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''Tianyuraptor'' is a
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 short-armed
dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
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 ...
('running lizard'; a type of small dinosaur considered to be closely related to birds) that lived during 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 ...
, about 122 million years ago. Its remains have been found in western
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
,
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 ...
. It was similar to other dromaeosaurids found in Liaoning, with the exception of being somewhat more primitive. 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 ...
, formally named in 2009, shows features not seen in previously known Northern Hemisphere (
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 ...
n) dromaeosaurids, but present in Southern Hemisphere (
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) species and early
birds 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 lightweigh ...
. Because of this, the scientists who first studied ''Tianyuraptor'' described it as a "
transitional species A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
", bridging the gap between northern and southern types of dromaeosaurid. ''Tianyuraptor'' also differs from previously known dromaeosaurids in that it possesses a relatively small
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
("wishbone"), and unusually short forelimbs.


Etymology

The
generic Generic or generics may refer to: In business * Generic term, a common name used for a range or class of similar things not protected by trademark * Generic brand, a brand for a product that does not have an associated brand or trademark, other ...
name of ''Tianyuraptor'' combines ''Tianyu'', referring to the Shandong Tianyu Museum of Nature where the
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 ...
specimen is stored, with ''raptor'', the Latin word for 'robber', referring to the action of grabbing prey, often used in naming dromaeosaurids. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
, ''ostromi'', is in honor of
John Ostrom John Harold Ostrom (February 18, 1928 – July 16, 2005) was an American paleontologist who revolutionized modern understanding of dinosaurs in the 1960s. As first proposed by Thomas Henry Huxley in the 1860s, Ostrom showed that dinosaurs were ...
, who contributed greatly to the study of dromaeosaurid fossils, including ''
Deinonychus ''Deinonychus'' ( ; ) is a genus of dromaeosaurid theropod dinosaur with one described species, ''Deinonychus antirrhopus''. This species, which could grow up to long, lived during the early Cretaceous Period, about 115–108 million ye ...
'' and
feathered dinosaurs A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been s ...
.


Description

''Tianyuraptor'' is a medium-sized dromaeosaurid that has several
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
features that separate it from other dromaeosaurids. These include the length of the middle caudal (tail) vertebrae being more than twice that of the dorsal (back) vertebrae, a small and extremely slender
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
, and an unusually long hindlimb that is roughly three times as long as the entire series of dorsal vertebrae. As in other dromaeosaurid fossils discovered in Liaoning, the tail is relatively long at , nearly 4.8 times as long as the femur. 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 ...
is STM1–3, a nearly complete and fully articulated skeleton that is only missing the extreme distal end of the tail. A total of 25 fully articulated
caudal vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
are preserved and at the end three at most are estimated to be missing. 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 preserved ...
was discovered in the Dawangzhangzi Bed of the
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its ex ...
( Jehol Group), located in Lingyuan in western
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
, China. The Yixian Formation is an
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 ...
rock unit, dated to between approximately 129.7 and 122.1 million years old, in the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
and
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
faunal stages In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convent ...
. The Dawangzhangzi Bed specifically has been dated to about 122 million years ago. STM1–3 is believed to be a sub-adult, with features including the incomplete fusion of skeletal parts during
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
. The holotype of ''Tianyuraptor'' preserves no soft tissues, unlike many other
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 ...
specimens from the Jehol Group.


Limbs

The forelimbs are comparatively short, being only 53% of the hindlimbs' length. This differs greatly from the known skeletal elements of other dromaeosaurids, most of which have relatively long forelimbs that are more than 70% of the hindlimbs' length. While ''Tianyuraptor'' is larger in size than all other known microraptorines, it also has relatively elongated lower hindlimbs, like other microraptorines. In this regard, it is different from most other dromaeosaurids, which have relatively short lower legs. For example, ''Tianyuraptor'' has a
tibiotarsus The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia. A similar structure also occurred in the Mesozoic Heterodontosauridae. These sm ...
/femur length ratio of greater than 1.30, while ''
Velociraptor mongoliensis ''Velociraptor'' (; ) is a genus of small dromaeosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago. Two species are currently recognized, although others have been assigned in the pa ...
'', a creature of similar size, exhibits a ratio of less than 1.10. Aside from elongate hind limbs, ''Tianyuraptor'' is different from other members of Microraptorinae in regards to the relative lengths of the forelimb elements. The forelimbs of Tianyuraptor are proportionally much shorter than those of larger dromaeosaurids. For example, a similar-sized ''
Velociraptor ''Velociraptor'' (; ) is a genus of small dromaeosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago. Two species are currently recognized, although others have been assigned in the pa ...
'' specimen shows an arm/leg length ratio of approximately 0.75., while ''Tianyuraptor'' has an arm/leg ratio of 0.53.


