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''Hexing'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of basal ornithomimosaur
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 ...
known from the Early Cretaceous of northeastern China. It contains a single species, ''Hexing qingyi''.


Discovery and naming

In the early twenty-first century, a local farmer at Xiaobeigou in Liaoning discovered the skeleton of a small
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 ...
. He prepared the fossil himself, trying to enhance its value by restoring damaged bones and adding fake parts. Eventually the specimen was obtained by the Geological Museum of Jilin University and more expertly prepared, during which process the added parts were again removed. In 2012, 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 specime ...
''Hexing qingyi'' was named and described by Jin Liyong, Chen Jun and
Pascal Godefroit Pascal Godefroit is a Belgian paleontologist. He discovered dinosaurs like '' Olorotitan'' in 2003. Godefroit is the director of earth and life sciences This list of life sciences comprises the branches of science that involve the scientific st ...
. The generic name means "like a crane" in
Chinese Chinese can refer to: * Something related to China * Chinese people, people of Chinese nationality, citizenship, and/or ethnicity **''Zhonghua minzu'', the supra-ethnic concept of the Chinese nation ** List of ethnic groups in China, people of ...
. The specific name means "with slender wings". 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 sever ...
, JLUM-JZ07b1, was found in fluvial deposits of the lower
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 exq ...
, which have a highest possible age of 139 million years and a lowest of 128 million years and thus date from some time in the early
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 Cretace ...
to early
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 precede ...
stage. It consists of a partial skeleton, containing the skull, the lower jaws, a series of five cervical vertebrae, the shoulder girdle and the majority of both forelimbs and hindlimbs. The remains have not been well preserved. The specimen represents a subadult or adult individual.


Description

The holotype specimen consists of the remains of a small individual. Because most of the vertebral column is absent, its body length cannot be directly determined, but a comparison can be made with the previously smallest known ornithomimosaur, the 1.6 metres long ''
Shenzhousaurus ''Shenzhousaurus'' (meaning "Shenzhou lizard") is a genus of basal ornithomimosaur from the Lower Cretaceous of China. The holotype (NGMC 97-4-002, National Geological Museum of China) was collected from near the bottom (fluvial beds) of the ...
'', which has a thighbone length of 191 millimetres, while the femur length of ''Hexing'' is 135 millimetres. Such a small body size was among ornithomimosaurs up till now only known from juveniles but the holotype is not a young animal as is shown by the complete fusion of the skull bones, the neck ribs, the
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 ...
and the ankle bones. The describers determined some
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 ...
of ''Hexing'', its unique derived traits. The snout tip reaches downwards in front of the lower jaws, so deeply that the roof of the mouth at this point is at a level with the bottom edge of the lower jaw. The ''fossa antorbitalis'', a depression on the side 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. T ...
, covers almost the entire outer surface of that bone. The
parietal bone The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is nam ...
s are joint at their midline in a crest. The large bone extensions at the back of the skull, the ''processus paroccipitales'', are hanging down below the level of the foramen magnum. In the lower jaw, the
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
has an opening in its side. The formula for the phalanges of the hand is 1-2-3-0-0 or 2-3-3-0-0 — or 0-1-2-3-0/0-2-2-3-0 if the three fingers of the hand are interpreted as the second, third and fourth. The upper phalanges of the second and third (or third and fourth) finger are elongated with more than 75% of the length of the corresponding metacarpal. The lower leg is relatively long with the
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 s ...
having 137% of the length of the thighbone. The skull of ''Hexing'' is relatively large with a length of 136 millimetres. It is elongated and triangular in side view. The snout is appending with a slight kink above the middle of the ''fossa antorbitalis'', which at this point is reinforced by a distinctive vertical bone strut, a ''pila interfenestralis'' dividing the depression in two halves: at its rear an oval skull opening, the ''fenestra antorbitalis'', pierces the surface and at its front the uniquely large and deep hollowing out of the maxilla side is visible, in which another opening, the ''fenestra maxillaris'', might have been present, though this is uncertain because of damage. The front of the snout consists of a small
praemaxilla 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 mammal has ...
, continuing the line of the nasals downwards and forming a small upper beak in front of the lower jaws. The beak is separated by a low notch from the lower maxilla edge, which is toothless. The front of the lower jaw is low and slightly upward curving. In the left dentary the remains of three or four low conical teeth are visible; of these damage obscures most detail. The dentary has a small opening in the side surface. The higher back of the lower jaw seems to show a much larger opening, but this is an artefact caused by the original inexpert preparation damaging the thin bone surface of an extensive mandibular fossa. A real and much smaller external mandibular fenestra is present in front of this. Neither the lower jaw nor the upper jaw form cutting edges. The cervical vertebrae are elongated with low spines. They are pneumatised and have large triangular diapophyses and
postzygapophyses The articular processes or zygapophyses ( Greek ζυγον = " yoke" (because it links two vertebrae) + απο = "away" + φυσις = " process") of a vertebra are projections of the vertebra that serve the purpose of fitting with an adjacent vert ...
. The shoulder blade is elongated and narrow, without expanded upper end. The humerus is somewhat shorter than the shoulder blade and is slender. Its shaft is not twisted. The bones of the lower arm are likewise elegant and straight. A large part of the over ten centimetres long hand has been preserved; this shows a configuration that is fundamentally different from that of related species. The describers have carefully checked whether this could have been caused by falsifications during the original preparation but could find no sign of any tampering. As a result, the identification of the respective parts is highly problematical. As preserved the hand shows three rows of elements: the first with two bones and the second and third with four bones. If these would represent both phalanges and metacarpals, these series should have three, four and five elements, however: from the first and third row a bone is missing. The authors considered it most likely that in the first finger the upper phalanx was completely reduced, that is: naturally absent. However, as this would imply that the claw attached directly to the metacarpal and this metacarpal would then be exceptionally long, they allowed for the alternative possibility that the visible element was the first phalanx and that the metacarpal was lacking because of an incompleteness of the fossil. In the third finger the number of phalanges seemed almost certainly reduced from four to three because the place of the two normal short upper phalanges is taken by a single long element. The description of this situation is complicated by the fact that the describers follow the hypothesis of Xu Xing that with
Tetanurae 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 ...
— including birds and ''Hexing'' — the first, second and third fingers are actually the second, third and fourth. This would in the standard terminology make the formula of the phalanges 1-3-3-0-0 or alternatively 2-3-3-0-0 and following the hypothesis of Xu 0-1-3-3-0 or 0-2-3-3-0. In general the phalanges are slender and elongated. The hand claws are relatively large and curved, with a flatter underside. The thighbone is strongly curved, with a convex anterior end. 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 ...
is 185 millimetres long and thus very elongated compared with the femur; only some juvenile ornithomimid specimens have relatively longer tibiotarsi. The first
metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the me ...
is half as long as the second, the longest of the foot. The foot claws are flatter than the hand claws.


