Parasuchus Temporal range: Late Triassic, | |
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The skull of ISI R42, the neotype of Parasuchus hislopi | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | † Phytosauria |
Genus: | † Parasuchus Lydekker, 1885 |
Type species | |
†Parasuchus hislopi Lydekker, 1885 |
Parasuchus is an extinct genus of basal phytosaur known from the Late Triassic (late Carnian to early Norian stage) of Andhra Pradesh and Madhya Pradesh, India. At its most restricted definition, Parasuchus contains a single species, Parasuchus hislopi. [1] [2] Parasuchus hislopi is one of several species belonging to a basal grade of phytosaurs, typified by the genus Paleorhinus . Historically, Paleorhinus has been known from better-described fossils, and many species have been lumped into that genus. Parasuchus hislopi, despite being described earlier than Paleorhinus, was considered an undiagnostic chimera until new neotype fossils were described in the late 1970s. Parasuchus hislopi and the two unambiguously valid species of Paleorhinus (P. bransoni and P. angustifrons) are all closely related; some authors have historically described them all under the species Paleorhinus, while others place the two Paleorhinus species into Parasuchus according to the principle of priority. [3]
The name Parasuchus was first used by Thomas Henry Huxley (1870) in a faunal list. Since a diagnosis wasn't provided, it would have been considered a nomen nudum at the time. Richard Lydekker (1885) formally described and named P. hislopi, and proposed the family name Parasuchidae. However, Lydekker's description was based on a chimeric syntype, combining fossils from multiple unrelated reptiles: a rhynchosaurian basicranium mixed with the partial snout of a phytosaur, scutes and some teeth. Friedrich von Huene (1940) identified the basicranium as belonging to Paradapedon huxleyi (now known as Hyperodapedon huxleyi) and the phytosaurian material to a newly named species, "aff." Brachysuchus maleriensis. Later, Edwin Harris Colbert (1958) designated all the Indian parasuchian material as Phytosaurus maleriensis while Gregory (1962) considered the material undiagnostic.
Sankar Chatterjee (1978) described many complete remains of the Indian parasuchian and showed that it is not assignable either to Brachysuchus (which is closely related to or synonymous with Angistorhinus ), or to Phytosaurus (a dubious name, probably the senior synonym of Nicrosaurus ). He argued that the rhynchosaur basicranium qualifies as neither the holotype of P. hislopi, nor the lectotype of Paradapedon huxleyi. He re-introduced P. hislopi, based on Lydekker's snout fossil and new well-preserved material. [1] To avoid additional confusion, the nondiagnostic holotype of P. hislopi was replaced by a neotype (ISI R 42) with approval from the ICZN (Opinion 2045) following the application of Chatterjee (2001). [4]
The partial premaxillary rostrum (snout) originally described by Lydekker, GSI H 20/11, was chosen as the lectotype of Parasuchus hislopi by Sankar Chatterjee. GSI H 20/11 was collected from the Lower Maleri Formation (Pranhita–Godavari Basin), near the Maleri village of Adilabad district, Andhra Pradesh. The lectotype was rendered obsolete when neotype fossils were approved for the genus in 2003. [4]
Sankar Chatterjee later described more comprehensive remains from the Lower Maleri Formation, as well as one nearly complete skull form the Tiki Formation that he also assigned to Parasuchus hislopi. Two complete and articulated skeletons that include complete skulls were collected from the Lower Maleri Formation in the vicinity of the Mutapyram village of Adilabad district. Both individuals were roughly 8 ft (2.4 m) in length, lying side by side. The left individual, ISI R 42, is perfectly preserved, and was designated as the neotype of the species. [4] The right individual, ISI R 43, is nearly complete and only missing part of the snout. [1] [3]
