List of fossil pinnipedimorphs

Pinnipeds are marine mammals that evolved from arctoid carnivorans that includes seals, eared seals, and walruses. There are 34 recent species of pinnipeds[1] and 102 species of fossil pinnipeds and their stem-relatives (Pinnipedimorpha), collectively referred to as pinnipedimorphs.[2] Scientists still debate on which lineage of arctoid carnviroans are the closest relatives to the pinnipedimorphs, being either more closely related to bears or to musteloids.[3][4][5][6][7] Two stem-pinniped families found outside of Pinnipedimorpha, Amphicynodontidae and Semantoridae, were in the past considered to be subfamilies of Ursidae and Mustelidae respectively.[8] In comparison to the two other major groups or marine or sea mammals, cetaceans and sirenians, pinnipeds are a relatively younger group having appeared about 24 to 38 million years ago and are still able to return on land to breed.

Life restoration of Gomphotaria pugnax.

The list of fossil taxa is based on mostly the historiographical data from Valenzuela-Toro and Pyenson (2019).[2] The two stem-pinniped arctoid families Amphicynodontidae and Semantoridae are included here as well, although neither family are members of Pinnipedimorpha.[9] The list does not include the recently extinct Caribbean monk seal (Neomonachus tropicalis) and the Japanese sea lion (Zalophus japonicus), as they became extinct within the last two centuries.

Phylogeny

Below is an overall phylogeny of the taxa covered in the article followed after a composite tree in Berta et al. (2018)[8] and a total-evidence (combined molecular-morphological) dataset in Paterson et al. (2020):[9]

 
 
 

†Amphicynodontidae

 
 
 

†Semantoridae

 
 
Pinnipedimorpha
 

†Enaliarctidae

 
 
Pinnipediformes
 

†Pinnarctidion

 
 
 
 

†Pacificotaria

 
 

†Pteronarctos

 
 
Pinnipedia
Phocoidea
 

Phocidae

 
 
Otarioidea
 

†Desmatophocidae

 
 
 
 

Odobenidae

 
 
Panotariidae
 

†Eotaria

 
 

Otariidae

 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

Stem-pinniped arctoids

†Amphicynodontidae Simpson, 1945
Artist's restoration of Kolponomos newportensis.
  • †Amphicticeps Matthew & Granger, 1924
    • †Amphicticeps makhchinus Wang et al., 2005
    • †Amphicticeps dorog Wang et al., 2005
    • †Amphicticeps shackelfordi Matthew & Granger, 1924
  • †Drassonax Galbreath, 1953
    • †Drassonax harpagops Galbreath, 1953
  • †Parictis Scott, 1893 [Campylocynodon Chaffee, 1954]
    • †Parictis primaevus Scott, 1893
    • †Parictis personi (Chaffee, 1954) [Campylocynodon personi Chaffee, 1954]
    • †Parictis montanus Clark & Guensburg, 1972
    • †Parictis parvus Clark & Beerbower, 1967
    • †Parictis gilpini Clark & Guensburg, 1972
    • †Parictis dakotensis Clark, 1936
  • †Kolponomos Stirton, 1960
    • †Kolponomos newportensis Tedford et al., 1994
    • †Kolponomos clallamensis Stirton, 1960
  • †Allocyon Merriam, 1930
    • †Allocyon loganensis Merriam, 1930
  • †Wangictis L. de Bonis et al., 2019
    • †Wangictis tedfordi (Wang & Qiu, 2003) [Pachycynodon tedfordi Wang & Qiu, 2003]
  • †Pachycynodon Schlosser, 1888
    • †Pachycynodon tenuis Teilhard de Chardin, 1915
    • †Pachycynodon filholi Schlosser, 1888
    • †Pachycynodon boriei (Filhol, 1876) [Cynodon robustus Filhol, 1876; Pachycynodon gryei (Filhol, 1876); Pachycynodon leymeriei (Filhol, 1876); Pachycynodon curvirostris (Filhol, 1876); Pachycynodon ferratus (Quenstedt, 1885)]
    • †Pachycynodon crassirostris Schlosser, 1888
  • †Amphicynodon Filhol, 1881 [Cynodon Aymard, 1848, name already taken by a fish]
    • †Amphicynodon mongoliensis Janovskaja, 1970
    • †Amphicynodon teilhardi (Matthew & Granger, 1924) [Cynodon teilhardi Matthew & Granger, 1924]
    • †Amphicynodon typicus Schlosser, 1888
    • †Amphicynodon chardini Cirot & De Bonis, 1992
    • †Amphicynodon cephalogalinus Teilhard, 1915
    • †Amphicynodon gracilis (Filhol, 1874) [Cynodon gracilis Filhol, 1874]
    • †Amphicynodon crassirostris (Filhol, 1876) [Cynodon crassirostris Filhol, 1874]
    • †Amphicynodon brachyrostris (Filhol, 1876) [Cynodon brachyrostris Filhol, 1876]
    • †Amphicynodon leptorhynchus (Filhol, 1874) [Cynodon leptorhynchus Filhol, 1874]
    • †Amphicynodon velaunus (Aymard, 1846)

