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''Eodromaeus'' (meaning "dawn runner") is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of probable basal
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 ...
dinosaurs from the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. ...
of
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. Like many other of the earliest-known dinosaurs, it hails from the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
-age (~230 Ma)
Ischigualasto Formation The Ischigualasto Formation is a Late Triassic fossiliferous formation and Lagerstätte in the Ischigualasto-Villa Unión Basin of the southwestern La Rioja Province and northeastern San Juan Province in northwestern Argentina. The formation ...
, within the
Ischigualasto-Villa Unión Basin The Ischigualasto-Villa Unión Basin ( es, Cuenca de Ischigualasto-Villa Unión) is a small sedimentary basin located in the Argentine Northwest, Argentina. It is located in the southwestern part of La Rioja Province and the northeastern part of ...
of northwestern
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. Upon its discovery, it was argued to be one of the oldest true theropods, supplanting its contemporary ''
Eoraptor ''Eoraptor'' () is a genus of small, lightly built, basal sauropodomorph. One of the earliest-known dinosaurs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwes ...
'', which was reinterpreted as a basal sauropodomorph.Martínez et al., 2011


Discovery

Fossils from ''Eodromaeus'' were first discovered in 1996 by Argentinean paleontologist Ricardo N. Martinez and Earthwatch volunteer Jim Murphy, and it was first believed that the fossils were a new species of ''
Eoraptor ''Eoraptor'' () is a genus of small, lightly built, basal sauropodomorph. One of the earliest-known dinosaurs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwes ...
''. However, as the researchers started to take a closer look at the fossils, they found that it had many skeletal features which were absent in ''Eoraptor'', and they understood that it came from a new genus. ''Eodromaeus'' is known from six specimens found at various levels of the
Ischigualasto Formation The Ischigualasto Formation is a Late Triassic fossiliferous formation and Lagerstätte in the Ischigualasto-Villa Unión Basin of the southwestern La Rioja Province and northeastern San Juan Province in northwestern Argentina. The formation ...
of San Juan Province,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. By far the most complete specimen is the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
, PVSJ 560, a crushed but nearly complete articulated skeleton recovered from the base of the Valle de la Luna Member. The Valle de la Luna Member is the thick third-oldest member of the formation. It overlies the thinner Cancha de Bochas and La Peña Members, the second-oldest and oldest members of the formation, respectively. Other referred specimens include PVSJ 561, 562, and 563 from the Valle de la Luna Member, PVSJ 534 from the La Peña Member, and PVSJ 877 from the Cancha de Bochas Member. Every ''Eodromaeus'' specimen lived within the humid ''
Scaphonyx ''Hyperodapedon'' is a genus of rhynchosaurs (beaked, archosaur-like reptiles) from the Late Triassic period (Carnian stage). Fossils of the genus have been found in Africa, Asia, Europe and North and South America. Its first discovery and naming ...
''-''
Exaeretodon ''Exaeretodon'' is an extinct genus of fairly large, low-slung traversodontid cynodonts from the southern parts of Pangea. Four species are known, from various formations. ''E. argentinus'' is from the Carnian-age (Late Triassic) Cancha de Boch ...
''-''
Herrerasaurus ''Herrerasaurus'' is a genus of saurischian dinosaur from the Late Triassic period. This genus was one of the earliest dinosaurs from the fossil record. Its name means "Herrera's lizard", after the rancher who discovered the first specimen in ...
'' biozone, which occupies the first half of the Ischigualasto Formation. A
bentonite Bentonite () is an absorbent swelling clay consisting mostly of montmorillonite (a type of smectite) which can either be Na-montmorillonite or Ca-montmorillonite. Na-montmorillonite has a considerably greater swelling capacity than Ca-m ...
deposit in the La Peña Member has been dated to 231 ± 0.4 Ma, indicating that the Ischigualasto Formation corresponds to the later part of the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
stage, the first stage in the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. ...
. ''Eodromaeus'' was named by Ricardo N. Martínez,
Paul C. Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic Society, National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several co ...
, Oscar A. Alcober, Carina E. Colombi, Paul R. Renne, Isabel P. Montañez and Brian S. Currie in 2011 and the
type species In 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 species that contains the biological type specime ...
is ''Eodromaeus murphi''. The generic name is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words ''Eos'' ("Dawn", "Early") and ''Dromaeus'' ("Runner"). The specific name honors Jim Murphy, who used to work the area nearby where the fossils were found.


