Eoabelisaurus
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''Eoabelisaurus'' () is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found ...
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 ...
from the
Lower Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma&nb ...
Cañadón Asfalto Formation The Cañadón Asfalto Formation is a Lower Jurassic to Late Jurassic geologic formation, from the Jurassic period of the Mesozoic Era. Its age is controversial, uranium-lead dating of the volcanic tuff beds having given various different ages. A ...
of the
Cañadón Asfalto Basin The Cañadón Asfalto Basin ( es, Cuenca de Cañadón Asfalto) is an irregularly shaped sedimentary basin located in north-central Patagonia, Argentina. The basin stretches from and partly covers the North Patagonian Massif in the north, a high for ...
in
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 ...
,
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
. The generic name combines a Greek ἠώς, (''eos''), "dawn", with the name '' Abelisaurus'', in reference to the fact it represents an early relative of the latter. Only one species is currently recognized, ''E. mefi''; the specific name honours the MEF, the Museo Paleontológico "Egidio Feruglio", where discoverer Diego Pol is active. It is characterized by reduced forelimb proportions that show primitive characteristics of the
Abelisauridae Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are fou ...
family.


Discovery and naming

In 2009, Argentinian
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Diego Pol discovered the skeleton of a theropod near the village of Cerro Cóndor in
Chubut Province Chubut ( es, Provincia del Chubut, ; cy, Talaith Chubut) is a province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa Cruz Province), the Andes ra ...
. The remains were found in the Jugo Loco locality that is placed in a series of fine beds of mudstone, marlstone, and limestone in the
Cañadón Asfalto Formation The Cañadón Asfalto Formation is a Lower Jurassic to Late Jurassic geologic formation, from the Jurassic period of the Mesozoic Era. Its age is controversial, uranium-lead dating of the volcanic tuff beds having given various different ages. A ...
. In 2012, based on these remains, 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 specimen ...
''Eoabelisaurus mefi'' was named and described by Pol and his German colleague Oliver Rauhut. Before discovery, the oldest known abelisaurids were represented only by fragmentary remains from the late Early Cretaceous of South America and Africa and older records of abelisauroids in general were questionable. With the discovery of ''Eoabelisaurus'', the temporal range of the clade was extended for more than 40 
million years The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
into the
Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toarcian ...
. The existence of a derived ceratosaur at the time indicated a rapid diversification of ceratosaurs during that time period, as all the major ceratosaurian lineages (ceratosaurids, noasaurids, and abelisaurids) had been established.


Description

It was a
biped Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
al
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
that was estimated to have reached in length, although a comprehensive analysis of abelisaur size conducted in 2016 yielded a size estimate of . Molina-Pérez and Larramendi gave a higher estimation of 7.5 meters (25.6 ft) and 445 kg (981 lbs). 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 several ...
specimen, MPEF PV 3990, was uncovered in a layer of the
Cañadón Asfalto Formation The Cañadón Asfalto Formation is a Lower Jurassic to Late Jurassic geologic formation, from the Jurassic period of the Mesozoic Era. Its age is controversial, uranium-lead dating of the volcanic tuff beds having given various different ages. A ...
, a lacustrine deposit dating originally from the
Aalenian The Aalenian () is a subdivision of the Middle Jurassic Epoch/Series of the geologic timescale that extends from about 174.1 Ma to about 170.3 Ma (million years ago). It was preceded by the Toarcian and succeeded by the Bajocian. Stratigraphi ...
-
Bajocian In the geologic timescale, the Bajocian is an age and stage in the Middle Jurassic. It lasted from approximately 170.3 Ma to around 168.3 Ma (million years ago). The Bajocian Age succeeds the Aalenian Age and precedes the Bathonian Age. Stra ...
, roughly 170 million years old, yet a recent advanced zircon datation constrained it´s age to the Middle-Late Toarcian (179-178 million years). It consists of a nearly complete skeleton with skull, of a subadult or adult individual. The posterior half of the skeleton was found in articulation and the anterior dorsal and cervical vertebrae and forelimbs were found partially disarticulated prior to burial. The skull was discovered slightly separated from the vertebral column. The skull and anterior presacrals were also exposed at the time of discovery and had been partially been destroyed by erosion. From the material known of the snout, only a small fragment of the right maxilla has been recovered and shows that the interdental plates are fused, but not striated. The known posterior part of the skull is high, with an oval orbit and an enlarged infratemporal fenestra, similar to other
ceratosaur Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with ''Ceratosaurus'' than with birds. The oldest known ceratosaur, ''Saltriovenator'', dates to the earlies ...
s. The skull roof is not notably thickened and no cranial ornamentation is present. Both humeri from the known material are poorly preserved, but show primitive characters. The articular head is slightly rounded, but not a globular shape that is commonly seen in
noasaurid Noasauridae is an extinct family of theropod dinosaurs belonging to the group Ceratosauria. They were closely related to the short-armed abelisaurids, although most noasaurids had much more traditional body types generally similar to other ther ...
s and
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found ...
s. The radius and ulna are short and the ulna has a large olecranon process. The manus is very foreshortened, retains four digits, and has short metacarpals. Metacarpal I is about half the length of metacarpal II and considerably more slender. The arm also bears a stout phalanx that is slightly longer than the metacarpal. The non-terminal manual phalanges are about as long as wide and lack any constriction between the articular ends, and manual unguals are reduced. It is these reduced limb proportions that demonstrate ''Eoabelisaurus'' was indeed a primitive abelisaurid. The exact number of vertebrae is unknown due to several gaps in the holotype's spine, but its cervical vertebrae are short and have two pneumatic foramina on either side of the centra. The length of vertebral centra remains constant over the preserved portion of the tail, but middle and posterior caudals are considerably lower.


