Therapsida Temporal range: | |
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From top to bottom and left to right, several examples of non-mammalian therapsids: Biarmosuchus (Biarmosuchia), Moschops (Dinocephalia), Lystrosaurus (Anomodontia), Inostrancevia (Gorgonopsia), Glanosuchus (Therocephalia) and Chiniquodon (Cynodontia) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Sphenacodontia |
Clade: | Pantherapsida |
Clade: | Sphenacodontoidea |
Clade: | Therapsida Broom, 1905 [1] |
Clades | |
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Therapsida [lower-alpha 1] 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.
Therapsids evolved from earlier synapsids commonly called "pelycosaurs", specifically within the Sphenacodontia, more than 279.5 million years ago. They replaced the pelycosaurs as the dominant large land animals in the Guadalupian through to the Early Triassic. In the aftermath of the Permian–Triassic extinction event, therapsids declined in relative importance to the rapidly diversifying archosaurian sauropsids (pseudosuchians, dinosaurs and pterosaurs, etc.) during the Middle Triassic.
The therapsids include the cynodonts, the group that gave rise to mammals (Mammaliaformes) in the Late Triassic around 225 million years ago, the only therapsid clade that survived beyond the end of the Triassic. The only other group of therapsids to have survived into the Late Triassic, the dicynodonts, became extinct towards the end of the period. The last surviving group of non-mammaliaform cynodonts were the Tritylodontidae, which became extinct during the Early Cretaceous.
Therapsids' temporal fenestrae were larger than those of the pelycosaurs. The jaws of some therapsids were more complex and powerful, and the teeth were differentiated into frontal incisors for nipping, great lateral canines for puncturing and tearing, and molars for shearing and chopping food.
Therapsid legs were positioned more vertically beneath their bodies than were the sprawling legs of reptiles and pelycosaurs. Also compared to these groups, the feet were more symmetrical, with the first and last toes short and the middle toes long, an indication that the foot's axis was placed parallel to that of the animal, not sprawling out sideways. This orientation would have given a more mammal-like gait than the lizard-like gait of the pelycosaurs. [2]
The physiology of therapsids is poorly understood. Most Permian therapsids had a pineal foramen, indicating that they had a parietal eye like many modern reptiles and amphibians. The parietal eye serves an important role in thermoregulation and the circadian rhythm of ectotherms, but is absent in modern mammals, which are endothermic. [3] Near the end of the Permian, dicynodonts, therocephalians and cynodonts show parallel trends towards loss of the pineal foramen, and the foramen is completely absent in probainognathian cynodonts. Evidence from oxygen isotopes, which are correlated with body temperature, suggests that most Permian therapsids were ectotherms and that endothermy evolved convergently in dicynodonts and cynodonts near the end of the Permian. [4] In contrast, evidence from histology suggests that endothermy is shared across Therapsida, [5] whereas estimates of blood flow rate and lifespan in the mammaliaform Morganucodon suggest that even early mammaliaforms had reptile-like metabolic rates. [6] Evidence for respiratory turbinates, which have been hypothesized to be indicative of endothermy, was reported in the therocephalian Glanosuchus , but subsequent study showed that the apparent attachment sites for turbinates may simply be the result of distortion of the skull. [7]
The evolution of integument in therapsids is poorly known, and there are few fossils that provide direct evidence for the presence or absence of fur. The most basal synapsids with unambiguous direct evidence of fur are docodonts, which are mammaliaforms very closely related to crown-group mammals. Two "mummified" juvenile specimens of the dicynodont Lystrosaurus murrayi preserve skin impressions; the skin is hairless, leathery, and dimpled, somewhat comparable to elephant skin. [8] [9] Fossilized facial skin from the dinocephalian Estemmenosuchus has been described as showing that the skin was glandular and lacked both scales and hair. [10]
Coprolites containing what appear to be hairs have been found from the Late Permian. [11] [12] Though the source of these hairs is not known with certainty, they may suggest that hair was present in at least some Permian therapsids.
The closure of the pineal foramen in probainognathian cynodonts may indicate a mutation in the regulatory gene Msx2, which is involved in both the closure of the skull roof and the maintenance of hair follicles in mice. [13] This suggests that hair may have first evolved in probainognathians, though it does not entirely rule out an earlier origin of fur. [13]
Whiskers probably evolved in probainognathian cynodonts. [13] [14] Some studies had inferred an earlier origin for whiskers based on the presence of foramina on the snout of therocephalians and early cynodonts, but the arrangement of foramina in these taxa actually closely resembles lizards, [15] which would make the presence of mammal-like whiskers unlikely. [14]
Therapsids evolved from a group of pelycosaurs called sphenacodonts. [17] [18] Therapsids became the dominant land animals in the Middle Permian, displacing the pelycosaurs. Therapsida consists of four major clades: the dinocephalians, the herbivorous anomodonts, the carnivorous biarmosuchians, and the mostly carnivorous theriodonts. After a brief burst of evolutionary diversity, the dinocephalians died out in the later Middle Permian (Guadalupian) but the anomodont dicynodonts as well as the theriodont gorgonopsians and therocephalians flourished, being joined at the very end of the Permian by the first of the cynodonts.
