Velociraptor 32529 225409586 2008-07-13T15:09:31Z 74.140.243.143 /* In popular culture */ {{Taxobox | name = ''Velociraptor'' | fossil_range = {{fossil range|83|70}}[[Late Cretaceous]] | image = Velociraptor skull crâne 2.png | image_width = 250px | image_caption = Replica of a ''Velociraptor mongoliensis'' skull. | regnum = [[Animal]]ia | phylum = [[Chordate|Chordata]] | classis = [[Reptile|Sauropsida]] | superordo = [[Dinosaur]]ia | ordo = [[Saurischia]] | subordo = [[Theropoda]] | familia = [[Dromaeosauridae]] | genus = '''''Velociraptor''''' | genus_authority = [[Henry Fairfield Osborn|Osborn]], 1924 | subdivision_ranks = [[Species]] | subdivision = *''V. mongoliensis'' <small>Osborn, 1924 ([[Type (biology)|type]])</small> *''V. osmolskae'' <small> Godefroit ''et al.'', 2008</small> }} '''''Velociraptor''''' ({{IPAEng|vɨˈlɒsɨræptɚ}}; meaning 'swift thief') is a [[genus]] of [[dromaeosaurid]] [[Theropoda|theropod]] [[dinosaur]] that existed approximately 83 to 70&nbsp;[[mega-annum|Ma]] ([[million years ago]]) during the later part of the [[Cretaceous]] [[Period (geology)|Period]]. Only two species are currently recognized, although others have been assigned in the past. The [[type species]] is ''V. mongoliensis''; [[fossil]]s of this species have been discovered in both [[Inner Mongolia|Inner]] and [[Mongolia|Outer Mongolia]] in [[central Asia]]. A second species, ''V. osmolskae'', was named in 2008 for skull material from Inner Mongolia. Smaller than other dromaeosaurids like ''[[Deinonychus]]'' and ''[[Achillobator]]'', the [[turkey (bird)|turkey]]-sized ''Velociraptor'' nevertheless shared many of the same [[anatomy|anatomical]] features. It was a [[biped]]al, [[feather]]ed [[carnivore]] with a long, stiffened tail and an enlarged sickle-shaped [[claw]] on each hindfoot, which is thought to have been used to kill its [[predation|prey]]. ''Velociraptor'' can be distinguished from other dromaeosaurids by its long and low [[skull]], with an upturned snout. ''Velociraptor'' (commonly shortened to 'raptor') is one of the dinosaur genera most familiar to the general public due to its prominent role in the ''[[Jurassic Park (film)|Jurassic Park]]'' motion picture series, although in the films it was shown much larger than it was in reality and without feathers as well as having other anatomical inaccuracies. It is also well-known to [[paleontology|paleontologist]]s, with over a dozen recovered fossil skeletons — the most of any dromaeosaurid. One particularly famous specimen preserves a ''Velociraptor'' locked in combat with a ''[[Protoceratops]]''. ==Description== [[Image:Vraptor-scale.svg|thumb|left|''Velociraptor'' compared in size to a human.]] ''Velociraptor'' was a mid-sized dromaeosaurid, with adults measuring up to {{convert|2.07|m|ft|abbr=on|lk=on}} long, {{convert|0.5|m|ft|abbr=on}} high at the hip, and weighing up to {{convert|15|kg|lb|abbr=on|lk=on}}.<ref name=paul1988>{{cite_book |last=Paul |first=Gregory S. |authorlink=Gregory S. Paul |year=1988 |title=Predatory Dinosaurs of the World |location=New York |publisher=Simon and Schuster |pages=464pp |isbn=978-0671619466}}</ref> The skull, which grew up to {{convert|25|cm|in|abbr=on|lk=on}} long, was uniquely up-curved, concave on the upper surface and convex on the lower. The jaws were lined with 26&ndash;28 widely-spaced teeth on each side, each more strongly serrated on the back edge than the front — possibly an adaptation that improved its ability to catch and hold fast-moving prey.<ref name=osborn1924a>{{cite_journal |last=Osborn |first=Henry F. |authorlink=Henry Fairfield Osborn |year=1924a |title=Three new Theropoda, ''Protoceratops'' zone, central Mongolia |journal=American Museum Novitates |volume=144 |pages=1–12 |url=http://hdl.handle.net/2246/3223}}</ref><ref name=barsboldosmolska1999>{{cite_journal |last=Barsbold |first=Rinchen |authorlink=Rinchen Barsbold |coauthors=& Osmólska, Halszka. |year=1999 |title=The skull of ''Velociraptor'' (Theropoda) from the Late Cretaceous of Mongolia |journal=Acta Palaeontologica Polonica |volume=44 |issue=2 |pages=189–219 |url=http://app.pan.pl/acta44-2.htm#Barsbold |format={{dead link|date=June 2008}} &ndash; <sup>[http://scholar.google.co.uk/scholar?hl=en&lr=&q=author%3ABarsbold+intitle%3AThe+skull+of+%27%27Velociraptor%27%27+%28Theropoda%29+from+the+Late+Cretaceous+of+Mongolia&as_publication=Acta+Palaeontologica+Polonica&as_ylo=1999&as_yhi=1999&btnG=Search Scholar search]</sup>}}</ref> [[Image:Velociraptor mongoliensis jmallon.jpg|thumb|''Velociraptor mongoliensis''.]] ''Velociraptor'', like other dromaeosaurids, had a large [[Manus (zoology)|manus]] ('hand') with three strongly-curved claws, which were similar in construction and flexibility to the wing bones of modern [[bird]]s. The second digit was the longest of the three digits present, while the first was shortest. The structure of the [[carpal]] (wrist) bones prevented [[pronation]] of the wrist and forced the 'hands' to be held with the [[palmar]] surface facing inwards ([[Anatomical terms of location#Left and right .28lateral.29.2C and medial|medially]]), not downwards.