Eodicynodon Temporal range: | |
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Restoration of Eodicynodon oosthuizeni | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Family: | † Eodicynodontidae Barry, 1974 |
Genus: | † Eodicynodon Barry, 1974 |
Type species | |
†E. oosthuizeni Barry, 1974 |
Eodicynodon (eo-, early or primitive, dicynodont) is an extinct genus of dicynodont therapsids, a highly diverse group of herbivorous synapsids that were widespread during the middle-late Permian and early Triassic. As its name suggests, Eodicynodon is the oldest and most primitive dicynodont yet identified, ranging from the middle to late Permian and possessing a mix of ancestral anomodont/therapsid features and derived dicynodont synapomorphies.
First described by paleontologist T. H. Barry in 1974, E. oosthuizeni is named after Roy Oosthuizen, the South African farmer who discovered the type specimen (a partial skull without the mandible) on his Cape Province farm between 1964 and 1970. [1]
The Eodicynodon Assemblage Zone in South Africa is named after its presence there.
Eodicynodon was a medium-sized dicynodont, at about 450 mm long and 150 mm high. [2] While it had many features common to all dicynodonts, such as canine tusks and jaw structures related to the "cheek pivot system" of mastication, it also displayed a number of ancestral features more similar to some of its primitive therapsid relatives, [2] [3] [1] [4] which are listed below.
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While the premaxillary bones are fused in more derived dicynodonts, a thin suture extending dorsally up from the palatal facet reveals that they are paired in Eodicynodon, an ancestral feature they share with their primitive relatives Venyukovia, Otsheria, and Pelycosauria. [1]
Similarly, while the vomers of later dicynodonts are fused, they are paired in Eodicynodon and together with the pterygoid border the intervomero-pterygoidal vacuity; in more derived dicynodonts, this vacuity is more posteriorly located and exclusively bordered by the pterygoid. These features are also present in more primitive relatives of dicynodonts, including Pristerodon, [5] sphenacodont pelycosaurs, cotylosaurs, and Venyukovia. [6] [1]
Dicynodonts were specialized herbivores that employed a unique “cheek pivot system” of mastication that created powerful shearing action upon closure of the jaw and subsequently ground mouth contents through a system of interlocking ridges and grooves formed from the palate and dentary. [3]
Two morphological features, present already in Eodicynodon, made this motion possible. The first was a double convex jaw joint, wherein both the quadrate and articular formed convex condyles. As the jaw closed, the articular condyle of the lower jaw slid anterio-dorsally along the quadrate condyle, resulting in closure of the mouth from back to front as the posterior end of the mandible was elevated dorsally relative to the anterior end. Forward slide of the lower jaw was limited by the second morphological feature unique to dicynodonts, a pivot point created between the dentary groove and palatal notch upon closure of the jaw. The lower jaw would then move so that the articular condyle slid anterio-ventrally along the quadrate condyle, which would cause the mandible to pivot in such a way that the front of the mouth closed and the back opened. [3]
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This section is empty. You can help by adding to it. (March 2018) |
The South African Karoo Supergroup is a fossil-rich series of bedded shales that was continuously deposited beginning in the late Carboniferous through the early Jurassic. Though a diverse assemblage of dicynodonts appears early on in the Beaufort Group, the immediately preceding Ecca was long understood to be barren of fossils, despite a lack of geological evidence for a change in paleoenvironment from Ecca to Beaufort. From 1964 to 1970, the farmer Roy Oosthuizen, whose land was located in an area firmly established as Upper Ecca (Middle Permian), collected a number of nodules containing the remains of several therapsids, including several small dicynodonts and the partial skull that is the type specimen of Eodicynodon. [1]
Below is a cladogram modified from Angielczyk and Rubidge (2010) showing the phylogenetic relationships of Dicynodontia: [7]
Dicynodontia |
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Robertia is an extinct genus of small herbivorous dicynodonts from the Middle to Late Permian of South Africa, between 260 and 265 million years ago. It is a monospecific genus, consisting of the type-species R. broomiana, which was classified by Lieuwe Dirk Boonstra in 1948 and named in honor of Robert Broom for his study of South African mammal-like reptiles.
