Carnosaur
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Carnosauria is an extinct large group of
predator 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 ...
y
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 ...
s that lived during the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
and
Cretaceous 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 ...
periods. Starting from the 1990s, scientists have discovered some very large carnosaurs in 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 ...
family, such as ''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period, approximately 99.6 to 95 million years ago. The holotype specimen was discovered in th ...
'', ''
Mapusaurus ''Mapusaurus'' () was a giant carcharodontosaurid carnosaurian dinosaur from the early Late Cretaceous (early Turonian stage), approximately 93.9 to 89.6 million years ago, of what is now Argentina. Discovery ''Mapusaurus'' was excavated b ...
'', ''
Carcharodontosaurus ''Carcharodontosaurus'' (; ) is a genus of large carcharodontosaurid theropod dinosaur that existed during the Cenomanian age of the Late Cretaceous in Northern Africa. The genus ''Carcharodontosaurus'' is named after the shark genus '' Carc ...
'' and ''
Tyrannotitan ''Tyrannotitan'' (; ) is a genus of huge bipedal carnivorous dinosaur of the carcharodontosaurid family from the Aptian stage of the early Cretaceous period, discovered in Argentina. It is closely related to other giant predators like ''Carchar ...
'' which are among the largest known predatory dinosaurs. While it originally contained a wide assortment of giant
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 ...
s that were not closely related, the group has since been defined to encompass only the allosaurs and their closest kin. However, with the description and publication in 2019 of '' Asfaltovenator vialidadi'', a basal allosauroid curiously displaying both primitive and derived features seen in
Tetanurae Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans (including birds). Tetanurans ar ...
, the new phylogenetic analysis has found
Megalosauroidea Megalosauroidea (meaning 'great/big lizard forms') is a superfamily (or clade) of tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period. The group is defined as '' Megalosaurus bucklandii'' and all taxa s ...
to be a basal
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also ref ...
of carnosaurs in respect to Allosauroidea; thus significantly expanding Carnosauria's inclusiveness towards its original context. Distinctive characteristics of carnosaurs include large eyes, a long narrow
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
and modifications of the
leg A leg is a weight-bearing and locomotive anatomical structure, usually having a columnar shape. During locomotion, legs function as "extensible struts". The combination of movements at all joints can be modeled as a single, linear element ca ...
s and pelvis such as the thigh (
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 ...
) being longer than the shin (
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 ...
). Carnosaurs first appeared in the Middle Jurassic, around 176 mya. The last definite known carnosaurs, the carcharodontosaurs, became extinct in the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded b ...
epoch of the Cretaceous, roughly 90 mya; reportedly later remains of carcharodontosaurids, from the late
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 ...
(70–66 Ma ago)
Bauru Group The Bauru Group is a geological group of the Bauru Sub-basin, Paraná Basin in Minas Gerais, São Paulo, General Salgado, Itapecuru-Mirim, Mato Grosso, Brazil whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils ...
in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, were later interpreted as those of
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 ...
. The phylogenetically problematic
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 ...
ns, which may or may not be carnosaurs, became extinct around 66 mya. '' Unquillosaurus'' is another possible Carnosaurian Theropod that has an extended range, hailing from rocks dated to 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 ...
, around 75-70 mya.


Systematics

Modern
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 defines Carnosauria as those tetanurans sharing a more recent common ancestor with ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'' than with modern birds.


Taxonomy

Carnosauria has traditionally been used as a dumping ground for all large theropods. Even non-dinosaurs, such as the
rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilians ...
n ''
Teratosaurus ''Teratosaurus'' is a genus of rauisuchians known from the Triassic Stubensandstein ( Löwenstein Formation - Norian stage) of Germany. It is estimated to be 6 meters (19.5 ft) long. Discovery In 1860, Sixt Friedrich Jakob von Kapff a ...
'', were once considered carnosaurs. However, analysis in the 1980s and 1990s revealed that other than size, the group shared very few characteristics, making it
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
. Most former carnosaurs (such as the megalosaurids, the spinosaurids, and the
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) were
reclassified ''Reclassified'' is the reissue of Australians, Australian rapper Iggy Azalea's debut studio album, ''The New Classic'' (2014). It was released internationally on 21 November 2014 by Virgin EMI Records, and in the United States on 24 November ...
as more primitive theropods. Others (such as the
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to thirteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
s) that were more closely related to birds were placed in
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, t ...
.


