Anomocephaloidea Temporal range: | |
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Skulls of Anomocephalus (left) and Tiarajudens (right) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Anomocephaloidea Cisneros et al., 2011 |
Genera | |
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 (the clade's namesake) 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. [1] [2]
Anomocephaloids are characterised by batteries of tightly occluding molariform teeth at the back of the jaws, the first example of such teeth in the therapsid fossil record. Uniquely to anomocephaloids, though, the upper molariforms are in fact expanded palatal teeth from the roof of the mouth that occlude against the marginal dentition of the lower jaw. Such occlusion between palatal and marginal teeth is not known in any other synapsids. The precise occlusion, heavy wear, and rapid tooth replacement of anomocephaloid teeth all suggests that they fed upon tough, high-fibre vegetation. The adaptations of anomocephaloids to herbivory were novel both within anomodonts, but also for therapsids and mammal evolution as a whole. [3] [4]
Anomocephalus and Tiarajudens are very similar to each other, and are only definitely distinguished by the large sabre-like caniniform teeth of Tiarajudens. They both had short and deep skulls resembling those of dicynodonts, but although their snouts are short they are still proportionally longer than those of other anomodonts (occupying ~45% of the total skull length). [5] Both species had skulls between 21–22.5 centimetres (8.3–8.9 in) long, a third longer than that of the next largest basal anomodont ( Ulemica ). [5]
From their limited post-cranial remains, anomocephaloids were bulky and dicynodont-like in proportions (although they were not especially closely related to them). They were unlike other, smaller basal anomodonts, such as the venyukovioids, which were slender and lightly built. Despite the limited material, however, both species are some of the few therapsids known to have preserved gastralia, or belly ribs. [2] [3]
The dentition of anomocephaloids is unusual and highly characteristic. The molariform palatal teeth are broad, three times wider than long, and when worn have a saddle-shaped crown with a raised labial (outer) edge and a broader, lower lingual (inner) surface. They are tightly packed and arranged en echelon across two bones of the palate, the pterygoid bone and, uniquely for therapsids, the ectopterygoid. Ectopterygoid teeth are found in 'pelycosaur'-grade synapsids, but are otherwise absent from therapsids except for anomocephaloids. These teeth were replaced frequently in waves, indicative of their heavy wear and use. These teeth tightly occluded with the similarly shaped rear teeth of the lower jaw, an adaptation to efficiently process high-fibre food in herbivorous tetrapods. Occlusion between palatal and marginal teeth is wholly unique to anomocephaloids amongst synapsids, with only the herbivorous 'pelycosaur' Edaphosaurus showing a comparable arrangement (in which palatal teeth contacted tooth-plates in the mandibles). [2] [3]
This configuration of the palatal teeth means that, unusually, the marginal upper dentition (i.e. the teeth of the premaxilla and maxilla) of both species are almost entirely incisiform. These teeth are leaf-shaped and coarsely serrated in Tiarajudens but are blunt and rounded in Anomocephalus, although this may just be due to heavy wear similar to the palatal teeth. The only other teeth are a tiny, peg-like pre-caniniform in both species and the sabre-like caniniforms themselves of Tiarajudens at the end of the marginal tooth row. The lower dentition likewise is made up of incisiforms at the front and molariforms at the rear. [3]
Although superficially similar to dicynodonts, anomocephaloids lacked the specialised slip of jaw muscle of dicynodonts (and some venyukovioids) that pulled the lower jaw backwards (palinally) to chew. Nonetheless, an incipient palinal jaw stroke may have been possible, as the lower jaw joint had sockets twice as long as they are wide and so potentially allowed the jaw joint to slide back-and-forth against quadrate bone. [5] At the same time, anomocephaloids may also be characterised by a much larger coronoid process on the mandible than other anomodonts (only definitively known in Anomocephalus) which would have supported large and powerful jaw musculature. [3]
Anomocephaloidea was defined by Cisneros et al. (2011) as the clade of all taxa closer to Anomocephalus than to Otsheria , a venyukovioid, and was named for the clade containing Anomocephalus and Tiarajudens. In this analysis, Anomocephaloidea was more derived than the basal-most anomodont Biseridens but outside of the clade of Venyukovioidea + Chainosauria. Subsequent analyses (such as Boos et al. 2016) [6] recovered similar results, however, a more recent analysis by Angielczyk and Kammerer in 2017 instead found Anomocephaloidea as a clade within Chainosauria, i.e. closer to dicynodonts than the venyukovioids are, and subsequent analyses (such as Angielczyk et al. 2021) have recovered the same result. [7] [8]
Two cladograms are shown below depicting the original phylogeny from Cisneros et al. (2011) (left) and that of Angielczyk & Kammerer (2017) (right), both simplified, highlighting the differing placement of Anomocephaloidea within anomodonts:
Cisneros et al. (2011): [2]
Anomodontia |
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Angielczyk and Kammerer (2017): [7]
Anomodontia |
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Anomocephaloids demonstrate that early anomodonts, although rare, were experimenting with diverse morphologies and ecologies for herbivory distinct from those of the highly successful dicynodonts. They also demonstrate that precise, mammalian levels of tooth occlusion evolved much earlier and independently of them in synapsid evolution. [2] [3] [4]
Anomocephalus was once considered the most basal known anomodont, and so was used to infer a Gondwanan origin for the whole group. The discovery of the closely related Tiarajudens forming the clade Anomocephaloidea suggested they may represent an endemic radiation of early Gondwanan anomodonts, comparable to the radiation of venyukovioids in Eurasia. [2]
A therapsid is a member of the clade Therapsida which is a major group of eupelycosaurian synapsids that includes mammals and their ancestors and 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, as opposed to the sprawling posture of many reptiles and salamanders.
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.
Suminia is an extinct genus of basal anomodont that lived during the Tatarian age of the late Permian, spanning approximately from 268-252 Ma. Suminia is recognized the youngest non-dicynodont anomodont. Its fossil localities are primarily derived from the Kotel’nich locality of the Kirov Oblast in Russia. However, there have been some isolated specimen found in a few different localities, all from eastern European regions of Russia.
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.
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.
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.
Eodicynodon 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.
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.
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.
Dimacrodon is an extinct genus of non-mammalian synapsid from the latest Early Permian San Angelo Formation of Texas. It is distinguished by toothless, possibly beaked jaw tips, large lower canines and a thin bony crest on top of its head. Previously thought to be an anomodont therapsid related to dicynodonts, it was later found to lack any diagnostic features of anomodonts or even therapsids and instead appears to be a 'pelycosaur'-grade synapsid of uncertain classification.
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.
Daptocephalus is an extinct genus of non-mammalian synapsid anomodont dicynodont, it 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.
Tiarajudens is an extinct genus of saber-toothed herbivorous anomodonts which lived during the Middle Permian period in what is now Rio Grande do Sul, Brazil. It is known from the holotype UFRGS PV393P, a nearly complete skull. The type species T. eccentricus was named in 2011.
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.
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.
Parasumina is an extinct genus of anomodont known from the late Capitanian age at the end of the middle Permian period of European Russia. The type and only species is Parasuminia ivakhnenkoi. It was closely related to Suminia, another Russian anomodont, and was named for its resemblance. Little is known about Parasuminia as the only fossils are of fragmentary pieces of the skull and jaw, but the known remains suggest that its head and jaws were deeper and more robust than those of Suminia, and with shorter, stouter teeth. However, despite these differences they appear to have been similar animals with a similarly complex method of processing vegetation.
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.