PBDB Taxon

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Grallator

Reptilia - Grallatoridae

Taxonomy

Grallator was named by Hitchcock (1858). It is considered to be a form taxon.

It was synonymized subjectively with Eubrontes by Rainforth (2007).

It was assigned to Pachydactyli by Quenstedt (1882); to Aves by Hitchcock (1898); to Theropoda by Lull (1917); to Coelurosauria by Lapparent and Montenat (1967); to Trisauropodia by Ellenberger (1972); to Coelurosauria by Taquet (1977) and Vignaud et al. (1994); to Anchisauripodidae by Kuhn (1963), Haubold (1969), Haubold (1971), Demathieu and Haubold (1972), Haubold (1974), Zhen et al. (1986), Haderer (1988), Zhen et al. (1989) and Leonardi and Mietto (2000); to Anchisauropodidae by Peng et al. (2005); to Theropoda by Madsen and Miller (1979), Lockley (1992), Hunt et al. (1998), Romero Molina et al. (2003) and Gierlinski et al. (2009); to Theropoda by Lucas et al. (2006), Weishampel (2006) and Niedźwiedzki (2011); and to Grallatoridae by Lull (1904), Lull (1915), Hitchcock and Grier (1927), Bock (1952), Lull (1953), Baird (1954), Baird (1957), Kuhn (1958), Olsen and Galton (1984), Gierlinski (1991), Gierlinski and Ahlberg (1994), Gierlinski (1996), Lucas et al. (2006), Dalman (2012), Tsukiji et al. (2019), Castanera et al. (2020) and Klein and Lucas (2021).

Species

Agialopous wyomingensis, Chongqingpus microiscus, Chuanchengpus wuhuangensis, G. (Grallator), G. andeolensis, G. angustidigitus, G. anietangensis, G. anserinum, G. crassidigitus, G. cursorius (syn. Ornithichnites parallelus, Atreipus sulcatus), G. damanei, G. deambulator, G. digitigradus, G. emeiensis, G. graciosus, G. lacunensis, G. lambereshei, G. leribeensis, G. lescurei, G. limnosus, G. limosus, G. matsiengensis, G. maximus, G. minimus, G. minor, G. mokanametsongensis, G. molapoi, G. morijensis, G. moshoeshoei, G. olonensis, G. palustris, G. pingchuanensis, G. princeps, G. rapidus, G. rectilineus, G. rectus, G. sauclierensis, G. socialis, G. ssatoi, G. tenuis (syn. G. cuneatus, G. formosus, Anchisauripus hitchcocki, Atreipus acadianus), G. tuberosus (syn. Atreipus metzneri, Brontozoum validum, Anchisauripus exsertus, Plesiornis mirabilis, Atreipus milfordensis, G. gracilis, Otouphepus magnificus, Brontozoum loxonyx), G. variabilis, G. yangi, G. yemiaoxiensis, Plastisauropus ingens, Platysauropus robustus, Prototrisauropus deambulator, Prototrisauropus gardonensis

Species lacking formal opinion data

G. pisanus

Synonyms

Qemetrisauropus was named by Ellenberger (1970). It is considered to be a form taxon. Its type is Qemetrisauropus princeps.

It was synonymized subjectively with Grallator by Olsen and Galton (1984); it was synonymized subjectively with Eubrontes by Lucas et al. (2006) and Klein and Lucas (2021).

It was assigned to Prosauropoda by Ellenberger (1970); to Trisauropodia by Ellenberger (1972) and Ellenberger (1974); and to Theropoda by Romero Molina et al. (2003).
Prototrisauropus was named by Ellenberger (1965). It is considered to be a form taxon. Its type is Prototrisauropus crassidigitus.

It was synonymized subjectively with Eubrontes by Lucas et al. (2006); it was synonymized subjectively with Grallator by Olsen and Galton (1984) and Klein and Lucas (2021).

It was assigned to Prosauropoda by Ellenberger (1970); to Trisauropodia by Ellenberger (1972); and to Theropoda by Romero Molina et al. (2003).
Platysauropus was named by Ellenberger (1970).

It was replaced with Plastisauropus by Ellenberger (1974).

It was assigned to Thecodontosauria by Ellenberger (1972).
Kainotrisauropus was named by Ellenberger (1970).

It was synonymized subjectively with Grallator by Olsen and Galton (1984).

