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Velociraptor

Velociraptor

Overview
Velociraptor is a genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of dromaeosaurid
Dromaeosauridae
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'...

 theropod
Theropoda
Theropoda is both a suborder of bipedal saurischian dinosaurs, and a clade consisting of that suborder and its descendants . Dinosaurs belonging to the suborder theropoda were primarily carnivorous, although a number of theropod groups evolved herbivory, omnivory, and insectivory...

 dinosaur
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...

 that existed approximately 75 to 71 million years ago during the later part of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 Period. Two species are currently recognized, although others have been assigned in the past. The type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 is V. mongoliensis; fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

s of this species have been discovered in Mongolia
Mongolia
Mongolia is a landlocked country in East and Central Asia. It is bordered by Russia to the north and China to the south, east and west. Although Mongolia does not share a border with Kazakhstan, its western-most point is only from Kazakhstan's eastern tip. Ulan Bator, the capital and largest...

. A second species, V. osmolskae, was named in 2008 for skull material from Inner Mongolia
Inner Mongolia
Inner Mongolia is an autonomous region of the People's Republic of China, located in the northern region of the country. Inner Mongolia shares an international border with the countries of Mongolia and the Russian Federation...

, China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

.

Smaller than other dromaeosaurids like Deinonychus
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

and Achillobator
Achillobator
Achillobator is a genus of dromaeosaurid theropod dinosaur from the late Cretaceous Period of what is now Mongolia, about 90 million years ago. It was probably an active bipedal predator, hunting with the large sickle-shaped claw on the second toe of each hind foot...

, Velociraptor nevertheless shared many of the same anatomical
Anatomy
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

 features.
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Encyclopedia
Velociraptor is a genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of dromaeosaurid
Dromaeosauridae
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'...

 theropod
Theropoda
Theropoda is both a suborder of bipedal saurischian dinosaurs, and a clade consisting of that suborder and its descendants . Dinosaurs belonging to the suborder theropoda were primarily carnivorous, although a number of theropod groups evolved herbivory, omnivory, and insectivory...

 dinosaur
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...

 that existed approximately 75 to 71 million years ago during the later part of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 Period. Two species are currently recognized, although others have been assigned in the past. The type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 is V. mongoliensis; fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

s of this species have been discovered in Mongolia
Mongolia
Mongolia is a landlocked country in East and Central Asia. It is bordered by Russia to the north and China to the south, east and west. Although Mongolia does not share a border with Kazakhstan, its western-most point is only from Kazakhstan's eastern tip. Ulan Bator, the capital and largest...

. A second species, V. osmolskae, was named in 2008 for skull material from Inner Mongolia
Inner Mongolia
Inner Mongolia is an autonomous region of the People's Republic of China, located in the northern region of the country. Inner Mongolia shares an international border with the countries of Mongolia and the Russian Federation...

, China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

.

Smaller than other dromaeosaurids like Deinonychus
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

and Achillobator
Achillobator
Achillobator is a genus of dromaeosaurid theropod dinosaur from the late Cretaceous Period of what is now Mongolia, about 90 million years ago. It was probably an active bipedal predator, hunting with the large sickle-shaped claw on the second toe of each hind foot...

, Velociraptor nevertheless shared many of the same anatomical
Anatomy
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

 features. It was a biped
Biped
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs, or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning "two feet"...

al, feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

ed carnivore
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

 with a long, stiffened tail and an enlarged sickle-shaped claw
Claw
A claw is a curved, pointed appendage, found at the end of a toe or finger in most mammals, birds, and some reptiles. However, the word "claw" is also often used in reference to an invertebrate. Somewhat similar fine hooked structures are found in arthropods such as beetles and spiders, at the end...

 on each hindfoot, which is thought to have been used to kill its prey
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...

. Velociraptor can be distinguished from other dromaeosaurids by its long and low skull
Skull
The skull is a bony structure in the head of many animals that supports the structures of the face and forms a cavity for the brain.The skull is composed of two parts: the cranium and the mandible. A skull without a mandible is only a cranium. Animals that have skulls are called craniates...

