''Vetusodon'' 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 of

cynodont The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variety ...

s belonging to the clade

Epicynodontia Epicynodontia is a clade of cynodont therapsids that includes most cynodonts, such as galesaurids, thrinaxodontids, and Eucynodontia (including mammals). It was erected as a stem-based taxon by Hopson and Kitching (2001) and defined as the mo ...

. It contains one species, ''Vetusodon elikhulu'', which is known from four specimens found in the

Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...

''Daptocephalus'' Assemblage Zone of South Africa. With a skull length of about , ''Vetusodon'' is the largest known cynodont from the Permian. Through

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...

, it possessed several unusual features reminiscent of the contemporary

therocephalia Therocephalia is an extinct suborder of eutheriodont therapsids (mammals and their close relatives) from the Permian and Triassic. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their te ...

n ''

Moschorhinus ''Moschorhinus'' is an extinct genus of therocephalian in the family Akidnognathidae with only one species: ''M. kitchingi''. It was a carnivorous synapsid which has been found in the Late Permian to Early Triassic of the South African Karoo Supe ...

'', including broad, robust jaws, large

incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...

s and

canines Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...

, and small, single- cusped postcanine teeth.

Discovery and naming

''Vetusodon'' is known from four specimens, all of which have been collected in South Africa from rocks belonging to the ''Daptocephalus'' Assemblage Zone. The

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

, BP/1/7971, consists of a well-preserved skull missing the lower jaw. It was discovered in 2010 in a

mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...

bed in the province of

KwaZulu-Natal KwaZulu-Natal (, also referred to as KZN and known as "the garden province") is a province of South Africa that was created in 1994 when the Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu) and Natal Province were merged. It is locate ...

by a team led by the palaeontologist Bruce S. Rubidge. The referred specimen CGP GHG141 was found in 1985 by Gideon Groenewald in the Thaba Nchu Mountain of the Free State province, and like the holotype, it consists of a partial skull missing the lower jaw. The specimen SAM-PK-K10596 is the least complete of the specimens, consisting only of a snout. It was found in 1958 or 1959 by the South African palaeontologist Alfred W. Crompton. The specimen SAM-PK-K10702 is the most complete of the four specimens, and comprises a complete skull with an intact lower jaw. It was discovered in 2009 or 2010 by Derik Wolvaardt at the Ripplemead Farm, close to the town of

Nieu-Bethesda Nieu-Bethesda (Afrikaans for ''New Bethesda'') is a village in the Eastern Cape at the foot of the Sneeuberge, approximately north of Graaff Reinet. It was founded in 1875 as a church town, like many other Karoo villages, and attained municipa ...

Eastern Cape The Eastern Cape is one of the provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha. The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in ...

province. ''Vetusodon elikhulu'' was first described in 2019 by the palaeontologists Fernando Abdala, Leandro C. Gaetano, Roger M. H. Smith and Bruce S. Rubidge. Its generic name is derived 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 ...

word , meaning "old", and the

Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...

word (), meaning "tooth". The

specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...

''elikhulu'' means "large" in the

Zulu language Zulu (), or isiZulu as an endonym, is a Southern Bantu language of the Nguni branch spoken in Southern Africa. It is the language of the Zulu people, with about 12 million native speakers, who primarily inhabit the province of KwaZulu-Natal o ...

, and alludes to the animal's size.

Description

''Vetusodon'' is the largest known Permian cynodont, based on its skull length of around . The temporal region (area behind the

eye socket In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...

s) made up around 60 percent of the skull length. The jaws were quite robust, and the snout was unusually broad compared to other cynodonts of its time. Between the

nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...

s was a bony internarial process that was formed by the

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 ...

e and

nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...

s. This process was somewhat forwards-pointing, giving the tip of the snout a pointed shape when seen from above. The back portion of the nostrils were formed by small bones known as the septomaxillae; these bones bore a thin extension (the intranarial process) that nearly divided each nostril into two halves. There was a small hole (the septomaxillary foramen) where the septomaxilla met with the

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 nasal bones made up the upper part of the snout, and extended backwards as far as the rear end of the

eye sockets In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...

