Grallator

{{short description|Ichnogenus of dinosaur footprints}} {{Ichnobox | fossil_range = Early Triassic-Early Cretaceous ~{{fossil range|250|100}} | image = Grallatorgrs.jpg | image_caption = Typical footprint form | taxon = Grallator | authority = Hitchcock, 1858 | type_ichnospecies = {{extinct}}''Grallator cursorius'' | type_ichnospecies_authority = Hitchcock, 1858 | subdivision_ranks = Ichnospecies | subdivision = Many, see text | synonyms = * ''Hunanpus'' }}

'''''Grallator''''' ({{respell|GRA|lə|tor}}) is an ichnogenus (form taxon based on footprints) which covers a common type of small, three-toed print made by a variety of bipedal theropod dinosaurs. ''Grallator''-type footprints have been found in formations dating from the Early Triassic through to the early Cretaceous periods. They are found in the United States, Canada, Europe, India, Australia, Brazil (Sousa and Santa Maria Formations) and China,<ref name=Fossilworks>[https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=64079 ''Grallator''] at Fossilworks.org</ref> but are most abundant on the east coast of North America, especially the Triassic and Early Jurassic formations of the northern part of the Newark Supergroup.<ref name="east coast">Weishampel, D. B. & L. Young. 1996. Dinosaurs of the East Coast. The Johns Hopkins University Press</ref><ref name="xingetal2009">Xing Li-da, Harris, J. D., Feng Xiang-yang, and Zhang Zhi-jun. (2009). "Theropod (Dinosauria: Saurischia) tracks from Lower Cretaceous Yixian Formation at Sihetun Village, Liaoning Province, China and possible track makers." ''Geological Bulletin of China'', '''28'''(6): 705–712.</ref> The name ''Grallator'' translates into "stilt walker", although the actual length and form of the trackmaking legs varied by species, usually unidentified. The related term "Grallae" is an ancient name for the presumed group of long-legged wading birds, such as storks and herons. These footprints were given this name by their discoverer, Edward Hitchcock, in 1858.<ref name="east coast"/>

''Grallator'' footprints are characteristically three-toed (tridactyl) and range from 10 to 20 centimeters (or 4 to 8 inches) long. Though the tracks show only three toes, the trackmakers likely had between four and five toes on their feet. While it is usually impossible to match these prints with the exact dinosaur species that left them, it is sometimes possible to narrow down potential trackmakers by comparing the proportions in individual ''Grallator'' ichnospecies with known dinosaurs of the same formation. For example, ''Grallator'' tracks identified from the Yixian Formation may have been left by ''Caudipteryx''.<ref name="xingetal2009"/>

==Species==

thumb|''Grallator toscanus'' from Monte Pisano (Italy) Source:<ref name="paleofile">{{cite web |title= Paleofile |url=http://www.paleofile.com/Demo/Mainpage/Taxalist/Ichnology.htm |access-date=5 April 2011}}</ref>

{{div col|colwidth=22em}} *Subgenus ''G. (Coelurosaurichnus)'' **''G. (C.) palmipes'' ***''G. (C.) p. exiguus'' (Ellenberger, 1970) *Subgenus ''G. (Grallator)'' **''G. (G.) zvierzi'' Gierlinski, 1991 *''G. andeolensis'' Gand, Vianey-Liaud, Demathieu, & Garric, 2000 *''G. angustidigitus'' (Ellenberger, 1970) *''G. angustus'' (Ellenberger, 1974) **''G. a. cursor'' Ellenberger, 1974 *''G. cursorius'' Hitchcock, 1858 (ichnotype) *''G. cuneatus'' Hitchcock, 1858 *''G. damanei'' Ellenberger, 1970 *''G. deambulator'' (Ellenberger, 1970) *''G. digitigradus'' (Ellenberger, 1974) *''G. emeiensis'' Zhen, Li, Han & Yang, 1995 *''G. formosus'' Hitchcock, 1858 *''G. gracilis'' Hitchcock, 1865 *''G. graciosus'' (Ellenberger, 1970) *''G. grancier'' (Courel & Demathieu, 2000) *''G. ingens'' (Ellenberger, 1970) *''G. jiuquwanensis'' (Zeng, 1982) =''Hunanpus'' *''G. kehli'' (Beurlen, 1950) *''G. kronbergeri'' (Rehnelt, 1959) *''G. lacunensis'' (Ellenberger, 1970) *''G. leribeensis'' (Ellenberger, 1970) *''G. limnosus'' Zhen, Li, & Rao, 1985 *''G. madseni'' Irby, 1995 *''G. magnificus'' (Ellenberger, 1970) *''G. matsiengensis'' Ellenberger, 1970 *''G. maximus'' Lapparent & Monetnat, 1967 *''G. minimus'' (Ellenberger, 1970) *''G. minor'' (Ellenberger, 1970) *''G. moeni'' (Beurlen, 1950) *''G. mokanametsongensis'' (Ellenberger, 1974) *''G. molapoi'' Ellenberger, 1974 *''G. morijiensis'' (Ellenberger, 1970) *''G. moshoeshoei'' (Ellenberger, 1970) *''G. olonensis'' Lapparent & Monetnat, 1967 *''G. palissyi'' (Gand, 1976) *''G. paulstris'' (Ellenberger, 1970) *''G. perriauxi'' (Demathieu & Gand, 1972) *''G. pingchuanensis'' Li et al., 2019)<ref name="G.pingchuanensis">{{cite journal |author1=Da-Qing Li |author2=Li-Da Xing |author3=Martin G. Lockley |author4=Anthony Romilio |author5=Jing-Tao Yang |author6=Long-Feng Li |year=2019 |title=The first theropod tracks from the Middle Jurassic of Gansu, Northwest China: new and rare evidence of quadrupedal progression in theropod dinosaurs |journal=Journal of Palaeogeography |volume=8 |issue=1 |article-number=10 |doi=10.1186/s42501-019-0028-4 |doi-access=free |bibcode=2019JPalG...8...10L }}</ref> *''G. princeps'' (Ellenberger, 1970) *''G. protocrassidigitus'' (Ellenberger, 1970) *''G. rapidus'' (Ellenberger, 1974) *''G. romanovskyi'' (Gabunia & Kurbatov) *''G. quthingensis'' (Ellenberger, 1974) *''G. rectilineus'' (Ellenberger, 1970) *''G. sabinensis'' (Gand & Pellier, 1976) *''G. sassendorfensis'' (Kuhn, 1958) *''G. sauclierensis'' Demathieu & Sciau, 1992 *''G. schlauersbachensis'' (Weiss, 1934) *''G. socialis'' (Ellenberger, 1970) *''G. ssatoi'' Yabe, Inai, & Shikama, 1940 *''G. tenuis'' Hitchcock, 1858 *''G. toscanus'' (Huene, 1941) *''G. variabilis'' Lapparent & Monetnat, 1967 {{div col end}}

