Pravusuchus

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Pravusuchus
Temporal range: Late Triassic, 228–209  Ma
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Phytosauria
Family: Parasuchidae
Clade: Leptosuchomorpha
Genus: Pravusuchus
Stocker, 2010
Type species
Pravusuchus hortus
Stocker, 2010

Pravusuchus is an extinct genus of leptosuchomorph parasuchid phytosaur known from the Late Triassic (Norian stage) of Arizona, United States. It contains a single species, Pravusuchus hortus, which is known from three specimens. These specimens were previously referred to Smilosuchus or to Leptosuchus , but Pravusuchus's autapomorphy, its phylogenetic position as well as a trait shared with mystriosuchins, justified the erection of a new taxon for the material. [1]

Contents

Discovery and naming

Pravusuchus was first described and named by Michelle R. Stocker in 2010 and the type species is Pravusuchus hortus. The generic name is derived from Latin, pravus, "evil" or "wicked", and Greek, souchus, for the Egyptian crocodile-headed god Sobek. The specific name, hortus, is the Latin word for park or grounds. The name refers to Devil's Playground, the locality in Petrified Forest National Park from which all specimens of this taxon were collected. Pravusuchus is known from the holotype AMNH FR 30646, mostly complete but partially disarticulated skull, and from the referred specimens PEFO 31218, a partial skull, and PEFO 34239, partial skull, partial mandible, and possible partial postcrania. Although the holotype is slightly crushed dorsoventrally, it is not as flattened as PEFO 31218. All specimens referable to Pravusuchus were collected from the Sonsela Sandstone Bed (later referred to as Jasper / Rainbow Forest Bed [2] ) of the Norian-aged Sonsela Member, Chinle Formation from Navajo County, Arizona. PEFO 31218 and PEFO 34239 were found in the same stratigraphic horizon just west of Colbert's 1946 locality where AMNH FR 30646 was collected, and only 2–3 m higher stratigraphically than the holotype. [1] Other phytosaur specimens from the Sonsela Member occur in other beds, stratigraphically lower (e.g. Protome batalaria , Smilosuchus adamanensis [1] and S. lithodendrorum ) or higher (e.g. Machaeroprosopus jablonskiae and M. pristinus ) then the white Rainbow Forest sandstone. [2]

Description

Stocker (2010) diagnosed Pravusuchus hortus by a single unambiguous autapomorphy (unique trait) and additionally by a unique combination of characters. Unlike all other phytosaurs, the lateral rim of its external naris (i.e. nostril) is formed by the "septomaxilla", and not by the nasal bone. The element in phytosaur skulls anterior to the external nares, located between the nasal, maxilla and premaxilla, has traditionally been referred to as the septomaxilla, however it probably is not homologous to the septomaxillae of squamates and synapsids. [1]

The antorbital fossa is absent in Pravusuchus, as in Pseudopalatus, Smilosuchus gregorii and S. adamanensis, however, unlike Pseudopalatus its interpremaxillary fossa (a depression on the premaxilla) is broad and rounded. In Pravusuchus, alveolar ridges visible in lateral view (from the sides), unlike the condition in S. gregorii and Pseudopalatus. Its rostrum is partially crested, as in Leptosuchus crosbiensis, L. studeri and S. adamanensis, [1] but it lacks a premaxillary crest, as also seen in Protome, Smilosuchus and L. crosbiensis. [2] Pravusuchus shares with Pseudopalatus, L. crosbiensis and L. studeri the presence of a long posterior process of the squamosal, unlike the condition seen in S. gregorii and S. adamanensis. [1] However, in contrast to the condition in Pseudopalatus, it is also greatly dorsoventrally expanded and rounded posteriorly as in Rutiodon, Protome, Leptosuchus, Smilosuchus and "Phytosaurus" doughtyi. As in "Machaeroprosopus" zunii, Protome, Smilosuchus and Pseudopalatus, Pravusuchus has basitubera, in front of the attachment point between the skull and the neck, that are connected and form a sharp ridge along their anterior border. [2] Its supratemporal fenestrae (upper temporal openings) partially depressed as in Rutiodon, Leptosuchus and S. gregorii, in contrast to the condition in Pseudopalatus, and are mostly visible in dorsal view as in Smilosuchus. In contrast to all other phytosaurs, Pravusuchus shares the presence of a subsidiary opisthotic process of the squamosal with Mystriosuchini, a possible synapomorphy. [1]

