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''Austroraptor'' ( ) is a
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 large
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 ...
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 ...
that lived during the
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'', ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
in what is now
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 ...
. ''Austroraptor'' was a large-sized, moderately-built, ground-dwelling,
bipedal 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' ...
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 s ...
, estimated at in length. It is one of the largest dromaeosaurids known, with only ''
Achillobator ''Achillobator'' ( ; meaning "Achilles hero") is a genus of large dromaeosaurid dinosaurs that lived in Asia during the Late Cretaceous period about 96 million to 89 million years ago in what is now the Bayan Shireh Formation. The genus is curren ...
'', ''
Dakotaraptor ''Dakotaraptor'' (meaning “thief from Dakota”) is a potentially chimaeric genus of large dromaeosaurid theropod dinosaur that lived in North America during the Late Cretaceous period. The remains have been found in the Maastrichtian stage of ...
'', and ''
Utahraptor ''Utahraptor'' (meaning "Utah's thief") is a genus of large dromaeosaurid dinosaur that lived in North America during the Early Cretaceous period. It was a heavy-built, ground-dwelling, bipedal carnivore. It contains a single species, ''Utah ...
'' approaching or surpassing it in length.


Discovery and naming

The type specimen of ''Austroraptor cabazai'',
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 ...
MML-195, was recovered in the Bajo de Santa Rosa locality of the
Allen Formation The Allen Formation is a geological formation in Argentina whose strata date back to the Late Cretaceous (middle Campanian to early Maastrichtian.Salgado et al., 2007 Dinosaur remains are among the fossils that have been recovered from the formati ...
, in Río Negro,
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 ...
. The specimen was collected in 2002 by the team of
Fernando Emilio Novas Fernando Emilio Novas (born 1960) is an Argentine paleontologist working for the Comparative Anatomy Department of the Bernardino Rivadavia Natural Sciences Museum in Buenos Aires, Argentina.
of the '' Museo Argentino de Ciencias Naturales''. It consists of a fragmentary skeleton including parts of the skull, lower jaw, a few neck and torso vertebrae, some ribs, a humerus, and assorted bones from both legs. The specimen was prepared by Marcelo Pablo Isasi and Santiago Reuil. In 2008, 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 ...
''Austroraptor cabazai'' was named and described by Fernando Emilio Novas, Diego Pol, Juan Canale, Juan Porfiri and Jorge Calvo. The genus name ''Austroraptor'' means "Southern thief," and is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word ''auster'' meaning "the south wind" and the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word ''raptor'' meaning "thief." The specific name ''cabazai'' was chosen in honor of Héctor "Tito" Cabaza, who founded the '' Museo Municipal de Lamarque'' where the specimen was partially studied. In 2012,
Phil Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
and Ariana Paulina-Carabajal referred a second specimen to ''Austroraptor cabazai'', MML-220, that was found in 2008. This specimen, a partial skeleton with the skull of an adult individual slightly smaller than the holotype, is also housed in the collection of the Museo Municipal de Lamarque in Argentina. It complements the holotype in several elements, mainly the lower arm, hand and foot.


Description

It is the largest
dromaeosaur 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 ...
to be discovered in the Southern Hemisphere; Novas ''et al.'' estimated that ''Austroraptor'' measured in length from head to tail.
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
later estimated its length at and weight of . However, Thomas R. Holtz Jr. has suggested that its weight is comparable to a lion, around .Supplementary Information 2012
/ref> The skull is low and elongated, much more so than that of other dromaeosaurs, and measures . ''Austroraptor'' has conical, non-serrated teeth, which Novas ''et al.'' compared to those of
spinosaurids The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
, based on how the enamel of the surface of its teeth is fluted. ''Austroraptor'' shares a trait that is unique to it and to '' Adasaurus'': the descending process of the lacrimal bones curves anteriorly to a large degree. ''Austroraptor'' has a bizarre morphology in its pedal phalanges, which are strangely disproportionate. Phalanx IV-2 is over twice the width of phalanx II-2, and nearly three times the expected width based on similarly sized members of its taxonomic family. This has suggested to some researchers that the holotype specimen is a paleontological chimera; however, there is no uncertainty about the affinity of the taxon, so a chimera hypothesis can not be assured. Several of ''Austroraptor'''s skull bones bear some resemblance to those of the smaller
troodontids Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
. The front limbs of ''Austroraptor'' were short for a dromaeosaur, with its humerus less than half the length of its
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 ...
. Among the Dromaeosauridae, only this genus, ''
Tianyuraptor ''Tianyuraptor'' is a genus of short-armed dromaeosaurid dinosaur ('running lizard'; a type of small dinosaur considered to be closely related to birds) that lived during the Early Cretaceous, about 122 million years ago. Its remains have bee ...
'', ''
Zhenyuanlong ''Zhenyuanlong'' (meaning ''"Zhenyuan's dragon"'', from Chinese Pinyin 龙/ 龍 lóng ''"dragon"'') is a genus of dromaeosaurid dinosaur from the Yixian Formation of Liaoning, China. It lived during the Aptian age of the early Cretaceous perio ...
'' and ''
Mahakala Mahākāla is a deity common to Hinduism and Tantric Buddhism. In Buddhism, Mahākāla is regarded as the sacred '' Dharmapāla'' ("Protector of the Dharma"), while in Hinduism, Mahākāla is a fierce manifestation of the Hindu god Shiva and th ...
'' have similarly reduced forelimbs. The relative length of its arms has caused ''Austroraptor'' to be compared to another, more famous short-armed dinosaur, ''
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 ...
,'' though there is no close relationship between the two taxa.


