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''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 past. The type species is ''V. mongoliensis''; fossils of this species have been discovered in the Djadochta Formation, Mongolia. A second species, ''V. osmolskae'', was named in 2008 for skull material from the Bayan Mandahu Formation, China.
Smaller than other dromaeosaurids like ''Deinonychus'' and ''Achillobator'', ''Velociraptor'' was about long with a body mass between . It nevertheless shared many of the same anatomy, anatomical features. It was a bipedal, feathered carnivore with a long tail and an enlarged sickle-shaped claw on each hindfoot, which is thought to have been used to tackle and restrain Predation, prey. ''Velociraptor'' can be distinguished from other dromaeosaurids by its long and low skull, with an upturned snout.
''Velociraptor'' (commonly referred to as "raptor") is one of the dinosaur genera most familiar to the general public due to its prominent role in the ''Jurassic Park'' films. In real life, however, ''Velociraptor'' was roughly the size of a Turkey (bird), turkey, considerably smaller than the approximately tall and reptiles seen in the novels and films (which were based on members of the related genus ''Deinonychus''). Today, ''Velociraptor'' is well known to paleontology, paleontologists, with over a dozen described fossil skeletons, the most of any dromaeosaurid. Fighting Dinosaurs, One particularly famous specimen preserves a ''Velociraptor'' locked in combat with a ''Protoceratops''.
Maxillae and a lacrimal bone, lacrimal (the main tooth-bearing bones of the upper jaw, and the bone that forms the anterior margin of the eye socket, respectively) recovered from the Bayan Mandahu Formation 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 colleagues named these bones ''V. osmolskae'' (for Polish paleontologist Halszka Osmólska) in 2008. However, the 2013 study noted that while "the elongate shape of the maxilla in ''V. osmolskae'' is similar to that of ''V. mongoliensis''," phylogenetic analysis found it to be closer to ''Linheraptor'', making the genus Paraphyly, paraphyletic; thus, ''V. osmolskae'' might not actually belong to the genus ''Velociraptor'' and requires reassessment.
Paleontologists Mark A. Norell and Peter J. Makovicky in 1997 described new and abundantly preserved specimens of ''V. mongoliensis'', namely MPC-D 100/985 collected from the Tugrik Shireh locality in 1993, and MPC-D 100/986 collected in 1993 from the Chimney Buttes locality. The team briefly mentioned another specimen, MPC-D 100/982, which by the time of this publication remained undescribed. In 1999 Norell and Makovicky provided more insights into the anatomy of ''Velociraptor'' with additional specimens. Among these, MPC-D 100/982 was partially described and figured, and referred to ''V. mongoliensis'' mainly based on cranial similarities with the holotype skull, although they stated that differences were present between the pelvic region of this specimen and other ''Velociraptor'' specimens. This relatively well-preserved specimen including the skull was discovered and collected in 1995 at the Bayn Dzak locality (more especifically at the "Volcano" sub-locality). Martin Kundrát in a 2004 abstract compared the neurocranium of MPC-D 100/982 to another ''Velociraptor'' specimen, MPC-D 100/976. He concluded that the overall morphology of the former was more derived (advanced) than the latter, suggesting that they could represent distinct taxa.
Mark J. Powers in his 2020 master thesis fully described MPC-D 100/982, which he concluded to represent a new and third species of ''Velociraptor''. This species, which he named ''"V. vadarostrum"'', was stated to mainly differ from other ''Velociraptor'' species in having a shallow maxilla morphology. Powers and colleagues also in 2020 used morphometric analyses to compare several dromaeosaurid maxillae, and found the maxilla of MPC-D 100/982 to strongly differ from specimens referred to ''Velociraptor''. They indicated that this specimen, based on these results, represents a different species. In 2021 Powers with team used Principal Component Analysis to separate dromaeosaurid maxillae, most notably finding that MPC-D 100/982 falls outside the instraspecific variability of ''V. mongoliensis'', arguing for a distinct species. They considered that both ''V. mongoliensis'' and this new species were ecologically separated based on their skull anatomy. The team in another 2021 abstract reinforced again the species-level separation, noting that additional differences can be found in the hindlimbs.
''Velociraptor'' was a small to medium-sized dromaeosaurid, with adults measuring between long, approximately high at the hips, and weighing about .
Prominent quill knobs—attachment site of "wing" feathers and direct indicator of a feather covering—have been reported from the ulna of a single ''Velociraptor'' specimen (IGM 100/981), which represents an animal of estimated long and in weight. The spacing of 6 preserved knobs suggests that 8 additional knobs may have been present, giving a total of 14 quill knobs that developed large Flight feather#Secondaries, secondaries ("wing" feathers stemming from the forearm). However, the specimen number has been corrected to IGM 100/3503 and its referral to ''Velociraptor'' may require reevaluation, pending further study. Nevertheless, there is strong Phylogenetic bracketing, phylogenetic evidence from other dromaeosaurid relatives that indicates the presence of feathers in ''Velociraptor'', including dromaeosaurids such as ''Microraptor'' or ''Zhenyuanlong''.