, 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
Jurassic Park (film)
Jurassic Park is a 1993 American science fiction adventure film directed by Steven Spielberg. The film is based on the novel of the same name by Michael Crichton. It stars Sam Neill, Laura Dern, Jeff Goldblum, Richard Attenborough, Martin Ferrero, and Bob Peck...

motion picture series. In the films it was shown with anatomical inaccuracies, including being much larger than it was in reality and without feathers. It is also well known to paleontologist
Paleontology
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...

s, with over a dozen described fossil skeletons—the most of any dromaeosaurid. One particularly famous specimen preserves a Velociraptor locked in combat with a Protoceratops
Protoceratops
Protoceratops is a genus of sheep-sized herbivorous ceratopsian dinosaur, from the Upper Cretaceous Period of what is now Mongolia. It was a member of the Protoceratopsidae, a group of early horned dinosaurs...

.

Description



Velociraptor was a mid-sized dromaeosaurid
Dromaeosauridae
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'...

, with adults measuring up to 2.07 m (6.8 ft) long, 0.5 m (1.6 ft) high at the hip, and weighing up to 15 kg (33.1 lb). The skull, which grew up to 25 cm (9.8 in) long, was uniquely up-curved, concave on the upper surface and convex on the lower. The jaws were lined with 26–28 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.

Velociraptor, like other dromaeosaurids, had a large manus
Manus (zoology)
The manus is the zoological term for the distal portion of the fore limb of an animal. In tetrapods, it is the part of the pentadactyl limb that includes the metacarpals and digits . During evolution, it has taken many forms and served a variety of functions...

 ('hand') with three strongly curved claws, which were similar in construction and flexibility to the wing bones of modern bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

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
Pronation
In anatomy, pronation is a rotational movement of the forearm at the radioulnar joint, or of the foot at the subtalar and talocalcaneonavicular joints. For the forearm, when standing in the anatomical position, pronation will move the palm of the hand from an anterior-facing position to a...

 of the wrist and forced the 'hands' to be held with the palmar
Palmar
Palmar may refer to:* a directional anatomical term of location* El Palmar de Troya, a small village near Utrera, in Andalusia, Spain* Palmar Sur , a town the Osa region, Puntarenas province, Costa Rica...

 surface facing inwards (medially), not downwards. The first digit of the foot, as in other theropods, was a small dewclaw
Dewclaw
A dewclaw is a vestigial digit on the foot of many mammals, birds, and reptiles . It commonly grows high on the leg so that in digitigrade species, when the animal is standing, it does not make contact with the ground...

. 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 6.5 cm (2.6 in) long around its outer edge, was most likely a predatory device used to tear into prey, possibly delivering a fatal blow.

Long bony projections (prezygapophyses) on the upper surfaces of the vertebrae, as well as ossified
Ossification
Ossification is the process of laying down new bone material by cells called osteoblasts. It is synonymous with bone tissue formation...

 tendon
Tendon
A tendon is a tough band of fibrous connective tissue that usually connects muscle to bone and is capable of withstanding tension. Tendons are similar to ligaments and fasciae as they are all made of collagen except that ligaments join one bone to another bone, and fasciae connect muscles to other...

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.

In 2007, paleontologists reported the discovery of quill knobs on a well-preserved Velociraptor mongoliensis forearm from Mongolia, confirming the presence of feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

s in this species.

History




During an American Museum of Natural History
American Museum of Natural History
The American Museum of Natural History , located on the Upper West Side of Manhattan in New York City, United States, is one of the largest and most celebrated museums in the world...

 expedition to the Outer Mongolian Gobi Desert
Gobi Desert
The Gobi is a large desert region in Asia. It covers parts of northern and northwestern China, and of southern Mongolia. The desert basins of the Gobi are bounded by the Altai Mountains and the grasslands and steppes of Mongolia on the north, by the Hexi Corridor and Tibetan Plateau to the...

, on 11 August 1923 Peter Kaisen 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
Henry Fairfield Osborn
Henry Fairfield Osborn, Sr. ForMemRS was an American geologist, paleontologist, and eugenicist.-Early life and career:...

