Teraterpeton Temporal range: Late Triassic, | |
---|---|
Life restoration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | † Allokotosauria |
Order: | † Trilophosauria |
Family: | † Trilophosauridae |
Genus: | † Teraterpeton Sues, 2003 |
Type species | |
†Teraterpeton hrynewichorum Sues, 2003 |
Teraterpeton (meaning "wonderful creeping thing" in Greek) is an extinct genus of trilophosaurid [1] archosauromorphs. It is known from a partial skeleton from the Late Triassic Wolfville Formation of Nova Scotia, described in 2003. It has many unique features seen in no other related form, including an elongated, toothless snout and large openings for the nostrils. Because of this, Teraterpeton was originally placed in its own family, Teraterpetidae, related to Trilophosaurus . [2] Newer studies generally place it within Trilophosauridae. [1]
Teraterpeton had an unusual appearance compared to other early archosauromorphs. Members of the genus had a long skull with no teeth at the ends of the upper and lower jaws. Over each eye socket is a bony projection formed by the lacrimal and prefrontal bones. At the back of the jaws are a set of small, sharp, closely spaced teeth. They continue below the level of the eye, an unusual trait among early archosauromorphs. The upper tooth row does not run along the edge of the jaw, but is inset closer to mouth. An additional tooth row on the palate runs alongside the maxillary tooth row of the upper jaw. The teeth of the upper jaws fit closely, or occlude, with the teeth of the lower jaw. The upper jaw teeth have sharp cusps with indentations in front of them, while the lower jaw teeth have cusps with indentations behind them. The cusps of the upper teeth fit into the indentations of the lower teeth, while the cusps of the lower teeth fit into the indentations of the upper teeth. [2]
Teraterpeton has an uncommon feature compared to most archosauromorphs: a euryapsid-type skull. Euryapsids have a single hole at the back of the skull called the supratemporal fenestra, which is located toward the top of the head. Teraterpeton evolved from diapsid reptiles with two holes at the back of their skull, the supratemporal fenestra and an infratemporal fenestra below it. Although it lacks an infratemporal fenestra, Teraterpeton still belongs to Diapsida because it is a descendant of true diapsids. Most other euryapsids such as the marine plesiosaurs and ichthyosaurs are not closely related to Teraterpeton. However, the euryapsid archosauromorph Trilophosaurus has been identified as a close relative. [2] [1]
Another unusual feature of Teraterpeton is the large size of its narial fenestra, a hole in the skull that serves as the opening for the nostril. This hole is positioned directly in front of the eyes and extends to the level of the toothless portion of the snout. It is positioned where the antorbital fenestra would normally be. In fact, Hans-Dieter Sues, the original describer of Teraterpeton, first considered the hole to be an antorbital fenestra before revising his hypothesis. The narial fenestra of Teraterpeton is approximately 1.5 times longer than the eye socket. [2]
A string of eight cervical (neck) vertebrae are preserved in the holotype. They have large neural spines, and the second to fourth vertebrae have additional blade-like structures directly behind the neural spines. Overall, the neck is similar to that of the early rhynchosaur Mesosuchus. The cervical ribs are completely fused to the large rib facets of the vertebrae. Isolated dorsal (back), sacral (hip), and caudal (tail) vertebrae are also preserved. Caudal vertebrae from a referred specimen had very long transverse processes (column-like rib facets). [3] The scapula (shoulder blade) is long and narrow. The unguals (claws) of both the hands and feet are deep and blade-like, a morphology described as "trenchant". [2]
The preacetabular process (i.e. the front blade of the ilium bone of the hip) was large, thin and had a convex upper edge. This is in contrast to the small, bump-like preacetabular process of Trilophosaurus, and more similar to that of advanced rhynchosaurs like Hyperodapedon . The pubis and ischium bones of the hip were vertically oriented and separated by a tall, rectangular gap likely homologous with the thyroid fenestra of lepidosauromorphs and tanystropheids, rather than the lack of separation present in other early archosauromorphs. Also like tanystropheids, the fifth metatarsal of the foot was thick, short, and straight. [3]
The simple femur and compact hip supports the interpretation that Teraterpeton had a sprawling gait. However, the modified and expanded ilium suggests that Teraterpeton's hindlimb musculature (as well as that of rhynchosaurs) was well-adapted for protracting and retracting during more erect forms of locomotion. Some paleontologists have argued that expanded leg musculature in rhynchosaurs was an adaptation for scratch-digging rather than an upright posture. While this may apply to thick-legged rhynchosaurs, digging was less likely for Teraterpeton, which had rather slender hindlimbs by comparison. [3]
