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Oviraptor

Oviraptor is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous period. The first remains were collected from the Djadokhta Formation of Mongolia in 1923 during a paleontological expedition led by Roy Chapman Andrews, and in the following year the genus and type species Oviraptor philoceratops were named by Henry Fairfield Osborn. The genus name refers to the initial thought of egg-stealing habits, and the specific name was intended to reinforce this view indicating a preference over ceratopsian eggs. Despite the fact that numerous specimens have been referred to the genus, Oviraptor is only known from a single partial skeleton regarded as the holotype, as well as a nest of about fifteen eggs and several small fragments from a juvenile.

History of discovery
of Mongolia in the Gobi Desert. The holotype remains of Oviraptor were found in this highly fossiliferous locality of the Djadokhta Formation The first remains of Oviraptor were discovered on reddish sandstones of the Late Cretaceous Djadokhta Formation of Mongolia, at the Bayn Dzak locality (also known as Flaming Cliffs), during the Third Central Asiatic expedition in 1923. This expedition was led by the North American naturalist Roy Chapman Andrews and ended in the discovery of three new-to-science theropod fossil remains—including those of Oviraptor. These were formally described by the North American paleontologist Henry Fairfield Osborn in 1924, who in the basis of the new material, named the genera Oviraptor, Saurornithoides and Velociraptor. The particular genus Oviraptor was erected with the type species O. philoceratops based on the holotype AMNH 6517, a partial individual lacking the back of the skeleton but including a badly crushed skull, partial cervical and dorsal vertebrae, pectoral elements including the furcula with the left arm and partial hands, the left ilium and some ribs. Accordingly, this specimen was found lying over a nest of approximately 15 eggs—a nest that has been catalogued as AMNH 6508—with the skull separated from the eggs by only of sediment. Given the close proximity of both specimens, Osborn interpreted Oviraptor as a dinosaur with egg-eating habits, and explained that the generic name, Oviraptor, is Latin for "egg seizer" or "egg thief", due to the association of the fossils. The specific name, philoceratops, is intended as "fondness for ceratopsian eggs" which is also given as a result of the initial thought of the nest pertaining to Protoceratops or another ceratopsian. However, Osborn suggested that the name Oviraptor could reflect an incorrect perception of this dinosaur. Furthermore, Osborn found Oviraptor to be similar to the unrelated—at the time, however, considered related—fast-running ornithomimids based on the toothless jaws, and assigned Oviraptor to the Ornithomimidae. Osborn had previously reported the taxon as "Fenestrosaurus philoceratops", but this was later discredited. After Osborn's initial description of Oviraptor, the egg nest associated with the holotype was accepted to have belonged to Protoceratops, and oviraptorids were largely considered to have been egg-eating theropods. Nevertheless, in the 1990s, the discovery of numerous nesting and nestling oviraptorid specimens proved that Osborn was correct in his caution regarding the name of Oviraptor. These findings showed that oviraptorids brooded and protected their nests by crouching on them. This new line of evidence showed that the nest associated with the holotype of Oviraptor belonged to it and the specimen was actually brooding the eggs at the time of death, not preying on them. Referred specimens After the naming of Oviraptoridae in 1976, Barsbold referred six additional specimens to Oviraptor, including two particular specimens under the number MPC-D 100/20 and 100/21. Most of the other specimens are also unlikely to belong to Oviraptor itself, and they have been assigned to other oviraptorids. A partial individual also with eggs from the Bayan Mandahu Formation of China was referred in 1996 by Dong Zhiming and Philip J. Currie, the specimen IVPP V9608. In 1981, Barsbold referred the specimen MPC-D 100/42 to Oviraptor, a very well-preserved and rather complete individual from the Djadokhta Formation. Since the known elements of Oviraptor were so fragmentary compared to other members, MPC-D 100/42 became the prime reference/depiction of this taxon being prominently labelled as Oviraptor philoceratops in scientific literature. In 1986, Barsbold described a second species of Oviraptor, "O. mongoliensis", based on specimen MPC-D 100/32a which hails from the Nemegt Formation. This was amended by the Polish paleontologist Halszka Osmólska and team in 2004 by formally naming the taxon Rinchenia mongoliensis. The North American paleontologist Mark A. Norell and colleagues in 2018 reported a new specimen of Oviraptor: AMNH 33092, which is composed of a tibia and two metatarsals of a nestling or very small juvenile. AMNH 33092 was found in association with the holotype and it was likely part of the nest. Oviraptor is now known from the holotype with associated eggs, and a juvenile/nestling. ==Description==
Description
