Sinovenator Changii
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''Sinovenator'' (meaning "Chinese hunter") 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
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
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 China. It is from the early
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 ...
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 ...
.


Discovery and naming

Two specimens of a
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
were described in
2002 File:2002 Events Collage.png, From left, clockwise: The 2002 Winter Olympics are held in Salt Lake City; Queen Elizabeth The Queen Mother and her daughter Princess Margaret, Countess of Snowdon die; East Timor gains East Timor independence, indepe ...
. They are both housed in the
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name suggest ...
, under the specimen numbers IVPP V 12615 and IVPP V 12583. Xu Xing,
Mark Norell Mark Allen Norell (born July 26, 1957) is an American paleontologist, acknowledged as one of the most important living vertebrate paleontologists. He is currently the chairman of paleontology and a research associate at the American Museum of Na ...
, and colleagues authored the study describing them, finding the specimens to represent a new
taxon In biology, a taxon (back-formation from ''taxonomy''; plural 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 nam ...
, for which was chosen the
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms * Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition ...
''Sinovenator changii''. The generic name was derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word ''Sinae'', for
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 ...
, and ''Venator'', or "hunter".
Meemann Chang Meemann Chang (; born 17 April 1936) also known as Zhang Miman, is a Chinese paleontologist at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP). She completed her undergraduate studies at Moscow University and completed her ...
is honoured by the species name for her contributions to the study of the Jehol Fauna. As Chang is a female researcher, the epithet should have been "changae"; however, such mistakes cannot be emended according to the rules of the
ICZN The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the I ...
and therefore forms such as "Sinovenator changae" or "Sinovenator changiae" that sometimes appear in the literature, are incorrect. 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 ...
or
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 ...
of ''Sinovenator changii'' is IVPP 12615, a partial skull and disarticulated skeleton. An additional specimen was by the original publication scientifically described and assigned as the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
of the species: an incomplete but articulated postcranial skeleton, numbered IVPP 12583. A third specimen was referred: IVPP V14322, a fragmentary skeleton. All three are in the collection of the IVPP in Beijing, China. However, ''Sinovenator'' fossils appear to be common in the Lujiatun Beds. In a 2006 survey of the
Jehol Biota The Jehol Biota includes all the living organisms – the ecosystem – of northeastern China between 133 and 120 million years ago. This is the Lower Cretaceous ecosystem which left fossils in the Yixian Formation and Jiufotang Formation. These d ...
, Xu and Norell reported that hundreds of undescribed specimens are known. The fossils have been preserved three-dimensionally, not strongly compressed on a slab. In 2014, the wrist of one specimen was described, IVPP V14009, an adult. In 2018, PMOL-AD00102, consisting of a partial skull and mandibles, and six cervical vertebrae, was referred to the taxon.


Description

''Sinovenator'' is a
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
, a group of small, bird-like,
gracile Gracility is slenderness, the condition of being gracile, which means slender. It derives from the Latin adjective ''gracilis'' (masculine or feminine), or ''gracile'' ( neuter), which in either form means slender, and when transferred for examp ...
maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
ns. All troodontids have many unique features of the skull, such as closely spaced
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 ...
in the lower jaw, and large numbers of teeth. Troodontids have sickle-claws and
raptorial The term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod's foreleg that make it function for the grasping of prey while it is consumed, where the gripping surfaces are formed from the o ...
hands A hand is a prehensile, multi-fingered appendage located at the end of the forearm or forelimb of primates such as humans, chimpanzees, monkeys, and lemurs. A few other vertebrates such as the koala (which has two opposable thumbs on each " ...
, and some of the highest non-
avian Avian may refer to: *Birds or Aves, winged animals *Avian (given name) (russian: Авиа́н, link=no), a male forename Aviation *Avro Avian, a series of light aircraft made by Avro in the 1920s and 1930s *Avian Limited, a hang glider manufacture ...
encephalization quotient Encephalization quotient (EQ), encephalization level (EL), or just encephalization is a relative brain size measure that is defined as the ratio between observed to predicted brain mass for an animal of a given size, based on nonlinear regressi ...
s, meaning they were behaviourally advanced and had keen senses. The holotype individual of ''Sinovenator'' was about the size of a
chicken The chicken (''Gallus gallus domesticus'') is a domesticated junglefowl species, with attributes of wild species such as the grey and the Ceylon junglefowl that are originally from Southeastern Asia. Rooster or cock is a term for an adult m ...
, less than a metre long. In 2010,
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 ...
estimated the weight of a one metre long individual at 2.5 kilogrammes.Paul, G.S., 2010, ''The Princeton Field Guide to Dinosaurs'', Princeton University Press p. 140 However, the holotype and paratype are subadults; specimens are known with about twice their length. This makes it one of the larger species of troodontids in the ecosystem 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 ...
, the formation where it was found. In the original description some distinguishing traits or
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 ...
were established. The front edge of the
antorbital fenestra 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 ...
, the large skull opening in front of the eye socket, is straight and vertically oriented. The
frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...
shows a vertical ridge touching the
lacrimal bone The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
. The
surangular The suprangular or surangular is a jaw bone found in most land vertebrates, except mammals. Usually in the back of the jaw, on the upper edge, it is connected to all other jaw bones: dentary, angular, splenial and articular The articular bone i ...
in the lower jaw has a T-shaped cross-section. The shinbone has a vertical ridge on the upper outer side, the ''crista cnemialis'', touching a lower vertical ridge, the ''crista fibularis''. There is a number of other notable traits. The upper branch of the praemaxilla excludes the maxilla from the nostril. The premaxillary teeth lack denticles; those on the maxillary teeth are small. The maxillary tooth row is positioned at some distance from the jaw rim. A ''fenestra promaxillaris'' is present. The branch of the jugal towards the postorbital is long, slender and inclined to the rear. The
foramen magnum The foramen magnum ( la, great hole) is a large, oval-shaped opening in the occipital bone of the skull. It is one of the several oval or circular openings (foramina) in the base of the skull. The spinal cord, an extension of the medulla oblon ...
is much larger than the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
. The braincase lacks a ''recessus suboticus'' and a ''crista otosphenoidalis''. The rear of the pterygoid has a secondary, rod-like, process. The front dentary teeth are closely packed. The rear of the surangular is extraordinarily deep. Five sacrals are present, those of the middle being clearly larger. The ilium is relatively small. The
ischium The ischium () form ...
is bird-like: small with a low obturator process and processes on the upper and lower rear edge. The shinbone is wide at the top and has a roughly rectangular lower joint surface. The fourth
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 med ...
is not strongly developed but the second metatarsal is small. The third metatarsal is pinched at the top but still visible at the upper front of the
metatarsus 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 med ...
, conforming to the subarctometatarsal condition.


