PBDB Taxon

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Grallator tuberosus

Reptilia - Grallatoridae

Taxonomy

Ornithichnites tuberosus was named by Hitchcock (1836). Its type specimen is AC 31/73, a footprint, and it is a trace fossil.

It was recombined as Ornithoidichnites tuberosus by Hitchcock (1837), Hitchcock (1841), Hitchcock (1843), Redfield (1843) and Hitchcock (1844); it was recombined as Eubrontes tuberosus by Hitchcock (1845), Hay (1902) and Mayor and Sarjeant (2001); it was synonymized subjectively with Brontozoum validum by Hitchcock (1858); it was synonymized subjectively with Brontozoum loxonyx by Miller (1879); it was synonymized subjectively with Brontozoum sillimani by Quenstedt (1882); it was recombined as Anchisauripus tuberosus by Lull (1904), Lull (1915), Lull (1917), Hitchcock and Grier (1927), Hay (1930), Lull (1953), Kuhn (1963), Nelson (1965), Haubold (1971), Welles (1971), Haubold (1974), Lockley (1986) and Fujita et al. (2008); it was considered a nomen dubium by Baird (1957); it was recombined as Grallator tuberosus by Weems (1992), Gierlinski (1996), Kordos (2000), Osi et al. (2011) and Li et al. (2019).

Sister species lacking formal opinion data

G. pisanus

Synonyms

Ornithichnites gracilis was named by Hitchcock (1837) [C. H. Hitchcock in E. Hitchcock (1865)]. It is considered to be a form taxon. Its type specimen is AC 17/2, a footprint, and it is a trace fossil. Its type locality is Ferry above Turners Falls, tracksite, which is in a Hettangian/Sinemurian fluvial-lacustrine sandstone in the Portland Formation of Massachusetts.

It was recombined as Grallator gracilis by Hitchcock (1865), Hitchcock (1871), Hitchcock (1889), Lyman (1894), Hitchcock (1898), Hay (1902), Lull (1904), Lull (1915), Lull (1917), Hitchcock and Grier (1927), Hay (1930), Lull (1953), Kuhn (1963), Demathieu (1970), Haubold (1971), Welles (1971) and Thulborn and Wade (1984); it was considered a nomen dubium by Baird (1957); it was synonymized subjectively with Grallator tuberosus by Weems (1992).
Plesiornis mirabilis was named by Hitchcock (1865). It is considered to be a form taxon. Its type specimen is AC 51/16, a footprint, and it is a trace fossil. Its type locality is Lily Pond Quarry, Gill, which is in a Hettangian fluvial-lacustrine siltstone/sandstone in the Turners Falls Formation of Massachusetts.

It was synonymized subjectively with Anchisauripus tuberosus by Haubold (1971).
Brontozoum exsertum was named by Hitchcock (1858). It is considered to be a form taxon. Its type specimen is AC 16/6, a footprint, and it is a trace fossil. Its type locality is Ashael Lyman's, Northampton tracksite, which is in a Hettangian/Sinemurian lacustrine - small sandstone/mudstone in the Portland Formation of Massachusetts.

It was recombined as Eubrontes exsertus by Hay (1902); it was recombined as Anchisauripus exsertus by Lull (1904), Lull (1915), Lull (1917), Hay (1930), Lull (1953), Kuhn (1963), Haubold (1969), Haubold (1971), Welles (1971), Haubold (1974), Thulborn and Wade (1984) and Weems (1987); it was considered a nomen dubium by Baird (1957); it was misspelled as Anchisauripus exertus by Courel and Demathieu (1976); it was synonymized subjectively with Grallator tuberosus by Weems (1992).
Otouphepus magnificus was named by Cushman (1904). It is considered to be a form taxon. Its type specimen is BSNH 12857, a footprint, and it is a trace fossil. Its type locality is Lily Pond Quarry, Gill, which is in a Hettangian fluvial-lacustrine siltstone/sandstone in the Turners Falls Formation of Massachusetts.

It was synonymized subjectively with Anchisauripus sillimani by Haubold (1971); it was recombined as Grallator magnificus by Olsen and Galton (1984); it was considered a nomen dubium by Baird (1957); it was synonymized subjectively with Grallator tuberosus by Weems (1992).
Brontozoum validum was named by Hitchcock (1858). It is considered to be a form taxon. It is a trace fossil.

