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Phytosauria

Reptilia

Taxonomy
Phytosauria was named by Jaeger (1828). It is not extant. It was considered monophyletic by Stocker (2010) and Nesbitt (2011).

It was reranked as the order Phytosauria by Osborn (1903) and Osborn (1903); it was synonymized subjectively with Phytosauriformes by Hay (1930); it was synonymized subjectively with Parasuchia by Kuhn (1946), Kuhn (1964), Kuhn (1966) and Charig (1967); it was synonymized subjectively with Parasuchida by Kuhn (1971); it was reranked as the unranked clade Phytosauria by Hungerbühler (2002), Stocker (2010), Nesbitt (2011) and Ezcurra (2016).

It was assigned to Diapsida by Osborn (1903) and Osborn (1903); to Parasuchia by Osborn (1904), Lull (1915), Lull (1917) and Roxo (1937); to Thecodontia by Baird (1957) and Carroll (1988); to Suchia by Weinbaum and Hungerbühler (2007); to Archosauria by Stocker (2010); to Crurotarsi by Hungerbühler (2002) and Nesbitt (2011); and to Pseudosuchia by Ezcurra (2016).

Synonyms
Synonymy list
YearName and author
1828Phytosauria Jaeger
1828Phytosaurus Jaeger p. 22–23
1828Phytosaurus cylindricodon Jaeger p. 23
1828Phytosaurus cubicodon Jaeger p. 33
1836Palaeosaurus Riley and Stutchbury p. 398
1836Palaeosaurus platyodon Riley and Stutchbury p. 398
1840Palaeosaurus Riley and Stutchbury p. 352
1840Palaeosaurus platyodon Riley and Stutchbury p. 352
1841Palaeosaurus Owen p. 267
1841Palaeosaurus platyodon Owen p. 268
1842Phytosaurus cylindricodon Owen p. 118
1842Palaeosaurus Owen p. 154
1842Palaeosaurus platyodon Owen p. 154
1844Belodon Meyer
1844Belodon plieningeri Meyer
1844Termatosaurus albertii Meyer and Plieninger
1844Phytosaurus cubicodon Meyer and Plieninger p. 92
1844Phytosaurus cylindricodon Meyer and Plieninger p. 92
1845Belodon Meyer p. 282
1845Palaeosaurus Meyer p. 282
1846Palaeosaurus platyodon Plieninger p. 151
1849Phytosaurus d'Orbigny p. 206
1849Palaeosaurus d'Orbigny p. 208
1850Palaeosaurus platyodon King p. 239
1851Clepsysaurus pennsylvanicus Lea p. 172
1851Clepsysaurus Lea p. 205
1851Clepsysaurus pennsylvanicus Lea p. 205
1852Palaeosaurus Quenstedt p. 109
1852Belodon Quenstedt p. 110
1852Belodon plieningeri Quenstedt p. 110
1855Belodon plieningeri Meyer p. 149
1855Palaeosaurus platyodon Meyer p. 158
1856Clepsisaurus Emmons
1856Clepsisaurus pennsylvanius Emmons p. 300
1856Clepsisaurus leai Emmons p. 313
1856Palaeosaurus platyodon Emmons p. 315
1856Centemodon Lea
1856Centemodon sulcatus Lea
1856Omosaurus Leidy
1856Compsosaurus Leidy p. 165 figs. not figured
1856Compsosaurus priscus Leidy p. 165 figs. not figured
1856Omosaurus perplexus Leidy p. 256
1856Palaeosaurus Roemer p. 788
1856Palaeosaurus platyodon Roemer p. 788
1857Omosaurus perplexus Emmons p. 67
1857Clepsisaurus pennsylvanicus Emmons p. 73
1857Clepsisaurus leai Emmons p. 79
1857Compsosaurus priscus Emmons p. 87
1857Palaeosaurus platyodon Emmons p. 87
1857Belodon plieningeri Rütimeyer p. 152
1858Termatosaurus albertii Quenstedt p. 33
1859Eurydorus Leidy
