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Metriacanthosauridae is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
allosauroid Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroidea, were among th ...
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 ...
s that lived from the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
to the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
. When broken down into its Greek roots, it means "moderately-spined lizards". The family is split into two subgroups: Metriacanthosaurinae, which includes dinosaurs closely related to ''
Metriacanthosaurus ''Metriacanthosaurus'' (meaning "moderately-spined lizard") is a genus of metriacanthosaurid dinosaur from the upper Oxford Clay of England, dating to the Late Jurassic period, about 160 million years ago (lower Oxfordian). History of discovery ...
'', and another group composed of the close relatives of ''
Yangchuanosaurus ''Yangchuanosaurus'' is an extinct genus of metriacanthosaurid theropod dinosaur that lived in China from the Middle to Late Jurassic periods (Bathonian to Oxfordian stages), and was similar in size and appearance to its North American and Eur ...
''. Metriacanthosaurids are considered
carnosaurs Carnosauria is an extinct large group of predatory dinosaurs that lived during the Jurassic and Cretaceous periods. Starting from the 1990s, scientists have discovered some very large carnosaurs in the carcharodontosaurid family, such as ''Gi ...
, belonging to the
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 ...
superfamily. The group includes species of large range in body size. Of their physical traits, most notable are their neural spines.Bailey, Jack Bowman. "Neural Spine Elongation in Dinosaurs: Sailbacks or Buffalo-Backs?" Journal of Paleontology, vol. 71, no. 06, 1997, pp. 1124–1146., doi:10.1017/s0022336000036076. The records of the group are mostly confined to Asia, though ''Metriacanthosaurus'' is known from Europe. Metriacanthosauridae is used as a senior synonym of Sinraptoridae.


Diagnostic traits

Metriacanthosaurids share the following unambiguous
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
among allosauroids: * A short or absent anterior ramus of the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
(also found in
carcharodontosaurids Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
and piatnitzkysaurids). * The laterosphenoid articulated on 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 ea ...
and
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
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal The quadratojugal is a skull bone present in many vertebrates, including ...
without constriction of the lower temporal fenestra. (also found in megalosauroids) * A flange on the squamosal covering the quadrate head laterally. * A well-defined longitudinal groove on the lateral side of the dentary housing a row of neurovascular foramina. * Broad, well developed spinopost-zygapophyseal lamina on the
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis *Axis (mathematics), a designator for a Cartesian-coordinate ...
. * A manus shorter than the forearm. * Subrectangular and sheet-like neural spines of middle caudal vertebrae. * A manus without digit V or the phalanges of digit IV (also found in neovenatorids). * A heart-shaped cross section of the ilium's paired midshafts (also found in coelophysids). * Fused distal end of the
ischium The ischium () form ...
(also found in neovenatorids and basal
tetanurans 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 ...
). Metriacanthosaurids share the following dental synapomorphies among theropods: * surface centrally positioned on the labial surface of the crown roughly flattened in lateral teeth * irregular and non-oriented enamel surface texture Metriacanthosaurines share the following synapomorphies among metriacanthosaurids: * The anteroventral border of the maxillary
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
being demarcated by a raised ridge (also found in ''
Eoraptor ''Eoraptor'' () is a genus of small, lightly built, basal sauropodomorph. One of the earliest-known dinosaurs, it lived approximately 231 to 228 million years ago, during the Late Triassic in Western Gondwana, in the region that is now northwest ...
'', coelophysids, ''
Masiakasaurus ''Masiakasaurus'' is a genus of small predatory noasaurid theropod dinosaurs from the Late Cretaceous of Madagascar. In Malagasy, ''masiaka'' means "vicious"; thus, the genus name means "vicious lizard". The type species, ''Masiakasaurus knopfl ...
'', ''
Marshosaurus ''Marshosaurus'' is a genus of medium-sized carnivorous theropod dinosaur, belonging to the Megalosauroidea, from the Late Jurassic Morrison Formation of Utah and possibly Colorado. Description ''Marshosaurus'' was medium-sized for a theropod. ...
,'' and ''
Compsognathus ''Compsognathus'' (; Greek ''kompsos''/κομψός; "elegant", "refined" or "dainty", and ''gnathos''/γνάθος; "jaw") is a genus of small, bipedal, carnivorous theropod dinosaur. Members of its single species ''Compsognathus longipes'' c ...
''). * A pronounced ventral keel on the anterior dorsal vertebrae (also found in ''
Condorraptor ''Condorraptor'' is an extinct genus of megalosauroid theropod dinosaur. Its genus name means 'robber from Cerro Condor', referencing a nearby village, while its species name, ''currumili'', is named after Hipolito Currumil, the landowner and di ...
'', ''
Piatnitzkysaurus ''Piatnitzkysaurus'' ( meaning "Piatnitzky's lizard") is a genus of megalosauroid theropod dinosaur that lived approximately 179 to 177 million years ago during the lower part of the Jurassic Period in what is now Argentina. ''Piatnitzkysaurus'' ...
'', ''
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 ''Carcha ...
'' and many megalosaurians). * A straight posterior margin of the iliac postacetabular process. * The angle between the long axes of the
pubic In vertebrates, the pubic region ( la, pubis) is the most forward-facing (ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior r ...
shaft and boot being less than 60 degrees. * A ventrally curved ischial shaft (also found in coelophysids, ''
Eustreptospondylus ''Eustreptospondylus'' ( ; meaning "true ''Streptospondylus''") is a genus of megalosaurid Theropoda, theropod dinosaur, from the Oxfordian stage of the Late Jurassic period (some time between 163 and 154 million years ago) in southern Eng ...
'', ''
Afrovenator ''Afrovenator'' (; "African hunter") is a genus of megalosaurid theropod dinosaur from the Middle Jurassic Period of northern Africa. Discovery and naming The remains of ''Afrovenator'' were discovered in 1993 in the Tiourarén Formation ...
'', ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years a ...
'' and ''Compsognathus''). * A bulbous fibular crest on the
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 ...
(also found in '' Sinosaurus'').