The holotype specimen has been estimated at in length with a weight ranging from . Though the holotype largely lacks the posterior region of the skeleton, it is likely that Oviraptor had two well-developed hindlimbs that ended in three functional toes with the first one being vestigial, as well as a relatively reduced tail. As evidenced in related oviraptorids, the arms were covered by elongated feathers, and the tail ended in a pygostyle, which is known to support a fan of feathers. The scapulocoracoid is fused in the holotype, however, the coracoid is badly damaged. The scapula is slightly bowed and measures in length. Oviraptor had a relatively elongated arm composed of the humerus, radius, ulna, and manus. The regarded juvenile Oviraptor AMNH 33092 preserves hindlimb material, comprising a right tibia with metatarsals III and IV. Its tibia is long, indicating a substantially smaller individual than the holotype. The nest AMNH 6508 preserves elongatoolithid eggs, with each egg being long (some are incomplete). Nevertheless, there is the possibility that taphonomical crushing may have compressed them by up to . ==Classification==
Classification
Oviraptor was originally allied with the ornithomimids by Osborn due to its toothless beak. Osborn also found similarities with Chirostenotes, which is still considered a close relative of Oviraptor. {{clade| style=font-size: 100%; line-height:100% ==Paleobiology==
Paleobiology
Feeding When first described in 1924 by Osborn, Oviraptor was originally presumed to have been ovivorous—an organism that has an egg-based diet—based on the association of the holotype with a nest thought to belong to Protoceratops. In a 1990 conference abstract, David K. Smith presented an osteological reevaluation of Oviraptor where he rejected the statements made by Barsbold. He found no evidence indicating a forelimb specialized in aquatic locomotion, and the jaws, rather than preserve a crushing mechanism, preserve shearing surfaces. As the skull is toothless, lightly built and lacks several strong muscle insertion areas, Smith suggested that leaves may have been an important part in the diet of Oviraptor. However, in 1995, Norell and colleagues reported the fragmented remains of a lizard in the body cavity of the holotype specimen, suggesting that Oviraptor was partially carnivorous. of several oviraptorosaur genera In 2008, Stig Olav K. Jansen compared the skull of several oviraptorid species to those of birds and turtles to investigate which properties can predict a rhamphotheca. He found the lower jaws of oviraptorids to be very similar to those of parrots, and the upper jaws to be more similar to those of turtles. Based on these observations, Jansen suggested that oviraptorids were omnivorous as the sharply developed rhamphotheca together with the prominent forelimbs would have been adapted to catch and tear small prey. Moreover, the pointed projections of the palate would have contributed in holding prey. Jansen pointed out that a fully herbivorous diet in oviraptorids seems unlikely as they lacked flat and wide tomia (cutting edges of the mandibles) to chew, and were unable to move the lower jaws sideways. However, he considered the lower jaws strong enough to have at least crushed elements like eggs, nuts or other hard seeds. Longrich and colleagues in 2010 also rejected a durophagous (animals that practise shell-crushing) hypothesis, given that such animals typically develop teeth with broad crushing surfaces. The pointed shape of the dentary bones in the lower jaws suggests that oviraptorids had a sharp-edged rhamphotheca used for shearing food instead. The symphyseal (bone union) region at the front of the dentary may have given some ability for crushing, but as this was a relatively small area, it was probably not the main function of the jaws. Another argument against them having been eaters of mollusks is the fact that most oviraptorids have been found in sediments that are interpreted to represent mostly arid or semi-arid environments, such as Oviraptor in the Djadokhta Formation. The team also found that oviraptorids and dicynodonts share cranial features such as short, deep, and toothless mandibles; elongated dentary symphyses; elongated mandibular openings; and a pointed palate. Modern animals with jaws that resemble those of oviraptorids include parrots and tortoises; the latter group also has tooth-like projections on the palatal region. Longrich and colleagues concluded that due to the similarities between oviraptorids and herbivorous animals, the bulk of their diet would most likely have been formed by plant material. The jaws of oviraptorids may have been specialised for processing food, such as xerophytic vegetation−a vegetation that is adapted for environments with little water—that would have grown in their arid environments, but this is not possible to demonstrate, as little is known about the paleoflora of the Gobi Desert. Reproduction Since the description of the embryonic Citipati specimen in 1994, oviraptorids became more understood: instead of having been egg-eating animals, they actually brooded and cared for the nests. This specimen showed that the holotype of Oviraptor was likely a sexually mature individual that perished incubating the associated nest with eggs. Thomas P. Hopp and Mark J. Orsen in 2004 analyzed the brooding behavior of extinct and extant dinosaur species, including oviraptorids, in order to evaluate the reason for the elongation and development of wing and tail feathers. Given that the most complete oviraptorid nesting specimen—at the time, the 1995 Citipati nesting specimen—was found in a very avian-like posture, with the forelimbs in a near-folded posture and the pectoral region, belly, and feet in contact with the eggs, Hopp and Orsen indicated that long pennaceous feathers and a feather covering were most likely