 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
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 words velox ('swift') and raptor ('robber' or 'plunderer') and refers to the animal's cursorial
Cursorial
Cursorial is a biological term that describes an organism as being adapted specifically to run. It is typically used in conjunction with an animal's feeding habits or another important adaptation. For example, a horse can be considered a "cursorial grazer", while a wolf may be considered a...

 nature and carnivorous diet. Osborn named the type species V. mongoliensis after its country of origin. 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
Oviraptor
Oviraptor is a genus of small Mongolian theropod dinosaur, first discovered by the paleontologist Roy Chapman Andrews, and first described by Henry Fairfield Osborn, in 1924...

). However, because the name "Ovoraptor" was not published in a scientific journal or accompanied by a formal description, it is considered a nomen nudum
Nomen nudum
The phrase nomen nudum is a Latin term, meaning "naked name", used in taxonomy...

('naked name'), and the name Velociraptor retains priority.

While North American teams were shut out of communist Mongolia during the Cold War
Cold War
The Cold War was the continuing state from roughly 1946 to 1991 of political conflict, military tension, proxy wars, and economic competition between the Communist World—primarily the Soviet Union and its satellite states and allies—and the powers of the Western world, primarily the United States...

, expeditions by Soviet
Soviet Union
The Soviet Union , officially the Union of Soviet Socialist Republics , was a constitutionally socialist state that existed in Eurasia between 1922 and 1991....

 and Polish
Poland
Poland , officially the Republic of Poland , is a country in Central Europe bordered by Germany to the west; the Czech Republic and Slovakia to the south; Ukraine, Belarus and Lithuania to the east; and the Baltic Sea and Kaliningrad Oblast, a Russian exclave, to the north...

 scientists, in collaboration with Mongolian colleagues, recovered several more specimens of Velociraptor. The most famous is part of the legendary "Fighting Dinosaurs" specimen (GIN
Mongolian Academy of Sciences
The Mongolian Academy of Sciences is Mongolia's first centre of modern sciences. It was founded in 1921, when the government established an Institute of Literature and Script, which was later upgraded to Institute of Science. Later, in 1961, it was reconstituted as the Mongolian Academy of Sciences...

 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. 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
New York City
New York is the most populous city in the United States and the center of the New York Metropolitan Area, one of the most populous metropolitan areas in the world. New York exerts a significant impact upon global commerce, finance, media, art, fashion, research, technology, education, and...

 for a temporary exhibition.

Between 1988 and 1990, a joint Chinese
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

-Canadian
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...

 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
Mongolian Academy of Sciences
The Mongolian Academy of Sciences is Mongolia's first centre of modern sciences. It was founded in 1921, when the government established an Institute of Literature and Script, which was later upgraded to Institute of Science. Later, in 1961, it was reconstituted as the Mongolian Academy of Sciences...

, turned up several well-preserved skeletons. 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
Washington Irving
Washington Irving was an American author, essayist, biographer and historian of the early 19th century. He was best known for his short stories "The Legend of Sleepy Hollow" and "Rip Van Winkle", both of which appear in his book The Sketch Book of Geoffrey Crayon, Gent. His historical works...

 character Ichabod Crane
Ichabod Crane
Ichabod Crane is a fictional character in Washington Irving's short story The Legend of Sleepy Hollow, first published in 1820.-Origin:...

). 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.

Maxilla
Maxilla
The maxilla is a fusion of two bones along the palatal fissure that form the upper jaw. This is similar to the mandible , which is also a fusion of two halves at the mental symphysis. Sometimes The maxilla (plural: maxillae) is a fusion of two bones along the palatal fissure that form the upper...

e and a lacrimal
Lacrimal
The term Lacrimal can refer to:*In typography:** A type of Stroke ending *In anatomy** Lacrimal apparatus** Lacrimal artery** Lacrimal bone...

 (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 colleagues named these bones V. osmolskae (for Polish paleontologist Halszka Osmólska
Halszka Osmólska
Halszka Osmólska was a Polish paleontologist who had specialized in Mongolian dinosaurs.She was born in Poznań. A member of the 1965 and 1970 Polish–Mongolian expeditions to the Gobi Desert, she described many finds from these rocks, often with Teresa Maryańska...

