar/images/d/d9/Afrotherian_mammals.pdf|archive-url=https://web.archive.org/web/20140517132203/http://phylodiversity.net/azanne/csfar/images/d/d9/Afrotherian_mammals.pdf|url-status=usurped|archive-date=May 17, 2014|first1=R.|last1=Tabuce| first2=R. J.|last2=Asher|first3=T.|last3=Lehmann|year=2008|title=Afrotherian mammals: a review of current data| journal=Mammalia|volume=72|pages=2–14|doi=10.1515/MAMM.2008.004|s2cid=46133294}}</ref>|cladogram={{clade|style=font-size:100%;width:640px;
|label1=[[Afrotheria]]
|1={{Clade
   |label1='''Paenungulata'''
   |1={{Clade
      |label1=[[Hyrax|Hyracoidea]]
      |1=[[Hyrax|Procaviidae]][[File:DendrohyraxEminiSmit white background.jpg|50 px]]
      |label2=[[Tethytheria]]
      |2={{Clade
         |label1=[[Sirenia]]
         |1={{Clade
            |1=[[Trichechidae]][[File:Manatee white background.jpg|60 px]]
            |2=[[Dugongidae]][[File:Dugong dugon Hardwicke white background.jpg|60 px]]
         }}
         |label2=[[Proboscidea]]
         |2=[[Elephantidae]][[File:Elephas africanus - 1700-1880 - Print - Iconographia Zoologica - (white background).jpg|70 px]]
         }}
      }}
   |label2=[[Afroinsectiphilia]]
   |2={{Clade
      |label1=[[Tubulidentata]]
      |1=[[Orycteropodidae]][[File:Aardvark2 (PSF) colourised.png|60 px]]
      |label2=[[Afroinsectivora]]
      |2={{Clade
         |label1=[[Macroscelidea]]
         |1=[[Macroscelididae]][[File:Rhynchocyon chrysopygus-J Smit white background.jpg|50 px]]
         |label2=[[Afrosoricida]]
         |2={{Clade
            |1=[[Chrysochloridae]][[File:The animal kingdom, arranged according to its organization, serving as a foundation for the natural history of animals (Pl. 18) (Chrysochloris asiatica).jpg|50 px]]
            |label2=[[Tenrecomorpha]]
            |2={{Clade
               |1=[[Potamogalidae]][[File:Potamogale velox illustration.jpg|50 px]]
               |2=[[Tenrecidae]][[File:Brehms Thierleben - Allgemeine Kunde des Thierreichs (1876) (Tenrec ecaudatus).jpg|50 px]]
               }}
            }}
         }}
      }}
   }}
}}
}}
{{clearleft}}

==Gallery==
{{Gallery
 |File:Ein klippschliefer.jpg|The rodent-like [[hyrax]] 
 |File:African Bush Elephant Mikumi.jpg|A bull [[bush elephant]]
 |File:Manatee with calf.PD.jpg|A [[manatee]] and her calf
 |File:Ocepeia_head_restoration_PLoS_ONE.png|''[[Ocepeia]]'', a basal species
 |File:Arsinoitherium by Antón.jpg|''[[Arsinoitherium]]'', a rhino-like [[Embrithopoda|embrithopod]]
 |File:Desmostylus 3 NT.jpg|''[[Desmostylus]]'', a member of  [[desmostylia]], the only extinct order of marine mammals.
}}

