Santacruzgnathus Temporal range: | |
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Photos and drawings of the holotype | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | Prozostrodontia |
Genus: | † Santacruzgnathus Martinelli et al., 2016 |
Species: | †S. abdalai |
Binomial name | |
†Santacruzgnathus abdalai Martinelli et al., 2016 | |
Santacruzgnathus is an extinct genus of small cynodonts from the Late Triassic (Carnian) Santacruzodon Assemblage Zone of Brazil. It contains one species, S. abdalai. Santacruzgnathus is known from a single partial lower jaw with four postcanine teeth, only one of which is well-preserved. Some features of the specimen, including the slender shape of the jaw and the incipiently double-rooted teeth, indicate that the animal was an early member of Prozostrodontia, a group that includes mammals and their close relatives.
The holotype and only known specimen of Santacruzgnathus (UFRGS-PV-1121-T) consists of a partial right dentary bone with a well-preserved final postcanine tooth and fragments of three other postcanines. The specimen was discovered at the Schoenstatt site near the town of Santa Cruz do Sul within the state of Rio Grande do Sul. The outcrop in which the specimen was found belongs to the Santa Cruz Sequence of the Santa Maria Supersequence, which has been dated to the early Carnian age of the Late Triassic. The specimen was first described in a 2011 paper by Soares et al. They interpreted it as a juvenile, and provisionally assigned it to cf. Probainognathus due to similarities with Probainognathus juveniles from the Chañares Formation of Argentina, as well as a supposed juvenile Probainognathus specimen from the Argentine Ischigualasto Formation. [1] [2]
The specimen was redescribed in a 2016 paper by Martinelli et al. They identified it as a new species of cynodont unrelated to Probainognathus, which was given the name Santacruzgnathus abdalai. The generic name derives from Santa Cruz do Sul where the holotype was found, and the Neo-Latin word gnathus meaning a lower jaw. The specific epithet abdalai references the Argentine palaeontologist Fernando Abdala. The authors reinterpreted the holotype as a likely subadult or adult; the last postcanine is fully erupted in the specimen, which is usually not the case in juvenile cynodonts. [2]
While only a small portion of the dentary is preserved, the bone appears to have been quite slender, as was the case with many early prozostrodontians. On the lingual (inner) side near the lower edge of the bone, a thin meckelian groove can be seen. Behind the tooth row, most of the coronoid process has broken off with only the base remaining. On the lingual side of the coronoid process, part of the coronoid bone has been preserved. [2]
Of the four teeth preserved in the specimen, the first two are represented only by fragmentary roots, while the third also preserves part of the base of the crown. Unlike the others, the fourth tooth is nearly complete. The crown bears a large central cusp (cusp a), a small front cusp (cusp b), a rear cusp (cusp c) somewhat larger than cusp b, and an even smaller cusp d behind cusp c. Cusp a is symmetrical and not recurved. The tip of cusp b is broken off. [1] Cusp b is shifted somewhat towards the labial (outer) side of the tooth, and is separated from cusp a by a deep groove. On the lingual side of the tooth, there is a discontinuous cingulum divided into mesiolingual (front) and distolingual (back) portions. The mesiolingual cingulum has a crenulated edge but no distinct cusps. The root is not fully divided, but has a prominent constriction on the labial side, as seen in most early prozostrodontians. [2]
Based on features of its jaw and teeth, Martinelli et al. (2016) interpreted Santacruzgnathus as a member of Prozostrodontia, a group of probainognathian cynodonts that includes mammals and close relatives of them. The genus was however not included in any phylogenetic analysis in that paper. [2] In 2017, the prozostrodontian Alemoatherium was described by Martinelli et al. Three phylogenetic analyses were performed, two of which included Santacruzgnathus. The resulting cladograms placed Santacruzgnathus within a large polytomy at the base of Prozostrodontia. The cladogram from the third analysis is shown below: [3]
Probainognathia |
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Probainognathidae is an extinct family of insectivorous cynodonts which lived in what is now South America during the Middle to Late Triassic. The family was established by Alfred Romer in 1973 and includes two genera, Probainognathus from the Chañares Formation of Argentina and Bonacynodon from the Dinodontosaurus Assemblage Zone of Brazil. Probainognathids were closely related to the clade Prozostrodontia, which includes mammals and their close relatives.
