Bidentalia Temporal range: Middle Permian - Norian, | |
---|---|
Skull of Dicynodon | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Clade: | † Therochelonia |
Clade: | † Bidentalia Owen, 1876 |
Bidentalia is a group of dicynodont therapsids. Bidentalia was one of the first names used to describe dicynodonts; the group was established in 1876, while the name "bidentals" dates back as far as 1845. With the increasing prominence of phylogenetics, the group was redefined as a clade in 2009. Bidentalia is now considered a stem-based taxon that includes all taxa more closely related to Aulacephalodon bainii and Dicynodon lacerticeps than Emydops arctatus .
In 1845, South African geologist Andrew Geddes Bain described the first known dicynodonts as "bidentals" for their two prominent tusks. [1] "Bidental" was the first name ever used for a group of non-mammalian synapsids. [2] The name Dicynodontia, which is more commonly used to refer to these animals, was erected by English paleontologist Richard Owen in 1860. [3] Realizing that Bain's "bidentals" predated his "dicynodonts", Owen named Bidentalia in 1876 as a replacement name for Dicynodontia. Owen described Bidentalia as reptiles with "a long ever-growing tusk in each maxillary; premaxillaries connate, forming with the lower jaw a beak-shaped mouth, probably sheathed with horn. Sacrum of more than two vertebrae; trunk-vertebrae amphicoelian; limbs ambulatory." [4] At this time, Bidentalia included three main species: Dicynodon lacerticeps , Dicynodon bainii , and Ptychognathus declivis . The two Dicynodon species were named in 1845, just before Bain described his bidentals. Ptychognathus was named in 1859, and is now called Lystrosaurus . In the following years Dicynodontia became the preferred name for these reptiles and Bidentalia quickly fell out of use. [2]
Bidentalia was reinstated as a clade in 2009. [2] It was used to include all therochelonians more closely related to Dicynodon than to emydopoids (a group of more basal dicynodonts). As a clade, Bidentalia forms a more inclusive group than it did under Owen's use. Owen's Bidentalia was equivalent to Dicynodontia, which today is used as a much larger group encompassing all dicynodonts. In its current use, Bidentalia includes two major subgroups, Cryptodontia and Dicynodontoidea.
Bidentalia originated in the Guadalupian epoch of the Permian and quickly spread worldwide, with early species known from both Brazil and Russia. However, they were conspicuously absent from Africa despite its well-known Permian fauna until the late Permian. [5] The oldest and most basal known bidentalian is Rastodon .
Bidentalia is divided into two major groups, the Cryptodontia and the Dicynodontoidea, as well as a small number of basal genera such as Elph and Rastodon [5] However, the Cryptodontia may be paraphyletic. Below is a cladogram showing the phylogeny of Bidentalia from a recent study, Kammerer et al. (2011): [6]
Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typically toothless beak, unique amongst all synapsids. Dicynodonts first appeared in Southern Pangaea during the mid-Permian, ca. 270–260 million years ago, and became globally distributed and the dominant herbivorous animals in the Late Permian, ca. 260–252 Mya. They were devastated by the end-Permian Extinction that wiped out most other therapsids ca. 252 Mya. They rebounded during the Triassic but died out towards the end of that period. They were the most successful and diverse of the non-mammalian therapsids, with over 70 genera known, varying from rat-sized burrowers to elephant-sized browsers.
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.
Cistecephalus is an extinct genus of dicynodont therapsid from the Late Permian of southern Africa. It was a small, specialised, burrowing dicynodont, possibly with habits similar to a modern mole. The head was flattened and wedge-shaped, the body long, and the forelimbs very strong, with similarities in structure to the forelimb of modern burrowing mammals.
Diictodon is an extinct genus of pylaecephalid dicynodont that lived during the Late Permian period, approximately 255 million years ago. Fossils have been found in the Cistecephalus Assemblage Zone of the Madumabisa Mudstone of the Luangwa Basin in Zambia and the Tropidostoma Assemblage Zone of the Teekloof Formation, Tapinocephalus Assemblage Zone of the Abrahamskraal Formation, Dicynodon Assemblage Zone of the Balfour Formation, Cistecephalus Assemblage Zone of the Middleton or Balfour Formation of South Africa and the Guodikeng Formation of China. Roughly half of all Permian vertebrate specimens found in South Africa are those of Diictodon. This small herbivorous animal was one of the most successful synapsids in the Permian period.
