Grallator
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''Grallator'' GRA-luh-tor"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. ''Ichnotaxa'' comes from the Greek ίχνος, ''ichnos'' meaning ''track'' and ταξις, ''taxis'' meaning ...
(form taxon based on footprints) which covers a common type of small, three-toed print made by a variety of bipedal
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
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 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a un ...
through to the
early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
periods. They are found in the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
( 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 oth ...
s) and
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
,''Grallator''
at
Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals ...
.org
but are most abundant on the east coast of North America, especially the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
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-J ...
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 rocks which outcrop intermittently along the United States East Coast. They were deposited in a series of Triassic basins app ...
.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 called Ciconiidae, and make up the order Ciconiiformes . Ciconiiformes previously included a number of other families, such as herons an ...
s and
heron The herons are long-legged, long-necked, freshwater and coastal birds in the family Ardeidae, with 72 recognised species, some of which are referred to as egrets or bitterns rather than herons. Members of the genera ''Botaurus'' and ''Ixobrychus ...
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 transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its ex ...
may have been left by ''
Caudipteryx ''Caudipteryx'' (which means "tail feather") is a genus of peacock-sized theropod dinosaurs that lived in the Barremian age of the early Cretaceous (about 124.6 million years ago). They were feathered and extremely birdlike in their overall appea ...
''.


Species

*Subgenus '' G. (Coelurosaurichnus)'' **'' G. (C.) palmipes'' ***'' G. (C.) p. exiguus'' ( Ellenberger, 1970) *Subgenus '' G. (Grallator)'' **'' G. (G.) zvierzi'' Gierlinski, 1991 *'' G. andeolensis''
Gand Gand may refer to: * Ghent (French: Gand), a city in Belgium * Gand (''Star Wars''), a fictional planet and its resident species in the ''Star Wars'' franchise * Gand., the standard author abbreviation for Michel Gandoger * ', Old Norse word for ...
, 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 filmmaker. 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 featur ...
, 1858 ( ichnotype)
*'' G. cuneatus''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English filmmaker. 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 featur ...
, 1858
*'' G. damanei'' Ellenberger, 1970 *'' G. deambulator'' ( Ellenberger, 1970) *'' G. digitigradus'' ( Ellenberger, 1974) *'' G. emeiensis'' Zhen, Li,
Han Han may refer to: Ethnic groups * Han Chinese, or Han People (): the name for the largest ethnic group in China, which also constitutes the world's largest ethnic group. ** Han Taiwanese (): the name for the ethnic group of the Taiwanese p ...
&
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 pr ...
, 1995
*'' G. formosus''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English filmmaker. 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 featur ...
, 1858
*'' G. gracilis''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English filmmaker. 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 featur ...
, 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, Chen or Cheng; in the Philippines, Chan; in Indonesia, Tjan; in Vietnam, Tăng. The surname Zeng is ...
, 1982) =''Hunanpus''
*'' G. kehli'' ( Beurlen, 1950) *'' G. kronbergeri'' ( Rehnelt, 1959) *'' G. lacunensis'' ( Ellenberger, 1970) *'' G. leribeensis'' ( Ellenberger, 1970) *'' G. limnosus'' Zhen, Li, &
Rao __NOTOC__ Rao may refer to: Geography * Rao, West Sumatra, one of the districts of West Sumatra, Indonesia * Råö, a locality in Kungsbacka Municipality, Halland County, Sweden Transport * Dr. Leite Lopes–Ribeirão Preto State Airport , IAT ...
, 1985
*'' G. madseni''
Irby Irby may refer to: * Irby (surname), a list of people * Irby, Merseyside, England, a village * Irby, Virginia, United States, an unincorporated community * Irby, Washington, United States, an unincorporated community * Irby in the Marsh, Lincolnsh ...
, 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 Gand may refer to: * Ghent (French: Gand), a city in Belgium * Gand (''Star Wars''), a fictional planet and its resident species in the ''Star Wars'' franchise * Gand., the standard author abbreviation for Michel Gandoger * ', Old Norse word for ...
, 1976)
*'' G. paulstris'' ( Ellenberger, 1970) *'' G. perriauxi'' ( Demathieu &
Gand Gand may refer to: * Ghent (French: Gand), a city in Belgium * Gand (''Star Wars''), a fictional planet and its resident species in the ''Star Wars'' franchise * Gand., the standard author abbreviation for Michel Gandoger * ', Old Norse word for ...
, 1972)
*'' 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 Gand may refer to: * Ghent (French: Gand), a city in Belgium * Gand (''Star Wars''), a fictional planet and its resident species in the ''Star Wars'' franchise * Gand., the standard author abbreviation for Michel Gandoger * ', Old Norse word for ...
& Pellier, 1976)
*'' G. sassendorfensis'' ( Kuhn, 1958) *'' G. sauclierensis'' Demathieu & Sciau, 1992 *'' G. schlauersbachensis'' ( Weiss, 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 Inai is a village in Saran district of Bihar state, India. It has the Ghaghara River to its south, the Chhapra- Ballia railway line to its north, a small irrigation and flood canal to its west, and Chhapra municipality to its east. It gains s ...
, & Shikama, 1940
*'' G. tenuis''
Hitchcock Sir Alfred Joseph Hitchcock (13 August 1899 – 29 April 1980) was an English filmmaker. 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 featur ...
, 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 sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
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 ei ...
dating to the
Norian The Norian is a division of the Triassic Period. It has the rank of an age (geochronology) or stage (chronostratigraphy). It lasted from ~227 to million years ago. It was preceded by the Carnian and succeeded by the Rhaetian. Stratigraphic defi ...
in southern
Wales Wales ( cy, Cymru ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by England to the Wales–England border, east, the Irish Sea to the north and west, the Celtic Sea to the south west and the ...
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. ''Ichnotaxa'' comes from the Greek ίχνος, ''ichnos'' meaning ''track'' and ταξις, ''taxis'' meaning ...
'' 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 Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is a un ...
through to the
early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
periods. They are found in the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
( 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 oth ...
s) and
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, but are most abundant on the east coast of North America, especially the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
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-J ...
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 rocks which outcrop intermittently along the United States East Coast. They were deposited in a series of Triassic basins app ...
.