†Semantoridae Orlov, 1931
A restored skeleton of the semantorid Puijila darwini.
  • †Necromites V. V. Bogachev, 1940
    • †Necromites nestoris V. V. Bogachev, 1940
  • †Potamotherium Geoffroy, 1833
    • †Potamotherium miocenicum Peters, 1868
    • †Potamotherium valletoni Geoffroy, 1833
  • †Puijila Rybczynski et al., 2009
    • †Puijila darwini Rybczynski et al., 2009
  • †Semantor Orlov, 1931
    • †Semantor macrurus Orlov, 1931

Stem-pinniped pinnipedimorphs

†Enaliarctidae Mitchell & Tedford, 1973
A fossil of Enaliarctos mealsi encased in the matrix.
  • †Enaliarctos Mitchell & Tedford, 1973
    • †Enaliarctos barnesi Berta, 1991
    • †Enaliarctos mealsi Mitchell & Tedford, 1973
    • †Enaliarctos tedfordi Berta, 1991
    • †Enaliarctos mitchelli Barnes, 1979
    • †Enaliarctos emlongi Berta, 1991

Stem-pinniped pinnipediforms

†Pinnarctidion Barnes, 1979
  • †Pinnarctidion rayi Berta, 1994
  • †Pinnarctidion bishopi Barnes, 1979

†Pacificotaria Barnes, 1992
  • †Pacificotaria hadromma Barnes, 1992

†Pteronarctos Barnes, 1989
  • †Pteronarctos piersoni Barnes, 1990
  • †Pteronarctos goedertae Barnes, 1989