It was synonymized subjectively with Eubrontes tuberosus by Hay (1902); it was synonymized subjectively with Anchisauripus tuberosus by Lull (1904), Lull (1915), Hay (1930) and Haubold (1971).
Brontozoum loxonyx was named by Hitchcock (1848). It is considered to be a form taxon. It is a trace fossil.

It was synonymized subjectively with Brontozoum validum by Hitchcock (1858); it was synonymized subjectively with Anchisauripus tuberosus by Lull (1904), Lull (1915) and Haubold (1971).
Coelurosaurichnus metzneri was named by Heller (1952). It is considered to be a form taxon. Its type specimen is Geologisches Institut Erlangen, a footprint, and it is a trace fossil. Its type locality is Altselingsback tracksite, which is in a Carnian terrestrial sandstone in the Middle Keuper Formation of Germany.

It was synonymized subjectively with Grallator tuberosus by Weems (1992); it was synonymized subjectively with Atreipus milfordensis by Hunt and Lucas (2007); it was recombined as Atreipus metzneri by Olsen and Baird (1986), Thulborn (1990), Thulborn (1993), Gand et al. (2000), D'Orazi Porchetti et al. (2008) and Klein and Lucas (2021).
Gigandipus milfordensis was named by Bock (1952) [originally assiged to Gigandipus? (Anchisauripus) milfordensis, although Anichisauripus is here meant as a possible alternative assignment, not a subgenus.]. It is considered to be a form taxon. Its type specimen is LC S488, a footprint, and it is a trace fossil. Its type locality is Smith Clark's quarry, Milford (level B), which is in a Norian terrestrial shale/sandstone in the Passaic Formation of New Jersey.

It was recombined as Anchisauripus milfordensis by Kuhn (1963), Demathieu (1970), Haubold (1971), Olsen (1980), Olsen (1981) and Lockley (1986); it was synonymized subjectively with Grallator tuberosus by Weems (1992); it was recombined as Atreipus milfordensis by Olsen and Baird (1986), Olsen and Baird (1986), Thulborn (1990), Thulborn (1993), Lockley and Hunt (1995), Gand et al. (2000), Szajna and Hartline (2003), Lucas and Sullivan (2006), Weishampel (2006), Thulborn (2006), Weems (2006), Klein and Haubold (2007), Hunt and Lucas (2007), Abbassi and Madanipour (2014) and Klein and Lucas (2021).