Phylogeny

In 2002 ''Sinovenator'' was placed in the
Troodontidae Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil disco ...
. ''Sinovenator'' is a basal or "primitive"
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
, and shares features with the most basal
dromaeosaur 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 ...
ids and
Avialae Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used ...
. It thus was one of the first fossil specimens conclusively demonstrating the close relation of these three groups in the overarching
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 ...
. A typical basal trait is the orientation of the
pubic bone In vertebrates, the pubic region ( la, pubis) is the most forward-facing (ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior ra ...
to the rear, similar to the situation in the Dromaeosauridae and basal birds. This proves that the orientation to the front with later troodontids, is a reversal and not a trait directly inherited from older theropods that also possess a forward pointing pubic bone. Another basal character is the lack of a ''bulla'', a swollen region on the underside of the braincase. This indicates that the similar ''bulla'' with 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 (geology), Period of Laurasia (now Asia ...
presents a case of
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and paral ...
and is not a proof for a
Bullatosauria Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings that contains ornithomimosaurs and maniraptorans. This group was named by Thomas Holtz, who defined it as "the most recent common ancestor of '' Ornithomimus'' and bir ...
, an earlier presumed close relationship between troodontids and ornithomimosaurs. Originally, the basal position of ''Sinovenator'' seemed to be corroborated by a very high age, the fossils being assumed to date from the
Hauterivian The Hauterivian is, in the geologic timescale, an age in the Early Cretaceous Epoch or a stage in the Lower Cretaceous Series. It spans the time between 132.9 ± 2 Ma and 129.4 ± 1.5 Ma (million years ago). The Hauterivian is preceded by the Va ...
. Later research however, indicated that the layers were younger. In 2017, a study by Shen ''et al.'', describing the genus ''
Daliansaurus ''Daliansaurus'' (meaning "Dalian reptile") is a genus of small troodontid theropod dinosaur, measuring approximately long, from the Early Cretaceous of China. It contains a single species, ''D. liaoningensis'', named in 2017 by Shen and colleag ...
'' found ''Sinovenator'' to form a
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 ...
group with other troodontids from 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 ...
, named
Sinovenatorinae Sinovenatorinae is a short-lived subfamily of the extinct bird-like troodontid dinosaurs known exclusively from the Early Cretaceous of China. It includes the genus ''Sinovenator ''Sinovenator'' (meaning "Chinese hunter") is a genus of troodo ...
. A wide, flattened fourth
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 med ...
was found to be a
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
uniting this group. The study recovered the following cladogram: A 2018 study by Yin ''et al.'' instead found the Chinese troodontids to form an intermediate grade:


See also

*
Timeline of troodontid research This timeline of troodontid research is a chronological listing of events in the History of paleontology, history of paleontology focused on the troodontids, a group of bird-like theropod dinosaurs including animals like ''Troodon''. Troodontid re ...


References

{{Portal bar, Dinosaurs, Cretaceous, China Early Cretaceous dinosaurs of Asia Troodontids Fossil taxa described in 2002 Taxa named by Xu Xing Yixian fauna