The back or anterior region of the skull was built by the frontal, lacrimal, postorbital, jugal, parietal, quadrate, and quadratojugal bones. The was large element, having a vaguely rectangular shape when seen from above. On its posterior end, this bone was in contact with the , and such elements were the main bodies of the skull roof. The was a T-shaped bone and its main body was thin and delicated. Its lower end meet the (often called cheek bone), which was a large, sub-triangular-shaped element. Its lower border was notably straight/horizontal. The was located just above the jugal: a stocky and strongly T-shaped bone. As a whole, the orbit or orbital fenestra (eye socket)—formed by the lacrimal, jugal, frontal, and postorbital—was large and near circular in shape, being longer than taller. When seen from above, a pair of large and markedly rounded holes were present near the rear of the skull (the temporal fenestrae), whose main components were the postorbital and squamosal. Behind the jugal, an inverted T-shaped bone (also seen in other dromaeosaurids), known as the , was developed. While the upper end of the quadratojugal joined the , an irregularly-shaped element, its inner side meet the . The latter was of great importance for the articulation with the lower jaw. The posteriormost bone was the and its projection the occipital condyle: a rounded and bulbous protuberance that meet the first vertebra of the neck.
The lower jaw of ''Velociraptor'' comprised mainly the dentary, splenial, angular, surangular, and articular bones. The was a very long, weakly curved, and narrow element that developed several alveoli on its top surface. On its posterior end, it meet the . It had a small hole near its posterior end, called surangular foramen or fenestra. Both bones were the largest elements of the lower jaw of ''Velociraptor'', contributing to virtually its entire length. Below them were the smaller and , closely articulated to each other. The , located on the inner side of the surangular, was a small element that joined the quadrate of the upper skull, enabling the articulation with the lower jaw. An elongated, near oval-shaped hole was developed in the center of the lower jaw (the mandibular fenestra), and it was produced by the joint of the dentary, surangular, and angular bones.
The teeth of ''Velociraptor'' were fairly homodont (equal in shape) and had several Denticle (tooth feature), denticles (serrations), each more strongly serrated on the back edge than the front. The premaxilla had 4 alveoli (meaning that 4 teeth were developed), and the maxilla had 11 alveoli. At the dentary, between 14-15 alveoli were present. All teeth present at the premaxilla were poorly curved, and the two first teeth were the longest, with the second having a characteristic large size. The maxillary teeth were more slender, recurved, and most notably, the lower end was strongly more serrated than the upper one.
The arm of ''Velociraptor'' was formed by the humerus (upper arm bone), Radius (bone), radius and ulna (forearm bones), and Manus (zoology), manus (hand). ''Velociraptor'', like other dromaeosaurids, had a large manus with three elongated Digit (anatomy), digits (fingers), which ended up in strongly curved unguals (claw bones) that were similar in construction and flexibility to the wing bones of modern birds. 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 manus to be held with the Anatomical terms of location, palmar surface facing inward (Lateral and medial, medially), not downward. The Pes (anatomy), pes (foot) anatomy of ''Velociraptor'' consisted of the metatarsus—a large element composed of three metatarsals of which the first one was extremely reduced in size—and four digits that developed large unguals. The first digit, as in other theropods, was a small dewclaw. The second digit, for which ''Velociraptor'' is most famous, was highly modified and held retracted off the ground, which caused ''Velociraptor'' and other dromaeosaurids to walk on only their third and fourth digits. It bore a relatively large, sickle-shaped claw, typical of dromaeosaurid and troodontid dinosaurs. This enlarged claw, which could grow to over long around its outer edge, was most likely a predatory device used to restrain struggling prey.
As in other dromaeosaurs, ''Velociraptor'' tails had prezygapophysis, prezygapophyses (long bony projections) on the upper surfaces of the vertebrae, as well as ossification, ossified tendons underneath. 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. These were once thought to fully stiffen the tail, forcing the entire tail to act as a single rod-like unit. However, at least one specimen has preserved a series of intact tail vertebrae curved sideways into an ''S''-shape, suggesting that there was considerably more horizontal flexibility than once thought.
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Below are the results for the Eudromaeosauria phylogeny based on the phylogenetics, phylogenetic analysis conducted by James G. Napoli and team in 2021 during the description of ''Kuru kulla, Kuru'', showing the position of ''Velociraptor'':
In 2007 Alan H. Turner and colleagues reported the presence of six quill knobs in the ulna of a referred ''Velociraptor'' specimen (IGM 100/981) from the Ukhaa Tolgod locality of the Djadochta Formation. 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. Because of the presence of another dromaeosaurid in Ukhaa Tolgod, ''Tsaagan'', Napoli and team have noted that the referral of this specimen to ''Velociraptor'' is currently subject to rexamination.
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. 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 preserved. The thick abdominal wall of skin and muscle of large prey species would have been difficult to slash without a specialized cutting surface. 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.
Remains of ''Deinonychus'', a closely related dromaeosaurid, have commonly been found in aggregations of several individuals. ''Deinonychus'' has also been found in association with the large ornithopod ''Tenontosaurus'', which has been cited as evidence of cooperative (pack) hunting. However, the only solid evidence for social behavior of any kind among dromaeosaurids comes from a Chinese trackway which shows six individuals of a large species moving as a group. Although many isolated fossils of ''Velociraptor'' have been found in Mongolia, none were closely associated with other individuals. 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. Dromeosaur footprints in China suggest that a few other raptor genera may have hunted in packs, but there have been no conclusive examples of pack behavior found.
