Utaherpeton | |
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Restoration of Utaherpeton franklini | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Subclass: | † Lepospondyli |
Genus: | † Utaherpeton Carroll et al., 1991 |
Type species | |
†Utaherpeton franklini Carroll et al., 1991 |
Utaherpeton is an extinct genus of lepospondyl amphibian from the Carboniferous of Utah. It is one of the oldest and possibly one of the most basal ("primitive") known lepospondyls. The genus is monotypic, including only the type species Utaherpeton franklini. Utaherpeton was named in 1991 from the Manning Canyon Shale Formation, which dates to the Mississippian-Pennsylvanian boundary. [1] It was originally classified within Microsauria, a group of superficially lizard- and salamander-like lepospondyls that is now no longer considered to be a valid clade or evolutionary grouping, but rather an evolutionary grade consisting of the most basal lepospondyls. Utaherpeton has been proposed as both the most basal lepospondyl and the oldest "microsaur", although more derived lepospondyls are known from earlier in the Carboniferous. However, its position within Lepospondyli remains uncertain due to the incomplete preservation of the only known specimen. The inclusion of Utaherpeton in various phylogenetic analyses has resulted in multiple phylogenies (hypotheses of evolutionary relationships) that are very different from one another, making it a significant taxon in terms of understanding the interrelationships of lepospondyls.
Like many other lepospondyls, Utaherpeton is salamander-like in appearance. It has an elongated body made up of 26 presacral (neck and back) vertebrae. The total body length excluding the tail is about 4 centimetres (1.6 in). [1] The forelimbs are markedly smaller than the hind limbs and the hind feet are unusual in that they are larger than the rest of the hind limbs. A unique anatomical feature that distinguishes Utaherpeton from other "microsaur" lepospondyls is a connection between the prefrontal and premaxilla bones at the front of the skull. [1]
When Utaherpeton was first described in 1991, it was tentatively classified within a group of "microsaurs" called Microbrachomorpha, which included the better-known Microbrachis . [1] A 2001 phylogenetic analysis found Utaherpeton to be the most basal lepospondyl, followed by the "microsaur" Hyloplesion , which has also been considered a microbrachomorph. However, Microbrachomorpha was not recognized as a true clade because Microbrachis was found to be more deeply nested within Lepospondyli, more closely related to derived lepospondyls like nectrideans and aïstopods than to other "microsaurs". [2] A 2004 analysis including Utaherpeton produced a very different phylogeny for lepospondyls, placing aïstopods and nectrideans as the most basal lepospondyls, followed by a paraphyletic assemblage of "microsaurs". Utaherpeton was still recovered as one of the most basal "microsaurs", but not as the most basal lepospondyl. Tentative support for a close relationship between Utaherpeton and Microbrachis was also found. [3] In a 2010 analysis, all "microsaurs" except Utaherpeton were recovered within a monophyletic group, but Utaherpeton was found to be the most basal member of a separate clade that included all other lepospondyls. [4]
Gymnophiona is the group of amphibians that includes the legless caecilians and all amphibians more closely related to them than to frogs or salamanders. The name derives from the Greek words γυμνος and οφις, as the caecilians were originally thought to be related to snakes. The body is cylindrical dark brown or bluish black in colour. The skin is slimy and bears grooves or ringlike markings; there are minute dermal scales.
Labyrinthodontia is an extinct amphibian subclass, which constituted some of the dominant animals of late Paleozoic and early Mesozoic eras. The group evolved from lobe-finned fishes in the Devonian and is ancestral to all extant landliving vertebrates. As such it constitutes an evolutionary grade rather than a natural group (clade). The name describes the pattern of infolding of the dentin and enamel of the teeth, which are often the only part of the creatures that fossilize. They are also distinguished by a heavily armoured skull roof, and complex vertebrae, the structure of which were used in older classifications of the group.
Lepospondyli is a diverse taxon of amphibian 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.
Lysorophia is an order of aquatic Carboniferous and Permian tetrapods within the extinct subclass Lepospondyli. Lysorophians resembled small snakes, as their bodies are extremely elongate. There is a single family, the Molgophidae. Currently there are around five genera included within Lysorophia, although many may not be valid.
Adelospondyli is an order of elongated, presumably aquatic, Carboniferous amphibians. They have a robust skull roofed with solid bone, and orbits located towards the front of the skull. The limbs were almost certainly absent, although some historical sources reported them to be present. Despite the likely absence of limbs, adelospondyls retained a large part of the bony shoulder girdle. Adelospondyls have been assigned to a variety of groups in the past. They have traditionally been seen as members of the subclass Lepospondyli, related to other unusual early tetrapods such as "microsaurs", "nectrideans", and aïstopods. Analyses such as Ruta & Coates (2007) have offered an alternate classification scheme, arguing that adelospondyls were actually far removed from other lepospondyls, instead being stem-tetrapod stegocephalians closely related to the family Colosteidae.
Aïstopoda is an order of highly specialised snake-like amphibians known from the Carboniferous and Early Permian of Europe and North America, ranging from tiny forms only 5 centimetres (2 in), to nearly 1 metre (3.3 ft) in length. The first appear in the fossil record in the Mississippian period and continue through to the Early Permian.
Lethiscus is the earliest known representative of the Aistopoda, a group of very specialised snake-like amphibians known from the early Carboniferous (Mississippian).
Reptiliomorpha is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians (lissamphibians). It was defined by Michel Laurin (2001) and Vallin and Laurin (2004) as the largest clade that includes Homo sapiens, but not Ascaphus truei.