, where they met with the

frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...

s. The sides of the snout were largely formed by the maxillae. The maxilla bore two rows of

foramina In anatomy and osteology, a foramen (;Entry "foramen"
in

(holes) with a diameter of around . The lower jaw was formed primarily by the

dentary bone 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

s, whereas the other (postdentary) bones were reduced in size to an extent not otherwise seen in cynodonts this primitive; as a result, the dentary stretched nearly as far back as the jaw joint. The

symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing together ...

(joint) between the two halves of the lower jaw was quite tall, creating a

chin The chin is the forward pointed part of the anterior mandible (List_of_human_anatomical_regions#Regions, mental region) below the lower lip. A fully developed human skull has a chin of between 0.7 cm and 1.1 cm. Evolution The presence of a we ...

-like structure. Abdala ''et al.'' (2019) interpreted the symphysis as being fused, but a 2020 study by Huttenlocker and Sidor reinterpreted it as unfused. The back portion of the dentary had a projection called the

coronoid process The Coronoid process (from Greek , "like a crown") can refer to: * The coronoid process of the mandible, part of the ramus mandibulae of the mandible * The coronoid process of the ulna The coronoid process of the ulna is a triangular process proj ...

, which extended as high as the upper half of the eye sockets. The coronoid process fit quite closely to the

postorbital bar The postorbital bar (or postorbital bone) is a bony arched structure that connects the frontal bone of the skull to the zygomatic arch, which runs laterally around the eye socket. It is a trait that only occurs in mammalian taxa, such as most streps ...

(the bony structure that divided the eye socket from 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, ...

). The teeth were conical, single-rooted and had no

serration Serration is a saw-like appearance or a row of sharp or tooth-like projections. A serrated cutting edge has many small points of contact with the material being cut. By having less contact area than a smooth blade or other edge, the applied pr ...

s. ''Vetusodon'' possessed four pairs of upper

s, which were long and recurved. There was a small gap (

diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...

) between the last incisor and the canine. The upper canines were large and had a round, somewhat elongated cross-section. Behind the canines were a set of relatively small postcanine teeth. The upper postcanines varied in number between specimens, ranging from 7 pairs in the holotype, to 11 in the specimen SAM-PK-K10596. Unlike most cynodonts, which have postcanines with multiple cusps, the postcanines of ''Vetusodon'' only bore one cusp each. The outer rims of the eye sockets (orbits) were formed by the

lacrimal bone The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...

s (in the front), prefrontal and

postorbital bone The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...

s (at the top), and the

jugal bone The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...

s (in the back and on the bottom). Between the eye sockets there was a large gap in the skull known as the interorbital vacuity. The

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 ...

(the cheek bones, which made up the outer edge of the temporal fenestrae) were formed by the jugal and

squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal The quadratojugal is a skull bone present in many vertebrates, including ...

bones, and were quite tall, particularly towards the back. The upper edge of the zygomatic arch was rather thick, whereas the lower edge was thinner. Like other non-

probainognathia Probainognathia is one of the two major subgroups of the clade Eucynodontia, the other being Cynognathia. The earliest forms were carnivorous and insectivorous, though some groups eventually also evolved herbivorous diets. The earliest and most ...

n cynodonts, ''Vetusodon'' possessed a hole known as the

pineal foramen A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhythm ...

at the top of the skull, between the temporal fenestrae. Running from the top of the orbits towards the pineal foramen, and bordering the temporal fenestrae, were two crests formed by the postorbital bones. Behind the pineal foramen there was a longitudinal ridge known as the

sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...

