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''Metoposaurus'' meaning "front lizard" is an extinct
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
stereospondyl The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
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 ...
s, known 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 a ...
of Germany, Italy, Poland, and Portugal. https://www.app.pan.pl/archive/published/app47/app47-535.pdf This mostly aquatic animal possessed small, weak limbs, sharp teeth, and a large, flat head. This highly flattened creature mainly fed on
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
, which it captured with its wide
jaw The jaws are a pair of opposable articulated structures at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth ...
s lined with needle-like
teeth A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
. Many ''Metoposaurus'' mass graves have been found, probably from creatures that grouped together in drying pools during
drought A drought is a period of drier-than-normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, ...
.


Discovery and species


Discovery

The earliest mention of Metoposauridae dates back to 1842 when Von Meyer described the dorsal view of the skull roof of a labyrinthodont from the
Keuper The Keuper is a lithostratigraphic unit (a sequence of rock strata) in the subsurface of large parts of west and central Europe. The Keuper consists of dolomite, shales or claystones and evaporites that were deposited during the Middle and Lat ...
Schilfsandstein of Feuerbacher Haide near
Stuttgart Stuttgart (; ; Swabian German, Swabian: ; Alemannic German, Alemannic: ; Italian language, Italian: ; ) is the capital city, capital and List of cities in Baden-Württemberg by population, largest city of the States of Germany, German state of ...
. Later, Meyer attempted a reconstruction of the same specimen and named it ''Metopias diagnosticus''. However, Lydekker later renamed the species as ''Metoposaurus diagnosticus'' in 1890 because the name ''Metopias'' was already in use.


Species

Based on the position of the lacrimal in the
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
, Metoposauridae is divided into two lineages. The group with the lacrimal external to the
orbit In celestial mechanics, an orbit (also known as orbital revolution) 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 ...
includes ''
Koskinonodon ''Anaschisma'' ("ripped up") is an extinct genus of large temnospondyls. These animals were part of the family called Metoposauridae, which filled the crocodile-like predatory Ecological niche, niches in the late Triassic. It had a large skull ab ...
bakeri'', ''
Dutuitosaurus ''Dutuitosaurus'' is a genus of metoposaurids, a group of temnospondyls that lived during the Late Triassic period. ''Dutuitosaurus'' was discovered in the early 1960s in Morocco and is known from the lower t5 units of the Timezgadiouine Formati ...
ouazzoui'', ''
Arganasaurus ''Arganasaurus'' is an extinct genus of temnospondyls belonging to the family Metoposauridae that lived in Morocco during the Late Triassic (Carnian). Taxonomy The type species of ''Arganasaurus'', ''A. lyazidi'', was originally described as ''M ...
lyazidi'', and ''
Apachesaurus ''Apachesaurus'' is an extinct genus of metoposaurid temnospondyls from western North America. Description and taxonomy ''Apachesaurus'' was described from the Late Triassic (late Norian-Rhaetian The Rhaetian is the latest age (geology), ...
gregorii''. They show a tendency towards a decreasing depth of the otic notch and a decrease in body size. ''Metoposaurus diagnosticus'', ''Metoposaurus diagnosticus krasiejowensis'' and ''Panthasaurus maleriensis'' fall in the category of the lacrimal forming the orbital margin. * ''Metoposaurus diagnosticus'' (Meyer, 1842) Fossils were found among other locations in the
Gres à Avicula contorta Formation The Gres a Avicula contorta is a geological formation in France. It dates back to the late Norian.Weishampel, David B; et al. (2004). "Dinosaur distribution (Late Triassic, Europe)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszka ...
of France, the
Weser Formation The Weser () is a river of Lower Saxony in north-west Germany. It begins at Hannoversch Münden through the confluence of the Werra and Fulda. It passes through the Hanseatic city of Bremen. Its mouth is further north against the ports of Br ...
of Germany and the Raibl Formation of Italy. The species has an anterior top of the lacrimal closer to the nares than the top of the prefrontal; interclavicle with posterior part longer than in ''Panthasaurus maleriensis''. ''Metoposaurus diagnosticus diagnosticus'' lived in the western
Germanic Basin The Germanic Basin () is a large region of sedimentation in Western and Central Europe that, during the Permian and Triassic periods, extended from England in the west to the eastern border of Poland in the east. To the south it is bounded by the ...
, from at least the Schilfsandstein to the
Lehrberg Lehrberg is a market town in the district of Ansbach (district), Ansbach, Mittelfranken, Bavaria, Germany. Villages There are districts mentioned below: References External links