In 2011, Denver Fowler and colleagues suggested a new method by which dromaeosaurs like ''Velociraptor'' and similar dromaeosaurs may have captured and restrained prey. This model, known as the "raptor prey restraint" (RPR) model of predation, proposes that dromaeosaurs killed their prey in a manner very similar to extant Accipitridae, accipitrid birds of prey: by leaping onto their quarry, pinning it under their body weight, and gripping it tightly with the large, sickle-shaped claws. These researchers proposed that, like accipitrids, the dromaeosaur would then begin to feed on the animal while it was still alive, and prey death would eventually result from blood loss and organ failure. This proposal is based primarily on comparisons between the morphology and proportions of the feet and legs of dromaeosaurs to several groups of extant birds of prey with known predatory behaviors. Fowler found that the feet and legs of dromaeosaurs most closely resemble those of eagles and hawks, especially in terms of having an enlarged second claw and a similar range of grasping motion. The short Tarsometatarsus, metatarsus and foot strength, however, would have been more similar to that of owls. The RPR method of predation would be consistent with other aspects of ''Velociraptor''s anatomy, such as their unusual jaw and arm morphology. The arms, which could exert a lot of force but were likely covered in long feathers, may have been used as flapping stabilizers for balance while atop a struggling prey animal, along with the stiff counterbalancing tail. The jaws, thought by Fowler and colleagues to be comparatively weak, would have been useful for row saw motion bites like the modern day Komodo dragon, which also has a weak bite, to finish off its prey if the kicks weren't powerful enough. These predatory adaptations working together may also have implications for the Origin of avian flight, origin of flapping in paravians.
In both Bayan Mandahu Formation, Bayan Mandahu and Djadochta formations many of the same genera were present, though they varied at the species level. These differences in species composition may be due a natural barrier separating the two formations, which are relatively close to each other geographically. However, given the lack of any known barrier which would cause the specific faunal compositions found in these areas, it is more likely that those differences indicate a slight time difference.
''V. osmolskae'' lived alongside the ankylosaurid ''Pinacosaurus mephistocephalus''; alvarezsaurid ''Linhenykus''; closely related dromaeosaurid ''Linheraptor''; oviraptorids ''Machairasaurus'' and ''Wulatelong''; protoceratopsids ''Bagaceratops'' and ''Protoceratops hellenikorhinus''; and troodontids ''Linhevenator'', ''Papiliovenator'', and ''Philovenator''. Sediments across the formation indicate a similar depositional environment to that of the Djadochta Formation.
Known specimens of ''Velociraptor mongoliensis'' have been recovered from the Djadochta Formation (also spelled Djadokhta), in the Mongolian province of Ömnögovi Province, Ömnögovi. This geological formation is estimated to date back to the Campanian Stage (stratigraphy), stage (between 75 million and 71 million years ago) of the Late Cretaceous Geologic time scale#Divisions of geologic time, epoch. The abundant sediments—sands, sandstones, or caliche—of the Djadochta Formation were deposited by Aeolian processes, eolian (wind) processes in arid settings with fields of sand dunes and only intermittent streams, as indicated by very sparse fluvial (river-deposited) sedimentation, under a semi-arid climate.
The Djadochta Formation is separated into a lower Bayn Dzak Member and upper Turgrugyin Member. ''V. mongoliensis'' is known from both members, represented by numerous specimens. The Bayn Dzak Member (mainly Bayn Dzak locality) has yielded the oviraptorid ''Oviraptor''; ankylosaurid ''Pinacosaurus grangeri''; protoceratopsid ''Protoceratops andrewsi''; and troodontid ''Saurornithoides''. The younger Turgrugyin Member (mainly Tugriken Shireh locality) has produced the bird ''Elsornis''; dromaeosaurid ''Mahakala omnogovae, Mahakala'': ornithomimid ''Aepyornithomimus''; and protoceratopsid ''Protoceratops andrewsi''.
''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 (sublocality of the larger Bayn Dzak locality/region), while the "Fighting Dinosaurs" were found at the Tugrik Shire locality (also known as Tugrugeen Shireh and many other spellings). The latter is notorious for its exceptional ''in situ'' fossil preservation. Based on deposits (such as structureless sandstones), it has been concluded that a large number of specimens were buried alive during powerful sand-bearing events, common to the these paleoenvironments.
