Niaftasuchus Temporal range: | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Genus: | † Niaftasuchus Ivakhnenko, 1990 |
Species: | †N. zekkeli |
Binomial name | |
†Niaftasuchus zekkeli Ivakhnenko, 1990 | |
Niaftasuchus is an extinct genus of therapsids. Its type and only named species is Niaftasuchus zekkeli.
Niaftasuchus had distinctive dentition. It has been interpreted as one of the earliest known herbivorous therapsids. It lived during the Guadalupian epoch of the Permian in what is now Russia, and inhabited an environment alongside some of the last non-therapsid synapsids, such as the caseasaur Ennatosaurus and the varanopid Mesenosaurus .
Niaftasuchus is one of several enigmatic early therapsids from Russia that may be based on juvenile material. Its phylogenetic affinities are controversial; it has been classified as a biarmosuchian, dinocephalian, or anomodont, and it has also been suggested to belong to a lineage of its own.
Niaftasuchus has a distinctive dentition composed of large, procumbent, leaf-shaped teeth. There are three pairs of incisors, but the canines are not particularly distinct from the other maxillary teeth. There are a few palatal teeth. Like biarmosuchians, Niaftasuchus has exceptionally large eye sockets, though this may be a juvenile character. The skull is low and broad.
The known specimens of Niaftasuchus are very small; the holotype skull is only 9 centimetres (3.5 in) long. [1] It has been interpreted as being based on probable juvenile material. [2]
The holotype specimen of Niaftasuchus zekkeli, an incomplete skull without the mandible, was collected near the Pyoza river in Arkhangelsk Oblast, Russia, near Nyafta (Russian : Няфта). It was named in 1990 by M. F. Ivakhnenko, with the name referring to the place where it was found and honoring the geologist I. D. Zekkel. Ivakhnenko initially interpreted it as a tapinocephalian, but subsequently proposed a monotypic order, Niaftasuchida, for it. In 2000, Battail and Surkov argued it was a biarmosuchian. In 2001, Ivakhnenko suggested it was an anomodont, but in 2003 returned to regarding it as a dinocephalian on the basis of new material. [3]
The classification of Niaftasuchus is considered problematic. [2] It is assigned to a family of its own, Niaftasuchidae. Ivakhnenko, who first described Niaftasuchus, regards it as a basal dinocephalian, whereas Battail and Surkov classified Niaftasuchus in Biarmosuchia. It is also possible that it belongs to neither lineage, and forms a distinctive therapsid group of its own. [4] [5]
There is only one named species, Niaftasuchus zekkeli, but a specimen that may represent a second species is known. [6] It differs from the type species in having a straight tooth row without enlarged maxillary teeth.
Niaftasuchus is interpreted as an herbivore, with its teeth adapted primarily for tearing off soft plant parts. Based on possible coprolites found in the mouth of a juvenile specimen, Ivakhnenko suggested that juvenile Niaftasuchus would ingest coprolites to develop their gut microbiota, as in many modern herbivores. [7]
Dinosaurus is an extinct genus of therapsid of controversial affinities. Its type and only species is Dinosaurus murchisonii. It is only known from a partial snout from the Permian of Russia. Its taxonomic history is intertwined with several other poorly-known Russian therapsids, particularly Rhopalodon, Brithopus, and Phthinosuchus.
Therapsida is a clade comprising a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close 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, resulting in a more "standing" quadrupedal posture, as opposed to the lower sprawling posture of many reptiles and amphibians.
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.
Dinocephalians are a clade of large-bodied early therapsids that flourished in the Early and Middle Permian between 279.5 and 260 million years ago (Ma), but became extinct during the Capitanian mass extinction event. Dinocephalians included herbivorous, carnivorous, and omnivorous forms. Many species had thickened skulls with many knobs and bony projections. Dinocephalians were the first non-mammalian therapsids to be scientifically described and their fossils are known from Russia, China, Brazil, South Africa, Zimbabwe, and Tanzania.
Suminia is an extinct genus of basal anomodont that lived during the Tatarian age of the late Permian, spanning approximately from 268-252 Ma. Suminia is recognized the youngest non-dicynodont anomodont. Its fossil localities are primarily derived from the Kotel’nich locality of the Kirov Oblast in Russia. However, there have been some isolated specimen found in a few different localities, all from eastern European regions of Russia.