Teraterpeton is primarily known from a single partial holotype skeleton referred to as NSM 999GF041. This skeleton was found in a sea cliff in Burntcoat Head, Nova Scotia. The deposit from which it was found is part of the late Carnian Wolfville Formation, comprising a section of the larger Newark Supergroup that stretches across Canada and the eastern United States. The formation is located in the Fundy Basin, a rift basin which, like other Newark basins, opened as Pangaea began to break apart at the end of the Triassic. The Wolfville Formation contains a diverse assemblage of Triassic tetrapods that includes temnospondyl amphibians, procolophonid reptiles, and traversodont cynodonts. Based on the presence of the temnospondyl Koskinonodon (a common Triassic index fossil), the assemblage is dated to the Carnian stage of the Late Triassic. [2]
The type species of Teraterpeton, T. hrynewichorum, was named in 2003. It is named after George P. Hrynewich and his son Sandy Hrynewich, fossil collectors who discovered the bones at Burntcoat Head in 1999. [2]
Postcranial remains from a referred specimen, NSM 018GF010.002, as well as an isolated cervical vertebra (NSM 018GF010.001) were described by Pritchard & Sues in 2019. [3]
A phylogenetic analysis was provided in the original description of Teraterpeton in 2003. It placed Trilophosaurus as the closest relative of the genus. Together, these two forms comprise an archosauromorph clade that is more derived than protorosaurians and more basal than rhynchosaurs. Below is a cladogram modified from the 2003 analysis: [2]
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.
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.
Tanystropheus is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs. Tanystropheus is one of the most well-described non-archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae, a clade collecting many long-necked Triassic archosauromorphs previously described as "protorosaurs" or "prolacertiforms".
Azendohsaurus is an extinct genus of herbivorous archosauromorph reptile from roughly the late Middle to early Late Triassic Period of Morocco and Madagascar. The type species, Azendohsaurus laaroussii, was described and named by Jean-Michel Dutuit in 1972 based on partial jaw fragments and some teeth from Morocco. A second species from Madagascar, A. madagaskarensis, was first described in 2010 by John J. Flynn and colleagues from a multitude of specimens representing almost the entire skeleton. The generic name "Azendoh lizard" is for the village of Azendoh, a local village near where it was first discovered in the Atlas Mountains. It was a bulky quadruped that unlike other early archosauromorphs had a relatively short tail and robust limbs that were held in an odd mix of sprawled hind limbs and raised forelimbs. It had a long neck and a proportionately small head with remarkably sauropod-like jaws and teeth.
Trilophosaurs are lizard-like Triassic allokotosaur reptiles related to the archosaurs. The best known genus is Trilophosaurus, a herbivore up to 2.5 metres long. It had a short, unusually heavily built skull, equipped with massive, broad flattened cheek teeth with sharp shearing surfaces for cutting up tough plant material. Teeth are absent from the premaxilla and front of the lower jaw, which in life were probably equipped with a horny beak.
Hyperodapedon is an extinct genus of rhynchosaur reptiles which lived during Late Triassic period. Like other rhynchosaurs, it was an heavily built archosauromorph, distantly related to archosaurs such as crocodilians and dinosaurs. Hyperodapedon in particular was part of the subfamily Hyperodapedontinae, a specialized rhynchosaurian subgroup with broad skulls, beaked snouts, and crushing tooth plates on the roof of the mouth.
Trilophosaurus is a lizard-like trilophosaurid allokotosaur known from the Late Triassic of North America. It was a herbivore up to 2.5 m long. It had a short, unusually heavily built skull, equipped with massive, broad flattened cheek teeth with sharp shearing surfaces for cutting up tough plant material. Teeth are absent from the premaxilla and front of the lower jaw, which in life were probably equipped with a horny beak.
Mesosuchus is an extinct genus of basal Rhynchosaur from early Middle Triassic deposits of Eastern Cape, South Africa. It is known from the holotype SAM 5882, a partial skeleton, and from the paratypes SAM 6046, SAM 6536, SAM 7416 and SAM 7701 from the Aliwal North Euparkeria site. Mesosuchus is quite small, spanning around 30 cm in length. Mesosuchus was discovered and named by David Meredith Seares Watson in 1912.