Probainognathus meaning “progressive jaw” is an extinct genus of cynodonts that lived around 235 to 221.5 million years ago, during the Late Triassic in what is now Argentina. Together with the genus Bonacynodon from Brazil, Probainognathus forms the family Probainognathidae. Probainognathus was a relatively small, carnivorous or insectivorous cynodont. Like all cynodonts, it was a relative of mammals, and it possessed several mammal-like features. Like some other cynodonts, Probainognathus had a double jaw joint, which not only included the quadrate and articular bones like in more basal synapsids, but also the squamosal and surangular bones. A joint between the dentary and squamosal bones, as seen in modern mammals, was however absent in Probainognathus.
Chiniquodon is an extinct genus of carnivorous cynodonts, which lived during the Late Triassic (Carnian) in South America and Africa. Chiniquodon was closely related to the genus Aleodon, and close to the ancestry of mammals.
Thrinaxodon is an extinct genus of cynodonts, including the species T. liorhinus which lived in what are now South Africa and Antarctica during the Early Triassic. Thrinaxodon lived just after the Permian–Triassic mass extinction event, its survival during the extinction may have been due to its burrowing habits.
Galesaurus is an extinct genus of carnivorous cynodont therapsid that lived between the Induan and the Olenekian stages of the Early Triassic in what is now South Africa. It was incorrectly classified as a dinosaur by Sir Richard Owen in 1859.
Prozostrodon is an extinct genus of probainognathian cynodonts that was closely related to mammals. The remains were found in Brazil and are dated to the Carnian age of the Late Triassic. The holotype has an estimated skull length of 6.7 centimetres (2.6 in), indicating that the whole animal may have been the size of a cat. The teeth were typical of advanced cynodonts, and the animal was probably a carnivore hunting reptiles and other small prey.
Brasilodon is an extinct genus of small, mammal-like cynodonts that lived in what is now Brazil during the Norian age of the Late Triassic epoch, about 225.42 million years ago. While no complete skeletons have been found, the length of Brasilodon has been estimated at 12 centimetres (4.7 in). Its dentition shows that it was most likely an insectivore. The genus is monotypic, containing only the species B. quadrangularis. Brasilodon belongs to the family Brasilodontidae, whose members were some of the closest relatives of mammals, the only cynodonts alive today. Two other brasilodontid genera, Brasilitherium and Minicynodon, are now considered to be junior synonyms of Brasilodon.
Cromptodon is an extinct genus of cynodonts from the Triassic of Cerro Bayo de Portrerillos, Cerro de las Cabras Formation, Argentina, South America. It is known only from PVL 3858, a mandible.
Pachygenelus is an extinct genus of tritheledontid cynodonts. Fossils have been found from the Karoo basin in South Africa and date back to the Early Jurassic.
Plinthogomphodon is an extinct genus of traversodontid cynodonts from the Late Triassic of the eastern United States. Fossils have been found from the Deep River basin of North Carolina, part of the larger Newark Supergroup. It is known from a single eroded snout. The type and only species is P. herpetairus.
Candelariodon is an extinct genus of carnivorous probainognathian cynodonts from the Middle to Late Triassic Santa Maria Formation of the Paraná Basin in Rio Grande do Sul state, Brazil. Candelariodon is known from a partial mandible having some complete teeth. It was first named by Téo Veiga de Oliveira, Cesar Leandro Schultz, Marina Bento Soares and Carlos Nunes Rodrigues in 2011 and the type species is Candelariodon barberenai.
Cricodon is an extinct genus of trirachodontid cynodonts that lived during the Early Triassic and Middle Triassic periods of Africa. A. W. Crompton named Cricodon based on the ring-like arrangement of the cuspules on the crown of a typical postcanine tooth. The epithet of the type species, C. metabolus, indicates the change in structure of certain postcanines resulting from replacement.