Biarmosuchia is an extinct clade of non-mammalian synapsids from the Permian. Biarmosuchians are the most basal group of the therapsids. They were moderately-sized, lightly built carnivores, intermediate in form between basal sphenacodont "pelycosaurs" and more advanced therapsids. Biarmosuchians were rare components of Permian ecosystems, and the majority of species belong to the clade Burnetiamorpha, which are characterized by elaborate cranial ornamentation.
Inostrancevia is an extinct genus of large carnivorous therapsids which lived during the Late Permian in what are now Siberia, Russia and Southern Africa. The first-known fossils of this gorgonopsian were discovered in the Northern Dvina, where two almost complete skeletons were exhumed. Subsequently, several other fossil materials were discovered in various oblasts, and these finds will lead to a confusion about the exact number of valid species in the country, before only three of them were officially recognized: I. alexandri, I. latifrons and I. uralensis. More recent research carried out in South Africa has discovered fairly well-preserved remains of the genus, being attributed to the species I. africana. An isolated left premaxilla suggests that Inostrancevia also lived in Tanzania during the earliest Lopingian age. The whole genus is named in honor of Alexander Inostrantsev, professor of Vladimir P. Amalitsky, the paleontologist who described the taxon.
Anteosaurus is an extinct genus of large carnivorous dinocephalian synapsid. It lived at the end of the Guadalupian during the Capitanian age, about 265 to 260 million years ago in what is now South Africa. It is mainly known by cranial remains and few postcranial bones. Measuring 5–6 m (16–20 ft) long and weighing about 600 kg (1,300 lb), 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.
Ennatosaurus is an extinct genus of caseid synapsid that lived during the Middle Permian in northern European Russia. The genus is only represented by its type species, Ennatosaurus tecton, which was named in 1956 by Ivan Antonovich Efremov. The species is known from at least six skulls associated with their lower jaws, as well as from the postcranial bones of several juvenile individuals. Ennatosaurus has the typical caseid skull with a short snout tilted forward and very large external nares. However, it differs from other derived caseids by its postcranial skeleton with smaller proportions compared to the size of the skull. As with other advanced caseids, the teeth of Ennatosaurus were well suited for slicing and cutting vegetation. The presence of a highly developed hyoid apparatus indicates the presence of a massive and mobile tongue, which had to work in collaboration with the palatal teeth during swallowing. With a late Roadian - early Wordian age, Ennatosaurus is one of the last known caseids.
Lemurosaurus is a genus of extinct biarmosuchian therapsids from the Late Permian of South Africa. The generic epithet Lemursaurus is a mix of Latin, lemures “ghosts, spirits”, and Greek, sauros, “lizard”. Lemurosaurus is easily identifiable by its prominent eye crests, and large eyes. The name Lemurosaurus pricei was coined by paleontologist Robert Broom in 1949, based on a single small crushed skull, measured at approximately 86 millimeters in length, found on the Dorsfontein farm in Graaff-Reinet. To date, only two skulls of the Lemurosaurus have been discovered, so body size is unknown. The second larger, more intact, skull was found in 1974 by a team from the National Museum, Bloemfontein.
Venyukovia is an extinct genus of venyukovioid therapsid, a basal anomodont from the Middle Permian of Russia. The type and sole species, V. prima, is known only by a partial lower jaw with teeth. Venyukovia has often been incorrectly spelt as 'Venjukovia' in English literature. This stems from a spelling error made by Russian palaeontologist Ivan Efremov in 1940, who mistakenly replaced the 'y' with a 'j', which subsequently permeated through therapsid literature before the mistake was caught and corrected. Venyukovia is the namesake for the Venyukovioidea, a group of small Russian basal anomodonts also including the closely related Otsheria, Suminia, Parasuminia and Ulemica, although it itself is also one of the poorest known. Like other venyukovioids, it had large projecting incisor-like teeth at the front and lacked canines, although the remaining teeth are simple compared to some other venyukovioids, but may resemble those of Otsheria.
Paraburnetia is an extinct genus of biarmosuchian therapsids from the Late Permian of South Africa. It is known for its species P. sneeubergensis and belongs to the family Burnetiidae. Paraburnetia lived just before the Permian–Triassic mass extinction event.
Megawhaitsia is an extinct genus of large therocephalian therapsids who lived during the Late Permian (Wuchiapingian) in what is now Eastern Europe. The only known species is M. patrichae, described in 2008 from several fossils discovered in various oblasts of European Russia. The fossils are representative of a large animal whose skull size is estimated to be 40–50 cm (16–20 in) long.