<ref name=paul2002>{{cite_book |last=Paul |first=Gregory S. |authorlink=Gregory S. Paul |year=2002 |title=Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds |location=Baltimore |publisher=Johns Hopkins University Press |pages=472pp |isbn=978-0801867637}}</ref> The first digit of the foot, as in other theropods, was a small [[dewclaw]]. However, whereas most theropods had feet with three digits contacting the ground, dromaeosaurids like ''Velociraptor'' walked on only their third and fourth digits. The second digit, for which ''Velociraptor'' is most famous, was highly modified and held retracted off of the ground. It bore a relatively large, sickle-shaped claw, typical of dromaeosaurid and [[troodontid]] dinosaurs. This enlarged claw, which could be over {{convert|6.5|cm|in|abbr=on}} long around its outer edge, was most likely a predatory device used to tear into prey, possibly delivering a fatal blow.<ref name=barsbold1983>{{cite_journal |last=Barsbold |first=Rinchen. |authorlink=Rinchen Barsbold |year=1983 |title=Carnivorous dinosaurs from the Cretaceous of Mongolia |journal=Transactions of the Joint Soviet-Mongolian Paleontological Expedition |volume=19 |pages=5–119}}</ref><ref name=norellmakovicky1999>{{cite_journal |last=Norell |first=Mark A. |coauthors=& Makovicky, Peter J. |year=1999 |title=Important features of the dromaeosaurid skeleton II: information from newly collected specimens of ''Velociraptor mongoliensis'' |journal=American Museum Novitates |volume=3282 |pages=1–45 |url=http://hdl.handle.net/2246/3025}}</ref> Long bony projections (prezygapophyses) on the upper surfaces of the [[vertebra]]e, as well as [[ossification|ossified]] [[tendon]]s underneath, stiffened the tail of ''Velociraptor''. The prezygapophyses began on the tenth tail (caudal) vertebra and extended forward to brace four to ten additional vertebrae, depending on position in the tail. The stiffening forced the entire tail to act as a single rod-like unit, preventing vertical motion between vertebrae. However, at least one specimen preserves a series of intact tail vertebrae curved sideways into an ''S''-shape, suggesting that there was considerably more horizontal flexibility. These adaptations of the tail probably provided balance and stability while turning, especially at high speeds.<ref name=barsbold1983/><ref name=norellmakovicky1999/> In 2007, paleontologists [[Alan Turner (paleontologist)|Alan Turner]], [[Peter Makovicky]], [[Mark Norell]] and colleagues reported the discovery of quill knobs on a well-preserved ''Velociraptor mongoliensis'' forearm from Mongolia, confirming the presence of feathers in this species.<ref name="turneretal2007b">{{cite journal |last=Turner |first=A.H. |coauthors=Makovicky, P.J.; Norell, M.A. |year=2007 |title=Feather quill knobs in the dinosaur ''Velociraptor'' |journal=Science |volume=317 |issue=5845 |pages=1721|doi=10.1126/science.1145076 |pmid=17885130 }}</ref> ==History== [[Image:Velociraptor mongoliensis type skull and jaws.jpg|thumb|A drawing of the [[holotype|type]] skull of ''Velociraptor mongoliensis''. From Osborn, 1924.]] An [[American Museum of Natural History]] expedition to the Outer Mongolian [[Gobi Desert]] in 1922 recovered the first ''Velociraptor'' fossil known to science: a crushed but complete skull, associated with one of the raptorial second toe claws ([[AMNH]] 6515). In 1924, museum president [[Henry Fairfield Osborn]] designated the skull and claw (which he assumed to come from the hand) as the [[type specimen]] of his new genus, ''Velociraptor''. This name is derived from the [[Latin]] words ''velox'' ('swift') and ''raptor'' ('robber' or 'plunderer') and refers to the animal's [[cursorial]] nature and carnivorous diet. Osborn named the type species ''V. mongoliensis'' after its country of origin.