1859Eurydorus serridens Leidy
1859Belodon Owen p. 141
1859Belodon plieningeri Owen p. 141
1860Palaeosaurus Owen p. 249
1860Palaeosaurus platyodon Owen p. 249
1860Belodon Owen p. 251
1860Belodon plieningeri Owen p. 251
1861Palaeosaurus Owen p. 276
1861Palaeosaurus platyodon Owen p. 276
1861Belodon Owen p. 278
1861Belodon plieningeri Owen p. 278
1861Belodon plieningeri Quenstedt p. 512
1866Belodon plieningeri Haeckel p. CXXXIV
1866Palaeosaurus Haeckel p. CXXXIV
1866Palaeosaurus platyodon Haeckel p. CXXXIV
1867Palaeosaurus Quenstedt p. 139
1869Belodon lepturus Cope p. 22
1869Clepsysaurus Cope p. 733
1869Clepsysaurus pennsylvanicus Cope p. 733
1870Belodon Cope
1870Belodon lepturus Cope
1870Clepsysaurus Cope
1870Clepsysaurus pennsylvanicus Cope pp. 122-A
1870Dicynodon rosmarus Cope p. 232
1870Clepsysaurus Cope p. 444
1870Palaeosaurus Huxley p. 34
1870Clepsysaurus Huxley p. 49
1875Parasuchia Cope p. 14
1875Belodon priscus Cope p. 34
1875Clepsysaurus leai Cope p. 36
1875Clepsysaurus Cope p. 39
1875Clepsysaurus pennsylvanicus Cope p. 39
1876Termatosaurus alberti Henry p. 404 figs. Pl 1, figs 5,5.1
1877Palaeoctonus Cope pp. 182-183
1877Palaeoctonus appalachianus Cope pp. 182-183
1877Clepsysaurus veatleianus Cope p. 184
1877Palaeoctonus aulacodus Cope p. 184
1877Suchoprion Cope p. 185
1877Suchoprion cyphodon Cope p. 185
1878Palaeosaurus fraserianus Cope
1878Suchoprion sulcidens Cope
1878Palaeosaurus fraserianus Cope p. 58
1879Clepsysaurus pennsylvanicus Miller p. 152
1881Belodon scolopax Cope p. 922
1881Palaeosaurus stricklandi Davis p. 421 figs. Pl. 22:6
1882Clepsysaurus Marsh
1882Palaeosaurus Marsh
1882Palaeosaurus Quenstedt p. 189
1882Clepsysaurus Sauvage pp. 35-36
1882Palaeosaurus Sauvage p. 36
1883Parasuchia Dollo p. 310
1883Belodon Dollo p. 311
1883Palaeosaurus stricklandi Sauvage p. 39
1884Clepsysaurus Dollo p. 298
1884Palaeosaurus Dollo p. 298
1884Clepsysaurus Marsh p. 69
1884Palaeosaurus Marsh p. 69
1884Clepsysaurus Marsh p. 338
1884Palaeosaurus Marsh p. 338
1884Clepsysaurus Vetter p. 363
1884Palaeosaurus Vetter p. 363
1885Clepsysaurus Marsh p. 765
1885Palaeosaurus Marsh p. 765
1887Belodon scolopax Cope p. 221
1890Palaeosaurus stricklandi Woodward and Sherborn p. 261
1890Thecodontosaurus platyodon Woodward and Sherborn p. 291
1890Parasuchia Zittel p. 450
1890Palaeosaurus Zittel p. 722
1890Palaeosaurus platyodon Zittel p. 722
1890Clepsysaurus Zittel p. 731
1893Belodon superciliosus Cope p. 12
1893Palaeoctonus orthodon Cope p. 15
1893Palaeoctonus dumblianus Cope p. 16
1893Belodon validus Marsh p. 170
1894Belodon priscus Lyman p. 207
1895Clepsysaurus Marsh
1895Palaeosaurus Marsh
1895Palaeosaurus platyodon Seeley p. 145
1896Belodon ingens Fraas
1896Rhytidodon rostratus Marsh
1896Clepsysaurus pennsylvanicus Marsh p. 147
1896Clepsysaurus Marsh p. 240
1896Palaeosaurus Marsh p. 240
1898Parasuchia Gadow p. 19
1898Heterodontosuchus Lucas
1898Heterodontosuchus ganei Lucas
1898Belodon ingens Spangenberg p. 408
1898Parasuchia Woodward p. 141
1901Clepsysaurus Nopcsa p. 196–197
