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''Yangchuanosaurus'' 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 ...
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
metriacanthosaurid Metriacanthosauridae is an extinct family (biology), family of Allosauroidea, allosauroid theropod dinosaurs that lived from the Middle Jurassic to the Early Cretaceous. When broken down into its Greek roots, it means "moderately-spined lizards". ...
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 ...
that lived in
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 ...
from the Middle to
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
periods (
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Strat ...
to Oxfordian stages), and was similar in size and appearance to its
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
n and
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 ...
an relative, ''
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" alluding to ...
''. ''Yangchuanosaurus'' hails from the
Upper Shaximiao Formation The Shaximiao Formation () is a Middle to Late Jurassic aged geological formation in Sichuan, China, most notable for the wealth of dinosaurs fossils that have been excavated from its strata. The Shaximiao Formation is exposed in and around the ...
and was the largest
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 ...
in a landscape that included the
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 ...
s ''
Mamenchisaurus ''Mamenchisaurus'' (or spelling pronunciation ) is a genus of sauropod dinosaur known for their remarkably long necks which made up nearly half the total body length. Numerous species have been assigned to the genus; however, many of these might ...
'' and ''
Omeisaurus ''Omeisaurus'' (meaning "Omei lizard") is a genus of sauropod dinosaur from the Middle Jurassic Period (Bathonian-Callovian stage) of what is now China. Its name comes from Mount Emei, where it was discovered in the lower Shaximiao Formation of ...
'' and the
stegosaurs Stegosauria is a group of Herbivore, herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous Period (geology), periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what i ...
''
Chialingosaurus ''Chialingosaurus'' (meaning "Chialing Lizard") is a genus of herbivorous stegosaurian dinosaur similar to ''Kentrosaurus'' from the Upper Shaximiao Formation, Late Jurassic beds in Sichuan Province in China. Its age makes it one of the oldest s ...
'', ''
Tuojiangosaurus ''Tuojiangosaurus'' (meaning "Tuo River lizard") is a genus of herbivorous stegosaurian dinosaur from the Late Jurassic Period, recovered from the Upper Shaximiao Formation of what is now Sichuan Province in China. Description ''Tuojiangosaurus ...
'' and ''
Chungkingosaurus ''Chungkingosaurus'', meaning "Chongqing Lizard", is a genus of herbivorous dinosaur from the Late Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of th ...
''. This theropod was named after the area in which was discovered,
Yongchuan Yongchuan () is a district of Chongqing, China, located by the north side of upper reach of Yangtze River, with a history of 1200 years. Yongchuan borders Sichuan province to the southwest and is away from Yuzhong District of central Chongqing a ...
, in China.


