Claudiosaurus Temporal range: Lopingian ~ | |
---|---|
Specimen of Claudiosaurus germaini, on display at the Redpath Museum, Montreal | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Neodiapsida |
Family: | † Claudiosauridae Carroll 1981 |
Genus: | † Claudiosaurus Carroll 1981 |
Type species | |
†Claudiosaurus germaini |
Claudiosaurus (claudus is Latin for 'lameness' and saurus means 'lizard') is an extinct genus of diapsid reptiles from the Late Permian Sakamena Formation of the Morondava Basin, Madagascar. It has been suggested to be semi-aquatic.
Claudiosaurus is known from the Sakamena Formation of Madagascar. Claudiosaurus is found from the Late Permian. Although a paper mentions that they have been also found in Early Triassic deposits of Madagascar, [1] citation does not mention that Claudiosaurus is from Triassic. [2]
Individuals of Claudiosaurus reached a body length of approximately 60 centimetres (2.0 ft). [3] The body form of Claudiosaurus is generally similar to those of other basal diapsids, although the neck of Claudiosaurus is somewhat elongated, with 8 cervical vertebrae, and had a proportionally small head. The body has 16 trunk vertebrae with gastralia present on the underside, and the tail has at least 45 caudal vertebrae. The jaws had numerous small teeth, with the roof of the mouth (palate) being covered in numerous denticles. [4] The sternum is unossified. [5] The pectoral girdle is similar to those of other primitive diapsids. [6] The phalanges of the hands show a reduction of length away from the base, with the exception of the distalmost phalange of the third digit, which is longer than the preceding phalange. The terminal phalanges are flattened. [4] The bones of Claudiosaurus show pachyostosis, suggested to possibly be an adaptation for aquatic life. [7]
Claudiosaurus is generally assumed to have been an amphibious animal, using its limbs for propulsion, though it was still likely capable of walking on land, and the skeleton shows only limited adaptations to aquatic life. It has been suggested to have fed on small invertebrates, such as crustaceans. [4]
Upon its original description, Robert L. Carroll suggested that Claudiosaurus belonged to Sauropterygia (which includes plesiosaurs). [4] Other later studies have generally recovered it as a basal neodiapsid, sometimes as a member of the Younginiformes. [8] [9] Claudiosaurus was recovered as a relative of turtles by Li et al. (2018), forming a clade with the basal neodiapsid Acerosodontosaurus . [10] Although another study in 2020 specifically disputed these conclusions. [11]
The Lower Sakamena Formation was deposited in a wetland environment situated within a North-South orientated rift valley, perhaps similar to Lake Tanganyika. The climate at the time of deposition was temperate, warm, and humid, with seasonal rainfall and possible monsoons. [12] Flora from the formation includes the equisetalean Schizoneura , the glossopterid gymnosperm Glossopteris , and seed fern Lepidopteris. Other vertebrates known from the Lower Sakamena Formation include the palaeoniscoid fish Atherstonia , the procolophonid parareptile Barasaurus , the gliding weigeltisaurid reptile Coelurosauravus , the neodiapsids Hovasaurus, Thadeosaurus , and Acerosodontosaurus , fragments of rhinesuchid temnospondyls, an indeterminate theriodont therapsid and the dicynodont Oudenodon . [13]
Diapsids are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ago during the late Carboniferous period. All diapsids other than the most primitive ones in the clade Araeoscelidia are sometimes placed into the clade Neodiapsida. The diapsids are extremely diverse, and include birds and all modern reptile groups, including turtles, which were historically thought to lie outside the group. Although some diapsids have lost either one hole (lizards), or both holes, or have a heavily restructured skull, they are still classified as diapsids based on their ancestry. At least 17,084 species of diapsid animals are extant: 9,159 birds, and 7,925 snakes, lizards, tuatara, turtles, and crocodiles.
