List of fossil pinnipedimorphs explained

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 of 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.

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. 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]

Stem-pinniped arctoids

†Amphicynodontidae Simpson, 1945

†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 [[Cynodon (fish)|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

†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

†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
†Pinnarctidion iverseni Everett, Deméré & Wyss, 2023

†Pacificotaria Barnes, 1992

†Pacificotaria hadromma Barnes, 1992

†Pteronarctos Barnes, 1989

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

Pinnipeds

Phocoidea J.E. Gray, 1821

Phocidae J. E. Gray, 1821
- †Noriphoca Dewaele et al., 2018
 - †Noriphoca gaudini (Guiscardi, 1870) [''Monotherium gaudini'' Guiscardi, 1870]
- †Devinophocinae Koretsky & Holec, 2002
 - †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 changorum Valenzuela‐Toro et al., 2016
 - †Virginiaphoca Dewaele et al., 2018
 - †Virginiaphoca magurai Dewaele et al., 2018
 - †Properiptychus Ameghino, 1897
 - †Properiptychus argentinus Ameghino, 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
 - †Magnotherium Rahmat et al., 2021
 - †Magnotherium johnsii Rahmat et al., 2021
 - †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
 - †Seronectes Hafed et al., 2024
 - †Seronectes meherrinensis Hafed et al., 2024
 - Monachini E. L. Trouessart, 1897
 - †Pliophoca Tavani, 1941
 - †Pliophoca etrusca Tavani, 1941
 - †Eomonachus Rule et al., 2020
 - †Eomonachus belegaerensis Rule et al., 2020
 - Miroungini Muizon, 1981
 - †Callophoca van Beneden, 1876
 - †Callophoca obscura van Beneden, 1876
 - Lobodontini J.E. Gray, 1869
 - †Sarcodectes Rule et al., 2020
 - †Sarcodectes magnus Rule et al., 2020
 - †Acrophoca Muizon, 1981
 - †Acrophoca longirostris Muizon, 1981
 - †Homiphoca Muizon & Hendey, 1980
 - †Homiphoca capensis (Hendey & Repenning, 1972) [''Prionodelphis capensis'' Hendey & Repenning, 1972]
 - †Homiphoca murfreesi Hafed et al., 2022
 - †Kawas Cozzuol, 2001
 - †Kawas benegasorum Cozzuol, 2001
 - †Piscophoca Muizon, 1981
 - †Piscophoca pacifica Muizon, 1981
 - †Hadrokirus Amson & De Muizon, 2013
 - †Hadrokirus martini Amson & De Muizon, 2013
 - †Hadrokirus novotini Hafed et al., 2022
 - †Magophoca Dewaele & de Muizon, 2024
 - †Magophoca brevirostris Dewaele & de Muizon, 2024
- 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
 - †Pachyphoca volkodavi Tarasenko, 2022
 - 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
 - †Planopusa Rahmat & Koretsky, 2021
 - †Planopusa semenovi Rahmat & Koretsky, 2021
 - †Praepusa Kretzoi, 1941
 - †Praepusa pannonica Kretzoi, 1941
 - †Praepusa boeska Koretsky et al., 2015
 - †Praepusa magyaricus Koretsky, 2003
 - †Praepusa tarchankutica Antoniuk & Koretsky, 1984
 - †Praepusa vindobonensis (Toula, 1897) [''Phoca vindobonensis'' Toula, 1897]
 - Phoca Linnaeus, 1758
 - †Phoca moori Newton, 1890

Otarioidea J.E. Gray, 1821

†Desmatophocidae Hay, 1930
- †Eodesmus Tate-Jones et al., 2020
  - †Eodesmus condoni Tate-Jones et al., 2020
- †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
 - †Pontolis kohnoi Biewer, Velez-Juarbe & Parham, 2020
 - †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
 - †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
 - †Valenictus sheperdi Boessenecker et al., 2024
 - 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
 †Neophoca palatina King, 1983
 - †Hydrarctos Muizon, 1978
 †Hydrarctos lomasiensis Muizon, 1978

Notes and References

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 . 10.1111/j.1365-2907.2011.00193.x.
Valenzuela-Toro . A. . Nicholas Pyenson . 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 . 10.1098/rsos.191394 . 31827869. 6894555 . 2019RSOS....691394V . free .
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 . 7877495 . 10.1093/oxfordjournals.molbev.a040189.
Hunt, R. M. Jr. . Barnes, L. G. . 1994. Basicranial evidence for ursid affinity of the oldest pinnipeds. Proceedings of the San Diego Society of Natural History. 29. 57–67. PDF.
Higdon, J. W. . Bininda-Emonds, O. R. . Beck, R. M. . Ferguson, S. H. . Phylogeny and divergence of the pinnipeds (Carnivora: Mammalia) assessed using a multigene dataset . 10.1186/1471-2148-7-216 . BMC Evolutionary Biology . 7 . 216 . 2007 . 1 . 17996107 . 2245807 . free . 2007BMCEE...7..216H .
Sato, J. J. . Wolsan, M. . Suzuki, H. . Hosoda, T. . Yamaguchi, Y. . Hiyama, K. . Kobayashi, M. . Minami, S. . 10.2108/zsj.23.125 . 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 . 2006 . 16603806. 2115/13508 . 25795496 . free .
Flynn, J. J. . Finarelli, J. A. . Zehr, S. . Hsu, J. . Nedbal, M. A. . Molecular phylogeny of the Carnivora (Mammalia): Assessing the impact of increased sampling on resolving enigmatic relationships . 10.1080/10635150590923326 . Systematic Biology . 54 . 2 . 317–37 . 2005 . 16012099. free .
Berta, A. . Churchill, M. . Boessenecker, R.W. . amp . 2018 . The Origin and Evolutionary Biology of Pinnipeds: Seals, Sea Lions, and Walruses . Annual Review of Earth and Planetary Sciences . 46 . 203–228 . 10.1146/annurev-earth-082517-010009. 2018AREPS..46..203B . free .
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 . 10.3389/fevo.2019.00457. free .
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 . 40 . 5 . e1833896 . 10.1080/02724634.2020.1833896 . 2020JVPal..40E3896B .