<ref name=osborn1924a/> Earlier that year, Osborn had mentioned the animal in a popular press article, under the name "Ovoraptor djadochtari" (not to be confused with the similarly named ''[[Oviraptor]]'').<ref name=osborn1924b>{{cite_journal |last=Osborn |first=Henry F. |authorlink=Henry Fairfield Osborn |year=1924b |title=The discovery of an unknown continent |journal=Natural History |volume=24 |pages=133–149}}</ref> However, because the name "Ovoraptor" was not published in a scientific journal or accompanied by a formal description, it is considered a ''[[nomen nudum]]'' ('naked name'), and the name ''Velociraptor'' retains priority. While North American teams were shut out of [[communist]] Mongolia during the [[Cold War]], expeditions by [[Soviet Union|Soviet]] and [[Poland|Polish]] scientists, in collaboration with Mongolian colleagues, recovered several more specimens of ''Velociraptor''. The most famous is part of the legendary "Fighting Dinosaurs" specimen ([[Mongolian Academy of Sciences|GIN]] 100/25), discovered by a Polish-Mongolian team in 1971. This fossil preserves a single ''Velociraptor'' in the midst of battle against a lone ''Protoceratops''.<ref name=barsbold1983/><ref name=kjbarsbold1972>{{cite_journal |last=Kielan-Jaworowska |first=Zofia |coauthors=& [[Rinchen Barsbold|Barsbold, Rinchen]]. |year=1972 |title=Narrative of the Polish-Mongolian Paleontological Expeditions |journal=Paleontologica Polonica |volume=27 |pages=5–13}}</ref><ref name=barsbold1974>{{cite_journal |last=Barsbold |first=Rinchen. |authorlink=Rinchen Barsbold |year=1974 |title=Saurornithoididae, a new family of theropod dinosaurs from Central Asia and North America |journal=Paleontologica Polonica |volume=30 |pages=5–22}}</ref> This specimen is considered a national treasure of Mongolia, although in 2000 it was loaned to the American Museum of Natural History in [[New York City]] for a temporary exhibition.<ref name=amnh>{{cite_web |url=http://www.amnh.org/exhibitions/fightingdinos/ex-fd.php |title=The Fighting Dinosaurs |author=[[American Museum of Natural History]] |accessdate=2007-06-13}}</ref> Between 1988 and 1990, a joint [[China|Chinese]]-[[Canada|Canadian]] team discovered ''Velociraptor'' remains in northern China.<ref name=jerzykiewiczetal1993>{{cite_journal |last=Jerzykiewicz |first=Tomasz |coauthors=[[Phil Currie|Currie, Philip J.]]; Eberth, David A.; Johnston, P.A.; Koster, E.H.; & Zheng J. |year=1993 |title=Djadokhta correlative strata in Chinese Inner Mongolia: An overview of the stratigraphy, sedimentary geology, and paleontology and comparisons with the type locality in the pre-Altai Gobi. |journal=Canadian Journal of Earth Sciences |volume=30 |pages=2180–2195}} [printed early 1994]</ref> American scientists returned to Mongolia in 1990, and a joint Mongolian-American expedition to the Gobi, led by the American Museum of Natural History and the [[Mongolian Academy of Sciences]], turned up several well-preserved skeletons.<ref name=norellmakovicky1999/><ref name=norellmakovicky1997>{{cite_journal |last=Norell |first=Mark A. |coauthors=& Makovicky, Peter J. |year=1997 |title=Important features of the dromaeosaur skeleton: information from a new specimen |journal=American Museum Novitates |volume=3215 |pages=1–28 |url=http://hdl.handle.net/2246/3557}}</ref> One such specimen, IGM 100/980, was nicknamed "Ichabodcraniosaurus" by Norell's team because the fairly complete specimen was found without its skull (an allusion to the [[Washington Irving]] character [[Ichabod Crane]]).<ref name="novacek1996">Novacek, M. (1996). ''Dinosaurs of the Flaming Cliffs'', Anchor Books.</ref> This specimen may belong to ''Velociraptor mongoliensis'', but Norell and Makovicky concluded that it was not complete enough to say for sure, and it awaits a formal description.<ref name=norellmakovicky1999/> [[Maxilla]]e and a [[lacrimal]] (the main tooth-bearing bones of the upper jaw, and the bone that forms the anterior margin of the eye socket, respectively) recovered in 1999 by the Sino-Belgian Dinosaur Expeditions were found to pertain to ''Velociraptor'', but not to the type species ''V. mongoliensis''. Pascal Godefroit and colleages named these bones ''V. osmolskae'' (for Polish paleontologist [[Halszka Osmólska]]) in 2008.