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==Taxonomy and evolution== ===Evolution=== [[File:Cranio Pliophoca etrusca, Orciano.JPG|thumb|Fossil ''[[Pliophoca]]'' skull]] The earliest known fossil earless seal is ''[[Noriphoca]] gaudini'' from the late [[Oligocene]] or earliest [[Miocene]] ([[Aquitanian (stage)|Aquitanian]]) of [[Italy]].<ref name=Noriphoca /> Other early fossil phocids date from the mid-Miocene, 15 million years ago in the north Atlantic.<ref name=Noriphoca /><ref>{{Cite journal|last1=Dewaele|first1=Leonard|last2=Lambert|first2=Olivier|last3=Louwye|first3=Stephen|date=2017-02-21|title=On ''Prophoca'' and ''Leptophoca'' (Pinnipedia, Phocidae) from the Miocene of the North Atlantic realm: redescription, phylogenetic affinities and paleobiogeographic implications|journal=PeerJ|language=en|volume=5|pages=e3024|doi=10.7717/peerj.3024|pmid=28243538|doi-access=free|pmc=5322758}}</ref> Until recently, many researchers believed that phocids evolved separately from [[otariids]] and [[Walrus|odobenids]]; and that they evolved from [[otter]]-like animals, such as ''[[Potamotherium]]'', which inhabited European freshwater lakes. Recent evidence strongly suggests a monophyletic origin for all pinnipeds from a single ancestor, possibly ''[[Enaliarctos]]'', most closely related to the [[Mustelidae|mustelids]] and [[bear]]s.<ref>{{Cite journal|last1=Paterson|first1=Ryan S.|last2=Rybczynski|first2=Natalia|last3=Kohno|first3=Naoki|last4=Maddin|first4=Hillary C.|date=2020|title=A Total Evidence Phylogenetic Analysis of Pinniped Phylogeny and the Possibility of Parallel Evolution Within a Monophyletic Framework|journal=Frontiers in Ecology and Evolution|language=en|volume=7|doi=10.3389/fevo.2019.00457|doi-access=free}}</ref> [[Monk seal]]s and [[elephant seal]]s were previously believed to have first entered the Pacific through the open straits between North and South America,<ref name="Fulton-2010" /> with the Antarctic true seals either using the same route or travelled down the west coast of Africa.<ref>{{cite book |author1=Savage, RJG |author2= Long, MR |name-list-style=amp|year=1986 |title= Mammal Evolution: an illustrated guide|url=https://archive.org/details/mammalevolutioni0000sava |url-access=registration |publisher= Facts on File|location=New York|pages= [https://archive.org/details/mammalevolutioni0000sava/page/94 94–95]|isbn= 978-0-8160-1194-0}}</ref> It is now thought that the [[monk seal]]s, [[elephant seal]]s, and [[Lobodontini|Antarctic seals]] all evolved in the southern hemisphere, and likely dispersed to their current distributions from more southern latitudes.<ref>{{Cite journal|last1=Rule|first1=James P.|last2=Adams|first2=Justin W.|last3=Marx|first3=Felix G.|last4=Evans|first4=Alistair R.|last5=Tennyson|first5=Alan J. D.|last6=Scofield|first6=R. Paul|last7=Fitzgerald|first7=Erich M. G.|date=2020-11-11|title=First monk seal from the Southern Hemisphere rewrites the evolutionary history of true seals|journal=Proceedings of the Royal Society B: Biological Sciences|volume=287|issue=1938|pages=20202318|doi=10.1098/rspb.2020.2318|pmid=33171079|pmc=7735288|doi-access=free}}</ref> ===Taxonomy=== {{Cladogram|align=left|title= |clade= {{clade |style=font-size:75%;line-height:75%;width:500px; |label1=[[Pinnipedia]] |1={{clade |label1='''Phocidae''' |1={{clade |label1=[[Phocinae]] |1={{clade |1=[[Bearded seal]]<!