Pinnipeds
See also: List of pinnipeds § Prehistoric pinnipeds

Phocoidea J. E. Gray, 1821
Skull of Piscophoca pacifica.
Skeletal mount of Acrophoca longirostris.
  • Phocidae J. E. Gray, 1821
    • †Noriphoca Dewaele et al., 2018
      • †Noriphoca gaudini Guiscardi, 1870 [Monotherium gaudini Guiscardi, 1870]
    • †Devinophocinae Koretsky & Holec, 2002
      • †Kawas Cozzuol, 2001
        • †Kawas benegasorum Cozzuol, 2001
      • †Devinophoca Koretsky & Holec, 2002
        • †Devinophoca emryi Koretsky & Rahmat, 2015
        • †Devinophoca claytoni Koretsky & Rahmat, 2002
    • Monachinae E. L. Trouessart, 1897
      • †Australophoca Valenzuela‐Toro et al., 2016
        • †Australophoca Valenzuela‐Toro et al., 2016
      • †Virginiaphoca Dewaele et al., 2018
        • †Virginiaphoca magurai Dewaele et al., 2018
      • †Properiptychus Ameghino, 1897
        • †Properiptychus argentinus Ameghino, 1897
      • †Homiphoca Muizon & Hendey, 1980
        • †Homiphoca capensis (Hendey & Repenning, 1972) [Prionodelphis capensis Hendey & Repenning, 1972]
      • †Acrophoca Muizon, 1981
        • †Acrophoca longirostris Muizon, 1981
      • †Hadrokirus Amson & De Muizon, 2013
        • †Hadrokirus martini Amson & De Muizon, 2013
      • †Piscophoca Muizon, 1981
        • †Piscophoca pacifica Muizon, 1981
      • Monachini E. L. Trouessart, 1897
        • †Pristiphoca Gervais, 1853
          • †Pristiphoca rugidens (Meyer, 1845) [Monachus rugidens Meyer, 1845]
          • †Pristiphoca occitana Gervais, 1853
        • †Terranectes Rahmat et al., 2017
          • †Terranectes parvus Rahmat et al., 2017
          • †Terranectes magnus Rahmat et al., 2017
        • †Afrophoca Koretsky & Domning, 2014
          • †Afrophoca libyca Koretsky & Domning, 2014
        • †Pontophoca Kretzoi, 1941
          • †Pontophoca jutlandica Koretsky et al., 2014
          • †Pontophoca sarmatica (Alekseev, 1924) [Phoca sarmatica Alekseev, 1924]
          • †Pontophoca simionescui Kretzoi, 1941
        • †Palmidophoca Ginsburg & Janvier, 1975
          • †Palmidophoca callirhoe Ginsburg & Janvier, 1975
        • †Messiphoca Muizon, 1981
          • †Messiphoca mauretanica Muizon, 1981
        • †Auroraphoca Dewaele et al., 2018
          • †Auroraphoca atlantica Dewaele et al., 2018
        • †Pliophoca Tavani, 1941
          • †Pliophoca etrusca Tavani, 1941
        • †Sarcodectes Rule et al., 2020
          • †Sarcodectes magnus Rule et al., 2020
        • †Eomonachus Rule et al., 2020
          • †Eomonachus belegaerensis Rule et al., 2020
      • Miroungini Muizon, 1981
        • †Callophoca van Beneden, 1876
          • †Callophoca obscura van Beneden, 1876
    • Phocinae J. E. Gray, 1869
      • †Leptophoca True, 1906
        • †Leptophoca amphiatlantica Koretsky et al., 2012
        • †Leptophoca proxima (van Beneden, 1876) [Phoca proxima van Beneden, 1876; Leptophoca lenis True 1906]
      • †Nanophoca Dewaele et al., 2017
        • †Nanophoca vitulinoides (van Beneden, 1871) [Phoca vitulinoides van Beneden, 1871]
      • †Frisiphoca Dewaele et al., 2018
        • †Frisiphoca affine (van Beneden, 1876) [Monotherium affine van Beneden, 1876]
        • †Frisiphoca aberratum (van Beneden, 1876) [Monotherium aberratum van Beneden, 1876)]
      • †Cryptophoca Koretsky & Ray, 1994
        • †Cryptophoca maeotica (Nordmann, 1860) [Monotherium maeotica Koretsky 2001]
      • †Monotherium van Beneden, 1876
        • †Monotherium delognii van Beneden, 1876
      • †Prophoca van Beneden, 1876
        • †Prophoca rousseaui van Beneden, 1876
      • †Monachopsis Kretzoi, 1941
        • †Monachopsis pontica (Eichwald, 1850) [Monachus pontica Eichwald, 1850]
      • †Sarmatonectes Koretsky, 2001
        • †Sarmatonectes sintsovi Koretsky, 2001
      • †Phocanella van Beneden, 1877
        • †Phocanella pumila van Beneden, 1877
      • Erignathini Chapski, 1955
        • †Platyphoca van Beneden, 1876
          • †Platyphoca danica Koretsky et al., 2014
          • †Platyphoca vulgaris van Beneden, 1876
      • Cystophorini Burns & Fay, 1970
        • †Miophoca Zapfe, 1937
          • †Miophoca vetusta Zapfe, 1937
        • †Pachyphoca Koretsky & Rahmat, 2013
          • †Pachyphoca chapskii Koretsky & Rahmat, 2013
          • †Pachyphoca ukrainica Koretsky & Rahmat, 2013
      • Phocini J. E. Gray, 1821
        • Histriophoca Gill, 1873
          • †Histriophoca alekseevi Koretsky, 2001
        • †Gryphoca van Beneden, 1877
          • †Gryphoca nordica Koretsky et al., 2014
          • †Gryphoca similis van Beneden, 1877
        • †Batavipusa Koretsky & Peters, 2008
          • †Batavipusa neerlandica Koretsky & Peters, 2008
        • †Praepusa Kretzoi, 1941
          • †Praepusa pannonica Kretzoi, 1941
          • †Praepusa boeska Koretsky et al., 2015
          • †Praepusa magyaricus Koretsky, 2003
          • †Praepusa archankutica Antoniuk & Koretsky, 1984
          • †Praepusa vindobonensis (Toula, 1897) [Phoca vindobonensis Toula, 1897]
        • Phoca Linnaeus, 1758
          • †Phoca moori Newton, 1890