==Extinct orders==
Each of the [[extinction|extinct]] orders, the [[Embrithopoda]] and [[Desmostylia]],{{efn|name=possible_exclusions|[[Desmostylia|Desmostylians]], however, have been placed in [[Perissodactyla]] by a 2014 [[cladistics|cladistic]] analysis,<ref name=Cooper2014>{{cite journal |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. |title=Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls |journal=PLOS ONE |volume=9 |issue=10 |article-number=e109232 |year=2014 |doi=10.1371/journal.pone.0109232 |pmid=25295875 |pmc=4189980 |bibcode=2014PLoSO...9j9232C |doi-access=free }}</ref> and the taxonomic placement of embrithopods has also been questioned<ref name=Erdal2016>{{cite journal |last1=Erdal |first1=O. |last2=Antoine |first2=P.-O. |last3=Sen |first3=S. |last4=Smith |first4=A. |title=New material of ''Palaeoamasia kansui'' (Embrithopoda, Mammalia) from the Eocene of Turkey and a phylogenetic analysis of Embrithopoda at the species level |journal=Palaeontology |volume=59 |issue=5 |year=2016 |pages=631–655 |doi=10.1111/pala.12247 |bibcode=2016Palgy..59..631E |s2cid=89418652 |url=https://hal.sorbonne-universite.fr/hal-01346066/file/Erdal_2016_New_material_of.pdf}}</ref> though recently supported.<ref name=Gheerbrant2018>{{cite journal |author1=E. Gheerbrant |author2=A. Schmitt |author3=L. Kocsis |year=2018 |title=Early African fossils elucidate the origin of embrithopod mammals |journal=Current Biology |volume=28 |issue=13 |pages=2167–2173.e2|doi=10.1016/j.cub.2018.05.032 |pmid=30008332 |doi-access=free |bibcode=2018CBio...28E2167G |url=https://hal.sorbonne-universite.fr/hal-01845698/file/Gheerbrant%20et%20al%202018%20Early%20African%20Fossils%20Elucidate%20the%20Origin%20ofof%20Embrithopod%20mammals%20-%20Stylolophus_sans%20marque.pdf }}</ref>}} was as unique in its members' ways of making a living as the three orders that survive. Embrithopods were [[rhinoceros]]-like herbivorous mammals with [[plantigrade]] feet, and desmostylians were [[hippopotamus]]-like amphibious animals. Their walking posture and diet have been the subject of speculation, but tooth wear indicates that desmostylians browsed on terrestrial plants and had a posture similar to other large hoofed mammals.<ref name="comparison">{{cite journal |author1=Kleinschmidt, Traute |author2=Czelusniak, John |author3=Goodman, Morris |author4=Braunitzer, Gerhard |title=Paenungulata: A comparison of the hemoglobin sequences from Elephant, Hyrax, and Manatee |journal=Mol. Biol. Evol. |volume=3 |issue=5 |pages=427–435 |year=1986 |url=http://mbe.library.arizona.edu/data/1986/0305/5klei.pdf |pmid=3444412 |doi=10.1093/oxfordjournals.molbev.a040411 |access-date=2010-07-24 |archive-url=https://web.archive.org/web/20100630022747/http://mbe.library.arizona.edu/data/1986/0305/5klei.pdf |archive-date=2010-06-30 |df=dmy-all|doi-access=free }}</ref>

==See also==
* [[Altungulata]]
* [[Meridiungulata]]

==Notes==
{{notelist|1}}

==References==
{{reflist|25em}}

== Sources==
<!-- please add inline references -->
* {{cite journal |last1=Kleinschmidt |first1=Traute |last2=Czelusniak |first2=John |last3=Goodman |first3=Morris |last4=Braunitzer |first4=Gerhard |title=Paenungulata: A comparison of the hemoglobin sequences from elephant, hyrax, and manatee |journal=Mol. Biol. Evol. |volume=3 |issue=5 |pages=427–435 |year=1986 |url=http://mbe.library.arizona.edu/data/1986/0305/5klei.pdf |pmid=3444412 |doi=10.1093/oxfordjournals.molbev.a040411 |access-date=2010-07-24 |archive-url=https://web.archive.org/web/20100630022747/http://mbe.library.arizona.edu/data/1986/0305/5klei.pdf |archive-date=2010-06-30 |df=dmy-all|doi-access=free }}

* {{cite book |editor1=McKenna, M.C. |editor2=Bell, S.K. |year=1997 |title=Classification of Mammals above the Species Level |place=New York, NY |publisher=Columbia University Press |isbn=0-231-11013-8}}

* {{cite journal |last1=Seiffert |first1=Erik |last2=Guillon |first2=J.M. |year=2007 |title=A new estimate of afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence |journal=BMC Evolutionary Biology |volume=7 |page=13 |doi=10.1186/1471-2148-7-224 |pmid=17999766 |pmc=2248600 |url= |doi-access=free |bibcode=2007BMCEE...7..224S }}

* {{cite journal |author=Simpson, G.G. |year=1945 |title=The principles of classification and a classification of mammals |journal=[[American Museum of Natural History]] |volume=85 |pages=1–350 |hdl=2246/1104 |url=http://digitallibrary.amnh.org/dspace/handle/2246/1104}}

==Further reading==
*{{cite book |last=Gheerbrant |first=E. |title=The Rise of Placental Mammals: Origins and relationships of the major extant clades |year=2005 |publisher=Johns Hopkins University Press |location=Baltimore, MD |isbn=0-8018-8022-X |pages=84–105 |chapter-url=https://books.google.com/books?id=DhchVG_rbQ8C&pg=PA84 |via=Google Books |editor1=Rose, Kenneth D. |editor2=Archibald, J. David |chapter=Paenungulata (Sirenia, Proboscidea, Hyracoidea, and Relatives)}}

==External links==
*{{cite web |url=http://www.palaeos.com/Vertebrates/Units/530Altungulata/530.100.html#Paenungulata |archive-url=https://web.archive.org/web/20080313100141/http://www.palaeos.com/Vertebrates/Units/530Altungulata/530.100.html#Paenungulata |archive-date=2008-03-13 |website=Paleos.com |title=Vertebrates: Paenungulata}}

{{Paenungulata}}
{{Taxonbar|from=Q131950}}

[[Category:Paenungulata| ]]
[[Category:Paenungulatomorpha]]
[[Category:Mammal taxonomy]]
[[Category:Mammals of Africa]]
[[Category:Extant Selandian first appearances]]
[[Category:Mammal unranked clades]]