Ekrixinatosaurus
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''Ekrixinatosaurus'' ('explosion-born reptile') 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
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 foun ...
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 ...
which lived approximately 100 to 97
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
period. Its fossils have been found in
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 ...
. Only one species is currently recognized, ''Ekrixinatosaurus novasi'', from which the specific name honors of Dr. Fernando Novas for his contributions to the study of abelisaurid theropods, while the genus name refers to the dynamiting of 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 ...
specimen. It was a large abelisaur, measuring between in length and weighing .


Discovery and naming

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 ...
, ''Ekrixinatosaurus novasi'', was first described in 2004 by Argentinian
paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Jorge Calvo, and Chilean paleontologists David Rubilar-Rogers and Karen Moreno. The fossils were found dispersed over an area of 15 m in the
Candeleros Formation The Candeleros Formation is a geologic formation that outcrop, crops out in the Rio Negro Province, Río Negro, Neuquen Province, Neuquén, and Mendoza Province, Mendoza provinces of northern Patagonia, Argentina. It is the oldest formation in the ...
, a geologic formation that outcrops in Río Negro,
Neuquén Neuquén (; ) is the capital city of the Argentine province of Neuquén and of the Confluencia Department, located in the east of the province. It occupies a strip of land west of the confluence of the Limay and Neuquén rivers which form t ...
and Mendoza provinces of Argentina. This formation dates from 100 to 97 mya, and consists of red beds where other famous vertebrate animals have been discovered, such as ''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period (geology), period, approximately 99.6 to 95 million years ago. The holotype specim ...
'', '' Rebbachisaurus'' and ''
Andesaurus ''Andesaurus'' ( ; "Andes lizard") is a genus of basal titanosaurian sauropod dinosaur which existed during the middle of the Cretaceous Period in South America. Like most sauropods, it would have had a small head on the end of a long neck and a ...
''. The discovery occurred due to excavations for building a gas pipeline conducted by Gasoducto del Pacífico Company in Bajo del Añelo. Before the discovery of ''Ekrixinatosaurus'', '' Carnotaurus sastrei'' and '' Aucasaurus garridoi'' were the most complete specimens 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. The remains of ''Ekrixinatosaurus'' helped fill in more information about abelisaur anatomy as it contained portions of the skeleton that were previously unknown, unpublished, or poorly preserved in other specimens. The holotype skeleton (MUCPv-294) was well preserved yet disarticulated. It contained elements including a left and partial right maxillae; basicranium; both dentaries; teeth; cervical, a dorsal, sacral and caudal vertebrae; haemal arches; ribs; ilia, pubis and proximal ischia; left and distal end of right femur; left tibia; left astragalus and calcaneum; proximal end of left fibula and right tibia; metatarsals; phalanges; and a pedal ungual.


Description

''Ekrixinatosaurus novasi'' is a large
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 foun ...
with a relatively large head and robust limbs. The combination of characteristics including a large skull, prominent supraorbital ridge above the dorsal aspect of the orbit, anteroposteriorly compressed cervical vertebrae, and robust, proportionally short hindlimbs with a tibia shorter than femur suggests that this was a massive animal with a great capacity to sustain injuries during intraspecific or interspecific combat. The only known specimen was previously estimated between in length and in body mass, and some suggested that this specimen actually represented the largest abelisaurid yet known at in length, surpassing the type of ''
Carnotaurus ''Carnotaurus'' (; ) is a genus of Theropoda, theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 72 and 69 million years ago. The only species is ''Carnotaurus sastrei''. Known from a si ...
''. However, it was later noted by other researchers that the latter length estimate of was based only on the absolute size of the skull, ignoring that limb bone comparisons clearly show ''Carnotaurus'' was larger, and thus ''Carnotaurus'' was larger than ''Ekrixinatosaurus'' but with a proportionally smaller head.


Skull

The skull of ''Ekrixinatosaurus'' was boxy and proportionally shorter and deeper than most other large carnivorous dinosaurs. The jaws also curved upward, a trait shared with some other abelisaurs. The skull is estimated to be approximately 83 cm long based on comparisons with ''
Carnotaurus ''Carnotaurus'' (; ) is a genus of Theropoda, theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 72 and 69 million years ago. The only species is ''Carnotaurus sastrei''. Known from a si ...
'' and '' Majungasaurus'', and while '' Abelisaurus'' does not have a complete maxilla its preserved size is similar to that of ''Ekrixinatosaurus''. As in other abelisaurids, the facial bones, especially the nasal bones, were sculptured with numerous small holes and spikes. In life, a wrinkled and possibly keratinous skin would have covered these bones. The maxilla was short and contained 16 alveoli, some with short teeth that were flattened laterally with anterior and posterior serrations. The maxilla of ''Ekrixinatosaurus'' also exhibits a dorsally projected ascending ramus and a short rostral ramus, suggesting a relatively high skull.


