Raranimus Temporal range: Middle Permian, | |
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Holotype skull from the sides | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Genus: | † Raranimus Liu et al., 2009 |
Type species | |
†Raranimus dashankouensis Liu et al., 2009 |
Raranimus is an extinct genus of therapsids of the Middle Permian. It was described in 2009 from a partial skull found in 1998 from the Dashankou locality of the Xidagou Formation, outcropping in the Qilian Mountains of Gansu, China. The genus is the most basal known member of the clade Therapsida, to which the later Mammalia belong. [1]
Raranimus shares a number of features with later therapsids and ancestral Sphenacodontia. The skull consists of a well preserved rostrum. The teeth suggest a carnivorous lifestyle for Raranimus, as the incisors are recurved and the second canines are serrated on their posterior edges. The incisors are morphologically similar to those seen in more derived theriodonts. The presence of two linguo−labially compressed canines is a diagnostic feature of Raranimus. The presence of two functional canines is characteristic of sphenacodontids, and this condition is seen in no other therapsid other than Rananimus. However, the slender, compressed shape of these canines is a derived characteristic of therapsids, with the canines of similarly sized sphenacodontids being more massively built. The precanines are small and anteriorly serrated, similar to what is seen in the synapsids Dimetrodon and Tetraceratops . [2] [3]
In the palate region of the skull, the anterior process of the vomer ventrally overlies the premaxilla at the anterior margin of the choana. This overlap is also seen in dinocephalians. However, unlike any other therapsid, the choanae are short and extend only from the level of the fourth incisor back to the first canine. [1]
According to a phylogenetic analysis conducted along with its initial description, Raranimus is considered to be the basalmost therapsid. [1] There has been some controversy as to whether Tetraceratops is a therapsid or a more basal pelycosaur. If Tetraceratops is a therapsid, as has recently been proposed, it would be the oldest and most basal one known, surpassing Raranimus in age by several million years. [3] However, later studies have questioned the placement of Tetraceratops within Therapsida, and the 2009 phylogenetic analysis using Raranimus places the genus outside of the clade. [1] [4] [5]
Raranimus occurs in strata that were deposited during the early Roadian stage of the Middle Permian. [6] The sphenacodontids were most diverse before the Roadian in the Early Permian, yet therapsids did not appear as a diverse group until near the Roadian-Wordian boundary. This has left a morphological and temporal gap in the fossil record during which the origin of therapsids must have occurred [7] called Olson’s Extinction. [8] [9]
With the general absence of therapsid remains from Olson's Extinction, different hypotheses have developed in order to explain the group's origins and initial diversification. One theory suggests that therapsids diversified quickly through rapid apomorphy accumulation sometime during the gap, while the other proposes that therapsids evolved gradually over the course of up to 35 Ma. [7] [10] Only recently have remains of basal therapsids such as Raranimus been found from China that occur during Olson's Extinction. Other therapsids that are known to have existed during the gap include Sinophoneus and Stenocybus . [11]
Below is a cladogram modified from Liu et al., 2009 depicting the phylogenetic relationships of Raranimus with therapsids that occur in Olson's Gap highlighted: [1]
Synapsida |
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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.
Dinocephalia is a clade of large-bodied early therapsids that flourished for a brief time in the Middle Permian between 270 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.
Sphenacodontidae is an extinct family of small to large, advanced, carnivorous, Late Pennsylvanian to middle Permian pelycosaurs. The most recent one, Dimetrodon angelensis, is from the late Kungurian or early Roadian San Angelo Formation. However, given the notorious incompleteness of the fossil record, a recent study concluded that the Sphenacodontidae may have become extinct as recently as the early Capitanian. Primitive forms were generally small, but during the later part of the early Permian these animals grew progressively larger, to become the top predators of their environments. Sphenacodontid fossils are so far known only from North America and Europe.
Sphenacodontia is a stem-based clade of derived synapsids. It was defined by Amson and Laurin (2011) as "the largest clade that includes Haptodus baylei, Haptodus garnettensis and Sphenacodon ferox, but not Edaphosaurus pogonias". They first appear during the Late Pennsylvanian epoch. From the end of the Carboniferous to the end of the Permian, most of them remained large, with only some secondarily becoming small in size.
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.
Anteosaurs are a group of large, primitive carnivorous dinocephalian therapsids with large canines and incisors and short limbs, that are known from the Middle Permian of South Africa, Russia, China, and Brazil. Some grew very large, with skulls 50–80 centimetres (20–31 in) long, and were the largest predators of their time. They died out at the end of the Middle Permian, possibly as a result of the extinction of the herbivorous Tapinocephalia on which they may have fed.
