Tropidostoma | |
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Tropidostoma dubium (SAM-PK-K11238) in (A) dorsal, (D) ventral, (G) right lateral, and (J) anterior views. Bulbasaurus phylloxyron (CGP/1/938) in (B) dorsal, (E) ventral, (H) right lateral, and (K) anterior views. Specimen has suffered some lateral compression. Scale bars equal 5 cm. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Family: | † Oudenodontidae |
Genus: | † Tropidostoma Owen, 1876 |
Species: | †T. dubium |
Binomial name | |
†Tropidostoma dubium Seeley, 1876 | |
Synonyms | |
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Tropidostoma is a medium-sized herbivorous oudenodontid dicynodont therapsid that lived during the Late Permian (Lopingian) period in South Africa. The first Tropidostoma fossil was described by Harry Govier Seeley in 1889. Later two subspecies were identified. Tropidostoma fossils are an index fossil in a biozone of the Karoo Basin known as the Tropidostoma Assemblage Zone. This biozone is characterized by the presence of this species in association with another dicynodont species, Endothiodon uniseries . [1] [2]
The first Tropidostoma fossil material was found during a field expedition in the Upper Permian-aged Teekloof Formation of the Beaufort Group. This material was later described by Seeley (1889) in a study in which he described two fossils which had been named Dicynodon microtrema and Tropidostoma dunni. In 1915, several years after Seeley's death, the paleontologist, Robert Broom, reexamined the same material and discovered that the fossil material in question was of the same species. This new single species was renamed Tropidostoma microtrema. [3] Some years later, the name T. microtrema was changed to Tropidostoma dubium, and Tropidostoma dunni is now considered to be the type species. [4] [5] [6] [7]
Two subspecies exist among the specimens referred to as Tropidostoma dubium and Tropidostoma dunni . T. dubium is observed to have two cranial morphs, one being robust form with a tall snout and large tusks and the other more gracile with a low snout and small or no tusks. The robust and gracile forms are considered to either represent sexual dimorphism or individual variation. [8]
Tropidostoma is currently classified as an oudenodontid within the larger dicynodont clade Bidentalia. This clade is characterized collectively by their reduced dentition with only their maxillary tusks being intact. However, many species in this clade sporadically lack tusks completely and their fossils only hold evidence that they retained their keratinous beaks. [9] Many Tropidostoma fossils previously collected have been misidentified as other species, such as of Oudenodon bainii due to their remarkable similarity. [7] In addition, the type fossil of the recently discovered Bulbasaurus phylloxyron was misidentified as a Tropidostoma fossil for several years. [10] [11]
Diictodon is an extinct genus of pylaecephalid dicynodont that lived during the Late Permian period, approximately 255 million years ago. Fossils have been found in the Cistecephalus Assemblage Zone of the Madumabisa Mudstone of the Luangwa Basin in Zambia and the Tropidostoma Assemblage Zone of the Teekloof Formation, Tapinocephalus Assemblage Zone of the Abrahamskraal Formation, Dicynodon Assemblage Zone of the Balfour Formation, Cistecephalus Assemblage Zone of the Middleton or Balfour Formation of South Africa and the Guodikeng Formation of China. Roughly half of all Permian vertebrate specimens found in South Africa are those of Diictodon. This small herbivorous animal was one of the most successful synapsids in the Permian period.
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 underlies the Stormberg Group. Based on stratigraphic position, lithostratigraphic and biostratigraphic correlations, palynological analyses, and other means of geological dating, the Beaufort Group rocks are considered to range between Middle Permian (Wordian) to Early Triassic (Anisian) in age.
Emydops is an extinct genus of dicynodont therapsids from the Middle Permian to Late Permian of what is now South Africa. The genus is generally small and herbivorous, sharing the dicynodont synapomorphy of bearing two tusks. In the following years, the genus grew to include fourteen species. Many of these species were erected on the basis of differences in the teeth and the positioning of the frontal and parietal bones. A 2008 study narrowed Emydops down to two species, E. arctatus and the newly described E. oweni.
