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

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.     

Summary of the fossil record of pinnipeds of Japan, and comparisons with that from the eastern North Pacific

Abstract The fossil pinniped record of the North Pacific Ocean includes both Phocidae and Otariidae ( sensu lato ), extends from the Late Oligocene to the Late Pleistocene, is taxonomically diverse, and is constantly becoming more complete owing to additional important discoveries. The earliest and most diverse fossil pinnipeds in the North Pacific are otariids, the phocids not appearing until the latest Pliocene. The theoretical center of otariid pinniped evolutionary history has been considered by some to be in the eastern North Pacific. New materials from the western North Pacific, however, including representatives of the subfamilies Enaliarctinae, Imagotariinae, Odobeninae and Otariinae, indicate that pinniped evolutionary patterns were basin-wide phenomena, and that a more complete record undoubtedly would reveal numerous trans-Pacific distributions. This would be expected considering the distributions of living species. The paucity of fossil Phocidae and their absence from pre-Pliocene deposits are consistent with theories that the family primarily evolved outside the North Pacific.     

A new species of Middle Miocene sperm whale of the genus Scaldicetus (Cetacea; Physeteridae) from Shiga-mura, Japan

Abstract A nearly complete skeleton of a fossil sperm whale from the Middle Miocene age Bessho Formation at Shiga-mura, Nagano Prefecture, the most complete fossil physeterid skeleton found in Japan, is here named Scaldicetus shigensis, new species. Its abruptly tapered rostrum, deep supracranial basin, and high occipital crest are typical of physeterids. Large teeth with crenulate enamel on conical crowns, present in both the palate and mandible, are consistent with the genus Scaldicetus du Bus, 1867, a genus originally based on fossils discovered in the Antwerp Basin, Belgium. Scaldicetus shigensis is relatively primitive, having a relatively long, slender rostrum, large zygomatic arches, large tympanic bullae, prominent occipital condyles, low tooth count and a sloping occipital shield that is deeply emarginated laterally by large temporal fossae. The probable mandibular tooth count is 12, only one more than the primitive eutherian mammalian dentition. Despite its abundant primitive characters, Scaldicetus shigensis has very asymmetrical external nares, comparable to the living sperm whale, Physeter catodon, and is in this way more derived than most of the contemporaneous fossil physeterids. At a time when other sperm whales were more highly evolved, Scaldicetus shigensis retained a primitive occipital shield, large tympanic bullae, and low tooth count, as in primitive Oligocene odontocetes such as Agorophiidae. It was a relict form in Middle Miocene time, and provides an indication of a primitive stage of sperm whale evolution heretofore undocumented. Fossils from elsewhere previously referred to as Scaldicetus, largely on the basis of tooth characters, range in age from Early Miocene to Plio-Pleistocene time. Cranial material that would clarify relationships is lacking for most of these, and this time range seems too long for a cetacean genus. Because of this simplistic approach to identifications, Scaldicetus is probably a grade taxon, not a natural biological grouping. However, the tooth morphology of Scaldicetus shigensis is correct for the genus Scaldicetus, so we provisionally assign it to this genus. Redefinition of the genus Scaldicetus and revision of the Physeteridae are beyond the scope of the present study.     

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

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.