Oviraptoridae is a group of bird-like, herbivorous and omnivorous maniraptoran dinosaurs. Oviraptorids are characterized by their toothless, parrot-like beaks and, in some cases, elaborate crests. They were generally small, measuring between one and two metres long in most cases, though some possible oviraptorids were enormous. Oviraptorids are currently known only from the Late Cretaceous of Asia, with the most well-known species and complete specimens found only in the Gobi Desert of Mongolia and northwestern China.
|Fossil skeleton of Khaan mckennai|
The most characteristic feature of this group is the skull structure. Oviraptorids had short snouts and very deep mandibles. Some taxa (Oviraptor, Citipati, Rinchenia) had a midline crest on top of the skull, resembling that of a cassowary. Other distinguishing characteristics include a bony spike intruding on the mandibular fenestra, nostrils placed very high and far back on the snout, an extremely thin bony bar beneath the eye, and highly pneumatized skull bones. Like their relatives the caenagnathids, the jaws were edentulous (with no teeth), having instead two small bony projections on the roof of the mouth.
The classification of the oviraptorids has been controversial. Most studies divide oviraptorosaurs into two primary sub-groups, the Caenagnathidae and the Oviraptoridae. However, some phylogenetic studies have suggested that many traditional members of the Caenagnathidae may more closely related to the crested oviraptorids. Because of this, and the fact that at least one study found that Caenagnathus itself may not have been part of the 'caenagnathid' group, Tom Holtz (2010) placed that group close to the Oviraptoridae and termed it the Elmisauridae, though this idea has not gained consensus among other researchers.
The Oviraptoridae itself is traditionally divided into two "subfamilies": the small, short-armed, and mainly crestless subfamily "Ingeniinae" (the name was preoccupied by an invertebrate group), defined as a clade (Ajancingenia + Conchoraptor), and the larger, crested, long-armed Oviraptorinae (Oviraptor + Citipati). Some phylogenetic studies have shown that Oviraptor is the most primitive known oviraptorid, thus making Citipati a closer relative of the "ingeniines" and this traditional division into crestless and crested forms artificial.
Other possible oviraptorids include Nomingia gobiensis, Gigantoraptor erlianensis, Jiangxisaurus ganzhouensis and Shixinggia oblita. All four have been suggested to be oviraptorids, caenagnathids, or more primitive than either group. The below cladogram was found in the description of the genus Anzu.
Almost all oviraptorids come from desert deposits of the Gobi Desert. Even in the late Cretaceous period, much of this area was desert, or at least very dry, habitat. In many of the localities where they are found, oviraptorids are among the most abundant dinosaurs present, second only to ankylosaurs and protoceratopsids. This is consistent with the idea that they were primarily herbivores, which tend to far outnumber carnivores in a given environment. Oviraptorids appear to have been far more abundant in arid habitats dominated mainly by small dinosaurs (such as those preserved in the Barun Goyot and Djadochta Formations) than in wetter ecosystems where large dinosaurs are common (such as the Nemegt Formation). The same pattern holds true for protoceratopsids, indicating that both groups preferred dry, desert-like habitat, and fed mainly on the types of tough, low-growing plant life that grows in arid climates.
The diet of oviraptorids is not fully understood. Though some appear to have been at least partially carnivorous, they were probably primarily herbivorous.
Originally, oviraptorids were thought to be specialized egg raiders, based on a Mongolian find showing Oviraptor on top of a nest erroneously attributed to the ceratopsian dinosaur Protoceratops. However, discoveries in the 1990s, including Citipati specimens clearly brooding (rather than preying on) the same types of nests, and a Citipati embryo inside the same type of egg preserved in these nests, showed that the "specialized egg thief" idea was incorrect. Still, some scientists have suggested that oviraptorids may have fed on shelled food items like eggs or shellfish. However, animals specialized for eating shelled food typically have broad, crushing beaks or teeth. In contrast, the jaws of oviraptorids had thin, sharp edges probably supporting shearing beaks, ill-suited for cracking shells. Among other known animals, the beaks of oviraptorids most closely resemble those of herbivorous dicynodont synapsids, which are usually considered herbivorous. Their beaks also share similarities with the beaks of herbivorous parrots and tortoises.