Otarioidea J. E. Gray, 1821
Skeletal mount of Allodesmus sp.
Reconstruction of Archaeodobenus akamatsui by Tatsuya Shinmura (Ashoro Museum of Paleontology).
Life restoration of Thalassoleon mexicanus..
  • †Desmatophocidae Hay, 1930
    • †Eodesmus Tate-Jones et al., 2020
      • †Eodesmus condoni Mitchell, 1968
    • †Allodesminae Mitchell, 1968
      • †Atopotarus Downs, 1956
        • †Atopotarus courseni Downs, 1956
      • †Allodesmus Kellogg, 1922
        • †Allodesmus packardi Barnes, 1972
        • †Allodesmus uraiporensis Tonomori et al., 2018
        • †Allodesmus naorai Kohno, 1996
        • †Allodesmus sinanoensis Nagao, 1941
        • †Allodesmus demerei Boessenecker & Churchill, 2018
        • †Allodesmus kernensis Kellogg, 1922
    • †Desmatophocinae Hay, 1930
      • †Desmatophoca Condon, 1906
        • †Desmatophoca brachycephala Barnes, 1987
        • †Desmatophoca oregonensis Condon, 1906
  • Odobenidae J. A. Allen, 1880
    • †Prototaria Takeyama & Ozawa, 1984
      • †Prototaria planicephala Kohno, 1994
      • †Prototaria primigena Takeyama & Ozawa, 1984
    • †Proneotherium Kohno et al., 1995
      • †Proneotherium repenningi Kohno et al., 1995
    • †Neotherium Kellogg, 1931
      • †Neotherium mirum Kellogg, 1931
    • †Kamtschatarctos Dubrovo, 1981
      • †Kamtschatarctos sinelnikovae Dubrovo, 1981
    • †Pseudotaria Kohno, 2006
      • †Pseudotaria muramotoi Kohno, 2006
    • †Archaeodobenus Tanaka & Kohno, 2015
      • †Archaeodobenus akamatsui Tanaka & Kohno, 2015
    • †Nanodobenus Velez & Salinas-Marquez, 2018
      • †Nanodobenus arandai Velez & Salinas-Marquez, 2018
    • †Pelagiarctos Barnes, 1988
      • †Pelagiarctos thomasi Barnes, 1988
    • †Titanotaria Magallanes et al., 2018
      • †Titanotaria orangensis Magallanes et al., 2018
    • †Imagotaria Mitchell, 1968
      • †Imagotaria downsi Mitchell, 1968
    • †Pontolis True, 1905
      • †Pontolis barroni Biewer, Velez-Juarbe & Parham, 2020[10]
      • †Pontolis kohnoi Biewer, Velez-Juarbe & Parham, 2020[10]
      • †Pontolis magnus (True, 1905) [Pontoleon magnus True, 1905]
    • †Osodobenus Biewer, Velez-Juarbe & Parham, 2020[10]
      • †Osodobenus eodon Biewer, Velez-Juarbe & Parham, 2020
    • Neodobenia Magallanes et al., 2018
      • †Dusignathinae Mitchell, 1968
        • †Gomphotaria Barnes & Rashke, 1991
          • †Gomphotaria pugnax Barnes & Rashke, 1991
        • †Dusignathinae Mitchell, 1968
          • †Dusignathus Kellogg, 1927
            • †Dusignathus seftoni Deméré, 1994
            • †Dusignathus santacruzensis Kellogg, 1927
      • Odobeninae J. A. Allen, 1880
        • †Aivukus Repenning & Tedford, 1977
          • †Aivukus cedrosensis Repenning & Tedford, 1977
        • †Protodobenus Horikawa, 1995
          • †Protodobenus japonicus Horikawa, 1995
        • †Pliopedia Kellogg, 1921
          • †Pliopedia pacifica Kellogg, 1921
        • †Ontocetus Leidy, 1859
          • †Ontocetus emmonsi Leidy, 1859
        • †Valenictus Mitchell, 1961
          • †Valenictus chulavistensis Deméré 1994
          • †Valenictus imperialensis Mitchell, 1961
        • Odobenus Brisson, 1762
          • †Odobenus mandanoensis Tomida, 1989
  • Panotariidae Velez-Juarbe, 2017
    • †Eotaria Boessenecker & Churchill, 2015
      • †Eotaria circa Velez-Juarbe, 2017
      • †Eotaria crypta Boessenecker & Churchill, 2015
    • Otariidae J. E. Gray, 1825
      • Callorhinae Muizon, 1978
        • †Pithanotaria Kellogg, 1925
          • †Pithanotaria starri Kellogg, 1925
        • Callorhinus J. E. Gray, 1859
          • †Callorhinus inouei (Kohno, 1992) [Thalassoleon inouei Kohno, 1992]
          • †Callorhinus macnallyae (Repenning & Tedford, 1977) [Thalassoleon macnallyae Repenning & Tedford, 1977]
          • †Callorhinus gilmorei Berta & Deméré, 1986
      • Otariinae J. E. Gray, 1825
        • †Thalassoleon Repenning & Tedford, 1977
          • †Thalassoleon mexicanus Repenning & Tedford, 1977
        • Zalophini J. E. Gray, 1869
          • †Proterozetes Barnes et al., 2006
            • †Proterozetes ulysses Barnes et al., 2006
          • †Oriensarctos Mitchell, 1968
            • †Oriensarctos watasei (Matsumoto, 1925) [Eumetopias watasei Matsumoto, 1925]
        • Otariini J. E. Gray, 1825
          • Otaria Péron, 1816
            • †Otaria fischeri Gervais & Ameghino, 1880
          • Neophoca J. E. Gray, 1866
          • †Hydrarctos Muizon, 1978
            • †Hydrarctos lomasiensis Muizon, 1978