, which was unusually short in ''Vetusodon''. Behind the sagittal crest, two occipital crests, formed by 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 squamosal bones, stretched backwards diagonally and joined the rear ends of the zygomatic arches. The

secondary palate The secondary palate is an anatomical structure that divides the nasal cavity from the oral cavity in many vertebrates. In human embryology, it refers to that portion of the hard palate that is formed by the growth of the two palatine shelves medi ...

(the structure making up the roof of the mouth in most cynodonts) was formed by the premaxillae, maxillae and

palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the throat. Together with the maxillae, they comprise the hard palate. (''Palate'' is derived from the Latin ''pa ...

s. In ''Vetusodon'', the two halves of the secondary palate were not joined together in the middle, rendering 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 maxill ...

prominently visible from below. Depending on the

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

position of ''Vetusodon'', this incomplete secondary palate may either be an

autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...

(derived trait) of this taxon, or a

plesiomorphy In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

(ancestral trait) shared with other primitive cynodonts.

Classification

When ''Vetusodon'' was first described in 2019, it was included in a

phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

to test its relationships to other cynodonts. It was found to be a derived member of the clade

, being more closely related to

Eucynodontia Eucynodontia ("true dog teeth") is a clade of cynodont therapsids including mammals and most non-mammalian cynodonts. The oldest eucynodonts are known from the Early Triassic and possibly Late Permian. Eucynodontia includes two major subgroups, ...

than several

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 ...

genera such as ''

Thrinaxodon ''Thrinaxodon'' is an extinct genus of cynodonts, most commonly regarded by its species ''T. liorhinus'' which lived in what are now South Africa and Antarctica during the Early Triassic. ''Thrinaxodon'' lived just after the Permian–Triassic m ...

''. As ''Thrinaxodon'' and its relatives had relatively well-developed secondary palates, this would imply that the incomplete secondary palate of ''Vetusodon'' was secondarily reduced from a more complete one. However, a 2020 study by Huttenlocker and Sidor, followed by a 2021 study by Pusch and colleagues, recovered ''Vetusodon'' in a more early-diverging position within Epicynodontia, as the sister taxon of the Permian genus ''

Cynosaurus ''Cynosaurus'' is an extinct genus of cynodonts. Remains have been found from the ''Dicynodon'' Assemblage Zone in South Africa.T. S. Kemp: ''The Origin and Evolution of Mammals'' Oxford University Press, 2005. ''Cynosaurus'' was first describe ...

''. If this placement is correct, the incomplete secondary palate of ''Vetusodon'' may be an ancestral trait inherited from earlier cynodonts. Below are two

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

s from Abdala ''et al.'' (2019) and Pusch ''et al.'' (2021), which illustrate the varying phylogenetic placements of ''Vetusodon'':

Palaeobiology

The large incisors and canines and small, simple postcanines of ''Vetusodon'' imply that the front teeth played a larger role than the postcanines in the processing of food. In this respect it closely resembled the

gorgonopsia Gorgonopsia (from the Greek Gorgon, a mythological beast, and 'aspect') is an extinct clade of sabre-toothed therapsids from the Middle to Upper Permian roughly 265 to 252 million years ago. They are characterised by a long and narrow skull, a ...

ns and

ns, two other groups of

synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...

s that existed at the time, but differed from most other cynodonts, whose postcanines are generally the most complex and important teeth. The placement of the coronoid process of the dentary close to the postorbital bar may be evidence of a well-developed

temporalis muscle In anatomy, the temporalis muscle, also known as the temporal muscle, is one of the muscles of mastication (chewing). It is a broad, fan-shaped convergent muscle on each side of the head that fills the temporal fossa, superior to the zygomati ...

, which would have allowed for a powerful bite by the front teeth. The skull shape and dentition of ''Vetusodon'' were in general similar to those of the therocephalian ''

'', which it coexisted with. As therocephalians were only distantly related to ''Vetusodon'', this is thought to be an instance of

, where different groups that inhabit the same

ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...

evolve similar traits.