<ref name=PGetal2008>{{cite journal |last=Godefroit |first=Pascal |coauthors=Currie, Philip J.; Li Hong; Shang Chang Yong; and Dong Zhi-ming |year=2008 |title=A new species of ''Velociraptor'' (Dinosauria: Dromaeosauridae) from the Upper Cretaceous of northern China |journal=Journal of Vertebrate Paleontology |volume=28 |issue=2 |pages=432–438}}</ref> ==Provenance== [[Image:Velociraptor 28-12-2007 15-06-24.jpg|thumb|left|Mounted skeleton of ''Velociraptor mongoliensis'', Museum voor Natuurwetenschappen, [[Brussels]].]] All known specimens of ''Velociraptor mongoliensis'' were discovered in the [[Djadochta Formation]] (also spelled Djadokhta), in both the Mongolian province of [[Ömnögovi Province|Ömnögovi]] and Chinese Inner Mongolia. A species of ''Velociraptor'', possibly ''V. mongoliensis'', is also preserved in the slightly younger [[Barun Goyot Formation]] of Mongolia.<ref name=weishampeletal2004>{{cite_book |last=Weishampel |first=David B. |authorlink=David B. Weishampel |coauthors=Barrett, Paul M., [[Rodolfo Coria|Coria, Rodolfo A.]], Le Loueff, Jean, [[Xu Xing]], [[Zhao Xijin]], Sahni, Ashok, Gomani, Emily M.P. & Noto, Christopher N. |year=2004 |chapter=Dinosaur distribution |editor=[[David B. Weishampel|Weishampel, David B.]], [[Peter Dodson|Dodson, Peter]] & Osmólska, Halszka (eds.). |title=The Dinosauria |edition=Second Edition |location=Berkeley |publisher=University of California Press |pages=Pp. 517–606. |isbn=0520242092}}</ref> These [[geologic formation]]s are estimated to date back to the [[Campanian]] [[faunal stage|stage]] (about 83 to 70&nbsp;million years ago<ref name=gradsteinetal2005>{{cite_book |last=Gradstein |first=Felix M. |coauthors=Ogg, James G.; & Smith, Alan G. |year=2005 |title=A Geologic Time Scale 2004 |location=Cambridge |publisher=Cambridge University Press |pages=500pp. |isbn=978-0521781428}}</ref>) of the [[Late Cretaceous]] [[Epoch (geology)|Epoch]].<ref name=jerzykiewiczrussell1991>{{cite_journal |last=Jerzykiewicz |first=Tomasz |coauthors=& [[Dale Russell|Russell, Dale A.]] |year=1991 |title=Late Mesozoic stratigraphy and vertebrates of the Gobi Basin |journal=Cretaceous Research |volume=12 |issue=4 |pages=345–377 |doi=10.1016/0195-6671(91)90015-5}}</ref> ''V. mongoliensis'' has been found at many of the most famous and prolific Djadochta localities. The type specimen was discovered at the [[Flaming Cliffs]] site (also known as Bayn Dzak and Shabarakh Usu),<ref name=osborn1924a/> while the "Fighting Dinosaurs" were found at the Tugrig locality (also known as Tugrugeen Shireh).<ref name=barsbold1974/> More recently, fossils of ''V. mongoliensis'' were recovered from Bayan Mandahu, a prolific site from the Djadochta of Inner Mongolia in China.<ref name=jerzykiewiczetal1993/> The well-known Barun Goyot localities of Khulsan and Khermeen Tsav have also produced remains which may belong to the genus ''Velociraptor''.<ref name=osmolska1997>{{cite_book |last=Osmólska |first=Halszka |year=1997 |chapter=Barun Goyot Formation |editors=[[Phil Currie|Currie, Philip J.]] & Padian, Kevin (eds.). |title=Encyclopedia of Dinosaurs |location=San Diego |publisher=Academic Press |pages=p.41 |isbn=0-12-226810-1}}</ref> All of these sites preserve an arid environment with fields of sand [[dune]]s and only intermittent [[stream]]s, although the younger Barun Goyot environment seems to have been slightly wetter than the older Djadochta.<ref name=jerzykiewiczrussell1991/> Aside from ''Protoceratops'', upon which it preyed, ''Velociraptor'' shared its environment with other [[Basal (phylogenetics)|basal]] [[ceratopsia]]ns like ''[[Udanoceratops]]'' and [[ankylosaurid]]s like ''[[Pinacosaurus]]'', along with several species of [[oviraptorid]], [[troodontid]], other [[dromaeosaurid]]s (such as ''[[Adasaurus]]''), and [[alvarezsaurid]] theropods.<ref name=weishampeletal2004/> ''V. osmolskae'' was found in the [[Bayan Mandahu Formation]], contemporaneous with the Djadochta Formation. As in the Djadochta, ''Pinacosaurus'', ''Protoceratops'', oviraptorid, and troodontid theropods were present.<ref name=PGetal2008/> ==Taxonomy== [[Image:Velociraptor BW.jpg|thumb|left|Illustration of ''Velociraptor mongoliensis''.]] ''Velociraptor'' is a member of the subfamily [[Velociraptorinae]], a [[derived]] sub-group of the larger family Dromaeosauridae. In [[phylogenetic taxonomy]], Velociraptorinae is usually defined as "all dromaeosaurs more closely related to ''Velociraptor'' than to ''[[Dromaeosaurus]]''." Dromaeosaurid classification is highly variable. Originally, the subfamily Velociraptorinae was erected solely to contain ''Velociraptor''.<ref name=barsbold1983/> Other analyses have included other genera, usually ''Deinonychus'' and ''[[Saurornitholestes]]''.