present in life. The "wings" and tail of oviraptorids would have granted protection for the eggs and hatchlings against climate factors like the sunlight, wind, and rainfalls. However, the arms of this specimen were not extremely folded as in some modern birds, instead, they are more extended resembling the style of large flightless birds like the ostrich. The extended position of the arm is also similar to the brooding behavior of this bird, which is known to nest in large clutches like oviraptorids. Based on the forelimb position of nesting oviraptorids, Hopp and Orsen proposed brooding as the ancestral reason behind wing and tail feather elongation, as there was a greater need to provide optimal protection for eggs and juveniles. of a nesting Oviraptor. Oviraptorids may have sat on their nests during incubation In 2005, Tamaki Sato and team reported an unusual oviraptorid specimen from the Nanxiong Formation. This new specimen was found preserving mainly the pelvic region with two eggs inside and thereby indicating a female. The size and position of the eggs suggest that oviraptorids retained two functional oviducts, but had reduced the number of eggs ovulated to one per oviduct. David J. Varricchio and colleagues in 2008 found that the relatively large egg clutch-size of oviraptorids and troodontids is most similar to those of modern birds that practice polygamous mating and extensive male parental care, such as ratite birds, suggesting similar habits. This reproductive system is most likely to represent the ancestral condition for modern birds, with biparental care (where both parents participate) being a later development. In 2014, W. Scott Persons and colleagues suggested that oviraptorosaurs were secondarily flightless and several of the traits in their tails may indicate a propensity for display behaviour, such as courtship display. The tail of several oviraptorosaurs and oviraptorids ended in pygostyles, a bony structure at the end of the tail that, at least in modern birds, is used to support a feather fan. Furthermore, the tail was notably muscular and had a pronounced flexibility, which may have aided in courtship movements. es from the Nanxiong Formation In 2018, Tzu-Ruei Yang and colleagues identified cuticle layers on several egg-shells of maniraptoran dinosaurs including those of oviraptorids. These particular layers are composed of proteins, polysaccharides and pigments, but mainly of lipids and hydroxyapatite. In modern birds they serve to protect the eggs from dehydration and invasion of microorganisms. As most oviraptorid specimens have been found in formations of caliche-based sedimentation, Yang and colleagues suggested that the cuticle-coated eggs would have been a reproductive strategy adapted for enhancing their hatching success in such arid climates and environments. In 2019 Yang and colleagues re-evaluated the hypothesis of thermoregulatory contact incubation using complete oviraptorid nests from the Nanxiong Formation, and provided a detailed reconstruction of the architecture of the oviraptorid clutch. They noted that adult oviraptorid specimens in association with nest were not necessarily incubating the eggs as they could represent a female in the process of laying eggs, and the multi-ring clutch prevented sufficient heat transfer from the parent to the inner rings of eggs. An average oviraptorid nest was built as a gently inclined mound with a highly organized architecture: the eggs were likely pigmented and arranged in pairs with each pair arranged in three to four elliptical rings. As the parent was likely operating from the nest center, this region was devoid of eggs. Yang and colleagues concluded the oviraptorid nesting style was so unique that they lack modern analogs, therefore, using oviraptorid reproduction may not be the best example to inform about the evolution of bird reproductive strategies. However, the team was unable to determinate if the juvenile Oviraptor AMNH 33092 had hatched from the nest associated with the holotype. ==Paleoenvironment==
Paleoenvironment
Oviraptor is known from the Bayn Dzak locality of the Djadokhta Formation in Mongolia, a formation that dates back to the Late Cretaceous about 71 million to 75 million years ago. The paleoenvironment of the Djadokhta Formation is interpreted as having a semiarid climate, with sand dune and alluvial settings similar to the modern Gobi Desert. The semiarid steppe landscape was drained by intermittent streams and was sometimes affected by dust and sandstorms, and moisture was seasonal. Though this formation is largely considered to preserved highly arid environments, several short-lived water bodies have been reported from the Ukhaa Tolgod locality, based on fluvial sedimentation. Furthermore, it is thought that later in the Campanian age and into the Maastrichtian, the climate would shift to the more humid fluvial environment seen in the Nemegt Formation. Further dinosaur fauna from this member includes that of the Ukhaa Tolgod locality, composed of Apsaravis, Byronosaurus, Citipati, Gobipteryx, Khaan, Kol, Shuuvuia, Tsaagan, Taphonomy The pose of the holotype of Oviraptor along with the association of eggs, suggest that it was trapped over the nest during a sandstorm, and burial was relatively rapid given that the body had no opportunity to become fully disarticulated or scavenged by predators. Among elements, the skull have become particularly flattened and distorted during the fossilization process. ==See also==
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