Editing Ungulate (section)

==Classifications==

===History===
Euungulata is a clade (or in some taxonomies, a grand order) of mammals. The two extant orders of ungulates are the [[Odd-toed ungulate|Perissodactyla (odd-toed ungulates)]] and [[Even-toed ungulate|Artiodactyla (even-toed ungulates)]]. [[Hyrax|Hyracoidea (hyraxes)]], [[Sirenia|Sirenia (sea cows, dugongs and manatees)]] and [[Proboscidea|Proboscidea (elephants)]] were in the past grouped within the clade "Ungulata", later found to be a [[polyphyletic]] and invalid clade. The three orders of Paenungulata are considered a clade and grouped in the Afrotheria clade, while Euungulata is grouped under the [[Laurasiatheria]] clade.<ref name="euungulata"/>

In 2009, morphological<ref>{{cite journal | last1 = Asher | first1 = RJ | last2 = Bennet | first2 = N | last3 = Lehmann | first3 = T | year = 2009 | title = The new framework for understanding placental mammal evolution | journal = BioEssays | volume = 31 | issue = 8| pages = 853–864 | doi=10.1002/bies.200900053 | pmid=19582725| s2cid = 46339675 | doi-access = free }}</ref><ref name="Tabuce">{{cite journal | last1 = Tabuce | first1 = R. | last2 = Marivaux | first2 = L. | last3 = Adaci | first3 = M. | last4 = Bensalah | first4 = M. | last5 = Hartenberger | first5 = J. L. | display-authors = etal   | year = 2007 | title = Early tertiary mammals from north Africa reinforce the molecular afrotheria clade | journal = Proceedings of the Royal Society B | volume = 274 | issue = 1614| pages = 1159–1166 | doi=10.1098/rspb.2006.0229 | pmid=17329227 | pmc=2189562}}</ref><ref name="Seiffert">{{cite journal | last1 = Seiffert | first1 = E | year = 2007 | title = A new estimate of afrotherian phylogeny based on simultaneous analysis of genomic, morphological, and fossil evidence | journal = BMC Evol Biol | volume = 7 | page = 13 | doi = 10.1186/1471-2148-7-224 | pmid = 17999766 | pmc = 2248600 | doi-access = free }}</ref><ref name="SanchezVillagra">{{cite journal | last1 = Sanchez-Villagra | first1 = M. R. | last2 = Narita | first2 = Y. | last3 = Kuratani | first3 = S. | year = 2007 | title = Thoracolumbar vertebral number: the first skeletal synapomorphy for afrotherian mammals | journal = Syst Biodivers | volume = 5 | pages = 1–17 | doi = 10.1017/S1477200006002258 | s2cid = 85675984 }}</ref> and molecular<ref>{{cite journal | last1 = Springer | first1 = MS | last2 = Stanhope | first2 = MJ | last3 = Madsen | first3 = O | last4 = de Jong | first4 = WW | year = 2004 | title = Molecules consolidate the placental mammal tree | journal = Trends Ecol Evol | volume = 19 | issue = 8| pages = 430–438 | doi=10.1016/j.tree.2004.05.006 | pmid=16701301| s2cid = 1508898 }}</ref><ref>{{cite journal | last1 = Robinson | first1 = M. A. Yang | last2 = Fu | first2 = T. J. | last3 = Ferguson-Smith | first3 = B. | year = 2004 | title = Cross-species chromosome painting in the golden mole and elephant-shrew: support for the mammalian clades Afrotheria and Afroinsectiphillia but not Afroinsectivora | journal = Proceedings of the Royal Society B | volume = 271 | issue = 1547| pages = 1477–1484 | doi=10.1098/rspb.2004.2754 | pmid=15306319 | pmc=1691750}}</ref> work found that aardvarks, hyraxes, sea cows, and elephants were more closely related to each other and to [[sengi]]s, [[tenrec]]s, and [[golden mole]]s than to the perissodactyls and artiodactyls, and form the clade [[Afrotheria]]. Elephants, sea cows, and hyraxes were grouped together in the clade [[Paenungulata]], while the aardvark has been considered as either a close relative to them or a close relative to sengis in the clade [[Afroinsectiphilia]].<ref name=Seiffert2007>{{cite journal | author = Seiffert, E.R. | 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 | pages = 13 | url= | doi = 10.1186/1471-2148-7-224 | pmid = 17999766 | last2 = Guillon | first2 = JM | pmc=2248600 | issue=1 | doi-access = free }}</ref> This is a striking example of [[convergent evolution]].<ref>{{cite book|last=Dawkins|first=Richard|title=The Ancestor's Tale|publisher=Mariner Books|location=Boston|year=2005|page=[https://archive.org/details/ancestorstale00rich_0/page/195 195]|isbn=978-0-618-61916-0|url=https://archive.org/details/ancestorstale00rich_0/page/195}}</ref>