<ref name=currie1995>{{cite_journal |last=Currie |first=Philip J. |authorlink=Phil Currie |year=1995 |title=New information on the anatomy and relationships of ''Dromaeosaurus albertensis'' (Dinosauria: Theropoda) |journal=Journal of Vertebrate Paleontology |volume=15 |issue=3 |pages=576–591 |url=http://vertpaleo.org/publications/jvp/15-576-591.cfm}}</ref> A recent [[cladistic]] analysis indicated a [[monophyletic]] Velociraptorinae containing ''Velociraptor'', ''Deinonychus'', ''Tsaagan'', and a closely related (but uncertainly positioned) ''Saurornitholestes''.<ref name=norelletal2006>{{cite_journal |last=Norell |first=Mark A. |coauthors=Clark, James M.; Turner, Alan H.; Makovicky, Peter J., [[Rinchen Barsbold|Barsbold, Rinchen]]; & Rowe, Timothy. |year=2006 |title=A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia) |journal=American Museum Novitates |volume=3545 |pages=1–51 |url=http://hdl.handle.net/2246/5823 |doi=10.1206/0003-0082(2006)3545[1:ANDTFU]2.0.CO;2}}</ref> In the past, other dromaeosaurid species, including ''Deinonychus antirrhopus'' and ''Saurornitholestes langstoni'', have sometimes been classified in the genus ''Velociraptor''. Since ''Velociraptor'' was the first to be named, these species were renamed ''Velociraptor antirrhopus'' and ''V. langstoni''.<ref name=paul1988>{{cite_book |last=Paul |first=Gregory S. |authorlink=Gregory S. Paul |year=1988 |title=Predatory Dinosaurs of the World |location=New York |publisher=Simon and Schuster |pages=464pp |isbn=978-0671619466}}</ref> However, the only currently recognized species of ''Velociraptor'' are ''V. mongoliensis''<ref name=barsboldosmolska1999/><ref name=paul2002/><ref name=norellmakovicky2004>{{cite_book |last=Norell |first=Mark A. |coauthors=& Makovicky, Peter J. |year=2004 |chapter=Dromaeosauridae |editor=[[David B. Weishampel|Weishampel, David B.]], [[Peter Dodson|Dodson, Peter]] & Osmólska, Halszka (eds.). |title=The Dinosauria |edition=Second Edition |location=Berkeley |publisher=University of California Press |pages=Pp. 196–209 |isbn=0520242092}}</ref> and ''V. osmolskae''.<ref name=PGetal2008/> When first described in 1924, ''Velociraptor'' was placed in the family [[Megalosauridae]], as was the case with most carnivorous dinosaurs at the time (Megalosauridae, like ''[[Megalosaurus]]'', functioned as a sort of 'wastebin' taxon, where many unrelated species were grouped together).<ref name=osborn1924a/> As dinosaur discoveries multiplied, ''Velociraptor'' was later recognized as a dromaeosaurid. All dromaeosaurids have also been referred to the family [[Archaeopterygidae]] by at least one author (which would, in effect, make ''Velociraptor'' a flightless bird).<ref name=paul2002/> ==Paleobiology== ===Predatory behavior=== [[Image:Fightingdinosamnh2.jpg|thumb|''Velociraptor'' and ''Protoceratops'' in combat]] The "Fighting Dinosaurs" specimen, found in 1971, preserves a ''Velociraptor'' and ''[[Protoceratops]]'' in combat and provides direct evidence of predatory behavior. When originally reported, it was hypothesized that the two animals drowned.<ref name=barsbold1974/> However, as the animals were preserved in ancient sand dune deposits, it is now thought that the animals were buried in sand, either from a collapsing dune or in a [[Dust storm|sandstorm]]. Burial must have been extremely fast, judging from the lifelike poses in which the animals were preserved. Both forelimbs and one hindlimb of the ''Protoceratops'' are missing, which has been seen as evidence of [[scavenger|scavenging]] by other animals.<ref name=carpenter1998>{{cite_journal |last=Carpenter |first=Ken. |authorlink=Kenneth Carpenter |year=1998 |title=Evidence of predatory behavior by theropod dinosaurs. |journal=Gaia |volume=15 |pages=135–144 |url=http://vertpaleo.org/publications/jvp/15-576-591.cfm}} [not printed until 2000]</ref> [[Image:Fighting dinosaurs.jpg|thumb|left|Close-up of a replica of the "Fighting Dinosaurs".]] The distinctive claw, on the second digit of dromaeosaurids, has traditionally been depicted as a slashing weapon; its assumed use being to cut and [[disembowel]] prey.