Casineria

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Casineria
Temporal range: Early Carboniferous, 340–334  Ma
Casineria kiddi.jpg
The counterslab of the holotype fossil
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Sarcopterygii
Clade: Tetrapodomorpha
Clade: Eotetrapodiformes
Clade: Elpistostegalia
Clade: Stegocephali
Genus: Casineria
Paton, Smithson & Clack, 1999
Type species
Casineria kiddi
Paton, Smithson & Clack, 1999

Casineria is an extinct genus of tetrapod which lived about 340-334 million years ago in the Mississippian epoch of the Carboniferous period. Its generic name, Casineria, is a latinization of Cheese Bay. The site near Edinburgh, Scotland where the holotype fossil was found. [1] When originally described in 1999, it was identified as a transitional fossil noted for its mix of basal (amphibian-like) and advanced (reptile-like) characteristics, putting it at or very near the origin of the amniotes, the group containing all mammals, birds, modern reptiles, and other descendants of their reptile-like common ancestor. However, the sole known fossil is lacking key elements such as a skull, making exact analysis difficult. [2] As a result, the classification of Casineria has been more controversial in analyses conducted since 1999. Other proposed affinities include a placement among the lepospondyls, [3] seymouriamorphs, [4] "gephyrostegids", [5] or as a synonym of Caerorhachis , another controversial tetrapod which may have been an early temnospondyl. [6]

Contents

Description

Restoration Casineria kiddi reconstruction.jpg
Restoration

Casineria was a small animal with a postcrania length estimated to have been 15 centimeters. Its small size would have made it ideal for hunting the invertebrates of the Carboniferous. [1]

Casineria had five fingers, with the bones at the finger tip being tapering and curved. This spurred the initial describers to argue that it was the oldest known animal with claws on each hand, and marks the earliest clawed foot. [1] Claws are extremely rare among amphibians, but ubiquitous among amniotes, so their supposed presence in Casineria has been considered to be evidence towards its amniote identity. Claws are also a feature intimately bound to the formation of keratinous scales in reptiles, so in life, Casineria would have borne scaly, reptilian-type skin, and would have resembled a small lizard (despite being only distantly related to true lizards). [7]

Under the hypothesis that it was among the first amniotes in the biological sense, it would have laid an amniotic egg not dependent on being laid in water to survive, possibly hiding them in damp vegetation or hollowed out tree stumps. This has been inferred from the fact that Casineria was found in rocks showing a rather dry environment. [1] In the early Carboniferous period before the appearance of Casineria, vertebrates were primarily aquatic, only spending part of their time on land. Casineria was believed to be among the first vertebrates to live and reproduce on land.

However, later studies have not consistently placed Casineria as an early amniote. [3] [4] [6] Even the presence of claws has been considered doubtful, as Marjanovic & Laurin (2019) noted that the finger tips were squared-off, rather than pointed. [5]

Discovery

Location of Cheese Bay in Great Britain Cheese Bay, Scotland.svg
Location of Cheese Bay in Great Britain

In 1992, an amateur fossil collector spotted the remnants of this four-legged creature on the shore of Cheese Bay, Scotland. [8] For the next five years, the fossil languished at the National Museum of Scotland in Edinburgh while researchers focused on other projects. Consisting of a slab and counterslab (compression fossil) of a single partial skeleton, the fossil was cataloged with the specimen designation NMS G.1993.54.1. Around 1997, work began to expose the remainder of the fossil from the surrounding matrix. The work revealed that the animal probably lived in an environment much drier than previously understood. The findings were first reported in the April 8, 1999 edition of Nature. [1]

Classification

While retaining a general build like those found in the amphibian reptiliomorph groups like Seymouriamorpha and Diadectomorpha, Casineria also shows features that tie it in with early reptiles, notably a generally gracile build with light leg bones, unfused ankles and toes terminating in claws. This would enable the animal to use their feet actively in traction, rather than as holdfasts, an indication of a primarily terrestrial lifestyle. [1] These traits have been argued to show that it was more closely related to amniotes than to other known reptiliomorph amphibians.