Dinocephalosaurus is a genus of long necked, aquatic protorosaur that inhabited the Triassic seas of China. The genus contains the type and only known species, D. orientalis, which was named by Chun Li in 2003. Unlike other long-necked protorosaurs, Dinocephalosaurus convergently evolved a long neck not through elongation of individual neck vertebrae, but through the addition of neck vertebrae that each had a moderate length. As indicated by phylogenetic analyses, it belonged in a separate lineage that also included at least its closest relative Pectodens, which was named the Dinocephalosauridae in 2021. Like tanystropheids, however, Dinocephalosaurus probably used its long neck to hunt, utilizing the fang-like teeth of its jaws to ensnare prey; proposals that it employed suction feeding have not been universally accepted. It was probably a marine animal by necessity, as suggested by the poorly-ossified and paddle-like limbs which would have prevented it from going ashore.
Mesosaurs were a group of small aquatic reptiles that lived during the early Permian period (Cisuralian), roughly 299 to 270 million years ago. Mesosaurs were the first known aquatic reptiles, having apparently returned to an aquatic lifestyle from more terrestrial ancestors. It is uncertain which and how many terrestrial traits these ancestors displayed; recent research cannot establish with confidence if the first amniotes were fully terrestrial, or only amphibious. Most authors consider mesosaurs to have been aquatic, although adult animals may have been amphibious, rather than completely aquatic, as indicated by their moderate skeletal adaptations to a semiaquatic lifestyle. Similarly, their affinities are uncertain; they may have been among the most basal sauropsids or among the most basal parareptiles.
Archosauromorpha is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs rather than lepidosaurs. Archosauromorphs first appeared during the late Middle Permian or Late Permian, though they became much more common and diverse during the Triassic period.
Neodiapsida is a clade, or major branch, of the reptilian family tree, typically defined as including all diapsids apart from some early primitive types known as the araeoscelidians. Modern reptiles and birds belong to the neodiapsid subclade Sauria.
Pareiasaurs are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with osteoderms which covered large areas of the body. They first appeared in southern Pangea during the Middle Permian, before becoming globally distributed during the Late Permian. Pareiasaurs were the largest reptiles of the Permian, reaching sizes equivalent to those of contemporary therapsids. Pareiasaurs became extinct in the Permian–Triassic extinction event.
Coelurosauravus is an extinct genus of gliding reptile, known from the Late Permian of Madagascar. Like other members of the family Weigeltisauridae, members of this genus possessed long, rod-like ossifications projecting outwards from the body. These bony rods were not extensions of the ribs but were instead a feature unique to weigeltisaurids. It is believed that during life, these structures formed folding wings used for gliding flight, similar to living gliding Draco lizards.
Avicephala is a potentially polyphyletic grouping of extinct diapsid reptiles that lived during the Late Permian and Triassic periods characterised by superficially bird-like skulls and arboreal lifestyles. As a clade, Avicephala is defined as including the gliding weigeltisaurids and the arboreal drepanosaurs to the exclusion of other major diapsid groups. This relationship is not recovered in the majority of phylogenetic analyses of early diapsids and so Avicephala is typically regarded as an unnatural grouping. However, the clade was recovered again in 2021 in a redescription of Weigeltisaurus, raising the possibility that the clade may be valid after all.
Younginiformes is a group of diapsid reptiles known from the Permian-Triassic of Africa and Madagascar. It has been used as a replacement for "Eosuchia". Younginiformes were historically suggested to be lepidosauromorphs, but were later suggested to be basal non-saurian neodiapsids. The group is sometimes divided into two families, Tangasauridae and Younginidae. The monophyly of the group is disputed. A 2009 study found them to be an unresolved polytomy at the base of Neodiapsida, while a 2011 study recovered the group as paraphyletic. A 2022 study recovered the Younginiformes as a monophyletic group of basal neodiapsid reptiles, also including Claudiosaurus and Saurosternon as part of the group. Some younginiforms like Hovasaurus and Acerosodontosaurus are thought to have had an amphibious lifestyle, while others like Kenyasaurus, Thadeosaurus and Youngina were probably terrestrial.
