Dromomeron Gregorii
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''Dromomeron'' (meaning "running
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
lagerpetid Lagerpetidae (; originally Lagerpetonidae) is a family of basal avemetatarsalians (early-diverging members of the reptile lineage leading to birds and other dinosaurs). Though traditionally considered the earliest-diverging dinosauromorphs (ar ...
avemetatarsalian Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Dinos ...
which lived around 220 to 211.9 ± 0.7 million years ago. The genus contains species known from
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
-age rocks of the
Southwestern United States The Southwestern United States, also known as the American Southwest or simply the Southwest, is a geographic and cultural list of regions of the United States, region of the United States that includes Arizona and New Mexico, along with adjacen ...
and northwestern
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. It is described as most closely related to the earlier ''
Lagerpeton ''Lagerpeton'' is a genus of lagerpetid avemetatarsalian, comprising a single species, ''L. chanarensis''. First described from the Chañares Formation of Argentina by A. S. Romer in 1971, ''Lagerpeton'' anatomy is somewhat incompletely k ...
'' of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
, but was found among remains of true dinosaurs like ''
Chindesaurus ''Chindesaurus'' ( ) is an extinct genus of basal saurischian dinosaur from the Late Triassic (213-210 million years ago) of the southwestern United States. It is known from a single species, ''C. bryansmalli'', based on a partial skeleton recov ...
'', indicating that the first dinosaurs did not immediately replace related groups. Based on the study of the overlapping material of ''Dromomeron'' and ''
Tawa hallae ''Tawa'' (named after the Hopi word for the Puebloan sun god) is a genus of possible basal theropod dinosaurs from the Late Triassic period. The fossil remains of ''Tawa hallae'', the type and only species were found in the Hayden Quarry of Gh ...
'', Christopher Bennett proposed that the two taxa were conspecific, forming a single growth series of ''Dromomeron''. However, noting prominent differences between their femurs which cannot be attributed to variation with age, Rodrigo Muller rejected this proposal in 2017. He further noted that, while ''D. romeri'' is known from juveniles only, it shares many traits in common with ''D. gigas'', which is known from mature specimens.


Description

It is known from partial remains, largely from the hindlimbs, which indicate an animal with an overall length of . The brain of ''D. gregorii'' is characterised by a floccular lobe of the cerebellum that projects within the space of the semicircular canals, which are large relative to other archosauromorphs. The anterior canal is circularly shaped.


Classification

The bones of ''Dromomeron'' are most similar to those of the older pterosauromorph ''Lagerpeton'', and the two animals have been classified together in a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Lagerpetonidae Lagerpetidae (; originally Lagerpetonidae) is a family (biology), family of basal (phylogenetics), basal Avemetatarsalia, avemetatarsalians (early-diverging members of the reptile lineage leading to birds and other dinosaurs). Though traditionall ...
.Baron, M.G., Norman, D.B., and Barrett, P.M. (2017). A new hypothesis of dinosaur relationships and early dinosaur evolution. ''Nature'', 543: 501–506.
Cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
simplified after Kammerer, Nesbitt & Shubin (2012):


Discovery and species


''Dromomeron romeri''

The species name ''romeri'' honors influential 20th-century vertebrate paleontologist
Alfred Sherwood Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
. ''Dromomeron'' and
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''D. romeri'' are based on GR 218, a complete left
thigh bone The femur (; : femurs or femora ), or thigh bone is the only bone in the thigh — the region of the lower limb between the hip and the knee. In many four-legged animals the femur is the upper bone of the hindleg. The top of the femur fits in ...
from the
Hayden Quarry Hayden may refer to: Places Inhabited places in the United States *Hayden, Alabama *Hayden, Arizona *Hayden's Ferry, former name of Tempe, Arizona *Hayden, California, former name of Hayden Hill, California *Hayden, Colorado *Hayden, Idaho *Hayden ...
at
Ghost Ranch Ghost Ranch is a retreat and education center in Rio Arriba County in north central New Mexico, United States. It is about 65 miles northwest of Santa Fe and 14 miles from Abiquiu, the nearest community. In the later 20th century, it was the ...
,
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
. The rocks there are in the lower portion of the Petrified Forest Member of the
Chinle Formation The Chinle Formation is an Upper Triassic continental geological formation of fluvial, lacustrine, and palustrine to eolian deposits spread across the U.S. states of Nevada, Utah, northern Arizona, western New Mexico, and western Colorado. In ...
, and are
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 age. Additional hindlimb bones, some probably from the same individual, are also known, and a partial skeleton has been recovered from Hayden Quarry, but has not yet been fully prepared. A few other specimens have been recovered from nearby localities, including the Snyder Quarry. Other specimens from the Chinle Formation of
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
and a roughly contemporaneous part of the
Dockum Group The Dockum is a Late Triassic (approximately late Carnian through Rhaetian, or 223–200 Ma) geologic group found primarily on the Llano Estacado of western Texas and eastern New Mexico with minor exposures in southwestern Kansas, eastern Color ...
of
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
also have been assigned to this genus. These have been assigned to a second species ''D. gregorii'', named in 2009.


