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===Evolution=== {{Further|List of fossil pinnipedimorphs}} [[File:Puijila BW.jpg|thumb|right|Restoration of ''Puijila'']] One popular hypothesis suggested that pinnipeds are [[polyphyletic|diphyletic]] (descended from two ancestral lines), with walruses and otariids sharing a [[Most recent common ancestor|recent common ancestor]] with [[bear]]s; and phocids sharing one with [[Musteloidea]]. However, morphological and molecular evidence support a [[monophyletic]] origin.<ref name="Berta">Berta, A. "Pinniped evolution" in {{harvnb|Perrin|Würsig|Thewissen|2009|pp=861–866}}</ref> A 2021 genetic study found that pinnipeds are more closely related to musteloids.<ref>{{cite journal|author1=Hassanin, A.|author2=Veron, G.|author3=Ropiquet, A.|author4=van Vuuren, B. J.|author5=Lécu, A.|author6=Goodman, S. M.|author7=Haider, J.|author8=Nguyen, T. T.|year=2021|title=Evolutionary history of Carnivora (Mammalia, Laurasiatheria) inferred from mitochondrial genomes|journal=PLOS ONE|volume=16|issue=2|page=e0240770|doi=10.1371/journal.pone.0240770|pmid=33591975 |pmc=7886153 |bibcode=2021PLoSO..1640770H |doi-access=free }}</ref> Pinnipeds [[Cladogenesis|split]] from other caniforms 50 million years ago ([[Mya (unit)|mya]]) during the [[Eocene]].<ref name=leptin>{{cite journal |vauthors=Hammond JA, Hauton C, Bennett KA, Hall AJ |year=2012 |title=Phocid seal leptin: Tertiary structure and hydrophobic receptor binding site preservation during distinct leptin gene evolution |journal=PLOS ONE |volume=7 |issue=4 |page=e35395 |doi=10.1371/journal.pone.0035395 |editor1-last=Nikolaidis |editor1-first=Nikolas |bibcode=2012PLoSO...735395H |pmid=22536379 |pmc=3334926|doi-access=free }}</ref> The earliest fossils of pinnipeds date back to the [[Chattian|Late Oligocene]].<ref>{{Cite journal |last=Valenzuela-Toro |first=Ana |last2=Pyenson |first2=Nicholas D. |date=27 November 2019 |title=What do we know about the fossil record of pinnipeds? A historiographical investigation |url=https://royalsocietypublishing.org/doi/10.1098/rsos.191394 |journal=[[Royal Society Open Science]] |language=en |volume=6 |issue=11 |pages=191394 |doi=10.1098/rsos.191394 |issn=2054-5703 |pmc=6894555 |pmid=31827869 |access-date=13 November 2024}}</ref> Fossil animals representing basal lineages include ''[[Puijila]]'', of the [[Early Miocene]] in Arctic Canada. It resembled a modern otter, but shows evidence of quadrupedal swimming—retaining a form of aquatic locomotion that led to those employed by modern pinnipeds. ''[[Potamotherium]]'', which lived in the same period in Europe, was similar to ''Puijila'' but more aquatic.<ref name="Dawson2009">{{Cite journal |author1=Rybczynski, N. |author2=Dawson, M. R. |author3=Tedford, R. H. |doi=10.1038/nature07985 |title=A semi-aquatic Arctic mammalian carnivore from the Miocene epoch and origin of Pinnipedia |journal=Nature |volume=458 |issue=7241 |pages=1021–24 |year=2009 |pmid=19396145|bibcode=2009Natur.458.1021R |s2cid=4371413 }}</ref> The braincase of ''Potamotherium'' shows evidence that it used its [[whiskers]] to hunt, like modern seals.<ref>{{Cite journal |last1=Lyras |first1=G. A. |last2=Werdelin |first2=L |last3=van der Geer |first3=B. G. M. |last4=van der Geer |first4=A. A. E. |year=2023 |title=Fossil brains provide evidence of underwater feeding in early seals |journal=Communications Biology |language=en |volume=6 |issue=1 |page=747 |doi=10.1038/s42003-023-05135-z |pmid=37591929 |issn=2399-3642|pmc=10435510 }}</ref> Both ''Puijila'' and ''Potamotherium'' fossils have been found in lake deposits, suggesting that seal ancestors were originally adapted for fresh water.<ref name="Dawson2009"/> [[File:Enaliarctos mealsi.JPG|left|thumb|Fossil of ''Enaliarctos'']] ''Enaliarctos'', a fossil species of late [[Oligocene]]/early Miocene (24–22 mya) [[California]], closely resembled modern pinnipeds; it was adapted to an aquatic life with flippers and a flexible spine. Its teeth were more like land predators in that they were more adapted for [[Carnassial|shearing]]. Its hind-flippers may have allowed it to walk on land, and it probably did not leave coastal areas as much as its modern relatives. ''Enaliarctos'' was likely more of a fore-flipper swimmer, but could probably swim with either pair.