1901Clepsysaurus pennsylvanicus Nopcsa p. 196–197
1901Palaeosaurus Nopcsa p. 197
1901Palaeosaurus frazerianus Nopcsa p. 197
1901Palaeosaurus platyodon Nopcsa p. 197
1901Palaeoctonus Nopcsa p. 201
1901Palaeoctonus appalachianus Nopcsa p. 201
1902Dicynodon rosmarus Hay p. 430
1902Palaeoctonus Hay p. 490
1902Palaeoctonus appalachianus Hay p. 490
1902Palaeoctonus dumblianus Hay p. 490
1902Palaeoctonus orthodon Hay p. 490
1902Suchoprion Hay p. 490
1902Suchoprion aulacodus Hay p. 490
1902Suchoprion cyphodon Hay p. 490
1902Suchoprion sulcidens Hay p. 490
1902Clepsysaurus Hay p. 492
1902Clepsysaurus pennsylvanicus Hay p. 492
1902Clepsysaurus veatleianus Hay p. 492
1902Thecodontosaurus fraserianus Hay p. 492
1902Parasuchia Hay p. 509
1902Phytosaurus Hay p. 509
1902Phytosaurus cylindricodon Hay p. 509
1902Phytosaurus leaii Hay p. 510
1902Phytosaurus lepturus Hay p. 510
1902Phytosaurus priscus Hay p. 510
1902Phytosaurus rostratus Hay p. 510
1902Phytosaurus scolopax Hay p. 510
1902Phytosaurus serridens Hay p. 510
1902Phytosaurus superciliosus Hay p. 510
1902Phytosaurus validus Hay p. 510
1902Heterodontosuchus Hay p. 515
1902Heterodontosuchus ganei Hay p. 515
1902Palaeoctonus Huene
1902Palaeosaurus Huene
1902Suchoprion Huene
1902Rileya bristolensis Huene pp. 62-63
1902Omosaurus Lucas p. 435
1902Omosaurus perplexus Lucas p. 435
1903Phytosauria Osborn p. 276
1904Phytosauria Osborn p. 100
1904Parasuchia Osborn p. 112
1907Palaeosaurus stricklandi Sauvage p. 14
1907Clepsysaurus Sauvage p. 15
1908Termatosaurus albertii Amthor p. 94 figs. Pl 2, figs 1,2
1908Rileya bristolensis Huene p. 16
1908Rileya platyodon Huene p. 16
1908Rileya bristolensis Huene p. 99
1908Rileya platyodon Huene p. 99
1910Belodonti Jaekel p. 340
1911Mystriosuchus ruetimeyeri Huene p. 31
1911Belodonti Jaekel p. 163
1913Rutiodon manhattanensis Huene p. 277
1914Palaeosaurus Huene p. 146
1915Parasuchia Lull p. 98
1915Belodon validus Lull pp. 109-110
1915Phytosauria Lull p. 109
1915Rutiodon manhattanensis Lull p. 113
1915Clepsysaurus Lull p. 119
1915Palaeosaurus Lull p. 119
1915Angistorhinus gracilis Mehl p. 146
1917Phytosauria Lull p. 110
1917Rutiodon validus Lull p. 110
1917Parasuchia Lull p. 113
1917Phytosaurus Watson p. 179
1921Palaeoctonus Huene p. 561
1921Palaeoctonus appalachianus Huene p. 561
1921Palaeoctonus aulacodus Huene p. 563
1921Suchoprion Huene p. 564
1921Clepsysaurus Huene p. 565
1921Clepsysaurus pennsylvanicus Huene p. 565
1921Suchoprion cyphodon Huene p. 565
1921Clepsysaurus veatleianus Huene p. 567
1922Leptosuchus imperfecta Case p. 68
1922Parasuchia Huene p. 22
1922Angistorhinopsis Huene p. 23
1922Phytosaurus Huene p. 23
1923Angistorhinopsis Huene
1923Angistorhinopsis ruetimeyeri Huene
1928Parasuchia Perrier p. 3118
1930Clepsysaurus Camp p. ix
1930Dicynodon rosmarus Hay p. 63
1930Phytosaurus Hay p. 142
1930Phytosaurus cylindricodon Hay p. 142
1930Phytosaurus leaii Hay p. 142
1930Phytosaurus lepturus Hay p. 142
1930Phytosaurus scolopax Hay p. 142
1930Phytosaurus superciliosus Hay p. 142
1930Phytosaurus validus Hay p. 142