) in 2008.

Provenance



All known specimens of Velociraptor mongoliensis were discovered in the Djadochta Formation
Djadochta Formation
The Djadochta Formation is situated in central Asia and dates from the Late Cretaceous Period. Laid down in the early Campanian, possibly starting in the latest Santonian, it is dated somewhat uncertainly at about 84-75 mya...

 (also spelled Djadokhta), in the Mongolian province of Ömnögovi
Ömnögovi Province
Ömnögovi is an aimag of Mongolia, located in the south of the country, in the Gobi Desert. Ömnögovi is Mongolia's largest aimag. The capital is Dalanzadgad....

. Species of Velociraptor, have also been reported from the slightly younger Barun Goyot Formation
Barun Goyot Formation
The Barun Goyot Formation , dating from the Late Cretaceous Period, is located within and is widely represented in the Gobi Desert basin, in the Ömnögovi Province of Mongolia....

 of Mongolia, though these are indeterminate and may belong to a related genus instead. These geologic formation
Geologic formation
A formation or geological formation is the fundamental unit of lithostratigraphy. A formation consists of a certain number of rock strata that have a comparable lithology, facies or other similar properties...

s are estimated to date back to the Campanian
Campanian
The Campanian is, in the ICS' geologic timescale, the fifth of six ages of the Late Cretaceous epoch . The Campanian spans the time from 83.5 ± 0.7 Ma to 70.6 ± 0.6 Ma ...

 stage
Faunal stage
In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convention have the same name, and the same boundaries.Rock...

 (between 83 and 70 million years ago) of the Late Cretaceous
Late Cretaceous
The Late Cretaceous is the younger of two epochs into which the Cretaceous period is divided in the geologic timescale. Rock strata from this epoch form the Upper Cretaceous series...

 Epoch
Epoch (geology)
An epoch is a subdivision of the geologic timescale based on rock layering. In order, the higher subdivisions are periods, eras and eons. We are currently living in the Holocene epoch...

.

V. mongoliensis has been found at many of the most famous and prolific Djadochta localities. The type specimen was discovered at the Flaming Cliffs
Flaming Cliffs
The Flaming Cliffs site, really Bayanzag , is a region of the Gobi Desert in the Ömnögovi Province of Mongolia, in which important fossil finds have been made. It was given this name by American paleontologist Roy Chapman Andrews, who visited in the 1920s. The area is most famous for yielding the...

 site (also known as Bayn Dzak and Shabarakh Usu), while the "Fighting Dinosaurs" were found at the Tugrig locality (also known as Tugrugeen Shireh). The well-known Barun Goyot localities of Khulsan and Khermeen Tsav have also produced remains which may belong to Velociraptor or a related genus. Teeth and partial remains attributed to juvenile V. mongoliensis have also been reported from the Bayan Mandahu Formation
Bayan Mandahu Formation
The Bayan Mandahu Formation is a geological unit of "redbeds" located near the village of Bayan Mandahu in Inner Mongolia, China Asia and dates from the Late Cretaceous Period...

, a prolific site in Inner Mongolia, China that is contemporaneous with the Djadochta Formation. However, these fossils had not been prepared or studied as of 2008. A partial adult skull from the Bayan Mandahu Formation has been assigned to a distinct species, Velociraptor osmolskae.

Paleoecology


All of the fossil sites that have yielded Velociraptor remains preserve an arid environment with fields of sand dune
Dune
In physical geography, a dune is a hill of sand built by wind. Dunes occur in different forms and sizes, formed by interaction with the wind. Most kinds of dunes are longer on the windward side where the sand is pushed up the dune and have a shorter "slip face" in the lee of the wind...

s and only intermittent stream
Stream
A stream is a body of water with a current, confined within a bed and stream banks. Depending on its locale or certain characteristics, a stream may be referred to as a branch, brook, beck, burn, creek, "crick", gill , kill, lick, rill, river, syke, bayou, rivulet, streamage, wash, run or...

