Ekrixinatosaurus Novasi
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''Ekrixinatosaurus'' ('explosion-born reptile') is a genus of abelisaurid theropod which lived approximately 100 to 97 million years ago during the Late Cretaceous period. Its fossils have been found in Argentina. 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. It was a large abelisaur, measuring between in length and weighing .


Discovery and naming

The type species, ''Ekrixinatosaurus novasi'', was first described in 2004 by Argentinian paleontologist Jorge Calvo, and Chilean paleontologists David Rubilar-Rogers and Karen Moreno. The fossils were found dispersed over an area of 15m in the
Candeleros Formation The Candeleros Formation (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
, a geologic formation that outcrops in Río Negro, Neuquén, and Mendoza provinces of Argentina. This formation dates from 100-97 mya., and consists of red beds where other famous vertebrate animals have been discovered, such as '' Giganotosaurus'', ''
Rebbachisaurus ''Rebbachisaurus'' (meaning "Khebbash, Aït Rebbach lizard") is a genus of sauropod dinosaur of the superfamily Diplodocoidea, that lived during the Late Cretaceous period in Africa and possibly also South America about 99-97 million years ago. R ...
'', and '' Andesaurus''. 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 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fill in more information about abelisaur anatomy as it contained portions of the skeleton that were previously unknown, unpublished, or poorly preserved in other specimens.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 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''. 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'' and '' Majungasaurus'', and while ''
Abelisaurus ''Abelisaurus'' (; "Abel's lizard") is a genus of predatory abelisaurid theropod dinosaur alive during the Late Cretaceous Period (Campanian) of what is now South America. It was a bipedal carnivore that probably reached about in length, althou ...
'' 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. The article by Calvo ''et al.'' in 2004, also contained a preliminary cladistic analysis of the phylogenetic 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 Carnotaurini is a tribe of the theropod dinosaur family Abelisauridae from the Late Cretaceous period of Patagonia. It includes the dinosaurs ''Carnotaurus sastrei''; the type species, ''Aucasaurus garridoi'', and ''Abelisaurus comahuensis''. Thi ...
than to '' Ceratosaurus''. In 2008, a study recovered it as the sister species of ''Skorpiovenator''.


Paleoecology

''Ekrixinatosaurus'' was found in the red beds of the
Candeleros Formation The Candeleros Formation (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
, which has yielded a wide variety of vertebrates. It shared its environment with the titanosaurian
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 bo ...
'' Andesaurus'' and the
rebbachisaurid Rebbachisauridae is a 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 McIntosh included t ...
sauropods ''
Limaysaurus ''Limaysaurus'' ("Limay lizard") is a genus represented by a single species of rebbachisaurid sauropod dinosaurs, which lived during the mid-Cretaceous period, about 99.6 to 97 million years ago, in the Cenomanian, in what is now South America ( ...
'' and '' Nopcsaspondylus''. Iguanodont ornithischian remains have reportedly also been found. The carcharodontosaurid '' Giganotosaurus'' was possibly the apex predator. Smaller predators also inhabited the area, including the
dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
'' Buitreraptor'', the
alvarezsauroid Alvarezsauroidea is a group of small maniraptoran dinosaurs. Alvarezsauroidea, Alvarezsauridae, and Alvarezsauria are named for the historian Gregorio Álvarez, not the more familiar physicist Luis Alvarez, or his son geologist Walter Alvarez ...
''
Alnashetri ''Alnashetri'' is an extinct genus of alvarezsaurid coelurosaurian theropod dinosaur known from the early Late Cretaceous ( Cenomanian stage) of La Buitrera, Río Negro Province, Argentina. It contains one known species, ''Alnashetri cerropolici ...
'', and the basal
coelurosaurian 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, tyrann ...
''
Bicentenaria ''Bicentenaria'' (meaning "two hundred year anniversary"; named after the 200th anniversary of the 1810 May Revolution in Argentina) is a genus of carnivorous coelurosaurian theropods which lived during the early Late Cretaceous (about 90 mi ...
''. Other primitive reptiles lived in the area, such as the primitive snake '' Najash'', the crocodile '' Araripesuchus'', along with turtles, fish, pipid frogs, and mammals. Pterosaurs 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 (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
, suggesting significant activity in the area. Recent studies on
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
n theropods have interpreted abelisauroids as modest and medium-sized dinosaurs that co-occurred with giant carcharodontosaurids during the Early and early Late Cretaceous. 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'' 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 a large igneous province (LIP) located on the Antarctic Plate, in the southern Indian Ocean. It is about to the southwest of Australia and is ...
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 focused on the ceratosaurs, a group of relatively primitive, often horned, predatory theropod dinosaurs that became the apex predators of ...


References

{{Portal bar, Dinosaurs, Cretaceous Brachyrostrans Cenomanian life Late Cretaceous dinosaurs of South America Cretaceous Argentina Fossils of Argentina Candeleros Formation Fossil taxa described in 2004