<ref name=ostrom1969>{{cite_journal |last=Ostrom |first=John H. |authorlink=John Ostrom |year=1969 |title=Osteology of ''Deinonychus antirrhopus'', an unusual theropod from the Lower Cretaceous of Montana |journal=Bulletin of the Peabody Museum of Natural History |volume=30 |pages=1–165}}</ref> In the "Fighting Dinosaurs" specimen, the ''Velociraptor'' lies underneath, with one of its sickle claws apparently embedded in the throat of its prey, while the beak of ''Protoceratops'' is clamped down upon the right forelimb of its attacker. This suggests ''Velociraptor'' may have used its sickle claw to pierce vital organs of the throat, such as the [[jugular vein]], [[carotid artery]], or [[vertebrate trachea|trachea]] (windpipe), rather than slashing the abdomen. The inside edge of the claw was rounded and not unusually sharp, which may have precluded any sort of cutting or slashing action, although only the bony core of the claw is known. The thick abdominal wall of [[skin]] and [[muscle]] of large prey species would have been difficult to slash without a specialized cutting surface.<ref name=carpenter1998/> The slashing [[hypothesis]] was tested during a 2005 [[BBC]] documentary, ''[[The Truth About Killer Dinosaurs]]''. The producers of the program created an artificial ''Velociraptor'' leg with a sickle claw and used a [[pork belly]] to simulate the dinosaur's prey. Though the sickle claw did penetrate the abdominal wall, it was unable to tear it open, indicating that the claw was not used to disembowel prey. However, this experiment has not been published or repeated by other scientists, so its results cannot be confirmed. Remains of ''[[Deinonychus]]'', a closely related dromaeosaurid, have commonly been found in aggregations of several individuals. ''Deinonychus'' has also been found in association with a large herbivore, ''[[Tenontosaurus]]'', which has been seen as evidence of cooperative hunting.<ref name=maxwellostrom1995>{{cite_journal |last=Maxwell |first=W. Desmond |coauthors=& [[John Ostrom|Ostrom, John H.]] |year=1995 |title=Taphonomy and paleobiological implications of Tenontosaurus-Deinonychus associations |journal=Journal of Vertebrate Paleontology |volume=15 |issue=4 |pages=707–712 |url=http://vertpaleo.org/publications/jvp/15-707-712.cfm}}</ref><ref name=brinkmanetal1998>{{cite_journal |last=Brinkman |first=Donald L. |coauthors=Cifelli, Richard L.; & Czaplewski, N.J. |year=1998 |title=First occurrence of ''Deinonychus antirrhopus'' (Dinosauria: Theropoda) in the Antlers Formation (Lower Cretaceous: Aptian-Albian) of Oklahoma |journal=Oklahoma Geological Survey Bulletin |volume=146 |pages=1–27}}</ref> The only solid evidence for social behavior among dromaeosaurids comes from a Chinese trackway of fossil footprints, which shows six individuals of a large species moving as a group, though no evidence of cooperative hunting was found.<ref name="Lietal2007">{{cite journal | first=Rihui | last=Li | coauthors= Lockley, M.G., Makovicky, P.J., Matsukawa, M., Norell, M.A., Harris, J.D. and Liu, M.| title=Behavioral and faunal implications of Early Cretaceous deinonychosaur trackways from China | date=2007 | publisher= | url =http://www.springerlink.com/content/v1u455854212404r/ | work =Naturwissenschaften | pages = | accessdate = | language = }}</ref> Although many isolated fossils of ''Velociraptor'' have been found in Mongolia, none were closely associated with any other individuals.<ref name=norellmakovicky2004/> Therefore, while ''Velociraptor'' is commonly depicted as a [[pack hunter]], as in ''Jurassic Park'', there is only limited fossil evidence to support this theory for dromaeosaurids in general, and none specific to ''Velociraptor'' itself. ===Metabolism=== ''Velociraptor'' was probably [[warm-blooded]] to some degree, as it required a significant amount of energy to hunt. Modern animals that possess feathery or furry coats, like ''Velociraptor'' did, tend to be warm-blooded, since these coverings function as insulation. However, bone growth rates in dromaeosaurids and some early birds suggest a more moderate [[metabolism]], compared with most modern warm-blooded mammals and birds. The [[kiwi]] is similar to dromaeosaurids in anatomy, feather type, bone structure and even the narrow anatomy of the nasal passages (usually a key indicator of metabolism). The kiwi is a highly active, if specialized, flightless bird, with a stable body temperature and a fairly low resting metabolic rate, making it a good model for the metabolism of primitive birds and dromaeosaurids.<ref name=paul2002/> ===Feathers=== [[Image:Velociraptor dinoguy2.jpg|thumb|''Velociraptor mongoliensis''.</center>]] Fossils of dromaeosaurids more primitive than ''Velociraptor'' are known to have had [[feather]]s covering their bodies, and fully-developed, feathered wings.