Dvinosauria Temporal range: Late Carboniferous - Early Triassic, | |
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Dvinosaurus primus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Temnospondyli |
Suborder: | † Dvinosauria Yates and Warren, 2000 |
Subgroups | |
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Dvinosaurs are one of several new clades of temnospondyls named in the phylogenetic review of the group by Yates and Warren 2000. They represent a group of primitive semi-aquatic to completely aquatic temnospondyls, and are known from the Late Carboniferous to the Early Triassic, being most common in the Permian period. Their distinguishing characteristics are a reduction of the otic notch; the loss of a flange on the rear side of the pterygoid; and 28 or more presacral vertebrae.
Trimerorhachidae is the most basal family of dvinosaurs. Most other dvinosaurs are placed in the superfamily Dvinosauroidea. Within Dvinosauroidea are two families, Eobrachyopidae and Tupilakosauridae, as well as dvinosaurs that cannot be placed in either family, such as Dvinosaurus and Kourerpeton . A 2008 phylogenetic analysis found Eobrachyopidae to be paraphyletic, representing a grade of basal dvinosauroids. Below is a cladogram showing the phylogenetic relationships of dvinosaurs from Englehorn et al. (2008): [2]
Dvinosauria |
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Temnospondyli or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carboniferous, Permian and Triassic periods, with fossils being found on every continent. A few species continued into the Jurassic and Early Cretaceous periods, but all had gone extinct by the Late Cretaceous. During about 210 million years of evolutionary history, they adapted to a wide range of habitats, including freshwater, terrestrial, and even coastal marine environments. Their life history is well understood, with fossils known from the larval stage, metamorphosis and maturity. Most temnospondyls were semiaquatic, although some were almost fully terrestrial, returning to the water only to breed. These temnospondyls were some of the first vertebrates fully adapted to life on land. Although temnospondyls are amphibians, many had characteristics such as scales and large armour-like bony plates (osteoderms) that generally distinguish them from the modern soft-bodied lissamphibians.
Trematosauria is one of two major groups of temnospondyl amphibians that survived the Permian-Triassic extinction event, the other being the Capitosauria. The trematosaurs were a diverse and important group that included many medium-sized to large forms that were semi-aquatic to fully aquatic. The group included long-snouted forms such as the trematosauroids and short, broad-headed forms such as the metoposaurs. Although most groups did not survive beyond the Triassic, one lineage, the brachyopoids, continued until the Cretaceous period. Trematosauria is defined as all stereospondyls more closely related to Trematosaurus than to Parotosuchus, a capitosaurian.
Euskelia is a proposed clade of extinct temnospondyl amphibians. The naming derives from the ancient Greek eu, meaning "true", and skelos, meaning "limb", in reference to well-ossified limb bones with crests to which muscles were attached. Members of this group have the most ossified skeleton of all temnospondyls.
Sclerocephalus is an extinct genus of temnospondyl amphibian from the lowermost Permian of Germany and Czech Republic with four valid species, including the type species S. haeuseri. It is one of the most completely preserved and most abundant Palaeozoic tetrapods. Sclerocephalus was once thought to be closely related to eryopoid temnospondyls, but it is now thought to be more closely related to archegosauroids. It is the only genus in the family Sclerocephalidae.
Dvinosaurus is an extinct genus of amphibious temnospondyls localized to regions of western and central Russia during the middle and late Permian, approximately 265-254 million years ago. Its discovery was first noted in 1921 by Russian paleontologist Vladimir Prokhorovich Amalitskii in a posthumously published paper that documents the findings of a site in Russia's Arkhangelsk District. Its name is derived from the proximity of this site to the Northern Dvina River.
Konzhukovia is an amphibian genus that belongs to an extinct family Konzhukoviidae of temnospondyls, the largest clade of basal tetrapods including about 198 genera, 292 species, and more than half of which were alive during the early Mesozoic period. The animal was a predator that lived about 260 million years ago, and could get up to about three meters in length. Specifically, Konzukovia lived during the Permian, between 252 and 270 million years ago according to the type of rock the fossil was found in. There are three species within this genus, K. vetusta, K. tarda, and K. sangabrielensis, the first two originating from Russia while the latest originating from Southern Brazil. The discovery of this specimen in Southern Brazil provided more evidence to support the idea that during this animals existence, there was a “biological corridor” because of the supercontinent Pangea, allowing these species to be found so far apart from each other. Konzhukovia belongs to the family Archegosauridae, a family consisted of large temnospondyls that most likely compare to modern day crocodiles. Since the discovery of the latest species, K. sangabrielensis, Pacheco proposes that there must be the creation of a new family, Konzhokoviidae, a monophyletic group in a sister-group relationship with Stereospondlyi in order to accommodate the three species. Konzhukovia skulls usually exhibit typical rhinesuchid features including an overall parabolic shape, small orbits located more posteriorly, and the pterygoids do not reach the vomer. These animals were long-snouted amphibians that had clear adaptations made for fish catching, as well as exemplifying aquatic features.
Brachyopoidea is a superfamily of temnospondyls that lived during the Mesozoic. It contains the families Brachyopidae and Chigutisauridae. The earliest records of brachyopids are from the Lower Triassic in Australia. The latest-surviving member of the superfamily is the chigutisaurid Koolasuchus from the Early Cretaceous of Australia.
Lydekkerinidae is a family of stereospondyl temnospondyls that lived in the Early Triassic period. During this time period, lydekkerinids were widely distributed, with putative remains reported from Russia, Greenland, India, South Africa, Madagascar, Australia, and Antarctica. In contrast to most other stereospondyls, lydekkerinids were relatively small-bodied. The type genus is Lydekkerina, the namesake of the family and the best-known lydekkerinid.
