Teraterpeton
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''Teraterpeton'' (meaning "wonderful creeping thing" in
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
) is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of trilophosaurid
archosauromorph Archosauromorpha (Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) rather than lepidosaurs (such as tuataras, liza ...
s. It is known from a partial skeleton from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
Wolfville Formation of
Nova Scotia Nova Scotia ( ; ; ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
, 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 Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
, Teraterpetidae, related to ''
Trilophosaurus ''Trilophosaurus'' (Greek for "lizard with three ridges") 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 wi ...
''. Newer studies generally place it within
Trilophosauridae Trilophosaurs are lizard-like Triassic allokotosaur reptiles related to the archosaurs. The best known genus is ''Trilophosaurus'', a herbivore up to long. It had a short, unusually heavily built skull, equipped with massive, broad flattened che ...
.


Description


Skull

''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 bone The prefrontal bone is a bone separating the lacrimal and frontal bones in many tetrapod skulls. It first evolved in the sarcopterygian clade Rhipidistia, which includes lungfish and the Tetrapodomorpha. The prefrontal is found in most modern and ...
s. 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 The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
runs alongside the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
ry 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. ''Teraterpeton'' has an uncommon feature compared to most archosauromorphs: a
euryapsid __NOTOC__ Euryapsida is a polyphyletic (unnatural, as the various members are not closely related) group of Sauropsida, sauropsids that are distinguished by a single temporal fenestra, an opening behind the orbit (anatomy), orbit, under which the ...
-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 Diapsids ("two arches") 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 ...
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
plesiosaur The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared ...
s and
ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
s are not closely related to ''Teraterpeton''. However, the euryapsid archosauromorph ''Trilophosaurus'' has been identified as a close relative. 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 A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
. 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 An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
would normally be. In fact,
Hans-Dieter Sues Hans-Dieter Sues (born January 13, 1956) is a German-born American paleontologist who is Senior Scientist and Curator of Vertebrate Paleontology at the National Museum of Natural History of the Smithsonian Institution in Washington, DC. He receiv ...
, 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.


Postcranial skeleton

A string of eight
cervical In anatomy, cervical is an adjective that has two meanings: # of or pertaining to any neck. # of or pertaining to the female cervix: i.e., the ''neck'' of the uterus. *Commonly used medical phrases involving the neck are **cervical collar **cervic ...
(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). The
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
(shoulder blade) is long and narrow. The
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulates have ungual phalanges, as did the sauropods and horned dinosaurs. A claw is a highly modified ungual ...
s (claws) of both the hands and feet are deep and blade-like, a morphology described as "trenchant". 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 ''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Triassic, Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery ...
''. The pubis and
ischium The ischium () form ...
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. 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.


History

''Teraterpeton'' is primarily known from a single partial
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
skeleton referred to as NSM 999GF041. This skeleton was found in a sea cliff in
Burntcoat Head, Nova Scotia Burntcoat Head (improperly known as Burncoat) is an unincorporated rural Canadian community in Hants County, Nova Scotia. The area is known for having the largest tidal range (the greatest difference in height between high tide and low tide) of ...
. The deposit from which it was found is part of the late
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
Wolfville Formation, comprising a section of the larger
Newark Supergroup The Newark Supergroup, also known as the Newark Group, is an assemblage of Upper Triassic and Lower Jurassic sedimentary rocks which outcrop intermittently along the United States East Coast. They were deposited in a series of Triassic basins app ...
that stretches across
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
and the eastern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
. The formation is located in the
Fundy Basin The Fundy Basin is a sediment-filled rift basin on the Atlantic coast of southeastern Canada. It contains three sub-basins; the Fundy sub-basin, the Minas Basin and the Chignecto Basin. These arms meet at the Bay of Fundy, which is contained with ...
, a rift basin which, like other Newark basins, opened as
Pangaea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
began to break apart at the end of the Triassic. The Wolfville Formation contains a diverse assemblage of Triassic tetrapods that includes
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
amphibians,
procolophonid Procolophonidae is an extinct family of small, lizard-like parareptiles known from the Late Permian to Late Triassic that were distributed across Pangaea, having been reported from Europe, North America, China, South Africa, South America, Antarc ...
reptiles, and traversodont
cynodont The cynodonts () (clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variety ...
s. Based on the presence of the temnospondyl ''
Koskinonodon ''Anaschisma'' is an extinct genus of large temnospondyl amphibians. These animals were part of the family called Metoposauridae, which filled the crocodile-like predatory niches in the late Triassic. It had large skull about long, and possi ...
'' (a common Triassic
index fossil Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock Stratum, strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictiona ...
), the assemblage is dated to the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followed by t ...
stage of the Late Triassic. 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. 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.


Classification

A phylogenetic analysis was provided in the original description of ''Teraterpeton'' in 2003. It placed ''
Trilophosaurus ''Trilophosaurus'' (Greek for "lizard with three ridges") 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 wi ...
'' as the closest relative of the genus. Together, these two forms comprise an archosauromorph clade that is more
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
than
protorosauria Protorosauria is an extinct polyphyletic group of archosauromorph reptiles from the latest Middle Permian (Capitanian stage) to the end of the Late Triassic (Rhaetian stage) of Asia, Europe and North America. It was named by the English anatomis ...
ns and more basal than
rhynchosaur Rhynchosaurs are a group of extinct herbivorous Triassic archosauromorph reptiles, belonging to the order Rhynchosauria. Members of the group are distinguished by their triangular skulls and elongated, beak like premaxillary bones. Rhynchosaurs ...
s. Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
modified from the 2003 analysis:


References


External links


A photograph of the skull, published by the Fundy Geological Museum twitter accountAnother view of the skull, published by Victoria Arbour's twitter accountA photograph of hip and vertebral elements, published by the Nova Scotia Museum twitter account
{{Taxonbar, from=Q7701615 Allokotosaurs Prehistoric reptile genera Late Triassic reptiles of North America Triassic Canada Fossils of Canada Paleontology in Nova Scotia Fossil taxa described in 2003 Taxa named by Hans-Dieter Sues