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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 zitteli (†Embrithopoda); bottom: Asian elephant (Proboscidea), West Indian manatee (Sirenia).

| taxon = Paenungulata

| authority = Simpson, 1945

| display_parents = 2

| subdivision_ranks = Subgroups

| subdivision = * Hyracoidea

  • †Sudamericungulata?{{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}}
  • Tethytheria

| range_map = Paenungulata map cropped.jpg

}}

Paenungulata (from Latin paene "almost" + ungulātus "having hoofs") is a clade of "sub-ungulates", which groups three extant mammal orders: Proboscidea (including elephants), Sirenia (sea cows, including dugongs and manatees), and Hyracoidea (hyraxes). At least two more possible orders are known only as fossils, namely Embrithopoda and Desmostylia.

Molecular evidence indicates that Paenungulata (or at least its extant members) is part of the cohort Afrotheria, an ancient assemblage of mainly African mammals of great diversity. The other members of this cohort are the orders Afrosoricida (tenrecs and golden moles), Macroscelidea (elephant shrews) and Tubulidentata (aardvarks).

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.{{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/|url-status=dead|archive-date=June 17, 2013}}

History

In 1945, George Gaylord Simpson used traditional taxonomic techniques to group these spectacularly diverse mammals in the superorder he named Paenungulata ("almost ungulates"), but there were many loose threads in unravelling their genealogy.{{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 |pages=13 |doi=10.1186/1471-2148-7-224 |pmid=17999766 |pmc=2248600 |url= |doi-access=free |bibcode=2007BMCEE...7..224S }} 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 parsimonious cladograms 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).

However, a 2014 cladistic analysis placed anthracobunids and desmostylians, two major extinct groups that have been considered to be non-African afrotheres, close to each other within Perissodactyla.

Phylogeny

{{cladogram|style=font-size:100%;margin-top:25px;margin-bottom:25px|align=left|caption=A cladogram of Afrotheria based on molecular evidence{{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}}|cladogram={{clade|style=font-size:100%;width:640px;

|label1=Afrotheria

|1={{Clade

|label1=Paenungulata

|1={{Clade

|label1=Hyracoidea

|1=Procaviidae50 px

|label2=Tethytheria

|2={{Clade

|label1=Sirenia

|1={{Clade

|1=Trichechidae60 px

|2=Dugongidae60 px

}}

|label2=Proboscidea

|2=Elephantidae70 px

}}

}}

|label2=Afroinsectiphilia

|2={{Clade

|label1=Tubulidentata

|1=Orycteropodidae60 px

|label2=Afroinsectivora

|2={{Clade

|label1=Macroscelidea

|1=Macroscelididae50 px

|label2=Afrosoricida

|2={{Clade

|1=Chrysochloridae50 px

|label2=Tenrecomorpha

|2={{Clade

|1=Potamogalidae50 px

|2=Tenrecidae50 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 embrithopod

|File:Desmostylus 3 NT.jpg|Desmostylus, a member of desmostylia, the only extinct order of marine mammals.

}}

Extinct orders

Each of the extinct orders, the Embrithopoda and Desmostylia,{{efn|name=possible_exclusions|Desmostylians, however, have been placed in Perissodactyla by a 2014 cladistic analysis,{{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 |pages=e109232 |year=2014 |doi=10.1371/journal.pone.0109232 |pmid=25295875 |pmc=4189980 |bibcode=2014PLoSO...9j9232C |doi-access=free }} and the taxonomic placement of embrithopods has also been questioned{{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}} though recently supported.{{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 }}}} 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.{{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 |url-status=dead |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 }}

See also

Notes

{{notelist|1}}

References

{{reflist|25em}}

Sources

  • {{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 |url-status=dead |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 |pages=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=080188022X |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 |url-status=dead |archive-date=2008-03-13 |website=Paleos.com |title=Vertebrates: Paenungulata}}

{{Paenungulata}}

{{Taxonbar|from=Q131950}}

Category:Paenungulatomorpha

Category:Mammal taxonomy

Category:Mammals of Africa

Category:Extant Selandian first appearances

Category:Mammal unranked clades