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''Biseridens'' ("two rows of teeth") is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
anomodont 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.Chinsamy-Turan, A. (2011) ''Forerunners of Mammals: Ra ...
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, 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 ...
, 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 The Qilian Mountains (, also romanized as Tsilien; Mongghul: Chileb), together with the Altyn-Tagh (Altun Shan) also known as Nan Shan (, literally "Southern Mountains"), as it is to the south of Hexi Corridor, is a northern outlier of the Kunlu ...
of Gansu,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, in 2009 that clarified its relationships to anomodonts, such as the
dicynodonts Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typicall ...
. Among
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
taxa, the therapsid clade
anomodont 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.Chinsamy-Turan, A. (2011) ''Forerunners of Mammals: Ra ...
ia in which ''Biseridens'' belong has one of the largest population sizes, highest levels of diversity, and longest stratigraphic range (Middle
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
to the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, possibly into the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
), as well as being one of the only clades known from all the continents. 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 In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
, and presence of a septomaxilla that distinctly lacks a facial exposure between the nasal and
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
. 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 A palatine or palatinus (in Latin; plural ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman times.
and
pterygoid Pterygoid, from the Greek for 'winglike', may refer to: * Pterygoid bone, a bone of the palate of many vertebrates * Pterygoid processes of the sphenoid bone ** Lateral pterygoid plate ** Medial pterygoid plate * Lateral pterygoid muscle * Medi ...
, articulation between the opisthotic bone and the
tabular Table may refer to: * Table (furniture), a piece of furniture with a flat surface and one or more legs * Table (landform), a flat area of land * Table (information), a data arrangement with rows and columns * Table (database), how the table data ...
bones on the posterior surface of the skull, the absence of the
mandibular foramen The mandibular foramen is an opening on the internal surface of the ramus of the mandible. It allows for divisions of the mandibular nerve and blood vessels to pass through. Structure The mandibular foramen is an opening on the internal surfa ...
on the lower jaw, and a pterygoid in which the transverse flange of the pterygoid has a laterally extending process but lacks posterior ramus.


Etymology

- is Latin for “double rows”; -dens originated from Greek “” for “teeth”


Geology and paleoenvironment

At a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
level, anomodonts were terrestrial
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
of all sizes that dominated the Late
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
. The most dominant sub-clade were
dicynodonts Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivorous animals with a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typicall ...
, more derived relatives of ''Biseridens''. Most anomodonts are postulated to have been
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
, and they were the only herbivorous taxa to survive the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
. 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. A feature that extends across most of the anomodont clade is jaw articulation that allowed for anterior-posterior movement of jaw for slicing. On the basis of converging faunal evidence and comparison to other regions, the Xidagou Formation has been considered to be Middle Permian and home to the oldest therapsid fauna, suggesting that anomodonts had a
Laurasian Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
rather than a
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
n origin.


Discovery

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 The Qingtoushan Formation is a Middle Permian-age geologic formation in the Qilian Mountains of Gansu, China. It is known for its diverse tetrapod fauna known as the Dashankou fauna, which likely dates to the Roadian, and includes some of the ol ...
of
Gansu, China Gansu (, ; alternately romanized as Kansu) is a province in Northwest China. Its capital and largest city is Lanzhou, in the southeast part of the province. The seventh-largest administrative district by area at , Gansu lies between the Tibe ...
. 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. The more recently discovered specimen includes a nearly complete skull (lacking the
occiput The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cereb ...
) with lower jaws, found with a set of 14 articulated
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
. This new specimen was preserved in
mudrock Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone, slate, and shale. Most of the particles of which the stone is composed are less than and are too ...
and also discovered in the upper part of Xidagou Formation in Dashankau, Yumen in Gansu.


Description

''Biseridens'' is known mostly from
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
and jaw material, as well as a series of 14
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
associated with one skull. Like other anomodonts, the region of the skull in front of the eyes is relatively short compared to other
therapsids Therapsida is a major group of eupelycosaurian synapsids that includes mammals, 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 mor ...
. The
orbits In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as a p ...
are large and rounded, and the
temporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets ...
are large and wide and broad
zygomatic arches In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygomati ...
, similar to but not as developed as those of later dicynodonts. The skull is estimated to be at least long, relatively small compared to other basal anomodonts.


Dentition

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 In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
and
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
, and a broad spread of teeth on the
pterygoid Pterygoid, from the Greek for 'winglike', may refer to: * Pterygoid bone, a bone of the palate of many vertebrates * Pterygoid processes of the sphenoid bone ** Lateral pterygoid plate ** Medial pterygoid plate * Lateral pterygoid muscle * Medi ...
and
palatine A palatine or palatinus (in Latin; plural ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman times.
. There are also denticles on the
vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right max ...
, palatine, and pterygoid. The postcanies 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.


Palate

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.


Skull

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 An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets ...
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 Table may refer to: * Table (furniture), a piece of furniture with a flat surface and one or more legs * Table (landform), a flat area of land * Table (information), a data arrangement with rows and columns * Table (database), how the table data ...
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 In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
. 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.


Classification

''Biseridens'' belong to the clade
anomodont 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.Chinsamy-Turan, A. (2011) ''Forerunners of Mammals: Ra ...
which is nested within the clade
therapsids Therapsida is a major group of eupelycosaurian synapsids that includes mammals, 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 mor ...
, often called  “mammal-like reptiles”. Therapsids are a group of advanced synapsids, one of the major branches of amniotes that gave rise to
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
. 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 Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
''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. 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, as well as those of Cisneros and colleagues (2011) and Kammerer and colleagues (2013): 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.


See also

*
List of therapsids This list of therapsids is an attempt to create a comprehensive listing of all genera that have ever been included in the Therapsida excluding mammals and purely vernacular terms. The list includes all commonly accepted genera, but also genera tha ...


References

{{Taxonbar, from=Q2154061 Anomodont genera Guadalupian synapsids Permian synapsids of Asia Fossil taxa described in 1997 Guadalupian genus first appearances Guadalupian genus extinctions