Urocordylidae

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Urocordylidae
Temporal range: Late Carboniferous
Sauropleura.jpg
Life restoration of Sauropleura pectinata
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Sarcopterygii
Clade: Tetrapodomorpha
Order: Nectridea
Family: Urocordylidae
Lydekker, 1889
Subfamilies

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. [1]

Contents

Fossils have been found from Ireland, France, and the eastern United States. The family was named by English naturalist Richard Lydekker in 1889 and includes the well-known genera Urocordylus and Sauropleura , as well as several others based on less-complete material. [2] The family Urocordylidae is divided into two subfamilies, the Urocordylinae and the Sauropleurinae. The two groups are distinguished by the shapes of their skulls; urocordylines have short, blunt skulls, and sauropleurines have longer, pointed skulls. [3]

Description

Life restoration of Urocordylus wandesfordii, an early urocordylid from Ireland Urocordylus.jpg
Life restoration of Urocordylus wandesfordii , an early urocordylid from Ireland

Urocordylids are distinguished by their elongated tails. Each tail vertebra has an upper crest of bone called a neural arch and a lower crest called a haemal arch, and each of these arches is square-shaped and laterally compressed (flattened side-to-side). These arches give the tails of urocordylids a paddle-like shape. [4] The urocordylids Sauropleura and Crossotelos have holes in their tail vertebrae that are thought to have served as passageways for spinal nerves, an unusual feature given that spinal nerves pass between successive vertebrae in most vertebrates. Salamanders also have spinal nerves passing through their vertebrae, although the holes are in different positions and do not indicate a close evolutionary relationship with urocordylids. [3]

Another distinguishing feature of urocordylids is the loose connection between the bones of the skull. The bones at the back of the lower jaw do not connect with the rest of the skull. Instead, a gap extends across the skull and closes in front of the eye sockets. This allows the upper jaw to flex upward, increasing the gape of the mouth. The bones at the back of the jaw may also be able to flex outward. [4] The flexibility of the jaws of urocordylids may have been an adaptation for feeding, allowing sharp teeth at the tip of the jaw to face outward so prey could be more easily captured. [1]

Relationships

Urocordylids belong to a large group of Paleozoic amphibians called lepospondyls, which are distinguished from other early land vertebrates by their simple spool-shaped vertebrae. The Urocordylidae belong to a group within the Lepospondyli called the Nectridea, which include forms with long tails and short bodies. Urocordylids have traditionally been considered the most primitive nectrideans because they lack some of the more unusual features of other members of the group, such as the tabular horns of diplocaulids. [4] However, some recent phylogenetic analyses of lepospondyls have not supported a primitive position for urocordylids among nectrideans. The analysis of Ruta et al. (2003), for example, places the Urocordylidae as the closest relatives of a group of legless lepospondyls called the Aistopoda, while other nectrideans fall into a more basal position on the evolutionary tree. This splits up the Nectridea, making them a polyphyletic taxon and not a true clade or evolutionary grouping. Below is part of the cladogram from Ruta et al.: [2]

Lepospondyli

Batropetes fritschi

Tuditanomorpha

Microbrachis pelikani

Hyloplesion longicostatum

Odonterpeton triangulare

Lysorophia

Adelospondyli

Holospondyli

Scincosauridae

Diplocaulidae

Urocordylidae

Aistopoda

Related Research Articles

<span class="mw-page-title-main">Labyrinthodontia</span> Paraphyletic group of tetrapodomorphs

"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.

<span class="mw-page-title-main">Lepospondyli</span> Polyphyletic group of tetrapodomorphs

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.

Lysorophia is an order of fossorial Carboniferous and Permian tetrapods within the Recumbirostra. 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.

<span class="mw-page-title-main">Adelospondyli</span> Extinct order of amphibians

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.

<span class="mw-page-title-main">Aistopoda</span> Extinct order of amphibians

Aistopoda is an order of highly specialised snake-like stegocephalians 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. They first appear in the fossil record in the Mississippian period and continue through to the Early Permian.

<span class="mw-page-title-main">Reptiliomorpha</span> Clade of reptile-like animals

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. Laurin and Reisz (2020) defined Pan-Amniota as the largest total clade containing Homo sapiens, but not Pipa pipa, Caecilia tentaculata, and Siren lacertina.

<span class="mw-page-title-main">Temnospondyli</span> Ancestors of modern amphibians adapted to 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.

<i>Westlothiana</i> Extinct genus of tetrapods

Westlothiana is a genus of reptile-like tetrapod that lived about 338 million years ago during the latest part of the Viséan age of the Carboniferous. Members of the genus bore a superficial resemblance to modern-day lizards. The genus is known from a single species, Westlothiana lizziae. The type specimen was discovered in the East Kirkton Limestone at the East Kirkton Quarry, West Lothian, Scotland in 1984. This specimen was nicknamed "Lizzie the lizard" by fossil hunter Stan Wood, and this name was quickly adopted by other paleontologists and the press. When the specimen was formally named in 1990, it was given the specific name "lizziae" in homage to this nickname. However, despite its similar body shape, Westlothiana is not considered a true lizard. Westlothiana's anatomy contained a mixture of both "labyrinthodont" and reptilian features, and was originally regarded as the oldest known reptile or amniote. However, updated studies have shown that this identification is not entirely accurate. Instead of being one of the first amniotes, Westlothiana was rather a close relative of Amniota. As a result, most paleontologists since the original description place the genus within the group Reptiliomorpha, among other amniote relatives such as diadectomorphs and seymouriamorphs. Later analyses usually place the genus as the earliest diverging member of Lepospondyli, a collection of unusual tetrapods which may be close to amniotes or lissamphibians, or potentially both at the same time.