Synonymy list

Year	Name and author
1836	Ornithichnites tuberosus Buckland p. 41
1836	Ornithichnites tuberosus Hitchcock p. 318 figs. 2, 5
1837	Ornithichnites gracilis Hitchcock p. 175
1837	Ornithichnites tuberosus Hitchcock p. 175
1837	Ornithoidichnites tuberosus Hitchcock p. 175
1841	Ornithoidichnites tuberosus Hitchcock p. 486
1843	Ornithichnites tuberosus Deane p. 179–180
1843	Ornithoidichnites tuberosus Hitchcock p. 254
1843	Ornithoidichnites tuberosus Redfield p. 136
1844	Ornithoidichnites tuberosus Hitchcock p. 303
1845	Eubrontes tuberosus Hitchcock p. 23
1848	Brontozoum loxonyx Hitchcock p. 172 figs. Pl. 2:1-2
1854	Brontozoum loxonyx Warren p. 27
1858	Brontozoum exsertum Hitchcock p. 67 figs. Pl. 12:1
1858	Brontozoum validum Hitchcock p. 67
1861	Brontozoum validum Deane p. 39
1861	Brontozoum exsertum Deane p. 41
1865	Grallator gracilis Hitchcock p. 8 figs. Pl. 9:7
1865	Plesiornis mirabilis Hitchcock p. 35 figs. Pl. 20
1871	Brontozoum exsertum Hitchcock p. xxi
1871	Brontozoum validum Hitchcock p. xxi
1871	Grallator gracilis Hitchcock p. xxi
1871	Plesiornis mirabilis Hitchcock p. xxi
1879	Brontozoum loxonyx Miller p. 147
1889	Brontozoum exsertum Hitchcock p. 118
1889	Brontozoum validum Hitchcock p. 118
1889	Grallator gracilis Hitchcock p. 118
1889	Plesiornis mirabilis Hitchcock p. 118
1894	Brontozoum exsertum Lyman p. 205
1894	Brontozoum validum Lyman p. 205
1894	Grallator gracilis Lyman p. 205
1894	Plesiornis mirabilis Lyman p. 205
1895	Plesiornis mirabilis Lyman figs. Pl. 608
1898	Brontozoum exsertum Hitchcock p. 401
1898	Brontozoum validum Hitchcock p. 401
1898	Grallator gracilis Hitchcock p. 401
1902	Eubrontes exsertus Hay p. 543
1902	Eubrontes tuberosus Hay p. 544
1902	Grallator gracilis Hay p. 545
1902	Plesiornis mirabilis Hay p. 550
1904	Otouphepus magnificus Cushman p. 155 figs. Pl. 6
1904	Plesiornis mirabilis Cushman p. 156
1904	Anchisauripus tuberosus Lull pp. 488-489 fig. 6
1904	Anchisauripus exsertus Lull p. 489 fig. 7
1904	Grallator gracilis Lull pp. 496-497
1904	Plesiornis mirabilis Lull p. 522
1915	Anchisauripus tuberosus Lull p. 182 fig. 40
1915	Anchisauripus exsertus Lull p. 184 fig. 41
1915	Otouphepus magnificus Lull p. 190 fig. 44
1915	Grallator gracilis Lull p. 202 fig. 55
1917	Anchisauripus exsertus Lull p. 114
1917	Anchisauripus tuberosus Lull p. 114
1917	Otouphepus magnificus Lull p. 115
1917	Grallator gracilis Lull p. 116
1927	Anchisauripus tuberosus Hitchcock and Grier p. 187
1927	Grallator gracilis Hitchcock and Grier p. 189
1930	Anchisauripus exsertus Hay p. 3
1930	Anchisauripus tuberosus Hay p. 3
1930	Grallator gracilis Hay p. 13
1930	Otouphepus magnificus Hay p. 17
1930	Plesiornis mirabilis Hay p. 19
1952	Gigandipus milfordensis Bock p. 403 figs. Pl. 43:3
1952	Coelurosaurichnus metzneri Heller pp. 135-137 figs. Pl. 9:2-3
1953	Grallator gracilis Lull p. 155
1953	Plesiornis mirabilis Lull p. 164
1953	Anchisauripus tuberosus Lull pp. 169-171
1953	Anchisauripus exsertus Lull p. 171
1953	Otouphepus magnificus Lull p. 176
1958	Coelurosaurichnus metzneri Kuhn p. 23
1959	Coelurosaurichnus metzneri Schmidt p. 103
1963	Grallator gracilis Kuhn p. 78
1963	Anchisauripus milfordensis Kuhn p. 80
1963	Anchisauripus exsertus Kuhn p. 81
1963	Anchisauripus tuberosus Kuhn p. 81
1963	Plesiornis mirabilis Kuhn p. 83
1963	Coelurosaurichnus metzneri Kuhn p. 87
1965	Anchisauripus tuberosus Nelson p. 7
1965	Otouphepus magnificus Nelson p. 7
1969	Anchisauripus exsertus Haubold p. 100 fig. 8
1970	Anchisauripus milfordensis Demathieu p. 184
1970	Grallator gracilis Demathieu p. 184
1970	Otouphepus magnificus Ellenberger p. 349
1971	Anchisauripus exsertus Haubold p. 65
1971	Anchisauripus milfordensis Haubold p. 65
1971	Anchisauripus tuberosus Haubold p. 66
1971	Grallator gracilis Haubold p. 67
1971	Coelurosaurichnus metzneri Haubold p. 69
1971	Anchisauripus exsertus Welles p. 30
1971	Anchisauripus tuberosus Welles p. 30
1971	Grallator gracilis Welles p. 31
1972	Coelurosaurichnus metzneri Demathieu and Haubold p. 806
1972	Otouphepus magnificus Ellenberger p. 152
1974	Anchisauripus exsertus Haubold p. 128
1974	Anchisauripus tuberosus Haubold p. 128
1976	Anchisauripus exertus Courel and Demathieu p. 209
1980	Anchisauripus milfordensis Olsen p. 29
1981	Anchisauripus milfordensis Olsen p. 558
1984	Grallator magnificus Olsen and Galton p. 109
1984	Anchisauripus exsertus Thulborn and Wade p. 449
1984	Grallator gracilis Thulborn and Wade p. 449
1986	Anchisauripus milfordensis Lockley p. 9
1986	Anchisauripus tuberosus Lockley p. 19
1986	Atreipus milfordensis Olsen and Baird p. 22
1986	Atreipus metzneri Olsen and Baird p. 69
1987	Anchisauripus exsertus Weems p. 15
1990	Atreipus metzneri Thulborn p. 157
1990	Atreipus milfordensis Thulborn p. 157
1992	Grallator tuberosus Weems p. 124
1993	Atreipus metzneri Thulborn p. 463
1993	Atreipus milfordensis Thulborn p. 463
1995	Atreipus milfordensis Lockley and Hunt p. 80
1996	Grallator tuberosus Gierlinski p. 120
2000	Atreipus metzneri Gand et al. p. 615
2000	Atreipus milfordensis Gand et al. p. 615
2000	Grallator tuberosus Kordos p. 16
2001	Eubrontes tuberosus Mayor and Sarjeant p. 151
2003	Atreipus milfordensis Szajna and Hartline p. 266
2006	Atreipus milfordensis Lucas and Sullivan p. 252
2006	Atreipus milfordensis Thulborn p. 291 fig. 8
2006	Atreipus milfordensis Weems p. 376
2006	Atreipus milfordensis Weishampel p. 134
2007	Atreipus milfordensis Hunt and Lucas p. 222
2007	Atreipus milfordensis Klein and Haubold p. 125–126
2008	Atreipus metzneri D'Orazi Porchetti et al. p. 282
2008	Anchisauripus tuberosus Fujita et al. p. 42
2011	Grallator tuberosus Osi et al. p. 91
2014	Atreipus milfordensis Abbassi and Madanipour p. 113
2019	Grallator tuberosus Li et al. p. 7
2021	Atreipus metzneri Klein and Lucas p. 6
2021	Atreipus milfordensis Klein and Lucas p. 6