See also
  • List of pinniped species

References
  1. Berta, A.; Churchill, M. (2012). "Pinniped taxonomy: review of currently recognized species and subspecies, and evidence used for their description". Mammal Review. 42 (3): 207–234. doi:10.1111/j.1365-2907.2011.00193.x.
  2. Valenzuela-Toro, A.; Pyenson, N. D. (2019). "What do we know about the fossil record of pinnipeds? A historiographical investigation". Royal Society Open Science. 6 (11): 207–234. Bibcode:2019RSOS....691394V. doi:10.1098/rsos.191394. PMC 6894555. PMID 31827869.
  3. Lento, G. M.; Hickson, R. E.; Chambers, G. K.; Penny, D. (1995). "Use of spectral analysis to test hypotheses on the origin of pinnipeds". Molecular Biology and Evolution. 12 (1): 28–52. doi:10.1093/oxfordjournals.molbev.a040189. PMID 7877495.
  4. Hunt, R. M. Jr.; Barnes, L. G. (1994). "Basicranial evidence for ursid affinity of the oldest pinnipeds" (PDF). Proceedings of the San Diego Society of Natural History. 29: 57–67.
  5. Higdon, J. W.; Bininda-Emonds, O. R.; Beck, R. M.; Ferguson, S. H. (2007). "Phylogeny and divergence of the pinnipeds (Carnivora: Mammalia) assessed using a multigene dataset". BMC Evolutionary Biology. 7: 216. doi:10.1186/1471-2148-7-216. PMC 2245807. PMID 17996107.
  6. Sato, J. J.; Wolsan, M.; Suzuki, H.; Hosoda, T.; Yamaguchi, Y.; Hiyama, K.; Kobayashi, M.; Minami, S. (2006). "Evidence from nuclear DNA sequences sheds light on the phylogenetic relationships of Pinnipedia: Single origin with affinity to Musteloidea". Zoological Science. 23 (2): 125–46. doi:10.2108/zsj.23.125. hdl:2115/13508. PMID 16603806. S2CID 25795496.
  7. Flynn, J. J.; Finarelli, J. A.; Zehr, S.; Hsu, J.; Nedbal, M. A. (2005). "Molecular phylogeny of the Carnivora (Mammalia): Assessing the impact of increased sampling on resolving enigmatic relationships". Systematic Biology. 54 (2): 317–37. doi:10.1080/10635150590923326. PMID 16012099.
  8. Berta, A.; Churchill, M. & Boessenecker, R.W. (2018). "The Origin and Evolutionary Biology of Pinnipeds: Seals, Sea Lions, and Walruses". Annual Review of Earth and Planetary Sciences. 0: 203–228. Bibcode:2018AREPS..46..203B. doi:10.1146/annurev-earth-082517-010009.
  9. Paterson, R. S.; Rybczynski, N.; Kohno, N.; Maddin, H. C. (2020). "Pinniped taxonomy: review of currently recognized species and subspecies, and evidence used for their description". Frontiers in Ecology and Evolution. 7 (457). doi:10.3389/fevo.2019.00457.
  10. Jacob N. Biewer; Jorge Velez-Juarbe; James F. Parham (2020). "Insights on the dental evolution of walruses based on new fossil specimens from California". Journal of Vertebrate Paleontology. in press: e1833896. doi:10.1080/02724634.2020.1833896.
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