Like all land animals, the therapsids were seriously affected by the Permian–Triassic extinction event, with the very successful gorgonopsians and the biarmosuchians dying out altogether and the remaining groups—dicynodonts, therocephalians and cynodonts—reduced to a handful of species each by the earliest Triassic. Surviving dicynodonts were represented by two families of disaster taxa (Lystrosauridae and Myosauridae), the scarcely known Kombuisia , and a single group of large stocky herbivores, the Kannemeyeriiformes, which were the only dicynodont lineage to thrive during the Triassic. [20] They and the medium-sized cynodonts (including both carnivorous and herbivorous forms) flourished worldwide throughout the Early and Middle Triassic. They disappear from the fossil record across much of Pangea at the end of the Carnian (Late Triassic), although they continued for some time longer in the wet equatorial band and the south.[ citation needed ]
Some exceptions were the still further derived eucynodonts. [21] At least three groups of them survived. They all appeared in the Late Triassic period. The extremely mammal-like family, Tritylodontidae, survived into the Early Cretaceous. Another extremely mammal-like family, Tritheledontidae, are unknown later than the Early Jurassic. Mammaliaformes was the third group, including Morganucodon and similar animals. Some taxonomists refer to these animals as "mammals", though most limit the term to the mammalian crown group.
The non-eucynodont cynodonts survived the Permian–Triassic extinction; Thrinaxodon , Galesaurus and Platycraniellus are known from the Early Triassic. By the Middle Triassic, however, only the eucynodonts remained.
The therocephalians, relatives of the cynodonts, managed to survive the Permian–Triassic extinction and continued to diversify through the Early Triassic period. Approaching the end of the period, however, the therocephalians were in decline to eventual extinction, likely outcompeted by the rapidly diversifying Saurian lineage of diapsids, equipped with sophisticated respiratory systems better suited to the very hot, dry and oxygen-poor world of the End-Triassic.
Dicynodonts were among the most successful groups of therapsids during the Late Permian, and survived through to near the end of the Triassic.
Mammals are the only living therapsids. The mammalian crown group, which evolved in the Early Jurassic period, radiated from a group of mammaliaforms that included the docodonts. The mammaliaforms themselves evolved from probainognathians, a lineage of the eucynodont suborder.
The Hopson and Barghausen paradigm for therapsid relationships |
Six major groups of therapsids are generally recognized: Biarmosuchia, Dinocephalia, Anomodontia, Gorgonopsia, Therocephalia and Cynodontia. A clade uniting therocephalians and cynodonts, called Eutheriodontia, is well supported, but relationships among the other four clades are controversial. [23] The most widely accepted hypothesis of therapsid relationships, the Hopson and Barghausen paradigm, was first proposed in 1986. Under this hypothesis, biarmosuchians are the earliest-diverging major therapsid group, with the other five groups forming the Eutherapsida, and within Eutherapsida, gorgonopsians are the sister taxon of eutheriodonts, together forming the Theriodontia. Hopson and Barghausen did not initially come to a conclusion about how dinocephalians, anomodonts and theriodonts were related to each other, but subsequent studies suggested that anomodonts and theriodonts should be classified together as the Neotherapsida. However, there remains debate over these relationships; in particular, some studies have suggested that anomodonts, not gorgonopsians, are the sister taxon of Eutheriodontia, other studies have found dinocephalians and anomodonts to form a clade, and both the phylogenetic position and monophyly of Biarmosuchia remain controversial.
In addition to the six major groups, there are several other lineages and species of uncertain classification. Raranimus from the early Middle Permian of China is likely to be the earliest-diverging known therapsid. [24] Tetraceratops from the Early Permian of the United States has been hypothesized to be an even earlier-diverging therapsid, [25] [26] but more recent study has suggested it is more likely to be a non-therapsid sphenacodontian. [27]
Biarmosuchia is the most recently recognized therapsid clade, first recognized as a distinct lineage by Hopson and Barghausen in 1986 and formally named by Sigogneau-Russell in 1989. Most biarmosuchians were previously classified as gorgonopsians. Biarmosuchia includes the distinctive Burnetiamorpha, but support for the monophyly of Biarmosuchia is relatively low. Many biarmosuchians are known for extensive cranial ornamentation.
Dinocephalia comprises two distinctive groups, the Anteosauria and Tapinocephalia.
