Nectridea

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Nectridea
Temporal range: Late Carboniferous–Lopingian
Diplocaulus - skeleton and model.jpg
Skeleton and model of Diplocaulus .
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Sarcopterygii
Clade: Tetrapodomorpha
Clade: Stegocephali
Order: Nectridea
Miall, 1875
Subgroups

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. [1] They were characterized by long, flattened tails to aid in swimming, as well as numerous features of the vertebrae.

Contents

Description

A life restoration of Urocordylus, a urocordylid Urocordylus.jpg
A life restoration of Urocordylus , a urocordylid

Nectrideans are a diverse group of tetrapods, including the aquatic Urocordylidae, the presumably terrestrial Scincosauridae, and the bizarre horned members of Diplocaulidae (also known as Keraterpetonidae), which includes the "boomerang-headed" Diplocaulus, one of the most famous genera of prehistoric amphibians (in the traditional sense of the word). By the time the earliest known nectrideans appeared in the Late Carboniferous fossil record, they had already diversified into these families, indicating that basal nectrideans are unknown. These different families are united primarily by features of the spinal column rather than the skull.

Vertebrae

In many groups of early tetrapods, each vertebra is formed by three parts: an intercentrum at the front, a pleurocentrum at the back, and a plate-like neural spine jutting out from the top of the vertebrae which may be positioned between the two or fused to the pleurocentrum. In nectrideans, only the pleurocentra are present, and the neural spines are completely fused to them. Their vertebrae also possess an extra set of joints connecting the vertebrae (apophyses), just above the typical set (zygapophyses). Furthermore, their caudal (tail) vertebrae also possess haemal spines which jut out from the bottom of each vertebra. While most early tetrapods have slanted haemal spines which are positioned between the vertebrae, the haemal spines of nectrideans are completely vertical in orientation and fused to the pleurocentra, directly opposite the neural spines. Both the neural spines and haemal spines of the tail are characteristically fan-shaped. As a whole, nectrideans have rather short bodies and long and paddle-like tails, adapted for swimming. [2] [1]

Other features

Almost all early tetrapods have three coronoid bones lining the inside edge of each side of the jaw. Nectrideans, however, only have one. Their clavicular blades (shoulder girdles) are wide and possess a short rear extension, while the scapulocoracoids (shoulder blades) are weakly ossified. [2]

They also had well-developed hind limbs, with a full set of five toes each. Their forelimbs were slightly reduced, but not to the same extent as in microsaurs and other lepospondyls. Although Urocordylus retained five fingers, most nectridieans had only four, similar to modern amphibians. [2] [3]

Life restoration of Scincosaurus, a scincosaurid Scincosaurus.jpg
Life restoration of Scincosaurus , a scincosaurid

Paleobiology

Both diplocaulids and urocordylids are considered to be completely aquatic, although only diplocaulids possess lateral line canals on their skulls. However, this does not mean that urocordylids lacked lateral lines, as several modern amphibians possess that organ without any skeletal indication. Scincosaurids have much more well-ossified limbs and thinner tails than other nectrideans, and are usually considered to be terrestrial. However, some have been reported to possess lines of pits on the skull, which may be an indication that they were amphibious rather than purely terrestrial. [4]

A trace fossil known as Hermundurichnus fornicatus of a tetrapod resting on a lake bed may have been attributed to Diplocaulus or an animal like it. This trace indicates that the underside of nectrideans was covered in small, diamond-shaped scales and that the "horns" of the skull were connected with the body by flaps of skin. [5]

Classification

There has been some controversy over the precise classification of Nectridea over the past century, and even whether it constitutes a valid monophyletic clade. A few older studies consider nectrideans to be very basal tetrapods, related to Ichthyostega or colosteids. However, most studies generally place nectrideans inside the subclass Lepospondyli. Aistopoda, a group of legless lepospondyls, have often been allied with particular families of nectrideans, but not the order as a whole. For example, Anderson (2001) positioned diplocaulids as close relatives of Aistopods with urocordylids and scincosaurids as progressively more primitive members of the group. On the other hand, Ruta et al. (2003) claimed that urocordylids were closer to aistopods. As aistopods do not possess most nectridean features, they have traditionally been excluded from the group. However, this would make Nectridea an invalid paraphyletic group as the aistopods, may have been descended from nectrideans, are not considered nectrideans themselves. [6]

The following is a cladogram from the Ruta et al. (2003):

Lepospondyli

Batropetes fritschi

Tuditanomorpha

Microbrachis pelikani

Hyloplesion longicostatum

Odonterpeton triangulare

Lysorophia

Adelospondyli

Holospondyli

Scincosauridae

Diplocaulidae

Urocordylidae

Aistopoda

"Nectridea"

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.

Lethiscus is the earliest known representative of the Aistopoda, a group of very specialised snake-like tetrapodomorphs known from the early Carboniferous (Mississippian).

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

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<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.

<span class="mw-page-title-main">Embolomeri</span> Extinct order of tetrapods

Embolomeri is an order of tetrapods or stem-tetrapods, possibly members of Reptiliomorpha. Embolomeres first evolved in the Early Carboniferous (Mississippian) Period and were the largest and most successful predatory tetrapods of the Late Carboniferous (Pennsylvanian) Period. They were specialized semiaquatic predators with long bodies for eel-like undulatory swimming. Embolomeres are characterized by their vertebral centra, which are formed by two cylindrical segments, the pleurocentrum at the rear and intercentrum at the front. These segments are equal in size. Most other tetrapods have pleurocentra and intercentra which are drastically different in size and shape.

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>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.

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.

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.

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

Caerorhachis is an extinct genus of early tetrapod from the Early Carboniferous of Scotland, probably from the Serpukhovian stage. Its placement within Tetrapoda is uncertain, but it is generally regarded as a primitive member of the group. The type species C. bairdi was named in 1977.

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

Holospondyli is a proposed clade of lepospondyls from the Early Carboniferous to the Late Permian that includes the aistopods, the paraphyletic nectrideans, and possibly also Adelospondyli. However, aistopods have since been recovered as stem-tetrapods more primitive than temnospondyls or other groups of lepospondyls.

<span class="mw-page-title-main">Urocordylidae</span> Extinct family of tetrapodomorphs

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.

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

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.

References

  1. 1 2 Milner, Andrew R. (1993). "The Paleozoic Relatives of Lissamphibians". Herpetological Monographs. 7: 8–27. doi:10.2307/1466948. JSTOR   1466948.
  2. 1 2 3 Carroll, R. L. (2009). The rise of amphibians. The Johns Hopkins University Press.
  3. Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. p. 54. ISBN   1-84028-152-9.
  4. Vallin, Grégoire; Laurin, Michel. "Cranial morphology and affinities ofMicrobrachis, and a reappraisal of the phylogeny and lifestyle of the first amphibians". Journal of Vertebrate Paleontology. 24 (1): 56–72. doi:10.1671/5.1.
  5. Lockey, Martin; Meyer, Christian (2000). Dinosaur Tracks and other Fossil Footprints of Europe. Columbia University Press.
  6. 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.