Description

''Eodromaeus'' was a relatively small dinosaur, like most
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
dinosaurs apart from herrerasaurids. A press release estimated that its total length reach about 1.2 meters (4.0 feet) from nose to tail, with a weight of about 4.5 to 6.8 kilograms (10-15 pounds). Benson ''et al.'' (2018) estimated that ''Eodromaeus'' had a mass of 7.1 kg (~16 lbs). The animal was lightly-built and had long hindlimbs, suggesting that it was well-built for running even by the standards of most early dinosaurs. Kubo & Kubo (2012) found that, among 23 sampled Triassic
archosaurs Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
, only ''
Marasuchus ''Marasuchus'' (meaning "Mara crocodile") is a genus of basal dinosauriform archosaur which is possibly synonymous with '' Lagosuchus''. Both genera lived during the Late Triassic in what is now La Rioja Province, Argentina. ''Marasuchus'' conta ...
'' exceeded ''Eodromaeus'' in adaptations for cursoriality. Paul Sereno has estimated that it could run about 32 km per hour (20 miles per hour).Weise E
"New dog-sized dinosaur discovered"
''www.usatoday.com'', 2011-01-23.
''Eodromaeus'' has been cited by Sereno as resembling a predicted
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
to all dinosaurs, the "Eve" of dinosaurs. The skull was low and rectangular, about 12 cm (4.7 inches) in length. It had an expansive
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, bird ...
edged from below by a sharp ridge. The antorbital fenestra was preceded a promaxillary fenestra, an additional hole in the skull characteristic of
theropods 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 ca ...
and ''
Herrerasaurus ''Herrerasaurus'' is a genus of saurischian dinosaur from the Late Triassic period. This genus was one of the earliest dinosaurs from the fossil record. Its name means "Herrera's lizard", after the rancher who discovered the first specimen in ...
''. The braincase had deep depressions on its side like many theropods, and connected to the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly separ ...
via thin and blade-like basipterygoid processes. The cranium had a relatively low number of knife-shaped teeth (fifteen in total) which were longest in the front half of the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
. There was also a row of tiny teeth on the
pterygoid bone The pterygoid is a paired bone forming part of the palate of many vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fis ...
of the palate. Palatal teeth are very rare in dinosaurs (and true archosaurs in general), though they have also been found in ''
Eoraptor ''Eoraptor'' () is a genus of small, lightly built, basal sauropodomorph. One of the earliest-known dinosaurs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwes ...
''. The lower jaw was slender, with widely spaced teeth extending to the tip of the snout as in early theropods. The cervicals (neck vertebrae) were significantly more elongated than those of ''Eoraptor'', and those near the shoulders had large pits which would have housed air sacs. The shorter but more numerous dorsals (trunk vertebrae) were reinforced by hyposphene-hypantrum articulations, while the caudals (tail vertebrae) were connected by elongated prezygapophyses, as seen in other theropods. The shoulder girdle had a deep
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
connecting to a long and rod-shaped
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eith ...
. The large forelimbs had closely-appressed forearm bones and strongly-developed wrist and elbow joints. The manus (hand) had five fingers, with the second and third elongating towards their tips. In contrast, the fourth and fifth fingers were very thin and short. The pelvis (hip) had a tall ilium which connected to three sacrals (hip vertebrae). It also included an elongated ischium and pubis, the latter of which tapers before expanding into a small pubic boot similar to that of other theropods. The hindlimb possessed several muscle scars characteristic of theropods, such as a depression on the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
for extensor muscles, and a flange for tibial ligaments on the
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity i ...
.


