Eutherocephalia Temporal range: Late Permian-Middle Triassic, | |
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Life restoration of the eutherocephalian Moschorhinus preying on the dicynodont Lystosaurus . | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | † Therocephalia |
Clade: | † Scylacosauria |
Clade: | † Eutherocephalia Hopson and Barghusen, 1986 |
Families and genera | |
Eutherocephalia ("true beast head") is an extinct clade of advanced therocephalian therapsids. Eutherocephalians are distinguished from the lycosuchids and scylacosaurids, two early therocephalian families. While lycosuchids and scyalosaurids became extinct by the end of the Permian period, eutherocephalians survived the Permian–Triassic extinction event. The group eventually became extinct in the Middle Triassic.
The Eutherocephalians evolved several mammal-like traits through convergent evolution with Cynodontia. Among those traits were the loss of palatine teeth and the reduction of the parietal eye. [1] The latter organ is instrumental in thermoregulation among lizards and snakes, indicating both eutherocephalians and cynodonts were evolving toward a more active, homeothermic lifestyle, though the eye never fully disappeared in the eutherocephalians. [2]
The clade Eutherocephalia contains the majority of therocephalians, yet the phylogenetic relations of the groups within it remain unclear. Eutherocephalia is supported as a true clade in many phylogenetic analyses, but the placement of groups like Akidnognathidae, Hofmeyriidae, Whaitsiidae, and Baurioidea, all of which lie within Eutherocephalia, remains debated. [1]
Therapsida is a clade composing of a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, resulting in a more "standing" quadripedal posture, as opposed to the lower sprawling posture of many reptiles and amphibians.
Eucynodontia is a clade of cynodont therapsids including mammals and most non-mammalian cynodonts. The oldest eucynodonts are known from the Early Triassic and possibly Late Permian. Eucynodontia includes two major subgroups, Cynognathia and Probainognathia.
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.
Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores. Anomodonts were very diverse during the Middle Permian, including primitive forms like Anomocephalus and Patranomodon and groups like Venyukovioidea and Dromasauria. Dicynodonts became the most successful and abundant of all herbivores in the Late Permian, filling ecological niches ranging from large browsers down to small burrowers. Few dicynodont families survived the Permian–Triassic extinction event, but one lineage (Kannemeyeriiformes) evolved into large, stocky forms that became dominant terrestrial herbivores right until the Late Triassic, when changing conditions caused them to decline, finally going extinct during the Triassic–Jurassic extinction event.
Therocephalia is an extinct clade of eutheriodont therapsids from the Permian and Triassic periods. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their teeth, suggest that they were carnivores. Like other non-mammalian synapsids, therocephalians were once described as "mammal-like reptiles". Therocephalia is the group most closely related to the cynodonts, which gave rise to the mammals, and this relationship takes evidence in a variety of skeletal features. Indeed, it had been proposed that cynodonts may have evolved from therocephalians and so that therocephalians as recognised are paraphyletic in relation to cynodonts.
Euchambersia is an extinct genus of therocephalian therapsids that lived during the Late Permian in what is now South Africa and China. The genus contains two species. The type species E. mirabilis was named by paleontologist Robert Broom in 1931 from a skull missing the lower jaw. A second skull, belonging to a probably immature individual, was later described. In 2022, a second species, E. liuyudongi, was named by Jun Liu and Fernando Abdala from a well-preserved skull. It is a member of the family Akidnognathidae, which historically has also been referred by as the synonymous Euchambersiidae.
Moschorhinus is an extinct genus of therocephalian synapsid in the family Akidnognathidae with only one species: M. kitchingi, which has been found in the Late Permian to Early Triassic of the South African Karoo Supergroup. It was a large carnivorous therapsid, reaching 1.5 m (4.9 ft) in total body length with the largest skull comparable to that of a lion in size, and had a broad, blunt snout which bore long, straight canines.
Theriognathus is an extinct genus of therocephalian therapsid belonging to the family Whaitsiidae, known from fossils from South Africa, Zambia, and Tanzania. Theriognathus has been dated as existing during the Late Permian. Although Theriognathus means mammal jaw, the lower jaw is actually made up of several bones as seen in modern reptiles, in contrast to mammals. Theriognathus displayed many different reptilian and mammalian characteristics. For example, Theriognathus had canine teeth like mammals, and a secondary palate, multiple bones in the mandible, and a typical reptilian jaw joint, all characteristics of reptiles. It is speculated that Theriognathus was either carnivorous or omnivorous based on its teeth, and was suited to hunting small prey in undergrowth. This synapsid adopted a sleek profile of a mammalian predator, with a narrow snout and around 1 meter long. Theriognathus is represented by 56 specimens in the fossil record.