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'' ( traditionally; or , as heard more commonly in recent decades) is an extinct genus of coelophysid theropod dinosaur that lived approximately 228 to 201.3 million years ago during the latter part of the Triassic Period from t ...
''. 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 An ichnotaxon (plural ichnotaxa) is "a taxon based on the fossilized work of an organism", i.e. the non-human equivalent of an artifact. ''Ichnotaxa'' comes from the Greek ίχνος, ''ichnos'' meaning ''track'' ...
''. The trackmaker of ''Atreipus'' prints was a quadrupedal
ornithischian Ornithischia () is an extinct order 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 s ...
. 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 Lily Wilder found a 215 to 220-million years old dinosaur footprint at Bendricks Bay in the
Vale of Glamorgan The Vale of Glamorgan ( cy, Bro Morgannwg ), often referred to as The Vale, is a county borough in the south-east of Wales. It borders Bridgend County Borough to the west, Cardiff to the east, Rhondda Cynon Taf to the north, and the Bristol C ...
,
Wales Wales ( cy, Cymru ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by England to the Wales–England border, east, the Irish Sea to the north and west, the Celtic Sea to the south west and the ...
. Experts believe 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, an archaeologist from
Amgueddfa Cymru – National Museum Wales Amgueddfa Cymru – Museum Wales, branded as simply Amgueddfa Cymru (formerly the National Museums and Galleries of Wales and legally National Museum of Wales), is a Welsh Government sponsored body that comprises seven museums in Wales: * N ...
, called it as the discovery of "international importance." Cindy Howells, the curator of the museum, described it as “one of the best-preserved examples from anywhere in the UK” and said 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 all ...


References


Further reading

* {{Taxonbar, from=Q23691966 Dinosaur trace fossils