-- Erignathus barbatus --> |2={{clade |1=[[Hooded seal]]<!-- Cystophora cristata --> |label2=Phocini |2={{clade |1={{clade |1={{clade |1={{clade |1={{clade |1=[[Ringed seal]]<!-- Pusa hispida --> |2=[[Baikal seal]]<!-- Pusa sibirica --> }} |2=[[Caspian seal]]<!-- Pusa caspica --> }} |2={{clade |1=[[Spotted seal]]<!-- Phoca largha --> |2=[[Harbor seal]]<!-- Phoca vitulina --> }} }} |2=[[Grey seal]]<!-- Halichoerus grypus--> }} |2={{clade |1=[[Ribbon seal]]<!-- Histriophoca fasciata) --> |2=[[Harp seal]]<!-- Pagophilus groenlandicus --> }} }} }} }} |label2=[[Monachinae]] |2={{clade |1={{clade |label1=Lobodontini |1={{clade |1={{clade |1={{clade |1=[[Weddell seal]]<!-- Leptonychotes weddellii --> |2=[[Leopard seal]]<!-- Hydrurga leptonyx--> }} |2=[[Crabeater seal]]<!-- Lobodon carcinophaga --> }} |2=[[Ross seal]]<!-- Ommatophoca rossii --> }} |label2=Miroungini|sublabel2=(elephant seals) |2={{clade |1=[[Southern elephant seal]]<!-- Mirounga leonina --> |2=[[Northern elephant seal]]<!-- Mirounga angustirostris --> }} }} |label2=Monachini |2={{clade |1=[[Mediterranean monk seal]]<!-- Monachus monachus --> |2=[[Hawaiian monk seal]]<!-- Neomonachus schauinslandi --> |3=[[Caribbean monk seal]] }} }} }} |2={{clade |1=[[Otariidae]] ([[eared seal]]s) |2=[[Odobenidae]] ([[Walrus]]) }} }} }} |caption=[[Cladogram]] showing relationships among the phocids, combining several phylogenetic analyses.<ref name="Berta Churchill Boessenecker 2018">{{cite journal | last1=Berta | first1=Annalisa | last2=Churchill | first2=Morgan | last3=Boessenecker | first3=Robert W. | title=The Origin and Evolutionary Biology of Pinnipeds: Seals, Sea Lions, and Walruses | journal=Annual Review of Earth and Planetary Sciences | publisher=Annual Reviews | volume=46 | issue=1 | date=2018-05-30 | issn=0084-6597 | doi=10.1146/annurev-earth-082517-010009 | pages=203–228| bibcode=2018AREPS..46..203B | s2cid=135439365 | doi-access=free }}</ref> }} In the 1980s and 1990s, morphological [[phylogenetics|phylogenetic]] analysis of the phocids led to new conclusions about the interrelatedness of the various genera. More recent molecular phylogenetic analyses have confirmed the [[monophyly]] of the two phocid subfamilies ([[Phocinae]] and Monachinae).<ref>{{Cite journal|last1=Árnason|first1=Úlfur|last2=Bodin|first2=Kristina|last3=Gullberg|first3=Anette|last4=Ledje|first4=Christina|last5=Suzette|first5=Mouchaty|date=1995|title=A molecular view of pinniped relationships with particular emphasis on the true seals|journal=Journal of Molecular Evolution|volume=40 |issue=1 |pages=78–85|doi=10.1007/BF00166598|pmid=7714914 |bibcode=1995JMolE..40...78A |s2cid=7537924}}</ref><ref>{{Cite journal|last1=Arnason |first1=Ulfur |last2=Gullberg |first2=Anette |last3=Janke|first3=Axel |last4=Kullberg|first4=Morgan |last5=Lehman |first5=Niles |last6=Petrov |first6=Evgeny A.