Two articulated and almost complete skeletons of Malerisaurus robinsonae (both designated as ISIR 150), an azendohsaurid archosauromorph, were found as presumable gastric contents of these two skeletons. [5] From the same locality as the neotype, three isolated conjoined basioccipital/basisphenoids (ISI R 45-47) were also recovered. A couple of miles north of that locality, near the Venkatapur village, two more excellently preserved skulls were found. ISI R 160 represents an isolated but nearly complete skull, while ISI R 161 represents partial skull and articulated postcranial remains. [1] Finally, a skull recovered from the Tikisuchus holotype site of the Tiki Formation (Gondwana Group), [6] about 4 miles west of Tiki village of Shadol District, Madhya Pradesh, is missing only the end of the snout and the squamosal. As the lectotype of the genus, it was found nearby Paradapedon remains. [1] Both formations date to the late Carnian to early Norian stage of the Late Triassic period, about 222.5-212 million years ago. [6] [7]
The two articulated skeletons, ISI R 42-43, alone represent the most complete phytosaurs known to date. Additionally, Parasuchus has been found as one of the most primitive phytosaurs in phylogenetic analyses of Phytosauria, making it very significant for understanding the origin of the Phytosauria. [2] Some studies, like Chatterjee (1978) and Lucas et al. (2007), synonymized Parasuchus with another basal phytosaur, Paleorhinus . If this is the case, the name Parasuchus would have precedence over Paleorhinus because Paleorhinus was named in 1904, nineteen years after Parasuchus was named. A phylogenetic analysis conducted by Kammerer et al. (2016) confirmed that Parasuchus hislopi is nested within the least inclusive clade containing the species Paleorhinus bransoni and Paleorhinus angustifrons. The authors thus considered the genus Parasuchus to be a senior synonym of the genus Paleorhinus (as well as Arganarhinus ), and referred the species Paleorhinus bransoni, Paleorhinus angustifrons and Arganarhinus magnoculus to the genus Parasuchus. [3] The cladogram below follows Kammerer et al. (2016). [3]
Parasuchidae |
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Parasuchus was first described and named by Richard Lydekker in 1885 and the type species is Parasuchus hislopi. The generic name is derived from the Greek para/παρα meaning "beside" or "near" and suchus from the Greek souchos, which refers to the Egyptian crocodile-headed god Sobek. The specific name, hislopi, honors Hislop who drawn attention in 1854 to the red clays near Maleri village in which the holotype (and later the neotype and other specimens) of P. hislopi was recovered. [8]
A holotype is a single physical example of an organism used when the species was formally described. It is either the single such physical example or one of several examples, but explicitly designated as the holotype. Under the International Code of Zoological Nomenclature (ICZN), a holotype is one of several kinds of name-bearing types. In the International Code of Nomenclature for algae, fungi, and plants (ICN) and ICZN, the definitions of types are similar in intent but not identical in terminology or underlying concept.
Phytosaurs are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. and are sometimes referred to as parasuchians. Phytosauria, Parasuchia, Parasuchidae, and Phytosauridae have often been considered equivalent groupings containing the same species. Some recent studies have offered a more nuanced approach, defining Parasuchidae and Phytosauridae as nested clades within Phytosauria as a whole. Phytosaurs were long-snouted and heavily armoured, bearing a remarkable resemblance to modern crocodilians in size, appearance, and lifestyle, as an example of convergence or parallel evolution. The name "phytosaur" means "plant reptile", as the first fossils of phytosaurs were mistakenly thought to belong to plant eaters.
In binomial nomenclature, a nomen dubium is a scientific name that is of unknown or doubtful application.
Paleorhinus is an extinct genus of widespread basal phytosaur known from the Late Triassic. The genus was named in 1904 based on the type species Paleorhinus bransoni, which is known from Wyoming and Texas in the United States. Another valid species, Paleorhinus angustifrons from Bavaria, Germany, is also commonly referred to the genus. Paleorhinus had a length of about 2.5 meters.