Stereospondylomorpha is a clade of temnospondyls. It includes the superfamily Archegosauroidea and the more diverse group Stereospondyli. Stereospondylomorpha was first proposed by Yates and Warren (2000), who found Archegosauroidea and Stereospondyli to be sister taxa in their phylogenetic analysis. A similar clade is Archegosauriformes, named by Schoch and Milner (2000), which includes Stereospondyli and some Permian temnospondyls that are similar in appearance to stereospondyls, including the archegosauroids. However, according to Schoch and Milner's phylogeny, Archegosauroidea is a paraphyletic group of taxa that are successively basal to Stereospondyli, rather than a monophyletic sister taxon.
Limnarchia is a clade of temnospondyls. It includes the mostly Carboniferous-Permian age Dvinosauria and the mostly Permian-Triassic age Stereospondylomorpha. The clade was named in a 2000 phylogenetic analysis of stereospondyls and their relatives. Limnarchia means "lake rulers" in Greek, in reference to their aquatic lifestyles and long existence over a span of approximately 200 million years from the Late Carboniferous to the Early Cretaceous. In phylogenetic terms, Limnarchia is a stem-based taxon including all temnospondyls more closely related to Parotosuchus than to Eryops. It is the sister group of the clade Euskelia, which is all temnospondyls more closely related to Eryops than to Parotosuchus. Limnarchians represent an evolutionary radiation of temnospondyls into aquatic environments, while euskelians represent a radiation into terrestrial environments. While many euskelians were adapted to life on land with strong limbs and bony scutes, most limnarchians were better adapted for the water with poorly developed limbs and lateral line sensory systems in their skulls.
Acroplous is an extinct genus of dvinosaurian Temnospondyli within the family Eobrachyopidae.
Broiliellus is an extinct genus of dissorophoid temnospondyl within the family Dissorophidae. Broiliellus is most closely related to the genus Dissorophus, and both have been placed in the subfamily Dissorophinae. Broiliellus is known from five species from the Early Permian: the type species is Broiliellus texensis, and the other species are Broiliellus brevis,Broiliellus olsoni, Broiliellus arroyoensis, and Broiliellus reiszi. An additional species, Broiliellus novomexicanus, which was originally named Aspidosaurus novomexicanus, is now thought to fall outside the genus as a member of the subfamily Eucacopinae.
Lapillopsis is an extinct genus of stereospondyl temnospondyl within the family Lapillopsidae. Fossils belonging to the genus have been found in the Arcadia Formation of Queensland, Australia.
Thoosuchus is an extinct genus of basal trematosauroid trematosaurian temnospondyl. Fossils have been found from Russia and date back to the Early Triassic. It is the type genus of the family Thoosuchidae, formerly called the subfamily Thoosuchinae and placed within Benthosuchidae. The benthosuchids were originally composed of the majority of basal trematosaurian forms regarded as the ancestors of the trematosaurids.
Capitosauria is an extinct group of large temnospondyl amphibians with simplified stereospondyl vertebrae. Mainly living as piscivores in lakes and rivers, the Capitosauria and its sister taxon Trematosauria were the only major labyrinthodonts that existed during the Mesozoic in ecological niches broadly similar to those of modern crocodiles, and some grew to very large sizes. At 6 meters in length, the Mid-Triassic Mastodonsaurus giganteus is not only thought to have been the largest capitosaur, but possibly also the largest amphibian to have lived. The latest known remains are from the Rhaetian of Germany and are referred to Cyclotosaurus.
Rhytidostea is a clade of stereospondyl temnospondyls. It was erected in 2000 to include several temnospondyl groups distinct from the "higher" group of capitosaurs, including lydekkerinids, brachyopoids, and rhytidosteids. Rhytidosteans first appeared in the Permian period and underwent an evolutionary radiation during the Induan stage of the Early Triassic. Along with capitosaurs, rhytidosteans comprise much of the larger suborder Stereospondyli. Rhytidostea has often been considered the sister group of the clade Capitosauria, but has been placed in various other phylogenetic positions. In many studies, members of Rhytidostea are split, with lydekkerinids having a more basal position among stereospondyls while rhytidosteids and brachyopoids form a group placed among the more derived trematosaurian stereospondyls.
Olsoniformes is an extinct clade of dissorophoid temnospondyls, including the families Dissorophidae and Trematopidae. Most members of the clade were highly adapted to a terrestrial lifestyle. The clade was named in 2008 and is defined as the least inclusive clade containing Dissorophus multicinctus and Acheloma cumminsi but not Amphibamus grandiceps, Micromelerpeton credneri, or Apateon pedestris. Olsoniforms share various features such as a stout and low ilium and a thin cultriform process. The earliest-branching olsoniform is Palodromeus bairdi, from the Late Carboniferous of Ohio.
Perryella is an extinct genus of dvinosaurian(?) temnospondyl from the Permian of Oklahoma.
Edopoidea is a clade of primitive temnospondyl amphibians including the genus Edops and the family Cochleosauridae. Edopoids are known from the Late Carboniferous and Early Permian of North America and Europe, and the Late Permian of Africa. They are among the most basal temnospondyls, and possess a number of primitive features that were lost in later members of the group.
Vigilius is an extinct genus of brachyopid temnospondyl amphibian from the Triassic of Arizona. It is known from the single type species Vigilius wellesi.