Classification

''Eoabelisaurus'' was assigned to the basalmost position in
Abelisauridae Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are fou ...
by its describers. It would, then, be the oldest abelisaurid species known by forty (40) million years. The describers indicated that in the
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis a difference of only a single trait would have resulted in a position lower in the evolutionary tree, basal in the
Abelisauroidea Abelisauroidea is typically regarded as a Cretaceous group, though the earliest abelisauridae remains are known from the Middle Jurassic of Argentina (classified as the species Eoabelisaurus mefi) and possibly Madagascar (fragmentary remains of ...
. 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 d ...
follows their analysis. In 2018, Rafael Delcourt placed ''Eoabelisaurus'' at the basic position within
Ceratosauridae Ceratosauridae is an extinct family of theropod dinosaurs belonging to the infraorder Ceratosauria. The family's type genus, ''Ceratosaurus'', was first found in Jurassic rocks from North America. Ceratosauridae is made up of the genera ''Cer ...
, noting that the topology of this genus is still problematic. The following cladogram demonstrates the relationships of ''Eoabelisaurus'' according to latter publication.


Paleobiology


Reduced forelimbs

Before the discovery of ''Eoabelisaurus'', abelisaurid anatomy was only known from a handful of
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
taxa that were aberrant in their morphology, such as their unusual skull structure and reduction of their forelimbs. ''Eoabelisaurus'' shows what was previously an unknown stage in the evolution of abelisaurids, having only some of the cranial modifications and a unique combination of features in its forelimbs. The manus of ''Eoabelisaurus'' have a derived morphology, with short and robust metacarpals, non-terminal phalanges, and reduced manual unguals. The humerus is unreduced, and the ulna and radius are shortened but do not differ from more basal ceratosaurs. Therefore, the modification of the forelimbs in the evolution of abelisaurids seems to have occurred in a modular fashion, starting with the distal elements such as the fingers, and only later affecting the more proximal elements. It is with this evidence that ''Eoabelisaurus'' confirms a recently proposed evolutionary scenario of abelisaurid limbs based on the incomplete evidence of Cretaceous abelisauroids.


See also

*
Timeline of ceratosaur research This timeline of ceratosaur research is a chronological listing of events in the history of paleontology focused on the ceratosaurs, a group of relatively primitive, often horned, predatory theropod dinosaurs that became the apex predators of ...


References


External links

{{Portal bar, Dinosaurs Abelisaurids Dinosaur genera Middle Jurassic dinosaurs of South America Jurassic Argentina Fossils of Argentina Cañadón Asfalto Formation Fossil taxa described in 2012