Distinguishing traits

However little of the entire skeleton was found, the bones that are available for analysis possess some distinct characteristics that differentiate ''Austroraptor'' from other dromaeosaurs. ''Austroraptor'' is particularly notable because of its relatively short forearms, which are much shorter in proportion when compared to the majority of the members of Dromaeosauridae. According to Novas ''et al.'' 2008, ''Austroraptor'' can be distinguished based on the following characteristics: *A lacrimal that is highly pneumatized, with the descending process strongly curved rostrally, and with a caudal process flaring out horizontally above the orbit. *The lack of a dorsomedial process on the postorbital bone for articulation with the
frontal Front may refer to: Arts, entertainment, and media Films * ''The Front'' (1943 film), a 1943 Soviet drama film * ''The Front'', 1976 film Music * The Front (band), an American rock band signed to Columbia Records and active in the 1980s and e ...
bone, and with the squamosal process extremely reduced. *The maxillary and dentary teeth are small, conical, devoid of serrations and fluted. *The humerus is short, at approximately 46% of length of the femur. *The pedal phalanx II-2 is transversely narrow, contrasting with the extremely robust phalanx IV-2. In 2012, comparison with a second specimen showed that the fourth toe was not especially broad; the purported second phalanx had in fact been a first phalanx.


Classification

A
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 of the holotype specimen's anatomical features by the describers placed ''Austroraptor'' within the subfamily
Unenlagiinae Unenlagiinae is a subfamily of long-snouted paravian theropods. They are traditionally considered to be members of Dromaeosauridae, though some authors place them into their own family, Unenlagiidae, alongside the subfamily Halszkaraptorinae. ...
of Dromaeosauridae. This assignment was based on characteristics observed in the bones of the skull, the teeth, and the geometry and formation of the specimen's vertebral elements. It was determined that ''Austroraptor'' was a close relative of the unenlagiine dromaeosaur ''
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 go ...
'', with which it shares certain derived characteristics of the neck vertebrae. 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 ...
is based on the phylogenetic analysis conducted by Turner, Makovicky and Norell in 2012, showing the relationships of ''Austroraptor'' among the other genera assigned to the taxon
Unenlagiinae Unenlagiinae is a subfamily of long-snouted paravian theropods. They are traditionally considered to be members of Dromaeosauridae, though some authors place them into their own family, Unenlagiidae, alongside the subfamily Halszkaraptorinae. ...
: Cau ''et al.'' 2017 published a phylogenetic analysis of the Dromaeosauridae during the description of
Halszkaraptor ''Halszkaraptor'' (; meaning " Halszka's seizer") is a genus of waterfowl-like dromaeosaurid dinosaurs from Mongolia that lived during the Late Cretaceous period. It contains only one known species, ''Halszkaraptor escuilliei''. The type speci ...
, in which members of the Unenlagiinae are classified as: In 2019, during the description of ''
Hesperornithoides ''Hesperornithoides'' (meaning "western bird form"; nicknamed "Lori") is a genus of troodontid theropod dinosaur that lived in North America during the Late Jurassic period. Discovery Although several Troodontid teeth were found, with the trood ...
'', many paravian groups were examined for the inclusion of the new
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 ...
. In this analysis, ''Austroraptor'' is found to be a more basal member of Unenlagiinae.


Paleobiology

It has been suggested that unenlagiines had better capacities for running and pursuit predation than other dromaeosaurids. While
Laurasian Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
dromaeosaurids (
Eudromaeosauria Eudromaeosauria ("true dromaeosaurs") is a subgroup of terrestrial dromaeosaurid theropod dinosaurs. They were relatively small to medium-sized, feathered hypercarnivores (with diets consisting almost entirely of other terrestrial vertebrates) ...
) were more stocky and had shorter legs and had an active predatory lifestyle, unenlagiines could likely maintain high speeds for extended amounts of time because they were more gracile. Unenlagiines have modified metatarsals that resemble those of microraptorines: they are relatively thin and lengthened. Based on these adaptations, it is likely that unenlagiines preyed on small, fast animals, although the exact animals are unknown. ''
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 go ...
'' features particular traits that can be attributed to specific hunting methods. Models for ''Buitreraptor'' propose that it hunted by traveling large distances in pursuit of prey, which may explain the long-legged trait shared by various genera of Unenlagiidae. ''Buitreraptor'' is characterized by its long forelimbs and hands; it likely relied on them to restrain prey and the curved claw of the second pedal digit would have injured or killed the victim. ''Buitreraptor'' probably swallowed its prey whole due to its lack of serrated teeth with flesh-tearing capabilities; the teeth functioned to simply hold prey. The same model was proposed for the much larger ''Austroraptor'' with few exceptions: *It would not have used its arms to handle prey, due to their relatively small size. *Its teeth were conical and probably stronger, so it may have been able to use them for hunting larger prey. The teeth of ''Austroraptor'' are conical and lacking denticles, similarly to those of
spinosaurids The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
. Since it probably had
subarctometatarsal
condition and similar hindlimb lengths compared to ''Buitreraptor'', ''Austroraptor'' likely had well-developed
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often us ...
capacities. An unpublished theory is that austroraptor may have had a piscivorous diet. Austroraptor had non-serrated conical teeth, similar to those of
spinosaurids The Spinosauridae (or spinosaurids) are a clade or family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous period. Spinosaurid fossils have been recovered worldwide, includi ...
, relatively short forearms, and proportionally long legs which may have aided it in wading.