Historically, carnivorous dinocephalians, including both anteosaurs and titanosuchids, were called titanosuchians and classified as members of Theriodontia, while the herbivorous Tapinocephalidae were classified as members of Anomodontia.
Anomodontia includes the dicynodonts, a clade of tusked, beaked herbivores, and the most diverse and long-lived clade of non-cynodont therapsids. Other members of Anomodontia include Suminia, which is thought to have been a climbing form.
Gorgonopsia is an abundant but morphologically homogeneous group of saber-toothed predators.
It has been suggested that Therocephalia might not be monophyletic, with some species more closely related to cynodonts than others. [28] However, most studies regard Therocephalia as monophyletic.
Cynodonts are the most diverse and longest-lived of the therapsid groups, as Cynodontia includes mammals. Cynodonts are the only major therapsid clade to lack a Middle Permian fossil record, with the earliest-known cynodont being Charassognathus from the Wuchiapingian age of the Late Permian. Non-mammalian cynodonts include both carnivorous and herbivorous forms.
Synapsida is one of the two major clades of vertebrate animals in the group Amniota, the other being the Sauropsida. The synapsids were the dominant land animals in the late Paleozoic and early Mesozoic, but the only group that survived into the Cenozoic are mammals. 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.
Cynodontia is a clade of eutheriodont therapsids that first appeared in the Late Permian, and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extinct ancestors and close relatives (Mammaliaformes), having evolved from advanced probainognathian cynodonts during the Late Triassic.
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.
Anomodontia is an extinct group of non-mammalian therapsids from the Permian and Triassic periods. By far the most speciose group are the dicynodonts, a clade of beaked, tusked herbivores. Anomodonts were very diverse during the Middle Permian, including primitive forms like Anomocephalus and Patranomodon and groups like Venyukovioidea and Dromasauria. Dicynodonts became the most successful and abundant of all herbivores in the Late Permian, filling ecological niches ranging from large browsers down to small burrowers. Few dicynodont families survived the Permian–Triassic extinction event, but one lineage (Kannemeyeriiformes) evolved into large, stocky forms that became dominant terrestrial herbivores right until the Late Triassic, when changing conditions caused them to decline, finally going extinct during the Triassic–Jurassic extinction event.
Gorgonopsia is an extinct clade of sabre-toothed therapsids from the Middle to the Upper Permian, roughly between 265 and 252 million years ago. They are characterised by a long and narrow skull, as well as elongated upper and sometimes lower canine teeth and incisors which were likely used as slashing and stabbing weapons. Postcanine teeth are generally reduced or absent. For hunting large prey, they possibly used a bite-and-retreat tactic, ambushing and taking a debilitating bite out of the target, and following it at a safe distance before its injuries exhausted it, whereupon the gorgonopsian would grapple the animal and deliver a killing bite. They would have had an exorbitant gape, possibly in excess of 90°, without having to unhinge the jaw.
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.
Therocephalia is an extinct clade of eutheriodont therapsids from the Permian and Triassic periods. 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, and this relationship takes evidence in a variety of skeletal features. Indeed, it had been proposed that cynodonts may have evolved from therocephalians and so that therocephalians as recognised are paraphyletic in relation to cynodonts.
The theriodonts are a major group of therapsids which appeared during the Middle Permian and which includes the gorgonopsians and the eutheriodonts, itself including the therocephalians and the cynodonts.
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.
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.
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 Qingtoushan Formation 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.
Lycaenodon is an extinct genus of biarmosuchian therapsids from the Late Permian of South Africa. It is known from a single species, Lycaenodon longiceps, which was named by South African paleontologist Robert Broom in 1925. Both are small-bodied biarmosuchians. Two specimens are known, and both preserve only the front portions of the skull. These specimens come from the Cistecephalus Assemblage Zone of the Karoo Basin. Broom attributed the back portion of a third skull to Lycaenodon, but subsequent examiners considered it to belong to a gorgonopsian or dinocephalian and not a biarmosuchian. Most of the distinguishing features of Lycaenodon come from its palate. As a member of Biarmosuchia, the most basal group of therapsids, Lycaenodon shares many features with earlier and less mammal-like synapsids like Dimetrodon.
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.
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.
Eutheriodontia is a clade of therapsids which appear during the Middle Permian and which includes therocephalians and cynodonts, this latter group including mammals and related forms.
Anomocephaloidea is a clade of basal anomodont therapsids related to the dicynodonts known from what is now South Africa and Brazil during the Middle Permian. It includes only two species, Anomocephalus africanus from the Karoo Basin of South Africa and Tiarajudens eccentricus from the Paraná Basin of Brazil. Anomocephaloidea was named in 2011 with the discovery of Tiarajudens, although Anomocephalus itself has been known since 1999.