Coelurosaurichnus
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''Grallator'' () is an
ichnogenus An ichnotaxon (plural ichnotaxa) is "a taxon based on the fossilized work of an organism", i.e. the non-human equivalent of an artifact. ''Ichnotaxon'' comes from the Ancient Greek (''íchnos'') meaning "track" and English , itself derived from ...
(form taxon based on footprints) which covers a common type of small, three-toed print made by a variety of bipedal
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s. ''Grallator''-type footprints have been found in formations dating from the
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...
through to the
early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
periods. They are found in the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
,
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
,
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
,
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
,
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
,
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
( Sousa and
Santa Maria Formation The Santa Maria Formation is a sedimentary rock formation found in Rio Grande do Sul, Brazil. It is primarily Carnian in age (Late Triassic), and is notable for its fossils of cynodonts, " rauisuchian" pseudosuchians, and early dinosaurs and othe ...
s) and
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
,''Grallator''
at
Fossilworks Fossilworks was a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database, a large relational database assembled by hundreds of paleontologists from around the world. History Fossilworks was cr ...
.org
but are most abundant on the east coast of North America, especially the
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
and
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic ...
formations of the northern part of the
Newark Supergroup The Newark Supergroup, also known as the Newark Group, is an assemblage of Upper Triassic and Lower Jurassic sedimentary and volcanic rocks which outcrop intermittently along the east coast of North America. They were deposited in a series of Tr ...
.Weishampel, D. B. & L. Young. 1996. Dinosaurs of the East Coast. The Johns Hopkins University PressXing 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. 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
stork Storks are large, long-legged, long-necked wading birds with long, stout bills. They belong to the family Ciconiidae, and make up the order Ciconiiformes . Ciconiiformes previously included a number of other families, such as herons and ibise ...
s and
heron Herons are long-legged, long-necked, freshwater and coastal birds in the family Ardeidae, with 75 recognised species, some of which are referred to as egrets or bitterns rather than herons. Members of the genus ''Botaurus'' are referred to as bi ...
s. These footprints were given this name by their discoverer,
Edward Hitchcock Edward Hitchcock (May 24, 1793 – February 27, 1864) was an American geologist and the third President of Amherst College (1845–1854). Life Born to poor parents, he attended newly founded Deerfield Academy, where he was later principal, ...
, in 1858. ''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 The Yixian Formation (; formerly Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
may have been left by ''
Caudipteryx ''Caudipteryx'' (meaning "tail feather") is a genus of small oviraptorosaur dinosaurs that lived in Asia during the Early Cretaceous, around 124.6 million years ago. They were feathered and extremely birdlike in their overall appearance, to the ...
''.


Species

Source: *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 Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English film director. He is widely regarded as one of the most influential figures in the history of cinema. In a career spanning six decades, he directed over 50 featu ...
, 1858 ( ichnotype)
*'' G. cuneatus''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English film director. He is widely regarded as one of the most influential figures in the history of cinema. In a career spanning six decades, he directed over 50 featu ...
, 1858
*'' G. damanei'' Ellenberger, 1970 *'' G. deambulator'' ( Ellenberger, 1970) *'' G. digitigradus'' ( Ellenberger, 1974) *'' G. emeiensis'' Zhen, Li, Han &
Yang Yang may refer to: * Yang, in yin and yang, one half of the two symbolic polarities in Chinese philosophy * Korean yang, former unit of currency of Korea from 1892 to 1902 * YANG, a data modeling language for the NETCONF network configuration p ...
, 1995
*'' G. formosus''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English film director. He is widely regarded as one of the most influential figures in the history of cinema. In a career spanning six decades, he directed over 50 featu ...
, 1858
*'' G. gracilis''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English film director. He is widely regarded as one of the most influential figures in the history of cinema. In a career spanning six decades, he directed over 50 featu ...
, 1865
*'' G. graciosus'' ( Ellenberger, 1970) *'' G. grancier'' ( Courel & Demathieu, 2000) *'' G. ingens'' ( Ellenberger, 1970) *'' G. jiuquwanensis'' (
Zeng Zeng (, ) is a Chinese family name. In Cantonese, it is Tsang; In Wade–Giles, such as those in Taiwan, Tseng or Tzeng; in Malaysia and Singapore, Tsen, Chen or Cheng; in the Philippines, Chan; in Indonesia, Tjan; in Vietnam, Tăng. The surname ...
, 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 Irby may refer to: People * Irby (given name), a list of people * Irby (surname), a list of people Places * Irby, Merseyside, England, a village * Irby, Virginia, United States, an unincorporated community * Irby, Washington, United States, a ...
, 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) *'' 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 Weiss or Weiß may refer to: People * Weiss (surname), including spelling Weiß * Weiss Ferdl (1883-1949), German actor Places * Mount Weiss, Jasper National Park, Alberta, Canada * Weiss Lake, Alabama * Weiß (Sieg), a river in North Rhine-Wes ...
, 1934)
*'' G. socialis'' ( Ellenberger, 1970) *'' G. ssatoi''
Yabe was a village located in Yame District, Fukuoka Prefecture, Japan. Yabe is located at 330m above sea level. The average temperature is 14 °C and the average annual precipitation is 2,706mm. As of 2006, the village had an estimated popula ...
, Inai, & Shikama, 1940
*'' G. tenuis''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English film director. He is widely regarded as one of the most influential figures in the history of cinema. In a career spanning six decades, he directed over 50 featu ...
, 1858
*'' G. toscanus'' ( Huene, 1941) *'' G. variabilis'' Lapparent & Monetnat, 1967


