Titanotaria Temporal range: Miocene | |
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Holotype skull of Titanotaria orangensis | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Carnivora |
Clade: | Pinnipedia |
Family: | Odobenidae |
Genus: | † Titanotaria Magallanes et al., 2018 |
Species: | †T. orangensis |
Binomial name | |
†Titanotaria orangensis Magallanes et al., 2018 | |
Titanotaria is a genus of late, basal walrus from the Miocene of Orange County, California. Unlike much later odobenids, it lacked tusks. Titanotaria is known from an almost complete specimen which serves as the holotype for the only recognized species, Titanotaria orangensis, it is the best preserved fossil walrus currently known.
Although the holotype specimen (OCPC 11141) of Titanotaria had been discovered in 1993 and represents one of the most complete fossil walrus known, little attention was given to the material for over 20 years. The first mention of the fossils in peer-reviewed literature came in 2017 with Barboza and colleagues [1] publishing a faunal list of the Oso Member of the Capistrano Formation, where Titanotaria had been found. Specifically, the fossilized bones were collected from the town of Lake Forest, Orange County, California, during the construction of the Saddleback Church. [2] A full description followed a year after its mention by Barboza and was led by Isaac Magallanes, who published a detail examination of the fossils alongside a phylogenetic analysis. [3] According to paleontologist Robert Boessenecker, the remains were unofficially known by the name "Waldo". [4]
The name Titanotaria honors the California State University, Fullerton, widely known as the Titans. This was meant to recognize the collaboration between the university and Orange County, which lead to the creation of the John D. Cooper Archaeological and Paleontological Center. The second part of the genus name, otaria, is a reference to the genus Otaria and a commonly used suffix for fossil pinnipeds. The species name means "coming from Orange County". [3]
The holotype skull of Titanotaria belongs to a male individual with an asymmetric skull, likely caused by a healed pathology. The rostrum of Titanotaria is elongated and widens at around the root of the first canine tooth. The premaxilla are triangular in outline and elevated slightly above the tooth row. The front-most tip of the premaxilla is marked by a knob-shaped prenarial process, which is immediately followed by a depression located above the incisors and canines that likely serves as an origin for the lateral nasalis muscle. The nasal bones are long (60% of the rostrum length) with parallel edges and a broad, V-shaped suture with the frontal bone. The zygomatic arch is broad and possesses an oval prominence on its ventral surface. The point of articulation between the jugal and the maxilla is largely fused and a small, triangular postorbital process is present on the jugal element of the zygomatic arch. The frontal bone is widest towards the front of the skull and bears two temporal crests, which fuse to form the sagittal crest. The crest is prominent and long, with a sinuous profile. This differs from the more sloping sagittal crests of other odobenids like Imagotaria and Neotherium . Towards the back of the skull the sagitall crest meets the nuchal crests, which is wide and obscures the occipital region in top view. [3]
The tooth formula of Titanotaria is . In the upper jaw the incisors are long and slender with an oval crosssection and a single root. The canines are robust, conical and larger than the incisors. While the first premolar likely only possesses a single root based on the morphology of the alveolous, the second is bi-lobed with a bulbous tooth crown. The following teeth also show two tooth roots and there is a decrease in size between the two molars. No incisors are preserved in the lower jaw and their alveoli are obscured by sediment. The mandibular tooth row is very short, only taking up 40% of the mandible. The lower canines are almost as large as their upper counterparts and like them they are robust and conical with a curve to them. Like in the upper jaw, the teeth starting with the second premolar of the mandible are double rooted with bulbous crowns. The last lower molar however appears to have been single rooted based on the anatomy of its tooth socket. [3]
Titanotaria preserves most of its postcranial material; however, only elements relevant to phylogenetic analysis were described. The holotype is only missing few ribs, parts of the right forelimb, most of the pelvis and some of the distal limb elements. [3] It reached a length of 10 ft (3.0 m) and weighed around 1,200 lb (540 kg). [2]
Phylogenetic analysis found that Titanotaria was a basal odobenid, nesting outside of the clade Neodobenia (named within the same publication as the genus). [3] The same placement was later recovered by Biewer and colleagues when they described Osodobenus . [5]
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Titanotaria is known from the Oso Member of the Capistrano Formation, which preserves a rich assemblage of fossil walrus species such as Gomphotaria pugnax , Pontolis magnus , Pontolis kohnoi and Osodobenus eodon . The eared seal Thalassoleon was also found in this formation, alongside giant sea cows, cetotheriid whales, the bizarre Desmostylus , various sharks and the remains of indetermined crocodiles. [1] [5] [3]
Odobenidae is a family of pinnipeds. The only extant species is the walrus. In the past, however, the group was much more diverse, and includes more than a dozen fossil genera.
Catopsbaatar is a genus of multituberculate, an extinct order of rodent-like mammals. It lived in what is now Mongolia during the late Campanian age of the Late Cretaceous epoch, about 72 million years ago. The first fossils were collected in the early 1970s, and the animal was named as a new species of the genus Djadochtatherium in 1974, D. catopsaloides. The specific name refers to the animal's similarity to the genus Catopsalis. The species was moved to the genus Catopsalis in 1979, and received its own genus in 1994. Five skulls, one molar, and one skeleton with a skull are known; the last is the genus' most complete specimen. Catopsbaatar was a member of the family Djadochtatheriidae.
