Insights on the Dental Evolution of Walruses Based on New Fossil Specimens from California

LITERATURE CITED

• Akiba, F. 1982. Taxonomy and biostratigraphic significance of a new diatom, Thalassionema schraderi. Bacillaria 5:43–61.
   Google Scholar
• Allen, J. A. 1880. History of North American pinnipeds, a monograph of the walruses, sea-lions, sea-bears and seals of North America. U.S. Geological and Geographical Survey of the Territories Miscellaneous Publications 12:1–785.
   Google Scholar
• Barboza, M. M., J. F. Parham, G-P. Santos, B. N. Kussman, and J. Velez-Juarbe. 2017. The age of the Oso Member, Capistrano Formation, and a review of fossil crocodylians from California. PaleoBios 34:1–16.
   Google Scholar
• Barnes, L. G. 1988. A new fossil pinniped (Mammalia: Otariidae) from the middle Miocene Sharktooth Hill Bonebed, California. Natural History Museum of Los Angeles County Contributions in Science 403:1–26.
   Google Scholar
• Barnes, L. G., and R. E. Raschke. 1991. Gomphotaria pugnax, a new genus and species of late Miocene dusignathine otariid pinniped (Mammalia: Carnivora) from California. Natural History Museum of Los Angeles County Contributions in Science 426:1–16.
   Google Scholar
• Berta, A., J. L. Sumich, and K. M. Kovacs. 2005. Marine Mammals: Evolutionary Biology. Academic Press, Burlington, 560 pp.
   Google Scholar
• de Blainville, H. D. 1820. Sur quelques crânes de phoques. Journal de Physique, de Chimie, d'Histoire Naturelle et des Arts 91:286–300.
   Google Scholar
• Blake, G. H. 1991. Review of the Neogene biostratigraphy and stratigraphy of the Los Angeles Basin and implications for basin evolution; pp. 135–184 in K. T. Biddle (ed.), Active Margin Basins. American Association of Petroleum Geologists Memoir 52.
   Google Scholar
• Boessenecker, R. W. 2017. A new early Pliocene record of the toothless walrus Valenictus (Carnivora, Odobenidae) from the Purisima Formation of Northern California. PaleoBios 34:1–6.
   Google Scholar
• Boessenecker, R. W., and M. Churchill. 2013. A reevaluation of the morphology, paleoecology, and phylogenetic relationships of the enigmatic walrus Pelagiarctos. PLoS ONE 8:e54311.
   PubMed Web of Science ®Google Scholar
• Boessenecker, R. W., and M. Churchill. 2015. The oldest known fur seal. Biology Letters 11:20140835.
   PubMed Web of Science ®Google Scholar
• Bowdich, T. E. 1821. An Analysis of the Natural Classification of Mammalia. Smith, Paris, 115 pp.
   Google Scholar
• Brisson, M. J. 1762. Regnum Animale in Classes IX Distributum Sive Synopsis Methodica. Jean-Baptiste Bauche, Paris, 296 pp.
   Google Scholar
• Cantino, P. D., and K. de Queiroz. 2020. International Code of Phylogenetic Nomenclature (PhyloCode). CRC Press, Boca Raton, FL, 189 pp.
   Google Scholar
• Cullen, T. M., D. Fraser, N. Rybczynski, and C. Schröder-Adams. 2014. Early evolution of sexual dimorphism and polygyny in Pinnipedia. Evolution 68:1469–1484.
   PubMed Web of Science ®Google Scholar
• Deméré, T. A. 1994a. Two new species of fossil walruses (Pinnipedia: Odobenidae) from the Upper Pliocene San Diego Formation, California. Proceedings of the San Diego Society of Natural History 29:77–98.
   Google Scholar
• Deméré, T. A. 1994b. The Family Odobenidae: a phylogenetic analysis of fossil and living taxa. Proceedings of the San Diego Society of Natural History 29:99–123.
   Google Scholar
• Deméré, T. A., and A. Berta. 2001. A reevaluation of Proneotherium repenningi from the Miocene Astoria Formation of Oregon and its position as a basal odobenid (Pinnipedia: Mammalia). Journal of Vertebrate Paleontology 21:270–310.
