Qianzhousaurus
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''Qianzhousaurus'' (meaning "Qianzhou lizard") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
that lived in
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
during the
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
age of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
period. There is currently only one species named, the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
''Qianzhousaurus sinensis'', which is a member of the tribe Alioramini and most closely related to '' Alioramus'', the only other known alioramin.


History of discovery

The holotype specimen, GM F10004, was unearthed in southern
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
,
Ganzhou Ganzhou (), alternately romanized as Kanchow, is a prefecture-level city in the south of Jiangxi province, China, bordering Fujian to the east, Guangdong to the south, and Hunan to the west. Its administrative seat is at Zhanggong District. His ...
, at the
Nanxiong Formation The Nanxiong Formation (also known as Yuanpu Formation) is a Late Cretaceous geologic Formation (geology), formation in Guangdong Province. Dinosaur remains are among the fossils that have been recovered from the formation. Description It cons ...
in the summer of 2010 during the construction of an industrial park and it was first described by paleontologists Junchang Lü, Laiping Yi, Stephen L. Brusatte, Ling Yang, Hua Li and Liu Chen in the journal ''
Nature Communications ''Nature Communications'' is a peer-reviewed, open access, scientific journal published by Nature Portfolio since 2010. It is a multidisciplinary journal that covers the natural sciences, including physics, chemistry, earth sciences, medic ...
'' in 2014. The genus is known from a partial sub-adult individual consisting of a nearly complete skull with the lower jaws missing all
teeth A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
(lost during fossilization), 9
cervical vertebrae In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
, 3 dorsal vertebrae, 18 caudal vertebrae, both
scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...
s, partial ilia, and the left leg compromising the
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
,
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
,
fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. ...
,
astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
with
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel; : calcanei or calcanea) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is t ...
, and metatarsals III and IV. The generic name, ''Qianzhousaurus'', is in reference to Qianzhou (the older name of Ganzhou) where the remains were discovered, and the
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, ''sinensis'', is derived from the
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
word Σῖναι (sin, sino, sinai) in reference to China. The
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 preserve ...
remains were discovered by workmen at a construction site near the city of Ganzhou, who then took them to a local museum. Lead author Lü Junchang from the Institute of Geology,
Chinese Academy of Geological Sciences The Chinese Academy of Geological Sciences (CAGS; ) is an institution that engages in geoscience research in the People's Republic of China. The academy was established in 1956 and reorganized in 1999. Administratively it is under the PRC Ministr ...
stated that "the new discovery is very important. Along with '' Alioramus'' from Mongolia, it shows that the long-snouted tyrannosaurids were widely distributed in Asia. Although we are only starting to learn about them, the long-snouted tyrannosaurs were apparently one of the main groups of predatory dinosaurs in Asia." The existence of long-snouted tyrannosaurs was previously suspected due to other inconclusive fossil finds which could be explained as the juveniles of short-snouted species, but co-author Stephen L. Brusatte from the
University of Edinburgh The University of Edinburgh (, ; abbreviated as ''Edin.'' in Post-nominal letters, post-nominals) is a Public university, public research university based in Edinburgh, Scotland. Founded by the City of Edinburgh Council, town council under th ...
reveals that the find "tells us pretty unequivocally that these long-snouted tyrannosaurs were a real thing. They were a different breed, living right at the end of the age of dinosaurs."


Description

''Qianzhousaurus'' was a medium-sized
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
estimated at in length, in hip height and in body mass. Higher estimates suggest that it could have reached a maximum length between . The
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
can be differentiated from other tyrannosaurids in having a highly narrowed
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
, a pneumatic opening on the upper extension of the
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
, and the lack of a vertical ridge-like structure on the lateral surface of the ilium. Unlike more "traditional" tyrannosaurids, which had prominent deep-set jaws and thick teeth, ''Qianzhousaurus'' had a particularly elongated snout, with (when restored) narrow teeth. The holotype specimen is notably larger and more mature than the holotypes of both species of '' Alioramus''. However, since some sutures between the cervical and dorsal vertebrae are partially fused, the holotype seems to have come from an immature animal, likely a sub-adult. ''Qianzhousaurus'' was a long-legged animal with a long
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
and long
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
.


Classification

The discovery of ''Qianzhousaurus'' led to a new branch of the tyrannosaur family being named, consisting of the long-snouted ''Q. sinensis'' and the two known species of ''Alioramus''. This clade, named the tribe Alioramini, had an uncertain placement relative to other members of the tyrannosaur branch in the initial analysis that discovered it. The primary
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis found Alioramini to be closer to ''Tyrannosaurus'' than to ''Albertosaurus'', and therefore a member of the group Tyrannosaurinae. However, a second analysis in the same paper found it to be located outside of the clade including Albertosaurinae and Tyrannosaurinae, and therefore the sister group of
Tyrannosauridae Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to fifteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
. Below is the first analysis found by the authors: Thomas D. Carr with colleagues in 2017 regarded ''Qianzhousaurus'' as a junior synonym of ''Alioramus'' based on the reasoning that they were recovered as
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
in their phylogenetic analysis. Despite their argument, they indicated that this synonymy does not necessarily constitute a taxonomic reevaluation of the Alioramini. However, Foster and his colleagues conducted a detailed redescription of the holotype skull of ''Qianzhousaurus'' in 2022, finding several traits unique to this genus, hence supporting the separation of ''Alioramus'' and ''Qianzhousaurus''.