Vancleavea is a genus of extinct, armoured, non-archosaurian archosauriforms from the Late Triassic of western North America. The type and only known species is V. campi, named by Robert Long & Phillip A Murry in 1995. At that time, the genus was only known from fragmentary bones including osteoderms and vertebrae. However, since then many more fossils have been found, including a pair of nearly complete skeletons discovered in 2002. These finds have shown that members of the genus were bizarre semiaquatic reptiles. Vancleavea individuals had short snouts with large, fang-like teeth, and long bodies with small limbs. They were completely covered with bony plates known as osteoderms, which came in several different varieties distributed around the body. Phylogenetic analyses by professional paleontologists have shown that Vancleavea was an archosauriform, part of the lineage of reptiles that would lead to archosaurs such as dinosaurs and crocodilians. Vancleavea lacks certain traits which are present in most other archosauriforms, most notably the antorbital, mandibular and supratemporal fenestrae, which are weight-saving holes in the skulls of other taxa. However, other features clearly support its archosauriform identity, including a lack of intercentra, the presence of osteoderms, an ossified laterosphenoid, and several adaptations of the femur and ankle bones. In 2016, a new genus of archosauriform, Litorosuchus, was described. This genus resembled both Vancleavea and more typical archosauriforms in different respects, allowing Litorosuchus to act as a transitional fossil linking Vancleavea to less aberrant archosauriforms.
Cosesaurus is a genus of archosauromorph reptiles likely belonging to the family Tanystropheidae. It is known from fossil imprints of a single small skeleton, MGB V1, which was found in Muschelkalk outcrops near the municipalities of Mont-ral and Alcover in Spain. These outcrops are dated to the Ladinian age of the middle Triassic about 242 to 237 million years ago. The specimen is stored at the Museu Martorell, which is now part of the Museu de Ciències Naturals de Barcelona. The poor preservation and likely juvenile nature of the specimen has led to the anatomy of Cosesaurus being misidentified by several different sources. For example, Paul Ellenberger claimed that it was an ancestor to birds in the 1970s, while David Peters claimed that it was a pterosaur ancestor in 2000. Both of these claims contrast with mainstream scientific theories on the origins of either group, and other paleontologists who study the specimen are unable to find the features which Ellenberger or Peters reported to be present. The Ellenberger and Peters hypotheses are thus considered fringe theories with questionable scientific soundness due to their low reproducibility. Mainstream hypotheses for the relations of Cosesaurus generally agree that it is a "protorosaur", specifically a tanystropheid closely related to long-necked reptiles such as Macrocnemus, Tanytrachelos, Tanystropheus, or Langobardisaurus.
Jesairosaurus is an extinct genus of early archosauromorph reptile known from the Illizi Province of Algeria. It is known from a single species, Jesairosaurus lehmani. Although a potential relative of the long-necked tanystropheids, this lightly-built reptile could instead be characterized by its relatively short neck as well as various skull features.
Protorosauria is an extinct, likely paraphyletic group of basal archosauromorph reptiles from the latest Middle Permian to the end of the Late Triassic of Asia, Europe and North America. It was named by the English anatomist and paleontologist Thomas Henry Huxley in 1871 as an order, originally to solely contain Protorosaurus. Other names which were once considered equivalent to Protorosauria include Prolacertiformes and Prolacertilia.
Jaxtasuchus is an extinct genus of armored doswelliid archosauriform reptile known from the Middle Triassic of the Erfurt Formation in Germany. The type species, Jaxtasuchus salomoni, was named in 2013 on the basis of several incomplete skeletons and other isolated remains. Like other doswelliids, members of the genus were heavily armored, with four longitudinal rows of bony plates called osteoderms covering the body. Jaxtasuchus is the first doswelliid known from Europe and is most closely related to Doswellia from the Late Triassic of the eastern United States. However, it was not as specialized as Doswellia, retaining several generalized archosauriform characteristics and having less armor. Jaxtasuchus fossils have been found in aquatic mudstones alongside fossils of temnospondyl amphibians, crustaceans, and mollusks, suggesting that Jaxtasuchus was semiaquatic like modern crocodilians.