"Dromasaurs" are an artificial grouping of small anomodont therapsids from the Middle and Late Permian of South Africa. They represent either a paraphyletic grade or a polyphyletic grouping of small non-dicynodont basal anomodonts rather than a clade, and as such are considered an invalid group today. "Dromasaurs" were historically united by their superficially similar appearances that were unlike other known anomodonts. They are all small in size with slender limbs and long tails, and have short skulls with very large eye sockets. "Dromasauria" traditionally includes three genera, all from the Karoo Supergroup of South Africa: Galepus, Galechirus, and Galeops. These genera have sometimes been divided into two subgroups, the monotypic family Galeopidae and the Galechiridae for Galechiris and Galepus.
Dicynodon is a genus of dicynodont therapsid that flourished during the Upper Permian period. Like all dicynodonts, it was an herbivorous animal. This synapsid was toothless, except for prominent tusks, hence the name. It probably cropped vegetation with a horny beak, much like a tortoise, while the tusks may have been used for digging up roots and tubers.
Emydops is an extinct genus of dicynodont therapsids from the Middle Permian to Late Permian of what is now South Africa. The genus is generally small and herbivorous, sharing the dicynodont synapomorphy of bearing two tusks. In the following years, the genus grew to include fourteen species. Many of these species were erected on the basis of differences in the teeth and the positioning of the frontal and parietal bones. A 2008 study narrowed Emydops down to two species, E. arctatus and the newly described E. oweni.
Dicynodontoides is a genus of small to medium-bodied, herbivorous, emydopoid dicynodonts from the Late Permian. The name Dicynodontoides references its “dicynodont-like” appearance due to the caniniform tusks featured by most members of this infraorder. Kingoria, a junior synonym, has been used more widely in the literature than the more obscure Dicynodontoides, which is similar-sounding to another distantly related genus of dicynodont, Dicynodon. Two species are recognized: D. recurvidens from South Africa, and D. nowacki from Tanzania.
Tropidostoma is a medium-sized herbivorous oudenodontid dicynodont therapsid that lived during the Late Permian (Lopingian) period in South Africa. The first Tropidostoma fossil was described by Harry Govier Seeley in 1889. Later two subspecies were identified. Tropidostoma fossils are an index fossil in a biozone of the Karoo Basin known as the Tropidostoma Assemblage Zone. This biozone is characterized by the presence of this species in association with another dicynodont species, Endothiodon uniseries.
Aulacephalodon is an extinct genus of medium-sized dicynodonts, or non-mammalian synapsids, that lived during late Permian period. Individuals of Aulacephalodon are commonly found in the Lower Beaufort Group of the Karoo Supergroup of South Africa. Rising to dominance during the Late Permian, Aulacephalodon was among the largest terrestrial vertebrate herbivores until its extinction at the end of the Permian. Two species have been named, the type species, A. bainii, and a second species, A. peavoti. However, debate exists among paleontologists if A. peavoti is a true member of the genus Aulacephalodon. Aulacephalodon belongs to the family Geikiidae, a family of dicynodonts generally characterized by their short, broad skulls and large nasal bosses. Sexual dimorphism has been identified in A. bainii.
Lystrosauridae is a family of dicynodont therapsids from the Permian and Triassic time periods. It includes two genera, Lystrosaurus and Kwazulusaurus. Kwazulusaurus includes a single species, K. shakai, from the Late Permian of South Africa and Lystrosaurus includes many species from the Late Permian and Early Triassic of South Africa, India, and Antarctica.
Venyukovioidea is an infraorder of anomodont therapsids related to dicynodonts from the Permian of Russia. They have also been known as 'Venjukovioidea', as well as by the similar names 'Venyukoviamorpha' or 'Venjukoviamorpha' in literature. This in part owes to a misspelling by Russian palaeontologist Ivan Efremov in 1940 when he mistakenly spelt Venyukovia, the namesake of the group, with a 'j' instead of a 'y', which permeated through subsequent therapsid literature before the mistake was caught and corrected. The order Ulemicia has also been coined for a similar taxonomic concept in Russian scientific literature, which notably excludes Suminia and Parasuminia.
