Tapinocephalus Temporal range: Guadalupian epoch, Capitanian stage | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Dinocephalia |
Family: | † Tapinocephalidae |
Subtribe: | † Tapinocephalina |
Genus: | † Tapinocephalus Owen, 1876 |
Species: | †T. atherstonei |
Binomial name | |
†Tapinocephalus atherstonei Owen, 1876 | |
Synonyms | |
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Tapinocephalus ("low, depressed head") is an extinct genus of large herbivorous dinocephalians that lived during the Middle Permian Period in what is now South Africa. Only the type species, Tapinocephalus atherstonei is now considered valid for this genus.
Fossils of Tapinocephalus atherstonii were collected and donated to the British Museum by William Guybon Atherstone. They were described by Richard Owen, who described and named the species in 1876. He initially considered it a close relative of Pareiasaurus and classified both as members of Dinosauria. Based on the only remains of the skull known at the time—a poorly-preserved partial snout—he believed it had a low, broad skull similar to labyrinthodonts. Owen accordingly named it Tapinocephalus, from Greek ταπεινός "low, depressed" and κεφαλή "head". [1]
These stocky, barrel-bodied animals were characterised by a massive bony skull roof and short weak snout. It is thought that, like other tapinocephalians, they engaged in intraspecific head-butting, possibly to compete for territory or mates. In life, they were over 3 metres (10 ft) in length and massed around 1.5 to 2 tonnes (3,300 to 4,400 lb), making them among the largest animals of their time.
Tapinocephalus atherstonei is known from a number of skulls and postcranial bones. The skull is large with a heavily pachyostotic skull roof, a massive bony frontals and a short weak Moschops -like snout.
The fossil remains (skull and postcranial elements) of Tapinocephalus are known from the Lower, Middle, and Upper part of the Tapinocephalus Assemblage Zone (Capitanian age) of the Lower Beaufort Beds of the South African Karoo. [2]
Taurops is considered as a synonym of Tapinocephalus. [2] Phocosaurus megischion is another synonym differing only in that the transition from the frontals to the snout is not abrupt.
Tapinocephalus fossils are found in the Diictodon–Styracocephalus subzone of the Tapinocephalus Assemblage Zone, of which it is the namesake. It is not particularly common. It was one of several tapinocephalids in the ecosystem. [3]
Dinocephalians are a clade of large-bodied early therapsids that flourished in the Early and Middle Permian between 279.5 and 260 million years ago (Ma), but became extinct during the Capitanian mass extinction event. Dinocephalians included herbivorous, carnivorous, and omnivorous forms. Many species had thickened skulls with many knobs and bony projections. Dinocephalians were the first non-mammalian therapsids to be scientifically described and their fossils are known from Russia, China, Brazil, South Africa, Zimbabwe, and Tanzania.
Moschops is an extinct genus of therapsids that lived in the Guadalupian epoch, around 265–260 million years ago. They were heavily built plant eaters, and they may have lived partly in water, as hippopotamuses do. They had short, thick heads and might have competed by head-butting each other. Their elbow joints allowed them to walk with a more mammal-like gait rather than crawling. Their remains were found in the Karoo region of South Africa, belonging to the Tapinocephalus Assemblage Zone. Therapsids, such as Moschops, are synapsids, the dominant land animals in the Permian period, which ended 252 million years ago.
Gorgonops is an extinct genus of gorgonopsian therapsid, of which it is the type genus. Gorgonops lived during the Late Permian (Wuchiapingian), about 260–254 million years ago in what is now South Africa.
Jonkeria is an extinct genus of dinocephalians. Jonkeria was a large and omnivorous animal, from the Tapinocephalus Assemblage Zone, Lower Beaufort Group, of the South African Karoo.
The Beaufort Group is the third of the main subdivisions of the Karoo Supergroup in South Africa. It is composed of a lower Adelaide Subgroup and an upper Tarkastad Subgroup. It follows conformably after the Ecca Group and unconformably underlies the Stormberg Group. Based on stratigraphic position, lithostratigraphic and biostratigraphic correlations, palynological analyses, and other means of geological dating, the Beaufort Group rocks are considered to range between Middle Permian (Wordian) to Early Triassic (Anisian) in age.
