{{Short description|Extinct genus of reptiles}} {{Automatic taxobox | fossil_range = [[Late Triassic]], {{fossil range|221.5|205.6}} | image = Parasuchus angustifrons.png | image_caption = Skull of the ''P. angustifrons'' holotype with CT scan | taxon = Paleorhinus | authority = [[Samuel Wendell Williston|Williston]], [[1904 in paleontology|1904]] | type_species = {{extinct}}'''''Paleorhinus bransoni''''' | type_species_authority = Williston, 1904 | subdivision_ranks = Species | subdivision = *{{extinct}}'''''P. angustifrons''''' <small>([[Oskar Kuhn|Kuhn]], 1936 [originally ''[[Francosuchus]] angustifrons''])</small> *{{extinct}}'''''P. bransoni''''' <small>Williston, 1904</small> }}
'''''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''''', 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.<ref name="Paleorhinus">{{cite journal | last1 = Williston | first1 = S. W. | year = 1904 | title = Notice of some new reptiles from the Upper Trias of Wyoming | url = https://zenodo.org/record/1431445| journal = Journal of Geology | volume = 12 | issue = 8| pages = 688–697 | doi=10.1086/621190 | bibcode=1904JG.....12..688W| s2cid = 140167568 }}</ref><ref name="StockerButler13">{{Cite journal | last1 = Stocker | first1 = M. R. | last2 = Butler | first2 = R. J. | title = Phytosauria | doi = 10.1144/SP379.5 | journal = Geological Society, London, Special Publications | year = 2013 | volume=379 | issue = 1 | pages=91–117| bibcode = 2013GSLSP.379...91S | s2cid = 219192243 }}</ref> ''Paleorhinus'' had a length of about {{convert|2.5|m|ft|abbr=off|sp=us}}.
''Paleorhinus'' has had a complicated [[Taxonomy|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 [[Lumpers and splitters|lump]] all basal phytosaurs into only one or two genera, rendering those genera [[Paraphyly|paraphyletic]] [[Evolutionary grade|evolutionary grades]] ancestral to later phytosaurs. More recently, these grades have been broken up into multiple genera.<ref name="StockerButler13" /> ''[[Arganarhinus magnoculus]]'' (from [[Morocco]]) and ''[[Wannia scurriensis]]'' (from Texas)<ref name="Wannia" /> 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.<ref name="StockerButler13" /><ref name=":1" />
''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 (paleontology)|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.<ref name="ICZN" /> 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''.<ref name=":0" /><ref name=":1" />
==Species==
===''P. bransoni''=== ''P. bransoni'' is the [[type species]] of ''Paleorhinus''. It was first described and named by [[Samuel Wendell Williston]] in [[1904 in paleontology|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).<ref name=Paleorhinus/><ref name=Pravusuchus>{{cite journal|author=Michelle R. Stocker |year=2010 |title=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 |journal=Palaeontology |volume=53 |issue=5 |pages=997–1022 |doi=10.1111/j.1475-4983.2010.00983.x |s2cid=83536253 |doi-access=free }}</ref> Hunt and Lucas (1991) referred [[Panhandle Plains Museum|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]].<ref name=HuntLucas91>{{cite journal | last1 = Hunt | first1 = A. P. | last2 = Lucas | first2 = S. G. | year = 1991 | title = The ''Paleorhinus'' biochron and the correlation of the non-marine Upper Triassic of Pangaea | url = https://www.scribd.com/doc/4652272/The-Paleorhinus-biochron-and-the-correlation-of-the-nonmarine-Upper-Triassic-of-Pangaea | journal = Palaeontology | volume = 34 | issue = 2| pages = 487–501 }}</ref> Later, Long and Murry (1995) referred additional material, including isolated [[postcrania]]l 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.<ref name=LongMurry95>{{cite journal | last1 = Long | first1 = R. A. | last2 = Murry | first2 = P. A. | year = 1995 | title = Late Triassic (Carnian and Norian) tetrapods from the southwestern United States | journal = New Mexico Museum of Natural History and Science Bulletin | volume = 4 | pages = 1–254 }}</ref>