s, although the younger Barun Goyot environment seems to have been slightly wetter than the older Djadochta. The posture of some complete fossils, as well as the mode of preservation most show within structureless sandstone
Sandstone
Sandstone is a sedimentary rock composed mainly of sand-sized minerals or rock grains.Most sandstone is composed of quartz and/or feldspar because these are the most common minerals in the Earth's crust. Like sand, sandstone may be any colour, but the most common colours are tan, brown, yellow,...

 deposits, may show that a number of specimens were buried alive during sandstorm
Dust storm
A dust / sand storm is a meteorological phenomenon common in arid and semi-arid regions. Dust storms arise when a gust front or other strong wind blows loose sand and dirt from a dry surface. Particles are transported by saltation and suspension, causing soil to move from one place and deposition...

 events common to the three environments.

Many of the same genera were present across these formations, though they varied at the species level. For example, the Djadochta was inhabited by Velociraptor mongoliensis, Protoceratops andrewsi, and Pinacosaurus grangeri, while the Bayan Mandahu was home to Velociraptor osmolskae, Protoceratops hellenikorhinus, and Pinacosaurus mephistocephalus. 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.

Other dinosaurs known from the same locality as V. mongoliensis include the troodontid Saurornithoides mongoliensis, the oviraptorid Oviraptor philoceratops, and the dromaeosaurid Mahakala omnogovae. V. osmolskae lived alongside the ceratopsia
Ceratopsia
Ceratopsia or Ceratopia is a group of herbivorous, beaked dinosaurs which thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. The earliest known ceratopsian, Yinlong downsi, lived between 161.2 and 155.7...

n species Magnirostris dodsoni, as well as the oviraptorid Machairasaurus leptonychus and the dromaeosaurid Linheraptor exquisitus.

Classification



Velociraptor is a member of the subfamily Velociraptorinae
Velociraptorinae
Velociraptorinae is a subfamily of the theropod group Dromaeosauridae. Velociraptorine remains are found mostly in northern hemisphere. In Late Cretaceous, they are mostly present in Mongolia, Romania and China,and in early, in USA. The earliest velociraptorines are probably Nuthetes from the...

, a derived
Derived
In phylogenetics, a derived trait is a trait that is present in an organism, but was absent in the last common ancestor of the group being considered. This may also refer to structures that are not present in an organism, but were present in its ancestors, i.e. traits that have undergone secondary...

 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
Dromaeosaurus
Dromaeosaurus was a genus of theropod dinosaur which lived during the Late Cretaceous period , sometime between 76.5 and 74.8 million years ago, in the western United States and Alberta, Canada. The name means 'running lizard'....

." Dromaeosaurid classification is highly variable. Originally, the subfamily Velociraptorinae was erected solely to contain Velociraptor. Other analyses have included other genera, usually Deinonychus and Saurornitholestes
Saurornitholestes
Saurornitholestes is a genus of carnivorous dromaeosaurid theropod dinosaur from the late Cretaceous of Alberta, Montana and New Mexico....

. A recent cladistic analysis indicated a monophyletic Velociraptorinae containing Velociraptor, Deinonychus, Tsaagan, and a closely related (but uncertainly positioned) Saurornitholestes.

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. However, the only currently recognized species of Velociraptor are V. mongoliensis and V. osmolskae.

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
Megalosaurus
Megalosaurus is a genus of large meat-eating theropod dinosaurs of the Middle Jurassic period of Europe...

, 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
Archaeopterygidae
The Archaeopterygidae is a group of maniraptoran dinosaurs that lived during the late Jurassic and period.-Taxonomy:The order Archaeopterygiformes was coined by Max Fürbringer in 1888 to contain the single family Archaeopterygidae and genus Archaeopteryx...

 by at least one author (which would, in effect, make Velociraptor a flightless bird).

Predatory behavior



The "Fighting Dinosaurs" specimen, found in 1971, preserves a Velociraptor mongoliensis and Protoceratops andrewsi
Protoceratops
Protoceratops is a genus of sheep-sized herbivorous ceratopsian dinosaur, from the Upper Cretaceous Period of what is now Mongolia. It was a member of the Protoceratopsidae, a group of early horned dinosaurs...