Phylogeny

The clade Allosauroidea was originally proposed by
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 Zhao (1993; p. 2079), and later used as an undefined
stem-based taxon Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...
by
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
(1997). Sereno (1998; p. 64) was the first to provide a stem-based definition for the Allosauroidea, defining the clade as "All neotetanurans closer to ''Allosaurus'' than to
Neornithes 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 ...
."
Kevin Padian Kevin Padian (born 1951) is a Professor of Integrative Biology at the University of California, Berkeley, Curator of Paleontology, University of California Museum of Paleontology and was President of the National Center for Science Education fro ...
(2007) used a node-based definition, defined the Allosauroidea as ''Allosaurus'', ''
Sinraptor ''Sinraptor'' is a genus of metriacanthosaurid theropod dinosaur from the Late Jurassic. The name ''Sinraptor'' comes from the Latin prefix "Sino", meaning Chinese, and "raptor" meaning robber. The specific name ''dongi'' honours Dong Zhiming. ...
'', their
most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
, and all of its descendants. Thomas R. Holtz and colleagues (2004; p. 100) and
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 Ken Carpenter (2000), among others, have followed this node-based definition. However, in some analyses (such as Currie & Carpenter, 2000), the placement of the carcharodontosaurids relative to the allosaurids and sinraptorids is uncertain, and therefore it is uncertain whether or not they are allosauroids (Currie & Carpenter, 2000). The
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 ...
presented here follows the 2010 analysis by Benson, Carrano and Brusatte. In 2019, Rauhut and Pol described '' Asfaltovenator vialidadi'', a basal allosauroid displaying a mosaic of primitive and derived features seen within
Tetanurae Tetanurae (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most theropod dinosaurs, including megalosauroids, allosauroids, tyrannosauroids, ornithomimosaurs, compsognathids and maniraptorans (including birds). Tetanurans ar ...
. Their phylogenetic analysis found traditional
Megalosauroidea Megalosauroidea (meaning 'great/big lizard forms') is a superfamily (or clade) of tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period. The group is defined as '' Megalosaurus bucklandii'' and all taxa s ...
to represent a basal
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also ref ...
of carnosaurs, paraphyletic with respect to Allosauroidea.


"Carnosaurus"

"Carnosaurus" is an
informal Formal, formality, informal or informality imply the complying with, or not complying with, some set of requirements (forms, in Ancient Greek). They may refer to: Dress code and events * Formal wear, attire for formal events * Semi-formal attir ...
generic name, attributed to Friedrich von Huene, ca. 1929. It is the result of a
typographical error A typographical error (often shortened to typo), also called a misprint, is a mistake (such as a spelling mistake) made in the typing of printed (or electronic) material. Historically, this referred to mistakes in manual type-setting (typography) ...
created by the translation of the von Huene monograph from German to Spanish. Von Huene himself intended to assign indeterminate remains to Carnosauria ''
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, uncertain ...
''.


See also

portal:dinosaur
Carnotaurus ''Carnotaurus'' (; ) is a genus of theropod dinosaur that lived in South America during the Late Cretaceous period, probably sometime between 71 and 69 million years ago. The only species is ''Carnotaurus sastrei''. Known from a single well-p ...


References


Sources

* (on "Carnosaurus") * * Holtz, T. R., Jr. and Osmólska H. 2004. Saurischia; pp. 21–24 in D. B. Weishampel, P. Dodson, and H. Osmólska (eds.), ''The Dinosauria'' (2nd ed.), University of California Press, Berkeley. * *


External links

* {{Taxonbar, from=Q131190 Aalenian first appearances Turonian extinctions Fossil taxa described in 1920 Taxa named by Friedrich von Huene