Dinogorgon is a genus of gorgonopsid from the Late Permian of South Africa and Tanzania. The generic name Dinogorgon is derived from Greek, meaning "terrible gorgon", while its species name rubidgei is taken from the surname of renowned Karoo paleontologist, Professor Bruce Rubidge, who has contributed to much of the research conducted on therapsids of the Karoo Basin. The type species of the genus is D. rubidgei.
The Tapinocephalus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the middle Abrahamskraal Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 2,000 metres (6,600 ft), occur from Merweville and Leeu-Gamka in its southernmost exposures, from Sutherland through to Beaufort West where outcrops start to only be found in the south-east, north of Oudshoorn and Willowmore, reaching up to areas south of Graaff-Reinet. Its northernmost exposures occur around the towns Fraserburg and Victoria West. The Tapinocephalus Assemblage Zone is the second biozone of the Beaufort Group.
The Cistecephalus Assemblage Zone is a tetrapod assemblage zone or biozone found in the Adelaide Subgroup of the Beaufort Group, a majorly fossiliferous and geologically important geological group of the Karoo Supergroup in South Africa. This biozone has outcrops located in the Teekloof Formation north-west of Beaufort West in the Western Cape, in the upper Middleton and lower Balfour Formations respectively from Colesberg of the Northern Cape to east of Graaff-Reinet in the Eastern Cape. The Cistecephalus Assemblage Zone is one of eight biozones found in the Beaufort Group, and is considered to be Late Permian in age.
The Daptocephalus Assemblage Zone is a tetrapod assemblage zone or biozone found in the Adelaide Subgroup of the Beaufort Group, a majorly fossiliferous and geologically important Group of the Karoo Supergroup in South Africa. This biozone has outcrops located in the upper Teekloof Formation west of 24°E, the majority of the Balfour Formation east of 24°E, and the Normandien Formation in the north. It has numerous localities which are spread out from Colesberg in the Northern Cape, Graaff-Reniet to Mthatha in the Eastern Cape, and from Bloemfontein to Harrismith in the Free State. The Daptocephalus Assemblage Zone is one of eight biozones found in the Beaufort Group and is considered Late Permian (Lopingian) in age. Its contact with the overlying Lystrosaurus Assemblage Zone marks the Permian-Triassic boundary.
The Lystrosaurus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the upper Adelaide and lower Tarkastad Subgroups of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. This biozone has outcrops in the south central Eastern Cape and in the southern and northeastern Free State. The Lystrosaurus Assemblage Zone is one of eight biozones found in the Beaufort Group, and is considered to be Early Triassic in age.
The Cynognathus Assemblage Zone is a tetrapod biozone utilized in the Karoo Basin of South Africa. It is equivalent to the Burgersdorp Formation, the youngest lithostratigraphic formation in the Beaufort Group, which is part of the fossiliferous and geologically important Karoo Supergroup. The Cynognathus Assemblage Zone is the youngest of the eight biozones found in the Beaufort Group, and is considered to be late Early Triassic (Olenekian) to early Middle Triassic (Anisian) in age. The name of the biozone refers to Cynognathus crateronotus, a large and carnivorous cynodont therapsid which occurs throughout the entire biozone.
The Pristerognathus Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the upper Abrahamskraal Formation and lowermost Teekloof Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching not more than 300 metres (980 ft), occur just east of Sutherland through to Beaufort West in the south and Victoria West in the north. Exposures are also found west of Colesberg and south of Graaff-Reinet. The Pristerognathus Assemblage Zone is the third biozone of the Beaufort Group.
The Tropidostoma Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the lower Teekloof Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 240 metres (790 ft), occur from east of Sutherland through to Beaufort West and Victoria West, to areas south of Graaff-Reinet. Its northernmost exposures occur west/north-west of Colesberg. The Tropidostoma Assemblage Zone is the fourth biozone of the Beaufort Group.
The Eodicynodon Assemblage Zone is a tetrapod assemblage zone or biozone which correlates to the Abrahamskraal Formation, Adelaide Subgroup of the Beaufort Group, a fossiliferous and geologically important geological Group of the Karoo Supergroup in South Africa. The thickest outcrops, reaching approximately 620 metres (2,030 ft), occur south-east of Sutherland, north of Prince Albert, and south-east of Beaufort West. The Eodicynodon Assemblage Zone is the lowermost biozone of the Beaufort Group.