There is some dispute as to whether this smaller Euungulata is a [[cladistic]] (evolution-based) group, or merely a [[phenetics|phenetic]] group ([[Form classification|form taxon]]) or [[folk taxon]] (similar, but not necessarily related). Some studies have indeed found the [[mesaxonia]]n ungulates and paraxonian ungulates to form a monophyletic lineage,<ref name=asher>{{cite journal|last1=Asher|first1=Robert J|last2=Helgen|first2=Kristofer M|title=Nomenclature and placental mammal phylogeny|journal=BMC Evolutionary Biology|volume=10|issue=1|year=2010|pages=102|doi=10.1186/1471-2148-10-102|pmid=20406454|pmc=2865478 |doi-access=free }}</ref><ref name=Spauldingetal2009>{{cite journal|pmc=2740860 | pmid=19774069 | doi=10.1371/journal.pone.0007062 | volume=4 | issue=9 | title=Relationships of Cetacea (Artiodactyla) among mammals: increased taxon sampling alters interpretations of key fossils and character evolution | year=2009 | journal=PLOS ONE | pages=e7062|bibcode = 2009PLoSO...4.7062S|editor1-last=Farke|editor1-first=Andrew Allen|last1=Spaulding|first1=Michelle|last2=O'Leary|first2=Maureen A.|last3=Gatesy|first3=John | doi-access=free }}</ref><ref name= "Nery">{{Cite journal | last1 = Nery | first1 = M. F. | last2 = González | first2 = D. M. J. | last3 = Hoffmann | first3 = F. G. | last4 = Opazo | first4 = J. C. | title = Resolution of the laurasiatherian phylogeny: Evidence from genomic data | doi = 10.1016/j.ympev.2012.04.012 | journal = Molecular Phylogenetics and Evolution | volume = 64 | issue = 3 | pages = 685–689 | year = 2012 | pmid =  22560954}}</ref> closely related to either the [[Ferae]] (the [[carnivora]]ns and the [[pangolin]]s)<ref>{{cite journal|last1=Beck|first1=Robin MD|last2=Bininda-Emonds|first2=Olaf RP|last3=Cardillo|first3=Marcel|last4=Liu|first4=Fu-Guo|last5=Purvis|first5=Andy|journal=BMC Evolutionary Biology|volume=6|issue=1|year=2006|pages=93|doi=10.1186/1471-2148-6-93|title=A higher-level MRP supertree of placental mammals|pmid=17101039|pmc=1654192 |doi-access=free }}</ref><ref>{{cite journal |author=Zhou, X. |year=2011 |title=Phylogenomic analysis resolves the interordinal relationships and rapid diversification of the Laurasiatherian mammals  |journal=[[Systematic Biology]] |doi=10.1093/sysbio/syr089 |volume=61 |pages=150–64 |pmid=21900649 |pmc=3243735 |issue=1 |first2=S. |last2=Xu |first3=J. |last3=Xu |first4=B. |last4=Chen |first5=K. |last5=Zhou |first6=G. |last6=Yang|display-authors=etal}}</ref> in the clade [[Fereuungulata]] or to the [[bat]]s.<ref>{{cite web | title = Researchers Greatly Improve Evolutionary Tree of Life for Mammals | url = http://newsroom.ucr.edu/2729 | url-status = dead | archive-url = https://web.archive.org/web/20111001174523/http://newsroom.ucr.edu/2729 | archive-date = 1 October 2011 | date = 22 September 2011 | website = UCR Newsroom | publisher = UC Riverside | access-date = 9 April 2021}}</ref> Other studies found the two orders not that closely related, as some place the perissodactyls as close relatives to bats and Ferae in [[Pegasoferae]]<ref>{{Cite journal | last1 = Nishihara | first1 = H.| last2 = Hasegawa | first2 = M.| last3 = Okada | first3 = N.| title = Pegasoferae, an unexpected mammalian clade revealed by tracking ancient retroposon insertions | doi = 10.1073/pnas.0603797103 | journal = [[Proceedings of the National Academy of Sciences of the United States of America|Proceedings of the National Academy of Sciences]]| volume = 103 | issue = 26 | pages = 9929–9934 | year = 2006 | pmid = 16785431| pmc = 1479866| bibcode = 2006PNAS..103.9929N| doi-access = free}}</ref> and others place the artiodactyls as close relatives to bats.<ref name=Gatesy>{{cite journal | doi=10.1016/j.ympev.2012.10.012 | volume=66 | issue=2 | title=A phylogenetic blueprint for a modern whale | journal=Molecular Phylogenetics and Evolution | pages=479–506 | pmid=23103570 | last1 = Gatesy | first1 = J | last2 = Geisler | first2 = JH | last3 = Chang | first3 = J | last4 = Buell | first4 = C | last5 = Berta | first5 = A | last6 = Meredith | first6 = RW | last7 = Springer | first7 = MS | last8 = McGowen | first8 = MR| date=February 2013 }}</ref>