Paleoecology

The holotype specimen was found in terrestrial sediments that were deposited during the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...
stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...
of the
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'', ...
period. Thomas R. Holtz Jr. has estimated that ''Austroraptor'' lived between 78 million and 66 million years ago, until the end of the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
era. ''Austroraptor'' shared its paleoenvironment in the
Allen Formation The Allen Formation is a geological formation in Argentina whose strata date back to the Late Cretaceous (middle Campanian to early Maastrichtian.Salgado et al., 2007 Dinosaur remains are among the fossils that have been recovered from the formati ...
with diverse dinosaurs and early mammals. The discovery of ''Austroraptor'' increases the understanding of ecological and morphological diversity among unenlagiines, demonstrating that members of the subfamily included giant short-armed and small long-armed members, and suggesting that during the end of the Cretaceous large
coelurosaurs 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, tyran ...
became common after the decreasing dominance of
carcharodontosaurids Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
. This genus represents the earliest record of
Gondwanan 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 stage ...
dromaeosaurids, and supports the fact that large dromaeosaurids took the role of large
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
alongside
abelisaurids 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 ...
such as ''
Quilmesaurus ''Quilmesaurus'' is a genus of carnivorous abelisaurid theropod dinosaur from the Patagonian Upper Cretaceous (Campanian stage) of Argentina. It was a member of Abelisauridae, closely related to genera such as ''Carnotaurus''. The only known rema ...
''. During his description of '' Pellegrinisaurus'', Salgado suggested that large
titanosaurs Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still th ...
and
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 ...
inhabited interior environments of the region, and contemporaneous hadrosaurids and the titanosaur ''
Aeolosaurus ''Aeolosaurus'' (; "Aeolus' lizard") is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now South America. Like most sauropods, it would have been a quadrupedal herbivore with a long neck and tail. ''Aeolos ...
'' inhabited coastal lowlands. The diversity among titanosaurs indicates that the
Allen Formation The Allen Formation is a geological formation in Argentina whose strata date back to the Late Cretaceous (middle Campanian to early Maastrichtian.Salgado et al., 2007 Dinosaur remains are among the fossils that have been recovered from the formati ...
had environments that supported a great range of
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
dinosaurs. Other contemporaneous paleofauna includes the titanosaurs '' Laplatasaurus'', '' Rocasaurus'' and ''
Saltasaurus ''Saltasaurus'' (which means "lizard from Salta") is a genus of saltasaurid dinosaur of the Late Cretaceous period of Argentina. Small among sauropods, though still heavy by the standards of modern creatures, ''Saltasaurus'' was characterized by ...
'', the
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
''
Limenavis ''Limenavis'' is a prehistoric bird genus from the Late Cretaceous. It lived about 70 million years ago, around the Campanian-Maastrichtian boundary. Known from several broken bones, the remains of the only known species ''Limenavis patagonica'' ...
'', and the hadrosaurids ''
Bonapartesaurus ''Bonapartesaurus'' (meaning "Bonaparte's lizard", named after José Bonaparte) is an extinct genus of herbivorous ornithopod dinosaur belonging to Hadrosauridae, which lived in the area of modern Argentina during the Campanian and Maastrichtia ...
'', ‘’ Kelumapusaura’’, and '' Lapampasaurus''.


See also

*
Allen Formation The Allen Formation is a geological formation in Argentina whose strata date back to the Late Cretaceous (middle Campanian to early Maastrichtian.Salgado et al., 2007 Dinosaur remains are among the fossils that have been recovered from the formati ...
*
Timeline of dromaeosaurid research This timeline of dromaeosaurid research is a chronological listing of events in the history of paleontology focused on the dromaeosaurids, a group of sickle-clawed, bird-like theropod dinosaurs including animals like ''Velociraptor''. Since the ...
*
2008 in paleontology Protozoa New taxa Plants Angiosperms Monocots Arthropoda Arachnids Insects Xiphosurans Fishes Bony fish Placoderms General research *Hilton & Grande redescribe the fossil mooneyes of western North America synonymizing ...


Notes


References


External links

* * {{Taxonbar, from=Q132257 Allen Formation Campanian life Cretaceous Argentina Fossil taxa described in 2008 Fossils of Argentina Late Cretaceous dinosaurs of South America Taxa named by Fernando Novas Unenlagiines