===''P. angustifrons''=== ''P. angustifrons'' was first described and named by [[Oskar Kuhn]] in [[1936 in paleontology|1936]] as a species of ''[[Francosuchus]]''. ''P. angustifrons'' is known exclusively from the [[holotype]] BSPG 1931 X 502 a partial skull lacking the [[rostrum (anatomy)|rostrum]] and [[mandible]]s. It was collected at Ebrach Quarry, bed number 9 of [[Bavaria]], southern Germany, from the late [[Carnian]]-aged [[Blasensandstein Member]] of the Hassberge Formation.<ref name=Kuhn36>{{cite journal | last1 = Kuhn | first1 = O | year = 1936 | title = Weitere Parasuchier und Labyrinthodonten aus dem Blasensandstein des mittleren Keuper von Ebrach | journal = Palaeontographica, Abteilung A | volume = 83 | pages = 61–98 }}</ref>
Hunt and Lucas (1991) mistakenly referred to ''F. angustifrons'' as ''[[Ebrachosuchus]] angustifrons'', and considered it and the other two ''[[Francosuchus]]'' species, ''[[Francosuchus broilii|F. broilii]]'' and ''[[Francosuchus latus|F. latus]]'', to be [[synonym (taxonomy)|synonym]]s of ''[[Paleorhinus neukami]]''.<ref name=HuntLucas91/> 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 [[nomen dubium|nomina dubia]] as their [[holotype]]s were destroyed during [[World War II]].<ref name=StockerButler13/>
===''P. cf. arenaceus''=== [[File:Paleorhinus.jpg|thumb|Life restoration of "''Paleorhinus" cf. arenaceus'']] [[Jerzy Dzik|Dzik]] & Sulej (2007) assigned several skulls, partial articulated postcranial skeletons, and numerous isolated phytosaur bones in various [[Ontogeny|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]]''.<ref name=DzikSulej07>{{cite journal|author1=Dzik, J. |author2=Sulej, T. |year=2007 |title=A review of the early Late Triassic Krasiejów biotafrom Silesia, Poland |journal=Palaeontologia Polonica |volume=64 |issue=1 |pages=3–27 |url=http://palaeontologia.pan.pl/PP64/Dzik.pdf }}</ref> ''"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]]'',<ref name="Hungerbühler01">{{Cite journal | last1 = Hungerbühler | first1 = A. | doi = 10.1007/BF03022600 | title = The status and phylogenetic relationships of "Zanclodon"arenaceus: The earliest known phytosaur? | journal = Paläontologische Zeitschrift | volume = 75 | pages = 97–112| year = 2001 | s2cid = 84966794 }}</ref> 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''.<ref name=DzikSulej07/>
===Other species=== [[File:Paleorhinus arenaceus skull and skeleton.jpg|thumb|Skull and skeleton from Poland]] [[File:MEPAN fitozaur.jpg|thumb|upright|Model based on the Polish remains]] 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:
*''[[Ebrachosuchus neukami]]'' Kuhn, 1936<ref name=Kuhn36/> - Hunt and Lucas (1991) reassigned the species as ''Paleorhinus neukami''.<ref name=HuntLucas91/> Subsequent researches accepted this referral. More recently, phylogenetic analysis found ''E. neukami'' to be more closely related to [[Phytosauridae]] than to ''Paleorhinus'' and thus the genus ''Ebrachosuchus'' was re-validated.<ref name=StockerButler13/> *''[[Francosuchus broilii]]'' and ''[[Francosuchus latus|F. latus]]'' Kuhn, 1933 - Gregory (1962)<ref name=Gregory62>{{cite journal | last1 = Gregory | first1 = J. T. | year = 1962 | title = The relationships of the American phytosaur ''Rutiodon'' | journal = American Museum Novitates | issue = 2095 | pages = 1–22 }}</ref> and Wastphal (1976) referred these species, as well as, ''E. neukami'' and ''P. angustifrons'' to ''Paleorhinus'' subgenus ''Francosuchus'', while Chatterjee (1978) placed the four species in ''Francosuchus'' in a different subfamily. Hunt and Lucas (1991) considered these species to be synonymous with ''Paleorhinus neukami''.<ref name=HuntLucas91/> More recently, ''F. broilii'' and ''F. latus'' were considered to be [[nomen dubium|nomina dubia]].<ref name=StockerButler13/> *''[[Francosuchus trauthi|"Francosuchus" trauthi]]'' Huene, 1939 - ''"F." trauthi'' was synonymized with ''Paleorhinus'', but a re-description of the species by Butler (2013) found no evidence to support the synonymy, or even a phytosaurian identification. Although it is based solely on a [[rostrum (anatomy)|rostrum]] fragment, ''"F." trauthi'' was found to possess a unique combination of characters that distinguish it from all other Triassic [[tetrapod]]s. Thus it was reassigned to a new genus, ''[[Dolerosaurus]]'',<ref name=Dolerosaurus>{{Cite journal | last1 = Butler | first1 = R. J. | title = ''{{'}}Francosuchus' trauthi'' is not ''Paleorhinus'': Implications for Late Triassic vertebrate biostratigraphy | doi = 10.1080/02724634.2013.740542 | journal = Journal of Vertebrate Paleontology | volume = 33 | issue = 4 | pages = 858–864 | year = 2013 | s2cid = 86386615 }}</ref> now referable to [[Tetrapoda]] ''[[incertae sedis]]''.<ref name=StockerButler13/> *''[[Paleorhinus magnoculus]]'' Dutuit, 1977b - Long and Murry (1995) reassigned this species to a new genus ''[[Arganarhinus]]''. Although several studies reassigned it back to ''Paleorhinus'', more recent studies disagree with this referral.<ref name=StockerButler13/><ref name=Wannia/> Kammerer et al. (2016) considered ''P. magnoculus'' to be a species of ''Parasuchus,'' alongside ''P. hislopi, P. bransoni,'' and ''P. angustifrons''.<ref name=":0">{{Cite journal|last1=Kammerer|first1=Christian F.|last2=Butler|first2=Richard J.|last3=Bandyopadhyay|first3=Saswati|last4=Stocker|first4=Michelle R.|date=2016|title=Relationships of the Indian phytosaur ''Parasuchus hislopi'' Lydekker, 1885|url=https://onlinelibrary.wiley.com/doi/abs/10.1002/spp2.1022|journal=Papers in Palaeontology|language=en|volume=2|issue=1|pages=1–23|doi=10.1002/spp2.1022|s2cid=83780331 |issn=2056-2802|url-access=subscription}}</ref> *''[[Paleorhinus parvus|"Paleorhinus" parvus]]'' Mehl, 1928 - Hunt and Lucas (1991) considered this species to be [[synonym (taxonomy)|synonymous]] with ''P. bransoni'',<ref name=HuntLucas91/> while Stocker and Butler (2013) suggested that ''"P." parvus'' may not be referable to ''Paleorhinus''.<ref name=StockerButler13/> *''[[Paleorhinus sawini|"Paleorhinus" sawini]]'' Long and Murry, 1995 - some studies considered this species to be [[synonym (taxonomy)|synonymous]] with ''P. bransoni'', although recent species-level [[phylogenetic]] analyses of [[phytosaur]]s suggest that this species is more closely related to [[Phytosauridae]] than to ''P. bransoni'' and thus can't be referred to ''Paleorhinus''.