Miocene pinnipeds of the otariid subfamily Allodesminae in the North Pacific Ocean: Systematics and relationships

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Abstract
Fossil pinnipeds in the extinct otariid subfamily Allodesminae are large, relatively highly evolved marine carnivores that became abundant and diverse in Middle Miocene time and were restricted to the North Pacific Ocean. Their record extends from early Middle Miocene through Late Miocene, with records from California, Oregon, Washington, Baja California and Japan. Allodesmines are characterized by extreme sexual dimorphism, a large orbit, retracted orbital margin of the zygomatic arch, a deeply mortised jugal‐squamosal junction, wide palate, bulbous cheek tooth crowns, nearly flat tympanic bulla with wrinkled ventral surface, a large tympanohyal fossa, large ear ossicles and deep mandible.Eleven allodesmine species are known (eight of which are named), in at least four genera, and most belong to the typical genus Allodesmus Kellogg, 1922. The earliest and most generalized allodesmine known is from the early Middle Miocene (ca 16 Ma) Astoria Formation in coastal Oregon. The last known records are from Late Miocene rocks (ca 10 Ma) in California and Washington. New taxa proposed here are: the genus Brachyallodesmus Barnes and Hirota, to contain Allodesmus packardi Barnes, 1972; the genus Megagomphos Hirota and Barnes, to contain Allodesmus sinanoensis (Nagao, 1941); the species Allodesmus sadoensis Hirota, (Middle Miocene, Japan); the species Allodesmus megallos Hirota (Middle Miocene, Japan); and the species Allodesmus gracilis Barnes (Middle Miocene, California). Additionally, the genus Atopotarus Downs, 1956, and the species Allodesmus kelloggi Mitchell, 1966, are resurrected.Allodesmines were apparently a rapidly evolving group, and most appear to have been adapted to roles later filled by otariine, dusignathine and imagotariine otariids, and the Phocidae (true seals). They became extinct in Late Miocene time and left no living descendants. Although some of their characters evolved convergently with various living species of the pinniped family Phocidae, Allodesminae are an otariid group and not part of the evolutionary history of Phocidae.