   Web of Science ®Google Scholar
• Domning, D. P. 1994. A phylogenetic analysis of the Sirenia. Proceedings of the San Diego Society of Natural History 29:177–189.
   Google Scholar
• Downs, T. 1955. A fossil sea lion from the Miocene of the San Joaquin Hills, Orange County, California. Bulletin, Southern California Academy of Sciences 54:49–56.
   Google Scholar
• Drehmer, C. J., M. E. Fabián, and J. O. Menegheti. 2004. Dental anomalies in the Atlantic population of South American sea lion, Otaria byronia (Pinnipedia, Otariidae): evolutionary implications and ecological approach. Latin American Journal of Aquatic Mammals 3:7–18.
   Google Scholar
• Drehmer, C. J., D. Sanfelice, and C. Loch. 2015. Dental anomalies in pinnipeds (Carnivora: Otariidae and Phocidae): occurrence and evolutionary implications. Zoomorphology 134:325–338.
   Web of Science ®Google Scholar
• Ehrenberg, C. G. 1837. Gesammtsitzung der Akademie. Monatsberichte der Berlin Königliche Pruessische Akademie Wissenschaften 1837:215–246.
   Google Scholar
• Ehrenberg, C. G. 1839. Über die Bildung der Kreidefelsen und des Kreidemergels durch unsichtbare Organismen. Abhandlungen der Königlichen Akademie der Wissenschaften zu Berlin 1838:59–147.
   Google Scholar
• Ehrenberg, C. G. 1843. Verbreitung und Einfluss des mikroskopischen Lebens in Süd-und Nord-Amerika. Abhandlungen der Königlichen Akademie der Wissenschaften zu Berlin 1841:291–466.
   Google Scholar
• Fay, F. H. 1982. Ecology and biology of the Pacific walrus, Odobenus rosmarus divergens Illiger. North American Fauna 74:1–279.
   Google Scholar
• Garlich-Miller, J., and R. Stewart. 1998. Growth and sexual dimorphism of Atlantic walruses (Odobenus rosmarus rosmarus) in Foxe Basin, Northwest Territories, Canada. Marine Mammal Science. 14:803–818.
   Web of Science ®Google Scholar
• Gloger, C. W. L. 1841 [1841–1842]. Gemeinnütziges Hand- und Hilfsbuch der Naturgeschichte. Für gebildete Leser aller Stände, besonders für die reifere Jugend und ihrer Lehrer. August Schulz und Comp., Breslau [Wrocław], 495 pp.
   Google Scholar
• Horikawa, H. 1995. A primitive odobenine walrus of Early Pliocene age from Japan. Island Arc 3:309–328.
   Google Scholar
• Howell, A. B. 1929. Contributions to the comparative anatomy of the eared and earless seals (Genera Zalophus and Phoca). Proceedings of the U.S. National Museum 73:1–142.
   Google Scholar
• Illiger, C. 1811. Prodromus Systematis Mammalium et Avium. Salfeld, Berlin, 301 pp.
   Google Scholar
• Kastelein, R. A., and M. A. van Gaalen. 1988. The sensitivity of the vibrissae of a Pacific walrus (Odobenus rosmarus divergens). Part 1. Aquatic Mammals 14:123–133.
   Google Scholar
• Kastelein, R. A., S. Stevens, and P. Mosterd. 1990. The tactile sensitivity of the mystacial vibrissae of a Pacific walrus (Odobenus rosmarus divergens). Part 2: masking. Aquatic Mammals 16:78–87.
   Google Scholar
• Kellogg, R. 1921. A new pinniped from the Upper Pliocene of California. Journal of Mammalogy 2:212–226
   Google Scholar
• Kellogg, R. 1927. Fossil pinnipeds from California. Carnegie Institution of Washington Publication 348:27–37.
   Google Scholar
• Kellogg, R. 1931. Pelagic mammals from the Temblor Formation of the Kern River region, California. Proceedings of the California Academy of Sciences 19:217–397.