<ref name=StockerButler13/><ref name=Pravusuchus/><ref name=Wannia/> *''[[Paleorhinus scurriensis|"Paleorhinus" scurriensis]]'' Langston, 1949 - Hunt and Lucas (1991) considered this species to be [[synonym (taxonomy)|synonymous]] with ''P. bransoni'',<ref name=HuntLucas91/> although recent species-level phylogenetic analyses of phytosaurs suggest that this species is [[basal (phylogenetics)|basal]] to the clade formed by ''P. bransoni'' and the [[Phytosauridae]], and thus can't be referred to ''Paleorhinus''.<ref name=StockerButler13/><ref name=Pravusuchus/> This species was reassigned to its own genus, ''[[Wannia]]'' by Stocker (2013), and represents the most [[basal (phylogenetics)|basal]] known phytosaur.<ref name=Wannia>{{Cite journal | last1 = Stocker | first1 = M. R. | title = A new taxonomic arrangement for ''Paleorhinus scurriensis'' | doi = 10.1017/S1755691013000340 | journal = Earth and Environmental Science Transactions of the Royal Society of Edinburgh | volume = 103 | issue = 3–4 | pages = 251–263 | year = 2013 | s2cid = 130567475 }}</ref> *''[[Parasuchus hislopi]]'' Lydekker, 1885 - ''P. hislopi'' was based on a chimeric [[syntype]] material - a [[rhynchosauria]]n [[basicranium]] mixed with [[phytosauria]]n partial snout, scutes and some teeth. [[Friedrich von Huene]] (1940) identified the basicranium as belonging to ''[[Paradapedon huxleyi]]'' (now known as ''Hyperodapedon huxleyi''), thus he assigned 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''. Gregory (1962) accepted this proposal. [[Sankar Chatterjee]] (1978), who described many complete remains of the Indian parasuchian, showed that it is not assignable either to ''Brachysuchus'' (which is closely related to or synonymous with ''[[Angistorhinus]]''), or to ''Phytosaurus'' (a [[nomen dubium|doubtful name]], probably the [[senior synonym]] of ''[[Nicrosaurus]]''). He noted that since the rhynchosaur basicranium is neither the [[holotype]] of ''P. hislopi'', nor the [[lectotype]] of ''Paradapedon huxleyi'', the suppression of ''P. hislopi'' should be avoided.<ref name=Chatterjee78>{{cite journal |author=Sankar Chatterjee |year=1978 |title=A primitive parasuchid (phytosaur) reptile from the Upper Triassic Maleri Formation of India |url=http://palaeontology.palass-pubs.org/pdf/Vol%2021/Pages%2083-127.pdf |journal=Palaeontology |volume=21 |issue=1 |pages=83–127 |access-date=2013-06-25 |url-status=usurped |archive-url=https://web.archive.org/web/20130810232427/http://palaeontology.palass-pubs.org/pdf/Vol%2021/Pages%2083-127.pdf |archive-date=2013-08-10 }}</ref> Nevertheless, Hunt and Lucas (1991) considered the species to be a [[nomen dubium]], and provisionally created the combination ''Paleorhinus hislopi'' for the diagnosable phytosaur specimens from the [[Lower Maleri Formation]], stating that the Indian species can be distinguished from other ''Paleorhinus'' species.<ref name=HuntLucas91/> To avoid additional confusion, the nondiagnostic holotype of ''P. hislopi'' was replaced by a [[neotype]] with approval from the [[International Commission on Zoological Nomenclature|ICZN]] (Opinion 2045) following the application of Chatterjee (2001).<ref name=ICZN>{{cite journal |year=2003 |title=''Parasuchus Hislopi'' Lydekker, 1885 (Reptilia, Archosauria): Lectotype Replaced By A Neotype |url=http://biostor.org/cache/pdf/b8/fc/68/b8fc688a53ce7a142c5951d1ed169396.pdf |journal=Bulletin of Zoological Nomenclature |volume=60 |pages=174–175 }}</ref> As a result, most subsequent studies referred all ''Paleorhinus'' species to ''Parasuchus'' in accordance with the rules of the [[International Code of Zoological Nomenclature|ICZN]], while others kept referring ''P. hislopi'' to ''Paleorhinus''. In light of the complex taxonomic history of ''Paleorhinus'', studies as of 2013 considered the synonymy between the genera to be premature.