===Taxonomy===
[[File:Zebras Ngorongoro Crater.jpg|thumb|[[Plains zebra]]]]
[[File:Diceros bicornis.jpg|thumb|[[Black rhinoceros]]]]
[[File:Elaphurus davidianus 001.jpg|thumb|[[Père David's deer]]]]
[[File:Hippopotamus - 04.jpg|thumb|[[Hippopotamus]]]]
[[File:Bluewhale877.jpg|thumb|[[Blue whale]]]]
[[File:Comdolph.jpg|thumb|[[Common dolphin]]]]
Below is a simplified taxonomy (assuming that ungulates do indeed form a natural grouping) with the extant families, in order of the relationships. Keep in mind that there were still some grey areas of conflict, such as the case with the relationship between the [[pecora]]n families and the [[baleen whale]] families. See each family for the relationships of the species as well as the controversies in their respective articles.

* '''Euungulata<ref name=asher/>'''
** '''[[Odd-toed ungulate|Perissodactyla]]''' (Mesaxonian ungulates)
*** '''Hippomorpha'''
**** [[Equidae]]: Horses, asses and zebras
*** '''[[Ceratomorpha]]'''
**** [[Tapiridae]]: Tapirs
**** [[Rhinocerotidae]]: Rhinoceroses
** '''[[Artiodactyla]] (= [[Cetartiodactyla]])''' (Paraxonian ungulates)
*** '''[[Tylopoda]]'''
**** [[Camelidae]]: Camels and llamas
*** '''Artiofabula'''
**** '''[[Suina]]'''
***** [[Tayassuidae]]: Peccaries
***** [[Suidae]]: Pigs
**** '''[[Cetruminantia]]'''
***** '''[[Ruminantia]]'''
****** [[Tragulidae]]: Chevrotains
****** '''[[Pecora|Cervoidea]]'''
******* [[Antilocapridae]]: Pronghorn
******* [[Giraffidae]]: Giraffes and okapi
******* [[Cervidae]]: Deer
******* [[Moschidae]]: Musk deer
******* [[Bovidae]]: Cattle and antelopes
***** '''[[Whippomorpha]]'''
****** [[Hippopotamidae]]: Hippopotamuses
****** '''[[Cetacea]]'''
******* '''[[Mysticeti]]'''
******** [[Balaenidae]]: Bowhead and right whales
******** [[Cetotheriidae]]: Pygmy right whale
******** [[Balaenopteridae]]: Rorquals
******* '''[[Odontoceti]]'''
******** '''[[Physeteroidea]]'''
********* [[Physeteridae]]: Sperm whale
********* [[Kogiidae]]: Lesser sperm whales
******** '''[[Platanistoidea]]'''
********* [[Platanistidae]]: Indian river dolphins
******** '''[[Ziphioidea]]'''
********* [[Ziphiidae]]: Beaked whales
******** '''[[Lipotidae|Lipotoidea]]'''
********* [[Lipotidae]]: Baiji (functionally extinct)
******** '''[[Inioidea]]'''
********* [[Iniidae]]: Amazonian river dolphins
********* [[Pontoporiidae]]: La Plata dolphin
******** '''[[Delphinoidea]]'''
********* [[Monodontidae]]: Beluga and narwhal
********* [[Phocoenidae]]: Porpoises
********* [[Delphinidae]]: Oceanic dolphins