   Google Scholar
• Kohno, N. 1994. A new Miocene pinniped in the genus Prototaria (Carnivora: Odobenidae) from the Moniwa Formation, Miyagi, Japan. Journal of Vertebrate Paleontology 14:414–426.
   Google Scholar
• Kohno, N. 2006. A new Miocene odobenid (Mammalia: Carnivora) from Hokkaido, Japan, and its implications for odobenid phylogeny. Journal of Vertebrate Paleontology 26:411–421.
   Web of Science ®Google Scholar
• Kohno, N., L. G. Barnes, and K. Hirota. 1995. Miocene fossil pinnipeds of the genera Prototaria and Neotherium (Carnivora; Otariidae; Imagotariinae) in the North Pacific Ocean: evolution, relationships and distribution. Island Arc 3:285–308.
   Google Scholar
• Leidy, J. 1859. Remarks on Dromatherium sylvestre and Ontocetus emmonsi. Proceedings of the Academy of Natural Sciences of Philadelphia 1859:162.
   Google Scholar
• Lesson, R. P. 1828. Histoire Naturelle des Phoques; pp. 400–426 in B. G. M. Bory de Saint Vincent (ed.), Dictionnaire Classique d’Histoire Naturelle. Vol. 13. Rey et Gravier, Paris.
   Google Scholar
• Lindenfors, P., B.S. Tullberg, and M. Biuw. 2002. Phylogenetic analyses of sexual selection and sexual size dimorphism in pinnipeds. Behavioral Ecology and Sociobiology 52:188–193.
   Web of Science ®Google Scholar
• Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio Decima 1:1–824
   Google Scholar
• Lyon, G. M. 1941. A Miocene sea lion from Lomita, California. University of California Publications in Zoology 47(2):22–41.
   Google Scholar
• Magallanes, I., J. F. Parham, G. Santos, and J. Velez-Juarbe. 2018. A new tuskless walrus from the Miocene of Orange County, California, with comments on the diversity and taxonomy of odobenids. PeerJ 6:e5708.
   PubMed Web of Science ®Google Scholar
• Mitchell, E. D., Jr. 1961. A new walrus from the Imperial Pliocene of Southern California: with notes on odobenid and otariid humeri. Los Angeles County Museum Contributions in Science 44:1–28.
   Google Scholar
• Mitchell, E. D. 1968. The Mio-Pliocene pinniped Imagotaria. Journal of the Fisheries Research Board of Canada 25:1843–1900.
   Web of Science ®Google Scholar
• Muchlinski, M. N. 2008. The relationship between the infraorbital foramen, infraorbital nerve, and maxillary mechanoreception: implications for interpreting the paleoecology of fossil mammals based on infraorbital foramen size. Anatomical Record 291:1221–1226.
   Google Scholar
• O'Reilly, J. E., M. N. Puttic, L. Parry, A. R. Tanner, J. E. Tarver, J. Fleming, D. Pisani, and P. C. J. Donoghue. 2016. Bayesian methods outperform parsimony but at the expense of precision in the estimation of phylogeny from discrete morphological data. Biology Letters 12:20160081.
   PubMed Web of Science ®Google Scholar
• Parham, J. F., and N. D. Pyenson. 2010. New sea turtle from the Miocene of Peru and the iterative evolution of feeding ecomorphologies since the Cretaceous. Journal of Paleontology 84:231–247.
   Web of Science ®Google Scholar
• Paterson, R. S., N. Rybczynski, N. Kohno, and H. C. Maddin. 2020. A total evidence phylogenetic analysis of pinniped phylogeny and the possibility of parallel evolution within a monophyletic framework. Frontiers in Ecology and Evolution 7:457.
   Web of Science ®Google Scholar
• Péron, F. 1816. Voyage de découvertes aux terres australes, exécuté sur les Corvettes le Géographe, le Naturaliste, et al Goëlette le Casuarina, pendent les années 1800, 1801, 1802, 1803 et 1804. L’Imprimerie Royale, Paris 2:471
   Google Scholar
• Powell, C. L., II, R. W. Boessenecker, N. A. Smith, R. J. Fleck, S. J. Carlson, J. R. Allen, D. J. Long, A. M. Sarna-Wojcicki, and R. B. Guruswami-Naidu. 2019. Geology and paleontology of the late Miocene Wilson Grove Formation at Bloomfield Quarry, Sonoma County, California: U.S. Geological Survey Scientific Investigations Report 5021:1–77.