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Rank	Name	Author
kingdom	Animalia	()
—	Bilateria
—	Eubilateria	Ax 1987
—	Deuterostomia
phylum	Chordata	Haeckel 1874
subphylum	Vertebrata
superclass	Gnathostomata
—	Osteichthyes	()
subclass	Sarcopterygii	()
subclass	Dipnotetrapodomorpha	(Nelson 2006)
subclass	Tetrapodomorpha	()
—	Tetrapoda
—	Reptiliomorpha
—	Anthracosauria
subclass	Amphibiosauria	Kuhn 1967
—	Cotylosauria	()
—	Amniota
—	Sauropsida

Rank	Name	Author
class	Reptilia
subclass	Eureptilia	()
—	Romeriida
—	Diapsida	()
—	Archosauromorpha	(Huene 1946)
—	Crocopoda
—	Archosauriformes	Gauthier 1986
—	Eucrocopoda
—	Archosauria	()
informal	Avemetatarsalia
—	Ornithodira
—	Dinosauromorpha
—	Dinosauriformes
—	Dinosauria	()
—	Theropoda	()
family	Grallatoridae
genus	Grallator
species	tuberosus	()

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.

†Grallator tuberosus Hitchcock 1836

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Invalid names: Anchisauripus exsertus Hitchcock 1858 [synonym], Atreipus metzneri Heller 1952 [synonym], Atreipus milfordensis Bock 1952 [synonym], Brontozoum loxonyx Hitchcock 1848 [synonym], Brontozoum validum Hitchcock 1858 [synonym], Grallator gracilis Hitchcock 1837 [synonym], Otouphepus magnificus Cushman 1904 [synonym], Plesiornis mirabilis Hitchcock 1865 [synonym]

Diagnosis

Reference	Diagnosis
R. S. Lull 1915	Pes: Length of digit II, 86 mm.; III, 128 mm.; IV, 120 mm.; foot exclusive of hallux, 168 mm., with hallux, ?195 mm. Divarication of digits I and II, ?135°; II and III, 14°; II and IV, 13°. Length of step, 440 mm.
R. S. Lull 1915 (Anchisauripus exsertus)	Pes: Based upon Cat. No. 54/1, Amherst collection (plesiotype). Length of digit I, 75 mm.; II, 117 mm.; III, 161 mm.; IV, 153 mm.; foot exclusive of hallux, 226 mm., including hallux, 265 mm. Divarication of digits I and II, 75°; II and III, 20°; II and IV, 20°. Length of step in type No. 16/6, a somewhat larger specimen, 922 mm.
R. S. Lull 1915 (Otouphepus magnificus)	Pes: Length of digit II, 83 mm.; III, 114 mm.; IV, 114 mm.; entire foot, 165 mm. Distance between lateral tips, 89 mm. Divarication of digits II and III, 22°; II and IV, 17° approximately. Length of step not known.