Tetracynodon is an extinct genus of therocephalian. Fossils of Tetracynodon have been found in the Karoo Basin of South Africa. Two species are known: the type species T. tenuis from the Late Permian and the species T. darti from the Early Triassic. Both species were small-bodied and probably fed on insects and small vertebrates. Although Tetracynodon is more closely related to mammals than to reptiles, its braincase is very primitive and more resembles that of modern amphibians and reptiles than of mammals.
Paraburnetia is an extinct genus of biarmosuchian therapsids from the Late Permian of South Africa. It is known for its species P. sneeubergensis and belongs to the family Burnetiidae. Paraburnetia lived just before the Permian–Triassic mass extinction event.
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.
Akidnognathidae is an extinct family of therocephalian therapsids from the Late Permian and Early Triassic of South Africa, Russia and China. The family includes many large-bodied therocephalians that were probably carnivorous, including Moschorhinus and Olivierosuchus. One akidnognathid, Euchambersia, may even have been venomous. Akidnognathids have robust skulls with a pair of large caniniform teeth in their upper jaws. The family is morphologically intermediate between the more basal therocephalian group Scylacosauridae and the more derived group Baurioidea.
Bauriidae is an extinct family of therocephalian therapsids. Bauriids were the latest-surviving group of therocephalians after the Permian–Triassic extinction event, going extinct in the Middle Triassic. They are among the most advanced eutherocephalians and possess several mammal-like features such as a secondary palate and wide postcanine teeth at the back of the jaws. Unlike other therocephalians, bauriids were herbivorous. They were also smaller than earlier members of the group. Two subfamilies are classified within Bauriidae: Nothogomphodontinae and Bauriinae.
Blattoidealestes is an extinct genus of therocephalian therapsid from the Middle Permian of South Africa. The type species Blattoidealestes gracilis was named by South African paleontologist Lieuwe Dirk Boonstra from the Tapinocephalus Assemblage Zone in 1954. Dating back to the Middle Permian, Blattoidealestes is one of the oldest therocephalians. It is similar in appearance to the small therocephalian Perplexisaurus from Russia, and may be closely related.
Pelictosuchus is an extinct genus of therocephalian therapsids from the Late Permian of South Africa. It is classified in the family Akidnognathidae. The type species Pelictosuchus paucidens was named by South African paleontologist Robert Broom in 1940 from the Dicynodon Assemblage Zone.
Eutheriodontia is a clade of therapsids which appear during the Middle Permian and which includes therocephalians and cynodonts, this latter group including mammals and related forms.
Lycideops is an extinct genus of therocephalians from the Late Permian of South Africa. The type species is Lycideops longiceps, named in 1931 by South African paleontologist Robert Broom. Fossils of Lycideops come from the Dicynodon Assemblage Zone of the Beaufort Group. Lycideops is a member of the family Lycideopidae. Like other lycideopids, Lycideops has a long snout.
Ichibengops is an extinct genus of therocephalian therapsids known from the type species Ichibengops munyamadziensis, which lived in what is now Zambia during the Late Permian. Ichibengops was named in 2015 on the basis of fossils found in the Wuchiapingian-age Madumabisa Mudstone Formation in the Luangwa Basin. Therocephalians have been known from the Luangwa Basin for decades, yet Ichibengops was the first endemic Zambian therocephalian to have been described in detail. Phylogenetic analysis indicates that it is a basal member of the clade Eutherocephalia, lying just outside a clade containing hofmeyriids, whaitsiids, and baurioids. Ichibengops is the sister taxon of the Russian therocephalian Chthonosaurus; together they form one of several known African-Russian sister taxon pairs of eutherocephalians, which indicate that eutherocephalians could freely disperse across most of Pangea during the Late Permian. Like the fellow therocephalian Euchambersia, Ichibengops might have had venom glands, as evidenced by grooves above its teeth.
Thliptosaurus is an extinct genus of small kingoriid dicynodont from the latest Permian period of the Karoo Basin in KwaZulu-Natal, South Africa. It contains the type and only known species T. imperforatus. Thliptosaurus is from the upper Daptocephalus Assemblage Zone, making it one of the youngest Permian dicynodonts known, living just prior to the Permian mass extinction. It also represents one of the few small bodied dicynodonts to exist at this time, when most other dicynodonts had large body sizes and many small dicynodonts had gone extinct. The unexpected discovery of Thliptosaurus in a region of the Karoo outside of the historically sampled localities suggests that it may have been part of an endemic local fauna not found in these historic sites. Such under-sampled localities may contain 'hidden diversities' of Permian faunas that are unknown from traditional samples. Thliptosaurus is also unusual for dicynodonts as it lacks a pineal foramen, suggesting that it played a much less important role in thermoregulation than it did for other dicynodonts.
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.