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Diictodon | |
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Adult males with distorted skull | |
Undistorted skull of Diictodon in top view (left) and side view (right) scale bar = 1 cm | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Family: | † Pylaecephalidae |
Genus: | † Diictodon Owen 1876 |
Species [3] [4] | |
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Synonyms | |
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Diictodon is an extinct genus of pylaecephalid dicynodont [5] 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. [6] 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. [7]
Diictodon had disproportionally large heads that ended in a horny beak. ‘There is a clear distinction between specimens having canine tusks and those lacking them, as tusked specimens are generally larger and more likely to develop a pineal boss. This probably reflects sexual dimorphism, with the tusked sex almost certainly being the male.’ Diictodon had strong arms and legs, as well as 5 sharp claws on each hand, and may have had keen senses of smell and sight. Their gait was similar to the 'high walk' of crocodiles. Their jaws were also simplified, with some of the bones dedicated instead to hearing, considered a key sign of mammalian adaptation. Diictodon also had many adaptations for digging, such as highly developed muscles, a cylindrical body, and wide hands. [8] Researchers Chinsmay and Rubridge analyzed seven other Dicynodonts species discovering the humeral bone microstructure in Diictidon showed no signs of growth marks indicating a variation in its growth strategy that further improved their ability to dig. [9]
As a therapsid, Diictodon shared many features with modern-day mammals. Most noticeably, they made burrows into the earth, but most reached up to 0.5 m (1.6 ft) in depth, suggesting that they might have been infrequent diggers and occupied abandoned burrows. [8] Still, many scientists believe that Diictodon lived like the modern gopher. Their burrows could have been used to escape the heat of the desert, which was the dominant environment on the continent of Pangaea in the Late Permian Period. Inside these burrows, nests have been found, where Diictodon skeletons are present. They constituted of quite a gregarious lifestyle with numerous burrows in 500 square meters of space. However, their burrows were unconnected and did not form any large colonies. Many Diictodon nested close to flood plains, and some specimens may have been killed as water flowed into the nests, drowning the animals. Diictidon’s primary utilization of humeral excursion rather than forearm extension aided in employing rotation thrusting when burrowing. [9] Diictodon had no known rival species competing in its niche, so they may have competed primarily with others of their species for the little plant material available. [8] Fossils of infant Diictodon discovered in brood chambers in some burrows suggest there was parental care in the genus, and that males seem to have been involved in raising the infants, based on the fact that some adults in said burrows had tusks. [10]
Like all dicynodonts, Diictodon were herbivorous. They used their beaks to break off pieces of the sparse desert shrubs. Like modern desert animals, Diictodon may have had unusually efficient digestive systems, due to the lack of nutrients present in desert plants. As burrowing animals, they may have fed off of water-rich plant tubers. [11]
Previously, tusked individuals were considered males, while tuskless individuals were considered females. Differences in pelvic structure may be the other evidence for sexual dimorphism. [8]
Diictodon in a cladogram modified from Angielczyk and Rubidge (2010) showing the phylogenetic relationships of Dicynodontia: [12]
Dicynodontia |
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Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, typically toothless beak, unique amongst all synapsids. Dicynodonts first appeared in Southern Pangaea during the mid-Permian, ca. 270–260 million years ago, and became globally distributed and the dominant herbivorous animals in the Late Permian, ca. 260–252 Mya. They were devastated by the end-Permian Extinction that wiped out most other therapsids ca. 252 Mya. They rebounded during the Triassic but died out towards the end of that period. They were the most successful and diverse of the non-mammalian therapsids, with over 70 genera known, varying from rat-sized burrowers to elephant-sized browsers.
Robertia is an extinct genus of small herbivorous dicynodonts from the Middle to Late Permian of South Africa, between 260 and 265 million years ago. It is a monospecific genus, consisting of the type-species R. broomiana, which was classified by Lieuwe Dirk Boonstra in 1948 and named in honor of Robert Broom for his study of South African mammal-like reptiles.
