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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 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 basal phytosaur 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 ...
(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 Andhra Pradesh (, abbr. AP) is a state in the south-eastern coastal region of India. It is the seventh-largest state by area covering an area of and tenth-most populous state with 49,386,799 inhabitants. It is bordered by Telangana to the ...
and
Madhya Pradesh Madhya Pradesh (, ; meaning 'central province') is a state in central India. Its capital is Bhopal, and the largest city is Indore, with Jabalpur, Ujjain, Gwalior, Sagar, and Rewa being the other major cities. Madhya Pradesh is the seco ...
,
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
. 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 (phylogenetics), basal phytosaur known from the Late Triassic (late Carnian stage). The genus was named in 1904 based on the type species ''Paleorhinus bransoni'', wh ...
''. 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 Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The storie ...
(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 Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
(1885) formally described and named ''P. hislopi'', and proposed the family name
Parasuchidae 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 ...
. However, Lydekker's description was based on a chimeric syntype, combining fossils from multiple unrelated reptiles: a
rhynchosauria Rhynchosaurs are a group of extinct herbivorous Triassic archosauromorph reptiles, belonging to the order Rhynchosauria. Members of the group are distinguished by their triangular skulls and elongated, beak like premaxillary bones. Rhynchosaurs ...
n
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 ''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Triassic, Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery ...
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 ''Phytosaurus'' (meaning "plant lizard") is a nomen dubium, dubious genus of Extinction, extinct Parasuchidae, parasuchid phytosaur found in an outcrop of the Keuper (likely the Exter Formation) in Germany. ''Phytosaurus'' was the first phytosaur ...
maleriensis'' while Gregory (1962) considered the material undiagnostic.
Sankar Chatterjee Sankar Chatterjee (born May 28, 1943) is a paleontologist, and is the Paul W. Horn Professor of Geosciences at Texas Tech University and Curator of Paleontology at the Museum of Texas Tech University. He earned his Ph. D. from the University of ...
(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 ''Phytosaurus'' (meaning "plant lizard") is a nomen dubium, dubious genus of Extinction, extinct Parasuchidae, parasuchid phytosaur found in an outcrop of the Keuper (likely the Exter Formation) in Germany. ''Phytosaurus'' was the first phytosaur ...
'' (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 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 ...
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 The Lower Maleri Formation is a sedimentary geological formation, rock formation found in Andhra Pradesh and Telangana, India. It is the lowermost member of the Pranhita–Godavari Basin. It is of late Carnian to early Norian age (Upper Triassi ...
(
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 The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
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 ''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 remain ...
'' 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'' 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 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 ...
period, about 222.5-212 
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (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 phytosaur, ''
Paleorhinus ''Paleorhinus'' (Greek: ''"Old Nose"'') is an extinct genus of widespread basal (phylogenetics), basal phytosaur known from the Late Triassic (late Carnian stage). The genus was named in 1904 based on the type species ''Paleorhinus bransoni'', wh ...
''. 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 Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe 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 In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
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