1930Angistorhinus gracilis Hay p. 143
1930Leptosuchus imperfectus Hay p. 143
1930Heterodontosuchus Hay p. 144
1930Heterodontosuchus ganei Hay p. 144
1930Rutiodon manhattanensis Hay p. 145
1930Palaeoctonus Hay p. 171
1930Palaeoctonus appalachianus Hay p. 171
1930Palaeoctonus dumblianus Hay p. 171
1930Clepsysaurus Hay p. 172
1930Clepsysaurus fraserianus Hay p. 172
1930Clepsysaurus pennsylvanicus Hay p. 172
1930Clepsysaurus veatleianus Hay p. 172
1930Palaeoctonus orthodon Hay p. 172
1930Suchoprion Hay p. 172
1930Suchoprion aulacodus Hay p. 172
1930Suchoprion cyphodon Hay p. 172
1930Suchoprion sulcidens Hay p. 172
1932Francosuchus Kuhn
1932Francosuchus broilii Kuhn p. 123
1932Francosuchus latus Kuhn p. 123
1934Thecodontosaurus platyodon Swinton p. 31
1937Parasuchia Roxo p. 44
1937Phytosauria Roxo p. 44
1938Palaeosaurus Berckhemer p. 196
1939Palaeosaurus Kuhn p. 57
1940Brachysuchus maleriensis Huene p. 21
1940Phytosaurus plieningeri Huene p. 23
1941Clepsysaurus Colbert and Chafee p. 12
1941Clepsysaurus pennsylvanicus Colbert and Chafee p. 12
1941Clepsysaurus manhattanensis Colbert and Chafee p. 14
1941Clepsysaurus validus Colbert and Chafee p. 14
1944Clepsysaurus Rapp p. 287
1944Clepsysaurus pennsylvanicus Rapp p. 287
1946Parasuchia Kuhn p. 64
1953Clepsysaurus Lull p. 89
1954Parasuchia Huene p. 44
1956Palaeosaurus Romer p. 614
1957Phytosauria Baird p. 486
1957Thecodontosaurus platyodon Robinson p. 262
1958Phytosaurus maleriensis Colbert p. 75
1958Parasuchia Huene p. 205
1961Rileyasuchus bristolensis Kuhn
1962Clepsysaurus pennsylvanicus Gregory p. 4
1962Mystriosuchus plieningeri Gregory p. 12
1963Termatosaurus alberti Persson pp. 21-22
1964Parasuchia Kuhn p. 317
1964Phytosaurus Kuhn p. 319
1964Rileyasuchus bristolensis Kuhn p. 319
1964Francosuchus Kuhn p. 324
1966Parasuchia Kuhn p. 77
1966Belodon Kuhn p. 81
1966Francosuchus Kuhn p. 81
1966Phytosaurus Kuhn p. 81
1967Parasuchia Charig p. 704
1967Phytosaurus Charig p. 704
1967Rutiodon rutimeyeri Charig p. 709
1970Palaeosauriscus platyodon Steel p. 25
1970Palaeosauriscus fraserianus Steel p. 26
1970Palaeosauriscus stricklandi Steel p. 26
1971Belodon plieningeri Kuhn p. 14
1971Francosuchus broilii Kuhn p. 14
1971Parasuchida Kuhn p. 14
1971Angistorhinopsis Kuhn p. 15
1971Angistorhinopsis ruetimeyeri Kuhn p. 15
1971Termatosaurus albertii Kuhn p. 20
1974Parasuchia Haubold p. 40
1985Parasuchia Galton
1985Parasuchia Gauthier and Padian pp. 187-189 fig. 1
1988Belodon Carroll
1988Phytosauria Carroll
1990Parasuchia Sereno and Arcucci p. 39 fig. 7
1991Parasuchia Sereno p. 27
1993Parasuchia Parrish p. 290 fig. 1
1995Parasuchia Long and Murry
2002Phytosauria Hungerbühler p. 380
2005Parasuchia Irmis pp. 71-72
2007Phytosauria Weinbaum and Hungerbühler p. 141
2010Phytosauria Stocker
2011Phytosauria Nesbitt p. 193 fig. 51
2016Phytosauria Ezcurra

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RankNameAuthor
kingdomAnimalia()
Bilateria
EubilateriaAx 1987
Deuterostomia
phylumChordataHaeckel 1874
subphylumVertebrata
superclassGnathostomata