Phylogeny

The describers assigned ''Hexing'' a basal position in the
Ornithomimosauria Ornithomimosauria ("bird-mimic lizards") are theropod dinosaurs which bore a superficial resemblance to the modern-day ostrich. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia (now Asia, Europe and Nor ...
. Their
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 ''Hexing'' to be more derived than ''
Pelecanimimus ''Pelecanimimus'' (meaning " pelican mimic") is an extinct genus of basal ("primitive") ornithomimosaurian dinosaur from the Early Cretaceous of Spain. It is notable for possessing more teeth than any other member of the Ornithomimosauria ( ...
'', but less derived than a clade consisting of '' Beishanlong'', ''
Harpymimus ''Harpymimus'' is a basal ornithomimosaurian theropod dinosaur from the Early Cretaceous Period of what is now Mongolia. Unlike later, more derived ornithomimosaurs, ''Harpymimus'' still possessed teeth, although they appear to have been restr ...
'', ''
Garudimimus ''Garudimimus'' (meaning "Garuda mimic") is a genus of ornithomimosaur that lived in Asia during the Late Cretaceous. The genus is known from a single specimen found in 1981 by a Soviet-Mongolian paleontological expedition in the Bayan Shireh Fo ...
'' and the
Ornithomimidae Ornithomimidae (meaning "bird-mimics") is a family of theropod dinosaurs which bore a superficial resemblance to modern ostriches. Ornithomimids were fast, omnivorous or herbivorous dinosaurs known mainly from the Late Cretaceous Period of Lauras ...
. ''
Shenzhousaurus ''Shenzhousaurus'' (meaning "Shenzhou lizard") is a genus of basal ornithomimosaur from the Lower Cretaceous of China. The holotype (NGMC 97-4-002, National Geological Museum of China) was collected from near the bottom (fluvial beds) of the ...
'', also from the Yixian Formation, was found in a
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 ...
with ''Hexing'' and this clade, as shown by the following
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 ...
: The small forms ''Hexing'' and ''Senzhousaurus'' are the oldest known ornithomimosaurians, as ''
Pelecanimimus ''Pelecanimimus'' (meaning " pelican mimic") is an extinct genus of basal ("primitive") ornithomimosaurian dinosaur from the Early Cretaceous of Spain. It is notable for possessing more teeth than any other member of the Ornithomimosauria ( ...
'' dates from the late
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 precede ...
. Because this latter species is more basal, the authors considered this a strong indication that a land bridge had formed between Europe and Asia long before the Aptian, the normally assumed date for this event.


See also

* Timeline of ornithomimosaur research


References

{{Portalbar, Dinosaurs, Cretaceous, China Fossil taxa described in 2012 Ornithomimosaurs Early Cretaceous dinosaurs of Asia Yixian fauna