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 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.
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.
Anomocephalus is an extinct genus of primitive anomodonts and belongs to the clade Anomocephaloidea. The name is said to be derived from the Greek word anomos meaning lawless and cephalos meaning head. The proper word for head in Greek is however κεφαλή (kephalē). It is primitive in that it retains a complete set of teeth in both jaws, in contrast to its descendants, the dicynodonts, whose dentition is reduced to only a single pair of tusks, with their jaws covered by a horny beak similar to that of a modern tortoise. However, they are in no way closely related.
Patranomodon is an extinct genus belonging to the group of Anomodontia. Rubidge and Hopson named this anomodont in 1990 after discovering its skull. Patranomodon is known to have ranged in the Karoo of Southern Africa.
Brachyprosopus is an extinct genus of dicynodont therapsid from the middle Permian Tapinocephalus Assemblage Zone in the Abrahamskraal Formation belonging to the Beaufort Group of the Karoo Basin, South Africa.
Biseridens is an extinct genus of anomodont therapsid, and one of the most basal anomodont genera known. Originally known from a partial skull misidentified as an eotitanosuchian in 1997, another well-preserved skull was found in the Qingtoushan Formation in the Qilian Mountains of Gansu, China, in 2009 that clarified its relationships to anomodonts, such as the dicynodonts.
Daptocephalus is an extinct genus of dicynodont synapsid, which was found in Late Permian strata, in a biozone known precisely for the presence of fossils of this dicynodont, the Daptocephalus Assemblage Zone, in the Karoo Basin in South Africa. An additional species, D. huenei, is known from the Usili Formation in Tanzania and was formerly assigned to the genus Dicynodon before a study in 2019 recognised that the type specimen belonged to Daptocephalus.
Eosimops is an extinct genus of pylaecephalid dicynodonts. They were small synapsids superficially resembling modern mammals. Eosimops is known from several skull specimens, as well as one complete skeleton. Eosimops lived during the Middle Permian of South Africa.
Kawingasaurus is an extinct genus of dicynodont therapsid from the Late Permian Usili Formation of Tanzania. It is a member of the family Cistecephalidae, and like other cistecephalids it is thought to have been fossorial. It is a member of the family Cistecephalidae. Cistephalidae includes genera Cisteceohalus, Cistecephaloides and Kawingasaurus. Greek for Saurus meaning “lizard” appears as a suffix denoting a reptilian origin. Living between 254.17 and 259.9 million years ago in the late Permian and believed to have the first and last recorded appearance in this time period. It lived in deep burrows as a suggested by most burrowing dicynodonts from evaluation of cranial sutures, vestibule inflation and enlarged stapes foot plates which are thought to be functionally correlated with bone-conduction hearing; all observed in fossorial vertebrates which use seismic signals as communication.
Pristerodon is an extinct genus of dicynodont therapsid from the Late Permian of South Africa, Zambia and India.
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.
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.
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.
Anomocephaloidea is a clade of basal anomodont therapsids related to the dicynodonts known from what is now South Africa and Brazil during the Middle Permian. It includes only two species, Anomocephalus africanus from the Karoo Basin of South Africa and Tiarajudens eccentricus from the Paraná Basin of Brazil. Anomocephaloidea was named in 2011 with the discovery of Tiarajudens, although Anomocephalus itself has been known since 1999.
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.
Nyaphulia is an extinct genus of dicynodont therapsid from the middle Permian of South Africa, containing only the type species N. oelofseni. The generic name is in honour of John Nyaphuli of the National Museum of Bloemfontein, who contributed extensively to South African palaeontology and discovered the holotype specimen of Nyaphulia in 1982. Nyaphulia was initially named as a second species of the basal dicynodont Eodicynodon by Professor Bruce Rubidge in 1990 as E. oelofseni, named after his mentor in palaeontology and geology Dr. Burger Oelofsen.