Osteichthyes()
subclassSarcopterygii()
subclassDipnotetrapodomorpha(Nelson 2006)
subclassTetrapodomorpha()
Tetrapoda
Reptiliomorpha
Anthracosauria
subclassAmphibiosauriaKuhn 1967
RankNameAuthor
Cotylosauria()
Amniota
Sauropsida
classReptilia
subclassEureptilia()
Romeriida
Diapsida()
Archosauromorpha(Huene 1946)
Crocopoda
ArchosauriformesGauthier 1986
Eucrocopoda
Archosauria()
Pseudosuchia(Zittel 1890)
Phytosauria(Jaeger 1828)
Phytosauria(Jaeger 1828)

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

Unr. †Phytosauria Jaeger 1828
show all | hide all
G. †Coburgosuchus Heller 1954
+
Coburgosuchus goeckeli Heller 1954
G. †Diandongosuchus Li et al. 2012
+
Diandongosuchus fuyuanensis Li et al. 2012
G. †Heterodontichnites Rinehart et al. 2006
+
Heterodontichnites hunti Rinehart et al. 2006
Fm. †Parasuchidae Lydekker 1885
+
G. †Apatopus Baird 1957
+
Apatopus lineatus Bock 1952
G. †Ebrachosuchus Kuhn 1936
+
Ebrachosuchus neukami Kuhn 1936
Subfm. †Mystriosuchinae von Huene 1915
+
G. †Angistorhinus Mehl 1913
+
Angistorhinus alticephalus Stovall and Wharton Jr. 1936
Angistorhinus grandis Mehl 1913
Angistorhinus maximus Mehl 1928
Angistorhinus parvus Mehl 1928
Angistorhinus talainti Dutuit 1977
G. †Brachysuchus Case 1929
+
Unr. †Leptosuchomorpha Stocker 2010
+
G. †Leptosuchus Case 1922
+
+
Invalid names: Phytosaurus doughtyi Case 1920 [synonym]
Leptosuchus studeri Case and White 1934
Unr. †Mystriosuchini Huene 1915
+
G. †Machaeroprosopus Mehl 1916
+
Invalid names: Arribasuchus Long and Murry 1995 [synonym], Pseudopalatus Mehl 1928 [synonym], Redondasaurus Hunt and Lucas 1993 [synonym]
G. †Mystriosuchus Fraas 1896
Invalid names: Pseudopalatinae Long and Murry 1995 [replaced]
G. †Pravusuchus Stocker 2010
+
Pravusuchus hortus Stocker 2010
G. †Smilosuchus Long and Murry 1995
+
+
Invalid names: Machaeroprosopus zunii Camp 1930 [synonym]
Smilosuchus gregorii Camp 1930
G. †Protome Stocker 2012
+
Protome batalaria Stocker 2012
G. †Rutiodon Emmons 1856
+
Rutiodon carolinensis Emmons 1856
Invalid names: Metarhinus Jaekel 1910 [synonym], Palaeonornis Emmons 1857 [nomen dubium], Palaeonornis struthionoides Emmons 1857 [nomen dubium]
G. †Volcanosuchus Datta et al. 2019
+
Volcanosuchus statisticae Datta et al. 2019
Invalid names: Phytosauridae Jaeger 1828 [replaced]
G. †Parasuchus Lydekker 1885
+
Parasuchus bransoni Williston 1904
Parasuchus hislopi Lydekker 1885
Parasuchus magnoculus Long and Murry 1995
Invalid names: Arganarhinus Long and Murry 1995 [synonym], Paleorhinus Williston 1904 [synonym]
G. †Wannia Stocker 2013
+
Wannia scurriensis Langston 1949
Invalid names: Parasuchinae Lydekker 1885 [empty]
G. †Promystriosuchus Case 1922
G. †Rectilinetopus Biron and Dutuit 1981
+
Rectilinetopus ourikensis Biron and Dutuit 1981
G. †Termatosaurus Plieninger 1844
+
Termatosaurus alberti Meyer and Plieninger 1844
Termatosaurus crocodilinus Quenstedt 1858
