Therapsid

Last updated

Therapsids
Temporal range: CisuralianHolocene 279.5–0 Ma(Range includes mammals)
Therapsida 3.jpg
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 OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Sphenacodontia
Clade: Pantherapsida
Clade: Sphenacodontoidea
Clade: Therapsida
Broom, 1905 [1]
Clades

A therapsid is a member of the clade Therapsida, [lower-alpha 1] which is 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, as opposed to the sprawling posture of many reptiles and salamanders.

Contents

Therapsids evolved from "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 reptiles 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. Of the non-mammalian therapsids, only cynodonts survived beyond the end of the Triassic, with the only other remaining group of therapsids to have survived into the Late Triassic, the dicynodonts, becoming 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.

Characteristics

Illustration of Alopecognathus, an early therocephalian therapsid Pristeroognathus DB.jpg
Illustration of Alopecognathus , an early therocephalian therapsid

Jaw and teeth

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.

Posture

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]

Physiology

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]

Integument

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. Fossilized facial skin from the dinocephalian Estemmenosuchus has been described as showing that the skin was glandular and lacked both scales and hair. [8]

Coprolites containing what appear to be hairs have been found from the Permian. [9] [10] 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. [11] This suggests that hair may have first evolved in probainognathians, though it does not entirely rule out an earlier origin of fur. [11]

Whiskers probably evolved in probainognathian cynodonts. [11] [12] 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, [13] which would make the presence of mammal-like whiskers unlikely. [12]

Evolutionary history

Holotype skull of Raranimus dashankouensis, the most basal known therapsid Raranimus dashankouensis.jpg
Holotype skull of Raranimus dashankouensis , the most basal known therapsid

Therapsids evolved from a group of pelycosaurs called sphenacodonts. [15] [16] 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.

Restoration of Euchambersia with dicynodont prey. Note that this South African therocephalian is suspected to be the oldest known venomous tetrapod. Euchambersia DB.jpg
Restoration of Euchambersia with dicynodont prey. Note that this South African therocephalian is suspected to be the oldest known venomous tetrapod.

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. The dicynodonts, now represented by a single group of large stocky herbivores, the Kannemeyeriiformes, 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.

Some exceptions were the still further derived eucynodonts. [18] 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.

Reconstruction of Bonacynodon schultzi, a probainognathian cynodont related to the ancestors of mammals Bonacynodon witout visible ears.png
Reconstruction of Bonacynodon schultzi , a probainognathian cynodont related to the ancestors of mammals

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.

Classification

Therapsida
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. [20] 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. [21] Tetraceratops from the Early Permian of the United States has been hypothesized to be an even earlier-diverging therapsid, [22] [23] but more recent study has suggested it is more likely to be a non-therapsid sphenacodontian. [24]

Biarmosuchia

Biarmosuchus, a biarmosuchian Biarmosuchus BW.jpg
Biarmosuchus , a biarmosuchian

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

Two genera of dinocephalians : Titanophoneus (an anteosaur) devouring a Ulemosaurus (a tapinocephalian) Titanophoneus 3.jpg
Two genera of dinocephalians  : Titanophoneus (an anteosaur) devouring a Ulemosaurus (a tapinocephalian)

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

Lystrosaurus, a dicynodont anomodont Lystr georg1DB.jpg
Lystrosaurus , a dicynodont anomodont

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

Inostrancevia, a gorgonopsian Inostrancevia BW.jpg
Inostrancevia , a gorgonopsian

Gorgonopsia is an abundant but morphologically homogeneous group of saber-toothed predators.

Therocephalia

Moschorhinus, a therocephalian Moschorhinus DB.jpg
Moschorhinus , a therocephalian

It has been suggested that Therocephalia might not be monophyletic, with some species more closely related to cynodonts than others. [25] However, most studies regard Therocephalia as monophyletic.

Cynodontia

Trucidocynodon, a non-mammalian cynodont Trucidocynodon riograndensis.jpg
Trucidocynodon , a non-mammalian cynodont

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.

See also

Notes

  1. Greek: 'beast-arch'

Related Research Articles

<span class="mw-page-title-main">Synapsida</span> Clade of tetrapods

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.

<span class="mw-page-title-main">Cynodont</span> Clade of therapsids

Cynodonts are 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.

<span class="mw-page-title-main">Dinocephalia</span> Extinct clade of stem-mammals

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.

<span class="mw-page-title-main">Gorgonopsia</span> Extinct group of saber-toothed therapsids from the Permian

Gorgonopsia is an extinct clade of sabre-toothed therapsids from the Middle to Upper Permian roughly 265 to 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.

<span class="mw-page-title-main">Biarmosuchia</span> Extinct suborder of therapsids

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.

<span class="mw-page-title-main">Anteosaur</span> Extinct clade of therapsids

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.