Yale’s legacy in ''Jurassic World''
" ''Yale News'', 18-Jun-2015. They portrayed the animals with the size, proportions, and snout shape of ''Deinonychus'' rather than ''Velociraptor''. Production on ''Jurassic Park'' began before the discovery of the large dromaeosaurid ''Utahraptor'' was made public in 1991, but as Jody Duncan wrote about this discovery: "Later, after we had designed and built the raptor, there was a discovery of a raptor skeleton in Utah, which they labeled 'super-slasher.' They had uncovered the largest Velociraptor to date and it measured five-and-a-half-feet tall, just like ours. So we designed it, we built it, and then they discovered it. That still boggles my mind." Spielberg's name was briefly considered for naming of the new dinosaur in exchange for funding of field work, but no agreement was reached. ''Jurassic Park (film), Jurassic Park'' and its sequel ''The Lost World: Jurassic Park'' were released before the discovery that dromaeosaurs had feathers, so the ''Velociraptor'' in both films were depicted as scaled and featherless. For ''Jurassic Park III,'' the male ''Velociraptor'' was given quill-like structures along the back of the head and neck, but these structures do not resemble the feathers that ''Velociraptor'' would have had in reality due to reasons of continuity. The ''Jurassic World'' sequel trilogy ignored the feathers of ''Velociraptor'', adhering to the designs from ''Jurassic Park''. However, the dromaeosaur ''Pyroraptor'' was feathered for ''Jurassic World Dominion'', along with other changes such as stiffening the tail to account for ossified tendons and de-pronating the hands.
3D skull model of ''Velociraptor mongoliensis''
at Sketchfab
Skeletal reconstruction of ''Velociraptor mongoliensis''
at Dr. Scott Hartman's Skeletal Drawing
at American Museum of Natural History (Wayback Machine) {{featured article Campanian genus first appearances Djadochta fauna Eudromaeosaurs Feathered dinosaurs Fossil taxa described in 1924 Late Cretaceous dinosaurs of Asia Taxa named by Henry Fairfield Osborn
History of discovery
During an American Museum of Natural History expedition to the Flaming Cliffs (Bayn Dzak or Bayanzag) of the Djadochta Formation, Gobi Desert, on 11 August 1923, Peter Kaisen discovered 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. Earlier that year, Osborn had informally mentioned the animal in a popular press article, under the name "Ovoraptor djadochtari" (not to be confused with the similarly named ''Oviraptor''), eventually changed into ''V. mongoliensis'' during its formal description. While North American teams were shut out of Mongolian People's Republic, 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 "Fighting Dinosaurs" specimen (Mongolian Paleontological Center, MPC-D 100/25; formerly IGM, GIN, or GI SPS), discovered by a Polish-Mongolian team in 1971. The fossil preserves a ''Velociraptor'' in battle against a ''Protoceratops''. It is considered a national treasure of Mongolia, and in 2000 it was loaned to the American Museum of Natural History in New York City for a temporary exhibition. Between 1988 and 1990, a joint China, Chinese-Canadians, Canadian team discovered ''Velociraptor'' remains in northern China. 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. One such specimen, MPC-D 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).Novacek, Michael J. (1996). ''Dinosaurs of the Flaming Cliffs''. New York: Anchor Books. . While Norell and Makovicky provisionally considered it a specimen of ''Velociraptor mongoliensis'', it was named as a new species ''Shri devi'' in 2021.Additional species
Maxillae and a lacrimal bone, lacrimal (the main tooth-bearing bones of the upper jaw, and the bone that forms the anterior margin of the eye socket, respectively) recovered from the Bayan Mandahu Formation 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 colleagues named these bones ''V. osmolskae'' (for Polish paleontologist Halszka Osmólska) in 2008. However, the 2013 study noted that while "the elongate shape of the maxilla in ''V. osmolskae'' is similar to that of ''V. mongoliensis''," phylogenetic analysis found it to be closer to ''Linheraptor'', making the genus Paraphyly, paraphyletic; thus, ''V. osmolskae'' might not actually belong to the genus ''Velociraptor'' and requires reassessment.
Paleontologists Mark A. Norell and Peter J. Makovicky in 1997 described new and abundantly preserved specimens of ''V. mongoliensis'', namely MPC-D 100/985 collected from the Tugrik Shireh locality in 1993, and MPC-D 100/986 collected in 1993 from the Chimney Buttes locality. The team briefly mentioned another specimen, MPC-D 100/982, which by the time of this publication remained undescribed. In 1999 Norell and Makovicky provided more insights into the anatomy of ''Velociraptor'' with additional specimens. Among these, MPC-D 100/982 was partially described and figured, and referred to ''V. mongoliensis'' mainly based on cranial similarities with the holotype skull, although they stated that differences were present between the pelvic region of this specimen and other ''Velociraptor'' specimens. This relatively well-preserved specimen including the skull was discovered and collected in 1995 at the Bayn Dzak locality (more especifically at the "Volcano" sub-locality). Martin Kundrát in a 2004 abstract compared the neurocranium of MPC-D 100/982 to another ''Velociraptor'' specimen, MPC-D 100/976. He concluded that the overall morphology of the former was more derived (advanced) than the latter, suggesting that they could represent distinct taxa.
Mark J. Powers in his 2020 master thesis fully described MPC-D 100/982, which he concluded to represent a new and third species of ''Velociraptor''. This species, which he named ''"V. vadarostrum"'', was stated to mainly differ from other ''Velociraptor'' species in having a shallow maxilla morphology. Powers and colleagues also in 2020 used morphometric analyses to compare several dromaeosaurid maxillae, and found the maxilla of MPC-D 100/982 to strongly differ from specimens referred to ''Velociraptor''. They indicated that this specimen, based on these results, represents a different species. In 2021 Powers with team used Principal Component Analysis to separate dromaeosaurid maxillae, most notably finding that MPC-D 100/982 falls outside the instraspecific variability of ''V. mongoliensis'', arguing for a distinct species. They considered that both ''V. mongoliensis'' and this new species were ecologically separated based on their skull anatomy. The team in another 2021 abstract reinforced again the species-level separation, noting that additional differences can be found in the hindlimbs.
