Gomphodonts Temporal range: Early - Late Triassic, | |
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Skull of the gomphodont Diademodon tetragonus | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Cynodontia |
Clade: | † Cynognathia |
Clade: | † Gomphodontia Seeley, 1895 |
Subgroups | |
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Gomphodontia is a clade of cynognathian cynodonts that includes the families Diademodontidae, Trirachodontidae, and Traversodontidae. Gomphodonts are distinguished by wide and closely spaced molar-like postcanine teeth, which are convergent with those of mammals. Other distinguishing characteristics of gomphodonts include deep zygomatic arches, upper postcanines with three or more cusps spanning their widths and lower postcanines with two cusps spanning their widths. [2] Gomphodonts first appeared in the Early Triassic and became extinct at the end of the Late Triassic. Fossils are known from southern Africa, Argentina and southern Brazil (Paleorrota geopark), eastern North America, Europe, China, and Antarctica.
Gomphodontia was first named by paleontologist Harry Seeley in 1895. [3] He considered it an order of wide-toothed therapsids (then called anomodonts) from South Africa, distinct from Cynodontia. By the 1930s Gomphodontia was considered a suborder of Cynodontia and included the families Diademodontidae, Trirachodontidae, Traversodontidae, and Tritylodontidae. [4] These four families have also been grouped in the superfamily Traversodontoidea, named by paleontologist Edward Drinker Cope in 1884. Tritylodontoidea has occasionally replaced Gomphodontia in various cynodont taxonomies. In 2001 Gomphodontia was defined as a stem-based clade including all cynodonts more closely related to Exaeretodon than to Cynognathus . [5] This placed it within the larger clade Cynognathia, one of the two main groups of eucynodonts (the other being Probainognathia). Since then most studies have removed Tritylodontidae from Gomphodontia and reclassified it within Probainognathia as a group more closely related to mammals than are the convergently similar gomphodonts. Tritylodontoidea has fallen into disuse while Gomphodontia continues to be used in many studies.
Below is a cladogram from Ruta, Botha-Brink, Mitchell and Benton (2013) showing one hypothesis of gomphodont relationships: [6]
Cynognathia |
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The cynodonts are a clade of eutheriodont therapsids that first appeared in the Late Permian, and extensively diversified after the Permian–Triassic extinction event. Cynodonts occupied a variety of ecologies, including as carnivores and herbivores. Mammals are cynodonts, as are their extinct ancestors and close relatives, having evolved from advanced probainognathian cynodonts during the Late Triassic. All other cynodont lines went extinct, with the last known non-mammalian cynodont group, the Tritylodontidae, having its youngest records in the Early Cretaceous.
Cynognathus is an extinct genus of large-bodied cynodontian therapsids that lived in the Middle Triassic. It is known from a single species, Cynognathus crateronotus. Cynognathus was a 1.2-metre long predator closely related to mammals and had a southern hemispheric distribution. Fossils have so far been recovered from South Africa, Argentina, Antarctica, and Namibia.
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.
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 mass extinction event, its survival during the extinction may have been due to its burrowing habits.
Tritylodon is an extinct genus of tritylodonts, one of the most advanced group of cynodont therapsids. They lived in the Early Jurassic and possibly Late Triassic periods along with dinosaurs. They also shared many characteristics with mammals, and were once considered mammals because of overall skeleton construction. That was changed due to them retaining the vestigial reptilian jawbones and a different skull structure. Tritylodons are now regarded as non-mammalian synapsids.
The theriodonts are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts.
Cynognathia is one of two major clades of cynodonts, the other being Probainognathia. Cynognathians included the large carnivorous genus Cynognathus and the herbivorous traversodontids. Cynognathians can be identified by several synapomorphies including a very deep zygomatic arch that extends above the middle of the orbit.
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 basal probainognathian is the Middle Triassic (Anisian) aged Lumkuia, from South Africa, though probainognathians would not become prominent until the mid Norian stage of the Late Triassic. Three groups survived the extinction at the end of Triassic: Tritheledontidae and Tritylodontidae, which both survived until the Jurassic—the latter even into the Cretaceous —and Mammaliaformes, which includes the mammals.
Tritylodontidae is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, bearing several mammalian traits like erect limbs, endothermy and details of the skeleton. They were the last-known family of the non-mammaliaform synapsids, persisting into the Early Cretaceous.
Traversodontidae is an extinct family of herbivorous cynodonts. Traversodonts were primarily Gondwanan, with many species known from Africa and South America. Recently, traversodonts have also been found from Europe and eastern North America. Traversodonts first appeared in the Middle Triassic and diversified in the Late Triassic before going extinct at the end of the epoch. The family Traversodontidae was erected by Friedrich von Huene in 1936 for cynodonts first found in São Pedro do Sul in Paleorrota, Brazil.
Lumkuia is an extinct genus of cynodonts, fossils of which have been found in the Cynognathus Assemblage Zone of the Beaufort Group in the South African Karoo Basin that date back to the early Middle Triassic. It contains a single species, Lumkuia fuzzi, which was named in 2001 on the basis of the holotype specimen BP/1/2669, which can now be found at the Bernard Price Institute in Johannesburg, South Africa. The genus has been placed in its own family, Lumkuiidae. Lumkuia is not as common as other cynodonts from the same locality such as Diademodon and Trirachodon.
Trirachodontidae is an extinct, possibly paraphyletic family of cynognathian cynodonts from the Triassic of China and southern Africa. Trirachodontids appeared during the Early Triassic soon after the Permian-Triassic extinction event and quickly spread over a wide geographic area in a comparatively brief amount of time from 250 to 237 million years ago.
Pascualgnathus is an extinct genus of traversodontid cynodonts from the Middle Triassic of Argentina. Fossils have been found from the Río Seco de la Quebrada Formation of the Puesto Viejo Group. The type species P. polanskii was named in 1966.
Arctotraversodon is an extinct genus of traversodontid cynodonts from the Late Triassic of Canada. Fossils first described from the Wolfville Formation in Nova Scotia in 1984 represented the first known traversodontid from North America. The type and only species is A. plemmyridon and is represented by teeth and several dentary bones.
Boreogomphodon is an extinct genus of traversodontid cynodonts from the Late Triassic of the eastern United States. Fossils have been found from the Turkey Branch Formation in Virginia.
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.
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.
Etjoia is an extinct genus of traversodontid cynodonts that lived during the Middle Triassic or Late Triassic period in southern Africa. This medium-sized omnivorous cynognathian provides important information on the dental evolution of early diverging gomphodonts and traversodontids.
Scalenodontoides is an extinct genus of traversodontidae, a family of herbivorous cynodonts. It lived during the Late Triassic in what is now South Africa. Its type species is Scalenodontoides macrodontes. It was named in 1957 by A. W. Crompton and F. Ellenberger. Arctotraversodon plemmyrodon was originally classified as a species of Scalenodontoides, but was given its own genus in 1992. It is found in the Scalenodontoides Assemblage Zone of the Elliot Formation, which is named for it. It is one of the geologically youngest traversodontids, alongside the putative traversodontid Boreogomphodon. It is closely related to Exaeretodon and Siriusgnathus, but is distinguished by the presence of a shelf-like expansion of its parietal called the nuchal table. Though the largest known complete skull is only 248 millimetres (9.8 in) long, it may have been the largest non-mammaliaform cynodont, as an incomplete snout would have belonged to a specimen with an estimated skull length of 617 millimetres (24.3 in).