Evidence of partial carnivory among some oviraptorines comes from a lizard skeleton preserved in the body cavity of the type specimen of Oviraptor and two hatchling Byronosaurus skulls found in a Citipati nest. Some scientists have also suggested that some oviraptorids (especially the small-handed, weak-clawed "ingeniines") fed mainly on plant material.
Although fossilized dinosaur eggs are generally rare, oviraptorid eggs are relatively well known. Several oviraptorid nests, eggs, and embryos are known, mostly uncovered in the Gobi Desert. Some specimens of Oviraptor philoceratops, Citipati osmolskae, Nemegtomaia and cf. Machairasaurus have been found in brooding positions in association with nests. All of the nesting specimens are situated on top of egg clutches, with their limbs spread symmetrically on each side of the nest, front limbs covering the nest perimeter. This brooding posture is found today only in birds and supports a behavioral link between birds and theropod dinosaurs.
Oviraptorid eggs are shaped like elongated ovals (elongatoolithid) and resemble the eggs of ratite birds (such as ostriches) in texture and shell structure. In the nest, eggs are typically found in pairs and arranged in concentric circles of up to three layers, with complete clutches consisting of as many of 22 eggs in some species. The eggs of Citipati are the largest known definitive oviraptorid eggs, at 18 cm. In contrast, eggs associated with Oviraptor are only up to 14 cm long.
The first oviraptorid eggs (of the genus Oviraptor, which mean "Egg thief") were found in close proximity to the remains of the ceratopsian dinosaur Protoceratops and it was assumed that the oviraptorids were preying upon the eggs of the ceratopsians. It was not until 1993, when a Citipati embryo was discovered inside an egg of the type assigned to Protoceratops, that the error was corrected. Norell et al., who recognized the embryo as oviraptorid, assigned it to the genus Citipati. The egg containing the embryo was smaller than most known Citipati eggs at only 12 cm, though it was partially eroded and broken into three pieces, making an accurate estimate of its original size difficult. The embryo-bearing egg was otherwise identical to other oviraptorid eggs in shell structure and was found in an isolated nest, again arranged in a circular pattern.
An oviraptorosaurian specimen from China described in 2005 was found to have two unlaid eggs within the pelvic canal. This suggests that, unlike modern crocodilians, oviraptorosaurs did not produce and lay many eggs at the same time. Rather, the eggs were produced within the reproductive organs in pairs, and laid two at a time, with the mother positioned in the center of the nest and rotating in a circle as each pair was laid. This behavior is supported by the fact that the eggs oval shape, with the more narrow end pointing backward from the birth canal, matching their orientation toward the center of the nest after being laid.
The presence of two shelled eggs within the birth canal shows that oviraptorosaurs were intermediate between the reproductive biology of crocodilians and modern birds. Like crocodilians, they had two oviducts. However, crocodilians produce multiple shelled eggs per oviduct at a time, whereas oviraptorosaurs, like birds, produced only one egg per oviduct at a time.
In 2017, paleontologists discovered colored pigments in some fossilized oviraptorid embryos of the egg-shell genus Macroolithus (which may represent eggs of Heyuannia). Examinations of eggs attributed to Heyuannia by Jasmina Wiemann and Tzu-Ruei Yang et al revealed the eggs preserved the blue-green pigment biliverdin and the reddish-brown pigment protoporphyrin, the same pigments found in many modern birds' eggshells. The eggs are thought to have been a blue-green color, because biliverdin is preserved in much greater abundance the photoporphyrin. In modern bird eggs, coloration can camouflage the eggs or help parents identify eggs, and it is correlated with more intensive parental care.
A study by Robert Eagle et al. of the University of California-Los Angeles indicates that from specimens of eggs found in Mongolia and examination of the isotopes carbon-13 and oxygen 18 found within, Oviraptorids had body temperatures that could be elevated higher than that of the surrounding environment but lower than that of birds. This is very different from the isotope ratios of sauropod dinosaurs like Brachiosaurus, which had body temperatures of up to 100 °F (38 °C) and were fully endothermic.