Pamelaria is an extinct genus of allokotosaurian archosauromorph reptile known from a single species, Pamelaria dolichotrachela, from the Middle Triassic of India. Pamelaria has sprawling legs, a long neck, and a pointed skull with nostrils positioned at the very tip of the snout. Among early archosauromorphs, Pamelaria is most similar to Prolacerta from the Early Triassic of South Africa and Antarctica. Both have been placed in the family Prolacertidae. Pamelaria, Prolacerta, and various other Permo-Triassic reptiles such as Protorosaurus and Tanystropheus have often been placed in a group of archosauromorphs called Protorosauria, which was regarded as one of the most basal group of archosauromorphs. However, more recent phylogenetic analyses indicate that Pamelaria and Prolacerta are more closely related to Archosauriformes than are Protorosaurus, Tanystropheus, and other protorosaurs, making Protorosauria a polyphyletic grouping.
Fuyuansaurus is an extinct genus of "protorosaur" reptiles known from the Middle Triassic Zhuganpo Formation of southern China. Fuyuansaurus was first named by Nicholas C. Fraser, Olivier Rieppel and Li Chun in 2013 and the type species is Fuyuansaurus acutirostris.
Ozimek is a genus of sharovipterygid archosauromorph reptile, known from Late Triassic deposits in Poland and closely related to the Kyrgyzstani Sharovipteryx. It contains one species, O. volans, named in 2016 by Jerzy Dzik and Tomasz Sulej. Like Sharovipteryx, Ozimek had long, slender limbs with the hindlimbs longer than the forelimbs; the hindlimbs likely supported gliding membranes as fossilized in Sharovipteryx. Another unusual characteristic was the shoulder girdle, where the massive coracoids formed a shield-like structure covering the bottom of the shoulder region that would have limited mobility. In other respects, such as its long neck, it was a typical member of the non-natural grouping Protorosauria. Phylogenetic analysis has indicated that it, possibly along with Sharovipteryx, may have been an unusual member of the protorosaur group Tanystropheidae, although further study of its anatomy is needed to resolve its precise relationships.
Shringasaurus is an extinct genus of archosauromorph reptile from the Middle Triassic (Anisian) of India. It is known from the type and only known species, S. indicus. Shringasaurus is known from the Denwa Formation in the state of Madhya Pradesh. Shringasaurus was an allokotosaur, a group of unusual herbivorous reptiles from the Triassic, and is most closely related to the smaller and better known Azendohsaurus in the family Azendohsauridae. Like some ceratopsid dinosaurs, Shringasaurus had two large horns over its eyes that faced up and forwards from its skull. Shringasaurus also bears convergent physical similarities to sauropodomorph dinosaurs, such as its long neck, its shoulders and forelimbs, and the shape of its teeth. Shringasaurus possibly occupied a similar ecological niche as a large browsing herbivore before such dinosaurs had evolved.
Boreopricea is an extinct genus of archosauromorph reptile from the Early Triassic of arctic Russia. It is known from a fairly complete skeleton discovered in a borehole on Kolguyev Island, though damage to the specimen and loss of certain bones has complicated study of the genus. Boreopricea shared many similarities with various other archosauromorphs, making its classification controversial. Various studies have considered it a close relative of Prolacerta, tanystropheids, both, or neither. Boreopricea is unique among early archosauromorphs due to possessing contact between the jugal and squamosal bones at the rear half of the skull.
Polymorphodon is an extinct genus of archosauriform reptile from the Middle Triassic of Germany. The only known species is Polymorphodon adorfi, discovered in Lower Keuper deposits at a quarry in Eschenau, Germany. Polymorphodon is notable for its heterodont dentition, with long and conical premaxillary teeth followed by thin maxillary teeth with large serrations. Maxillary teeth near the back of the mouth are short and leaf-shaped, similar to some living and extinct reptiles with a herbivorous or omnivorous diet. This may suggest that Polymorphodon had some reliance on plants in its diet, a rarity among basal archosauriforms, most of which are carnivores.
Oryctorhynchus is an extinct genus of rhynchosaur from the Late Triassic (Carnian-Norian)-aged Wolfville Formation of Nova Scotia, Canada that may have been the same animal as Beesiiwo. The type species, O. bairdi, was named and described in 2020. It was originally seen as a species of Hyperodapedon until 2020.