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''Eodromaeus'' (meaning "dawn runner") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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 probable basal
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 from the
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 ...
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 ...
. 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 227.3 ...
-age (~230 Ma) Ischigualasto Formation, within the Ischigualasto-Villa Unión Basin of 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 ...
. 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 (phylogenetics), basal sauropodomorpha, sauropodomorph dinosaur. One of the earliest-known dinosaurs and one of the earliest sauropodomorphs, it lived approximately 231 to 228 million yea ...
'', 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 (phylogenetics), basal sauropodomorpha, sauropodomorph dinosaur. One of the earliest-known dinosaurs and one of the earliest sauropodomorphs, it lived approximately 231 to 228 million yea ...
''. 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 of San Juan Province,
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 ...
. By far the most complete specimen is the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
, PVSJ 560, a crushed but nearly complete articulated
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
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''-'' Exaeretodon''-'' Herrerasaurus'' biozone, which occupies the first half of the Ischigualasto Formation. A
bentonite Bentonite ( ) is an Absorption (chemistry), 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 swelli ...
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 227.3 ...
stage, the first stage in the
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 ...
. ''Eodromaeus'' was named by Ricardo N. Martínez, Paul C. Sereno, Oscar A. Alcober, Carina E. Colombi, Paul R. Renne, Isabel P. Montañez and Brian S. Currie in
2011 The year marked the start of a Arab Spring, series of protests and revolutions throughout the Arab world advocating for democracy, reform, and economic recovery, later leading to the depositions of world leaders in Tunisia, Egypt, and Yemen ...
and the
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 ...
is ''Eodromaeus murphi''. The generic name is derived from the
Greek Greek may refer to: 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 of all kno ...
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 227.3 ...
dinosaurs apart from herrerasaurids. A press release estimated that its total length reach about from nose to tail, with a weight of about . Benson ''et al.'' (2018) estimated that ''Eodromaeus'' had a mass of . 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
archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...
s, only ''
Marasuchus ''Marasuchus'' (meaning "Mara (mammal), Mara crocodile") is a genus of basal Dinosauriformes, dinosauriform archosaur which is possibly synonymous with ''Lagosuchus''. Both genera lived during the Late Triassic in what is now La Rioja Province, A ...
'' 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. According to modern evolutionary biology, all living beings could be descendants of a unique ancestor commonl ...
to all dinosaurs, the "Eve" of dinosaurs. The skull was low and rectangular, about 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 Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
edged from below by a sharp ridge. The antorbital fenestra was preceded a promaxillary fenestra, an additional hole in the skull characteristic of theropods and '' Herrerasaurus''. The
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calv ...
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 sep ...
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 In vertebrates, the maxilla (: maxillae ) 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. The two maxil ...
. There was also a row of tiny teeth on the pterygoid bone 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 (phylogenetics), basal sauropodomorpha, sauropodomorph dinosaur. One of the earliest-known dinosaurs and one of the earliest sauropodomorphs, it lived approximately 231 to 228 million yea ...
''. 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 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 present as part of the scapula, but this is n ...
connecting to a long and rod-shaped
scapula The scapula (: 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 either side ...
. 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 The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
(hip) had a tall ilium which connected to three sacrals (hip vertebrae). It also included an elongated
ischium The ischium (; : is ...
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 (; : 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 ...
for extensor muscles, and a flange for tibial ligaments on the
fibula The fibula (: fibulae or fibulas) 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. ...
.


Classification

When originally described in 2011, ''Eodromaeus'' was regarded as the earliest member of
Theropoda Theropoda (; from ancient Greek iktionary:θηρίον, , (''therion'') "wild beast"; wiktionary:πούς, , wiktionary:ποδός, (''pous, podos'') "foot" is one of the three major groups (Clade, clades) of Dinosaur, dinosaurs, alon ...
. Theropods include the vast majority of carnivorous dinosaurs, such as ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
'', ''
Allosaurus ''Allosaurus'' ( ) is an extinct genus of theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic period ( Kimmeridgian to late Tithonian ages). The first fossil remains that could definitively be ascribed to th ...
'', and ''
Velociraptor ''Velociraptor'' (; ) is a genus of small dromaeosaurid dinosaurs 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 th ...
'', as well as the only living dinosaurs,
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s. ''Eodromaeus'' was considered one of the most basal theropod and placed outside of the group Neotheropoda, 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 (phylogenetics), basal sauropodomorpha, sauropodomorph dinosaur. One of the earliest-known dinosaurs and one of the earliest sauropodomorphs, it lived approximately 231 to 228 million yea ...
. 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, the group that includes animals like '' Apatosaurus''. 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 recov ...
.'' 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
saurischia 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 Ornithi ...
n. 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'', 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 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 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 ...
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 Saurischia Dinosaur genera Carnian dinosaurs Ischigualasto Formation Fossil taxa described in 2011 Taxa named by Paul Sereno Dinosaurs of Argentina