Palaeoenvironment

''Vetusodon'' belongs to the ''Daptocephalus'' Assemblage Zone, a

biozone In biostratigraphy, biostratigraphic units or biozones are intervals of geological strata that are defined on the basis of their characteristic fossil taxa, as opposed to a lithostratigraphic unit which is defined by the lithological properties of ...

that is part of the

Beaufort Group The Beaufort Group is the third of the main subdivisions of the Karoo Supergroup in South Africa. It is composed of a lower Adelaide Subgroup and an upper Tarkastad Subgroup. It follows conformably after the Ecca Group and unconformably underlie ...

, which itself is part of the larger

Karoo Supergroup The Karoo Supergroup is the most widespread stratigraphic unit in Africa south of the Kalahari Desert. The supergroup consists of a sequence of units, mostly of nonmarine origin, deposited between the Late Carboniferous and Early Jurassic, a perio ...

. ''Vetusodon'' belongs to a subzone of the ''Daptocephalus'' AZ known as the ''Lystrosaurus maccaigi-Moschorhinus'' Subzone. In addition to ''Vetusodon'', several other

s are known from this subzone. These include the fellow cynodonts ''

'' and '' Nanictosaurus'', the

biarmosuchian Biarmosuchians are an extinct clade of non-mammalian synapsids from the Permian. They are the most basal group of the therapsids. All of them were moderately-sized, lightly-built carnivores, intermediate in form between basal sphenacodont "pelyco ...

Ictidorhinus ''Ictidorhinus'' is an extinct genus of biarmosuchian therapsids. Fossils have been found from the ''Dicynodon'' Assemblage Zone of the Beaufort Group in the Karoo Basin, South Africa and are of Late Permian age. It had a short snout and proport ...

'', the

dicynodont 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 ...

s ''

Aulacephalodon ''Aulacephalodon'' is an extinct genus of medium-sized dicynodonts, or non-mammalian synapsids, that lived during Permian period, about 299-252 million years ago. Individuals of ''Aulacephalodon'' are commonly found in the Lower Beaufort Group o ...

'', ''

Daptocephalus ''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'' Zone, in the Ka ...

'', ''

Dicynodon ''Dicynodon'' ("two dog-teeth") is a genus of dicynodont therapsid that flourished during the Upper Permian period. Like all dicynodonts, it was herbivorous animal. This reptile was toothless, except for prominent tusks, hence the name. It probab ...

'', ''

Dicynodontoides ''Dicynodontoides'' is a genus of small to medium-bodied, herbivorous, emydopoid dicynodonts from the Late Permian. The name ''Dicynodontoides'' references its “dicynodont-like” appearance (dicynodont = two-dog-tooth) due to the caniniform t ...

'', ''

Diictodon ''Diictodon'' is an extinct genus of pylaecephalid dicynodont. These mammal-like synapsids lived during the Late Permian period, approximately 255 million years ago. Fossils have been found in the ''Cistecephalus'' Assemblage Zone of the Mad ...

'', ''

Dinanomodon ''Dinanomodon'' is a genus of dicynodont from Late Permian (Changhsingian) of the ''Lystrosaurus'' Assemblage Zone, Katberg Formation, and ''Cistecephalus'' Assemblage Zone, Balfour Formation Beaufort Group, Karoo Basin of South Africa ...

'', ''

Emydops ''Emydops'' is an extinct genus of dicynodont therapsid from the Permian of South Africa. ''Emydops'' was first named by South African paleontologist Robert Broom in 1912 when he described ''Emydops minor''. In the following years, the genus gre ...

'', '' Emydorhinus'', '' Kwazulusaurus'', ''

Lystrosaurus ''Lystrosaurus'' (; 'shovel lizard'; proper Greek is λίστρον ''lístron'' ‘tool for leveling or smoothing, shovel, spade, hoe’) is an extinct genus of herbivorous dicynodont therapsids from the late Permian and Early Triassic epochs (a ...

