Biseridens Temporal range: Middle Permian, | |
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Life illustration of Biseridens qilianicus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Genus: | † Biseridens Li and Cheng, 1997 |
Species: | †B. qilianicus |
Binomial name | |
†Biseridens qilianicus Li and Cheng, 1997 | |
Biseridens ("two rows of teeth") 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. [1]
Among tetrapod taxa, the therapsid clade anomodontia in which Biseridens belong has one of the largest population sizes, highest levels of diversity, and longest stratigraphic range (Middle Permian to the Triassic, possibly into the Cretaceous), as well as being one of the only clades known from all the continents. [2] [3] Primarily understood from the recently discovered, most-well preserved specimen, Biseridens is most notably distinguishable as an anomodont due to its short snout, dorsally elevated zygomatic arch, and presence of a septomaxilla that distinctly lacks a facial exposure between the nasal and maxilla. [1] The features that distinguish Biseridens from other anomodonts include the presence of heterodont dentition, or differentiated teeth rows in which different teeth have distinct morphology (ex. precanines, canines, molars, etc.), small toothlike projections, or denticles, located on the palatine and pterygoid, articulation between the opisthotic bone and the tabular bones on the posterior surface of the skull, the absence of the mandibular foramen on the lower jaw, and a pterygoid in which the transverse flange of the pterygoid has a laterally extending process but lacks posterior ramus. [1]
Biseri- is Latin for "double rows"; -dens originated from Greek "odons" for "teeth"
At a clade level, anomodonts were terrestrial vertebrates of all sizes that dominated the Late Permian. [4] The most dominant sub-clade were dicynodonts, more derived relatives of Biseridens. Most anomodonts are postulated to have been herbivorous, [5] and they were the only herbivorous taxa to survive the Permian–Triassic extinction event. [4] The rich diversity within different anomodont genera indicates a range of dental specializations that implies they utilized a variety of food processing mechanisms including modified jaw muscles, jaw joints, and teeth morphology and distribution. [1] A feature that extends across most of the anomodont clade is jaw articulation that allowed for anterior-posterior movement of jaw for slicing. [4]
On the basis of converging faunal evidence and comparison to other regions, the Xidagou Formation has been considered to be Middle Permian [6] and home to the oldest therapsid fauna, suggesting that anomodonts had a Laurasian rather than a Gondwanan origin. [7]
Until the more recent discovery of a new specimen, Biseridens was primarily known from fragmented skull and jaw material discovered in the 1990s in the upper part of the upper Xidagou Formation of Gansu, China. [8] The specimen included an incomplete skull with the posterior portion of the left ramus of the lower jaw and the anterior part of the lower jaws including well-preserved teeth. [8] The more recently discovered specimen includes a nearly complete skull (lacking the occiput) with lower jaws, found with a set of 14 articulated vertebrae. This new specimen was preserved in mudrock and also discovered in the upper part of Xidagou Formation in Dashankau, Yumen in Gansu. [1]
Biseridens is known mostly from skull and jaw material, as well as a series of 14 vertebrae associated with one skull. Like other anomodonts, the region of the skull in front of the eyes is relatively short compared to other therapsids. The orbits are large and rounded, and the temporal fenestra are large and wide and broad zygomatic arches, similar to but not as developed as those of later dicynodonts. The skull is estimated to be at least 17 centimetres (6.7 in) long, relatively small compared to other basal anomodonts.
There are several diagnostic features that characterize this specimen as a Biseridens. Related to its heterodont dentition, Biseridens is distinct from other anomodonts because of a differentiated tooth row that includes two rows of teeth on both the jaws, precanine teeth on the dentary and premaxilla, and a broad spread of teeth on the pterygoid and palatine. There are also denticles on the vomer, palatine, and pterygoid. The postcanines possess oval cross-sections and have grinding surfaces, and the canines have a basal diameter of 10 mm. There are no serrations on the marginal teeth, which aligns Biseridens within the anomodont clade. [1]
The paired pterygoid of Biseridens is the most distinctly visible bone on its palate. It contains an anterior process, a quadrate ramus and a ventromedial process. There is a prominent posterolaterally curving transverse curving process. Similar to other basal therapsids, this process reaches much lower than the level of the palate. [1]
Regarding its skull structure, the features that distinguish Biseridens from other anomodonts include having an intertemporal region that is wider than the interorbital region, and a temporal fenestra larger than the orbits. Additionally, the mandibular fenestra is absent from the lower jaw, the transverse flange of the pterygoid extends laterally, and there is contact between the tabular and opisthotic, or the posterior part of the otic capsule. There is a concavity on the posterior region of the postorbital bone as well as a lateral process on the postorbital that fails to reach the zygomatic arch. Characteristic of other basal therapsids, the prominent maxilla constitutes the majority of the snout and the nasal bones are the longest bones of the skull roof. [1]
Biseridens belong to the clade anomodont which is nested within the clade therapsids, often called "mammal-like reptiles". Therapsids are a group of advanced synapsids, one of the major branches of amniotes that gave rise to mammals.