<ref name="Berta"/> One species, ''Enaliarctos emlongi'', exhibited notable [[sexual dimorphism]], suggesting that this physical characteristic may have been an important driver of pinniped evolution.<ref name=Cullenetal2014>{{cite journal|last1=Cullen|first1=T. M.|last2=Fraser|first2=D.|last3=Rybczynski|first3=N.|last4=Shroder-Adams|first4=C.|title=Early evolution of sexual dimorphism and polygyny in Pinnipeda|journal=Evolution|date=2014|volume=68|issue=5|pages=1469–1484|doi=10.1111/evo.12360|pmid=24548136|s2cid=10389875|doi-access=free}}</ref> A closer relative of extant pinnipeds was ''[[Pteronarctos]]'', which lived in [[Oregon]] 19–15 mya. As in modern seals, the [[maxilla]] or upper jaw bone of ''Pteroarctos'' intersects with the [[Orbit (anatomy)|orbital wall]]. The extinct family [[Desmatophocidae]] lived 23–10 mya in the North Pacific. They had long skulls that with large orbits, interlocked [[zygomatic bone]]s and rounded [[molar (tooth)|molar]]s and [[premolars]]. They also were sexually dimorphic and may have been capable of swimming with both or either pair of flippers.<ref name="Berta"/> They are grouped with modern pinnipeds, but there is debate as to whether they are more closely related to phocids or to otariids and walruses.<ref>{{cite journal|last1=Boessenecker|first1=R. W.|last2=Churchill|first2=M.|year=2018|title=The last of the desmatophocid seals: a new species of ''Allodesmus'' from the upper Miocene of Washington, USA, and a revision of the taxonomy of Desmatophocidae|journal=Zoological Journal of the Linnean Society|volume=184|issue=1|pages=211–235|doi=10.1093/zoolinnean/zlx098}}</ref><ref name="Berta Churchill Boessenecker 2018"/> A 2024 study places them within Phocidae, specifically Phocinae.<ref name=Park2024/> [[Image:Archaeodobenus restoration.PNG|thumb|Reconstruction of ''[[Archaeodobenus]] akamatsui'' family Odobenidae]] The ancestors of the Otarioidea and Phocoidea diverged around 25 mya.<ref name=Nyakatura>{{cite journal|last1=Nyakatura|first1=K|last2=Bininda-Emonds|first2=O. R. P.|year=2012|title=Updating the evolutionary history of Carnivora (Mammalia): a new species-level supertree complete with divergence time estimates|journal=BMC Biology|volume=10|page=12|doi=10.1186/1741-7007-10-12|doi-access=free|pmid=22369503|pmc=3307490}}</ref> Phocids are known to have existed for at least 15 million years,<ref name="Berta"/> and molecular evidence supports a divergence of the Monachinae and Phocinae lineages around this time.<ref name="Berta Churchill Boessenecker 2018"/> The fossil genera ''[[Monotherium]]'' and ''[[Leptophoca]]'' of southeastern North America represent the earliest members of Monachinae and Phocinae respectively.<ref name="Berta"/> Both lineages may have originated in the North Atlantic, and likely reached the Pacific via the [[Central American Seaway]]. Phocines mainly stayed in the Northern Hemisphere, while the monachines diversified southward.<ref name="Berta Churchill Boessenecker 2018"/> The [[Lineage (evolution)|lineages]] of Otariidae and Odobenidae split around 20 mya.<ref name=Nyakatura/> The earliest fossil records of otariids are in North Pacific and dated to around 11 mya. Early fossil genera include ''[[Pithanotaria]]'' and ''[[Thalassoleon]]''.<ref name="Berta"/> The ''Callorhinus'' lineage split the earliest, followed by the ''[[Eumetopias]]''/''[[Zalophus]]'' lineage and then the rest, which colonized the Southern Hemisphere.<ref name="Berta"/><ref name="Berta Churchill Boessenecker 2018"/> The earliest fossils of Odobenidae—''[[Prototaria]]'' of Japan and ''[[Proneotherium]]'' of Oregon—date to 18–16 mya. These primitive walruses had normal sized canines and fed on fish instead of mollusks. Later taxa like ''[[Gomphotaria]]'', ''[[Pontolis]]'' and ''[[Dusignathus]]'' had longer canines on both the upper and lower jaw. The familiar long upper tusks developed in the genera ''[[Valenictus]]'' and ''Odobenus''. The lineage of the modern walrus may have spread from the North Pacific to the North Atlantic through the Caribbean and Central American Seaway 8–5 mya, and then back to the North Pacific via the Arctic 1 mya, or to the Arctic and subsequently the North Atlantic during the [[Pleistocene]].<ref name="Berta"/>
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