   Google Scholar
• Rambaut, A, A. J., Drummond, D. Xie, G. Baele, and M. A. Suchard. 2018. Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67:901–904.
   PubMed Web of Science ®Google Scholar
• Reep, R. L., C. D. Marshall, M. L. Stoll, and D. M. Whitaker. 1998. Distribution and innervation of facial bristles and hairs in the Florida manatee (Trichechus manatus latirostris). Marine Mammal Science 14:257–273.
   Web of Science ®Google Scholar
• Reep, R. L., M. L. Stoll, C. D. Marshall, B. L. Homer, and D. A. Samuelson. 2001. Microanatomy of facial vibrissae in the Florida manatee: the basis for specialized sensory function and oripulation. Brain, Behaviour and Evolution 58:1–14.
   PubMed Web of Science ®Google Scholar
• Repenning, C. A., and R. H. Tedford. 1977. Otarioid seals of the Neogene. U.S. Geological Survey Professional Paper 992:1–93.
   Google Scholar
• Ronquist, F., M. Teslenko, P. Van Der Mark, D. L. Ayres, A. Darling, S. Höhna, B. Larget, L. Liu, M. A. Suchard, and J. P. Huelsenbeck. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61:539–542.
   PubMed Web of Science ®Google Scholar
• Schulz, P., 1928. Beitrage zur Kenntnis fossiler und rezenter Silicoflagellaten. Botanischen Archiv 21:225–292.
   Google Scholar
• Swofford, D. L. 2002. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sunderland: Sinauer Associates.
   Google Scholar
• Takeyama, K., and T. Ozawa. 1984. A new Miocene otarioid from Japan. Proceedings of the Japan Academy, Series B 60:36–39.
   Google Scholar
• Tanaka, Y., and N. Kohno. 2015. A new Late Miocene odobenid (Mammalia: Carnivora) from Hokkaido, Japan suggests rapid diversification of basal Miocene odobenids. PLoS ONE 10:e0141406.
   PubMed Web of Science ®Google Scholar
• Tomida, 1989 is missing
   Google Scholar
• True, F. W. 1905a. Diagnosis of a new genus and species of fossil sea-lion from the Miocene of Oregon. Smithsonian Miscellaneous Collections 48:47–49.
   Google Scholar
• True, F. W. 1905b. New name for Pontoleon. Proceedings of the Biological Society of Washington 17:253–253.
   Google Scholar
• Velez-Juarbe, J. 2017. Eotaria citrica, sp. nov., a new stem otariid from the “Topanga” Formation of Southern California. PeerJ 5:e3022.
   PubMed Web of Science ®Google Scholar
• Velez-Juarbe, J. 2018. New data on the early odobenid Neotherium mirum Kellogg, 1931, and other pinniped remains from the Sharktooth Hill Bonebed, California. Journal of Vertebrate Paleontology 38:e1481080.
   Web of Science ®Google Scholar
• Velez-Juarbe, J., and F. M. Salinas-Márquez. 2018. A dwarf walrus from the Miocene of Baja California Sur, Mexico. Royal Society Open Science 5:180423.
   PubMed Web of Science ®Google Scholar
• Velez-Juarbe, J., and A. M. Valenzuela-Toro. 2019. Oldest record of monk seals from the North Pacific and biogeographic implications. Biology letters 15:20190108.
   PubMed Web of Science ®Google Scholar
• Woodring, W. P., M. N. Bramlette, and W. S. W. Kew. 1946. Geology and paleontology of Palos Verdes Hills, California. U.S. Geological Survey Professional Paper 207, 145 pp.
   Google Scholar
• Yanagisawa, Y., and F. Akiba. 1990. Taxonomy and phylogeny of the three marine diatom genera Crucidenticula, Denticulopsis and Neodenticula. Bulletin of the Geological Survey of Japan 41:197–301.
   Google Scholar