<ref name=StockerButler13/><ref name=Wannia/> However, later studies argued that ''Paleorhinus bransoni'' and ''Paleorhinus angustifrons'' formed a strong clade with ''Parasuchus hislopi''. This clade would receive the name ''Parasuchus'' under ICZN rules, formally synonymizing the two genera in this more restricted sense.<ref name=":0" /><ref name=":1">{{Cite journal|last1=Jones|first1=Andrew S.|last2=Butler|first2=Richard J.|date=2018-12-10|title=A new phylogenetic analysis of Phytosauria (Archosauria: Pseudosuchia) with the application of continuous and geometric morphometric character coding|journal=PeerJ|language=en|volume=6|article-number=e5901|doi=10.7717/peerj.5901|pmid=30581656 |pmc=6292387 |issn=2167-8359 |doi-access=free }}</ref> *''[[Promystriosuchus ehlersi]]'' Case, 1922<ref>{{cite journal | last1 = Case | first1 = E. C. | year = 1922 | title = New reptiles and stegocephalians from the Upper Triassic of western Texas | journal = Carnegie Institution of Washington Publication | volume = 321 | pages = 1–84 }}</ref> - Gregory (1962)<ref name=Gregory62/> and Hunt and Lucas (1991)<ref name=HuntLucas91/> referred this taxon to ''Paleorhinus'' on the genus level, and considered it to be a [[nomen dubium]] on the species level, due to the poor preservation of it holotype and only known specimen. Stocker and Butler (2013) suggested that ''P. ehlersi'' may not be referable to ''Paleorhinus'' and referred it to [[Phytosauria]] ''incertae sedis''.<ref name=StockerButler13/>
=== Phylogeny === 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''.<ref name=":0" />
{{clade|{{clade |1=''[[Wannia scurriensis]]'' |2={{clade |1={{clade |1='''''Parasuchus (Paleorhinus) bransoni''''' |2={{clade |1='''''Parasuchus (Paleorhinus) angustifrons''''' |2=''[[Parasuchus hislopi]]'' }} }} |2={{clade |1=''[[Ebrachosuchus neukami]]'' |label2=[[Mystriosuchinae]] |2={{clade |1=''[[Angistorhinus]]'' |2=''"Paleorhinus" sawini'' |3=''[[Brachysuchus megalodon]]'' |4={{clade |1=''[[Protome batalaria]]'' |2=''"Machaeroprosopus" zunii'' |3=''[[Rutiodon carolinensis]]'' |label4=[[Leptosuchomorpha]] |4={{clade |1=TMM 31173-120 |2=''"Phytosaurus" doughtyi'' |DUP-3={{clade |1=''[[Leptosuchus studeri]]'' |2={{clade |1=''[[Leptosuchus imperfecta]]'' |2=''[[Leptosuchus crosbiensis]]'' }} }} |3={{clade |1={{clade |1=''[[Smilosuchus lithodendrorum]]'' |2={{clade |1=''[[Smilosuchus gregorii]]'' |2=''[[Smilosuchus adamanensis]]'' }} }} |2={{clade |1=''[[Pravusuchus hortus]]'' |label2=[[Mystriosuchini]] |2={{clade |1=''[[Machaeroprosopus mccauleyi]]'' |2={{clade |1=''[[Machaeroprosopus pristinus]]'' |2={{clade |1=''[[Machaeroprosopus jablonskiae]]'' |2=''[[Mystriosuchus westphali]]''}} }} }} }} }} }} }} }} }} }} }}|style=font-size:100%;line-height:85%|label1=[[Parasuchidae]] }}
==References== {{Reflist}} {{Portal|paleontology}}
{{Phytosauria}} {{Taxonbar|from=Q21711013}}
[[Category:Phytosauria]] [[Category:Prehistoric reptile genera]] [[Category:Carnian genera]] [[Category:Late Triassic reptiles of Europe]] [[Category:Late Triassic reptiles of North America]] [[Category:Chinle fauna]] [[Category:Fossil taxa described in 1904]] [[Category:Taxa named by Samuel Wendell Williston]]