Ansbach (district) {{Ansbachdistrict-geo-stub ...
Beds sedimentation. * ''Metoposaurus krasiejowensis'' (Sulej, 2002) Fossils were found in the Drawno Beds Formation of Poland. The species has a very short perineal part of the parietal with the high value of the expansion angle of the sutures separating partially from the supratemporal (mean angle value of 21.81). Some skulls possess a large quadrate
foramen In anatomy and osteology, a foramen (; : foramina, or foramens ; ) is an opening or enclosed gap within the dense connective tissue (bones and deep fasciae) of extant and extinct amniote animals, typically to allow passage of nerves, artery, ...
and small paraquadrate foramen. It had an estimated skull length up to . * ''Metoposaurus algarvensis'' (Brusatte et al. 2015) From the Late Triassic Grès de Silves Formation in the Algarve, Portugal. It has a broader skull than any other ''Metoposaurus''. It was named by
Stephen Brusatte Stephen Louis "Steve" Brusatte FRSE (born April 24, 1984) is an American author and evolutionary biologist who specializes in the anatomy and evolution of dinosaurs. He was educated at the University of Chicago for his Bachelor's degree, at the ...
, Richard Butler,
Octávio Mateus Octávio Mateus (born 1975) is a Portugal, Portuguese dinosaur paleontologist and biologist Professor of Paleontology at the Faculdade de Ciências e Tecnologia da NOVA University Lisbon, Universidade Nova de Lisboa. He graduated in University of � ...
and Sebastien Steyer. It had an estimated mandible length up to .


Formerly assigned species

* ''Metoposaurus maleriensis'' (Chowdhury, 1965) Described from the Maleri Formation of Central India,Chowdhury, T.R. 1965. A new metoposaurid amphibian from the Upper Triassic Maleri Formation of Central India. Philosophical Transactions of the Royal Society of London B 250, 1–52. was renamed '' Panthasaurus maleriensis'' by Chakravorti and Sengupta (2018). * ''Metoposaurus azerouali'' (Dutuit, 1976) Described from the Argana Group of Morocco. Reassigned to ''Arganasaurus'' by Buffa et al. (2019).


Synonyms and ''nomina dubia''

# ''Metoposaurus stuttgartensis'': synonymous, is first described by Fraas (1913) from the Keuper Lehbergstufe of Sonnenberg, near Stuttgart. Fraas identified the species based on the interclavicle and left clavicle, vertebrae and rib fragments which are now located in the Stuttgart Museum. # ''Metoposaurus santaecrucis'' ''Nomina dubia'' it was described by Koken (1913) based on the partial skull found in Heiligenkreuz and the specimen is now located in the University Museum of Tübingen. # ''Metoposaurus heimi'' synonymous it was described by Kuhn (1932) based on a complete skull from the middle Keuper Blasensandstein in Upper Franconia. The specimen is currently located in the Museum of Paleontology and Historical Geology, Munich.


Description


Skull


Lacrimal

The lacrimal contacts the nasal medially, the
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
laterally, the prefrontal posteromedially, and the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
posteriorly. Metoposaur taxonomy was based on the position of the
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
, and differing opinions have been published. According to a photograph published by Hunt (1993), it is noted that the lacrimal enters the orbit, contrary to the previous finding by Fraas (1889). According to Lucas, close examination of the skull and other metoposaur skulls does not support this claim, and it has been noted that the misidentification was possibly due to the poor preservation of the fossil. In 2007, Sulej noted that the variability in the position of the lacrimal is narrow enough to be used for phylogenic analysis, but with caution.


Parietal

A study conducted by Sulej (2007) shows that the parietal contacts the frontal anteriorly, the postfrontal anterolaterally, the supratemporal laterally, and the postparietal posteriorly. The pineal foramen is in the posterior region of the parietal. An interesting feature pointed out by Sulej on examining the skull of ''Metoposaurus diagnosticus krasiejowensis'' is that it has a shorter prepineal region of the parietal than ''Metoposaurus diagnosticus diagnosticus'' and the expansion angle of the suture separating the parietal from the supratemporal has a lower value.


Maxilla

The
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
forms a large, completely dentigerous shelf bearing 83 to 107 teeth. The first teeth are large, and tooth size decreases markedly further posteriorly. On the ventral side, the maxilla contacts the ectopterygoid, palatine and vomer. In the choanal region, the maxilla is slightly broadening medially on the palatal side where it borders the choana. The margin of the choana is variable. In most skulls, it is weakly distinguished and rounded, but in a few cases it is more solid and sharply outlined. palaeontologia.pan.pl/PP64/Sulej.pdf