==Paleopathology== Fossil tracks can be informative about theropod pathologies but apparently pathological traits may be due to unusual behaviors. Sandstone stratum dating to the Norian in southern Wales preserves tracks of an individual with a deformed digit III attributed to the ichnogenus ''Anchisauripus''. The distal end of the digit was consistently flexed. However, this apparent pathology could be caused by the animal rotating the tip of that digit when lifting the foot.<ref name="molnar-pathology">Molnar, R. E., 2001, Theropod paleopathology: a literature survey: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 337-363.</ref>

==Occurrences== ''Grallator''-type footprints have been found in formations dating from the Early Triassic through to the early Cretaceous periods. They are found in the United States, Canada, Europe, Australia, Brazil (Sousa and Santa Maria Formations) and China,<ref name=Fossilworks/> but are most abundant on the east coast of North America, especially the Triassic and Early Jurassic formations of the northern part of the Newark Supergroup.<ref name="east coast"/><ref name="xingetal2009"/>

===Newark Supergroup tracks=== thumb|125px|left|Negative footprint of ''G. cuneatus'' showing skin impressions The most famous, and archetypal tracks that conform to the ''Grallator'' type are those found on the East Coast of North America, specifically from the Late Triassic to Early Jurassic Newark Supergroup. These footprints were likely made by an unidentified, primitive dinosaur similar to ''Coelophysis''.<ref name="east coast"/> The Newark Supergroup footprints show digits II, III and IV, but no trace of the shorter digits I and V which would likely have been present in a dinosaur of this stage. The outer two digits would have been stubby and ineffective, not touching the ground during walking or running.<ref name="east coast"/> Despite losing most of their effectiveness, dinosaur evolution had not yet removed these digits to fully streamline the foot. This is known because rare specimens are found with traces of these outer digits. Digits II, III and IV have 3, 4 and 5 phalanges respectively, giving ''Grallator'' a ?-3-4-5-? digital formula.

Although the Newark Supergroup ''Grallator'' tracks were made by a bipedal saurischian dinosaur, they can easily be mistaken for those of the late Triassic ichnogenus ''Atreipus''.<ref>Safran, J. and Rainforth, E. C. (2004). [http://gsa.confex.com/gsa/2004NE/finalprogram/abstract_70257.htm "Distinguishing the tridactyl dinosaurian ichnogenus ''Atreipus'' and ''Grallator'': where are the latest Triassic Ornithischia in the Newark Supergroup?"] {{Webarchive|url=https://web.archive.org/web/20160307095344/http://gsa.confex.com/gsa/2004NE/finalprogram/abstract_70257.htm |date=7 March 2016 }} Abstracts with Programs, Geological Society of America, '''36'''(2): 96.</ref> The trackmaker of ''Atreipus'' prints was a quadrupedal ornithischian. The reason for this similarity is a lack of divergence in the foot evolution of the two distinct groups of dinosaurs: ornithischians and saurischians. [[File:Eubrontes01.JPG|thumb|''Eubrontes'' (= ''Grallator'') in the Lower Jurassic Moenave Formation at the St. George Dinosaur Discovery Site at Johnson Farm, southwestern Utah.]]