Rutiodon is an extinct genus of mystriosuchine phytosaurs from the Late Triassic of the eastern United States. The type species of Rutiodon, Rutiodon carolinensis, encompasses a large number of skulls and assorted postcranial fossils discovered in the Cumnock Formation of North Carolina. Fossils referable to the species are also known from Pennsylvania, New Jersey, and Virginia. Rutiodon carolinensis is the most well-described species of phytosaur in eastern North America, though its validity as a natural taxon has been questioned. Some paleontologists also recognize a larger and more robust species, Rutiodon manhattanensis, which is known from teeth and postcranial fossils from New Jersey and Pennsylvania.
Phytosaurus is a dubious genus of extinct parasuchid phytosaur found in an outcrop of the Keuper in Germany. Phytosaurus was the first phytosaur to be described, being done so by Georg Friedrich von Jaeger in 1828. The type species is P. cylindricodon and a second species, P. cubicodon, is also known.
Doswellia is an extinct genus of archosauriform from the Late Triassic of North America. It is the most notable member of the family Doswelliidae, related to the proterochampsids. Doswellia was a low and heavily built carnivore which lived during the Carnian stage of the Late Triassic. It possesses many unusual features including a wide, flattened head with narrow jaws and a box-like rib cage surrounded by many rows of bony plates. The type species Doswellia kaltenbachi was named in 1980 from fossils found within the Vinita member of the Doswell Formation in Virginia. The formation, which is found in the Taylorsville Basin, is part of the larger Newark Supergroup. Doswellia is named after Doswell, the town from which much of the taxon's remains have been found. A second species, D. sixmilensis, was described in 2012 from the Bluewater Creek Formation of the Chinle Group in New Mexico; however, this species was subsequently transferred to a separate doswelliid genus, Rugarhynchos. Bonafide Doswellia kaltenbachi fossils are also known from the Chinle Formation of Arizona.
Redondasaurus is an extinct genus or subgenus of phytosaur from the Late Triassic of the southwestern United States. It was named by Hunt & Lucas in 1993, and contains two species, R. gregorii and R. bermani. It is the youngest and most evolutionarily-advanced of the phytosaurs.
Mystriosuchus is an extinct genus of phytosaur that lived in the Late Triassic in Europe and Greenland. It was first named by Eberhard Fraas in 1896, and includes four species: M. planirostris, M. westphali, M. steinbergeri, and M. alleroq.
Malerisaurus is an extinct genus of archosauromorph known from Andhra Pradesh of India and Texas of the USA.
Mystriosuchini, historically known as Pseudopalatinae, is an extinct tribe of derived phytosaurs in the clade Leptosuchomorpha. As with all other phytosaurs, mystriosuchins lived during Late Triassic. The name is derived from the genus Mystriosuchus.
Ebrachosuchus is an extinct genus of basal phytosaur known from the Late Triassic of Bavaria, southern Germany. It is known only from the holotype BSPG 1931 X 501, a complete skull missing both mandibles. It was collected at Ebrach Quarry, bed number 9 from the late Carnian-aged Blasensandstein Member of the Hassberge Formation. It was first named by Oskar Kuhn in 1936 and the type species is Ebrachosuchus neukami.
Machaeroprosopus is an extinct genus of mystriosuchin leptosuchomorph phytosaur from the Late Triassic of the southwestern United States. M. validus, once thought to be the type species of Machaeroprosopus, was named in 1916 on the basis of three complete skulls from Chinle Formation, Arizona. The skulls have been lost since the 1950s, and a line drawing in the original 1916 description is the only visual record of the specimen. Another species, M. andersoni, was named in 1922 from New Mexico, and the species M. adamanensis, M. gregorii, M. lithodendrorum, M. tenuis, and M. zunii were named in 1930. Most species have been reassigned to the genera Smilosuchus, Rutiodon, or Phytosaurus. Until recently, M. validus was considered to be the only species that has not been reassigned. Thus, Machaeroprosopus was considered to be a nomen dubium or "doubtful name" because of the lack of diagnostic specimens that can support its distinction from other phytosaur genera. However, a taxonomic revision of Machaeroprosopus, conducted by Parker et al. in 2013, revealed that UW 3807, the holotype of M. validus, is not the holotype of Machaeroprosopus, while the species Machaeroprosopus buceros, Machaeroprosopus being a replacement name, with a fixed type species, for Metarhinus, is the combinatio nova of the type species of the genu: Belodon buceros. Therefore, the name Pseudopalatus must be considered a junior synonym of Machaeroprosopus, and all species of the former must be reassigned to the latter. This revised taxonomy was already accepted in several studies, including Stocker and Butler (2013). Stocker and Butler (2013) also treated M. andersoni as a valid species, and not a junior synonym of Machaeroprosopus buceros as was previously suggested by Long and Murry (1995).