It was assigned to Plateosauridae by Ellenberger (1970); to Trisauropodia by Ellenberger (1974); and to Theropoda by Knoll (2005).
Saurichnium anserinum was named by Gürich (1926). It is considered to be a form taxon. It is a trace fossil. Its type locality is Otjihaene-Maparero 92 Farm tracksite (ONP I), which is in a Jurassic eolian sandstone in the Etjo Formation of Namibia.

It was considered a nomen dubium by Wagensommer et al. (2016).
Aetonychopus was named by Ellenberger (1972). It is considered to be a form taxon.

It was synonymized subjectively with Grallator by Olsen and Galton (1984).

It was assigned to Coelurosauria by Ellenberger (1972); to Paleosauridae by Ellenberger (1974); and to Theropoda by Romero Molina et al. (2003).
Plastisauropus was named by Ellenberger (1972). It is considered to be a form taxon.

It was synonymized subjectively with Grallator by Olsen and Galton (1984).

It was assigned to Carnosauria by Ellenberger (1972); to Paleosauridae by Ellenberger (1974); and to Theropoda by Romero Molina et al. (2003).
Chongqingpus was named by Yang and Yang (1987). It is considered to be a form taxon.

It was synonymized subjectively with Kayentapus by Xing et al. (2013); it was synonymized subjectively with Grallator by Xing et al. (2021).

It was assigned to Anchisauripodidae by Yang and Yang (1987).
Chongqingpus microiscus was named by Yang and Yang (1987). It is considered to be a form taxon. Its type specimen is CFZW176, a footprint, and it is a trace fossil. Its type locality is Wu Ma Cun, site B (large slab), which is in an Aalenian/Bajocian terrestrial horizon in the Xintiangou Formation of China.

It was considered a nomen dubium by Xing et al. (2021); it was recombined as Grallator microiscus by Lockley et al. (2013), Xing et al. (2013), Xing et al. (2017) and Li et al. (2022).
Chuanchengpus was named by Yang and Yang (1987). It is considered to be a form taxon.

It was synonymized subjectively with Grallator by Lockley et al. (2013) and Xing et al. (2021).

It was assigned to Anomoepodidae by Yang and Yang (1987).
Chuanchengpus wuhuangensis was named by Yang and Yang (1987). It is considered to be a form taxon. Its type specimen is CFZW101, a footprint, and it is a trace fossil. Its type locality is Wu Ma Cun, site A (Rice cleaning), which is in an Aalenian/Bajocian terrestrial sandstone in the Xintiangou Formation of China.

It was considered a nomen dubium by Xing et al. (2021); it was recombined as Grallator wuhuangensis by Lockley et al. (2013), Xing et al. (2017) and Li et al. (2021).
Agialopous was named by Branson and Mehl (1932). Its type is Agialopous wyomingensis.

It was synonymized subjectively with Grallator by Hunt et al. (1998), Heckert and Lucas (2000), Hunt and Lucas (2007) and Milner et al. (2009).

It was assigned to Grallatoridae by Branson and Mehl (1932); to Saurischia by Kuhn (1958) and Kuhn (1963); to Parachirotheriidae by Haubold (1969), Haubold (1971) and Haubold (1974); and to Theropoda by Romero Molina et al. (2003).