Paleobiology

The stark difference in arm length compared to other dromaeosaurids implies that the function of the arms in ''Tianyuraptor'' was different from that of other dromaeosaurs. Members of the subfamily Microraptorinae, such as ''
Microraptor ''Microraptor'' (Greek, μικρός, ''mīkros'': "small"; Latin, ''raptor'': "one who seizes") is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China. They ...
'', have been suggested to have been aerodynamic and may have glided. Microraptorines are usually noted for their long and robust forelimbs and large, asymmetrical flight feathers. However, the shortened forelimbs, small furcula, and the transversely wide
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
in ''Tianyuraptor'' suggest that it was not suited for aerodynamic gliding or flight.


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 performed by Zheng ''et al.'' showed ''Tianyuraptor'' to be a basal member of a group containing
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 ...
n dromaeosaurids. ''Tianyuraptor'' seems to possess several features that are unknown in other Laurasian dromaeosaurids, but which are seen in basal avialans and
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 dromaeosaurids, including ''
Austroraptor ''Austroraptor'' ( ) is a genus of large dromaeosaurid dinosaur that lived during the Late Cretaceous Period in what is now Argentina. ''Austroraptor'' was a large-sized, moderately-built, ground-dwelling, bipedal carnivore, estimated at in len ...
'', ''
Buitreraptor ''Buitreraptor'' (meaning "La Buitrera seizer") is a genus of dromaeosaurid dinosaurs that lived during the Late Cretaceous of Argentina at the Candeleros Formation. ''Buitreraptor'' was described in 2005 and the type species is ''Buitreraptor go ...
'', '' Neuquenraptor'', ''
Rahonavis ''Rahonavis'' is a genus of bird-like theropods from the Late Cretaceous (Maastrichtian, about 70 mya) of what is now northwestern Madagascar. It is known from a partial skeleton ( UA 8656) found by Catherine Forster and colleagues in Maevaran ...
'', and ''
Unenlagia ''Unenlagia'' (meaning "half-bird" in Latinized Mapudungun) is a genus of dromaeosaurid theropod dinosaur that lived in South America during the Late Cretaceous period. The genus ''Unenlagia'' has been assigned two species: ''U. comahuensis'', ...
''. Zheng and colleagues also noted that ''Tianyuraptor'' shares some features with the
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 ...
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Microraptorinae Microraptoria (Greek, μίκρος, ''mīkros'': "small"; Latin, ''raptor'': "one who seizes") is a clade of basal dromaeosaurid theropod dinosaurs. The first microraptorians appeared 125 million years ago in China. Many are known for long feather ...
, though they went on to say that this mixture of features suggests a basal placement for ''Tianyuraptor'' within Microraptorinae, as evidenced by their phylogenetic analysis which indicated
maximum parsimony In phylogenetics, maximum parsimony is an optimality criterion under which the phylogenetic tree that minimizes the total number of character-state changes (or miminizes the cost of differentially weighted character-state changes) is preferred. ...
in six of the 30 results recovered by the analysis. The authors then go on to suggest that since ''Tianyuraptor'' is considered a short-armed microraptorine, more
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
long-armed microraptorines might have independently evolved flight capability. However, it is also equally possible, as argued by Zheng ''et al.'', that ''Tianyuraptor'' may in fact be a basal member of a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
containing all other
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 ...
n dromaeosaurids with the exception of Microraptorinae. This is indicated by the other 24 out of 30 most parsimonious trees recovered from the analysis. The discovery of ''Tianyuraptor'' sheds new light on the early evolution of dromaeosaurs and further exemplifies the great diversity this group enjoyed at an early stage.


See also

*
Timeline of dromaeosaurid research This timeline of dromaeosaurid research is a chronological listing of events in the history of paleontology focused on the dromaeosaurids, a group of sickle-clawed, bird-like theropod dinosaurs including animals like ''Velociraptor''. Since the ...
*
Paraves Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the avial ...


References

{{Taxonbar, from=Q133038 Early Cretaceous dinosaurs of Asia Yixian fauna Microraptorians Fossil taxa described in 2009 Taxa named by Xu Xing