Invalid names: Angistorhinopsis Huene 1922 [nomen dubium], Angistorhinopsis ruetimeyeri Huene 1911 [nomen dubium], Angistorhinus gracilis Mehl 1915 [nomen dubium], Belodon Meyer 1844 [nomen dubium], Belodon ingens Fraas 1896 [nomen dubium], Belodon lepturus Cope 1869 [nomen dubium], Belodon plieningeri Meyer 1844 [nomen dubium], Belodon scolopax Cope 1881 [nomen dubium], Belodon superciliosus Cope 1893 [nomen dubium], Belodon validus Marsh 1893 [nomen dubium], Brachysuchus maleriensis Huene 1940 [nomen dubium], Centemodon Lea 1856 [nomen dubium], Centemodon sulcatus Lea 1856 [nomen dubium], Clepsysaurus Lea 1851 [nomen dubium], Clepsysaurus leai Emmons 1856 [nomen dubium], Clepsysaurus pennsylvanicus Lea 1851 [nomen dubium], Clepsysaurus veatleianus Cope 1877 [nomen dubium], Compsosaurus Leidy 1856 [nomen dubium], Compsosaurus priscus Leidy 1856 [nomen dubium], Dicynodon rosmarus Cope 1870 [nomen dubium], Eurydorus Leidy 1859 [nomen dubium], Eurydorus serridens Leidy 1859 [nomen dubium], Francosuchus Kuhn 1932 [nomen dubium], Francosuchus broilii Kuhn 1932 [nomen dubium], Francosuchus latus Kuhn 1932 [nomen dubium], Heterodontosuchus Lucas 1898 [nomen dubium], Heterodontosuchus ganei Lucas 1898 [nomen dubium], Leptosuchus imperfecta Case 1922 [nomen dubium], Omosaurus Leidy 1856 [nomen dubium], Omosaurus perplexus Leidy 1856 [nomen dubium], Palaeoctonus Cope 1877 [nomen dubium], Palaeoctonus appalachianus Cope 1877 [nomen dubium], Palaeoctonus aulacodus Cope 1877 [nomen dubium], Palaeoctonus dumblianus Cope 1893 [nomen dubium], Palaeoctonus orthodon Cope 1893 [nomen dubium], Palaeosaurus Riley and Stutchbury 1836 [nomen dubium], Palaeosaurus fraserianus Cope 1878 [nomen dubium], Palaeosaurus platyodon Riley and Stutchbury 1836 [nomen dubium], Palaeosaurus stricklandi Davis 1881 [nomen dubium], Parasuchia Huxley [synonym], Phytosaurus Jaeger 1828 [nomen dubium], Phytosaurus cubicodon Jaeger 1828 [nomen dubium], Phytosaurus cylindricodon Jaeger 1828 [nomen dubium], Rhytidodon rostratus Marsh 1896 [nomen dubium], Rileya bristolensis Huene 1902 [nomen dubium], Rutiodon manhattanensis Huene 1913 [nomen dubium], Suchoprion Cope 1877 [nomen dubium], Suchoprion cyphodon Cope 1877 [nomen dubium], Suchoprion sulcidens Cope 1878 [nomen dubium], Termatosaurus albertii Meyer and Plieninger 1844 [nomen dubium]
Diagnosis
ReferenceDiagnosis
E. Perrier 1928 (Parasuchia)Des dents sur toute la longueur des prémaxillaires qui sont très longs. Narines placées très en arrière. Vertèbres biconcaves ou biplanes.
M. R. Stocker 2010A stem-based group that is the most inclusive clade containing Rutiodon carolinensis (Emmons 1856) but not Aetosaurus ferratus (Fraas 1877), Rauisuchus tiradentes (Huene 1942), Prestosuchus chiniquensis (Huene 1942), Ornithosuchus woodwardi (Newton 1894) or Crocodylus niloticus (Laurenti 1768).
S. J. Nesbitt 2011The most inclusive clade containing Rutiodon carolinensis(Emmons, 1856) but not Aetosaurus ferratus Fraas, 1877, Rauisuchus tiradentes Huene, 1942, Prestosuchus chiniquensis Huene, 1942, Ornithosuchus longidens Huxley, 1877, and Crocodylus niloticus Laurenti, 1768 (sensu Sereno, 2005).