Phylogeny

To test the evolutionary relationships of Pravusuchus, Stocker (2010) performed the most inclusive phylogenetic analysis of Phytosauria up to that time. Pravusuchus was coded in the analysis using solely its holotype, as both referred specimen had identical codings for applicable characters. The analysis placed Pravusuchus outside the clade containing species of Mystriosuchus and Pseudopalatus (now Machaeroprosopus ), suggesting that it is the sister taxon of Mystriosuchini. [1] However, to confirm its exclusion from Mystriosuchini, Nicrosaurus kapffi must be included in the analysis, as Mystriosuchini is a node-based taxon and Nicrosaurus usually occupies the most basal position in it. [3] Furthermore, Pravusuchus was found to occupy a more derived position then Leptosuchus and even Smilosuchus . This justified the erection of a new genus and species for Pravusuchus, as its holotype was previously referred to these genera. [1] Below is a cladogram showing the phylogenetic relationships of Pravusuchus from Stocker (2010) which was also recovered in Stocker (2012): [2]

Phytosauria  

"Paleorhinus" scurriensis

Paleorhinus bransoni

"Paleorhinus" sawini

Phytosauridae

Brachysuchus megalodon

Angistorhinus

Rutiodon carolinensis

"Machaeroprosopus" zunii

Protome batalaria

Leptosuchomorpha

"Phytosaurus" doughtyi

TMM 31173-120

Leptosuchus crosbiensis

Leptosuchus studeri

Smilosuchus lithodendrorum

Smilosuchus adamanensis

Smilosuchus gregorii

Pravusuchus hortus

Pseudopalatinae

Pseudopalatus mccauleyi

Mystriosuchus westphali

Pseudopalatus pristinus

Related Research Articles

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.

<span class="mw-page-title-main">Chinle Formation</span> Geological formation in the western US

The Chinle Formation is an Upper Triassic continental geological formation of fluvial, lacustrine, and palustrine to eolian deposits spread across the U.S. states of Nevada, Utah, northern Arizona, western New Mexico, and western Colorado. In New Mexico, it is often raised to the status of a geological group, the Chinle Group. Some authors have controversially considered the Chinle to be synonymous to the Dockum Group of eastern Colorado and New Mexico, western Texas, the Oklahoma panhandle, and southwestern Kansas. The Chinle Formation is part of the Colorado Plateau, Basin and Range, and the southern section of the Interior Plains. A probable separate depositional basin within the Chinle is found in northwestern Colorado and northeastern Utah. The southern portion of the Chinle reaches a maximum thickness of a little over 520 meters (1,710 ft). Typically, the Chinle rests unconformably on the Moenkopi Formation.

<i>Paleorhinus</i> Extinct genus of reptiles

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.

<i>Rutiodon</i> Extinct genus of reptiles

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.

<i>Nicrosaurus</i> Extinct genus of reptiles

Nicrosaurus (/nɪkroʊˈsɔrəs/) is an extinct genus of phytosaur reptile existing during the Late Triassic period. Although it looked like a crocodile, it was not closely related to these creatures, instead being an example of parallel evolution. The main difference between Nicrosaurus and modern crocodiles is the position of the nostrils – Nicrosaurus's nostrils, or external nares, were placed directly in front of the forehead, whereas in crocodiles, the nostrils are positioned on the end of the snout. A 2013 study has also found that ilium of Nicrosaurus is quite distinctive from all other phytosaurs.

<i>Acaenasuchus</i> Genus of reptiles

Acaenasuchus is an extinct genus of pseudosuchian, endemic to what would be presently be known as Arizona during the Late Triassic, specifically during the Carnian and Norian stages of the Triassic. Acaenasuchus had a stratigraphic range of approximately 11.5 million years. Acaenasuchus is further categorized as one of the type fauna that belong to the Adamanian LVF, based on the fauna of the Blue Mesa Member of the Chinle Petrified Forest Formation of Arizona, where Acaenasuchus was initially discovered.

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.