Probainognathus meaning “progressive jaw” is an extinct genus of cynodonts that lived around 235 to 221.5 million years ago, during the Late Triassic in what is now Argentina. Together with the genus Bonacynodon from Brazil, Probainognathus forms the family Probainognathidae. Probainognathus was a relatively small, carnivorous or insectivorous cynodont. Like all cynodonts, it was a relative of mammals, and it possessed several mammal-like features. Like some other cynodonts, Probainognathus had a double jaw joint, which not only included the quadrate and articular bones like in more basal synapsids, but also the squamosal and surangular bones. A joint between the dentary and squamosal bones, as seen in modern mammals, was however absent in Probainognathus.
Thrinaxodon is an extinct genus of cynodonts, most commonly regarded by its species T. liorhinus which lived in what are now South Africa and Antarctica during the Early Triassic. Thrinaxodon lived just after the Permian–Triassic mass extinction event, its survival during the extinction may have been due to its burrowing habits.
Adamantinasuchus is an extinct genus of notosuchian crocodylomorph from and named after the Late Cretaceous Adamantina Formation of Brazil. It is known from only one fossil, holotype UFRJ-DG 107-R, collected by William Nava. The fossil consists of a partial skull, fragmentary limb bones and a few broken vertebrae, and was found 25 km SW of the town of Marilia, near a reservoir dam. Adamantinasuchus was approximately 60 cm long from nose to tail, and would have only weighed a few kilograms.
Cynosaurus is an extinct genus of cynodonts. Remains have been found from the Dicynodon Assemblage Zone in South Africa. Cynosaurus was first described by Richard Owen in 1876 as Cynosuchus suppostus. Cynosaurus has been found in the late Permian period. Cyno- is derived from the Greek word kyon for dog and –sauros in Greek meaning lizard.
Scymnosaurus is an extinct genus of therocephalian therapsids, first described by Robert Broom in 1903. There are three species that still take the name Scymnosaurus, S. ferox, S. watsoni and S. major, with a fourth, S. warreni, now identified as Moschorhinus warreni. Each of these have now been reclassified into Lycosuchidae incertae sedis.
Eritherium is an extinct genus of early Proboscidea found in the Ouled Abdoun basin, Morocco. It lived about 60 million years ago. It was first named by Emmanuel Gheerbrant in 2009 and the type species is Eritherium azzouzorum. Eritherium is the oldest, smallest and most primitive known elephant relative.
Karenites is an extinct genus of therocephalian therapsids from the Late Permian of Russia. The only species is Karenites ornamentatus, named in 1995. Several fossil specimens are known from the town of Kotelnich in Kirov Oblast.
The Capistrano Formation is a geologic formation in coastal southern Orange County, California. It preserves fossils dating back to the late Miocene to early Pliocene, with the Oso Member representing a near-shore environment. Fifty-nine species and varieties of foraminifera are recognized from the Capistrano Formation alongside a diverse array of marine mammals including up to five species of walrus.
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 of North America along what is now the western coasts of California and Oregon between 11.608 and 5.332 million years ago, during the Miocene and Pliocene.
Lentiarenium was an early sea cow from the Late Oligocene (Chattian) Linz-Melk Formation of Austria. Known since the mid 19th century, Lentiarenium was long considered to be a species of Halitherium until a 2016 analysis showed it to be distinct.
Abdalodon is an extinct genus of late Permian cynodonts, known by its only species A. diastematicus.Abdalodon together with the genus Charassognathus, form the clade Charassognathidae. This clade represents the earliest known cynodonts, and is the first known radiation of Permian cynodonts.
Archaeodobenus is an extinct genus of pinniped that lived during the Late Miocene of what is now Japan. It belonged to the Odobenidae family, which is today only represented by the walrus, but was much more diverse in the past, containing at least 16 genera.
Nanokogia is an extinct genus of pygmy sperm whale that lived off the coast of Panama during the Late Miocene.
Ankylorhiza is an extinct genus of toothed whale that lived in what is now the United States during the Oligocene epoch, between 29 and 23.5 million years ago. The type and only known species is A. tiedemani, though two fossil skeletons may represent an additional, second species within the genus. Ankylorhiza was about 4.8 meters (16 ft) long, with a long, robust skull bearing conical teeth that were angled forwards at the tip of the snout.
Pseudotherium is an extinct genus of prozostrodontian cynodonts from the Late Triassic of Argentina. It contains one species, P. argentinus, which was first described in 2019 from remains found in the La Peña Member of the Ischigualasto Formation in the Ischigualasto-Villa Unión Basin.
Vetusodon is an extinct genus of cynodonts belonging to the clade Epicynodontia. It contains one species, Vetusodon elikhulu, which is known from four specimens found in the Late Permian Daptocephalus Assemblage Zone of South Africa. With a skull length of about 18 centimetres (7.1 in), Vetusodon is the largest known cynodont from the Permian. Through convergent evolution, it possessed several unusual features reminiscent of the contemporary therocephalian Moschorhinus, including broad, robust jaws, large incisors and canines, and small, single-cusped postcanine teeth.
Osodobenus is an extinct genus of walrus from the Miocene to Pliocene of California. 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.