Miocene fossil pinnipeds of the genera Prototaria and Neotherium (Carnivora; Otariidae; Imagotariinae) in the North Pacific Ocean: Evolution, relationships and distribution

Abstract

Abstract Fossil otariid pinnipeds of the extinct genera Prototaria Takeyama and Ozawa, 1984, and Neotherium Kellogg, 1931, known from Middle Miocene deposits bordering the North Pacific Ocean, are small, primitive pinnipeds in the subfamily Imagotariinae. They have a small supraorbital process of the frontal or have lost it entirely, a three‐rooted first molar, small paroccipital process, and ear morphology indicating that they belong in the subfamily Imagotariinae. Their unique derived characters include extreme intertemporal constriction and highly modified cheek teeth, the premolars having become molarized by the addition of protocones and lingual cingula. Prototaria Takeyama and Ozawa, 1984, the most primitive known imagotariine genus, contains two species, P. primigena Takeyama and Ozawa, 1984, and P. planicephala Kohno, 1994, both of early Middle Miocene age from Japan. Prototaria has a few derived characters, including a large antorbital process, narrow intertemporal region, and large orbit, but its primitive characters apparently were inherited from enaliarctine ancestors. The long enigmatic Neotherium mirum Kellogg, 1931, of Middle Miocene age from California, USA, is related to Prototaria, but differs by having an elongate skull, very slender zygomatic arch, ventrally exposed median lacerate foramen, and smaller but more molarized premolars. A more primitive new genus and species, Proneotherium repenningi Barnes, related to N. mirum, is from the early Middle Miocene Astoria Formation, coastal Oregon, USA. It shares some derived characters with Prototaria, and shares many other important derived characters with N. mirum. Imagotariines probably arose from some species of Early Miocene enaliarctines, became diverse in Middle and Late Miocene time, and are only known from the North Pacific realm. Although they might include the ancestors of true walruses of the subfamily Odobeninae, no known imagotariines appear to have been adapted for mollusk feeding as are the highly evolved modern walruses. Instead, imagotariines appear to have retained a primitive piscivorous diet, as did the fur seals and sea lions of the subfamily Otariinae.