Classification

When originally described in 2011, ''Eodromaeus'' was regarded as the earliest member of Theropoda. Theropods include the vast majority of carnivorous dinosaurs, such as ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
'', ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'', and ''
Velociraptor ''Velociraptor'' (; ) is a genus of small dromaeosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago. Two species are currently recognized, although others have been assigned in the p ...
'', as well as the only living dinosaurs, birds. ''Eodromaeus'' was considered one of most basal theropod and placed outside of the group
Neotheropoda Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and is the only group of theropods that survived the Triassic–Jurassic extinction event. All neotheropods became extinct by the ...
, which contains coelophysoids and post-Triassic theropods. The only putative theropods found to be more basal were the herrerasaurids, a group of relatively large early carnivorous dinosaurs with an unstable relationship to other dinosaurs. The discovery of ''Eodromaeus'' also lead to scrutiny regarding the contemporary early dinosaur ''
Eoraptor ''Eoraptor'' () is a genus of small, lightly built, basal sauropodomorph. One of the earliest-known dinosaurs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwes ...
. Eoraptor'' had previously been regarded as a theropod, but Martinez ''et al.'' (2011) instead concluded that it likely represented one of the most basal
sauropodomorphs Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...
, the group that includes animals like ''
Apatosaurus ''Apatosaurus'' (; meaning "deceptive lizard") is a genus of herbivorous sauropod dinosaur that lived in North America during the Late Jurassic period. Othniel Charles Marsh described and named the first-known species, ''A. ajax'', in 1877, ...
''. There is still a small amount of debate over the position of ''Eoraptor'', with a few studies reclaiming it as a theropod alongside ''Eodromaeus''.Bergman D.S., Sues H-D. (2011)
"A late-surviving basal theropod dinosaur from the latest Triassic of North America"
''
Proceedings of the Royal Society B ''Proceedings of the Royal Society'' is the main research journal of the Royal Society. The journal began in 1831 and was split into two series in 1905: * Series A: for papers in physical sciences and mathematics. * Series B: for papers in life s ...
'', published online 2011-04-13.
Most studies since 2011 have continued to place ''Eodromaeus'' as a basal theropod akin to the results of Martinez ''et al''. There are some disagreements; many subsequent studies consider ''Eodromaeus'' to lie crownward (i.e. more 'advanced') relative to '' Tawa'' and/or ''
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 recovere ...
.'' A few also reject the notion of herrerasaurids being theropods. An alternative hypothesis, first presented by Cabreira ''et al.'' (2016)'','' argues that ''Eodromaeus'' was not a theropod, but rather a more basal
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 ...
. This means that its lineage would have evolved prior to the split between sauropodomorphs and true theropods. Other putative theropods such as ''Tawa'', ''Chindesaurus'', ''
Daemonosaurus ''Daemonosaurus'' is an extinct genus of possible theropod dinosaur from the Late Triassic of New Mexico. The only known fossil is a skull and neck fragments from deposits of the latest Triassic Chinle Formation at Ghost Ranch. ''Daemonosaurus'' ...
'', and herrerasaurids fall even further stemwards than ''Eodromaeus'' according to this study and derived works. A few other unusual placements have been found in studies relating to the controversial
Ornithoscelida Ornithoscelida () is a proposed clade that includes various major groupings of dinosaurs. An order Ornithoscelida was originally proposed by Thomas Henry Huxley but later abandoned in favor of Harry Govier Seeley's division of Dinosauria into Sau ...
hypothesis of Baron ''et al''. (2017). Although Baron ''et al''.'s original paper did place ''Eodromaeus'' as a theropod, a critique and recoding by Langer ''et al.'' (2017) recovered Saurischia and moved ''Eodromaeus'' out of Theropoda. A subsequent response to that critique, by Baron ''et al.'' (2017), placed ''Eodromaeus'' as a basal herrerasaurian and simultaneously shifted the entire family Herrerasauridae outside of Dinosauria. The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
represents the original theropodan placement of ''Eodromaeus'' found by Martinez ''et al.'', 2011: The following cladogram represents the alternative hypothesis of Cabreira ''et al.'' (2016), placing ''Eodromaeus'' as a basal saurischian:


References


Bibliography

*


External links


''Eodromaeus'' skeleton

''Eodromaeus'' skull

Video: “Video: Fast, "Nasty" Little Dinosaur Discovered.” National Geographic January 13, 2011.
{{Portal bar, Dinosaurs Prehistoric theropods Dinosaur genera Carnian genera Late Triassic dinosaurs of South America Fossils of Argentina Triassic Argentina Ischigualasto Formation Fossil taxa described in 2011 Taxa named by Paul Sereno