Alemoatherium is an extinct genus of prozostrodontian cynodont which lived in the Late Triassic of Brazil. It contains a single species, A. huebneri, named in 2017 by Agustín Martinelli and colleagues. The genus is based on UFSM 11579b, a left lower jaw (dentary) found in the Alemoa Member of the Santa Maria Formation, preserving the late Carnian-age Hyperodapedon Assemblage Zone. Alemoatherium was among the smallest species of cynodonts found in the rich synapsid fauna of the Santa Maria Formation. Its blade-like four-cusped postcanine teeth show many similarities with those of dromatheriids, an obscure group of early prozostrodontians.
Aleodon is an extinct genus of cynodonts that lived from the Middle to the Late Triassic. Relatively few analyses have been conducted to identify the phylogenetic placement of Aleodon, however those that have place Aleodon as a sister taxon to Chiniquodon. Two species of Aleodon are recognized: A. brachyramphus which was discovered in Tanzania, and A. cromptoni which was discovered most recently in Brazil.
Abdalodon is an extinct genus of late Permian cynodonts, known by its only species A. diastematicus.Abdalodon together with the genus Charassognathus, form the clade Charassognathidae. This clade represents the earliest known cynodonts, and is the first known radiation of Permian cynodonts.
Dromatheriidae is an extinct family of prozostrodontian cynodonts, closely related to mammals. Members of the family are known from the Late Triassic of India, Europe and North America. Apart from a few jaw fragments, dromatheriids are mainly known from their sectorial (flesh-slicing) postcanine teeth. The teeth were fairly typical among early prozostrodontians, as they were labiolingually compressed, with a single root and crown hosting a longitudinal row of sharp cusps. Dromatheriids in particular have a very narrow and symmetrical crown without a prominent cingulum.
Pseudotherium is an extinct genus of prozostrodontian cynodonts from the Late Triassic of Argentina. It contains one species, P. argentinus, which was first described in 2019 from remains found in the La Peña Member of the Ischigualasto Formation in the Ischigualasto-Villa Unión Basin.
Bonacynodon is an extinct genus of cynodonts that lived in what is now southern Brazil during the Triassic period. The genus is monotypic, containing only the type species Bonacynodon schultzi. B. schultzi is known from two specimens, consisting of two partial skulls and some badly preserved parts of the postcranium. Both specimens were recovered from the Pinheiros-Chiniquá Sequence, part of the Santa Maria Supersequence of the Paraná Basin. This sequence preserves a faunal association known as the Dinodontosaurus Assemblage Zone, which contains numerous other species of cynodonts, dicynodonts and reptiles. Bonacynodon was a small, likely insectivorous cynodont, whose length has been estimated at around 30 centimetres (12 in). It can be distinguished from other cynodonts by its large, serrated (saw-like) canine teeth. Together with the genus Probainognathus of Argentina, it made up the family Probainognathidae, one of the earliest-diverging lineages of the clade Probainognathia. It was a fairly close relative of mammals, the only group of cynodonts alive today.
Vetusodon is an extinct genus of cynodonts belonging to the clade Epicynodontia. It contains one species, Vetusodon elikhulu, which is known from four specimens found in the Late Permian Daptocephalus Assemblage Zone of South Africa. With a skull length of about 18 centimetres (7.1 in), Vetusodon is the largest known cynodont from the Permian. Through convergent evolution, it possessed several unusual features reminiscent of the contemporary therocephalian Moschorhinus, including broad, robust jaws, large incisors and canines, and small, single-cusped postcanine teeth.
Kalaallitkigun is an extinct genus of haramiyidan mammaliaforms from the Late Triassic of Greenland. It contains a single species, Kalaallitkigun jenkinsi, which was described in 2020 from a partial dentary found in the Fleming Fjord Formation. More specifically, it was found in the mid-late Norian Carlsberg Fjord beds of the Ørsted Dal Member. It is the oldest of several mammaliaform species discovered in the Late Triassic sediments of Greenland. It is also the oldest mammaliaform with double-rooted teeth, and its pattern of tooth cusps help to clarify the evolution of haramiyidan teeth relative to their morganucodont-like ancestors.