''Orkoraptor'' 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 medium-sized
megaraptora
Megaraptora is a clade of carnivorous Tetanurae, tetanuran theropod dinosaurs with controversial relations to other theropods. Its Derived (phylogenetics), derived members, the Megaraptoridae are noted for their elongated hand claws and proporti ...
n
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 late
Cretaceous Period
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of t ...
of
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 ...
. It is known from incomplete
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
remains including parts of the skull, teeth, tail vertebrae, and a partial
tibia
The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
. The specialized teeth resemble those of some
maniraptoriform
Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings that contains ornithomimosaurs and maniraptorans. This group was named by Thomas Holtz, who defined it as "the most recent common ancestor of ''Ornithomimus'' and bir ...
theropods, namely the
deinonychosaurs
Deinonychosauria is a clade of paravian dinosaurs which lived from the Late Jurassic to the Late Cretaceous periods. Fossils have been found across the globe in North America, Europe, Africa, Asia, South America, and Antarctica,Case, J.A., Martin ...
and
compsognathids
Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other d ...
. This and other anatomical features led the authors who described it (Novas, Ezcurra, and Lecuona) to suggest that it was a
maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
n
coelurosaur
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 ...
. However, subsequent studies found it to be a
megaraptora
Megaraptora is a clade of carnivorous Tetanurae, tetanuran theropod dinosaurs with controversial relations to other theropods. Its Derived (phylogenetics), derived members, the Megaraptoridae are noted for their elongated hand claws and proporti ...
n.
Found in the
Cerro Fortaleza Formation
The Cerro Fortaleza Formation, in older literature described as Pari Aike Formation, is a Late Cretaceous geologic formation of Campanian to Maastrichtian age (although it has formerly been reported to be Cenomanian to Santonian)Varela et al., 20 ...
of Southern
Patagonia
Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
, it is one of the southernmost carnivorous dinosaurs known from South America.
Etymology
The name ''Orkoraptor'' means "Toothed River thief", and was derived from the
Aoniken "Orr-Korr", the local name for the
La Leona River
The La Leona River is a river of Patagonia, Argentina. It flows in the eastern part of the Los Glaciares National Park. It has its origin to the south of Lake Viedma and winds for about 50 kilometers (30 miles) through the Andes before flowing int ...
, located near the original fossil site. The species name honors Coleman Burke, an amateur paleontologist who supported the expedition that collected the original fossils.
Description
''Orkoraptor'' was a medium-sized theropod. In 2010,
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 ...
estimated its length as and mass as 500 kilograms.
In 2016 Molina-Pérez and Larramendi gave a higher estimation of 8.4 meters (27.5 ft) and 1.4 tonnes (1.5 short tons) for the holotype specimen. 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 ...
, MPM-Pv 3457 was excavated in 2001 and consists of a right
postorbital
The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
, a right
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians.
Anatomy and function
In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
, a probable right coronoid, eight isolated teeth, the intercentrum of the
atlas
An atlas is a collection of maps; it is typically a bundle of maps of Earth or of a region of Earth.
Atlases have traditionally been bound into book form, but today many atlases are in multimedia formats. In addition to presenting geographic ...
and its right neurapophysis, two proximal caudal vertebrae, the proximal half of the right
tibia
The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, eight fragmentary ribs, and three incomplete chevrons. It was initially believed to be a
coelurosaur
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 ...
due to the upturned anterior process of the postorbital as well as specialized teeth which are unserrated on the mesial (front) side and have an 8-shaped cross section, similar to those of
dromaeosaurids
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 ...
and
compsognathids
Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other d ...
. However, many other features contradict those of these families and other coelurosaurs, so its original describers were unable to conclusively assign it to any specific family. They did note its similarity to a then-undescribed Patagonian theropod now known as ''
Murusraptor
''Murusraptor'' ("wall thief") is a genus of carnivorous megaraptoran theropod dinosaur from the Sierra Barrosa Formation, part of the Neuquén Group of Patagonia, in Argentina, South America. It is known from a single specimen that consists of ...
