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''Dyoplax'' 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
pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
n
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avi ...
, possibly an
erpetosuchid Erpetosuchidae is an extinct family of pseudosuchian archosaurs. Erpetosuchidae was named by D. M. S. Watson in 1917 to include ''Erpetosuchus''. It includes the type species '' Erpetosuchus granti'' from the Late Triassic of Scotland, ''Erpetos ...
.
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s have been found from the
type locality Type locality may refer to: * Type locality (biology) * Type locality (geology) See also * Local (disambiguation) * Locality (disambiguation) {{disambiguation ...
within the upper Schilfsandstein Formation in Stuttgart,
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 ...
. 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 was a natural cast of a nearly complete skeleton that lacked only parts of the tail and limb bones.


Classification

Oscar Fraas Oscar Friedrich von Fraas (17 January 1824, in Lorch (Württemberg) – 22 November 1897, in Stuttgart) was a German clergyman, paleontologist and geologist. He was the father of geologist Eberhard Fraas (1862–1915). Biography He studied theo ...
, the original describer of ''Dyoplax,'' described the specimen as having "the head of a
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia al ...
and the armor of a gavial". When the taxon
Pseudosuchia Pseudosuchia is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely related to crocodilians than to birds. Pseudosuchians are also informally known as "crocodilian-line archosaurs". Prior to ...
was first proposed in 1890, ''Dyoplax'' was considered one of the three genera within the clade, and was included within the family " Aetosauridae". Several other papers published in later years have also placed the genus within Pseudosuchia. In 1956 the genus was referred to Notochampsidae, now known as Protosuchidae. It was suggested to be a possible
erpetosuchid Erpetosuchidae is an extinct family of pseudosuchian archosaurs. Erpetosuchidae was named by D. M. S. Watson in 1917 to include ''Erpetosuchus''. It includes the type species '' Erpetosuchus granti'' from the Late Triassic of Scotland, ''Erpetos ...
in 1966, but was later classified as one of the earliest protosuchids in 1994. In that same year a paper was published that identified several synamomorphies characteristic of the clade Aetosauria. This confirmed that ''Dyoplax'' was not within the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Aetosauria as had been previously speculated because it lacked four out of five of the synapomorphies associated with Aetosauria. A paper published in 1998 by Spencer ''et al.'' considered Dyoplax to be within
Sphenosuchia Sphenosuchia is a suborder of basal crocodylomorphs that first appeared in the Triassic and occurred into the Middle Jurassic. Most were small, gracile animals with an erect limb posture. They are now thought to be ancestral to crocodyliforms, a ...
, a
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also r ...
of basal
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
. The authors claimed that all synapomorphies present within Crocodylomorpha, as defined in 1992 by Sereno & Wild, were present in ''Dyoplax''. They further concluded that the genus had most of the synapomorphies common to Sphenosuchia, lacking only the forked posterior process of the
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral co ...
. On this basis, the authors concluded that there was enough evidence to place ''Dyoplax'' within Sphenosuchia. The age of the specimen within the strata would make ''Dyoplax'' the oldest sphenosuchian known at the time, predating other genera from the late Carnian that were once thought to be the oldest members of Sphenosuchia such as ''
Hesperosuchus ''Hesperosuchus'' is an extinct genus of crocodylomorph reptile that contains a single species, ''Hesperosuchus agilis''. Remains of this pseudosuchian have been found in Late Triassic (Carnian) strata from Arizona and New Mexico.Colbert, E. H. ...
'' and ''
Parrishia ''Parrishia'' is an extinct genus of sphenosuchian crocodylomorph known from the Late Triassic Chinle, Dockum, and Santa Rosa Formations in Arizona and New Mexico. Discovery and naming The genus was named in 1995 from fossils found from t ...
''. However, Maisch, Matzke and Rathgeber (2013) questioned the placement of ''Dyoplax'' within Crocodylomorpha, and argued that it shared important cranial and postcranial features with '' Erpetosuchus''; the authors tentatively reassigned ''Dyoplax'' to Erpetosuchidae.


Paleoecology

The Shilfsandstein Formation was deposited during the early
Carnian The Carnian (less commonly, Karnian) is the lowermost stage of the Upper Triassic Series (or earliest age of the Late Triassic Epoch). It lasted from 237 to 227 million years ago (Ma). The Carnian is preceded by the Ladinian and is followe ...
stage of 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 ...
(~ 228 million years ago) in a
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into '' coastal lagoons'' (or ''barrier lagoons' ...
al paleoenvironment. Numerous
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biv ...
s, chondrichthyean
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...
such as '' Palaeobates'', trematosaurian temnospondyls such as ''
Metoposaurus ''Metoposaurus'' meaning "front lizard" is an extinct genus of stereospondyl temnospondyl amphibian, known from the Late Triassic of Germany, Italy, Poland, and Portugal. https://www.app.pan.pl/archive/published/app47/app47-535.pdf This mostly ...
'',Hunt, A. P. (1993). A revision of the Metoposauridae (Amphibia: Temnospondyli) of the Late Triassic with description of a new genus from the western United States. Museum of Northern Arizona Bulletins 59:67-97; Flagstaff a
phytosaur Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalent gr ...
, and plants such as ''
Neocalamites ''Neocalamites'' is an extinct genus of Equisetales, equisetalean plant. ''Neocalamites'' thrived during the Permian and Triassic, and occurs worldwide. Description According to Elgorriaga et al. 2018, characteristics of ''Neocalamites'' include ...
'' and ''
Equisetites Equisetaceae, sometimes called the horsetail family, is the only extant family of the order Equisetales, with one surviving genus, ''Equisetum'', which comprises about twenty species. Evolution and systematics Equisetaceae is the only survivin ...
'' were also present in the paleoenvironment that existed at the time.


References


External links


''Dyoplax''
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=Q5319247 Prehistoric pseudosuchian genera Triassic archosaurs Late Triassic reptiles of Europe