The holotype and paratype of Biseridens were initially referred to Eotitanosuchia, a former suborder of therapsids that included various groups of early therapsids (although the range inclusion varied between authors). The genus was named from the Latin biseri- for "double rows" and -dens ("teeth") to refer to the paired double rows of cheek teeth in each jaw, while the species was named after the Qilian mountains where the fossils were discovered. They also assigned Biseridens to a distinct new family of eotitanosuchians, the Biseridensidae, although they did not provide a diagnosis for this family outside of the genus itself. [8]
Although relatively primitive, a number of shared traits (synapomorphies) ally Biseridens with anomodonts including the shortened snout, raised zygomatic arch and exclusion of the septomaxilla between the maxilla and nasals. However, it retains a number of primitive traits that exclude it from the more derived anomodonts, including the differentiated tooth row, palatal teeth, contact between tabular and opisthotic; lateral process of transverse flange of pterygoid free of posterior ramus and absence of mandibular foramen. Several cladistic analyses indicate that Biseridens is the most basal anomodont known, including that of Liu and colleagues (2009) shown below, [1] as well as those of Cisneros and colleagues (2011) [9] and Kammerer and colleagues (2013): [10]
The most recent analysis performed by Liu et al. in 2009 indicates a significantly stable placement of Biseridens as the most basal anomodont based on a matrix study performed on 75 cranial characters over 15 synapsid genera and conclusions of primitive and derived features. Based on the geographic discoveries of Biseridans, the most basal anomodont, Raranimus, the most basal therapsid, with basal dinocephalians, bolosaurids, and dissorophids, there is now strong evidence that the oldest therapsids resided in China. [1]
Therapsida |
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Therapsida is a clade comprising 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" quadrupedal posture, as opposed to the lower sprawling posture of many reptiles and amphibians.
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.
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.
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.
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.
Endothiodon is an extinct genus of medium to large dicynodont from the Late Permian. Like other dicynodonts, Endothiodon was an herbivore, but it typically lacked the two tusks that characterized most other dicynodonts and instead had long rows of teeth inset in the jaws that replaced in waves. The anterior portion of the upper and lower jaw are curved upward, creating a distinct beak that is thought to have allowed them to be specialized grazers.
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.
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.
Myosaurus is a genus of dicynodont synapsids. Myosaurus was a small, herbivorous synapsid that existed around the early Triassic period. All of the fossils found of this species were found in Antarctica and South Africa. Compared to other fossils found from species that existed during this time, the Myosaurus is not common in the fossil record. This is due to a shortage of discovered fossils that possess characteristics unique to the Myosaurus. Notably, under 130 fossil fragments have been found that have been classified as Myosauridae, and almost all have been skulls. These skulls can be classified as Myosaurus because this species, unlike other dicynodonts, do not possess tusks or postfrontal teeth. The only species identified in the family Myosauridae is the Myosaurus gracilis, or M. gracilis. It should be recognized that the Myosaurus is almost always referred to as the M. gracilis in scientific research.
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.
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.
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.
Sinophoneus is an extinct genus of carnivorous dinocephalian therapsid belonging to the family Anteosauridae. It lived 272 to 270 million years ago at the beginning of the Middle Permian in what is now the Gansu Province in northern China. It is known by a skull of an adult individual, as well as by many skulls of juvenile specimens. The latter were first considered as belonging to a different animal, named Stenocybus, before being reinterpreted as immature Sinophoneus. Sinophoneus shows a combination of characters present in other anteosaurs. Its bulbous profile snout and external nostrils located in front of the canine are reminiscent of the basal anteosaur Archaeosyodon, while its massive transverse pterygoids processes with enlarged distal ends are more similar to the more derived anteosaurs Anteosaurus and Titanophoneus. First phylogenetic analyzes identified Sinophoneus as the most basal Anteosaurinae. A more recent analysis positioned it outside the Anteosaurinae and Syodontinae subclades, and recovers it as the most basal Anteosauridae.
Kunpania is an extinct genus of dicynodont therapsid from the Quanzijie Formation of Xinjiang, China. The type and only species is K. scopulusa, and it is known only by a single incomplete specimen including parts of the skull and forelimb. Since its initial description in 1978 by palaeontologist Ailing Sun, it has sometimes been considered to be another species of Dicynodon by other researchers, or potentially undiagnostic. However, a redescription in 2021 reaffirmed its distinctiveness, including a uniquely well developed muscle attachment on the humerus. Kunpania is perhaps the oldest known member of the derived dicynodont group Dicynodontoidea, potentially dating to the Middle Permian period during the Capitanian, and so may fill a knowledge gap in the history of dicynodont evolution.
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.
Taoheodon is an extinct genus of dicynodont therapsid from the Sunjiagou Formation in the Shanxi province of China, dated to the Wuchiapingian age of the Late Permian. Its type and only known species is T. baizhijuni. Taoheodon was a close relative of the well known Dicynodon, and may represent a biogeographical link between the South African Dicynodon and similar dicynodonts found in Laos.
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. 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 specimen from the older underlying Eodicynodon Assemblage Zone. 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.
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.