<span class="mw-page-title-main">Therocephalia</span> Extinct order of therapsids

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.

<span class="mw-page-title-main">Theriodontia</span> Clade of therapsids

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.

<i>Anteosaurus</i> Extinct genus of anteosaurid synapsid from the Permian

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. 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.

<i>Tapinocephalus</i> Assemblage Zone

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.

<i>Eodicynodon</i> Extinct genus of dicynodonts

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.

<i>Paraburnetia</i> Extinct genus of therapsids

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.

<i>Biseridens</i> Extinct genus of therapsids

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.

<i>Dimacrodon</i> Extinct genus of synapsids

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.

<span class="mw-page-title-main">Venyukovioidea</span> Extinct infraorder of therapsids

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.

<span class="mw-page-title-main">Abrahamskraal Formation</span> Geological formation of the Beaufort Group in South Africa

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.

<span class="mw-page-title-main">Eutheriodontia</span> Clade of therapsids

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.

<span class="mw-page-title-main">Anomocephaloidea</span> Extinct clade of therapsids

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.

References

  1. Broom, R. (1905). "On the use of the term Anomodontia". Records of the Albany Museum. 1 (4): 266–269.
  2. Carroll, R. L. (1988). Vertebrate Paleontology and Evolution. New York: W. H. Freeman and Company. pp.  698. ISBN   978-0-7167-1822-2.
  3. Benoit, Julien; Abdala, Fernando; Manger, Paul; Rubidge, Bruce (2016). "The sixth sense in mammalians forerunners: variability of the parietal foramen and the evolution of the pineal eye in South African Permo-Triassic eutheriodont therapsids". Acta Palaeontologica Polonica. doi: 10.4202/app.00219.2015 . ISSN   0567-7920. S2CID   59143925.
  4. Rey, Kévin; Amiot, Romain; Fourel, François; Abdala, Fernando; Fluteau, Frédéric; Jalil, Nour-Eddine; Liu, Jun; Rubidge, Bruce S; Smith, Roger MH; Steyer, J Sébastien; Viglietti, Pia A; Wang, Xu; Lécuyer, Christophe (18 July 2017). "Oxygen isotopes suggest elevated thermometabolism within multiple Permo-Triassic therapsid clades". eLife. 6: –28589. doi: 10.7554/eLife.28589 . ISSN   2050-084X. PMC   5515572 . PMID   28716184.
  5. Faure-Brac, Mathieu G.; Cubo, Jorge (2 March 2020). "Were the synapsids primitively endotherms? A palaeohistological approach using phylogenetic eigenvector maps". Philosophical Transactions of the Royal Society B: Biological Sciences. 375 (1793): 20190138. doi:10.1098/rstb.2019.0138. ISSN   1471-2970. PMC   7017441 . PMID   31928185.
  6. Newham, Elis; Gill, Pamela G.; Brewer, Philippa; Benton, Michael J.; Fernandez, Vincent; Gostling, Neil J.; Haberthür, David; Jernvall, Jukka; Kankaanpää, Tuomas; Kallonen, Aki; Navarro, Charles; Pacureanu, Alexandra; Richards, Kelly; Brown, Kate Robson; Schneider, Philipp; Suhonen, Heikki; Tafforeau, Paul; Williams, Katherine A.; Zeller-Plumhoff, Berit; Corfe, Ian J. (2020). "Reptile-like physiology in Early Jurassic stem-mammals". Nature Communications. 11 (1): 5121. Bibcode:2020NatCo..11.5121N. doi:10.1038/s41467-020-18898-4. ISSN   2041-1723. PMC   7550344 . PMID   33046697.
  7. Hopson, James A. (18 October 2012). "The Role of Foraging Mode in the Origin of Therapsids: Implications for the Origin of Mammalian Endothermy". Fieldiana Life and Earth Sciences. 5: 126–148. doi:10.3158/2158-5520-5.1.126. ISSN   2158-5520. S2CID   84471370.
  8. Chudinov, P. K. (1968). "Structure of the integuments of theromorphs". Doklady Akad. Nauk SSSR. 179: 226–229.
  9. Smith, Roger M.H.; Botha-Brink, Jennifer (2011). "Morphology and composition of bone-bearing coprolites from the Late Permian Beaufort Group, Karoo Basin, South Africa". Palaeogeography, Palaeoclimatology, Palaeoecology. 312 (1–2): 40–53. Bibcode:2011PPP...312...40S. doi:10.1016/j.palaeo.2011.09.006. ISSN   0031-0182.