Daptocephalus is an extinct genus of dicynodont synapsid, which was found in Late Permian strata, in a biozone known precisely for the presence of fossils of this dicynodont, the Daptocephalus Assemblage Zone, in the Karoo Basin in South Africa. An additional species, D. huenei, is known from the Usili Formation in Tanzania and was formerly assigned to the genus Dicynodon before a study in 2019 recognised that the type specimen belonged to Daptocephalus.
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. Specimens of O. luangwensis have been found from Zambia. One species, O. sakamenensis, is the only Permian therapsid yet described from Madagascar. It is the type genus of the family Oudenodontidae, which includes members such as Tropidostoma.
Aulacephalodon is an extinct genus of medium-sized dicynodonts, or non-mammalian synapsids, that lived during late Permian period. Individuals of Aulacephalodon are commonly found in the Lower Beaufort Group of the Karoo Supergroup of South Africa. Rising to dominance during the Late Permian, Aulacephalodon was among the largest terrestrial vertebrate herbivores until its extinction at the end of the Permian. Two species have been named, the type species, A. bainii, and a second species, A. peavoti. However, debate exists among paleontologists if A. peavoti is a true member of the genus Aulacephalodon. Aulacephalodon belongs to the family Geikiidae, a family of dicynodonts generally characterized by their short, broad skulls and large nasal bosses. Sexual dimorphism has been identified in A. bainii.
Pylaecephalidae is a family of dicynodont therapsids that includes Diictodon, Robertia, and Prosictodon from the Permian of South Africa. Pylaecephalids were small burrowing dicynodonts with long tusks. The family was first named in 1934 and was redefined in 2009. Diictodontidae and Robertiidae are considered junior synonyms of Pylaecephalidae; although Pylaecephalus itself is considered a junior synonym of Diictodon, the name Pylaecephalidae predates these names and therefore takes priority.
The Abrahamskraal Formation is a geological formation and is found in numerous localities in the Northern Cape, Western Cape, and the Eastern Cape of South Africa. It is the lowermost formation of the Adelaide Subgroup of the Beaufort Group, a major geological group that forms part of the greater Karoo Supergroup. It represents the first fully terrestrial geological deposits of the Karoo Basin. Outcrops of the Abrahamskraal Formation are found from the small town Middelpos in its westernmost localities, then around Sutherland, the Moordenaarskaroo north of Laingsburg, Williston, Fraserburg, Leeu-Gamka, Loxton, and Victoria West in the Western Cape and Northern Cape. In the Eastern Cape outcrops are known from Rietbron, north of Klipplaat and Grahamstown, and also southwest of East London.
Bulbasaurus is an extinct genus of dicynodont that is known from the Lopingian epoch of the Late Permian period of what is now South Africa, containing the type and only species B. phylloxyron. It was formerly considered as belonging to Tropidostoma; however, due to numerous differences from Tropidostoma in terms of skull morphology and size, it has been reclassified the earliest known member of the family Geikiidae, and the only member of the group known from the Tropidostoma Assemblage Zone. Within the Geikiidae, it has been placed close to Aulacephalodon, although a more basal position is not implausible.
The Teekloof Formation is a geological formation that forms part of the Beaufort Group, one of the five geological groups that comprises the Karoo Supergroup in South Africa. The Teekloof Formation is the uppermost formation of Adelaide Subgroup deposits West of 24ºE and contains Middle to Late Permian-aged deposits and four biozones of the Beaufort Group. It overlies the Abrahamskraal Formation. The Teekloof Formation does not underlie other units other than the younger Karoo dolerites and sills that relate to the emplacement of the Early Jurassic Drakensberg Group to the east. Outcrops and exposures of the Teekloof Formation range from Sutherland through the mountain escarpments between Fraserburg and Beaufort West. The northernmost localities of the Teekloof Formation are found by Loxton, Victoria West and Richmond.
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.