Parasuchus Hislopi
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''Parasuchus'' 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 of
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
phytosaur known from the Late Triassic (late Carnian to early
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
stage) of Andhra Pradesh and Madhya Pradesh, India. At its most restricted definition, ''Parasuchus'' contains a single species, ''Parasuchus hislopi''. ''Parasuchus hislopi'' is one of several species belonging to a basal grade of phytosaurs, typified by the genus ''
Paleorhinus ''Paleorhinus'' (Greek: ''"Old Nose"'') is an extinct genus of widespread basal phytosaur known from the Late Triassic (late Carnian stage). The genus was named in 1904 based on the type species ''Paleorhinus bransoni'', which is known from Wyom ...
''. 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 270px, '' valid name. Priority is a fundamental principle of modern botanical nomenclature and zoological nomenclature. Essentially, it is the principle of recognising the first valid application of a name to a plant or animal. There are two asp ...
.


History

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 In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
'' 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 The base of skull, also known as the cranial base or the cranial floor, is the most inferior area of the skull. It is composed of the endocranium and the lower parts of the calvaria. Structure Structures found at the base of the skull are for ...
mixed with the partial snout of a phytosaur, scutes and some teeth.
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
(1940) identified the basicranium as belonging to ''
Paradapedon huxleyi ''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery and namin ...
'' (now known as '' Hyperodapedon huxleyi'') and the phytosaurian material to a newly named species, "aff." ''
Brachysuchus ''Brachysuchus'' (meaning "wide crocodile") is an extinct genus of phytosaur known from the late Triassic period (Carnian stage) of Dockum Group in Texas, United States. It is known from the holotype UMMP 10336 is composed of a skull, lower jaws ...
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 ''Brachysuchus'' (meaning "wide crocodile") is an extinct genus of phytosaur known from the late Triassic period (Carnian stage) of Dockum Group in Texas, United States. It is known from the holotype UMMP 10336 is composed of a skull, lower jaws ...
'' (which is closely related to or synonymous with ''
Angistorhinus ''Angistorhinus'' (meaning "narrow snout" or "hook snout") is an extinct genus of phytosaur known from the Late Triassic period of Texas and Wyoming, United States. It was first named by Mehl in 1913 and the type species is ''Angistorhinus grand ...
''), or to '' Phytosaurus'' (a
dubious name In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
, probably the senior synonym of ''
Nicrosaurus ''Nicrosaurus'' (/nɛkroʊˈsɔrəs/) is an extinct genus of phytosaur reptile existing during the Late Triassic period. Although it looked like a crocodile (and probably lived like the more terrestrial crocodylomorphs), it was not closely rela ...
''). 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. To avoid additional confusion, the nondiagnostic holotype of ''P. hislopi'' was replaced by a neotype (ISI R 42) with approval from the
ICZN The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the I ...
(Opinion 2045) following the application of Chatterjee (2001).


Fossil material

The partial
premaxillary The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
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 The Pranhita–Godavari Basin is a northwest–southeast striking geological structural basin (rift basin) in eastern India. The basin contains up to 7 kilometres of sedimentary strata of late Carboniferous/Early Permian to Cretaceous age. The b ...
), near the Maleri village of Adilabad district, Andhra Pradesh. The lectotype was rendered obsolete when neotype fossils were approved for the genus in 2003. Sankar Chatterjee later described more comprehensive remains from the Lower Maleri Formation, as well as one nearly complete skull form the
Tiki Formation The Tiki Formation is a Late Triassic (Carnian to Norian) geologic formation in Madhya Pradesh, northern India.skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s 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 in length, lying side by side. The left individual,
ISI ISI or Isi may refer to: Organizations * Intercollegiate Studies Institute, a classical conservative organization focusing on college students * Ice Skating Institute, a trade association for ice rinks * Indian Standards Institute, former name of ...
 R 42, is perfectly preserved, and was designated as the neotype of the species. The right individual, ISI R 43, is nearly complete and only missing part of the snout. Two articulated and almost complete
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s of '' Malerisaurus robinsonae'' (both designated as ISIR 150), an azendohsaurid archosauromorph, were found as presumable gastric contents of these two skeletons. 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. Finally, a skull recovered from the '' Tikisuchus'' holotype site of the Tiki Formation ( Gondwana Group), 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 ''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Late Triassic period ( Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery and na ...
'' remains. Both formations date to the late Carnian to early
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
stage of the Late Triassic period, about 222.5-212  million years ago.


Classification

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. Some studies, like Chatterjee (1978) and Lucas ''et al.'' (2007), synonymized ''Parasuchus'' with another
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
phytosaur, ''
Paleorhinus ''Paleorhinus'' (Greek: ''"Old Nose"'') is an extinct genus of widespread basal phytosaur known from the Late Triassic (late Carnian stage). The genus was named in 1904 based on the type species ''Paleorhinus bransoni'', which is known from Wyom ...
''. 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 ''Arganarhinus'' (meaning "Argana (Morocco) snout") is an extinct genus of phytosaur known from the late Triassic period (geology), period (Middle Carnian stage) of Argana Basin in Morocco. It is known from a skull which is housed at the Muséum ...
''), and referred the species ''Paleorhinus bransoni'', ''Paleorhinus angustifrons'' and ''Arganarhinus magnoculus'' to the genus ''Parasuchus''. The cladogram below follows Kammerer ''et al.'' (2016).


Etymology

''Parasuchus'' was first described and named by Richard Lydekker in
1885 Events January–March * January 3– 4 – Sino-French War – Battle of Núi Bop: French troops under General Oscar de Négrier defeat a numerically superior Qing Chinese force, in northern Vietnam. * January 4 – ...
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 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 ...
, ''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.


References

{{Taxonbar, from=Q251814 Phytosaurs Prehistoric reptile genera Late Triassic reptiles of Asia Triassic India Fossils of India Taxa named by Richard Lydekker Fossil taxa described in 1885