Sinovenator (update)
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''Sinovenator'' (meaning "Chinese hunter") 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
troodontid Troodontidae is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinos ...
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 ...
from China. It is from the early
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
Period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
.


Discovery and naming

Two specimens of a
troodontid Troodontidae is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinos ...
were described in
2002 The effects of the September 11 attacks of the previous year had a significant impact on the affairs of 2002. The war on terror was a major political focus. Without settled international law, several nations engaged in anti-terror operation ...
. They are both housed in the
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of People's Republic of China, China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian For ...
, 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 vertebrate paleontologist. He is currently the chairman of paleontology and a research associate at the American Museum of Natural History. He is best known as the discoverer of the first ...
, and colleagues authored the study describing them, finding the specimens to represent a new
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 ...
, 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 languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word ''Sinae'', for
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 ...
, 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 Jehol, an irregular romanization of Chinese (熱河兒), Mongolian, or Manchu, may refer to: * Jehol Province Rehe, previously romanized as Jehol, was a former Chinese special administrative region and province centered on the city of Rehe, n ...
. 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 formal author, t ...
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 wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...
or
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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 (biology), isotype ...
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 from the Late Jurassic to Late Cretaceous. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinos ...
, 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, Dromaeosauridae, Troodontidae, ...
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 In biology (specifically the anatomy of arthropods), the term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod leg, arthropod's foreleg that make it function for the grasping of prey whi ...
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 "han ...
, and some of the highest non- avian
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 and predicted brain mass for an animal of a given size, based on nonlinear regre ...
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 subspecies of the red junglefowl (''Gallus gallus''), originally native to Southeast Asia. It was first domesticated around 8,000 years ago and is now one of the most common and w ...
, 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. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
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 Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
, 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 ...
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 Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
, the large skull opening in front of the eye socket, is straight and vertically oriented. The
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
shows a vertical ridge touching the
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
. The
surangular The surangular or suprangular 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 bone, angular, splenial and articular. It is o ...
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 () 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 oblongata, passes thro ...
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 (anatomy), atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape ...
. 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 (; : is ...
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 (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
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 (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
, conforming to the subarctometatarsal condition.


Phylogeny

In 2002 ''Sinovenator'' was placed in the
Troodontidae Troodontidae is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinos ...
. ''Sinovenator'' is a basal or "primitive"
troodontid Troodontidae is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinos ...
, and shares features with the most basal
dromaeosaur Dromaeosauridae () is a family of feathered coelurosaurian theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Gr ...
ids and
Avialae Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds, and their closest relatives. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to Deinonychosauria, deinonychosaurs, though ...
. 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 dromaeosauridae, dromaeosaurids, troodontidae, troodontids, Anchiornithidae, anchiornithids, and possibly the scansor ...
. A typical basal trait is the orientation of the
pubic bone In vertebrates, the pubis or pubic bone () forms the lower and anterior part of each side of the hip bone. The pubis is the most forward-facing (ventral and anterior) of the three bones that make up the hip bone. The left and right pubic bones ar ...
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 of Laurasia (now Asia, Europe and No ...
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 pa ...
and is not a proof for a Bullatosauria, 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 132.6 ± 2 Ma and 125.77 (million years ago). The Hauterivian is preceded by the Valangi ...
. 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group with other troodontids from the
Yixian Formation The Yixian Formation (; formerly Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
, named
Sinovenatorinae Troodontidae is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinos ...
. A wide, flattened fourth
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
was found to be a
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
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 r ...


References

{{Portal bar, Dinosaurs, China Troodontidae Dinosaur genera Barremian dinosaurs Yixian Formation Taxa named by Xu Xing Taxa named by Mark Norell Fossil taxa described in 2002 Dinosaurs of China