Classification

In 2004, ''Ekrixinatosaurus'' was placed in 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 ...
. The article by Calvo ''et al.'' in 2004, also contained a preliminary
cladistic Cladistics ( ; from Ancient Greek 'branch') 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 ...
analysis of the
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
relationships between ''Ekrixinatosaurus'' and those abelisaurids of which at the time relatively complete skeletons were known, finding it to be more closely related to Carnotaurini than to ''
Ceratosaurus ''Ceratosaurus'' (from Greek 'horn' and 'lizard') is a genus of carnivorous theropod dinosaur that lived in the Late Jurassic period (Kimmeridgian to Tithonian ages). The genus was first described in 1884 by American paleontologist Othni ...
''. In 2008, a study recovered it as the
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''Skorpiovenator''.


Paleoecology

''Ekrixinatosaurus'' was found in the red beds of the
Candeleros Formation The Candeleros Formation is a geologic formation that outcrop, crops out in the Rio Negro Province, Río Negro, Neuquen Province, Neuquén, and Mendoza Province, Mendoza provinces of northern Patagonia, Argentina. It is the oldest formation in the ...
, which has yielded a wide variety of vertebrates. It shared its environment with the
titanosauria Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of Sauropoda, sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with tax ...
n
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
''
Andesaurus ''Andesaurus'' ( ; "Andes lizard") is a genus of basal titanosaurian sauropod dinosaur which existed during the middle of the Cretaceous Period in South America. Like most sauropods, it would have had a small head on the end of a long neck and a ...
'' and the
rebbachisaurid Rebbachisauridae is a Family (biology), family of sauropod dinosaurs known from fragmentary fossil remains from the Cretaceous of South America, Africa, North America, Europe and possibly Central Asia. Taxonomy In 1990 sauropod specialist Jack M ...
sauropods '' Limaysaurus'' and '' Nopcsaspondylus''.
Iguanodont Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivore, herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and rel ...
ornithischia Ornithischia () is an extinct clade 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 ...
n remains have reportedly also been found. The
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period (geology), period, approximately 99.6 to 95 million years ago. The holotype specim ...
'' was possibly the
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
. Smaller predators also inhabited the area, including the
dromaeosaurid Dromaeosauridae () is a family (biology), family of feathered coelurosaurian Theropoda, theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous period (geology), Period. The name Drom ...
''
Buitreraptor ''Buitreraptor'' (meaning "La Buitrera seizer") is a genus of dromaeosaurid dinosaurs that lived during the Late Cretaceous of Argentina at the Candeleros Formation. ''Buitreraptor'' was described in 2005 and the type species is ''Buitreraptor g ...
'', the alvarezsauroid '' Alnashetri'', and the basal
coelurosauria Coelurosauria (; from Greek, meaning "hollow-tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyra ...
n '' Bicentenaria''. Other primitive reptiles lived in the area, such as the primitive snake '' Najash'', the crocodile ''
Araripesuchus ''Araripesuchus'' is a genus of extinct crocodyliform that existed during the Cretaceous period of the late Mesozoic era some 125 to 66 million years ago. ''Araripesuchus'' is generally considered to be a notosuchian (belonging to the clade Mes ...
'', along with turtles, fish, pipid frogs and mammals.
Pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s also lived in the area, as evidenced by pterosaur tracks. A wide variety of dinosaur trackways have also been found in the
Candeleros Formation The Candeleros Formation is a geologic formation that outcrop, crops out in the Rio Negro Province, Río Negro, Neuquen Province, Neuquén, and Mendoza Province, Mendoza provinces of northern Patagonia, Argentina. It is the oldest formation in the ...
, suggesting significant activity in the area. Recent studies on
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
n theropods have interpreted abelisauroids as modest and medium-sized dinosaurs that co-occurred with giant
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
s during the Early and early Late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
. It has been hypothesized that it was only after extinction of these carcharodontosaurids that abelisaurids were able to diversify into more robust forms that occupied the niche of top predators of their ecosystems. However, it has been observed that both ''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period (geology), period, approximately 99.6 to 95 million years ago. The holotype specim ...
'' and ''Ekrixinatosaurus'' were among the largest of their respective clades yet existed at the same time, which refutes this hypothesis. Both these animals occupied the role of the largest carnivores; however, it is uncertain whether they played different ecological roles (such as active predation vs. scavenging). In addition, the known distribution of abelisaurids in South America, Madagascar and India brings the hypothesis of a dispersion route between these areas by a terrestrial bridge called the
Kerguelen Plateau The Kerguelen Plateau (, ), also known as the Kerguelen–Heard Plateau, is an oceanic plateau and large igneous province (LIP) located on the Antarctic Plate, in the southern Indian Ocean. It is about to the southwest of Australia and is near ...
that formed prior to the separation of Africa and South America.


See also

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


References

{{Taxonbar, from=Q132571 Abelisauridae Dinosaur genera Cenomanian dinosaurs Candeleros Formation Fossil taxa described in 2004 Dinosaurs of Argentina