=== Wales === In January 2021, while walking with her father Richard Wilder, a four-year-old girl called Lily Wilder found a 215- to 220-million year-old dinosaur footprint at Bendricks Bay in the Vale of Glamorgan, Wales.<ref name=":0">{{Cite web|title=Paleontologists stunned by a perfectly preserved dinosaur footprint discovered by a 4-year-old girl|url=https://www.businessinsider.in/science/news/paleontologists-stunned-by-a-perfectly-preserved-dinosaur-footprint-discovered-by-a-4-year-old-girl/articleshow/80616363.cms|access-date=1 February 2021|website=Business Insider}}</ref><ref>{{Cite web|last1=Wakefield|first1=Owen|last2=Shaw|first2=Amelia|date=30 January 2021|title=Girl, 4 finds 215 million-year-old dinosaur footprint along the Welsh coast|url=https://www.dailypost.co.uk/news/north-wales-news/girl-4-finds-215-million-19735002|access-date=1 February 2021|website=North Wales Live|language=en}}</ref><ref name=":1">{{Cite web|title=Britain's biggest turn-offs named|url=https://www.theweek.co.uk/news/world-news/951837/britains-biggest-turn-offs-named|access-date=1 February 2021|website=The Week UK|date=February 2021 |language=en}}</ref>

Experts believe that the footprint was most likely left by a dinosaur (grallator) that stood about 75 centimeters (29.5 inches) tall and 2.5 meters (about 8 feet) long and walked on its two hind feet.<ref name=":3">{{Cite web|title=Lily's Dinosaur: 4yo girl discovers 220 million year old dinosaur footprint|url=http://thescienceinsight.com/lilys-dinosaur-4yo-girl-discovers-220-million-year-old-dinosaur-footprint/|access-date=1 February 2021|website=The Science Insight|date=February 2021 |language=en-US}}</ref><ref>{{Cite web|last=Papenfuss|first=Mary|date=30 January 2021|title=Girl Discovers 220-Million-Year-Old Dinosaur Footprint On Beach Walk With Dad|url=https://www.huffpost.com/entry/lily-winkler-dinosaur-footprint-south-wales-beach_n_6016089ec5b6aa4bad34990c|access-date=1 February 2021|website=HuffPost|language=en}}</ref>

The scientists called the girl's discovery "the finest impression of a 215 million-year-old dinosaur print found in Britain in a decade".<ref name=":0" /> Karl-James Langford of Archaeology Cymru considered the find to be "internationally important".<ref name=":0" /><ref>{{Cite web|last=Wakefield|first=Owen|date=29 January 2021|title=Perfect 215-million-year-old dinosaur print found by girl, 4, on beach|url=https://www.walesonline.co.uk/news/wales-news/perfect-215-million-year-old-19731097|access-date=1 February 2021|website=WalesOnline|language=en}}</ref> Cindy Howells, a palaeontologist at Amgueddfa Cymru – National Museum Wales, described it as "one of the best-preserved examples from anywhere in the UK" and said that it "will really aid palaeontologists to get a better idea about how these early dinosaurs walked".<ref name=":1" /><blockquote> ==== Slovakia ==== </blockquote>In the 70s, in Tichá Dolina (English: Silent Valley) two paleontologists found tracks from the late triassic and named them "Coelurosaurichnus Tatricus" but later it was renamed as "Eubrontes Tatricus" (Eubrontes is probably a synonym of Grallator) and size estimates suggest it was made by a smaller carnivorous theropod, with indirect evidence suggesting it was Liliensternus, but we do not know for sure. <ref>{{Cite web |last=Horák |first=Otakar |date=2018-06-08 |title=Objavil stopy "tatranského dinosaura": Tak ako dinosaury, aj ľudstvo má svoj vymedzený čas a raz príde koniec |url=https://dennikn.sk/1134864/objavil-stopy-tatranskeho-dinosaura-tak-ako-dinosaury-aj-ludstvo-ma-svoj-vymedzeny-cas-a-raz-pride-koniec/ |access-date=2025-06-29 |website=Denník N |language=sk-SK}}</ref>

== See also == {{Portal|Dinosaurs|Paleontology}} * List of dinosaur ichnogenera

== References == {{Reflist}}

== Further reading == * {{cite journal|last1=Calvo|first1=Jorge Orlando|last2=Rivera|first2=Cynthia|title=Huellas de dinosaurios en la costa oeste del embalse Ezequiel Ramos Mexía y alrededores (Cretácico Superior, Provincia de Neuquén, República Argentina)|journal=Boletín de la Sociedad Geológica Mexicana|date=2018|volume=70|number=2|pages=449 ‒ 497|doi=10.18268/BSGM2018v70n2a11|doi-access=free |bibcode=2018BoSGM..70..449C }}

Category:Theropod trace fossils Category:Ichnotaxa Category:Fossil taxa described in 1858