Description
''Velociraptor'' was a small to medium-sized dromaeosaurid, with adults measuring between long, approximately high at the hips, and weighing about .
Prominent quill knobs—attachment site of "wing" feathers and direct indicator of a feather covering—have been reported from the ulna of a single ''Velociraptor'' specimen (IGM 100/981), which represents an animal of estimated long and in weight. The spacing of 6 preserved knobs suggests that 8 additional knobs may have been present, giving a total of 14 quill knobs that developed large Flight feather#Secondaries, secondaries ("wing" feathers stemming from the forearm). However, the specimen number has been corrected to IGM 100/3503 and its referral to ''Velociraptor'' may require reevaluation, pending further study. Nevertheless, there is strong Phylogenetic bracketing, phylogenetic evidence from other dromaeosaurid relatives that indicates the presence of feathers in ''Velociraptor'', including dromaeosaurids such as ''Microraptor'' or ''Zhenyuanlong''.
Skull
The skull of ''Velociraptor'' was rather elongated and grew up to long. It was uniquely up-curved at the snout region, concave on the upper surface, and convex on the lower surface. The snout, which occupied about 60% of the entire skull length, was notably narrow and mainly formed by the nasal, premaxilla, and maxilla bones. The was the anteriormost bone in the skull, and it was longer than taller. While its posterior end joined the nasal, the main body of the premaxilla touched the maxilla. The was nearly triangular in shape and the largest element of the snout. On its center or main body, there was a depression developing a small oval to circular-shaped hole, called maxillary fenestra. Though in front of this fenestra were two small openings, referred to as promaxillary fenestrae. The posterior border of the maxilla formed (predominantly) the antorbital fenestra, one of the several large holes in the skull. Both premaxilla and maxilla had several Dental alveolus, alveoli (tooth sockets) on their bottom surfaces. Above the maxilla and making contact with the premaxilla, there was the bone. It was a thin/narrow and elongated bone contributing to the top surface of the snout. Together, both premaxilla and nasal bones gave form to the naris or narial fenestra (nostril opening), which was relatively large and circular. The posterior end of the nasal was joined by the frontal and lacrimal bones.
The back or anterior region of the skull was built by the frontal, lacrimal, postorbital, jugal, parietal, quadrate, and quadratojugal bones. The was large element, having a vaguely rectangular shape when seen from above. On its posterior end, this bone was in contact with the , and such elements were the main bodies of the skull roof. The was a T-shaped bone and its main body was thin and delicated. Its lower end meet the (often called cheek bone), which was a large, sub-triangular-shaped element. Its lower border was notably straight/horizontal. The was located just above the jugal: a stocky and strongly T-shaped bone. As a whole, the orbit or orbital fenestra (eye socket)—formed by the lacrimal, jugal, frontal, and postorbital—was large and near circular in shape, being longer than taller. When seen from above, a pair of large and markedly rounded holes were present near the rear of the skull (the temporal fenestrae), whose main components were the postorbital and squamosal. Behind the jugal, an inverted T-shaped bone (also seen in other dromaeosaurids), known as the , was developed. While the upper end of the quadratojugal joined the , an irregularly-shaped element, its inner side meet the . The latter was of great importance for the articulation with the lower jaw. The posteriormost bone was the and its projection the occipital condyle: a rounded and bulbous protuberance that meet the first vertebra of the neck.
The lower jaw of ''Velociraptor'' comprised mainly the dentary, splenial, angular, surangular, and articular bones. The was a very long, weakly curved, and narrow element that developed several alveoli on its top surface. On its posterior end, it meet the . It had a small hole near its posterior end, called surangular foramen or fenestra. Both bones were the largest elements of the lower jaw of ''Velociraptor'', contributing to virtually its entire length. Below them were the smaller and , closely articulated to each other. The , located on the inner side of the surangular, was a small element that joined the quadrate of the upper skull, enabling the articulation with the lower jaw. An elongated, near oval-shaped hole was developed in the center of the lower jaw (the mandibular fenestra), and it was produced by the joint of the dentary, surangular, and angular bones.
The teeth of ''Velociraptor'' were fairly homodont (equal in shape) and had several Denticle (tooth feature), denticles (serrations), each more strongly serrated on the back edge than the front. The premaxilla had 4 alveoli (meaning that 4 teeth were developed), and the maxilla had 11 alveoli. At the dentary, between 14-15 alveoli were present. All teeth present at the premaxilla were poorly curved, and the two first teeth were the longest, with the second having a characteristic large size. The maxillary teeth were more slender, recurved, and most notably, the lower end was strongly more serrated than the upper one.
