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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 n ...
of dissorophoid temnospondyl 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. It is known from the
Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * 01'' (Richard Müller album), 2001 * ''01'' (Son of Dave album), 2000 * ''01'' (Urban Zakapa album), 2011 * ''O1'' (Hiroyuki Sawano ...
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 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 (FRG),, is a country in Central Europe. It is the most populous member state of the European Union. Germany lies between the Baltic and North Sea to the north and the Alps to the sou ...
. ''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 U.S. state, states, a Washington, D.C., federal district, five ma ...
.


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 seve ...
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 ...
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 characterist ...
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 mining, open-pit mine in which dimension stone, rock (geology), rock, construction aggregate, riprap, sand, gravel, or slate is excavated from the ground. The operation of quarries is regulated in some juri ...
that is well known for
tetrapod Tetrapods (; ) are four-limb (anatomy), limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant taxon, extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (p ...
trackways and articulated skeletons of terrestrial and semiterrestrial amphibians and reptiles. MNG 7722 was found in red-bed
fluvial In geography and geology, fluvial processes are associated with rivers and streams and the deposits and landforms created by them. When the stream or rivers are associated with glaciers, ice sheets, or ice caps, the term glaciofluvial or fluv ...
deposits consisting of well
consolidated Consolidated may refer to: *Consolidated (band) **'' ¡Consolidated!'', a 1989 extended play *Consolidated Aircraft (later Convair), an aircraft manufacturer *Consolidated city-county *Consolidated Communications * Consolidated school district *Co ...
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 carpals, tarsals, and endochondral portion of the braincase suggest that MNG 7722 represents an early stage of development, while the pitted
skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
, with tightly closed sutures between bones, indicates a mature individual. The name ''Tambachia'' refers to the Tambach Formation. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
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 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, 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 similar to those seen in modern reptiles. The subdivision of the
external naris External may refer to: * External (mathematics), a concept in abstract algebra * Externality, in economics, the cost or benefit that affects a party who did not choose to incur that cost or benefit * Externals, a fictional group of X-Men antagon ...
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 Manus may refer to: * Manus (anatomy), the zoological term for the distal portion of the forelimb of an animal (including the human hand) * ''Manus'' marriage, a type of marriage during Roman times Relating to locations around New Guinea * Man ...
, or hand, and five digits on the
pes Pes (Latin for "foot") or the acronym PES may refer to: Pes * Pes (unit), a Roman unit of length measurement roughly corresponding with a foot * Pes or podatus, a * Pes (rural locality), several rural localities in Russia * Pes (river), a river ...
, 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, 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. The ...
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 ''Fedexia'' is an extinct genus of carnivorous temnospondyl within the family Trematopidae. It lived 300 million years ago during the late Carboniferous period. It is estimated to have been long, and likely resembled a salamander.
'', 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 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 Paleo ...
{{Taxonbar, from=Q7681002 Dissorophids Trematopids Permian temnospondyls of Europe Permian Germany Fossil taxa described in 1998 Prehistoric amphibian genera