Paleorhinus Temporal range: Late Triassic, | |
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Skull of the P. angustifrons holotype with CT scan | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Order: | † Phytosauria |
Genus: | † Paleorhinus Williston, 1904 |
Type species | |
†Paleorhinus bransoni Williston, 1904 | |
Species | |
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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 Wyoming and Texas in the United States. Another valid species, Paleorhinus angustifrons from Bavaria, Germany, is also commonly referred to the genus. [1] [2] Paleorhinus had a length of about 2.5 meters (8.2 feet).
Paleorhinus has had a complicated taxonomic history involving frequent synonymy between diagnostic and undiagnostic material. This is mainly due to the fact that it is a quintessential basal phytosaur, mostly distinguished by a lack of specializations rather than unique traits. Historically, it was common practice to lump all basal phytosaurs into only one or two genera, rendering those genera paraphyletic evolutionary grades ancestral to later phytosaurs. More recently, these grades have been broken up into multiple genera. [2] Arganarhinus magnoculus (from Morocco) and Wannia scurriensis (from Texas) [3] were two phytosaur genera originally considered species of Paleorhinus. "Paleorhinus" sawini (from Texas) and "Paleorhinus" parvus (from Wyoming) are two more phytosaur species informally referred to Paleorhinus, though likely closer to more advanced phytosaurs. [2] [4]
P. angustifrons was originally considered a species of Francosuchus (a dubious phytosaur genus from the same area), and has also been compared to Ebrachosuchus neukami , another Paleorhinus-like Bavarian phytosaur. "Paleorhinus" cf. arenaceus (formerly " Zanclodon " arenaceus), fragmentary phytosaur remains from Poland, may represent Paleorhinus fossils.
Parasuchus hislopi , a basal phytosaur species named in 1885 from fossils discovered in India, was often lumped into Paleorhinus or considered a dubious chimera of phytosaur and rhynchosaur fossils. More complete neotype fossils for Parasuchus hislopi have helped to re-establish it as a valid genus and species. [5] Some phylogenetic analyses suggest that Parasuchus hislopi forms a clade with Paleorhinus bransoni and angustifrons. If this is the case, the valid species of Paleorhinus may instead be considered species of Parasuchus, since that genus name has priority over Paleorhinus. [6] [4]
P. bransoni is the type species of Paleorhinus. It was first described and named by Samuel Wendell Williston in 1904 on the basis of the holotype FMNH UC 632, a complete but fragmentary skull that has been extensively reconstructed with plaster. It was collected at Squaw Creek of Fremont County, Wyoming, from the Carnian-aged Popo Agie Formation of the Chugwater Group. The holotype was thoroughly described by Lees (1907). [1] [7] Hunt and Lucas (1991) referred PPM P217 to P. bransoni. It is an incomplete skull that was collected at Palo Duro Canyon, Randall County, west Texas, from the Carnian-aged Camp Springs Member of the Tecovas Formation. [8] Later, Long and Murry (1995) referred additional material, including isolated postcranial remains, to P. bransoni from the Carnian-aged Cooper Canyon Formation of the Dockum Group, Howard County, Texas. TMM 31025-172, a complete skull, was collected at Otis Chalk Quarry 1 (also known as SMU 122), TMM 31100-8, 101, 175, 418, 419, 453 were collected at Otis Chalk Quarry 3, and TMM 31185-11, 38 were collected at Otis Chalk Quarry 3A. [9]
P. angustifrons was first described and named by Oskar Kuhn in 1936 as a species of Francosuchus . P. angustifrons is known exclusively from the holotype BSPG 1931 X 502 a partial skull lacking the rostrum and mandibles. It was collected at Ebrach Quarry, bed number 9 of Bavaria, southern Germany, from the late Carnian-aged Blasensandstein Member of the Hassberge Formation. [10]
Hunt and Lucas (1991) mistakenly referred to F. angustifrons as Ebrachosuchus angustifrons, and considered it and the other two Francosuchus species, F. broilii and F. latus , to be synonyms of Paleorhinus neukami . [8] More recently, P. neukami was found to be more closely related to Mystriosuchinae than to Paleorhinus and thus the genus Ebrachosuchus was re-validated, while F. angustifrons was reassigned as P. angustifrons as it shares unique synapomorphies with P. bransoni and was found to be its sister taxon. The other two Francosuchus species, were not referred to P. angustifrons because they were found to be nomina dubia as their holotypes were destroyed during World War II. [2]
Dzik & Sulej (2007) assigned several skulls, partial articulated postcranial skeletons, and numerous isolated phytosaur bones in various ontogenetic stages from Krasiejów, Poland to Paleorhinus cf. arenaceus. They found some similarities between the material and P. bransoni and also Ebrachosuchus neukami or F. angustifrons (mistakenly referred to as E. broili), as well as some apparent differences with Parasuchus . The shape of the external mandibular fenestra of the material resembled that of "Zanclodon" arenaceus but it is also shared with the proterochampsid Chanaresuchus bonapartei . [11] "Z." arenaceus was suggested to represent the oldest reliably dated phytosaur, and was reassigned to various phytosaur species over the years, including Belodon , Mystriosuchus and Phytosaurus . Although Hungerbühler (2001) redescribed "Z." arenaceus as not belonging to Phytosauria and referred it to Archosauria incertae sedis , [12] Dzik & Sulej (2007) noted that its holotype "does not differ significantly from corresponding parts of the juvenile Krasiejów Paleorhinus, which is clearly a phytosaur". Furthermore, as the Feuerbacher Heide Schilfsandstein, from which "Z." arenaceus was collected, and Krasiejów share species of Metoposaurus , it might be possible that they also share the same species of phytosaur. Even though they agreed that due to the very fragmentary nature of "Z." arenaceus holotype the Krasiejów Paleorhinus can't be referred to it, they tentatively used the name Paleorhinus cf. arenaceus for the Krasiejów Paleorhinus. [11]
Through the years, various species have been referred to as Paleorhinus. The species that are no longer considered to belong to Paleorhinus, are summarised in the list below:
The following cladogram, from Kammerer et al., 2016, shows the relationships of P. bransoni, P. angustifrons, and "P." sawini to other phytosaurs. The authors placed P. bransoni and P. angustifrons into the genus Parasuchus, owing to their close relationship with Parasuchus hislopi. [6]
Parasuchidae |
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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.
Parasuchus is an extinct genus of basal phytosaur known from the Late Triassic 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. 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 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.
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.
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.
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.
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.
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.
Smilosuchus is an extinct genus of leptosuchomorph parasuchid from the Late Triassic of North America.
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.
Dyoplax is an extinct genus of pseudosuchian archosaur, possibly an erpetosuchid. Fossils have been found from the type locality within the upper Schilfsandstein Formation in Stuttgart, Germany. The holotype specimen was a natural cast of a nearly complete skeleton that lacked only parts of the tail and limb bones.
Ebrachosaurus is an extinct genus of aetosaur. It was named after the town of Ebrach, Germany, near an outcrop of the Blasensandstein Formation where the original fossils have been found. Other Blasensandstein fauna include the temnospondyl Metoposaurus and the phytosaur Francosuchus. The genus has often been considered synonymous with the closely related Stagonolepis. The holotype specimen was lost during World War II, so its relationships within Stagonolepididae remain indeterminant.
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.
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.
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).
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.
Dolerosaurus is an extinct genus of diapsid known from the early Late Triassic upper Lunz Formation of Austria. Dolerosaurus was first named by Richard J. Butler in 2013 and the type species is Francosuchus trauthi.
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.