Therocephalia is an extinct suborder of eutheriodont therapsids from the Permian and Triassic. The therocephalians ("beast-heads") are named after their large skulls, which, along with the structure of their teeth, suggest that they were carnivores. Like other non-mammalian synapsids, therocephalians were once described as "mammal-like reptiles". Therocephalia is the group most closely related to the cynodonts, which gave rise to the mammals. This relationship takes evidence in a variety of skeletal features.
Tetraceratops insignis is an extinct synapsid from the Early Permian that was formerly considered the earliest known representative of Therapsida, a group that includes mammals and their close extinct relatives. It is known from a single 90-millimetre-long (3.5 in) skull, discovered in Texas in 1908. According to a 2020 study, it should be classified as a primitive non-therapsid sphenacodont rather than a genuine basal therapsid.
Anteosaurus is an extinct genus of large carnivorous synapsids. It lived during the Capitanian age of the Guadalupian epoch in what is now South Africa. Like the well-known Moschops, they were dinocephalians, a clade of large-bodied therapsids that flourished from 270 to 260 million years ago and then went extinct without descendants.
Theriognathus is an extinct genus of therocephalian therapsid belonging to the family Whaitsiidae, known from fossils from South Africa, Zambia, and Tanzania. Theriognathus has been dated as existing during the Late Permian. Although Theriognathus means mammal jaw, the lower jaw is actually made up of several bones as seen in modern reptiles, in contrast to mammals. Theriognathus displayed many different reptilian and mammalian characteristics. For example, Theriognathus had canine teeth like mammals, and a secondary palate, multiple bones in the mandible, and a typical reptilian jaw joint, all characteristics of reptiles. It is speculated that Theriognathus was either carnivorous or omnivorous based on its teeth, and was suited to hunting small prey in undergrowth. This synapsid adopted a sleek profile of a mammalian predator, with a narrow snout and around 1 meter long. Theriognathus is represented by 56 specimens in the fossil record.
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.
Biseridens is an extinct genus of anomodont therapsid, and one of the most basal anomodont genera known. Originally known from a partial skull misidentified as an 'eotitanosuchian' in 1997, another well-preserved skull was found in the Xidagou Formation, an outcropping in the Qilian Mountains of Gansu, China, in 2009 that clarified its relationships to anomodonts, such as the dicynodonts.
Dimacrodon is an extinct genus of non-mammalian synapsid from the latest Early Permian San Angelo Formation of Texas. It is distinguished by toothless, possibly beaked jaw tips, large lower canines and a thin bony crest on top of its head. Previously thought to be an anomodont therapsid related to dicynodonts, it was later found to lack any diagnostic features of anomodonts or even therapsids and instead appears to be a 'pelycosaur'-grade synapsid of uncertain classification.
Eosyodon is a dubious genus of extinct non-mammalian synapsids from the Permian of Texas. Its type and only species is Eosyodon hudsoni. Though it was originally interpreted as an early therapsid, it is probably a member of Sphenacodontidae, the family of synapsids that includes Dimetrodon.
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 Qingtoushan Formation is a Middle Permian-age geologic formation in the Qilian Mountains of Gansu, China. It is known for its diverse tetrapod fauna known as the Dashankou fauna, which likely dates to the Roadian, and includes some of the oldest known therapsids. This formation was previously erroneously named as the Xidagou Formation, a name which applies to otherwise Triassic strata in the northern Qillian Mountains. The formation is over 300 metres thick, and primarily consists of purple-red coarse sandstones, with minor purple mudstone.
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.
Gorgodon is an extinct genus of basal synapsids. The genus is monotypic, known only from the type species Gorgodon minutus from the Early Permian of the southwestern United States. The only known remains of Gorgodon are two fossils consisting of fragments of the skull. Gorgodon was described and named by paleontologist Everett C. Olson in 1962 from the San Angelo Formation in Knox County, Texas. Based on what is known of Gorgodon—the squamosal, quadrate, and pterygoid bones of the back of the skull, the maxilla and premaxilla bones that make up the front of the skull, and several teeth—Gorgodon had a relatively large temporal fenestra and a pair large, conical caniniform teeth at the front of the jaw. Other distinguishing features of Gorgodon include the fused connection between the quadrate and squamosal bones and a long transverse process of the pterygoid.
Abdalodon is an extinct genus of late Permian cynodonts, known by its only species A. diastematicus.Abdalodon together with the genus Charassognathus, form the clade Charassognathidae. This clade represents the earliest known cynodonts, and is the first known radiation of Permian cynodonts.