Postcranial skeleton
The arm of ''Velociraptor'' was formed by the humerus (upper arm bone), Radius (bone), radius and ulna (forearm bones), and Manus (zoology), manus (hand). ''Velociraptor'', like other dromaeosaurids, had a large manus with three elongated Digit (anatomy), digits (fingers), which ended up in strongly curved unguals (claw bones) that were similar in construction and flexibility to the wing bones of modern birds. 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 manus to be held with the Anatomical terms of location, palmar surface facing inward (Lateral and medial, medially), not downward. The Pes (anatomy), pes (foot) anatomy of ''Velociraptor'' consisted of the metatarsus—a large element composed of three metatarsals of which the first one was extremely reduced in size—and four digits that developed large unguals. The first digit, as in other theropods, was a small dewclaw. The second digit, for which ''Velociraptor'' is most famous, was highly modified and held retracted off the ground, which caused ''Velociraptor'' and other dromaeosaurids to walk on only their third and fourth digits. It bore a relatively large, sickle-shaped claw, typical of dromaeosaurid and troodontid dinosaurs. This enlarged claw, which could grow to over long around its outer edge, was most likely a predatory device used to restrain struggling prey.
As in other dromaeosaurs, ''Velociraptor'' tails had prezygapophysis, prezygapophyses (long bony projections) on the upper surfaces of the vertebrae, as well as ossification, ossified tendons underneath. 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. These were once thought to fully stiffen the tail, forcing the entire tail to act as a single rod-like unit. However, at least one specimen has preserved a series of intact tail vertebrae curved sideways into an ''S''-shape, suggesting that there was considerably more horizontal flexibility than once thought.
Classification
''Velociraptor'' is a member of the group Eudromaeosauria, a Synapomorphy, derived sub-group of the larger family Dromaeosauridae. It is often placed within its own subfamily, Velociraptorinae. In phylogenetic taxonomy, Velociraptorinae is usually defined as "all dromaeosaurs more closely related to ''Velociraptor'' than to ''Dromaeosaurus''." However, dromaeosaurid classification is highly variable. Originally, the subfamily Velociraptorinae was erected solely to contain ''Velociraptor''. Other analyses have often included other genera, usually ''Deinonychus'' and ''Saurornitholestes'', and more recently ''Tsaagan''. Several studies published during the 2010s, including expanded versions of the analyses that found support for Velociraptorinae, have failed to resolve it as a distinct group, but rather have suggested it is a paraphyletic grade which gave rise to the Dromaeosaurinae. 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). 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). 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''. As of 2008, the only currently recognized species of ''Velociraptor'' are ''V. mongoliensis'' and ''V. osmolskae''. However, several studies have found "''V.''" ''osmolskae'' to be distantly related to ''V. mongoliensis''. Material was copied from this source, which is available under Creative Commons Attribution 4.0 International License
Below are the results for the Eudromaeosauria phylogeny based on the phylogenetics, phylogenetic analysis conducted by James G. Napoli and team in 2021 during the description of ''Kuru kulla, Kuru'', showing the position of ''Velociraptor'':
Paleobiology
Feathers
In 2007 Alan H. Turner and colleagues reported the presence of six quill knobs in the ulna of a referred ''Velociraptor'' specimen (IGM 100/981) from the Ukhaa Tolgod locality of the Djadochta Formation. 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. Because of the presence of another dromaeosaurid in Ukhaa Tolgod, ''Tsaagan'', Napoli and team have noted that the referral of this specimen to ''Velociraptor'' is currently subject to rexamination.
Senses
Examinations of the endocranium of ''Velociraptor'' indicate that it was able to detect and hear a wide range of sound frequencies (2,368–3,965 Hz) and could track prey with ease as a result. The endocranium examinations also further cemented the theory that the dromaeosaur was an agile, swift predator. Fossil evidence suggesting ''Velociraptor'' scavenged also indicates that it was an opportunistic and actively predatory animal, feeding on carrion during times of drought or famine, if in poor health, or depending on the animal's age.Feeding
In 2020, Powers and colleagues re-examined the maxillae of several eudromaeosaur taxa concluding that most Asian and North American eudromaeosaurs were separated by snout morphology and ecological strategies. They found the maxilla to be a reliable reference when inferring the shape of the premaxilla and overall snout. For instance, most Asian species have elongated snouts based on the maxilla (namely velociraptorines), indicating a selective feeding in ''Velociraptor'' and relatives, such as picking up small, fast prey. In contrast, most North American eudromaeosaurs, mostly dromaeosaurines, feature a robust and deep maxillar morphology. However, the large dromaeosurine ''Achillobator'' is a unique exception to Asian taxa with its deep maxilla.Predatory behavior
The "Fighting Dinosaurs" specimen, found in 1971, preserves a ''Velociraptor mongoliensis'' and ''Protoceratops, Protoceratops andrewsi'' in combat and provides direct evidence of predatory behavior. When originally reported, it was hypothesized that the two animals drowned. 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. Parts of the ''Protoceratops'' are missing, which has been seen as evidence of scavenger, scavenging by other animals. Comparisons between the sclerotic ring, scleral rings of ''Velociraptor'', ''Protoceratops'', and modern birds and reptiles indicates that ''Velociraptor'' may have been nocturnal, while ''Protoceratops'' may have been cathemeral, active throughout the day during short intervals, suggesting that the fight may have occurred at twilight or during low-light conditions.
