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 armour-like bony plates that distinguish them from the modern soft-bodied lissamphibians.
Eryopidae were a group of medium to large amphibious temnospondyls, known from North America and Europe. They are defined as all eryopoids with interpterygoid vacuities that are rounded at the front; and large external nares. Not all of the genera previously included in the Eryopidae are retained under the cladistic revisions.
The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecological niche to modern crocodilians prior to the diversification of pseudosuchian archosaurs.
Sclerothorax is an extinct genus of temnospondyl from the Early Triassic of Germany. It is distinguished from other temnospondyls by its short and very wide skull and the elongated neural spines that form a ridge along its back. Sclerothorax is a basal member of Capitosauria, a large clade of temnospondyls that lived throughout the Triassic.
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.
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.
Intasuchus is an extinct genus of temnospondyl amphibian from the Middle Permian of Russia. It is known from a single species, Intasuchus silvicola, which was named in 1956. Intasuchus belongs to the family Intasuchidae and is probably its sole member, although other taxa such as Syndyodosuchus and Cheliderpeton have been assigned to the family in the past. Intasuchus most likely belongs to the group Archegosauroidea, Permian relatives of the large, mostly Mesozoic temnospondyl clade Stereospondyli.
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.
Rhinesuchidae is a family of tetrapods that lived primarily in the Permian period. They belonged to the broad group Temnospondyli, a successful and diverse collection of semiaquatic tetrapods which modern amphibians are probably descended from. Rhinesuchids can be differentiated from other temnospondyls by details of their skulls, most notably the interior structure of their otic notches at the back of the skull. They were among the earliest-diverging members of the Stereospondyli, a subgroup of temnospondyls with flat heads and aquatic habits. Although more advanced stereospondyls evolved to reach worldwide distribution in the Triassic period, rhinesuchids primarily lived in the high-latitude environments of Gondwana during the Guadalupian and Lopingian epochs of the Permian. The taxonomy of this family has been convoluted, with more than twenty species having been named in the past; a 2017 review recognized only eight of them to be valid. While several purported members of this group have been reported to have lived in the Triassic period, most are either dubious or do not belong to the group. However, at least one valid genus of rhinesuchid is known from the early Triassic, a small member known as Broomistega. The most recent formal definition of Rhinesuchidae, advocated by Mariscano et al. (2017) is that of a stem-based clade containing all taxa more closely related to Rhinesuchus whaitsi than to Lydekkerina huxleyi or Peltobatrachus pustulatus. A similar alternate definition is that Rhinesuchidae is a stem-based clade containing all taxa more closely related to Uranocentrodon senekalensis than to Lydekkerina huxleyi, Trematosaurus brauni, or Mastodonsaurus giganteus.
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.
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.
Rhineceps is an extinct genus of temnospondyl amphibian in the family Rhinesuchidae. Rhineceps was found in Northern Malawi in Southern Africa known only from its type species R. nyasaensis. Rhineceps was a late Permian semi-aquatic carnivore that lived in streams, rivers, lakes or lagoons. Rhineceps is an early divergent Stereopondyl within the family Rhinesuchidae, which only existed in the late Permian (Lopingian) and failed to survive the Permian-Triassic extinction unlike other stereospondyl families.
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.
Palatinerpeton is an extinct genus of temnospondyl amphibian. Fossils have been found in the Lauterecken-Odernheim Formation of Germany.
Rhachitomi is a group of temnospondyl amphibians that includes all temnospondyls except edopoids and dendrerpetontids. It was established as a clade name by German paleontologist Rainer R. Schoch in 2013, although the name had first been established in 1919 by British paleontologist D. M. S. Watson to encompass an evolutionary grade of temnospondyls leading to the group Stereospondyli. American paleontologist Alfred Romer used the term in a similar sense, grouping most Permian and Triassic temnospondyls under Rhachitomi. A similar name that appeared earlier in the scientific literature is Rachitomi, which was named by American paleontologist Edward Drinker Cope in 1882. Rachitomi was commonly used in the late nineteenth and early twentieth century to include early amphibians such as Eryops and Archegosaurus that had rhachitomous vertebrae. Many early tetrapods have vertebrae that are split into two parts below the notochord: a pleurocentrum and an intercentrum. In rhachitomous vertebrae, the intercentrum is large and semicircular, while the pleurocentrum divided into two smaller paired elements. Schoch defined Rhachitomi as a node-based taxon to include four major and well-supported clades of temnospondyls: Dvinosauria, Eryopidae, Stereospondyli and a clade formed by Zatracheidae and Dissorophoidea. Not all members of Rhachitomi have rhachitomous vertebrae; the largest subgroup, Stereospondyli, lacks pleurocentra. Below is a cladogram from Schoch's analysis showing the placement of Rhachitomi within Temnospondyli:
Eryopiformes is a group of rhachitomi temnospondyls. Eryopiformes was named by the German paleontologist Rainer R. Schoch in 2013. He defined it as a node-based taxon to include Eryopidae and Stereospondylomorpha. Unlike previous analyses, a phylogenetic analysis he performed found no support for a monophyletic Euskelia, that is defined to include Dissorophoidea and Eryopidae, in relation to Stereospondylomorpha. In this analysis, Eryopidae and Stereospondylomorpha usually formed a monophyletic clade that excludes the clade formed by Zatracheidae and Dissorophoidea. Thus, he named this newly identified group as Eryopiformes.
Chinlestegophis is a diminutive Late Triassic stereospondyl that has been interpreted as a putative stem caecilian, a living group of legless burrowing amphibians. If Chinlestegophis is indeed both an advanced stereospondyl and a relative of caecilians, this means that stereospondyls survived to the present day; historically the group was thought to have gone extinct by the early Cretaceous. Chinlestegophis jenkinsi, the type and only species, is known from two partial skulls discovered in the Chinle Formation in Colorado.