With its advanced features, Casineria may have been one of the first true amniotes, and thus the first reptile under traditional classification. In phylogenetic parlance it would have been a stem amniote, close to, but outside the crown group Amniota (the group consisting of the last common ancestor of synapsids and sauropsids and all its descendants). Casineria pushes back the origin of amniote lineages much earlier than was previously assumed. [1] However, like with much of the basal tetrapod stock, the phylogenetic position is uncertain, and it has also been suggested Casineria is a lepospondyl, [3] seymouriamorph, [4] synonymous with Caerorhachis (which is possibly a basal temnospondyl amphibian), [6] or part of a grade of small tetrapods traditionally considered the family Gephyrostegidae. [5]

See also

Related Research Articles

<span class="mw-page-title-main">Tetrapod</span> Superclass of the first four-limbed vertebrates and their descendants

A tetrapod is any four-limbed vertebrate animal of the superclass Tetrapoda. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids and synapsids. Some tetrapods such as snakes, legless lizards, and caecilians had evolved to become limbless via mutations of the Hox gene, although some do still have a pair of vestigial spurs that are remnants of the hindlimbs.

<span class="mw-page-title-main">Amniote</span> Clade of tetrapods including reptiles, birds and mammals

Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibian ancestors during the Carboniferous period and further diverged into two groups, namely the sauropsids and synapsids. They are distinguished from the other living tetrapod clade — the non-amniote lissamphibians — by the development of three extraembryonic membranes, thicker and keratinized skin, and costal respiration.

<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">Batrachomorpha</span> Clade of amphibians

The Batrachomorpha are a clade containing recent and extinct amphibians that are more closely related to modern amphibians than they are to mammals and reptiles. According to many analyses they include the extinct Temnospondyli; some show that they include the Lepospondyli instead. The name traditionally indicated a more limited group.

<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.

<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">Anthracosauria</span> Paraphyletic group of tetrapodomorphs

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<i>Diadectes</i> Extinct genus of reptiles

Diadectes is an extinct genus of large reptiliomorphs or synapsids that lived during the early Permian period. Diadectes was one of the first herbivorous tetrapods, and also one of the first fully terrestrial vertebrates to attain large size.

<span class="mw-page-title-main">Diadectomorpha</span> Extinct clade of tetrapods

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<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>Eucritta</i> Extinct genus of tetrapods

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

Tulerpeton is an extinct genus of Devonian four-limbed vertebrate, known from a fossil that was found in the Tula Region of Russia at a site named Andreyevka. This genus and the closely related Acanthostega and Ichthyostega represent the earliest tetrapods.

<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.

<i>Gephyrostegus</i> Extinct genus of reptile-like amphibians

Gephyrostegus is a genus of extinct gephyrostegid reptiliomorph amphibian. It was a small animal at 22 cm snout-vent length, of generally lizard-like build and presumably habit. It had large eyes and a large number of small, pointed teeth, indicating it was an active insectivorous hunter. The remains have been found in Nýřany, Czech Republic, dating from around 310 million years ago.

<i>Solenodonsaurus</i> Extinct genus of reptiles

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

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<span class="mw-page-title-main">Evolution of reptiles</span> Origin and diversification of reptiles through geologic time

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<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.

References

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  2. Monastersky, R. (1999): Out of the Swamps, How early vertebrates established a foothold—with all 10 toes—on land, Science News vol. 155, No. 21, p. 328
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  5. 1 2 3 Marjanović, David; Laurin, Michel (2019-01-04). "Phylogeny of Paleozoic limbed vertebrates reassessed through revision and expansion of the largest published relevant data matrix". PeerJ. 6: e5565. doi: 10.7717/peerj.5565 . ISSN   2167-8359. PMC   6322490 . PMID   30631641.
  6. 1 2 3 Chapter 6: "Walking with early tetrapods: evolution of the postcranial skeleton and the phylogenetic affinities of the Temnospondyli (Vertebrata: Tetrapoda)." In: Kat Pawley (2006). "The postcranial skeleton of temnospondyls (Tetrapoda: temnospondyli)." PhD Thesis. La Trobe University, Melbourne.
  7. Alibardi, L. (2008): Microscopic analysis of lizard claw morphogenesis and hypothesis on its evolution. Acta Zoologica: Morphology and Evolution , vol 89 (2): pp 169–178. abstract
  8. "Archived copy" (PDF). Archived from the original (PDF) on 2008-07-28. Retrieved 2008-03-22.{{cite web}}: CS1 maint: archived copy as title (link)

Further reading