Nectridea is the name of an extinct order of lepospondyl tetrapods from the Carboniferous and Permian periods, including animals such as Diplocaulus. In appearance, they would have resembled modern newts or aquatic salamanders, although they are not close relatives of modern amphibians. They were characterized by long, flattened tails to aid in swimming, as well as numerous features of the vertebrae.
Microsauria is an extinct, possibly polyphyletic order of tetrapods from the late Carboniferous and early Permian periods. It is the most diverse and species-rich group of lepospondyls. Recently, Microsauria has been considered paraphyletic, as several other non-microsaur lepospondyl groups such as Lysorophia seem to be nested in it. Microsauria is now commonly used as a collective term for the grade of lepospondyls that were originally classified as members of Microsauria.
Gerobatrachus is an extinct genus of amphibamid temnospondyl that lived in the Early Permian, approximately 290 million years ago (Ma), in the area that is now Baylor County, Texas. When it was first described in 2008, Gerobatrachus was announced to be the closest relative of Batrachia, the group that includes modern frogs and salamanders. It possesses a mixture of characteristics from both groups, including a large frog-like head and a salamander-like tail. These features have led to it being dubbed a frogamander by the press. Some more recent studies place Gerobatrachus as the closest relative of Lissamphibia, the group that contains all modern amphibians including frogs, salamanders, and caecilians, or place modern amphibians far from Gerobatrachus within a group called Lepospondyli.
Acherontiscus is an extinct genus of stegocephalians that lived in the Early Carboniferous of Scotland. The type and only species is Acherontiscus caledoniae, named by paleontologist Robert Carroll in 1969. Members of this genus have an unusual combination of features which makes their placement within amphibian-grade tetrapods uncertain. They possess multi-bone vertebrae similar to those of embolomeres, but also a skull similar to lepospondyls. The only known specimen of Acherontiscus possessed an elongated body similar to that of a snake or eel. No limbs were preserved, and evidence for their presence in close relatives of Acherontiscus is dubious at best. Phylogenetic analyses created by Marcello Ruta and other paleontologists in the 2000s indicate that Acherontiscus is part of Adelospondyli, closely related to other snake-like animals such as Adelogyrinus and Dolichopareias. Adelospondyls are traditionally placed within the group Lepospondyli due to their fused vertebrae. Some analyses published since 2007 have argued that adelospondyls such as Acherontiscus may not actually be lepospondyls, instead being close relatives or members of the family Colosteidae. This would indicate that they evolved prior to the split between the tetrapod lineage that leads to reptiles (Reptiliomorpha) and the one that leads to modern amphibians (Batrachomorpha). Members of this genus were probably aquatic animals that were able to swim using snake-like movements.
Rhynchonkos is an extinct genus of microsaur. It is the only known member of the family Rhynchonkidae. Originally known as Goniorhynchus, it was renamed in 1981 because the name had already been given to another genus; the family, likewise, was originally named Goniorhynchidae but renamed in 1988. The type and only known species is R. stovalli, found from the Early Permian Fairmont Shale in Cleveland County, Oklahoma. Rhynchonkos shares many similarities with Eocaecilia, an early caecilian from the Early Jurassic of Arizona. Similarities between Rhynchonkos and Eocaecilia have been taken as evidence that caecilians are descendants of microsaurs. However, such a relationship is no longer widely accepted.
Trihecaton is an extinct genus of microsaur from the Late Pennsylvanian of Colorado. Known from a single species, Trihecaton howardinus, this genus is distinctive compared to other microsaurs due to possessing a number of plesiomorphic ("primitive") features relative to the rest of the group. These include large intercentra, folded enamel, and a large coronoid process of the jaw. Its classification is controversial due to combining a long body with strong limbs, features which typically are not present at the same time in other microsaurs. Due to its distinctiveness, Trihecaton has been given its own monospecific family, Trihecatontidae.
Scincosaurus is an extinct genus of nectridean lepospondyl within the family Scincosauridae.
Stegotretus is an extinct genus of lepospondyl microsaur referred to the Pantylidae. It is known from the Carboniferous–Permian boundary Cutler Formation exposures of New Mexico.
Altenglanerpeton is an extinct genus of microsaur amphibian from the Late Carboniferous or Early Permian of Germany. Altenglanerpeton was named in 2012 after the Altenglan Formation in which it was found. The type and only species is A.. schroederi.
Recumbirostra is a clade of lepospondyl amphibians from the Carboniferous and Permian that includes the families Pantylidae, Gymnarthridae, Ostodolepidae, Goniorhynchidae, Brachystelechidae, and Microbrachidae. Recumbirostra was erected as a clade in 2007. It includes many lepospondyls traditionally grouped in "Microsauria", which has since been shown to be a paraphyletic grouping. Not all phylogenetic analyses recognize Recumbirostra as a valid grouping. An alternative clade called Tuditanomorpha is occasionally supported and includes many of the same taxa. Below is a cladogram showing the phylogenetic relationships of recumbirostrans from Glienke (2012):
The Urocordylidae are an extinct family of nectridean lepospondyl amphibians. Urocordylids lived during the Late Carboniferous and Early Permian in what is now Europe and North America and are characterized by their very long, paddle-like tails. In life, they were probably newt-like and aquatic.
Andersonerpeton is an extinct genus of aïstopod from the Bashkirian of Nova Scotia, Canada. It is known from a single jaw, which shares an unusual combination of features from both other aistopods and from stem-tetrapod tetrapodomorph fish. As a result, Andersonerpeton is significant for supporting a new classification scheme which states that aistopods evolved much earlier than previously expected. The genus contains a single species, A. longidentatum, which was previously believed to have been a species of the microsaur Hylerpeton.