''Dromomeron gregorii''

''D. gregorii'', named for
Joseph T. Gregory Dr. Joseph Tracy Gregory (July 28, 1914 – November 18, 2007) was an American paleontologist and professor. Joseph Tracy Gregory was born in Eureka, California, the only child of Frank Gregory, a civil engineer, and Edith Tracy, a high school ...
, is based on TMM 31100–1306, a right femur (thigh bone) from the Otis Chalk Quarry,
Colorado City Formation The Colorado City Formation is a Late Triassic Formation (geology), geologic formation in the Dockum Group of Texas, United States. It has previously been known as the Iatan Member, Colorado City Member or 'Pre-Tecovas Horizon' (as it is assumed ...
, (Dockum Group), near Otis Chalk, Texas. Several other limb bones from the quarry, and a partial femur (thigh bone) from the ''
Placerias ''Placerias'' (meaning 'broad body') is an extinct genus of dicynodonts that lived during the Carnian to the Norian age of the Triassic Period (geology), Period (230–215 million years ago). ''Placerias'' belongs to a group of dicynodonts called ...
'' Quarry of eastern
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
have been assigned to this species. The rocks that ''D. gregorii'' is known from are older than those ''romeri'' has been found in. As with the Hayden Quarry, the Otis Chalk Quarry has at least one specimen of a
herrerasaurid Herrerasauridae is a family of carnivorous dinosaurs, possibly basal to either theropods or even all of saurischians, or even their own branching from Dracohors, separate from Dinosauria altogether. They are among the oldest known dinosaurs, fi ...
.


''Dromomeron gigas''

A third species, ''D. gigas'', was described by Martínez ''et al''. (2016) on the basis of fossils recovered from the Norian
Quebrada del Barro Formation The Quebrada del Barro Formation is a geological formation of the Marayes-El Carrizal Basin in San Juan Province, Argentina. This formation is the most fossiliferous portion of the Triassic Marayes Group, and is also the youngest unit of the gr ...
in northwestern Argentina.


Paleoecology

Also found at the
Hayden Quarry Hayden may refer to: Places Inhabited places in the United States *Hayden, Alabama *Hayden, Arizona *Hayden's Ferry, former name of Tempe, Arizona *Hayden, California, former name of Hayden Hill, California *Hayden, Colorado *Hayden, Idaho *Hayden ...
are the remains of
phytosaur Phytosaurs (Φυτόσαυροι in Greek, meaning 'plant lizard') are an extinct group of large, mostly semiaquatic Late Triassic archosauriform or basal archosaurian reptiles. Phytosaurs belong to the order Phytosauria and are sometimes ref ...
s,
aetosaur Aetosaurs () are heavily armored reptiles belonging to the extinct order (biology), order Aetosauria (; from Ancient Greek, Greek, (aetos, "eagle") and (, "lizard")). They were medium- to large-sized Omnivore, omnivorous or Herbivore, herbivor ...
s,
rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians ...
ns, and several types of dinosaurs and dinosaur relatives, including a ''
Silesaurus ''Silesaurus'' is a genus of silesaurid dinosauriform from the Late Triassic, of what is now Poland. Discovery and naming The Krasiejów claypit near Opole, Poland, was first discovered as a fossil locality in the 1980s after quarrying for ...
''-like animal, the
herrerasaur Herrerasauridae is a family of carnivorous dinosaurs, possibly basal to either theropods or even all of saurischians, or even their own branching from Dracohors, separate from Dinosauria altogether. They are among the oldest known dinosaurs, ...
id ''
Chindesaurus ''Chindesaurus'' ( ) is an extinct genus of basal saurischian dinosaur from the Late Triassic (213-210 million years ago) of the southwestern United States. It is known from a single species, ''C. bryansmalli'', based on a partial skeleton recov ...
'', and the basal theropod ''Tawa''. Finding the remains of four types of dinosaurs and dinosaur relatives (including ''Dromomeron'' itself) is noteworthy because it shows that dinosaurs did not immediately replace their dinosauromorph predecessors; that some of these groups, like the lagerpetonids, persisted (for longer than previously known) and diversified; and that dinosaurian replacement may have occurred at different times in different areas.


References


External links


Online supplementary material
for the Irmis ''et al.''. article (PDF).

for the Irmis ''et al.''. article. {{Taxonbar, from=Q2566615 Lagerpetidae Late Triassic archosaurs of North America Late Triassic reptiles of South America Fossil taxa described in 2007 Triassic Argentina