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. 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 preserved. The thick abdominal wall of skin and muscle of large prey species would have been difficult to slash without a specialized cutting surface. 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.
Remains of ''Deinonychus'', a closely related dromaeosaurid, have commonly been found in aggregations of several individuals. ''Deinonychus'' has also been found in association with the large ornithopod ''Tenontosaurus'', which has been cited as evidence of cooperative (pack) hunting. However, the only solid evidence for social behavior of any kind among dromaeosaurids comes from a Chinese trackway which shows six individuals of a large species moving as a group. Although many isolated fossils of ''Velociraptor'' have been found in Mongolia, none were closely associated with other individuals. 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. Dromeosaur footprints in China suggest that a few other raptor genera may have hunted in packs, but there have been no conclusive examples of pack behavior found.
In 2011, Denver Fowler and colleagues suggested a new method by which dromaeosaurs like ''Velociraptor'' and similar dromaeosaurs may have captured and restrained prey. This model, known as the "raptor prey restraint" (RPR) model of predation, proposes that dromaeosaurs killed their prey in a manner very similar to extant Accipitridae, accipitrid birds of prey: by leaping onto their quarry, pinning it under their body weight, and gripping it tightly with the large, sickle-shaped claws. These researchers proposed that, like accipitrids, the dromaeosaur would then begin to feed on the animal while it was still alive, and prey death would eventually result from blood loss and organ failure. This proposal is based primarily on comparisons between the morphology and proportions of the feet and legs of dromaeosaurs to several groups of extant birds of prey with known predatory behaviors. Fowler found that the feet and legs of dromaeosaurs most closely resemble those of eagles and hawks, especially in terms of having an enlarged second claw and a similar range of grasping motion. The short Tarsometatarsus, metatarsus and foot strength, however, would have been more similar to that of owls. The RPR method of predation would be consistent with other aspects of ''Velociraptor''s anatomy, such as their unusual jaw and arm morphology. The arms, which could exert a lot of force but were likely covered in long feathers, may have been used as flapping stabilizers for balance while atop a struggling prey animal, along with the stiff counterbalancing tail. The jaws, thought by Fowler and colleagues to be comparatively weak, would have been useful for row saw motion bites like the modern day Komodo dragon, which also has a weak bite, to finish off its prey if the kicks weren't powerful enough. These predatory adaptations working together may also have implications for the Origin of avian flight, origin of flapping in paravians.
Scavenging behavior
In 2010, Hone and colleagues published a paper on their 2008 discovery of shed teeth of what they believed to be a ''Velociraptor'' near a tooth-marked jaw bone of what they believed to be a ''Protoceratops'' in the Bayan Mandahu Formation. The authors concluded that the find represented "late-stage carcass consumption by ''Velociraptor''" as the predator would have eaten other parts of a freshly killed ''Protoceratops'' before biting in the jaw area. The evidence was seen as supporting the inference from the "Fighting Dinosaurs" fossil that ''Protoceratops'' was part of the diet of ''Velociraptor''. In 2012, Hone and colleagues published a paper that described a ''Velociraptor'' specimen with a long bone of an azhdarchid pterosaur in its gut. This was interpreted as showing scavenging behaviour.Metabolism
''Velociraptor'' was 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 (bird), 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.Paleopathology
Norell with colleagues in 1995 reported one ''V. mongoliensis'' skull bearing two parallel rows of small punctures on its frontal bones that, upon closer examination, match the spacing and size of ''Velociraptor'' teeth. They suggested that the wound was likely inflicted by another ''Velociraptor'' during a Intraspecific competition, fight between the species. Because its bone structure shows no sign of Bone healing, healing near the bite wounds and the overall specimen was not scavenged, this individual was likely killed by this fatal wound. In 2001 Molnar and team noted that this specimen is MPC-D 100/976 hailing from the Tugrik Shireh locality, which has also yielded the Fighting Dinosaurs specimen. In 2012 David Hone and team reported another injured ''Velociraptor'' specimen (MPC-D 100/54, roughly a sub-adult individual) found with the bones of an Azhdarchidae, azhdarchid pterosaur within its stomach cavity, was carrying or recovering from an injury sustained to one broken rib. From evidence on the pterosaur bones, which were devoid of pitting or deformations from digestion, the ''Velociraptor'' died shortly after, possibly from the earlier injury. Nevertheless, the team noted that this broken ribs shows signs of bone healing.Paleoenvironment
Bayan Mandahu Formation
In both Bayan Mandahu Formation, Bayan Mandahu and Djadochta formations many of the same genera were present, though they varied at the species level. These differences in species composition may be due a natural barrier separating the two formations, which are relatively close to each other geographically. However, given the lack of any known barrier which would cause the specific faunal compositions found in these areas, it is more likely that those differences indicate a slight time difference.
''V. osmolskae'' lived alongside the ankylosaurid ''Pinacosaurus mephistocephalus''; alvarezsaurid ''Linhenykus''; closely related dromaeosaurid ''Linheraptor''; oviraptorids ''Machairasaurus'' and ''Wulatelong''; protoceratopsids ''Bagaceratops'' and ''Protoceratops hellenikorhinus''; and troodontids ''Linhevenator'', ''Papiliovenator'', and ''Philovenator''. Sediments across the formation indicate a similar depositional environment to that of the Djadochta Formation.