<ref name=xuetal2003>{{cite_journal |last=Xu Xing |authorlink=Xu Xing |coauthors=Zhou Zhonghe, Wang Xiaolin, Kuang Xuewen, Zhang Fucheng & Du Xiangke |year=2003 |title=Four-winged dinosaurs from China |journal=Nature |issue=421 |pages=335–340 |doi=10.1038/nature01342 | volume = 421}}</ref> The fact that the ancestors of ''Velociraptor'' were feathered and possibly capable of flight long suggested to paleontologists that ''Velociraptor'' bore feathers as well, since even flightless birds today retain most of their feathers. In September 2007, researchers found [[feathers|quill knobs]] on the forearm of a ''Velociraptor'' found in Mongolia.<ref name="turneretal2007b"/> These bumps on bird wing bones show where feathers anchor, and their presence on ''Velociraptor'' indicate it too had feathers. According to paleontologist Alan Turner, <blockquote>A lack of quill knobs does not necessarily mean that a dinosaur did not have feathers. Finding quill knobs on ''Velociraptor'', though, means that it definitely had feathers. This is something we'd long suspected, but no one had been able to prove.<ref name=sciencedaily2007feathers>American Museum of Natural History. "Velociraptor Had Feathers." ''ScienceDaily'' [[20 September]] [[2007]]. [[23 January]] [[2008]] http://www.sciencedaily.com/releases/2007/09/070920145402.htm</ref></blockquote> Co-author Mark Norell, Curator-in-Charge of fossil reptiles, amphibians and birds at the [[American Museum of Natural History]], also weighed in on the discovery, saying: <blockquote>The more that we learn about these animals the more we find that there is basically no difference between birds and their closely related dinosaur ancestors like velociraptor. Both have wishbones, brooded their nests, possess hollow bones, and were covered in feathers. If animals like velociraptor were alive today our first impression would be that they were just very unusual looking birds.<ref name=sciencedaily2007feathers/></blockquote> According to Turner and co-authors Norell and Peter Makovicky, quill knobs are not found in all prehistoric birds, and their absence does not mean that an animal was not feathered – [[flamingo]]s for example have no quill knobs. However, their presence confirms that ''Velociraptor'' bore modern-style wing feathers, with a rachis and vane formed by barbs. The forearm specimen on which the quill knobs were found (specimen number IGM 100/981) represents an animal 1.5 meters in length (5 ft) and 15 kilograms (33 lbs) in weight. Based on the spacing of the six preserved knobs in this specimen, the authors suggested that ''Velociraptor'' bore 14 secondaries (wing feathers stemming from the forearm), compared with 12 or more in ''[[Archaeopteryx]]'', 18 in ''[[Microraptor]]'', and 10 in ''[[Rahonavis]]''. This type of variation in the number of wing feathers between closely related species, the authors asserted, is to be expected, given similar variation among modern birds. Turner and colleagues interpreted the presence of feathers on ''Velociraptor'' as evidence against the idea that the larger, flightless maniraptorans lost their feathers secondarily due to larger body size. Furthermore, they noted that quill knobs are almost never found in flightless bird species today, and that their presence in ''Velociraptor'' (presumed to have been flightless due to its relatively large size and short forelimbs) is evidence that the ancestors of dromaeosaurids could fly, making ''Velociraptor'' and other large members of this family secondarily flightless, though it is possible the large wing feathers inferred in the ancestors of ''Velociraptor'' had a purpose other than flight. The feathers of the flightless ''Velociraptor'' may have been used for display, for covering their nests while brooding, or for added speed and thrust when running up inclined slopes.<ref name="turneretal2007b"/> ==In popular culture== {{Seealso|Biological issues in Jurassic Park#Velociraptor}} ''Velociraptor'' is well-known from its role as a vicious and cunning killer in the 1990 novel ''[[Jurassic Park]]'' by [[Michael Crichton]] and its 1993 [[Jurassic Park (film)|film adaptation]], directed by [[Steven Spielberg]]. The "raptors" portrayed in ''Jurassic Park'' were modeled after a larger relative, ''[[Deinonychus]]'', which [[Gregory Paul]] at the time called ''Velociraptor antirrhopus''.