Synonymy list

Year	Name and author
1858	Grallator Hitchcock p. 72
1882	Grallator Quenstedt p. 131
1898	Grallator Hitchcock p. 401
1904	Grallator Lull p. 494
1915	Grallator Lull p. 200
1917	Grallator Lull p. 116
1926	Saurichnium anserinum Gürich p. 117
1927	Grallator Hitchcock and Grier p. 186
1932	Agialopous Branson and Mehl
1952	Grallator Bock p. 402
1953	Grallator Lull p. 153
1954	Grallator Baird p. 182
1957	Grallator Baird p. 453
1958	Grallator Kuhn p. 22
1958	Agialopous Kuhn p. 24
1963	Grallator Kuhn p. 77
1963	Agialopous Kuhn p. 89
1965	Prototrisauropus Ellenberger
1967	Grallator Lapparent and Montenat p. 33
1969	Agailopous Haubold p. 98
1969	Grallator Haubold p. 99
1970	Qemetrisauropus Ellenberger p. 344
1970	Prototrisauropus Ellenberger p. 345
1970	Platysauropus Ellenberger p. 348
1970	Kainotrisauropus Ellenberger p. 350
1971	Agialopous Haubold p. 64
1971	Grallator Haubold p. 66
1971	Saurichnium anserinum Haubold p. 82
1972	Grallator Demathieu and Haubold p. 806
1972	Aetonychopus Ellenberger p. 152
1972	Grallator Ellenberger p. 152
1972	Plastisauropus Ellenberger p. 152
1972	Platysauropus Ellenberger p. 152
1972	Prototrisauropus Ellenberger p. 152
1972	Qemetrisauropus Ellenberger p. 152
1974	Qemetrisauropus Ellenberger p. 16
1974	Aetonychopus Ellenberger p. 17
1974	Plastisauropus Ellenberger p. 17
1974	Kainotrisauropus Ellenberger p. 73
1974	Agialopous Haubold p. 40
1974	Grallator Haubold p. 43
1977	Grallator Taquet p. 326
1979	Grallator Madsen and Miller p. 36
1984	Grallator Olsen and Galton p. 97
1986	Grallator Zhen et al. p. 2
1987	Chongqingpus Yang and Yang p. 13 fig. 7
1987	Chongqingpus microiscus Yang and Yang p. 15 fig. 9
1987	Chuanchengpus Yang and Yang p. 23–24 fig. 21
1987	Chuanchengpus wuhuangensis Yang and Yang p. 23–24 fig. 21
1988	Grallator Haderer p. 8
1989	Grallator Zhen et al. p. 187
1991	Grallator Gierlinski p. 137
1992	Grallator Lockley p. 272
1994	Grallator Gierlinski and Ahlberg p. 101
1994	Grallator Vignaud et al. p. 497
1996	Grallator Gierlinski p. 120
1998	Grallator Hunt et al. p. 522
2000	Grallator Leonardi and Mietto p. 197
2003	Aetonychopus Romero Molina et al. p. 24
2003	Agialopous Romero Molina et al. p. 24
2003	Grallator Romero Molina et al. p. 24
2003	Plastisauropus Romero Molina et al. p. 24
2003	Prototrisauropus Romero Molina et al. p. 24
2003	Qemetrisauropus Romero Molina et al. p. 24
2005	Kainotrisauropus Knoll p. 84
2005	Grallator Peng et al.
2006	Grallator Lucas et al. p. 86–87
2006	Grallator Lucas et al. p. 114
2006	Grallator Weishampel p. 132
2009	Grallator Gierlinski et al. p. 227
2009	Chongqingpus microiscus Xing et al. p. 23
2011	Grallator Niedźwiedzki p. 295
2012	Grallator Dalman p. 336
2013	Grallator microiscus Lockley et al. p. 6
2013	Grallator wuhuangensis Lockley et al. p. 6
2013	Grallator microiscus Xing et al. p. 119
2017	Grallator microiscus Xing et al. p. 120
2017	Grallator wuhuangensis Xing et al. p. 120
2019	Grallator Tsukiji et al. p. 4
2020	Grallator Castanera et al. p. 279
2021	Grallator Klein and Lucas p. 13
2021	Grallator wuhuangensis Li et al. p. 14
2022	Grallator microiscus Li et al. p. 4233

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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

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.

G. †Grallator Hitchcock 1858

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Subg. †Grallator (Grallator) Olsen and Galton 1984

†Grallator andeolensis Gand et al. 2000

†Grallator angustidigitus Ellenberger 1970

†Grallator anietangensis Li et al. 2022

†Grallator crassidigitus Ellenberger 1970

†Grallator cursorius Hitchcock 1858

Invalid names: Ornithichnites parallelus Hitchcock 1837 [synonym]

