Velociraptorinae
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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. 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 Lulworth ...
'' 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 ...
, and possibly '' Deinonychus'' from North America. However, several indeterminate velociraptorines have also been discovered, dating to the Kimmeridgian 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 Mi ...
,
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.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 Dromaeosaurinae is a subfamily of the theropod group Dromaeosauridae. The earliest dromaeosaurine is ''Utahraptor'', dating back to the Early Cretaceous period in North America, however, some isolated teeth seems to represent an indeterminate spe ...
. 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 Is ...
,
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 ...
. 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 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 th ...
'' 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 by a thin bar of bone; the dorsal tympanic recess is present as a deep, posterolaterally directed concavity; pleurocoels 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 th ...
'', '' Deinonychus'', and material that was later named '' Tsaagan''. 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'' 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 th ...
'', 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 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 ...
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 ...
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 ...
below follows the phylogenetic analysis performed by Jasinski ''et al.'' 2020 during the description of '' Dineobellator''.


See also

* Timeline of dromaeosaurid research


References

{{Taxonbar, from=Q1066137 Eudromaeosaurs