Elliotsmithia Temporal range: Middle Permian | |
---|---|
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Family: | † Varanopidae |
Subfamily: | † Varanopinae |
Genus: | † Elliotsmithia Broom, 1937 |
Type species | |
†Elliotsmithia longiceps Broom, 1937 |
Elliotsmithia is a small varanopid synapsid found from the late Middle Permian of South Africa. It is the sole basal synapsid "pelycosaur" known from the supercontinent Gondwana and only two specimens have been yielded to date. Its species name longiceps is derived from Latin, meaning "long head". Both known Elliotsmithia fossils were recovered from Abrahamskraal Formation rocks—within the boundaries of the Tapinocephalus Assemblage Zone—of the lower Beaufort Group. [1] [2] It was named for the late Sir Grafton Elliot Smith in 1937. [3]
In 1917 Dr. Van Hoepen discovered the fossil holotype of Elliotsmithia. The holotype consisted of a skull, mandibles with most of the front region missing, the first four cervical vertebrae complete with atlas-axis complex, rib fragments, and dermal ossifications. Van Hoepen found the fossil in rocks associated with the Tapinocephalus Assemblage Zone near the town Prince Albert in the Western Cape. The specimen remained undescribed until renowned paleontologist and physician, Dr. Robert Broom, briefly described and named the fossil in 1937. [3] However, Broom did not go into great detail in his description. Several researchers authored papers over several decades, providing passing descriptions of the fossil. [4] [5] [6] [7] In the 1990s, renewed research on the holotype fossil classified Elliotsmithia as a varanopseid "pelycosaur" synapsid. [8] Another specimen was found in 2001. [9]
A possible third specimen was recovered from the Pristerognathus Assemblage Zone in 2010; however, this specimen requires further research. [10]
Elliotsmithia is considered to have been a small carnivore that hunted insects and small vertebrates. It had a long, slender snout complete with recurved, mediolaterally flattened serrated teeth for hooking its prey.
Both known Elliotsmithia fossils were recovered from rock deposits of the Middle Permian Tapinocephalus Assemblage Zone, Abrahamskraal Formation of the Beaufort Group. Another possible specimen of Elliotsmithia was recovered from the lower Teekloof Formation of the latest Middle Permian-aged Pristerognathus Assemblage Zone. Elliotsmithia is an extremely important fossil species as it provides proof of the cosmopolitan range of basal varanopseid synapsids, especially within Gondwana where therapsids were the dominant fauna.
It is currently accepted that Elliotsmithia is a basal varanopid eupelycosaur synapsid and part of the informal "pelycosaur" group. Recent paleobiology analyses of its cranial bones confidently places this basal synapsid as a sister taxon to the varanopid clade composed of Varanops , Aerosaurus , and Varanodon from North America.
Little research has been endeavored on the fossils of Elliotsmithia and this species warrants further study.
Synapsids are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptiles and birds. The group includes mammals and every animal more closely related to mammals than to sauropsids. Unlike other amniotes, synapsids have a single temporal fenestra, an opening low in the skull roof behind each eye orbit, leaving a bony arch beneath each; this accounts for their name. The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, when synapsids and sauropsids diverged, but was subsequently merged with the orbit in early mammals.
Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308 million years ago during the Early Pennsylvanian epoch, with the fossils of Echinerpeton and perhaps an even earlier genus, Protoclepsydrops, representing just one of the many stages in the evolution of mammals, in contrast to their earlier amniote ancestors.
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.
Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an Archaeothyris-like synapsid in the Late Carboniferous. However, some recent studies have recovered them being taxonomically closer to diapsid reptiles. A varanopid from the latest Middle Permian Pristerognathus Assemblage Zone is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.
Mycterosaurus is an extinct genus of synapsids belonging to the family Varanopidae. It is classified in the varanopid subfamily Mycterosaurinae. Mycterosaurus is the most primitive member of its family, existing from 290.1 to 272.5 MYA, known to Texas and Oklahoma. It lacks some features that its advanced relatives have.
Emydops is an extinct genus of dicynodont therapsids from the Middle Permian to Late Permian of what is now South Africa. The genus could be described as 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.