Paleobiology


Ontogeny

A 2022 study of the three known species of the two known alioramin genera, ''Qianzhousaurus'' and '' Alioramus'' (''A. altai'' and ''A. remotus''), respectively, suggests that the variation seen between the various species is consistent with the growth trends seen in other tyrannosaurid genera, though specimens that could constitute a full growth series from infant to adult for each species have not been recovered for any of these theropods. Of these, ''Qianzhousaurus'' represents the largest and most mature animal found within Alioramini thus far and represents the adult level of maturity. One part of the growth series across all specimens in this study was discovered to remain unique to alioramin tyrannosaurs, this being the rugose process of the jugal starts out small and conical, but becomes massive and indistinct as the animals grow. This same study also suggests Alioramins did not undergo a secondary metamorphosis from slender juveniles to robust adults like other tyrannosaurs, but maintained a unique physiology better suited to pursuit of fast prey.


Feeding

In 2022, Foster and his colleagues indicated that ''Qianzhousaurus'' and other alioramins, due to their slim and gracile build, may have been hunters of small, particularly fast and nimble prey, which would have allowed these tyrannosaurids to avoid competition with larger, robust relatives that specialized in killing larger animals. The long and delicate snouts of alioramins like ''Alioramus'' and ''Qianzhousaurus'' may have prevented them from killing the same prey that juvenile and adult tyrannosaurids like '' Tarbosaurus'' hunted, though these larger tyrannosaurs themselves may have hunted alioramins as prey on occasion. Alioramins may also have had a different feeding strategy than other tyrannosaurids, as their jaws seem to have been weaker than those of the larger genera and even juveniles of larger species have proportionately higher bite forces than alioramins of equivalent size. Furthermore, Alioramini seemingly remained confined to Asia, suggesting that some factor prevented them from colonizing the better-sampled fossil deposits from North America. What this may be remains a mystery at the present moment. Studies of the skulls of various tyrannosauroids in 2024 suggest that ''Qianzhousaurus'' and other alioramin tyrannosaurs experienced lower stresses when biting and feeding. Additional evidence in the same study suggests that ''Qianzhousaurus'' did not utilize the "puncture-and-pull" feeding method utilized by other, larger genera of tyrannosaurids.


Paleoenvironment

''Qianzhousaurus'' is known from the
Nanxiong Formation The Nanxiong Formation (also known as Yuanpu Formation) is a Late Cretaceous geologic Formation (geology), formation in Guangdong Province. Dinosaur remains are among the fossils that have been recovered from the formation. Description It cons ...
, which has been dated to the latest
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
age of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
period, about 66.7 ± 0.3
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
based on
argon–argon dating Argon–argon (or 40Ar/39Ar) dating is a radiometric dating method invented to supersede Potassium-argon dating, potassiumargon (K/Ar) dating in accuracy. The older method required splitting samples into two for separate potassium and argon measur ...
. The main
lithology The lithology of a rock unit is a description of its physical characteristics visible at outcrop, in hand or core samples, or with low magnification microscopy. Physical characteristics include colour, texture, grain size, and composition. Lit ...
of this formation is composed by purplish
mudstones Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from ''shale'' by its lack of fissility.Blatt, H., and R.J. Tracy, 1996, ''Petrology.'' New York, New York ...
and
siltstones Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt Silt is granular material of a size between sand and clay and composed mostly of broken grains of quartz. Silt may occur as a soil (often mixed ...
, deposited in a
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
environment under a relatively warm,
humid subtropical climate A humid subtropical climate is a subtropical -temperate climate type, characterized by long and hot summers, and cool to mild winters. These climates normally lie on the southeast side of all continents (except Antarctica), generally between ...
. Oviraptorid
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
s are particularly common across the formation with numerous well-preserved egg clutches or nests, as well as nesting individuals. The Nanxiong Formation has been noted for its abundant oviraptorosaur genera. It is most likely that, given the poor
stratigraphic Stratigraphy is a branch of geology concerned with the study of rock layers (strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
analysis of the formation, the extremely large oviraptorosaur diversity of this formation was temporally separated. The presence of the smaller and more robustly built tyrannosaurine '' Asiatyrannus'' also suggests that ''Qianzhousaurus'' was filling the role of apex predator in the formation's ecosystem, with ''Asiatyrannus'' occupying a different niche and hunting different prey than its larger relative. Other vertebrates in the Nanxiong Formation include numerous oviraptorosaurs, such as '' Banji'', '' Ganzhousaurus'', '' Corythoraptor'', ''
Nankangia ''Nankangia'' is an extinct genus of caenagnathoid oviraptorosaurian dinosaur known from the Upper Cretaceous Nanxiong Formation of Nankang, Jiangxi, Nankang County, Ganzhou City of Jiangxi Province, southeastern China. It contains a single speci ...
'', '' Huanansaurus'', '' Shixinggia'', or '' Tongtianlong''; the hadrosaurid '' Microhadrosaurus'' (may be ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
''); the sauropod '' Gannansaurus''; the therizinosaurid '' Nanshiungosaurus''; the crocodilian '' Jiangxisuchus''; the squamates '' Chianghsia'' and '' Tianyusaurus''; and the turtles '' Jiangxichelys'' and '' Nanhsiungchelys''.


See also

* Timeline of tyrannosaur research


References

{{Taxonbar, from=Q16777007 Tyrannosauridae Dinosaur genera Maastrichtian dinosaurs Taxa named by Lü Junchang Taxa named by Stephen L. Brusatte Fossil taxa described in 2014 Dinosaurs of China