Discovery and species

Dong ''et al.'' (1978) named ''Yangchuanosaurus shangyouensis'' on the basis of CV 00215, a complete
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, the ...
and
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
which was collected from the
Shangshaximiao Formation The Shaximiao Formation () is a Middle to Late Jurassic aged geological formation in Sichuan, China, most notable for the wealth of dinosaurs fossils that have been excavated from its strata. The Shaximiao Formation is exposed in and around the ...
, near Yongchuan,
Yongchuan District Yongchuan () is a district of Chongqing, China, located by the north side of upper reach of Yangtze River, with a history of 1200 years. Yongchuan borders Sichuan province to the southwest and is away from Yuzhong District of central Chongqing ...
,
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 ...
. It dates to the Oxfordian stage of the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
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 ...
, about 161.2 to 157.3 
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. It was discovered in June 1977 by a construction worker during the construction of the Shangyou Reservoir Dam. A second species from the same locality, ''Y. magnus'', was named by Dong ''et al.'' (1983) on the basis of CV 00216, another complete skull and skeleton. A detailed revision of
tetanuran 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 ...
phylogeny by Carrano, Benson & Sampson (2012) revealed that both species are conspecific. Dong ''et al.'' (1978) and Dong ''et al.'' (1983) differentiated these species primarily on the basis of size. In addition, Dong ''et al.'' (1983) noted that the maxilla of ''Y. magnus'' has an additional fenestra within the antorbital fossa, whereas ''Y. shangyouensis'' possessed only a fossa in this location. However, it is considered to be an intraspecific, possibly ontogenetic, variation. Furthermore, the apparent difference in cervical vertebral morphology can be explained by comparing different positions within the column. Hence, the holotypes of the two species of ''Yangchuanosaurus'' are effectively identical, and their codings are identical in Carrano ''et al.'' (2012) matrix.
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 ...
(1988) regarded this genus as a synonym of ''
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 ...
'', but this has not been supported. Carrano ''et al.'' (2012) assigned a third specimen to ''Y. shangyouensis''. CV 00214 is represented by a partial
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l skeleton lacking the skull. It was collected in the Wujiaba Quarry, near Zigong city,
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 ...
, from the lower part of the Shangshaximiao Formation. CV 00214 was initially listed by Dong ''et al.'' (1978) in a faunal list as a new species of ''
Szechuanosaurus ''Szechuanosaurus'' (" Szechuan lizard") is an extinct Chure, D.J. (2000) ''A new species of'' Allosaurus ''from the Morrison Formation of Dinosaur National Monument (UT–CO) and a revision of the theropod family Allosauridae''. Unpublished Ph.D ...
'', ''Szechuanosaurus'' "yandonensis". There is no description or illustration of it, making ''S.'' "yandonensis" a ''
nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
''. Later, Dong ''et al.'' (1983) described it, and assigned it to ''
Szechuanosaurus ''Szechuanosaurus'' (" Szechuan lizard") is an extinct Chure, D.J. (2000) ''A new species of'' Allosaurus ''from the Morrison Formation of Dinosaur National Monument (UT–CO) and a revision of the theropod family Allosauridae''. Unpublished Ph.D ...
campi'', a
dubious Doubt is a mental state in which the mind remains suspended between two or more contradictory propositions, unable to be certain of any of them. Doubt on an emotional level is indecision between belief and disbelief. It may involve uncertainty, ...
species which is known only from four teeth. Carrano ''et al.'' (2012) noted that CV 00214 can't be assigned to ''S. campi'' because the holotype materials of ''S. campi'' (IVPP V.235, V.236, V.238, V.239; teeth) are non-diagnostic and no teeth are preserved in CV 00214. A recent restudy of CV 00214 by Daniel Chure (2001) concluded that it represented a new taxon, informally named "Szechuanoraptor dongi", into which ''Szechuanosaurus zigongensis'' should also be subsumed. However, the most recent revision (by Carrano ''et al.'' (2012)) suggested that CV 00214 and "S." ''zigongensis'' cannot be cospecific as there are no
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
shared between them, and the latter derives from the underlying
Xiashaximiao Formation The Shaximiao Formation () is a Middle to Late Jurassic aged geological formation in Sichuan, China, most notable for the wealth of dinosaurs fossils that have been excavated from its strata. The Shaximiao Formation is exposed in and around th ...
. A phylogenetic analysis found CV 00214 to be most closely related to ''Y. shangyouensis'', and thus the former is assignable to it. Furthermore, ''Szechuanosaurus zigongensis'' was found to be closely related to ''Y. shangyouensis'' and therefore was designated as the second species of ''Yangchuanosaurus''. ''Yangchuanosaurus zigongensis'' is known from four specimens including ZDM 9011 (
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 ...
), a partial postcranial skeleton; ZDM 9012, a left
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 ...
; ZDM 9013, two teeth and ZDM 9014, a right hind limb. It was first described by Gao (1993), and all specimens were collected 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 ...
Xiashaximiao Formation The Shaximiao Formation () is a Middle to Late Jurassic aged geological formation in Sichuan, China, most notable for the wealth of dinosaurs fossils that have been excavated from its strata. The Shaximiao Formation is exposed in and around th ...
in the Dashanpu Dinosaur Quarry of Zigong, Sichuan. A third, informal species, ''Yangchuanosaurus'' "longqiaoensis", was briefly mentioned in a faunal listing of the
Penglaizhen Formation The Penglaizhen Formation (), is a geological formation in Sichuan, China. It was formerly thought to be Late Jurassic in age. However, the underlying Suining Formation has been dated to the Mid Cretaceous, so the formation must be this age or l ...
(initially believed to date to the Late Jurassic but is probably Early Cretaceous (
Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ago) ...
Valanginian In the geologic timescale, the Valanginian is an age or stage of the Early or Lower Cretaceous. It spans between 139.8 ± 3.0 Ma and 132.9 ± 2.0 Ma (million years ago). The Valanginian Stage succeeds the Berriasian Stage of the Lower Cretaceou ...
) in age) by Li, Zhang and Cai (1999). However, since it was published solely in a faunal list and not described in detail, it is a ''nomen nudum'' and is questionably referable to ''Yangchuanosaurus''.


Description

The
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
of ''Y. shangyouensis'' had a
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, the ...
long, and its total body length was estimated at . Another specimen, assigned to the new species ''Y. magnus'', was even larger, with a skull length of . It may have been up to long, and weighed as much as . Gregory S. Paul suggested that these are the same species and gave a length of 11 meters (36 feet) and a weight of 3 metric tons (3.3 short tons). There was a bony ridge on its nose and multiple hornlets and ridges, similar to ''
Ceratosaurus ''Ceratosaurus'' (from Ancient Greek, Greek κέρας/κέρατος, ' meaning "horn" and wikt:σαῦρος, σαῦρος ' meaning "lizard") was a carnivorous Theropoda, theropod dinosaur in the Late Jurassic Period (geology), period (Kim ...
''. ''Yangchuanosaurus'' was a large, powerful meat-eater. It walked on two large, muscular legs, had short arms, a strong, short neck, a big head with powerful jaws, and large, serrated teeth. It had a long, massive tail that was about half of its length. Its arms were short. The first digit of its foot was a small
dewclaw A dewclaw is a digit – vestigial in some animals – on the foot of many mammals, birds, and reptiles (including some extinct orders, like certain theropods). It commonly grows higher on the leg than the rest of the foot, such that in digit ...
. The three outer toes were used to bear weight and each was equipped with a large claw.


Classification

A
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 by Carrano ''et al.'' (2012) found ''Yangchuanosaurus'' to be the
basalmost In phylogenetics, basal is the direction of the ''base'' (or root) of a rooted phylogenetic tree or cladogram. The term may be more strictly applied only to nodes adjacent to the root, or more loosely applied to nodes regarded as being close to th ...
known
metriacanthosaurid Metriacanthosauridae is an extinct family (biology), family of Allosauroidea, allosauroid theropod dinosaurs that lived from the Middle Jurassic to the Early Cretaceous. When broken down into its Greek roots, it means "moderately-spined lizards". ...
and the only non-metriacanthosaurine metriacanthosaurid. 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 below follows their study. 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 below follows Zanno & Makovicky (2013).


References

* * *''Fantastic Facts About Dinosaurs'' () {{Portalbar, Dinosaurs, Paleontology Late Jurassic dinosaurs of Asia Metriacanthosaurids Taxa named by Dong Zhiming Fossil taxa described in 1978 Paleontology in Sichuan