Theriodonts | |
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Restoration of three genus of the theriodont clade: Arctognathus (Gorgonopsia), Lycosuchus (Therocephalia) and Progalesaurus (Cynodontia). | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Clade: | Therapsida |
Clade: | Theriodontia Owen, 1876 |
Groups | |
The theriodonts (clade Theriodontia [lower-alpha 1] ) 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.
In 1876, Richard Owen named a suborder Theriodontia, which he divided into the Cynodontia and the Gomphodontia. [1]
The modern clade concept was devised by James Allen Hopson. In his system, Theriodontia fall into two main groups: the Gorgonopsia and the Eutheriodontia. The latter consist of the Therocephalia and Cynodontia. [2]
Theriodonts appeared at the same time as their sister group within the Neotherapsida, the Anomodontia, about 270 million years ago, in the Middle Permian. Even these early theriodonts were more mammal-like than their anomodont and dinocephalian contemporaries.
Early theriodonts may have been endothermic. Early forms were carnivorous, but several later groups became herbivorous during the Triassic. Theriodont jaws were more mammal-like than was the case of other therapsids, because their dentary was larger, which gave them more efficient chewing ability. Furthermore, several other bones that were on the lower jaw (found in reptiles), moved into the ears, allowing the theriodonts to hear better and their mouths to open wider. This made the theriodonts the most successful group of synapsids.
Eutheriodontia refers to all theriodonts except the gorgonopsians (the most "primitive" group). They included the therocephalians and the cynodonts. The cynodonts include the mammals. The eutheriodonts have larger skulls, accommodating larger brains and improved jaw muscles.
The eutheriodontian theriodonts are one of the two synapsid survivors of the great Permian–Triassic extinction event, the other being the dicynodonts. Therocephalians included both carnivorous and herbivorous forms; both died out after the Early Triassic. The remaining theriodonts, the cynodonts, also included carnivores, such as Cynognathus , as well as newly evolved herbivores (Traversodontidae). While traversodontids for the most part remained medium-sized to reasonably large (the length of the largest species was up to two meters), the carnivorous forms became progressively smaller as the Triassic progressed. They "miniaturised". By the Late Triassic, the small cynodonts included the rodent-like Tritylodontidae (possibly related to or descended from traversodontids), and the tiny, shrew-like, Tritheledontidae, related to the Mammaliaformes. The tritheledontids died out during the Jurassic, and the tritylodontids survived into the Cretaceous, but their relatives, the mammals, continued to evolve. Many mammal groups managed to survive the Cretaceous–Paleogene extinction event, which wiped out the non-avian dinosaurs, allowing the mammals to diversify and dominate the Earth.
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.
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.
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.
Ericiolacerta is an extinct genus of small therocephalian therapsids from the early Triassic of South Africa and Antarctica. Ericiolacerta, meaning "hedgehog lizard", was named by D.M.S. Watson in 1931. The species E. parva is known from the holotype specimen which consists of a nearly complete skeleton found in the Lystrosaurus Assemblage Zone within the Katberg Formation of the Beaufort Group in South Africa, and from a partial jaw found in the Lower Triassic Fremouw Formation in Antarctica. Ericiolacerta was around 20 centimetres (7.9 in) in length, with long limbs and relatively small teeth. It probably ate insects and other small invertebrates. The therocephalians – therapsids with mammal-like heads – were abundant in Permian times, but only a few made it into the Triassic. Ericiolacerta was one of those. It is possible that they gave rise to the cynodonts, the only therapsid group to survive into post-Triassic times. Cynodonts gave rise to mammals.
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.
Eutherocephalia is an extinct clade of advanced therocephalian therapsids. Eutherocephalians are distinguished from the lycosuchids and scylacosaurids, two early therocephalian families. While lycosuchids and scyalosaurids became extinct by the end of the Permian period, eutherocephalians survived the Permian–Triassic extinction event. The group eventually became extinct in the Middle Triassic.
Bauria is an extinct genus of the suborder Therocephalia that existed during the Early and Middle Triassic period, around 246-251 million years ago. It belonged to the family Bauriidae. Bauria was probably a herbivore or omnivore. It lived in South Africa, specifically in the Burgersdorp Formation in South Africa.
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.
Moschorhinus is an extinct genus of therocephalian synapsid in the family Akidnognathidae with only one species: M. kitchingi, which has been found in the Late Permian to Early Triassic of the South African Karoo Supergroup. It was a large carnivorous therapsid, reaching 1.1–1.5 metres (3.6–4.9 ft) in total body length with the largest skull comparable to that of a lion in size, and had a broad, blunt snout which bore long, straight canines.
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.
Scylacops is an extinct genus of Gorgonopsia. It was first named by Broom in 1913, and contains two species, S. bigendens, and S. capensis. Its fossils have been found in South Africa and Zambia. It is believed to be closely related to the Gorgonopsian Sauroctonus progressus. Scylacops was a moderately sized Gorgonopsid.
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.
Akidnognathidae is an extinct family of therocephalian therapsids from the Late Permian and Early Triassic of South Africa, Russia and China. The family includes many large-bodied therocephalians that were probably carnivorous, including Moschorhinus and Olivierosuchus. One akidnognathid, Euchambersia, may even have been venomous. Akidnognathids have robust skulls with a pair of large caniniform teeth in their upper jaws. The family is morphologically intermediate between the more basal therocephalian group Scylacosauridae and the more derived group Baurioidea.
Bauriidae is an extinct family of therocephalian therapsids. Bauriids were the latest-surviving group of therocephalians after the Permian–Triassic extinction event, going extinct in the Middle Triassic. They are among the most advanced eutherocephalians and possess several mammal-like features such as a secondary palate and wide postcanine teeth at the back of the jaws. Unlike other therocephalians, bauriids were herbivorous. They were also smaller than earlier members of the group. Two subfamilies are classified within Bauriidae: Nothogomphodontinae and Bauriinae.
Blattoidealestes is an extinct genus of therocephalian therapsid from the Middle Permian of South Africa. The type species Blattoidealestes gracilis was named by South African paleontologist Lieuwe Dirk Boonstra from the Tapinocephalus Assemblage Zone in 1954. Dating back to the Middle Permian, Blattoidealestes is one of the oldest therocephalians. It is similar in appearance to the small therocephalian Perplexisaurus from Russia, and may be closely related.
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.
Gomphodontia is a clade of cynognathian cynodonts that includes the families Diademodontidae, Trirachodontidae, and Traversodontidae. Gomphodonts are distinguished by wide and closely spaced molar-like postcanine teeth, which are convergent with those of mammals. Other distinguishing characteristics of gomphodonts include deep zygomatic arches, upper postcanines with three or more cusps spanning their widths and lower postcanines with two cusps spanning their widths. They are thought to have been herbivorous or omnivorous. Gomphodonts first appeared in the Early Triassic and became extinct at the end of the Late Triassic. Fossils are known from southern Africa, Argentina and southern Brazil, eastern North America, Europe, China, and Antarctica.