|last7=Väinölä|first7=Risto|date=2006-11-01 |title=Pinniped phylogeny and a new hypothesis for their origin and dispersal|journal=Molecular Phylogenetics and Evolution |volume=41|issue=2 |pages=345–354|doi=10.1016/j.ympev.2006.05.022 |pmid=16815048|bibcode=2006MolPE..41..345A }}</ref><ref>{{Cite journal|last1=Fulton|first1=Tara Lynn|last2=Strobeck|first2=Curtis |date=2010|title=Multiple markers and multiple individuals refine true seal phylogeny and bring molecules and morphology back in line |journal=Proceedings of the Royal Society B: Biological Sciences |volume=277 |issue=1684 |pages=1065–1070 |doi=10.1098/rspb.2009.1783|pmc=2842760|pmid=19939841}}</ref><ref name="Fulton-2010">{{Cite journal |last1=Fulton |first1=Tara L. |last2=Strobeck|first2=Curtis |date=2010|title=Multiple fossil calibrations, nuclear loci and mitochondrial genomes provide new insight into biogeography and divergence timing for true seals (Phocidae, Pinnipedia) |journal=Journal of Biogeography |volume=37|issue=5 |pages=814–829 |doi=10.1111/j.1365-2699.2010.02271.x|bibcode=2010JBiog..37..814F |s2cid=59436229 }}</ref> The Monachinae (known as the "southern" seals), is composed of three tribes; the Lobodontini, Miroungini, and Monachini. The four [[Antarctic]] genera ''[[leopard seal|Hydrurga]]'', ''[[Weddell seal|Leptonychotes]]'', ''[[crabeater seal|Lobodon]]'', and ''[[Ross seal|Ommatophoca]]'' are part of the [[tribe (biology)|tribe]] [[Lobodontini]]. Tribe Miroungini is composed of the [[elephant seal]]s. The [[Monk seal]]s (''[[Mediterranean monk seal|Monachus]]'' and ''[[Neomonachus]]'') are all part of the tribe Monachini.<ref>{{Cite journal |last1=Scheel |first1=Dirk-Martin|last2=Slater|first2=Graham J.|last3=Kolokotronis|first3=Sergios-Orestis |last4=Potter |first4=Charles W.|last5=Rotstein|first5=David S.|last6=Tsangaras|first6=Kyriakos |last7=Greenwood|first7=Alex D.|last8=Helgen|first8=Kristofer M.|date=2014|title=Biogeography and taxonomy of extinct and endangered monk seals illuminated by ancient DNA and skull morphology |journal=ZooKeys|issue=409 |pages=1–33|doi=10.3897/zookeys.409.6244|pmc=4042687|pmid=24899841|doi-access=free|bibcode=2014ZooK..409....1S }}</ref> Likewise, subfamily Phocinae (the "northern" seals) also includes three tribes; Erignathini (''[[bearded seal|Erignathus]])'', Cystophorini (''[[hooded seal|Cystophora)]]'', and [[Phocini]] (all other phocines). More recently, five species have been split off from ''Phoca'', forming three additional genera.<ref>{{Cite journal|last1=Berta|first1=Annalisa |last2=Churchill|first2=Morgan|date=2012 |title=Pinniped taxonomy: review of currently recognized species and subspecies, and evidence used for their description|journal=Mammal Review |volume=42 |issue=3|pages=207–234 |doi=10.1111/j.1365-2907.2011.00193.x|bibcode=2012MamRv..42..207B }}</ref> Alternatively the three monachine tribes have been evaluated to familiar status, which elephant seals and the Antarctic seals are more closely related to the phocines.