<ref name=paul1988/> The paleontologists in the film and the novel excavate a so-called ''Velociraptor'' skeleton in [[Montana]], far from the central Asian range of ''Velociraptor'' but well within the range of ''Deinonychus''. A character in Crichton's novel also states that "…''Deinonychus'' is now considered one of the velociraptors", indicating that Crichton used Paul's [[taxonomy]], though the "raptors" in the novel are referred to as ''V. mongoliensis''.<ref name=crichton1990>{{cite_book |last=Crichton |first=Michael. |authorlink=Michael Crichton |year=1990 |title=Jurassic Park |location=New York |publisher=Alfred A. Knopf |pages=pg. 114–115 |isbn=0-394-58816-9}}</ref> Steven Spielberg may also have increased the size of the film's ''Velociraptor'' for dramatic reasons.<ref name=bakker1995>{{cite_book |last=Bakker |first=Robert T. |authorlink=Bob Bakker |year=1995 |title=Raptor Red |location=New York |publisher=Bantam Books |pages=pg. 4 |isbn=0-553-57561-9}}</ref> Additionally, the forelimbs of the film animals differed in structure and posture from those of real dromaeosaurids and their tails were too short and flexible, anatomical errors which directly contradict fossil evidence. The film version of ''Velociraptor'' was also covered in [[Scale (zoology)|scales]]. In life, ''Velociraptor'', like many other maniraptoran theropods, was covered in feathers. In ''[[Jurassic Park III]]'', the ''Velociraptor'' are depicted with quill-like structures along the back of the head and neck, although these do not resemble the down-like feathers known from real-life dromaeosaurids, and the quill knobs on some ''Velociraptor'' specimens show that they had fully-developed feathers akin to those of modern birds.<ref name="turneretal2007b"/> Also in ''Jurassic Park III'', [[Alan Grant (Jurassic Park character)|Dr. Alan Grant]], played by [[Sam Neill]], states that ''Velociraptor'' were smarter than dolphins, whales and some primates. Based on fossil evidence, this is highly unlikely. It is more probable that, while intelligent by dinosaur standards, they were less intelligent than modern [[big cats]].<ref name=larssonetal2000>{{cite_journal |last=Larson |first=Hans C.E. |coauthors=[[Paul Sereno|Sereno, Paul C.]]; & [[Jeffrey A. Wilson|Wilson, Jeffrey A.]] |year=2000 |title=Forebrain enlargement among nonavian theropod dinosaurs |journal=Journal of Vertebrate Paleontology |volume=20 |issue=3 |pages=615–618 |url=http://vertpaleo.org/publications/jvp/contents-20-3.cfm |doi=10.1671/0272-4634(2000)020[0615:FEANTD]2.0.CO;2}}</ref> Due to the success of most ''Jurassic Park''-related products, ''Velociraptor'' has become a ubiquitous representation of dinosaurs in popular culture. It has been featured in numerous [[toy|toy lines]], [[animated films]], video games and [[List of animated television series|television series]] for children, along with several recent [[Documentary film|television documentaries]]. In 1995, the city of [[Toronto]] was awarded a [[National Basketball Association]] [[expansion team]], which was named the [[Toronto Raptors]]. ==References== {{reflist|2}} ==External links== {{portalpar|Dinosaurs}} {{Wikispecies|Velociraptor}} {{commons}} {{Wikibooks|Wikijunior Dinosaurs/Velociraptor}} *[http://paleo.amnh.org/fossil/show.html?cat_num=FR%206515 AMNH 6515 (holotype)] in the [http://paleo.amnh.org/fossil/seek.html Online Collections Database] at the American Museum of Natural History. *[http://www.skeletaldrawing.com/velociraptor/dromaeosaurpage.htm Several artistic renditions of ''Velociraptor''.] *[http://www.amnh.org/exhibitions/fightingdinos/videos.html A video of ''Protoceratops'' vs ''Velociraptor''] from the [http://www.amnh.org American Museum of Natural History]. {{featured article}} [[Category:Cretaceous dinosaurs]] [[Category:Dinosaurs of Asia]] [[Category:Dromaeosaurs]] [[Category:Feathered dinosaurs]] {{Link FA|es}} {{Link FA|hu}} [[ast:Velociraptor]] [[br:Velosiraptor]] [[ca:Velociraptor]] [[cs:Velociraptor]] [[da:Velociraptor]] [[de:Velociraptor]] [[es:Velociraptor]] [[fr:Velociraptor]] [[id:Velociraptor]] [[is:Snareðla]] [[it:Velociraptor mongoliensis]] [[he:ולוצירפטור]] [[lv:Velosireptors]] [[lt:Velociraptorius]] [[hu:Velociraptor]] [[ml:വെലോസിറാപ്റ്റര്‍]] [[ms:Velociraptor]] [[nl:Velociraptor]] [[ja:ヴェロキラプトル]] [[no:Velociraptor]] [[pl:Welociraptor]] [[pt:Velociráptor]] [[ru:Велоцираптор]] [[simple:Velociraptor]] [[sk:Velociraptor]] [[sl:Velociraptor]] [[fi:Velociraptor]] [[sv:Velociraptor]] [[ta:வெலாசிராப்டர்]] [[th:เวโลซีแรปเตอร์]] [[vi:Velociraptor]] [[bat-smg:Veluoceraptuorius]] [[zh:伶盜龍]]