PBDB Taxon

Basic info	Taxonomic history	Classification	Included Taxa
Morphology	Ecology and taphonomy	External Literature Search	Age range and collections

Grallatoridae

Reptilia - Grallatoridae

Taxonomy

Grallatoridae was named by Lull (1904). It is considered to be a form taxon.

It was synonymized subjectively with Anchisauripodidae by Kuhn (1963), Haubold (1971).

It was assigned to Coelurosauria by Bock (1952), Lull (1953), Baird (1957); to Coelurosauria by Kuhn (1958), Welles (1971); to Trisauropodia by Ellenberger (1974); to Theropoda by Hitchcock and Grier (1927), Kuhn (1966), Gierlinski (1991), Mietto et al. (2000); to Carnosauria by Li et al. (2010); and to Theropoda by Lull (1904), Lull (1915), Carrano (2025).

Subtaxa

Anatopus, Anchisauripus, Atreipus (syn. Tridactylus), Banisterobates, Dilophosauripus, Eubrontes (syn. Paracoelurosaurichnus, Gigandipus, Chonglongpus, Gigantitherium, Tuojiangpus, Fulicopus, Amblonyx, Ichnites, Ornithichnites, Youngichnus, Brontozoum, Lufengopus, Weiyuanpus, Hyphepus, Megaichnites, Zizhongpus, Otouphepus), Grallator (syn. Aetonychopus, Prototrisauropus, Qemetrisauropus, Plastisauropus, Platysauropus, Chuanchengpus, Agialopous, Kainotrisauropus, Chongqingpus), Iguanodonopus, Pengxianpus

Synonyms

Iguanodonopus was named by Zhen et al. (1996). It is considered to be a form taxon.

It was considered a nomen nudum by Xing et al. (2009); it was considered a nomen dubium by Lockley et al. (2013), Lockley et al. (2014).
Iguanodonopus xingfuensis was named by Zhen et al. (1995). It is considered to be a form taxon. Its type specimen is CFEC-E-1a, b, a footprint, and it is a trace fossil.

It was recombined as Iguanodontipus xingfuensis by Xing et al. (2009), Xing et al. (2011); it was considered a nomen dubium by Lockley et al. (2013), Díaz-Martínez et al. (2015).
Anchisauropodidae was named by Lull (1904). It is considered to be a form taxon.

It was synonymized subjectively with Grallatoridae by Baird (1957).
Anatopus was named by Lapparent and Montenat (1967). It is considered to be a form taxon. Its type is Anatopus palmatus.

It was considered a nomen dubium by Moreau et al. (2024).

It was assigned to Saurischia by Haubold (1971); to Ornithischia by Haubold (1974); to Ornithopoda by Lapparent and Montenat (1967), Demathieu and Haubold (1972), Vignaud et al. (1994); and to Theropoda by Romero Molina et al. (2003).
Anatopus palmatus was named by Lapparent and Montenat (1967). It is considered to be a form taxon. Its type specimen is ULB-04C15_B, a footprint, and it is a trace fossil. It is the type species of Anatopus.

It was considered a nomen dubium by Moreau et al. (2024).

Synonymy list

Year	Name and author
1904	Anchisauropodidae Lull
1904	Grallatoridae Lull p. 494
1915	Grallatoridae Lull p. 200
1927	Grallatoridae Hitchcock and Grier p. 186
1952	Grallatoridae Bock p. 402
1953	Grallatoridae Lull p. 152
1957	Grallatoridae Baird p. 453
1958	Grallatoridae Kuhn p. 22
1966	Grallatoridae Kuhn p. 96
1967	Anatopus Lapparent and Montenat pp. 27-28 fig. 16
1967	Anatopus palmatus Lapparent and Montenat p. 28 fig. 16
1967	Anatopus Lapparent and Montenat p. 33
1968	Anatopus palmatus Courel et al. p. 277
1971	Anatopus Haubold p. 81
1971	Anatopus palmatus Haubold p. 81
1971	Grallatoridae Welles p. 29
1972	Anatopus Demathieu and Haubold p. 807
1972	Anatopus palmatus Demathieu and Haubold p. 807
1974	Grallatoridae Ellenberger p. 73
1974	Anatopus Haubold p. 46
1974	Anatopus palmatus Haubold p. 119
1991	Grallatoridae Gierlinski p. 137
1994	Anatopus Vignaud et al. p. 497
1994	Anatopus palmatus Vignaud et al. p. 497
1995	Iguanodonopus xingfuensis Zhen and et al.
1996	Iguanodonopus Zhen and et al.
2000	Anatopus palmatus Gand et al. p. 617
2000	Grallatoridae Mietto et al. p. 66
2003	Anatopus tersi Demathieu p. 59
2003	Anatopus Romero Molina et al. p. 24
2003	Anatopus tersi Viaud and Duclous p. 4
2009	Iguanodontipus xingfuensis Xing et al. pp. 832-833
2010	Grallatoridae Li et al. p. 729
2011	Iguanodontipus xingfuensis Xing et al. p. 478
2025	Grallatoridae Carrano
2025	Anatopus palmatus Foster et al. p. 279

