''Trematosaurus'' is an
extinct
Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
trematosaurid temnospondyl
Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
amphibian
Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
found in
Germany
Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
and
Russia
Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
.
It was first named by Hermann Burmeister in 1849 and the type species is ''Trematosaurus brauni''.
History of study
''Trematosaurus'' was one of the first temnospondyls to be described. The type locality, called Merkel's Quarry, is in east-central Germany
Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
at Bernburg an der Saale within the Bausandstein (Olenekian
In the geologic timescale, the Olenekian is an age (geology), age in the Early Triassic epoch (geology), epoch; in chronostratigraphy, it is a stage (stratigraphy), stage in the Lower Triassic series (stratigraphy), series. It spans the time betw ...
) and was collected for several decades from the 1840s into the early 20th century, producing extensive cranial remains, although the majority of these are preserved as internal molds (steinkerns) or natural molds. The name ''Trematosaurus'' was in fact coined in 1842 by Carl von Braun, a frequent collector who used the Greek suffix ''trema'' ('hole') in reference to the pineal foramen to form the generic epithet, but as he provided no formal description, the name was not considered valid until the work of Burmeister, who named the type species after Braun. Burmeister's work was largely reconstructive and frequently omitted references to which specimens were being described, and as a result, it remains unknown which specimens exactly were covered in his description. This was remedied by subsequent workers who provided specimen illustrations for material reposited at numerous museums across Europe. However, the complete osteology and ontogeny of this taxon remained poorly documented until the work of Schoch (2019).
Two other species were described from equivalent horizons in Thuringia
Thuringia (; officially the Free State of Thuringia, ) is one of Germany, Germany's 16 States of Germany, states. With 2.1 million people, it is 12th-largest by population, and with 16,171 square kilometers, it is 11th-largest in area.
Er ...
, ''T. fuchsi'' and ''T. thuringiensis''. ''Trematosaurus fuchsi'' is widely regarded as a junior synonym of ''T. brauni,'' while ''T. thuringiensis'' is tentatively considered to be valid with the caveat that poor preservation of the single specimen may account for some of the purported differences. ''Trematosaurus madagascariensis'' was synonymized with '' Tertremoides ambilobensis'' from the same horizon in Madagascar
Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
by Schoch & Milner (2000), with the species epithet of the former (named first) taking precedence by priority and the genus epithet of the latter taking precedence by distinction from ''Trematosaurus''. ''Trematosaurus sobeyi'' from Cynognathus Assemblage Zone of South Africa
South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
was placed in the novel genus '' Trematosuchus'' by Watson (1919). ''Trematosaurus kannemeyeri'' from the same zone is thought to belong to '' Aphaneramma'', but other workers do not consider it to be diagnostic and thus to be invalid. ''Trematosaurus yakovlevi'' from Russia was placed in the novel genus '' Thoosuchus'' by Efremov (1940). ''Trematosaurus weidenbaumi'' was first removed to ''Thoosuchus'' and then placed in the novel genus '' Angusaurus'' by Getmanov (1989). Various reports of ''Trematosaurus'' sp. from Russia
Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
belong to '' Inflectosaurus'', but ''Trematosaurus galae'' is a recently described Russian species that remains valid to date.
Description
The overwhelming majority of specimens of ''T. brauni'' are partial to complete skulls, with 75 such specimens known (size range from 10.9 to 41 cm in length). Postcrania are restricted to the clavicle and interclavicle, two large plate-like elements of the pectoral girdle. In part, this relates to the occurrence of other large stereospondyls at the type locality such as the capitosaur '' Parotosuchus nasutus'', but postcrania are generally scarce, which may be due to either collection bias or taphonomic bias. Therefore, all diagnostic features of ''T. brauni'' are based on the skull. Autapomorphies include (1) temporal sulcus of lateral line with two portions, (2) occipital sulcus continuous, (3) pre-orbital region slightly shorter than half skull length (0.43–0.49), and (4) interorbital distance wide (interorbital distance/ skull length: 0.15–0.2). The description is based only on referred specimens, as no holotype
A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
was ever designated and the syntypes described by Burmeister cannot be identified based on the scant details of his description; however, because they may exist, either within public or private collections, no neotype designation is warranted at present. ''Trematosaurus galae'' differs from ''T. brauni'' in (1) presence of well-pronounced lacrimal and zygomatic flexures of canals of the lateral line, (2) connection of both branches of the temporal groove at the ossification center of the supratemporal, and (3) in the presence of a supplementary groove (sulcus buccalis transversus) on the squamosal. The more questionably valid ''T. thuringiensis'' differs only in having larger orbits and a narrower skull table than the type species.
Given the large sample size of ''T. brauni'', its ontogeny can be characterized, exhibiting features such as decreased orbit size and skull width but not other features such as broadening of the interorbital region or increased preorbital length, which occur in other temnospondyls. Observations of more fragmentary material of ''T. galae'' indicate that specifically the prenarial region and the postorbital regions become longer, along with increased development of other features such as the basioccipital and ornamentation on the parasphenoid replacing a shagreen of denticles.
Classification
Below is a cladogram from Steyer (2002) showing the phylogenetic
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
relationships of trematosaurids:
''Trematosaurus'' in a cladogram after Novikov (2018) with only Early Triassic Eastern Europe taxa included:[ ]
References
{{Taxonbar, from=Q3776858
Triassic temnospondyls of Europe
Trematosaurines
Taxa named by Hermann Burmeister
Prehistoric amphibian genera