Shaochilong
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''Shaochilong'' (meaning "shark toothed dragon") 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
carcharodontosaurid Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
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 mid-
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 th ...
(Late
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 the end of 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 by t ...
stage)
Ulansuhai Formation The Ulansuhai Formation () is a geological formation in Inner Mongolia, north China. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, David B; et al. (2004). "Dinosaur distribution (Early Cretaceous, As ...
of
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 ...
(about 92 million years ago). 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 specimen ...
, ''S. maortuensis'', was originally named ''
Chilantaisaurus ''Chilantaisaurus'' (" lizard") is a genus of large theropod dinosaur, possibly a neovenatorid or a primitive coelurosaur, from the Late Cretaceous Ulansuhai Formation of China (Turonian age, about 92 million years ago). The type species, ''C. ...
maortuensis'', but was re-described and reclassified in 2009. It was one of the last known carcharodontosaurids to walk the earth. Alongside ''
Ulughbegsaurus ''Ulughbegsaurus'' (meaning "Ulugh Beg's lizard") is a dubious genus of theropod dinosaur from the Late Cretaceous aged Bissekty Formation, Uzbekistan. The type species is ''Ulughbegsaurus uzbekistanensis''. Discovery and naming ''Ulughbegsaur ...
'' from
Uzbekistan Uzbekistan (, ; uz, Ozbekiston, italic=yes / , ; russian: Узбекистан), officially the Republic of Uzbekistan ( uz, Ozbekiston Respublikasi, italic=yes / ; russian: Республика Узбекистан), is a doubly landlocked cou ...
and ''
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 be ...
'' from
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 ...
, they were the only members of the family to live until the end of 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 by t ...
epoch.


History

The material referred to ''Shaochilong'', IVPP V.2885.1-7, consisted of skull fragments (a braincase, partial skull roof, quadrates, and a right maxilla),
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-coordinat ...
and six caudal vertebrae. A fragmentary left maxilla was also referred to the species, although it has apparently gone missing as of 2009. Although these are believed to belong to a single individual, a
lectotype In biology, a type is a particular 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 anchor or centralizes the ...
was established in 2010 to accommodate for the possibility that the specimens came from multiple individuals. The lectotype consists of the braincase (IVPP V.2885.1) and partial skull roof (IVPP V.2885.2). All of these specimens were discovered in Inner Mongolia and described by Hu in 1964 as a species of ''
Chilantaisaurus ''Chilantaisaurus'' (" lizard") is a genus of large theropod dinosaur, possibly a neovenatorid or a primitive coelurosaur, from the Late Cretaceous Ulansuhai Formation of China (Turonian age, about 92 million years ago). The type species, ''C. ...
''.Hu, S.-Y. (1964). "Carnosaurian remains from Alashan, Inner Mongolia." ''Vertebrata PalAsiatica'', 8: 42–63. n Chinese, with English summary/ref> The genus was informally named "Alashansaurus" by Chure in 2000. Chure (2002) and Rauhut (2001) suggested that it did not belong to that genus, and was probably a primitive
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, tyran ...
.Glut, D. (2003). ''Dinosaurs - The Encyclopedia - Supplement 3''. McFarland Press: Jefferson, NC. However, a re-description by Brusatte and colleagues in 2009 found that it was a carcharodontosaurid, the first recognized from Asia. This re-description also provided a new genus name. A more comprehensive description and discussion was published the following year.


Description

The individual which IVPP V2885.1 belonged to was probably adult or nearly adult individual, due to the fusion of many elements of the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
. ''Shaochilong'' length – based on the length of the maxillary tooth row – is estimated at . Estimated length of the
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 with ...
is 61.5 cm which suggests the whole animal weighed approximately . This made ''Shaochilong'' an uncharacteristically small carcharodontosaurid, in contrast with other members of the family, which were among the largest carnivorous animals on earth. Nevertheless, as an unambiguous example of a fairly large
carnosaur 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 ''Gig ...
in middle Cretaceous Asia, it provides information on the ecosystem of this setting and is evidence that carcharodontosaurids persisted through the middle cretaceous and that large
tyrannosaurids Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family (biology), family of coelurosaurian Theropoda, theropod dinosaurs that comprises two subfamilies containing up to thirteen genus, genera, including the eponymous ''Tyrannos ...
(such as ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, 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. ''Tyrannosa ...
'') did not dominate
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 ...
until the very end of the Cretaceous. According to the proportions of its maxilla, ''Shaochilong'' was a relatively short-faced carcharodontosaur. In combination with its comparatively small size, it could have had a unique ecological role in comparison to other members of the family. ''Shaochilong'' can be distinguished from other carcharodontosaurids due to having the following
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 ...
: * A reduced and nearly absent maxillary
antorbital fossa 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 ...
. * No paradental groove on the medial surface 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 ...
. * Deep vertical grooves located dorsally on the maxillary
interdental plate The interdental plate refers to the bone-filled mesial-distal region between the teeth. The word "''interdental''" is a combination of "''inter''" + "''dental''" (meaning "''between the teeth''") which originated in approximately 1870. In paleobi ...
s. * A pneumatic recess which penetrates to the posterior end of the nasal. * A deep
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
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 e ...
. * A large pneumatic foramen in the anterodorsal corner of the prootic's dorsal tympanic recess.


Classification

Phylogenetic analysis performed by Brusatte and coworkers indicate that ''Shaochilong'' is deeply nested within the carchorodontosaurids, the most derived group among the
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 ...
s. ''Shaochilong'' appears to be more closely related to the
Gondwana 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 stages ...
n carcharodontosaurids (''
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 ''Carcharo ...
'', ''
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 ...
'', ''
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 be ...
'', ''
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 the ...
'') than the Laurasian ones (such as ''
Neovenator ''Neovenator'' (nee-o-ven-a-tor meaning "new hunter") is a genus of carcharodontosaurian theropod dinosaur. It is known from several skeletons found in the Early Cretaceous (Barremian~130-125 million years ago) Wessex Formation on the south coa ...
'' and ''
Acrocanthosaurus ''Acrocanthosaurus'' ( ; ) is a genus of carcharodontosaurid dinosaur that existed in what is now North America during the Aptian and early Albian stages of the Early Cretaceous, from 113 to 110 million years ago. Like most dinosaur genera, ' ...
''). ''Shaochilong'' is the youngest known Laurasian allosauroid suggesting that basal tetanurans not
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, were still the dominant group of large-bodied theropods in Laurasian during the Mid-Cretaceous and that the rise of tyrannosaurids as the dominant group of large terrestrial predators was sudden and confined to the very end of the Cretaceous. The following cladogram after Novas et al., 2013, shows the position of ''Shaochilong'' within Carcharodontosauridae.


References


External links


Shark toothed theropods in Asia – introducing ''Shaochilong''
Discoverers' blog {{Taxonbar, from=Q134476 Carcharodontosaurids Late Cretaceous dinosaurs of Asia Cretaceous animals of Asia Fossils of China Fossil taxa described in 2009 Taxa named by Stephen L. Brusatte Taxa named by Xu Xing Turonian life