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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
family	Grallatoridae

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.

Fm. †Grallatoridae Lull 1904

show all | hide all

G. †Anchisauripus Lull 1904

Subg. †Anchisauripus (Grallator) Olsen 1981

†Anchisauripus australis Lull 1942

†Anchisauripus bibractensis Demathieu 1971

†Anchisauripus gwyneddensis Bock 1952

†Anchisauripus poolei Bock 1952

†Anchisauripus thomasi Sollas 1879

G. †Atreipus Olsen and Baird 1986

Invalid names: Tridactylus Biron and Dutuit 1981 [synonym]

G. †Banisterobates Fraser and Olsen 1996

†Banisterobates boisseaui Fraser and Olsen 1996

G. †Dilophosauripus Welles 1971

†Dilophosauripus williamsi Welles 1971

G. †Eubrontes Hitchcock 1845

Subg. †Eubrontes (Grallator) Olsen 1981

†Eubrontes euskelosauroides Huene 1941

†Eubrontes expansus Hitchcock 1841

†Eubrontes fieldi Hitchcock 1858

†Eubrontes giganteus Hitchcock 1836

Invalid names: Eubrontes approximatus Hitchcock 1865 [synonym], Eubrontes divaricatus Hitchcock 1865 [synonym], Eubrontes veillonensis Lapparent and Montenat 1967 [synonym]

†Eubrontes glenrosensis Shuler 1935

†Eubrontes lyellii Hitchcock 1843

†Eubrontes minusculus Hitchcock 1837

Invalid names: Eubrontes tuberatus Hitchcock 1858 [synonym], Grallator maximus Lapparent and Montenat 1967 [synonym], Plesiornis giganteus Hitchcock 1889 [synonym]

†Eubrontes monax Zhen et al. 1986

†Eubrontes nianpanshanensis Yang and Yang 1987

†Eubrontes nobitai Xing et al. 2021

†Eubrontes pareschequier Xing et al. 2009

†Eubrontes platypus Hitchcock 1858

†Eubrontes sillimani Hitchcock 1843

Invalid names: Anchisauripus dananus Hitchcock 1845 [synonym], Ornithichnites cuneatus Hitchcock 1837 [synonym], Otouphepus minor Lull 1915 [synonym]

†Eubrontes tatricus Michalík and Sykora 1976

†Eubrontes zigongensis Gao 2007

Invalid names: Brontozoum Hitchcock 1847 [synonym], Eubrontes xiyangensis Zhen et al. 1986 [nomen dubium], Fulicopus Hitchcock 1845 [synonym], Gigandipus Hitchcock 1855 [synonym], Hyphepus Hitchcock 1858 [synonym], Ichnites Haubold 1971 [synonym], Lufengopus Lü et al. 2006 [synonym], Lufengopus dongi Lü et al. 2006 [nomen dubium], Megaichnites Yang and Yang 1987 [synonym], Megaichnites jizhaoshiensis Yang and Yang 1987 [nomen dubium], Ornithichnites Hitchcock 1836 [synonym], Otouphepus Cushman 1904 [synonym], Paracoelurosaurichnus Zhen et al. 1986 [synonym], Tuojiangpus Yang and Yang 1987 [synonym], Tuojiangpus shuinanensis Yang and Yang 1987 [nomen dubium], Weiyuanpus Gao 2007 [synonym], Youngichnus Zhen et al. 1986 [synonym], Zizhongpus Yang and Yang 1987 [synonym], Zizhongpus wumaensis Yang and Yang 1987 [nomen dubium]

G. †Grallator Hitchcock 1858

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 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]

G. †Pengxianpus Yang and Yang 1987

†Pengxianpus cifengensis Yang and Yang 1987

†Pengxianpus yulinensis Wang et al. 2016

Invalid names: Anatopus Lapparent and Montenat 1967 [nomen dubium], Anchisauropodidae Lull 1904 [synonym], Iguanodonopus Zhen 1996 [nomen dubium]

Diagnosis

Reference	Diagnosis
R. S. Lull 1904	Small bipedal forms; footprint tridactyl, limbs very long; no manus nor caudal impression.