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 sandstorm
Dust storm
A dust / sand storm is a meteorological phenomenon common in arid and semi-arid regions. Dust storms arise when a gust front or other strong wind blows loose sand and dirt from a dry surface. Particles are transported by saltation and suspension, causing soil to move from one place and deposition...

. 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 scavenging
Scavenger
Scavenging is both a carnivorous and herbivorous feeding behavior in which individual scavengers search out dead animal and dead plant biomass on which to feed. The eating of carrion from the same species is referred to as cannibalism. Scavengers play an important role in the ecosystem by...

 by other animals. Comparisons between the scleral ring
Sclerotic ring
Sclerotic rings are rings of bone found in the eyes of several groups of vertebrate animals, except for mammals and crocodilians. They can be made up of single bones or small bones together. They are believed to have a role in supporting the eye, especially in animals whose eyes are not spherical,...

s of Velociraptor, Protoceratops, and modern birds and reptiles indicates that Velociraptor may have been nocturnal, while Protoceratops may have been cathemeral
Cathemeral
A cathemeral organism is one that has sporadic and random intervals of activity during the day or night in which food is acquired, socializing with other organisms occurs, and any other activities necessary for livelihood are performed...

, 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
Jugular vein
The jugular veins are veins that bring deoxygenated blood from the head back to the heart via the superior vena cava.-Internal and external:There are two sets of jugular veins: external and internal....

, carotid artery
Carotid artery
Carotid artery can refer to:* Common carotid artery* External carotid artery* Internal carotid artery...

, or trachea
Vertebrate trachea
In tetrapod anatomy the trachea, or windpipe, is a tube that connects the pharynx or larynx to the lungs, allowing the passage of air. It is lined with pseudostratified ciliated columnar epithelium cells with goblet cells that produce mucus...

 (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
Skin
-Dermis:The dermis is the layer of skin beneath the epidermis that consists of connective tissue and cushions the body from stress and strain. The dermis is tightly connected to the epidermis by a basement membrane. It also harbors many Mechanoreceptors that provide the sense of touch and heat...

 and muscle
Muscle
Muscle is a contractile tissue of animals and is derived from the mesodermal layer of embryonic germ cells. Muscle cells contain contractile filaments that move past each other and change the size of the cell. They are classified as skeletal, cardiac, or smooth muscles. Their function is to...

 of large prey species would have been difficult to slash without a specialized cutting surface. The slashing hypothesis
Hypothesis
A hypothesis is a proposed explanation for a phenomenon. The term derives from the Greek, ὑποτιθέναι – hypotithenai meaning "to put under" or "to suppose". For a hypothesis to be put forward as a scientific hypothesis, the scientific method requires that one can test it...

 was tested during a 2005 BBC
BBC
The British Broadcasting Corporation is a British public service broadcaster. Its headquarters is at Broadcasting House in the City of Westminster, London. It is the largest broadcaster in the world, with about 23,000 staff...

 documentary, The Truth About Killer Dinosaurs
The Truth About Killer Dinosaurs
The Truth About Killer Dinosaurs is a two-part BBC documentary, presented by Bill Oddie, in which a group of scientists test out the strength of dinosaur weaponry using biomechanics. The first episode determines the winner of a battle between Tyrannosaurus and Triceratops, and second compares the...

. The producers of the program created an artificial Velociraptor leg with a sickle claw and used a pork belly
Pork belly
Pork belly is a boneless cut of fatty meat derived from the belly of a pig. Pork belly is popular in Asian cuisine, and forms a part of many traditional European dishes such as the Alsatian Choucroute garnie, the Swiss Berner Platte, and the German Schlachtplatte...

 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
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

, 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
Tenontosaurus
Tenontosaurus is a genus of medium- to large-sized ornithopod dinosaur. The genus is known from the late Aptian to Albian ages of the middle Cretaceous period sediments of western North America, dating between 115 to 108 million years ago...