Chainosauria is a large and speciose clade of anomodont therapsid that includes the highly diverse dicynodonts and a small number of closely related basal genera —although the total composition and taxonomic scope of Chainosauria is in flux. Chainosauria was named in 1923 to group together the dicynodonts and their close relatives, namely three small anomodont genera from South Africa that made up the now defunct group 'Dromasauria'. The name soon fell into disuse, however, as it was functionally replaced by Anomodontia. Chainosauria was later revived cladistically in 2009, preserving the association of dicynodonts and the 'dromasaurs' and has since served in effect as both a cladistic and a biogeographic counterpart to the Laurasian venyukovioids, with early chainosaurs appearing to have been a Gondwanan radiation.
Pylaecephalidae is a family of dicynodont therapsids that includes Diictodon, Robertia, and Prosictodon from the Permian of South Africa. Pylaecephalids were small burrowing dicynodonts with long tusks. The family was first named in 1934 and was redefined in 2009. Diictodontidae and Robertiidae are considered junior synonyms of Pylaecephalidae; although Pylaecephalus itself is considered a junior synonym of Diictodon, the name Pylaecephalidae predates these names and therefore takes priority.
Syops is an extinct genus of dicynodont therapsid. The type species S. vanhoepeni was first named in 1938 as Dicynodon vanhoepeni. Fossils of the genus have been found in the Cistecephalus Assemblage Zone in the Usili Formation of the Ruhuhu Basin, Tanzania and the Upper Madumabisa Mudstone Formation of the Luangwa Basin, Zambia. Its phylogenetic placement is somewhat uncertain, with multiple different studies finding it as either a basal geikiid, rhachiocephalid a dicynodontoid more derived than the most basal genera but less derived than Lystrosauridae, or a lystrosaurid.
Therochelonia is a group of dicynodont therapsids. The group was named by British paleontologist Harry Seeley in 1894 and fell into disuse in the following century. Therochelonia was redefined as a node-based clade in 2009. It is defined as the last common ancestor of Cistecephalus microrhinus and Dicynodon lacerticeps, and all of its descendants. Below is a simplified cladogram from Kammerer et al. (2011) showing the phylogenetic placement of Therochelonia:
Sintocephalus is an extinct genus of dicynodont therapsid from the Late Permian of South Africa. Fossils are known from the Cistecephalus Assemblage Zone of the Beaufort Group. The type species of Sintocephalus, S. alticeps, was first named in 1913 as a species of Dicynodon. The genus was erected in 1934, but in subsequent years its species were often regarded as members of other dicynodont genera.
Gordonia is an extinct genus of dicynodont therapsid from the Late Permian of Scotland. Fossils have been found from the Elgin sandstone of Cutties Hillock Sandstone in Elgin, Moray. These are among the many amniote fossils referred to as the Elgin Reptiles. Gordonia was named in 1893 with four species: G. traquairi, G. duffiana, G. huxleyana, and G. juddiana. Currently, the only recognized species is the type G. traquairi. All other species are considered synonyms of the type.
Jimusaria is an extinct genus of dicynodont therapsid from the Late Permian (Changhsingian) of China. The type species J. sinkianensis from the Guodikeng Formation in Xinjiang, was originally named as a species of Dicynodon, the first from Asia, but was given its own genus in 1963 before being sunk back into Dicynodon in 1988. The genus was resurrected in 2011 by palaeontologist Christian Kammerer in a taxonomic revision of the genus Dicynodon. Jimusaria was a mid-sized dicynodont, and was similar in appearance to the South African Dicynodon, but differed from it in features such as its narrower snout. A second species, Jimusaria monanensis was described from the Naobaogou Formation of northern China in 2023.
Turfanodon is an extinct genus of dicynodont therapsid from the Late Permian Sunan, Guodikeng, and Naobaogou Formations of China. The holotype of T. bogdaensis was discovered between 1963-1964 and was originally named in 1973 by A. Sun with the type species Turfanodon bogdaensis, Turfanodon was reclassified as a junior synonym of the related Dicynodon in 1988 by G. M. King. T. bogdaensis remained a species of Dicynodon for over two decades before the genus was reinstated in 2011 in a revision of the taxonomy of Dicynodon by palaeontologist Christian Kammerer. A second species from Inner Mongolia, T. jiufengensis, was named in 2021 by palaeontologist Jun Liu from a nearly complete skeleton and other referred bones. Turfanodon was a relatively large dicynodont, and similar in appearance to the related Daptocephalus from South Africa.