Djadochta Formation
Known specimens of ''Velociraptor mongoliensis'' have been recovered from the Djadochta Formation (also spelled Djadokhta), in the Mongolian province of Ömnögovi Province, Ömnögovi. This geological formation is estimated to date back to the Campanian Stage (stratigraphy), stage (between 75 million and 71 million years ago) of the Late Cretaceous Geologic time scale#Divisions of geologic time, epoch. The abundant sediments—sands, sandstones, or caliche—of the Djadochta Formation were deposited by Aeolian processes, eolian (wind) processes in arid settings with fields of sand dunes and only intermittent streams, as indicated by very sparse fluvial (river-deposited) sedimentation, under a semi-arid climate.
The Djadochta Formation is separated into a lower Bayn Dzak Member and upper Turgrugyin Member. ''V. mongoliensis'' is known from both members, represented by numerous specimens. The Bayn Dzak Member (mainly Bayn Dzak locality) has yielded the oviraptorid ''Oviraptor''; ankylosaurid ''Pinacosaurus grangeri''; protoceratopsid ''Protoceratops andrewsi''; and troodontid ''Saurornithoides''. The younger Turgrugyin Member (mainly Tugriken Shireh locality) has produced the bird ''Elsornis''; dromaeosaurid ''Mahakala omnogovae, Mahakala'': ornithomimid ''Aepyornithomimus''; and protoceratopsid ''Protoceratops andrewsi''.
''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 (sublocality of the larger Bayn Dzak locality/region), while the "Fighting Dinosaurs" were found at the Tugrik Shire locality (also known as Tugrugeen Shireh and many other spellings). The latter is notorious for its exceptional ''in situ'' fossil preservation. Based on deposits (such as structureless sandstones), it has been concluded that a large number of specimens were buried alive during powerful sand-bearing events, common to the these paleoenvironments.
Cultural significance
''Velociraptor'' is commonly perceived as a vicious and cunning killer thanks to their portrayal in the 1990 novel ''Jurassic Park (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 actually modeled after the closely related Dromaeosauridae, dromaeosaurid ''Deinonychus''. Paleontologists in both the novel and film excavate a skeleton in Montana, far from the central Asian range of ''Velociraptor'' but characteristic of the ''Deinonychus'' range. Crichton met with the discoverer of ''Deinonychus'', John Ostrom, several times at Yale University to discuss details of the animal's possible range of behaviors and appearance. Crichton at one point apologetically told Ostrom that he had decided to use the name ''Velociraptor'' in place of ''Deinonychus'' because the former name was "more dramatic." According to Ostrom, Crichton stated that the ''Velociraptor'' of the novel was based on ''Deinonychus'' in almost every detail, and that only the name had been changed. The ''Jurassic Park'' film-makers also requested all of Ostrom's published papers on ''Deinonychus'' during production.Cummings, M.Yale’s legacy in ''Jurassic World''
" ''Yale News'', 18-Jun-2015. They portrayed the animals with the size, proportions, and snout shape of ''Deinonychus'' rather than ''Velociraptor''. Production on ''Jurassic Park'' began before the discovery of the large dromaeosaurid ''Utahraptor'' was made public in 1991, but as Jody Duncan wrote about this discovery: "Later, after we had designed and built the raptor, there was a discovery of a raptor skeleton in Utah, which they labeled 'super-slasher.' They had uncovered the largest Velociraptor to date and it measured five-and-a-half-feet tall, just like ours. So we designed it, we built it, and then they discovered it. That still boggles my mind." Spielberg's name was briefly considered for naming of the new dinosaur in exchange for funding of field work, but no agreement was reached. ''Jurassic Park (film), Jurassic Park'' and its sequel ''The Lost World: Jurassic Park'' were released before the discovery that dromaeosaurs had feathers, so the ''Velociraptor'' in both films were depicted as scaled and featherless. For ''Jurassic Park III,'' the male ''Velociraptor'' was given quill-like structures along the back of the head and neck, but these structures do not resemble the feathers that ''Velociraptor'' would have had in reality due to reasons of continuity. The ''Jurassic World'' sequel trilogy ignored the feathers of ''Velociraptor'', adhering to the designs from ''Jurassic Park''. However, the dromaeosaur ''Pyroraptor'' was feathered for ''Jurassic World Dominion'', along with other changes such as stiffening the tail to account for ossified tendons and de-pronating the hands.
See also
* Timeline of dromaeosaurid research *Fighting DinosaursReferences
External links
* * * *3D skull model of ''Velociraptor mongoliensis''
at Sketchfab
Skeletal reconstruction of ''Velociraptor mongoliensis''
at Dr. Scott Hartman's Skeletal Drawing
at American Museum of Natural History (Wayback Machine) {{featured article Campanian genus first appearances Djadochta fauna Eudromaeosaurs Feathered dinosaurs Fossil taxa described in 1924 Late Cretaceous dinosaurs of Asia Taxa named by Henry Fairfield Osborn