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''Tambachia'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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
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 th ...
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 ...
within the
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 ...
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 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), 01'' (Richard Müller album), 2001 * 01 (Son of Dave album), ''01'' (Son of Dave album), 2000 * 01 (Urban ...
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 larg ...
,
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 territorie ...
.


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 several ...
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 characteristic ...
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 envi ...
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 theraps ...
trackways and articulated skeletons of terrestrial and semiterrestrial
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s and
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s. 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 fluviog ...
deposits consisting of well consolidated
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
s 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 t ...
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, th ...
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 In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
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 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 reptile ...
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, 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 ''Acheloma'' (also known as ''Trematops milleri'') is an extinct genus of temnospondyl that lived during the Early Permian. The type species is ''A. cumminsi''. History of study ''Acheloma'' was named by Edward Drinker Cope in 1882 based o ...
'', 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 The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
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 ''Anconastes'' is an extinct genus of dissorophoid temnospondyl within the family Trematopidae. It is known from two specimens from the Late Carboniferous Cutler Formation of north-central New Mexico in the southwestern United States. The genus n ...
'' 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. The 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 d ...
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 t ...
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