Anteosaurus is an extinct genus of large carnivorous dinocephalian synapsid. It lived at the end of the Guadalupian during the Capitanian stage, about 265 to 260 million years ago in what is now South Africa. It is mainly known by cranial remains and few postcranial bones. With its skull reaching 80–90 cm (31–35 in) in length and a body size estimated at more than 5 m (16 ft) in length, and 500 to 600 kg in weight, Anteosaurus was the largest known carnivorous non-mammalian synapsid and the largest terrestrial predator of the Permian period. Occupying the top of the food chain in the Middle Permian, its skull, jaws and teeth show adaptations to capture large prey like the giants titanosuchids and tapinocephalids dinocephalians and large pareiasaurs.
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 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 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.
Aerosaurus is an extinct genus within Varanopidae, a family of non-mammalian synapsids. It lived between 252-299 million years ago during the Early Permian in North America. The name comes from Latin aes (aeris) “copper” and Greek sauros “lizard,” for El Cobre Canyon in northern New Mexico, where the type fossil was found and the site of former copper mines. Aerosaurus was a small to medium-bodied carnivorous synapsid characterized by its recurved teeth, triangular lateral temporal fenestra, and extended teeth row. Two species are recognized: A. greenleeorum (1937) and A. wellesi (1981).
Eriphostoma is an extinct genus of gorgonopsian therapsids known from the Middle Permian of Tapinocephalus Assemblage Zone, South Africa. It has one known species, Eriphostoma microdon, and was first named by Robert Broom in 1911. It is the oldest known gorgonopsian and among the smallest and most basal members of the clade.
Scymnosaurus is an extinct genus of therocephalian therapsids, first described by Robert Broom in 1903. There are three species that still take the name Scymnosaurus, S. ferox, S. watsoni and S. major, with a fourth, S. warreni, now identified as Moschorhinus warreni. Each of these have now been reclassified into Lycosuchidae incertae sedis.
Heleosaurus scholtzi is an extinct species of basal synapsids, known as pelycosaurs, in the family of Varanopidae during the middle Permian. At first H. scholtzi was mistakenly classified as a diapsid. Members of this family were carnivorous and had dermal armor, and somewhat resembled monitor lizards. This family was the most geologically long lived, widespread, and diverse group of early amniotes. To date only two fossils have been found in the rocks of South Africa. One of these fossils is an aggregation of five individuals.
Owenetta is an extinct genus of owenettid procolophonian parareptile. Fossils have been found from the Beaufort Group in the Karoo Basin of South Africa. Although most procolophonians lived during the Triassic, Owenetta existed during the Wuchiapingian and Changhsingian stages of the Late Permian as well as the early Induan stage of the Early Triassic. It is the type genus of the family Owenettidae, and can be distinguished from other related taxa in that the posterior portion of the supratemporal bears a lateral notch and that the pineal foramen is surrounded by a depressed parietal surface on the skull table.
Olson's Extinction was a mass extinction that occurred 273 million years ago in the late Cisuralian or early Guadalupian of the Permian period and which predated the Permian–Triassic extinction event. It is named after Everett C. Olson. There was a sudden change between the early Permian and middle/late Permian faunas. Some authors also place a hiatus in the continental fossil record around that time, but others disagree. Since then this event has been realized across many groups, including plants, marine invertebrates, and tetrapods.
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.
Homodontosaurus is an extinct genus of therocephalian therapsids from the Late Permian of South Africa. The type species Homodontosaurus kitchingi was named by South African paleontologist Robert Broom in 1949. Broom based his description on a small skull found in the Cistecephalus Assemblage Zone near Graaff-Reinet. The skull is very small, at about 55 millimetres (2.2 in) long and 20 millimetres (0.79 in) wide. Homodontosaurus has large eye sockets and an elongated snout. The lower jaw is long, thin, and curved. Numerous small teeth line the upper jaw and are long, pointed, and round in cross-section.
The Middleton Formation is a geological formation that extends through the Northern Cape, Western Cape, and Eastern Cape provinces of South Africa. It overlies the lower Abrahamskraal Formation, and is the eastern correlate, East of 24ºE, of the Teekloof Formation. Outcrops and exposures of the Middleton Formation range from Graaff-Reinet in the Eastern Cape onwards. The Middleton Formation's type locality lies near the small hamlet, Middleton, approximately 25 km south of Cookhouse. Other exposures lie in hillsides along the Great Fish River in the Eastern Cape. The Middleton Formation forms part of the Adelaide Subgroup of the Beaufort Group, which itself forms part of the Karoo Supergroup.
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