  10. Bajdek, Piotr; Qvarnström, Martin; Owocki, Krzysztof; Sulej, Tomasz; Sennikov, Andrey G.; Golubev, Valeriy K.; Niedźwiedzki., Grzegorz (2016). "Microbiota and food residues including possible evidence of pre-mammalian hair in Upper Permian coprolites from Russia". Lethaia. 49 (4): 455–477. doi:10.1111/let.12156.
  11. 1 2 3 Benoit, J.; Manger, P. R.; Rubidge, B. S. (9 May 2016). "Palaeoneurological clues to the evolution of defining mammalian soft tissue traits". Scientific Reports. 6 (1): 25604. Bibcode:2016NatSR...625604B. doi:10.1038/srep25604. ISSN   2045-2322. PMC   4860582 . PMID   27157809.
  12. 1 2 Benoit, Julien; Ruf, Irina; Miyamae, Juri A.; Fernandez, Vincent; Rodrigues, Pablo Gusmão; Rubidge, Bruce S. (2020). "The Evolution of the Maxillary Canal in Probainognathia (Cynodontia, Synapsida): Reassessment of the Homology of the Infraorbital Foramen in Mammalian Ancestors". Journal of Mammalian Evolution. 27 (3): 329–348. doi:10.1007/s10914-019-09467-8. ISSN   1573-7055. S2CID   156055693.
  13. Estes, Richard (1961). "Cranial anatomy of the cynodont reptile Thrinaxodon liorhinus". Bulletin of the Museum of Comparative Zoology. 125: 165–180.
  14. Liu, J.; Rubidge, B; Li, J. (2009). "New basal synapsid supports Laurasian origin for therapsids" (PDF). Acta Palaeontologica Polonica. 54 (3): 393–400. doi: 10.4202/app.2008.0071 . Retrieved 25 September 2009.
  15. Synapsid Classification & Apomorphies
  16. Huttenlocker, Adam. K.; Rega, Elizabeth (2012). "Chapter 4. The Paleobiology and Bone Microstructure of Pelycosauriangrade Synapsids". In Chinsamy-Turan, Anusuya (ed.). Forerunners of Mammals: Radiation, Histology, Biology. Indiana University Press. pp. 90–119. ISBN   978-0253005335.
  17. Benoit, J.; Norton, L.A.; Manger, P.R.; Rubidge, B.S. (2017). "Reappraisal of the envenoming capacity of Euchambersia mirabilis (Therapsida, Therocephalia) using μCT-scanning techniques". PLOS ONE. 12 (2): e0172047. Bibcode:2017PLoSO..1272047B. doi: 10.1371/journal.pone.0172047 . PMC   5302418 . PMID   28187210.
  18. Padian, Kevin (4 September 2013). "A Review of "Forerunners of Mammals: Radiation, Histology, Biology"". Journal of Vertebrate Paleontology . 33 (5): 1250–1251. doi:10.1080/02724634.2013.763814. ISSN   0272-4634.
  19. Geggel, L. (2016). "Meet the Ancient Reptile that Gave Rise to Mammals". Scientific American.
  20. Angielczyk, Kenneth D.; Kammerer, Christian F. (22 October 2018). "Non-Mammalian synapsids: the deep roots of the mammalian family tree". In Zachos, Frank; Asher, Robert (eds.). Mammalian Evolution, Diversity and Systematics. De Gruyter. pp. 117–198. doi:10.1515/9783110341553-005. ISBN   978-3-11-034155-3. S2CID   92370138.
  21. Duhamel, A.; Benoit, J.; Rubidge, B. S.; Liu, J. (August 2021). "A re-assessment of the oldest therapsid Raranimus confirms its status as a basal member of the clade and fills Olson's gap". The Science of Nature. 108 (4): 26. Bibcode:2021SciNa.108...26D. doi:10.1007/s00114-021-01736-y. ISSN   0028-1042. PMID   34115204. S2CID   235403632.
  22. Laurin, M.; Reisz, R. R. (1996). "The osteology and relationships of Tetraceratops insignis, the oldest known therapsid". Journal of Vertebrate Paleontology. 16 (1): 95–102. doi:10.1080/02724634.1996.10011287.
  23. Amson, Eli; Laurin, Michel (2011). "On the Affinities of Tetraceratops insignis, an Early Permian Synapsid". Acta Palaeontologica Polonica. 56 (2): 301–312. doi: 10.4202/app.2010.0063 . ISSN   0567-7920. S2CID   56425905.
  24. Spindler, Frederik (2020). "The skull of Tetraceratops insignis (Synapsida, Sphenacodontia)". Palaeovertebrata. 43 (1): e1. doi:10.18563/pv.43.1.e1. S2CID   214247325.
  25. Abdala, Fernando; Rubidge, Bruce S.; van den Heever, Juri (2008). "The oldest therocephalians (Therapsida, Eutheriodontia) and the early diversification of Therapsida". Palaeontology. 51 (4): 1011–1024. doi: 10.1111/j.1475-4983.2008.00784.x . ISSN   1475-4983. S2CID   129791548.

Further reading