, which has been seen as evidence of cooperative hunting. 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. Although many isolated fossils of Velociraptor have been found in Mongolia, none were closely associated with any other individuals. Therefore, while Velociraptor is commonly depicted as a pack hunter
Pack hunter
A pack hunter is a predator belonging to the animal kingdom, which has evolved to hunt its prey by working together with other members of its species. Normally, such animals are closely related. The most commonly known pack hunter is the Gray Wolf, the ancestor of all breeds of domesticated dogs...

, as in Jurassic Park, there is only limited fossil evidence to support this theory for dromaeosaurids in general, and none specific to Velociraptor itself. The pack hunting theory was based on a discovery of several specimens of Deinonychus
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

found around the remains of a Tenontosaurus
Tenontosaurus
Tenontosaurus is a genus of medium- to large-sized ornithopod dinosaur. The genus is known from the late Aptian to Albian ages of the middle Cretaceous period sediments of western North America, dating between 115 to 108 million years ago...

. No other group of dromaeosaurids has been found in close association.

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.

Metabolism


Velociraptor was probably warm-blooded
Warm-blooded
The term warm-blooded is a colloquial term to describe animal species which have a relatively higher blood temperature, and maintain thermal homeostasis primarily through internal metabolic processes...

 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
Metabolism
Metabolism is the set of chemical reactions that happen in the cells of living organisms to sustain life. These processes allow organisms to grow and reproduce, maintain their structures, and respond to their environments. Metabolism is usually divided into two categories...

, compared with most modern warm-blooded mammals and birds. The kiwi
Kiwi
Kiwi are flightless birds endemic to New Zealand, in the genus Apteryx and family Apterygidae.At around the size of a domestic chicken, kiwi are by far the smallest living ratites and lay the largest egg in relation to their body size of any species of bird in the world...

 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.

Feathers



Fossils of dromaeosaurids more primitive than Velociraptor are known to have had feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

s covering their bodies, and fully developed, feathered wings. 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 quill knobs on the forearm of a Velociraptor found in Mongolia. 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,
Co-author Mark Norell, Curator-in-Charge of fossil reptiles, amphibians and birds at the American Museum of Natural History
American Museum of Natural History
The American Museum of Natural History , located on the Upper West Side of Manhattan in New York City, United States, is one of the largest and most celebrated museums in the world...

, also weighed in on the discovery, saying:
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
Flamingo
Flamingos or flamingoes are gregarious wading birds in the genus Phoenicopterus , the only genus in the family Phoenicopteridae...

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
Archaeopteryx
Archaeopteryx , sometimes referred to by its German name Urvogel , is a genus of theropod dinosaur that is closely related to birds. The name derives from the Ancient Greek meaning "ancient", and , meaning "feather" or "wing"...

, 18 in Microraptor
Microraptor
Microraptor is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China...

, and 10 in Rahonavis
Rahonavis
Rahonavis is a genus of bird-like dinosaur from the Late Cretaceous of what is now northwestern Madagascar. It is known from a partial skeleton found in Maevarano Formation rocks at a quarry near Berivotra, Mahajanga Province...

. 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.

Pathology



One Velociratoptor mongoliensis skull bears two parallel rows of small punctures that match the spacing and size of Velociraptor teeth. Scientists believe that the wound was likely inflicted by another Velociraptor during a fight. Further, because the fossil bone shows no sign of healing near the bite wounds, the injury probably killed it.

In popular culture




Velociraptor are well known for their role as vicious and cunning killers in the 1990 novel Jurassic Park by Michael Crichton
Michael Crichton
John Michael Crichton , best known as Michael Crichton, was an American best-selling author, producer, director, and screenwriter, best known for his work in the science fiction, medical fiction, and thriller genres. His books have sold over 200 million copies worldwide, and many have been adapted...

 and its 1993 film adaptation
Jurassic Park (film)
Jurassic Park is a 1993 American science fiction adventure film directed by Steven Spielberg. The film is based on the novel of the same name by Michael Crichton. It stars Sam Neill, Laura Dern, Jeff Goldblum, Richard Attenborough, Martin Ferrero, and Bob Peck...