Classification

Carrano, Benson & Sampson (2012) noted that the name Metriacanthosauridae should be used as it has priority over Sinraptoridae.
Cladistically 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 chara ...
, Sinraptoridae had been latest defined in 2005 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 ...
as the most inclusive
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group that contains '' Sinraptor dongi'' and all species closer to ''Sinraptor'' than to either ''
Allosaurus fragilis ''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" alluding to ...
'', ''
Carcharodontosaurus saharicus ''Carcharodontosaurus'' (; ) is a genus of large Carcharodontosauridae, carcharodontosaurid Theropoda, theropod dinosaur that existed during the Cenomanian age of the Late Cretaceous in Northern Africa. The genus ''Carcharodontosaurus'' is na ...
'', or the
house sparrow The house sparrow (''Passer domesticus'') is a bird of the sparrow family Passeridae, found in most parts of the world. It is a small bird that has a typical length of and a mass of . Females and young birds are coloured pale brown and grey, a ...
(''Passer domesticus''). Furthermore, the 2012 study named a new subfamily Metriacanthosaurinae to include all metriacanthosaurids more closely related to ''
Metriacanthosaurus ''Metriacanthosaurus'' (meaning "moderately-spined lizard") is a genus of metriacanthosaurid dinosaur from the upper Oxford Clay of England, dating to the Late Jurassic period, about 160 million years ago (lower Oxfordian). History of discovery ...
'' than to ''
Yangchuanosaurus ''Yangchuanosaurus'' is an extinct genus of metriacanthosaurid theropod dinosaur that lived in China from the Middle to Late Jurassic periods (Bathonian to Oxfordian stages), and was similar in size and appearance to its North American and Eur ...
''. A much larger
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis found ''
Xuanhanosaurus ''Xuanhanosaurus'' (meaning "Xuanhan lizard") is a genus of theropod dinosaur that lived during the Middle Jurassic of China, around 167.7 to 161.2 million years ago. Discovery The type species ''Xuanhanosaurus qilixiaensis'' was named by Don ...
'', previously considered a basal
megalosauroid 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 sha ...
, to be the basalmost metriacanthosaurid.Supporting Information
/ref> Both ''
Poekilopleuron ''Poekilopleuron'' (meaning "varied ribs") is a genus of tetanuran dinosaur, which lived during the middle Bathonian of the Jurassic, about 168 to 166 million years ago. The genus has been used under many different spelling variants, although on ...
'' and ''
Lourinhanosaurus ''Lourinhanosaurus'' (meaning "Lourinhã lizard") was a genus of carnivorous theropod dinosaur that lived during the Late Jurassic Period (Kimmeridgian/Tithonian) in Portugal. It is one of many large predators discovered at the Lourinhã Formati ...
'' were recovered outside the family, and many taxa within the Metriacanthosauridae were in
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
. However, the positions of ''Xuanhanosaurus'' and ''Poekilopleuron'' were very unstable, and their exclusion from the analysis gave a more resolved and stable cladogram. 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 d ...
presented here follows that study.