''.
More recent analyses have explained that these coelurosaur-like traits are also present in some megaraptorans. For example, ''Orkoraptor''
's postorbital is almost identical to that of ''
Aerosteon
''Aerosteon'' is a genus of megaraptoran dinosaur from the Late Cretaceous period of Argentina. Its remains were discovered in 1996 in the Anacleto Formation, which is from the late Campanian. The type and only known species is ''A. riocolorade ...
''
and the juvenile ''
Megaraptor
''Megaraptor'' () is a genus of large theropod dinosaur that lived in the ages of the Late Cretaceous. Its fossils have been discovered in the Patagonian Portezuelo Formation of Argentina, South America. Initially thought to have been a giant dr ...
'' described in 2014 has teeth similar to those of ''Orkoraptor''.
Classification
''Orkoraptor'' was originally classified as
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
''
incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''. While the majority of phylogenetic trees recovered in its describers' analysis recovered it as a
compsognathid
Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other ...
, the describers considered this unlikely based on its much larger size and presence in much younger strata.
''Orkoraptor'' was subsequently recovered in
Megaraptora
Megaraptora is a clade of carnivorous Tetanurae, tetanuran theropod dinosaurs with controversial relations to other theropods. Its Derived (phylogenetics), derived members, the Megaraptoridae are noted for their elongated hand claws and proporti ...
, a clade including several other enigmatic medium to large theropods, which has variously been considered to be a member of
Allosauroidea
Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four family (biology), families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroid ...
and
Tyrannosauroidea
Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent b ...
.
The phylogenetic trees in the publication of ''Gualicho shinyae
''Gualicho'' (named in reference to the gualichu) is a genus of theropod dinosaur. The type species is ''Gualicho shinyae''. It lived in what is now northern Patagonia, on what was then a South American island continent split off from the superco ...
'' have found megaraptorans to be either allosauroids or basal coelurosaurs.
Provenance
All known specimens of ''Orkoraptor'' were collected from the Pari Aike Formation
The Cerro Fortaleza Formation, in older literature described as Pari Aike Formation, is a Late Cretaceous geologic formation of Campanian to Maastrichtian age (although it has formerly been reported to be Cenomanian to Santonian)Varela et al., 20 ...
, which is now considered by different authors to be either the middle section of the Mata Amarilla Formation
The Mata Amarilla Formation is a fossiliferous formation of the Austral Basin in southern Patagonia, Argentina. The formation consists of sediments deposited during the Middle Cenomanian, dated to 96.94 to 95.52 Ma. The middle section of the fo ...
, or merely an alternative name for the Cerro Fortaleza Formation
The Cerro Fortaleza Formation, in older literature described as Pari Aike Formation, is a Late Cretaceous geologic formation of Campanian to Maastrichtian age (although it has formerly been reported to be Cenomanian to Santonian)Varela et al., 20 ...
. It was originally identified as coming from the 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 from ...
, and thus the youngest known megaraptora
Megaraptora is a clade of carnivorous Tetanurae, tetanuran theropod dinosaurs with controversial relations to other theropods. Its Derived (phylogenetics), derived members, the Megaraptoridae are noted for their elongated hand claws and proporti ...
n, but more recently is considered to be from the Cenomanian
The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
to Santonian
The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...
. The middle section of the Mata Amarilla Formation, whence ''Orkoraptor'' was collected, contains a tuff layer that has been dated to 96.2 ± 0.7 Ma, during the Cenomanian
The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
. However, the site where ''Orkoraptor'' is known from, Cerro Hornos, was recently reinterpreted as coming from 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. Campanian s ...
.
References
{{Taxonbar, from=Q2101463
Megaraptorans
Late Cretaceous dinosaurs of South America
Cretaceous Argentina
Fossils of Argentina
Cenomanian genus first appearances
Santonian genus extinctions
Fossil taxa described in 2008
Taxa named by Fernando Novas