Anteosaurus is an extinct genus of large carnivorous dinocephalian synapsid. It lived at the end of the Guadalupian during the Capitanian stage, about 265 to 260 million years ago in what is now South Africa. It is mainly known by cranial remains and few postcranial bones. Measuring 5–6 m (16–20 ft) long and weighing about 600 kg (1,300 lb), Anteosaurus was the largest known carnivorous non-mammalian synapsid and the largest terrestrial predator of the Permian period. Occupying the top of the food chain in the Middle Permian, its skull, jaws and teeth show adaptations to capture large prey like the giants titanosuchids and tapinocephalids dinocephalians and large pareiasaurs.
The Tapinocephalus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the middle Abrahamskraal Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 2,000 metres (6,600 ft), occur from Merweville and Leeu-Gamka in its southernmost exposures, from Sutherland through to Beaufort West where outcrops start to only be found in the south-east, north of Oudshoorn and Willowmore, reaching up to areas south of Graaff-Reinet. Its northernmost exposures occur around the towns Fraserburg and Victoria West. The Tapinocephalus Assemblage Zone is the second biozone of the Beaufort Group.
The Pristerognathus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the upper Abrahamskraal Formation and lowermost Teekloof Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching not more than 300 metres (980 ft), occur just east of Sutherland through to Beaufort West in the south and Victoria West in the north. Exposures are also found west of Colesberg and south of Graaff-Reinet. The Pristerognathus Assemblage Zone is the third biozone of the Beaufort Group.
The Eodicynodon Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the Abrahamskraal Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 620 metres (2,030 ft), occur south-east of Sutherland, north of Prince Albert, and south-east of Beaufort West. The Eodicynodon Assemblage Zone is the lowermost biozone of the Beaufort Group.
Struthiocephalus is an extinct genus of dinocephalian therapsids from the Permian of South Africa. It was a large animal, reaching 288 kg (635 lb) in body mass.
Tapinocaninus is an extinct genus of therapsids in the family Tapinocephalidae, of which it is the most basal member. Only one species is known, Tapinocaninus pamelae. The species is named in honor of Rubidge's mother, Pam. Fossils have been found dating from the Middle Permian.
Styracocephalus platyrhynchus is an extinct genus of dinocephalian therapsid that existed during the mid-Permian throughout South Africa, but mainly in the Karoo Basin. It is often referred to by its single known species Styracocephalus platyrhynchus. The Dinocephalia clade consisted of the largest land vertebrates and herbivores during the early to mid-Permian. This period is often also referred to as the Guadalupian epoch, approximately 270 to 260 million years ago.
Keratocephalus is an extinct genus of tapinocephalian therapsids from the early Capitanian age of South Africa. It was found in the Lower and Middle Tapinocephalus Assemblage Zone of the Karoo deposits, in the Lower Beaufort Beds in Beaufort West
Aelurosaurus is a small, carnivorous, extinct genus of gorgonopsian therapsids from the Late Permian of South Africa. It was discovered in the Karoo Basin of South Africa, and first named by Richard Owen in 1881. It was named so because it appeared to be an ancestor for cat-like marsupials, but not yet a mammal itself. It contains five species, A. felinus, A. whaitsi, A. polyodon, A. wilmanae, and A.? watermeyeri. A. felinus, the type species, is generally well described with established features, while the other four species are not due to their poorly preserved holotypes.