'', ''

Oudenodon ''Oudenodon'' is an extinct genus of dicynodont. It was common throughout southern Africa during the Late Permian. Several species of ''Oudenodon'' are known. Both ''O. bainii'', the type species, and ''O. grandis'' are known from South Africa ...

'', ''

Pelanomodon ''Pelanomodon'' is an extinct genus of dicynodont therapsids that lived in the Late Permian period. Fossil evidence of this genus is principally found in the Karoo Basin of South Africa, in the ''Dicynodon'' Assemblage Zone.Kammerer, C. F., K. D ...

'' and ''

Thliptosaurus ''Thliptosaurus'' (meaning "compressed lizard") 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 ...

'', the gorgonopsians ''

Arctognathus ''Arctognathus'' is an extinct genus of gorgonopsids that throve during the Late Permian in the Karoo basin of what is now South Africa. Discovery A carnivore, like all gorgonopsid, ''Arctognathus'' was given its name ("Bear jaw") in referen ...

'', '' Cyonosaurus'', ''

Lycaenops ''Lycaenops'' ("wolf-face") is a genus of carnivorous therapsids. It lived during the Middle Permian to the early Late Permian, about 260 mya, in what is now South Africa. Description ''Lycaenops'' measured about and weighed up to . Like the ...

'' and ''

Rubidgea ''Rubidgea'' is a genus of Gorgonopsia, gorgonopsid from the upper Permian of South Africa and Tanzania, containing the species ''Rubidgea atrox''. The generic name ''Rubidgea'' is sometimes believed to be derived from the surname of renowned Kar ...

'' and the therocephalians ''

Ictidosuchoides ''Ictidosuchoides'' is an extinct genus of ictidosuchid therocephalians. Fossils have been found from the Karoo Basin in South Africa. The genus is known to have been one of the few therocephalians to have survived the Permian-Triassic extinct ...

'', '' Lycideops'', ''Moschorhinus'', '' Nanictidops'', '' Promoschorhynchus'', '' Polycynodon'' and '' Scaloporhinus''. Other animals include the

bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...

Palaeanodonta Palaeanodonta ("ancient toothless animals") is an extinct clade of stem- pangolins. They were insectivorous, possibly fossorial, and lived from the Early Paleocene to Early Oligocene in North America, Europe and East Asia. While the taxonomic ...

'', the

ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species. The ray-finned fishes are so called because their fins are webs of skin supported by bony or hor ...

'' Atherstonia'' and '' Namaichthys'', the

temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...

s ''

Rhinesuchus ''Rhinesuchus'' (meaning "rasp crocodile" for the ridged surface texture on its skull bones) is a large temnospondyl amphibian. Remains of the genus are known from the Permian of the South African Karoo Basin's ''Tapinocephalus'' and ''Cistec ...

'' and ''

Uranocentrodon ''Uranocentrodon'' is an extinct genus of the family Rhinesuchidae. Known from a skull, ''Uranocentrodon'' was a large predator with a length up to . Originally named ''Myriodon'' by van Hoepen in 1911, it was transferred to a new genus on acco ...

'' and the

reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...

s ''

Owenetta ''Owenetta'' is an extinct genus of owenettid procolophonian parareptile. Fossils have been found from the Beaufort Group in the Karoo Basin of South Africa. Although most procolophonians lived during the Triassic, ''Owenetta'' existed during the ...

'', ''

Pareiasaurus ''Pareiasaurus'' is an extinct genus of pareiasauromorph reptile from the Permian period. It was a typical member of its family, the pareiasaurids, which take their name from this genus. Fossils have been found in the Beaufort Group. Descript ...

'' and '' Spondylolestes''.

References

{{Taxonbar, from=Q104093847 Prehistoric cynodont genera Changhsingian genera Lopingian synapsids of Africa Permian South Africa Fossils of South Africa Fossil taxa described in 2019