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Velociraptorinae is a subfamily of the
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 ...
group
Dromaeosauridae 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 ...
. The earliest velociraptorines are probably ''
Nuthetes ''Nuthetes'' is the name given to a genus of theropod dinosaur, likely a dromaeosaurid, known only from fossil teeth and jaw fragments found in rocks of the middle Berriasian ( Early Cretaceous) age in the Cherty Freshwater Member of the Lulwor ...
'' from the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
, and possibly ''
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 ...
'' from North America. However, several indeterminate velociraptorines have also been discovered, dating to the
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
stage, in the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
Period. These fossils were discovered in the Langenberg quarry, Oker near
Goslar Goslar (; Eastphalian: ''Goslär'') is a historic town in Lower Saxony, Germany. It is the administrative centre of the district of Goslar and located on the northwestern slopes of the Harz mountain range. The Old Town of Goslar and the Mines ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
.van der Lubbe, T., Richter, U. and Knotschke, N. (2009).
Velociraptorine dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany
" ''Acta Palaeontologica Polonica'', 54(3): 401-408.


Description

While most velociraptorines were generally small animals, at least one species may have achieved gigantic sizes comparable to those found among the dromaeosaurines. So far, this unnamed giant velociraptorine is known only from isolated teeth found on the
Isle of Wight The Isle of Wight ( ) is a county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the largest and second-most populous island of England. Referred to as 'The Island' by residents, the Isle of ...
,
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
. The teeth belong to an animal the size of dromaeosaurines of the genus ''
Utahraptor ''Utahraptor'' (meaning "Utah's thief") is a genus of large dromaeosaurid dinosaur that lived in North America during the Early Cretaceous period. It was a heavy-built, ground-dwelling, bipedal carnivore. It contains a single species, ''Utahrap ...
'', but they appear to belong to a velociraptorine, judging by the shape of the teeth and the anatomy of their serrations.Naish, D. Hutt, and Martill, D.M. (2001). "Saurischian dinosaurs: theropods." in Martill, D.M. and Naish, D. (eds). ''Dinosaurs of the Isle of Wight''. The Palaeontological Association, Field Guides to Fossils. 10, 242–309. In 2007
paleontologists Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
studied front limb bones of ''
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 ...
'' and discovered small bumps on the surface, known as quill knobs. The same feature is present in some bird bones, and represents the attachment point for strong secondary wing feathers. This finding provided the first direct evidence that velociraptorines, like all other maniraptorans, had feathers.


Distinguishing anatomical features

According to Currie (1995), Velociraptorinae can be distinguished based on the following characteristics: dromaeosaurids with maxillary and dentary teeth possessing denticles on the anterior carinae that are significantly smaller than the posterior denticles, and which have a second premaxillary tooth that is significantly larger than the third and fourth premaxillary teeth; dromaeosaurids with nasals that appear depressed, when observed in lateral view. According to Turner ''et al.'' (2012), Velociraptorinae can be distinguished based on the following unambiguous characteristics: the posterior opening of the basisphenoid recess is divided into two small, circular
foramina In anatomy and osteology, a foramen (;Entry "foramen"
in
by a thin bar of bone; the dorsal tympanic recess is present as a deep, posterolaterally directed concavity;
pleurocoels Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is h ...
are present in all of the dorsal vertebrae.


Classification

When erected by Barsbold in 1983, Velociraptorinae was conceived as a group containing ''Velociraptor'' and supposed closely related species. It was not until 1998 that this group was defined as a clade by Paul Sereno. Sereno defined the group as all dromaeosaurids more closely related to ''Velociraptor'' than to ''Dromaeosaurus''. While several studies have since recovered a group of dromaeosaurids closely related to ''Velociraptor'', they vary widely regarding which species are actually velociraptorines and which are either more basal or closer to ''Dromaeosaurus''. Novas and Pol (2005) found a distinct velociraptorine clade close to the traditional view, which included ''
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 ...
'', ''
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 material that was later named ''
Tsaagan ''Tsaagan'' (meaning "white") is a genus of dromaeosaurid dinosaur from the Djadokhta Formation of the Late Cretaceous of Mongolia. Discovery and naming The holotype of ''Tsaagan'' was discovered in 1996 and first identified as a specimen of '' ...
''. A cladistic analysis conducted by Turner ''et al.'' (2012) also supported a traditional, monophyletic of Velociraptorinae. However, some studies found a very different group of dromaeosaurids in velociraptorinae, such as Longrich and Currie (2009), which found ''
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 ...
'' to be a non-velociraptorine, non-dromaeosaurine eudromaeosaur, and ''Saurornitholestes'' to be a member of a more basal group they named Saurornitholestinae. A larger analysis in 2013 found some traditional velociraptorines, such as ''Tsaagan'', to be more basal than ''
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 ...
'', while others to be more closely related to ''Dromaeosaurus'', making them dromaeosaurines. This study found ''Balaur'', previously found to be a velociraptorine by most analyses, to be an
avialan 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 ...
instead. The
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 d ...
below follows a 2009 analysis by paleontologists Nicholas Longrich and
Philip J. Currie Philip John Currie (born March 13, 1949) is a Canadian palaeontologist and museum curator who helped found the Royal Tyrrell Museum of Palaeontology in Drumheller, Alberta and is now a professor at the University of Alberta in Edmonton. In the ...
, using a dataset of 114 characters scored for 23 taxa. The
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 d ...
below follows a 2012 analysis by Turner, Makovicky and Norell, using a dataset of 474 characters scored for 111 taxa. The
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 d ...
below follows the phylogenetic analysis performed by Jasinski ''et al.'' 2020 during the description of ''
Dineobellator ''Dineobellator'' (meaning Diné warrior, pronounced ) is a genus of dromaeosaurid theropod dinosaur that lived in North America during the Late Cretaceous period 68 million years ago. The remains have been found in the Maastrichtian stage of the ...
''.


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 ...


References

{{Taxonbar, from=Q1066137 Eudromaeosaurs