<i>Angistorhinus</i> Extinct genus of reptiles

Angistorhinus 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 grandis. Other species from Texas and Wyoming, A. alticephalus, A. gracilis and A. maximus, are cospecific with the type species. Angistorhinus is known from the holotype UC 631, partial skull and lower jaws recovered from the Popo Agie Formation, Chugwater Group, Wyoming and from the associated paratype UM 531, a partial skull, TMM 31098-1, skull and lower jaws and ROM 7977, partial skull and lower jaws, recovered from the 'Pre-Tecovas Horizon' in the Dockum Group, Texas. A possible second species, A. talainti is known from the Triassic of Morocco. In 1995, Long and Murry created the new combination, Angistorhinus megalodon by synonymy for Brachysuchus. Hungerbühler and Sues (2001) hypothesised that Angistorhinus is a junior synonym of Rutiodon. However, in 2010 Michelle R. Stocker retained the validity of Brachysuchus and of A. grandis.

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.

<i>Smilosuchus</i> Extinct genus of reptiles

Smilosuchus is an extinct genus of leptosuchomorph parasuchid from the Late Triassic of North America.

<span class="mw-page-title-main">Mystriosuchini</span> Extinct tribe of reptiles

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.

<i>Leptosuchus</i> Extinct genus of reptiles

Leptosuchus is an extinct genus of leptosuchomorph phytosaur with a complex taxonomical history. Fossils have been found from the Dockum Group and lower Chinle Formation outcropping in Texas, New Mexico, and Arizona, USA, and date back to the Carnian stage of the Late Triassic.

<i>Machaeroprosopus</i> Extinct genus of reptiles

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).

<i>Brachysuchus</i> Extinct genus of reptiles

Brachysuchus is an extinct genus of phytosaur known from the late Triassic period of Dockum Group in Texas, United States. It is known from the holotype UMMP 10336 is composed of a skull, lower jaws and partial postcranium and from the associated paratype UMMP 14366, nearly complete skull, recovered from the 'Pre-Tecovas Horizon' in the Dockum Group. It was first named by Case in 1929 and the type species is Brachysuchus megalodon. Its closest relative was Angistorhinus. However, its rostral crest was much smaller than that of Angistorhinus, and the rostrum as a whole is shorter and thicker.

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.

<i>Protome</i> (genus) Extinct genus of reptiles

Protome is an extinct genus of parasuchid phytosaur from the Late Triassic of Arizona, represented by a single species, Protome batalaria. It is known from a single holotype incomplete, partially disarticulated skull and left lower jaw called PEFO 34034 from the Upper Lot's Wife beds, Sonsela Member of the Chinle Formation in Petrified Forest National Park. The skull was discovered in 2004 by Michelle Stocker and Bill Parker and was described by them as a specimen of Smilosuchus adamanensis. It was placed in the new genus Protome in 2012. The genus name Protome is the Greek word for an animal's face. The specific name batalaria is the Latin word for battleship, which is a reference to Battleship NW, the locality within Petrified Forest where the skull was found.

<span class="mw-page-title-main">Leptosuchomorpha</span> Extinct clade of reptiles

Leptosuchomorpha is a clade of phytosaurs. It is a node-based taxon defined as the last common ancestor of Leptosuchus studeri and Pseudopalatus pristinus and all of its descendants.

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.

<i>Vivaron</i> Extinct genus of reptiles

Vivaron is a genus of rauisuchid known from the Late Triassic Chinle Formation in New Mexico. It is the second rauisuchid known from the southwestern United States, and it highlights the wide biogeographic range similar rauisuchid taxa occupied during the Late Triassic across Pangaea, despite the varied faunal assemblages at different latitudes.

References

  1. 1 2 3 4 5 6 7 8 9 Stocker, M. R. (2010). "A new taxon of phytosaur (Archosauria: Pseudosuchia) from the Late Triassic (Norian) Sonsela Member (Chinle Formation) in Arizona, and a critical reevaluation of Leptosuchus Case, 1922". Palaeontology. 53 (5): 997–1022. doi: 10.1111/j.1475-4983.2010.00983.x .
  2. 1 2 3 4 5 Stocker, M. R. (2012). "A new phytosaur (Archosauriformes, Phytosauria) from the Lot's Wife beds (Sonsela Member) within the Chinle Formation (Upper Triassic) of Petrified Forest National Park, Arizona". Journal of Vertebrate Paleontology. 32 (3): 573–586. doi:10.1080/02724634.2012.649815. S2CID   129527672.
  3. Parker, W. G.; Irmis, R. B. (2006). "A new species of the Late Triassic phytosaur Pseudopalatus (Archosauria: Pseudosuchia) from Petrified Forest National Park, Arizona" (PDF). Museum of Northern Arizona Bulletin. 62: 126–143.