Eosimops Temporal range: Middle Permian | |
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A life restoration of an adult Eosimops newtoni | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Suborder: | † Anomodontia |
Clade: | † Dicynodontia |
Family: | † Pylaecephalidae |
Genus: | † Eosimops Broom, 1921 |
Species | |
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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. [1]
Eosimops was named in by South African paleontologist Robert Broom. [1] Its name is derived from the Ancient Greek Ēṓs (“dawn”) and ópsis (“appearance”).
Eosimops was discovered in 1921 by Robert Broom, based on a single skull. [1] It was found in the Tapinocephalus Assemblage Zone strata of South Africa, an area which provides a rich variety of early therapsids and covers a period of almost 100 million years ranging from the Permian to the Jurassic. [2] Later, other skull specimens as well as a complete skeleton were found. Eosimops is found above the stratigraphic range of the early dicynodont Eodycynodon . Also found in the Pristerognathus Assemblage Zone, Eosimops occurs in association with animals like Eunotosaurus and scylacosaurid therocephalans. [1]
Eosimops was a small, non-mammalian synapsid. It was around 34 cm in length (13.4 in.), roughly the size of a prairie dog. [1] It had a long cylindrical body, with sprawling, clawed limbs which it probably used for digging. Eosimops had two long tusks on its upper jaw, [3] and a cutting keratinized beak [4] for processing vegetation. It likely fed on leaves, stems, roots, and fleshy parts of plants. It has been suggested that some dicynodonts had hair, so Eosimops may have sported small hairs for insulation and tactile sensation. Its short and stocky proportions could have also aided in heat retention. [5]
Like other pylaecephalids, Eosimops had a roughly square skull that sported caniform tusks. These tusks have been interpreted as representing sexual dimorphism within the closely related genus Diictodon , [6] so may have been used for sexual selection. At least one postcanine tooth was present on the dentary blade. Its skull shape is described as being similar to that of its fellow pylaecephalid Robertia , but likely attaining larger sizes. [1]
Eosimops had indistinguishably fused premaxillae, with the single element forming the anterior portion of the snout and alveolar margin along with the anterior edge of the external nares. The premaxilla forms part of the secondary pallet, and bears two sets of paired anterior ridges as well as a single median posterior palatal ridge. A dorsally directed portion of the premaxilla with a rounded edge projects between the nasals, diagnostic of dicynodonts thanks to the narrow groove along its midline. [1] The left and right dentaries of Eosimops’ mandible were fused, and the anterior surface sported a vascular foramina which was likely associated with a keratinaceous beak. [4] Mandibular teeth were present, along with a well-developed dentary table. [4] The dorsal margin of the symphysis is upturned, forming a cutting edge at the front of the lower jaw. [1] The jaw joint facilitated a fore and aft sliding motion, allowing the animal to process vegetation effectively. [7]
Eosimops had a cylindrical body with 47 vertebrae, much like its other dicynodont relatives. Six of these were cervical, and 23 were dorsal. In the one full specimen recovered, no atlantal or axial ribs were observed. Whether or not this represents a true absence or incomplete preservation is uncertain. The dorsal ribs of Eosimops were long and relatively thin, and there was a ventral component to the curvature of the thorax. The sacral ribs were laterally expanded and robust. Its body likely resembled that of its close relatives Robertia and Diictodon. Typical of dicynodonts, the humerus of Eosimops sported expanded proximal and distal ends. It had a typical anomodont phalangeal formula of 2-3-3-3-3 on its forelimbs. While the hind limbs were not preserved well enough to know for certain, it appears that this formula was present on the hind limbs as well. [1]
Both the fore and hind limbs possessed extended phalanges with long, flattened claws, which suggests that Eosimops was a digger. It had short limbs, and it likely had a sprawling posture similar to its close relative Diictodon. Like Diictodon, Eosimops likely used its forelimbs for postural support and digging and its hind limbs for impact loading. [8]
Like all other known dicynodonts Eosimops was herbivorous, using its horny beak to process plant matter. As it didn't have a well-developed mastication system in comparison to modern vertebrates and lacked a gastric mill, Eosimops likely had a well-developed digestive tract and focused on feeding on high-quality forage. This likely included gymnosperm plants, evidence of which has been found in dicynodont coprolites. [9]
Dicynodonts, including Eosimops, have been suspected for some time to be endothermic. [10] In a histological study of the closely related Diictodon, another pylaecephalid, rapid bone growth is shown to be part of their early ontogeny. Continued growth during adult stages was also observed. [11] This rapid growth as well as moderately vascularized bones suggests that Diictodon could have been an endotherm, and that Eosimops could have been as well. This would be consistent with the hypothesis that more derived dicynodonts were endotherms, as endothermy would likely have been evolved early in the taxon's history.
Eosimops also potentially had hair. The discovery of hair remains in coprolites from carnivorous species that had consumed dicynodonts suggests that the hair was of dicynodont origin, [5] so hair could well be present in basal forms such as pylaecephalids. [12] This, along with Eosimops’ stocky body proportions, would allow the animal to conserve generated heat. [5]
The genus Eosimops represents a single known species, Eosimops newtoni [1] .
Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, as opposed to the sprawling posture of many reptiles and salamanders.
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.
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 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.
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.
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.
Wadiasaurus is an extinct genus of dicynodont from the family Kannemeyeria, that lived in herds from the early to Middle Triassic. Substantial fossorial evidence of W. indicus was recovered from Yerrapalli Formation of the Pranhita-Godavari valley, India, and it is so far the only Kannemeyeriid known for certain from India. The Kannemeyeriiformes underwent a significant diversification during the middle Triassic, with roughly 40 known species distributed worldwide. All Kannemeyeriiformes were medium to large bodied, graviportal herbivores with relatively erect posture and gait. Wadiasaurus indicus is currently the only known species of Wadiasaurus.
Myosaurus is a genus of Anomodontia in the order Therapsida. They are also classified as Dicynodontia, which is a subclade of Anomodontia. The Mysosaurus was a small, herbivorous reptile 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.
Kawingasaurus is an extinct genus of dicynodont therapsid from the Late Permian Usili Formation of Tanzania. It is a member of the family Cistecephalidae, and like other cistecephalids it is thought to have been fossorial. It is a member of the family Cistecephalidae. Cistephalidae includes genera Cisteceohalus, Cistecephaloides and Kawingasaurus. Greek for Saurus meaning “lizard” appears as a suffix denoting a reptilian origin. Living between 254.17 and 259.9 million years ago in the late Permian and believed to have the first and last recorded appearance in this time period. It lived in deep burrows as a suggested by most burrowing dicynodonts from evaluation of cranial sutures, vestibule inflation and enlarged stapes foot plates which are thought to be functionally correlated with bone-conduction hearing; all observed in fossorial vertebrates which use seismic signals as communication. Kawingasaurus lives in a terrestrial environment and is known as one of the smallest Cistecephaloides according to skull size when compared to other well-known taxa.
Sangusaurus is an extinct genus of large dicynodont synapsid with two recognized species: S. edentatus and S. parringtonii. Sangusaurus is named after the Sangu stream in eastern Zambia near to where it was first discovered + ‘saur’ which is the Greek root for lizard. Sangusaurus fossils have been recovered from the upper parts of the Ntawere Formation in Zambia and of the Lifua Member of the Manda Beds in Tanzania. The earliest study considered Sangusaurus a kannemeyeriid dicynodont, but more recent phylogenetic analyses place Sangusaurus within the stahleckeriid clade of Dicynodontia. Until recently, little work had been done to describe Sangusaurus, likely due to the fact that only four incomplete fossil specimens have been discovered.
Tetragonias is an extinct genus of dicynodont from the Anisian Manda Beds of Tanzania. With tetra meaning “four,” and goni meaning “angle,” the name references the square shape of the Tetragonias skull when viewed dorsally. Not to be confused with the plant Tetragonia,Tetragonias were dicynodont anomodonts discovered in the late 1960s by paleontologist A. R. I. Cruickshank in the Manda Formation. Only the type species, T. njalilus, has been recognized.
Chainosauria is a large and speciose clade of anomodont therapsid that includes the highly diverse dicynodonts and a small number of closely related basal genera —although the total composition and taxonomic scope of Chainosauria is in flux. Chainosauria was named in 1923 to group together the dicynodonts and their close relatives, namely three small anomodont genera from South Africa that made up the now defunct group 'Dromasauria'. The name soon fell into disuse, however, as it was functionally replaced by Anomodontia. Chainosauria was later revived cladistically in 2009, preserving the association of dicynodonts and the 'dromasaurs' and has since served in effect as both a cladistic and a biogeographic counterpart to the Laurasian venyukovioids, with early chainosaurs appearing to have been a Gondwanan radiation.
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.
Syops is an extinct genus of dicynodont therapsid. The type species S. vanhoepeni was first named in 1938 as Dicynodon vanhoepeni. Fossils of the genus have been found in the Cistecephalus Assemblage Zone in the Usili Formation of the Ruhuhu Basin, Tanzania and the Upper Madumabisa Mudstone Formation of the Luangwa Basin, Zambia. Its phylogenetic placement is somewhat uncertain, with multiple different studies finding it as either a basal geikiid, rhachiocephalid a dicynodontoid more derived than the most basal genera but less derived than Lystrosauridae, or a lystrosaurid.
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 Aulacocephalodon, although a more basal position is not implausible.
Thliptosaurus 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. Thliptosaurus is from the upper Daptocephalus Assemblage Zone, making it one of the youngest Permian dicynodonts known, living just prior to the Permian mass extinction. It also represents one of the few small bodied dicynodonts to exist at this time, when most other dicynodonts had large body sizes and many small dicynodonts had gone extinct. The unexpected discovery of Thliptosaurus in a region of the Karoo outside of the historically sampled localities suggests that it may have been part of an endemic local fauna not found in these historic sites. Such under-sampled localities may contain 'hidden diversities' of Permian faunas that are unknown from traditional samples. Thliptosaurus is also unusual for dicynodonts as it lacks a pineal foramen, suggesting that it played a much less important role in thermoregulation than it did for other dicynodonts.