Thadeosaurus is an extinct genus of diapsid reptile belonging to the family Younginidae. Fossils have been found in the Lower Sakamena Formation of the Morondava Basin, Madagascar in 1981, and date to the late Permian to the early Triassic period.
Hovasaurus is an extinct genus of basal diapsid reptile. It lived in what is now Madagascar during the Late Permian and Early Triassic, being a survivor of the Permian–Triassic extinction event and the paleontologically youngest member of the Tangasauridae. Fossils have been found in the Permian Lower and Triassic Middle Sakamena Formations of the Sakamena Group, where it is amongst the commonest fossils. Its morphology suggests an aquatic ecology.
Drepanosaurs are a group of extinct reptiles that lived between the Carnian and Rhaetian stages of the late Triassic Period, approximately between 230 and 210 million years ago. The various species of drepanosaurid were characterized by specialized grasping limbs and often prehensile tails, adaptions for arboreal (tree-dwelling) and fossorial (digging) lifestyles, with some having also been suggested to be aquatic. Fossils of drepanosaurs have been found in Arizona, New Mexico, New Jersey, Utah, England, and northern Italy. The name is taken from the family's namesake genus Drepanosaurus, which means "sickle lizard," a reference to their strongly curved claws.
Sakamena is a village near Betroka in the region of Anosy in Madagascar.
Acerosodontosaurus is an extinct genus of neodiapsid reptiles that lived during the Late Permian of Madagascar. The only species of Acerosodontosaurus, A. piveteaui, is known from a natural mold of a single partial skeleton including a crushed skull and part of the body and limbs. The fossil was discovered in deposits of the Lower Sakamena Formation. Based on skeletal characteristics, it has been suggested that Acerosodontosaurus individuals were at least partially aquatic.
Tangasauridae is an extinct family of diapsids known from fossil specimens from Madagascar, Kenya and Tanzania that are Late Permian to Early Triassic in age. Fossils have been found of numerous specimens of common members of this family such as Hovasaurus in different stages of ontogenic development. Recent material from the Middle Sakamena Formation of the Morondava Basin of Madagascar that dates back to the early Triassic period suggests that the Tangasauridae were relatively unaffected by the Permian-Triassic extinction event.
Hupehsuchia is an order of diapsid reptiles closely related to ichthyosaurs. The group was short-lasting, with a temporal range restricted to the late Olenekian age, spanning only a few million years of the Early Triassic. The order gets its name from Hubei Province, China, from which many specimens have been found. They are probable members of the clade Ichthyosauromorpha.
Weigeltisauridae is a family of gliding neodiapsid reptiles that lived during the Late Permian, between 259.51 and 251.9 million years ago. Fossils of weigeltisaurids have been found in Madagascar, Germany, Great Britain, and Russia. They are characterized by long, hollow rod-shaped bones extending from the torso that probably supported wing-like membranes. Similar membranes are also found in several other extinct reptiles such as kuehneosaurids and Mecistotrachelos, as well as living gliding lizards, although each group evolved these structures independently.
Velosauria is a group of pareiasaur reptiles that existed in the late Permian period. They ranged in size from the 50-centimeter-long Pumiliopareia to the 3-meter-long Scutosaurus. Velosaurs were some of the largest reptiles of their time.
Reptiles arose about 320 million years ago during the Carboniferous period. Reptiles, in the traditional sense of the term, are defined as animals that have scales or scutes, lay land-based hard-shelled eggs, and possess ectothermic metabolisms. So defined, the group is paraphyletic, excluding endothermic animals like birds that are descended from early traditionally-defined reptiles. A definition in accordance with phylogenetic nomenclature, which rejects paraphyletic groups, includes birds while excluding mammals and their synapsid ancestors. So defined, Reptilia is identical to Sauropsida.
Pantestudines or Pan-Testudines is the group of all reptiles more closely related to turtles than to any other living animal. It includes both modern turtles and all of their extinct relatives. Pantestudines with a complete shell are placed in the clade Testudinata.
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