Palaeobiogeography

Metriacanthosauridae fossils have only been found in modern
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, parts of the prehistoric landmass
Laurasia 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 Pan ...
. From those found in Asia, most are from
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, with a few discoveries in
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
and other Asian countries. The group is thought to have originated in Asia and spread westward to Europe. ''
Yangchuanosaurus ''Yangchuanosaurus'' is an extinct genus of metriacanthosaurid theropod dinosaur that lived in China from the Middle to Late Jurassic periods (Bathonian to Oxfordian stages), and was similar in size and appearance to its North American and Eur ...
'' has only been found in Yangchuan,
Sichuan Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
, China. ''
Metriacanthosaurus ''Metriacanthosaurus'' (meaning "moderately-spined lizard") is a genus of metriacanthosaurid dinosaur from the upper Oxford Clay of England, dating to the Late Jurassic period, about 160 million years ago (lower Oxfordian). History of discovery ...
'' has been found in the
Oxford Clay The Oxford Clay (or Oxford Clay Formation) is a Jurassic marine sedimentary rock formation underlying much of southeast England, from as far west as Dorset and as far north as Yorkshire. The Oxford Clay Formation dates to the Jurassic, specifical ...
in Southern England. Similar to ''
Yangchuanosaurus ''Yangchuanosaurus'' is an extinct genus of metriacanthosaurid theropod dinosaur that lived in China from the Middle to Late Jurassic periods (Bathonian to Oxfordian stages), and was similar in size and appearance to its North American and Eur ...
'', ''
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. De ...
'' has been found in Sichuan, but also in
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
. In 2008, Xu and Clarke described a large tooth found in the Junggard Basin in Xinjian, China, belonging to a large sinraptorid.Xu, X., and J. M. Clark. "The Presence of a Gigantic Theropod in the Jurassic Shishugou Formation, Junggar Basin, Western China." Vertebrata PalAsiatica, 46: 158–160. www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200811/W020090813369286986655.pdf

The Late Jurassic
Qigu Formation The Qigu Formation is a Late Jurassic ( Oxfordian) geologic formation in the Southern Junggar Basin in China. Indeterminate Dinosaur remains are among the fossils that have been recovered from the formation, including theropod teeth and a fibula. ...
in
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
has also yielded evidence of metriacanthosaurid habitation in the form of sauropod bones with bitemarks most likely belonging to a metriacanthosaurid. ''
Siamotyrannus ''Siamotyrannus'' (meaning "Siamese tyrant") is a genus of theropod dinosaur from the early Cretaceous of Thailand. Discovery and naming In 1993, Somchai Traimwichanon found a partial skeleton of a large theropod at the Phu Wiang 9 site in Khon K ...
'' has been found exclusively in Thailand. ''
Shidaisaurus ''Shidaisaurus'' is a genus of metriacanthosaurid dinosaur. Its fossil was found in early Middle Jurassic-age rocks of the Chuanjie Formation in Yunnan, China. It is known from a partial skeleton, holotype DML-LCA 9701-IV, found at the bottom of ...
'' has been found in
Yunnan, China Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the C ...
. Fragmentary remains of allosauroids from the late Middle-early Late Jurassic
Marnes de Dives The Marnes de Dives is a geological formation in Normandy, France. It dates back to the upper part of the Callovian stage of the Middle Jurassic.Weishampel, David B; et al. (2004). "Dinosaur distribution (Middle Jurassic, Europe)." In: Weishampel, ...
in northern France bear close similarities to metriocanthosaurids, and may belong to members of the group.


References


External links

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