Measurements

No measurements are available

Composition:

hydroxyapatiteo

Entire body:

yeso

Adult length:

10 to < 100o

Adult width:

1.0 to < 10o

Adult height:

1.0 to < 10o

Architecture:

compact or denseo

Ontogeny:

accretion, modification of partso

Grouping:

solitaryo

Environment:

terrestrialo

Locomotion:

actively mobileo

Life habit:

ground dwellingo

Diet:

carnivoreo

Reproduction:

oviparouso

Dispersal:

direct/internalo

Dispersal 2:

mobileo

Created:

2004-12-14 12:50:08

Modified:

2004-12-14 14:50:08

Source: o = order

Reference: Marsh 1875

Age range: base of the Smithian to the top of the Campanian or 249.90000 to 72.20000 Ma

Collections (462 total)

Time interval	Ma	Country or state	Original ID and collection number
Oldest occurrences
Smithian	249.9 - 248.1	USA (Utah)	Grallator sp., Eubrontes sp. (105542)
Anisian	246.7 - 241.464	Morocco (Marrakech-Safi)	Atreipus sp. (199901)
Anisian - Ladinian	246.7 - 237.0	France (Rhône-Alpes)	Grallator sp. (199929)
Anisian - Ladinian	246.7 - 237.0	France (Bourgogne-Franche-Comté)	Anchisauripus bibractensis (62183 68222 68906 199909 199928) Anchisauripus sp. (28462) Anchisauripus sp., Anchisauripus bibractensis (62182 68226) Dinosauria indet. (24834)
Anisian - Ladinian	246.7 - 237.0	France (Auvergne-Rhône-Alpes)	Anchisauripus sp. (31310)
Middle Triassic	246.7 - 237.0	France (Bourgogne-Franche-Comté)	Anchisauripus bibractensis (28092)
Carnian	237.0 - 227.3	Canada (Nova Scotia)	Atreipus acadianus, Anchisauripus milfordensis (55416)
Carnian	237.0 - 227.3	Morocco (Marrakech-Safi)	Dinosauria indet. (34701) Eubrontes sp. (205518 230183) Grallator sp. (230179 230180 230181 230182) Grallatoridae indet. (195710 205534) Grallatoridae indet., Eubrontes sp. (205528)
Carnian	237.0 - 227.3	Italy (Toscana)	Grallator pisanus (68779)
Carnian	237.0 - 227.3	France	Grallator sp. (55316 68224)
Youngest occurrences
Time interval	Ma	Country or state	PBDB collection number
Early/Lower Albian	113.2 - 110.1	USA (Texas)	Dinosauria indet. (52540) Eubrontes sp. (51421) Irenesauripus glenrosensis (57674) Irenesauripus sp. (52148) Theropoda indet. (52339 52532 52535 52537 52538)
Albian	113.2 - 100.5	South Korea (Gyeongsangnam-do)	Grallator sp. (219899)
Albian	113.2 - 100.5	USA (New Mexico)	Theropoda indet. (52161)
Albian	113.2 - 100.5	China (Shandong)	Grallator sp. (219854)
Albian - Early/Lower Cenomanian	113.2 - 93.9	United Kingdom (England)	Ornithoidichnites giganteus (232275)
Albian - Coniacian	113.2 - 85.7	China (Fujian)	Grallatoridae indet. (229370)
Early/Lower Cenomanian	100.5 - 93.9	China (Jilin)	Theropoda indet. (55193)
Late/Upper Cenomanian - Turonian	100.5 - 89.8	Argentina (Neuquén)	Coelurosauria indet. (85571)
Campanian	83.6 - 72.2	USA (Colorado)	Grallator sp. (156036)
Late/Upper Campanian - Early/Lower Maastrichtian	83.6 - 66.0	USA (Alaska)	Grallator sp. (44848)