<ref name="Bonner1994">{{cite book | last1 = Bonner | first1 = N. | year = 1994| title = Seals and Sea Lions of the World | publisher = Blandford |location= United Kingdom | pages = 1–224 |isbn = 9780816057177}}</ref> {{-}} ===Extant genera=== {| class="wikitable" |- ! Subfamily!! Tribe !! Image !! Genus !! species |- ! rowspan="7" style="text-align:center;"| '''Subfamily [[Monachinae]]''' ! rowspan="2" style="text-align:center;"| '''Tribe [[Monachini]]''' |[[File:Monachus monachus.jpg|175px]] ||''[[Monachus]]'' <small>Fleming, 1822</small> || * [[Mediterranean monk seal]], ''Monachus monachus'' |- |[[File:Monachus schauinslandi.jpg|175px]] ||''[[Neomonachus]]'' <small>Slater & Helgen, 2014</small> || *[[Hawaiian monk seal]], ''Neomonachus schauinslandi'' * †[[Caribbean monk seal]], ''Neomonachus tropicalis'' (probably extinct around 1952) |- ! rowspan="1" style="text-align:center;"| '''Tribe Miroungini''' |[[File:Mirounga angustirostris, Point Reyes (cropped).jpg|175px]] ||''[[Mirounga]]'' <small>Gray, 1827</small> || * [[Northern elephant seal]], ''Mirounga angustirostris'' * [[Southern elephant seal]], ''Mirounga leonina'' |- ! rowspan="4" style="text-align:center;"| '''Tribe [[Lobodontini]]''' |[[File:Ross-seal.jpg|175px]] ||''[[Ommatophoca]]'' <small>Gray, 1844</small> || * [[Ross seal]], ''Ommatophoca rossi'' |- |[[File:Antarctica 2013 Journey to the Crystal Desert (8369556693).jpg|175px]] ||''[[Lobodon]]'' <small>Gray, 1844</small> || * [[Crabeater seal]], ''Lobodon carcinophagus'' |- |[[File:Leopard seal basking on Iceberg.jpg|175px]] ||''[[Hydrurga]]'' <small>Gistel, 1848</small> || * [[Leopard seal]], ''Hydrurga leptonyx'' |- |[[File:Weddell Seal in Antarctica (8381225741).jpg|175px]] ||''[[Leptonychotes]]'' <small>Gill, 1872</small> || * [[Weddell seal]], ''Leptonychotes weddellii'' |- ! rowspan="7" style="text-align:center;"| '''Subfamily [[Phocinae]]''' ! rowspan="1" style="text-align:center;"| '''Tribe Cystophorini''' |[[File:2016-klappmützemuseumkoenig.jpg|175px]] ||''[[Cystophora (mammal)|Cystophora]]'' <small>Nilsson, 1820</small> || * [[Hooded seal]], ''Cystophora cristata'' |- ! rowspan="1" style="text-align:center;"| '''Tribe Erignathini''' |[[File:Bearded Seal at Svalbard (cropped).jpg|175px]] ||''[[Erignathus]]'' <small>Gill, 1866</small> || * [[Bearded seal]], ''Erignathus barbatus'' |- ! rowspan="5" style="text-align:center;"| '''Tribe Phocini''' |[[File:Seehund2cele4.jpg|175px]] ||''[[Phoca]]'' <small>Linnaeus, 1758</small> || * [[Harbor seal]] or common seal, ''Phoca vitulina'' * [[Spotted seal]], ''Phoca largha'' |- |[[File:Pusa hispida hispida NOAA 1.jpg|175px]] ||''[[Pusa]]'' <small>Scopoli, 1771</small> || * [[Ringed seal]], ''Pusa hispida'' (formerly ''Phoca hispida'') * [[Baikal seal]], ''Pusa sibirica'' (formerly ''Phoca sibirica'') * [[Caspian seal]], ''Pusa caspica'' (formerly ''Phoca caspica'') |- |[[File:Harp seal at False Cape (cut).jpg|175px]] ||''[[Pagophilus]]'' <small>Gray, 1844</small> || * [[Harp seal]], ''Pagophilus groenlandicus'' (formerly ''Phoca groenlandica'') |- |[[File:Ribbon-seal-male Josh London NOAAedit (16086029928) (cropped).jpg|175px]] ||''[[Histriophoca]]'' <small>Gill, 1873</small> || * [[Ribbon seal]], ''Histriophoca fasciata'' (formerly ''Phoca fasciata'') |- |[[File:Halichoerus grypus He3.jpg|175px]] ||''[[Halichoerus]]'' <small>Nilsson, 1820</small> || * [[Grey seal]], ''Halichoerus grypus'' |- |}
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