, directed by Steven Spielberg
Steven Spielberg
Steven Allan Spielberg KBE is an American film director, screenwriter, producer, video game designer, and studio entrepreneur. In a career of more than four decades, Spielberg's films have covered many themes and genres. Spielberg's early science-fiction and adventure films were seen as an...

, in which they served as the main antagonists. The "raptors" portrayed in Jurassic Park were modeled after a larger relative, Deinonychus
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

, which Gregory Paul at the time called Velociraptor antirrhopus. The paleontologists in the film and the novel excavate a so-called Velociraptor skeleton in Montana
Montana
Montana is a state in the Western United States. The western third of Montana contains numerous mountain ranges. Smaller, "island ranges" are found in the central third of the state, for a total of 77 named ranges of the Rocky Mountains. This geographical fact is reflected in the state's name,...

, 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
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

, though the "raptors" in the novel are referred to as V. mongoliensis.

The filmmakers also had the size of the film's Velociraptor increased for dramatic reasons, and changed the shape of the snout. 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. Another noticeable difference is the lack of feathers for the novel and film version of Velociraptor. In real life, Velociraptor, like many other maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...

n theropods, was covered in feathers. Jurassic Park
Jurassic Park (film)
Jurassic Park is a 1993 American science fiction adventure film directed by Steven Spielberg. The film is based on the novel of the same name by Michael Crichton. It stars Sam Neill, Laura Dern, Jeff Goldblum, Richard Attenborough, Martin Ferrero, and Bob Peck...

and The Lost World: Jurassic Park
The Lost World: Jurassic Park
The Lost World: Jurassic Park is a 1997 science fiction thriller film, directed by Steven Spielberg. The film was produced by Bonnie Curtis, Kathleen Kennedy, Gerald R. Molen and Colin Wilson...

, however, were released before this was known. In Jurassic Park III
Jurassic Park III
Jurassic Park III is a 2001 American science fiction film and the third of the Jurassic Park franchise. It is the only film in the series that is neither directed by Steven Spielberg nor based on a book by Michael Crichton, though numerous scenes in the movie were taken from Crichton's two books,...

, 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. Also in Jurassic Park III, Dr. Alan Grant, played by Sam Neill
Sam Neill
Nigel John Dermot "Sam" Neill, DCNZM, OBE is a New Zealand actor. He is well known for his starring role as paleontologist Dr Alan Grant in Jurassic Park and Jurassic Park III....

, states that Velociraptor were smarter than dolphins, whales and some primates. Given only fossil evidence, this is impossible to ascertain and is purely conjecture.

Due to the success of many Jurassic Park-related products, Velociraptor has become a ubiquitous representation of dinosaurs in popular culture. It has been featured in numerous toy lines
Toy
A toy is any object that can be used for play. Toys are associated commonly with children and pets. Playing with toys is often thought to be an enjoyable means of training the young for life in human society. Different materials are used to make toys enjoyable and cuddly to both young and old...

, animated films, video games and television series for children, along with several television documentaries
Documentary film
Documentary films constitute a broad category of nonfictional motion pictures intended to document some aspect of reality, primarily for the purposes of instruction or maintaining a historical record...

. In 1995, the city of Toronto
Toronto
Toronto is the provincial capital of Ontario and the largest city in Canada. It is located in Southern Ontario on the northwestern shore of Lake Ontario. A relatively modern city, Toronto's history dates back to the late-18th century, when its land was first purchased by the British monarchy from...

 was awarded a National Basketball Association
National Basketball Association
The National Basketball Association is the pre-eminent men's professional basketball league in North America. It consists of thirty franchised member clubs, of which twenty-nine are located in the United States and one in Canada...

 expansion team
Expansion team
An expansion team is a brand new team in a sports league. The term is most commonly used in reference to the North American major professional sports leagues, but is applied to sports leagues worldwide that use a closed franchise system of league membership. The term comes from the expansion of the...

, which was named the Toronto Raptors
Toronto Raptors
The Toronto Raptors are a professional basketball team based in Toronto, Ontario, Canada. They are part of the Atlantic Division of the Eastern Conference in the National Basketball Association . The team was established in 1995, along with the Vancouver Grizzlies, as part of the NBA's re-expansion...

.

External links