Vertebral column

According to Sulej (2007), the intercentra of cervical and
thoracic The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main ...
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
are fully ossified. The pleurocentra are not preserved and no evidence is found that they were present as cartilages. The
atlas An atlas is a collection of maps; it is typically a bundle of world map, maps of Earth or of a continent or region of Earth. Advances in astronomy have also resulted in atlases of the celestial sphere or of other planets. Atlases have traditio ...
,
axis An axis (: axes) may refer to: Mathematics *A specific line (often a directed line) that plays an important role in some contexts. In particular: ** Coordinate axis of a coordinate system *** ''x''-axis, ''y''-axis, ''z''-axis, common names ...
, and third and fourth cervical
vertebrae Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
are characteristic and similar to those in other stereospondyls. The morphology of the atlas, axis, and third and fourth vertebrae suggests that the neck of ''Metoposaurus'' was relatively flexible. The intercentra of the region where the vertebral column contacts the shoulder girdle are flat anteriorly and posteriorly. The neural arches have almost vertically set prezygapophyses (vide postcervical vertebrae). This suggests that in this region the lateral bending of the vertebral column was very limited. It was probably connected through articulation of the vertebral column with the shoulder girdle. A stiffening of the vertebral column, in the region of contacting limbs, was apparently essential for swimming. He also describes that in ''Metoposaurus diagnosticus krasiejowensis'', the parapophyses become shorter posteriorly, similarly to plagiosaurids. The intercentra of dorsal and sacral vertebrae are fully ossified and form quite short disks, not connected with the neural arches. In the dorsal and sacral region, they have anterior and posterior surfaces that are concave, or the posterior surface is almost flat. This condition resembles that in the trunk of plesiosaurs and, to some degree, the ichthyosaurs, confirming the aquatic mode of life.


Geography and history

Metoposaurids are known from the early Late Triassic (
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...
) Keuper of
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
Austria Austria, formally the Republic of Austria, is a landlocked country in Central Europe, lying in the Eastern Alps. It is a federation of nine Federal states of Austria, states, of which the capital Vienna is the List of largest cities in Aust ...
. There have also been unconfirmed notifications reported from
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 ...
(Dutuit 1978) and
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
(Yang 1978). ''Metoposaurus diagnosticus krasiejoviensis'' is the most abundant metoposaurid amphibian of the
Krasiejów Krasiejów is a village in the administrative district of Gmina Ozimek, within Opole County, Opole Voivodeship, in south-western Poland. It lies approximately east of Ozimek and east of the regional capital Opole. Paleontological localities ...
site (the species name comes from the site) located in southern
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
. Among stereospondyls, metoposaurs seem to have been one of the latest survivors. However, a variety of other temnospondyl lines carried into the Jurassic, the latest of which was another stereospondyl, the chigutisaurid ''
Koolasuchus ''Koolasuchus'' is an extinct genus of brachyopoid temnospondyl in the family Chigutisauridae. Fossils have been found from Victoria, Australia and date back 125-120 million years ago to Barremian-Aptian stages of the Early Cretaceous. ''Koola ...
'', discovered in modern-day Australia, where it was supported by a colder mid-Cretaceous climate.


Paleobiology


Locomotion

Examination of the vertebral column and limb articulations of ''Metoposaurus'' suggests that they used their limbs as flippers and swam by making simultaneous and symmetrical limb movements similar to plesiosaurs. A recent study conducted in
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
suggests that the broad, flat head and arm bones, wide hands, and large tail of ''Metoposaurus diagnosticus'' are significant characteristics which led the researchers to conclude that they swam in ephemeral lakes during the wet season and used their heads and forearms to burrow under the ground when the dry season began. The study found that the medullary region is filled with well-developed trabecular bone. The growth marks in all bones are organized as thick layers of highly vascularized zones and thick compact annuli with numerous rest lines, which may correspond with favorably wet and long, unfavorably dry seasons.


Predators

The exact predators of ''Metoposaurus'' are unknown, but
phytosaurs Phytosaurs (Φυτόσαυροι in Greek, meaning 'plant lizard') are an extinct group of large, mostly semiaquatic Late Triassic archosauriform or basal archosaurian reptiles. Phytosaurs belong to the order Phytosauria and are sometimes ...
were found closely associated in
bone bed A bone bed is any Geology, geological stratum or deposition (geology), deposit that contains bones of whatever kind. Inevitably, such deposits are Sedimentary rock, sedimentary in nature. Not a formal term, it tends to be used more to describe esp ...
s.Mateus, O., Butler R. J., Brusatte S. L., Whiteside J. H., & Steyer S. J. (2014). The first phytosaur (Diapsida, Archosauriformes) from the Late Triassic of the Iberian Peninsula. Journal of Vertebrate Paleontology. 34(4), 970-975


References


Further reading

* Sengupta, D. P. (2002). Indian metoposaurid amphibians revised. Paleontological Research, 6(1), 41-65.


External links


Wrong number of fingers leads down wrong track
on: EurekAlert!. Source: News Release 24-Jul-2020, University of Bonn {{Taxonbar, from=Q2326287 Metoposauridae Norian genera Triassic temnospondyls of Europe Triassic France Fossils of France Triassic Germany Fossils of Germany Fossils of Italy Fossils of Poland Fossils of Portugal Fossil taxa described in 1890 Taxa named by Richard Lydekker Late Triassic amphibians of Europe Prehistoric amphibian genera