†Grallator damanei Ellenberger 1970

†Grallator deambulator Ellenberger 1970

†Grallator digitigradus Ellenberger 1972

†Grallator emeiensis Zhen et al. 1994

†Grallator graciosus Ellenberger 1970

†Grallator lacunensis Ellenberger 1970

†Grallator lambereshei Ellenberger 1970

†Grallator leribeensis Ellenberger 1970

†Grallator lescurei Demathieu 1990

†Grallator limnosus Zhen et al. 1986

†Grallator limosus Zhen et al. 1986

†Grallator matsiengensis Ellenberger 1970

†Grallator maximus Lapparent and Montenat 1967

†Grallator minimus Ellenberger 1970

†Grallator minor Lull 1915

†Grallator mokanametsongensis Ellenberger 1974

†Grallator molapoi Ellenberger 1970

†Grallator morijensis Ellenberger 1970

†Grallator moshoeshoei Ellenberger 1970

†Grallator olonensis Lapparent and Montenat 1967

†Grallator palustris Ellenberger 1970

†Grallator pingchuanensis Li et al. 2019

†Grallator princeps Ellenberger 1970

†Grallator rapidus Ellenberger 1972

†Grallator rectilineus Ellenberger 1970

†Grallator rectus Hitchcock and Grier 1927

†Grallator sauclierensis Demathieu and Sciau 1992

†Grallator socialis Ellenberger 1970

†Grallator ssatoi Yabe et al. 1940

†Grallator tenuis Hitchcock 1858

Invalid names: Anchisauripus hitchcocki Lull 1904 [synonym], Atreipus acadianus Olsen and Baird 1986 [synonym], Grallator cuneatus Hitchcock 1858 [synonym], Grallator formosus Hitchcock 1858 [synonym]

†Grallator tuberosus Hitchcock 1836

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]

†Grallator variabilis Lapparent and Montenat 1967

†Grallator yangi Lii and Zhang 2000

†Grallator yemiaoxiensis Yang and Yang 1987

Invalid names: Aetonychopus Ellenberger 1972 [synonym], Agialopous Branson and Mehl 1932 [synonym], Chongqingpus Yang and Yang 1987 [synonym], Chongqingpus microiscus Yang and Yang 1987 [nomen dubium], Chuanchengpus Yang and Yang 1987 [synonym], Grallator anserinum Gürich 1926 [nomen dubium], Kainotrisauropus Ellenberger 1970 [synonym], Plastisauropus Ellenberger 1972 [synonym], Prototrisauropus Ellenberger 1965 [synonym], Qemetrisauropus Ellenberger 1970 [synonym]

Diagnosis

Reference	Diagnosis
P. Ellenberger 1972 (Qemetrisauropus)	Nous groupons au sein de ce genre nouveau plusieurs types de pistes appartenant à des bipèdes tridactyles de taille moyenne à grande, tous de l'époque du Molteno Inférieur. Leur démarche est rectiligne; ils se caractérisent par leur sole pédieuse quasi-symétrique courte et arrondie à l'arrière par suite d'un raccourcissement du IVème doigt dont les 4 phalanges télescopées dessinent un arc de cercle à la convexité tournée vers l'arrière externe du pied la corde faisant un angle de 35° avec l'axe du déplacement; les phalanges du doigt médian forment de leur côté un arc de cercle régulier à la convexité tournée vers l'extérieur de la piste, légèrement à l'arrière, la corde faisant un angle de 6° avec l'axe du déplacement, - Le IIème doigt se montre fort renflé et étalé avec une convexité v,ers l'intérieur de la piste. Ces trois doigts forment un éventail bien ouvert sur l'articulation nettement moins large des métatarsiens qui paraissent ainsi resserrés entre eux.
G. Peng et al. 2005	Three-toed footprints made by small, bipedal coelurosaurs. Each footprint is narrow and pentagonal in outline, with clear impressions of pads and claws. Digit III is much longer than digit II and IV. No impressions of hallux and tail.
S. G. Dalman 2012	Small (average length 2.3 cm), bipedal, functionally tridactyl ichnite. Digit III projects anteriorly beyond the lateral digits, footprint narrower than that of the ichnogenera Anchisauripus, Eubrontes and Kayentapus (average L/W < 2). Metatarsal and hallux impressions rarely but occasionally pres- ent. Average total divarication angle of digits II to IV is 62°. Average pace angle in continuous trackways, up to 156°; average stride length, up to 24.6 cm. Short digit III extension or weak mesaxony with average L/W ratio for anterior triangle between 0.49 and 0.58. Digits moderately long with average length ratios of 1.27 for digits III/II and 1.10 for digits III/IV. Average digit III projection ratio is 1.91.
H. Klein and S. G. Lucas 2021	Small (<150 mm long), bipedal, functionally tridactyl ichnite. Digit III projects relatively farther anteriorly, and the foot is more narrow than in Eubrontes and Anchisauripus (length/width ratio close to or greater than 2). Hallux rarely impressed. Divarication of outer digits 10° to 30°.