<i>Diplocaulus</i> Extinct genus of amphibians

Diplocaulus is an extinct genus of lepospondyl amphibians which lived from the Late Carboniferous to the Late Permian of North America and Africa. Diplocaulus is by far the largest and best-known of the lepospondyls, characterized by a distinctive boomerang-shaped skull. Remains attributed to Diplocaulus have been found from the Late Permian of Morocco and represent the youngest-known occurrence of a lepospondyl.

<span class="mw-page-title-main">Nectridea</span> Extinct order of amphibians

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.

<span class="mw-page-title-main">Microsauria</span> Extinct order of amphibians

Microsauria is an extinct, possibly polyphyletic order of amphibians 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.

Arizonerpeton is an extinct genus of nectridean tetrapodomorphs. It contains a single species, Arizonerpeton wellsi. It lived in what is now the Swisshelm Mountains of modern-day Arizona, United States. This locality belongs to the Black Prince Limestone Formation, which is dated to the middle Pennsylvanian sub-period of the Carboniferous period.

<i>Acherontiscus</i> Extinct genus of amphibians

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.

<i>Euryodus</i> Extinct genus of amphibians

Euryodus is an extinct genus of microsaur within the family Gymnarthridae. Euryodus is a Lepospondyl from the clade Microsauria that lived during the Lower Permian. The name comes from Greek, meaning ‘broad-tooth’. It has been found in the southern half of North America, from its original discovery in Texas up to Utah.

Doleserpeton is an extinct, monospecific genus of dissorophoidean temnospondyl within the family Amphibamidae that lived during the Upper Permian, 285 million years ago. Doleserpeton is represented by a single species, Doleserpeton annectens, which was first described by John R. Bolt in 1969. Fossil evidence of Doleserpeton was recovered from the Dolese Brothers Limestone Quarry in Fort Sill, Oklahoma. The genus name Doleserpeton is derived from the initial discovery site in Dolese quarry of Oklahoma and the Greek root "herp-", meaning "low or close to the ground". This transitional fossil displays primitive traits of amphibians that allowed for successful adaptation from aquatic to terrestrial environments. In many phylogenies, lissamphibians appear as the sister group of Doleserpeton.

<i>Odonterpeton</i> Extinct genus of amphibians

Odonterpeton is an extinct genus of "microsaur" from the Late Carboniferous of Ohio, containing the lone species Odonterpeton triangulare. It is known from a single partial skeleton preserving the skull, forelimbs, and the front part of the torso. The specimen was found in the abandoned Diamond Coal Mine of Linton, Ohio, a fossiliferous coal deposit dated to the late Moscovian stage, about 310 million years ago.

<i>Rhynchonkos</i> Extinct genus of tetrapods

Rhynchonkos is an extinct genus of rhynchonkid microsaur. 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.

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. 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 that are very different from one another, making it a significant taxon in terms of understanding the interrelationships of lepospondyls.

<i>Scincosaurus</i> Extinct genus of tetrapodomorphs

Scincosaurus is an extinct genus of nectridean tetrapodomorphs within the family Scincosauridae.

<span class="mw-page-title-main">Sauropleurinae</span> Extinct subfamily of tetrapodomorphs

Sauropleurinae is an extinct subfamily of nectridean tetrapodomorphs. Along with the subfamily Urocordylinae, Sauropleurinae is part of the family Urocordylidae. Like other urocordylids, sauropleurines have long, flattened tails and superficially resemble aquatic newts. They differ from urocordylines in having narrow, pointed skulls. Sauropleurinae includes the genera Sauropleura, Crossotelos, Lepterpeton, and Montcellia, which lived during the Late Carboniferous and Early Permian in what is now North America and Europe.

References

  1. 1 2 Carroll, R.L.; Bossy, K.A.; Milner, A.C.; Andrews, S.M.; Wellstead, C.F. (1998). "Lepospondyli". In P. Wellnhofer (ed.). Handbuch der Paläoherpetologie. Vol. 1. Munich: Verlag Dr. Friedrich Pfeil. p. 216. ISBN   978-3-931516-26-0.
  2. 1 2 Marcello Ruta, Michael I. Coates and Donald L. J. Quicke (2003). "Early tetrapod relationships revisited" (PDF). Biological Reviews. 78 (2): 251–345. doi:10.1017/S1464793102006103. PMID   12803423.
  3. 1 2 Carlson, K. J. (1999). "Crossotelos, an Early Permian nectridian amphibian". Journal of Vertebrate Paleontology. 19 (4): 623–631. doi:10.1080/02724634.1999.10011176.
  4. 1 2 3 Carroll, R.L. (2009). "The Radiation of Carboniferous Amphibians". The Rise of Amphibians: 365 Million Years of Evolution . Baltimore: Johns Hopkins University Press. pp.  61–143.