The Tiki Formation is a Late Triassic geologic formation in Madhya Pradesh, northern India. Dinosaur remains are among the fossils that have been recovered from the formation, although none have yet been referred to a specific genus. Phytosaur remains attributable to the genus Volcanosuchus have also been found in the Tiki Formation.
Tikisuchus is an extinct genus of rauisuchid archosauromorph. It is known from the Late Triassic Tiki Formation in the Shahdol District of central India and was the first rauisuchid to have been found in Asia. The horizon from which remains have been found is Carnian in age. The type species is T. romeri, named in honor of American paleontologist Alfred Romer. Romer was present at the Tiki locality during the excavation of the fossil, but died before the description of the genus in 1987. Tikisuchus is known only from one specimen, called ISI R 305, which consists of the skull and some postcranial elements of a young individual.
Nambalia is a genus of basal sauropodomorph dinosaur. It lived during the Late Triassic period in what is now Telangana, central India. It is known from the holotype ISI R273, parts 1–3, partially articulated postcranial material and from the paratypes ISI R273, parts 4-29, including partial postcrania of at least two individuals of different sizes found closely associated and one of them is nearly the same size as the holotype.
Francosuchus is a dubious genus of probably basal phytosaur known from the Late Triassic of Bavaria, southern Germany. It was named by Oskar Kuhn in 1933 and the type species is Francosuchus broilii. In the same article Kuhn also named a second species Francosuchus latus. Both species were known solely from their holotypes, two partial skulls that were housed at the Bavarian State Collection for Palaeontogy and Geology. Both specimens were collected at Ebrach Quarry, bed number 13 from the late Carnian-aged Blasensandstein Member of the Hassberge Formation. As the holotypes were destroyed during World War II and poorly documented, Francosuchus and its species are usually considered to be nomina dubia.
Mesorhinosuchus is an extinct genus of basal phytosaur possibly known from the Early Triassic of Saxony-Anhalt, central-eastern Germany. It was first named by Otto Jaekel in 1910 and the type species is Mesorhinus fraasi. The generic name Mesorhinus was preoccupied by Mesorhinus piramydatus Ameghino, 1885, a macraucheniid mammal, which is now considered to be a junior synonym of Oxyodontherium. Thus, an alternative generic name, Mesorhinosuchus, was proposed by Oskar Kuhn in 1961. The genus is occasionally misspelled as Mesorhinosaurus, while Stocker and Butler (2013) recently misspelled its original generic name as Mesosuchus.
Parasuchidae is a clade of phytosaurs more derived than Diandongosuchus, a basal phytosaur. It encompasses nearly all phytosaurs, include early Parasuchus-grade forms as well as a more restricted clade of more specialized phytosaurs. This more restricted clade is traditionally known as the family Phytosauridae and more recently as the subfamily Mystriosuchinae.
Wannia is an extinct genus of basal phytosaur reptile known from the Late Triassic of Texas, southern United States. It contains a single species, Wannia scurriensis, which is known from a single specimen. This species was originally named as a species referred to Paleorhinus and later was considered as a possible junior synonym of Paleorhinus bransoni. However its re-description revealed five autapomorphies, and a phylogenetic position as the most basal known phytosaur, justifying the erection of a new generic name for the species.