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''Tambachia'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
dissorophoid Dissorophoideans are a clade of medium-sized, temnospondyl amphibians that appeared during the Moscovian in Euramerica, and continued through to the Late Permian and the Early Triassic of Gondwana. They are distinguished by various details of the ...
temnospondyl Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carbo ...
within the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Trematopidae Trematopidae is a family of dissorophoid temnospondyl spanning the late Carboniferous to the early Permian. Together with Dissorophidae, the family forms Olsoniformes, a clade comprising the medium-large terrestrial dissorophoids. Trematopids a ...
. It is known from the Early Permian
Tambach Formation The Tambach Formation is an Early Permian-age geologic formation in central Germany. It consists of red to brown-colored sedimentary rocks (red beds) such as conglomerate, sandstone, and mudstone, and is the oldest portion of the Upper Rotliegend ...
(the lowermost unit of the Upper Rotliegend) near the town of
Tambach-Dietharz Tambach-Dietharz is a town in the district of Gotha, in Thuringia, Germany. It is situated in the Thuringian Forest, 19 km south of Gotha. Mayor Since 2012, Marco Schütz (independent) is the mayor. His predecessor was the former lieutenan ...
in
Thuringia Thuringia (; german: Thüringen ), officially the Free State of Thuringia ( ), is a state of central Germany, covering , the sixth smallest of the sixteen German states. It has a population of about 2.1 million. Erfurt is the capital and lar ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. ''Tambachia'' is the first trematopid to have been discovered outside the
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 territori ...
.


Discovery

The
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 sever ...
specimen of ''Tambachia trogallas'', known as MNG 7722, has been found from an outcrop of the Tambach Formation at the Bromacker locality in the
Thuringian Forest The Thuringian Forest (''Thüringer Wald'' in German), is a mountain range in the southern parts of the German state of Thuringia, running northwest to southeast. Skirting from its southerly source in foothills to a gorge on its north-west side i ...
of central Germany. It consists of a skull and much of the postcranial skeleton. The only major portion of the skeleton that is missing is the presacral
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
l column. The Bromacker locality is a
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates ...
quarry A quarry is a type of open-pit mine in which dimension stone, rock, construction aggregate, riprap, sand, gravel, or slate is excavated from the ground. The operation of quarries is regulated in some jurisdictions to reduce their envir ...
that is well known for
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
trackways and articulated skeletons of terrestrial and semiterrestrial amphibians and reptiles. MNG 7722 was found in red-bed fluvial deposits consisting of well consolidated mudstones in flat-bedded channel fills. Although it is difficult to determine the maturity of MNG 7722, it is thought to be an early adult. Poor
ossification Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in ...
of the
carpal The carpal bones are the eight small bones that make up the wrist (or carpus) that connects the hand to the forearm. The term "carpus" is derived from the Latin carpus and the Greek καρπός (karpós), meaning "wrist". In human anatomy, the ...
s, tarsals, and
endochondral Endochondral ossification is one of the two essential processes during fetal development of the mammalian skeletal system by which bone tissue is produced. Unlike intramembranous ossification, the other process by which bone tissue is produced, ...
portion of the braincase suggest that MNG 7722 represents an early stage of development, while the pitted skull roof, with tightly closed sutures between bones, indicates a mature individual. The name ''Tambachia'' refers to the Tambach Formation. The specific name of ''T. trogallas'' is derived from the
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 ...
''trogo'', meaning munch or nibble, and ''allas'', meaning
sausage A sausage is a type of meat product usually made from ground meat—often pork, beef, or poultry—along with salt, spices and other flavourings. Other ingredients, such as grains or breadcrumbs may be included as fillers or extenders. ...
. The name refers to the Thuringian
bratwurst Bratwurst () is a type of German sausage made from pork or, less commonly, beef or veal. The name is derived from the Old High German ''Brätwurst'', from ''brät-'', finely chopped meat, and ''Wurst'', sausage, although in modern German it is o ...
that was frequently eaten in the Bromacker Quarry by the authors who described the species.


Description

The skull of MNG 7722 is severely crushed dorsoventrally, making it hard to determine the shape of the skull in lateral view. It is in length, which is small for a trematopid. The
orbits In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as a p ...
, or eye sockets, are centrally placed so that the preorbital length of the skull is equal to that of the postorbital length. They make up around 30% of the skull length. The skull table seems to extend posteriorly past the level of the jaw articulation in ''Tambachia'', which is unique among trematopids. However, this is likely to have been the result of crushing of the skull, which displaced the skull table. ''Tambachia'', like other trematopids, likely possessed a
salt gland The salt gland is an organ for excreting excess salts. It is found in the cartilaginous fishes subclass elasmobranchii (sharks, rays, and skates), seabirds, and some reptiles. Salt glands can be found in the rectum of sharks. Birds and reptiles ...
similar to those seen in modern reptiles. The subdivision of the external naris into anterior and posterior portions is evidence of the presence of salt glands. The anterior portion is sub-circular and is thought to be the true narial opening. The posterior portion likely held the salt gland. The expansion of the posterior portion may not be an accommodation for the salt gland, but rather a means of coping with cranial stresses during feeding. The postcranial skeleton is well known in ''Tambachia'', given the completeness of the holotype specimen. The relatively short tail, preserved only as a faint impression in MNG 7722, is long. The limbs are short, with four digits on the manus, or hand, and five digits on the pes, or foot. It is possible that the manus possessed five digits, since '' Acheloma'', another closely related trematopid, has been described as having five distal carpals. In the pes of the holotype specimen, there are only four digits preserved. However, since the third preserved digit is the longest, it likely represents the fourth digit, with the first digit absent due to imperfect preservation. The terminal phalanges of ''Tambachia'' were narrow, and would have been the core supports of claws.


Classification

In the original description of ''Tambachia'', Sumida ''et al.'' (1998) concluded that '' Anconastes'' was the closest relative of ''Tambachia''. Characteristics that supported this relationship included the absence of an internarial fenestra (which is also seen in
dissorophids Dissorophidae is an extinct family of medium-sized, temnospondyl amphibians that flourished during the late Carboniferous and early Permian periods. The clade is known almost exclusively from North America. History of study Dissorophidae is a ...
, but was interpreted as a convergent trait), the reduction of the suborbital process of the lacrimal, and a contribution to the ventral orbital rim (the rim of the eye socket) by 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. T ...
rather than the
palatine A palatine or palatinus (in Latin; plural ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman times.
. 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 ...
modified from Sumida ''et al.'' (1998) showing the proposed relationships of ''Tambachia'' and other trematopids: In their description of '' Fedexia'', Berman ''et al.'' (2010) placed ''Fedexia'' as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of the clade containing ''Tambachia'' and ''Anconastes''. ''Tambachia'', ''Anconastes'', and ''Fedexia'' all possess a shallow and broadly concave
occipital The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cereb ...
margin. In other trematopids, the margin is biconcave. Below is a cladogram modified from Berman ''et al.'' (2010):


References


External links


''Tambachia''
in the
Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database (PBDB) originated in the NCEAS-funded Phanerozoic Marine Pale ...
{{Taxonbar, from=Q7681002 Dissorophids Trematopids Permian temnospondyls of Europe Permian Germany Fossil taxa described in 1998 Prehistoric amphibian genera