Bullacephalus is an extinct genus of biarmosuchian therapsids belonging to the family Burnetiidae. The type species B. jacksoni was named in 2003. It is known from a relatively complete skull and lower jaw, discovered in the Late Permian Tapinocephalus Assemblage Zone of the Beaufort Group of South Africa. This genus of therapsida lived during the Late Permian period, approximately 250 million years ago. The name Bullacephalus comes from the Latin words "bullatus," meaning "bossed" or "knobbed," and "cephalus," meaning "head." This name refers to the distinctive bony knob on the top of the therapsid's skull, which contributes to the history of this genus. This stem based taxon includes Ictidorhinus or Hippasaurs. Bullacephalus can even be characterized as having a, “skull moderately to greatly pachyostotic; swollen boss present above the postorbital bar formed by the postfrontal and postorbital; deep linear sculpturing of the snout; exclusion of the jugal from the lateral temporal fenestra”. These Therapsids have spongy bone skull roof, palatal process of premaxilla are long, diverticulum of naris adding them to the Burnetiamorph. Furthermore, the discovery of Bullacephalus has helped to refine the taxonomic classification of therapsids. Prior to its discovery, there was uncertainty regarding the relationship between different groups of therapsids, particularly the Burnetiamorpha and the Biarmosuchia. However, the distinctive features of Bullacephalus suggest that it is a member of the Burnetiamorpha, and provides a bridge between this group and the Biarmosuchia. The discovery of Bullacephalus has also highlighted the importance of continued exploration and excavation in areas that have yielded few therapsid fossils. The Beaufort Group of South Africa, where Bullacephalus was discovered, has been an important site for therapsid fossils, but much of the area remains unexplored. Further discoveries in this region and other areas around the world may provide new insights into the evolution and diversification of therapsids, as well as other groups of extinct animals. These discoveries will also help to refine our understanding of the history of life on Earth and the processes that have shaped the diversity of organisms that exist today.
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.
Moschognathus is an extinct genus of dinocephalian therapsid in the family Tapinocephalidae. The genus includes only the type species M. whaitsi, named by palaeontologist Robert Broom in 1914. It was a short-snouted tapinocephalid, closely related to and resembling the well-known genus Moschops, but its skull is less thickened overall has a relatively longer and shallower snout by comparison. Indeed, Moschognathus has typically been regarded as a junior synonym of Moschops since 1969 after Lieuwe Dirk Boonstra sunk Moschognathus into Moschops, albeit retained as its own doubtfully valid species. However, researchers in the 21st century have expressed doubt over this synonymy and suggested that Moschognathus is a distinct taxon after all, including first by Christian Kammerer in a 2009 Ph.D. thesis and formally in 2015 by Alessandra D. S. Boos and colleagues in 2015. Moschognathus has since began to re-enter scientific literature of dinocephalians as a valid name and treated distinct from Moschops.
Criocephalosaurus is an extinct genus of tapinocephalian therapsids that lived in Southern Africa during the Guadalupian epoch of the Permian. They are the latest surviving dinocephalians, extending past the Abrahamskraal Formation into the lowermost Poortjie Member of the Teekloof Formation in South Africa. They are also regarded as the most derived of the dinocephalians, alongside Tapinocephalus, and the most abundant in the fossil record.
The Abrahamskraal Formation is a geological formation and is found in numerous localities in the Northern Cape, Western Cape, and the Eastern Cape of South Africa. It is the lowermost formation of the Adelaide Subgroup of the Beaufort Group, a major geological group that forms part of the greater Karoo Supergroup. It represents the first fully terrestrial geological deposits of the Karoo Basin. Outcrops of the Abrahamskraal Formation are found from the small town Middelpos in its westernmost localities, then around Sutherland, the Moordenaarskaroo north of Laingsburg, Williston, Fraserburg, Leeu-Gamka, Loxton, and Victoria West in the Western Cape and Northern Cape. In the Eastern Cape outcrops are known from Rietbron, north of Klipplaat and Grahamstown, and also southwest of East London.
Phorcys is an extinct genus of gorgonopsian that lived during the Middle Permian period (Guadalupian) of what is now South Africa. It is known from two specimens, both portions from the back of the skull, that were described and named in 2022 as a new genus and species P. dubei by Christian Kammerer and Bruce Rubidge. The generic name is from Phorcys of Greek mythology, the father of the Gorgons from which the gorgonopsians are named after, and refers to its status as one of the oldest representatives of the group in the fossil record. Phorcys was recovered from the lowest strata of the Tapinocephalus Assemblage Zone (AZ) of the Beaufort Group, making it one of the oldest known gorgonopsians in the fossil record—second only to fragmentary remains of an indeterminate gorgonopsian from the older underlying Eodicynodon Assemblage Zone.