Pachydectes is an extinct genus of biarmosuchian therapsids from the Middle Permian of South Africa known from a single skull. The etymology of the name Pachydectes is derived from the Greek word pakhus, meaning "thick" or "thickened", and dektes, meaning "biter". In conjunction this name is representative of the unique pachyostotic bone present above the maxillary canine tooth found in the skull of the specimen. There is only one known species within the genus, Pachydectes elsi which is named in honor of the person who discovered the fossil.
Venyukovioidea is an infraorder of anomodont therapsids related to dicynodonts from the Permian of Russia. They have also been known as 'Venjukovioidea', as well as by the similar names 'Venyukoviamorpha' or 'Venjukoviamorpha' in literature. This in part owes to a misspelling by Russian palaeontologist Ivan Efremov in 1940 when he mistakenly spelt Venyukovia, the namesake of the group, with a 'j' instead of a 'y', which permeated through subsequent therapsid literature before the mistake was caught and corrected. The order Ulemicia has also been coined for a similar taxonomic concept in Russian scientific literature, which notably excludes Suminia and Parasuminia.
Sinophoneus is an extinct genus of carnivorous dinocephalian therapsid belonging to the family Anteosauridae. It lived 272 to 270 million years ago at the beginning of the Middle Permian in what is now the Gansu Province in northern China. It is known by a skull of an adult individual, as well as by many skulls of juvenile specimens. The latter were first considered as belonging to a different animal, named Stenocybus, before being reinterpreted as immature Sinophoneus. Sinophoneus shows a combination of characters present in other anteosaurs. Its bulbous profile snout and external nostrils located in front of the canine are reminiscent of the basal anteosaur Archaeosyodon, while its massive transverse pterygoids processes with enlarged distal ends are more similar to the more derived anteosaurs Anteosaurus and Titanophoneus. First phylogenetic analyzes identified Sinophoneus as the most basal Anteosaurinae. A more recent analysis positioned it outside the Anteosaurinae and Syodontinae subclades, and recovers it as the most basal Anteosauridae.
Novocynodon is an extinct genus of therapsids from the Middle Permian of Russia. Fossils have been found in Alexandrovsky District, Orenburg Oblast. The type and only species is Novocynodon kutorgai. It was classified as a thrinaxodontid cynodont when first described in 2012, but this was disputed by Kammerer (2014), who argued that it might instead be a juvenile anomodont or dinocephalian.
Phthinosuchia is an extinct group of therapsids including two poorly known species, Phthinosuchus discors and Phthinosaurus borrisiaki, from the Middle Permian of Russia. Phthinthosuchus is known a partial crushed skull and Phthinosaurus is known from an isolated lower jaw. The two species have traditionally been grouped together based on their shared primitive characteristics, but more recent studies have proposed that they are more distantly related. Phthinosuchus is either a carnivorous gorgonopsian relative or an anteosaurian dinocephalian while Phthinosaurus is either a herbivorous rhopalodont dinocephalian or a therocephalian.
Phthinosaurus is an extinct genus of therapsids from the Middle Permian of Russia. The type species Phthinosaurus borrisiaki was named by Soviet paleontologist Ivan Yefremov in 1940 on the basis of an isolated lower jaw. Because this jaw provides few distinguishing characteristics, the evolutionary relationships of Phthinosaurus are poorly known. Yefremov named the family Phthinosuchidae in 1954 to include Phthinosaurus and the newly named Phthinosuchus, which was described on the basis of a crushed partial skull. American paleontologist Everett C. Olson placed both of these therapsids in the larger infraorder Phthinosuchia in 1961. In 1974 Leonid Tatarinov named the family Phthinosauridae to include Phthinosaurus alone, retaining Phthinosuchus within Phthinosuchidae.
Parasumina is an extinct genus of anomodont known from the late Capitanian age at the end of the middle Permian period of European Russia. The type and only species is Parasuminia ivakhnenkoi. It was closely related to Suminia, another Russian anomodont, and was named for its resemblance. Little is known about Parasuminia as the only fossils are of fragmentary pieces of the skull and jaw, but the known remains suggest that its head and jaws were deeper and more robust than those of Suminia, and with shorter, stouter teeth. However, despite these differences they appear to have been similar animals with a similarly complex method of processing vegetation.