Paleopathology

Fossil tracks can be informative about theropod pathologies but apparently pathological traits may be due to unusual behaviors.
Sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
stratum In geology and related fields, a stratum (: strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as ...
dating to the
Norian The Norian is a division of the Triassic geological period, Period. It has the rank of an age (geology), age (geochronology) or stage (stratigraphy), stage (chronostratigraphy). It lasted from ~227.3 to Mya (unit), million years ago. It was prec ...
in southern
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
preserves tracks of an individual with a deformed digit III attributed to the
ichnogenus An ichnotaxon (plural ichnotaxa) is "a taxon based on the fossilized work of an organism", i.e. the non-human equivalent of an artifact. ''Ichnotaxon'' comes from the Ancient Greek (''íchnos'') meaning "track" and English , itself derived from ...
'' 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.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.


Occurrences

''Grallator''-type footprints have been found in formations dating from the
Early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between 251.9 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which ...
through to the
early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
periods. They are found in the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
,
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
,
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
,
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
,
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
( Sousa and
Santa Maria Formation The Santa Maria Formation is a sedimentary rock formation found in Rio Grande do Sul, Brazil. It is primarily Carnian in age (Late Triassic), and is notable for its fossils of cynodonts, " rauisuchian" pseudosuchians, and early dinosaurs and othe ...
s) and
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, but are most abundant on the east coast of North America, especially the
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
and
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic ...
formations of the northern part of the
Newark Supergroup The Newark Supergroup, also known as the Newark Group, is an assemblage of Upper Triassic and Lower Jurassic sedimentary and volcanic rocks which outcrop intermittently along the east coast of North America. They were deposited in a series of Tr ...
.


Newark Supergroup tracks

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 ''Coelophysis'' ( Traditional English pronunciation of Latin, traditionally; or , as heard more commonly in recent decades) is a genus of coelophysid Theropoda, theropod dinosaur that lived Approximation, approximately 215 to 201.4 million y ...
''. 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. 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 ''Atreipus'' is an ichnogenus or trace fossil attributed to early Ornithischian dinosaurs. Its significance for Triassic biostratigraphy has earned it some fame. Reptile footprint faunules from the early Mesozoic Newark Supergroup of eastern No ...
''. The trackmaker of ''Atreipus'' prints was a quadrupedal
ornithischian Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek st ...
. The reason for this similarity is a lack of divergence in the foot evolution of the two distinct groups of dinosaurs: ornithischians and
saurischian Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithis ...
s.


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 The Vale of Glamorgan ( ), locally referred to as ''The Vale'', is a Principal areas of Wales, county borough in the South East Wales, south-east of Wales. It borders Bridgend County Borough to the west, Cardiff to the east, Rhondda Cynon Taf t ...
,
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
. 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. The scientists called the girl's discovery "the finest impression of a 215 million-year-old dinosaur print found in Britain in a decade". Karl-James Langford of Archaeology Cymru considered the find to be "internationally important". 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".


See also

*
List of dinosaur ichnogenera This list of dinosaur ichnogenera is a comprehensive listing of all ichnogenera that have been attributed to dinosaurs, excluding class Aves (birds, both living and those known only from fossils) and purely vernacular terms. The list includes al ...


References


Further reading

* {{Taxonbar, from=Q23691966 Theropod trace fossils Fossil taxa described in 1858