Euskelia Temporal range: Pennsylvanian - Early Triassic, | |
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Mounted skeleton of Eryops , American Museum of Natural History | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Temnospondyli |
Suborder: | † Euskelia Yates and Warren, 2000 |
Superfamilies | |
Synonyms | |
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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. [1] Members of this group have the most ossified skeleton of all temnospondyls. [2]
Euskelia is a stem-based taxon including all temnospondyls more closely related to Eryops (an eryopoid) than to Parotosuchus (a stereospondyl). The clade was named by Yates & Warren (2000), whose phylogenetic analysis argued that eryopoids were more closely related to dissorophoids than to stereospondyls. Euskelia was intended to encompass the terrestrial eryopoid+dissorophoid clade, opposite to the clade Limnarchia, which included aquatic groups such as dvinosaurs and stereospondylomorphs. [1]
Other studies propose a different structure of the temnospondyl family tree. For example, Schoch (2013) considered eryopoids to be closer to stereospondylomorphs than to dissorophoids. That study offered the name Eryopiformes for the eryopoid+stereospondylomorph clade, excluding dissorophoids. [3]
Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco, lepospondyls lived from the Early Carboniferous (Mississippian) to the Early Permian and were geographically restricted to what is now Europe and North America. Five major groups of lepospondyls are known: Adelospondyli; Aïstopoda; Lysorophia; Microsauria; and Nectridea. Lepospondyls have a diverse range of body forms and include species with newt-like, eel- or snake-like, and lizard-like forms. Various species were aquatic, semiaquatic, or terrestrial. None were large, and they are assumed to have lived in specialized ecological niches not taken by the more numerous temnospondyl amphibians that coexisted with them in the Paleozoic. Lepospondyli was named in 1888 by Karl Alfred von Zittel, who coined the name to include some tetrapods from the Paleozoic that shared some specific characteristics in the notochord and teeth. Lepospondyls have sometimes been considered to be either related or ancestral to modern amphibians or to Amniota. It has been suggested that the grouping is polyphyletic, with aïstopods being primitive stem-tetrapods, while recumbirostran microsaurs are primitive reptiles.
Dissorophoidea is a clade of medium-sized, temnospondyl amphibians that appeared during the Moscovian in Euramerica, and continued through to the Late Permian and the Early Triassic of Gondwana. They are distinguished by various details of the skull, and many species seem to have been well adapted for life on land.
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.
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.
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.
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.
Almasaurus is an extinct genus of trematosaurian temnospondyl within the family Latiscopidae. It is known from several skulls and some postcranial material found from the Argana Formation in Morocco, which dates back to the Late Triassic.
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.
Rileymillerus is an extinct genus of temnospondyl amphibian from the Late Triassic Post Quarry in the Dockum Group of Texas that was described by John Bolt and Sankar Chatterjee in 2000. The holotype, a nearly complete skull with articulated jaws, is housed at the Museum of Texas Tech University. The genus is named for Riley Miller, who allowed Chatterjee to work on the Post Quarry, and the species is named for the paleontologist John Cosgriff.
Stegops is an extinct genus of euskelian temnospondyl from the Late Carboniferous of the eastern United States. Fossils are known from the Pennsylvanian coal deposits of Linton, Ohio. It was once classified in the eryopoid family Zatrachydidae because it and other zatrachydids have spikes extending from the margins of its skull, but it is now classified as a dissorophoid that independently evolved spikes. Stegops was first named by American paleontologist Edward Drinker Cope in 1885, with his description of the type species Stegops divaricata. Cope had also named a species of Sauropleura from Linton in 1875, which he called Sauropleura newberryi. This species was later synonymized with Stegops divaricata when the type specimen of S. newberryi was prepared and found to be a large specimen of Stegops.
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.
Perryella is an extinct genus of dvinosaurian(?) temnospondyl from the Permian of Oklahoma.
Eutemnospondyli is a clade of temnospondyl amphibians that includes most temnospondyls except edopoids. Eutemnospondyli was named by German paleontologist Rainer R. Schoch in 2013. He defined it as a stem-based taxon including all temnospondyls more closely related to Stereospondyli than to Edopoidea. In his phylogenetic analysis, Eutemnospondyli included dendrerpetontids and a clade he referred to as Rhachitomi. Rhachitomi is defined to include four major and well-supported clades of temnospondyls: Dvinosauria, Eryopidae, Stereospondyli and a clade formed by Zatracheidae and Dissorophoidea. Below is a cladogram from Schoch's analysis:
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.
Manubrantlia was a genus of lapillopsid temnospondyls from the Early Triassic Panchet Formation of India. This genus is only known from a single holotype left jaw, given the designation ISI A 57. Despite the paucity of remains, the jaw is still identifiable as belonging to a relative of Lapillopsis. For example, all three of its coronoid bones possessed teeth, the articular bone is partially visible in lateral (outer) view, and its postsplenial does not contact the posterior meckelian foramen. However, the jaw also possesses certain unique features which justify the erection of a new genus separate from Lapillopsis. For example, the mandible is twice the size of any jaws referred to other lapillopsids. The most notable unique feature is an enlarged "pump-handle" shaped arcadian process at the back of the jaw. This structure is responsible for the generic name of this genus, as "Manubrantlia" translates from Latin to the English expression "pump-handle". The type and only known species of this genus is Manubrantlia khaki. The specific name refers to the greenish-brown mudstones of the Panchet Formation, with a color that had been described as "khaki" by the first British geologists who studied the formation.