Osodobenus
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''Osodobenus'' is an extinct genus of
walrus The walrus (''Odobenus rosmarus'') is a large pinniped marine mammal with discontinuous distribution about the North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere. It is the only extant species in the family Odobeni ...
from the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
to
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
. ''Osodobenus'' may have been the first tusked walrus and shows several adaptations that suggest it was a suction feeder, possibly even a benthic feeder like modern species. Three skulls are known showing pronounced sexual dimorphism, with the female lacking the same tusks as the male. Only a single species, ''Osodobenus eodon'', is currently recognized.


Discovery and naming

''Osodobenus'' is known from three specimens including an adult male, adult female and a juvenile specimen preserving skulls and some postcranial material. All the material was collected from the Late
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
to Early
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Messinian The Messinian is in the geologic timescale the last age or uppermost stage of the Miocene. It spans the time between 7.246 ± 0.005 Ma and 5.333 ± 0.005 Ma (million years ago). It follows the Tortonian and is followed by the Zanclean, the fir ...
to
Zanclean The Zanclean is the lowest stage or earliest age on the geologic time scale of the Pliocene. It spans the time between 5.332 ± 0.005 Ma (million years ago) and 3.6 ± 0.005 Ma. It is preceded by the Messinian Age of the Miocene Epoch, and f ...
) Capistrano Formation,
Orange County, California Orange County (officially the County of Orange; often initialized O.C.) is a county (United States), county located in the Los Angeles metropolitan area in Southern California, United States. As of the 2020 United States census, the population ...
alongside the remains of several other early odobenids. In 2020, Biewer and colleagues published a detailed description of the material, establishing ''Osodobenus'' as a new genus while also erecting two new species of ''
Pontolis ''Pontolis'' is an extinct genus of large walrus. It contained three species, ''P. magnus'', ''P. barroni'', and ''P. kohnoi''. Like all pinnipeds, ''Pontolis'' was a heavily built amphibious carnivore. ''Pontolis'' lived along the Pacific coast ...
''. The genus name of ''Osodobenus'' is a combination of the genus name for the modern
walrus The walrus (''Odobenus rosmarus'') is a large pinniped marine mammal with discontinuous distribution about the North Pole in the Arctic Ocean and subarctic seas of the Northern Hemisphere. It is the only extant species in the family Odobeni ...
"''Odobenus''" and the Oso Member, a Messinian strata within the Capistrano Formation. The
species name In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
''eodon'' translates to "dawn tooth" or "dawn tusk", a reference to this animal's key role in the evolution of walrus tusks.


Description

Compared to other basal odobenids, the rostrum of ''Osodobenus'' is short and robust, bulging notably around the canines and with a forward-projecting
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
. The nares are large with thick borders and roofed by the
nasal bones The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
. The
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
is arched and a pair of
infraorbital foramen In human anatomy, the infraorbital foramen is one of two small holes in the skull's upper jawbone ( maxillary bone), located below the eye socket and to the left and right of the nose. Both holes are used for blood vessels and nerves. In anatomic ...
enlarged. The later suggest that much like the modern species, ''Osodobenus'' had a mustache-like collection of whiskeres covering the upper lip. Behind the canines, the teeth are single-rooted with inflated bulbous crowns showing little wear. The first two
incisors Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
of ''Osodobenus'' are transversely compressed with minor wear of the apices. The third incisor is notably longer and slender and there is no cingulum on the inner side of the jaws as in other odobenids. The prominent canines are robust, conical and recurved. Compared to the width of the skull, these canines are approximately 50% larger than those of other basal walrus genera. The pulp cavity is open which indicates that the canines were continuously growing, something only known from tusked odobenids. This however only applies to the males, while the female has smaller canines more proportional to the skull width. This indicates that the presence of tusks in ''Osodobenus'' was
sexually dimorphic Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
. The teeth of ''Osodobenus'' show a somewhat transitional form between the less specialised odobenids with a tooth specialisation score of 0.4 and lower and the more derived and specialised members of the group (score >0.5). This indicates that odobenids rapidly specialised during the late
Tortonian The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago). It follows the Serravallian and is followed by the Messinian. The Tort ...
to early
Messinian The Messinian is in the geologic timescale the last age or uppermost stage of the Miocene. It spans the time between 7.246 ± 0.005 Ma and 5.333 ± 0.005 Ma (million years ago). It follows the Tortonian and is followed by the Zanclean, the fir ...
of the Miocene, which included the development of tusks in later taxa. This again corresponds to what is observed in ''Osodobenus'', whose canines form an intermediate form between what has previously been considered tusked and tuskless. Biewer and colleagues argue that the continuous nature of tusk growth in an evolutionary context makes a rigid separation futile and that ''Osodobenus'' represents the first walrus with canines that could be considered tusks.


Phylogeny

''Osodobenus'' was recovered as a sister taxon to the Neodobenia in both parsimony phylogenetic analysis and Bayesian analyses, which makes it the basalmost walrus with canines that can be considered tusks. The following phylogenetic tree was recovered by the study of Biewer ''et al.'' (2020).


Paleobiology

Although the presence of tusks is sexually dimorphic in ''Osodobenus'', it is suggested that feeding likewise played a part in the evolution of these enlarged canine teeth. ''Osodobenus'' and other tusked odobenids share an enlarged
infraorbital foramen In human anatomy, the infraorbital foramen is one of two small holes in the skull's upper jawbone ( maxillary bone), located below the eye socket and to the left and right of the nose. Both holes are used for blood vessels and nerves. In anatomic ...
, which correspond with the amount of facial nerves that connect to musculature and
Vibrissae Whiskers, also known as vibrissae (; vibrissa; ) are a type of stiff, functional hair used by most therian mammals to sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are coarser as t ...
(whiskers), the later being used by extant seals and
sirenians The Sirenia (), commonly referred to as sea cows or sirenians, are an order (biology), order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The extant Sirenia comprise tw ...
in foraging. Another trait that suggests that ''Osodobenus'' may habe been a
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
feeder is the anatomy of the palate, which indicates that this genus much like extant walrus was a specialised suction feeder. Biewer and his colleagues note that these traits alone are not enough to fully establish ''Osodobenus'' as a benthic feeder, but argue that its feeding behavior may have been similar regardless. As some more derived odobenids lack these adaptations, it is possible that this ecology was evolved independently from crown walrus. Fossils of ''Osodobenus'' were discovered in the Californian Capistrano Formation, which also yielded the fossils of the walrus ''Pontolis magnus'', ''Pontolis kohnoi'' and ''Titanotaria orangensis''. During the Miocene to Pliocene these waters were relatively warm compared to the habitat of the modern walrus and housed a great diversity of mammals including
pinnipeds Pinnipeds (pronounced ), commonly known as seals, are a widely distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammals. They comprise the extant families Odobenidae (whose only living member is the walr ...
,
desmostylians The Desmostylia (from Ancient Greek, Greek δεσμά ''desma'', "bundle", and στῦλος ''stylos'', "pillar") are an extinct order (biology), order of aquatic mammals native to the North Pacific from the early Oligocene (Rupelian) to the lat ...
,
cetaceans Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
and
sirenians The Sirenia (), commonly referred to as sea cows or sirenians, are an order (biology), order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The extant Sirenia comprise tw ...
like '' Hydrodamalis cuestae''. Crocodylians and sabertoothed salmons are also known to have inhabited the waters of the Capistrano Formation.


References

{{Taxonbar, from=Q111620850 Miocene pinnipeds Mammals described in 2020 Fossil taxa described in 2020 Prehistoric pinnipeds of North America Odobenids Monotypic prehistoric carnivoran genera