===Phylogeny===
Below is the general consensus of the phylogeny of the ungulate families.<ref name=Gatesy/><ref>{{cite journal | last1 = Kim | first1 = S. L. | last2 = Thewissen | first2 = J. G. | last3 = Churchill | first3 = M. M. | last4 = Suydam | first4 = R. S. | last5 = Ketten | first5 = D. R. | last6 = Clementz | first6 = M. T. | year = 2014 | title = Unique biochemical and mineral composition of whale ear bones | url = http://darchive.mblwhoilibrary.org/bitstream/1912/6709/1/Kim_2014_Physiol_Biochem_Zool.pdf| journal = Physiological and Biochemical Zoology | volume = 87 | issue = 4| pages = 576–584 | doi=10.1086/676309| pmid = 24940922 | hdl = 1912/6709 | s2cid = 27076616 | hdl-access = free }}</ref>
{{clade|style=font-size:100%
|label1='''Euungulata'''
|1={{Clade
   |label1=[[Perissodactyla]]
   |1={{Clade
      |1=[[Equidae]][[File:Equus quagga (white background).jpg|50 px]]
      |label2=
      |2={{Clade
         |1=[[Tapiridae]][[File:Tapir white background.jpg|50 px]]
         |2=[[Rhinocerotidae]][[File:Rhino white background.jpg|50 px]]
         }}
      }}
   |label2=[[Artiodactyla]]
   |2={{Clade
      |label1=[[Tylopoda]]
      |1=[[Camelidae]][[File:Cladogram of Cetacea within Artiodactyla (Camelus bactrianus).png|50 px]]
      |label2=[[Artiofabula]]
      |2={{Clade
         |label1=[[Suina]]
         |1={{Clade
            |1=[[Tayassuidae]][[File:Pecari tajacu white background.jpg|50 px]]
            |2=[[Suidae]][[File:Recherches pour servir à l'histoire naturelle des mammifères (Pl. 80) (white background).jpg|50 px]]
            }}
         |label2=[[Cetruminantia]]
         |2={{Clade
            |label1=[[Ruminantia]]
            |1={{Clade
               |1=[[Tragulidae]][[File:Tragulus napu - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|50 px]]
               |2={{Clade
                  |1=[[Antilocapridae]][[File:Antilocapra white background.jpg|50 px]]
                  |2={{Clade
                     |1=[[Giraffidae]][[File:Giraffa camelopardalis Brockhaus white background.jpg|50 px]]
                     |2={{Clade
                        |1=[[Cervidae]][[File:The deer of all lands (1898) Hangul white background.png|50 px]]
                        |2={{Clade
                           |1=[[Moschidae]][[File:Moschus chrysogaster white background.jpg|50 px]]
                           |2=[[Bovidae]][[File:Birds and nature (1901) (14562088237) white background.jpg|50 px]]
                           }}
                        }}
                     }}
                  }}
               }}
            |label2=[[Whippomorpha]]
            |2={{Clade
               |1=[[Hippopotamidae]][[File:Voyage en Abyssinie Plate 2 (white background).jpg|50 px]]
               |2={{Clade 
                  |label1=[[Cetacea]]
                  |1={{Clade 
                     |label1=[[Mysticeti]]
                     |1={{Clade
                        |1=[[Balaenidae]][[File:Balaena mysticetus NOAA.jpg|50 px]]
                        |2={{Clade
                           |1=[[Cetotheriidae]][[File:Caperea marginata 3 flipped.jpg|50 px]]
                           |2=[[Balaenopteridae]] [[File:Megaptera novaeangliae NOAA.jpg|50 px]]
                           }}
                        }}
                     |label2=[[Odontoceti]]
                     |2={{Clade
                        |1={{Clade
                           |1=[[Physeteridae]][[File:Physeter macrocephalus NOAA.jpg|50 px]]
                           |2=[[Kogiidae]][[File:Kogia_sima_(transparent_background).png|50 px]]
                           }}
                        |2={{Clade 
                           |1=[[Platanistidae]][[File:Ganges.png|50 px]]
                           |2={{Clade
                              |1=[[Ziphiidae]][[File:Ziphius cavirostris NOAA.jpg|50 px]]
                              |2={{Clade
                                 |1=†[[Lipotidae]][[File:Lipotes vexillifer.png|50 px]]
                                 |2={{Clade 
                                    |1={{Clade
                                       |1=[[La Plata dolphin|Pontoporiidae]][[File:Pontoporia blainvillei.jpg|50 px]]
                                       |2=[[Iniidae]][[File:Inia_geoffrensis.jpg|50 px]]
                                       }}
                                    |2={{Clade
                                       |1=[[Delphinidae]] [[File:Orcinus orca NOAA 2.jpg|50 px]]
                                       |2={{Clade
                                          |1=[[Phocoenidae]] [[File:Vaquita swimming through the void. (Phocoena Sinus).png|50 px]]
                                          |2=[[Monodontidae]] [[File:Delphinapterus leucas NOAA.jpg|50 px]]
                                          }}
                                       }}
                                    }}
                                 }}
                              }}
                           }}
                        }}
                     }}
                  }}
               }}
            }}
         }}
      }}
   }}
}}