Cistecephalus is an extinct genus of dicynodont therapsid from the Late Permian of southern Africa. It was a small, specialised, burrowing dicynodont, possibly with habits similar to a modern mole. The head was flattened and wedge-shaped, the body long, and the forelimbs very strong, with similarities in structure to the forelimb of modern burrowing mammals.
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.
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 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.
Eodicynodon is an extinct genus of dicynodont therapsids, a highly diverse group of herbivorous synapsids that were widespread during the middle-late Permian and early Triassic. As its name suggests, Eodicynodon is the oldest and most primitive dicynodont yet identified, ranging from the middle to late Permian and possessing a mix of ancestral anomodont/therapsid features and derived dicynodont synapomorphies.
Myosaurus is a genus of dicynodont synapsids. Myosaurus was a small, herbivorous synapsid that existed around the early Triassic period. All of the fossils found of this species were found in Antarctica and South Africa. Compared to other fossils found from species that existed during this time, the Myosaurus is not common in the fossil record. This is due to a shortage of discovered fossils that possess characteristics unique to the Myosaurus. Notably, under 130 fossil fragments have been found that have been classified as Myosauridae, and almost all have been skulls. These skulls can be classified as Myosaurus because this species, unlike other dicynodonts, do not possess tusks or postfrontal teeth. The only species identified in the family Myosauridae is the Myosaurus gracilis, or M. gracilis. It should be recognized that the Myosaurus is almost always referred to as the M. gracilis in scientific research.
Brachyprosopus is an extinct genus of dicynodont therapsid from the middle Permian Tapinocephalus Assemblage Zone in the Abrahamskraal Formation belonging to the Beaufort Group of the Karoo Basin, South Africa.
Dicynodontoides is a genus of small to medium-bodied, herbivorous, emydopoid dicynodonts from the Late Permian. The name Dicynodontoides references its “dicynodont-like” appearance due to the caniniform tusks featured by most members of this infraorder. Kingoria, a junior synonym, has been used more widely in the literature than the more obscure Dicynodontoides, which is similar-sounding to another distantly related genus of dicynodont, Dicynodon. Two species are recognized: D. recurvidens from South Africa, and D. nowacki from Tanzania.
Eosimops is an extinct genus of pylaecephalid dicynodonts. They were small synapsids superficially resembling modern mammals. Eosimops is known from several skull specimens, as well as one complete skeleton. Eosimops lived during the Middle Permian of South Africa.
Pristerodon is an extinct genus of dicynodont therapsid from the Late Permian of South Africa, Zambia and India.
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.
Prosictodon is an extinct genus of pylaecephalid dicynodont from Middle Permian of South Africa. It was first named by Kenneth D. Angielczyk and Bruce S. Rubidge in 2010 and the type species is Prosictodon dubei.
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.
Pristerodontia is a group of dicynodont therapsids that includes cryptodontids, geikiids, lystrosaurids, kannemeyeriids, and other related forms. Pristerodontians were one of the few groups of dicynodonts to survive the Permian–Triassic extinction event, diversifying in the Triassic.
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.
Cistecephalidae is an extinct family of dicynodont therapsids from the Late Permian of South Africa, India and Zambia. It includes the genera Cistecephalus, Cistecephaloides, and Kawingasaurus. Cistecephalids are thought to have had a fossorial or burrowing lifestyle, with adaptations such as broad skulls, strong forelimbs, and squat bodies. A similar group of dicynodonts called the pylaecephalids were also fossorial, although to a lesser extent than cistecephalids. Cistecephalids showed a high level of endemism, with each of the five known species unique to a single region.
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.
Kembawacela is an extinct genus of cistecephalid dicynodont from the Late Permian of East Africa. The genus contains two known species, the type species Kembawacela kitchingi from the Madumabisa Mudstone Formation of Zambia described in 2019, and a second species, K. yajuwayeyi, from the Chiweta Beds of Malawi described in 2022. Like other cistecephalids, Kembawacela was specialised for a fossorial, burrowing lifestyle similar to modern day moles. It is unique amongst cistecephalids for the presence of a pair of tusks in the upper jaw, characteristic of many other dicynodonts but lost in other cistecephalids. It is likely that Kembawacela was a locally endemic species of cistecephalid in the Luangwa Basin of Zambia.