Oviraptorids were probably feathered, since some close relatives were found with feathers preserved (including species of Caudipteryx, Protarchaeopteryx and Similicaudipteryx). Another finding pointing to this is the discovery in Nomingia of a pygostyle, a bone that results from the fusion of the last tail vertebrae and is responsible in birds to hold a fan of feathers in the tail. Finally, the arm position of the brooding Citipati would have been far more effective if feathers were present to cover the eggs.
The brooding oviraptorid specimen IGM 100/979 showed a callus and possible longitudinal groove left over from a healed fracture of the right ulna. Other oviraptorids have had pathological features reported in their phalanges but these have not been described in detail in the scientific literature.
Caenagnathidae is a family of bird-like maniraptoran theropod dinosaurs from the Late Cretaceous of North America and Asia. They are a member of the Oviraptorosauria, and close relatives of the Oviraptoridae. Like other oviraptorosaurs, caenagnathids had specialized beaks, long necks, and short tails, and would have been covered in feathers. The relationships of caenagnathids were long a puzzle. The family was originally named by Charles Hazelius Sternberg in 1940 as a family of flightless birds. The discovery of skeletons of the related oviraptorids revealed that they were in fact non-avian theropods, and the discovery of more complete caenagnathid remains revealed that Chirostenotes pergracilis, originally named on the basis of a pair of hands, and "Ornithomimus" elegans, named from a foot, were caenagnathids as well.Caenagnathoidea
Caenagnathoidea ("recent jaw forms") is a group of advanced oviraptorosaurian dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, often with bony crests atop the head. They ranged in size from Caudipteryx, which was the size of a turkey, to the 8 meter long, 1.4 ton Gigantoraptor. The group (along with all maniraptoran dinosaurs) is close to the ancestry of birds. The most complete specimens have been found in Asia, representing members of the sub-group Oviraptorinae. Notable but fragmentary remains are also known from North America, almost all of which belong to the subgroup Elmisaurinae.The earliest definitive caenangnathoid is Microvenator celer, which dates to the late Aptian age of the early Cretaceous period, though the slightly earlier Caudipteryx from the lower Yixian Formation of China, may also be a member of this group.Caudipteryx
Caudipteryx (which means "tail feather") is a genus of peacock-sized theropod dinosaurs that lived in the Aptian age of the early Cretaceous Period (about 124.6 million years ago). They were feathered and remarkably birdlike in their overall appearance.
Two species have been described; C. zoui (the type species), in 1998, and C. dongi, in 2000.Caudipteryx fossils were first discovered in the Yixian Formation of the Sihetun area of Liaoning Province, northeastern China in 1997.Conchoraptor
Conchoraptor (meaning "conch plunderer") is a genus of oviraptorid dinosaur from the late Cretaceous Period of what is now Asia.Corythoraptor
Corythoraptor (meaning "crested raptor") is a genus of crested oviraptorid theropod dinosaur from the Nanxiong Formation of China. It is known from one species, C. jacobsi, named after palaeontologist Louis L. Jacobs. Including it, there are seven oviraptorids known from the Nanxiong Formation, showing a high level of diversity in the area, and that the different taxa may have occupied different ecological niches.Dinosaur behavior
Dinosaur behavior is difficult for paleontologists to study since much of paleontology is dependent solely on the physical remains of ancient life. However, trace fossils and paleopathology can give insight into dinosaur behavior. Interpretations of dinosaur behavior are generally based on the pose of body fossils and their habitat, computer simulations of their biomechanics, and comparisons with modern animals in similar ecological niches. As such, the current understanding of dinosaur behavior relies on speculation, and will likely remain controversial for the foreseeable future. However, there is general agreement that some behaviors which are common in crocodiles and birds, dinosaurs' closest living relatives, were also common among dinosaurs. Gregarious behavior was common in many dinosaur species. Dinosaurs may have congregated in herds for defense, for migratory purposes, or to provide protection for their young. There is evidence that many types of dinosaurs, including various theropods, sauropods, ankylosaurians, ornithopods, and ceratopsians, formed aggregations of immature individuals. Nests and eggs have been found for most major groups of dinosaurs, and it appears likely that dinosaurs communicated with their young, in a manner similar to modern birds and crocodiles. The crests and frills of some dinosaurs, like the marginocephalians, theropods and lambeosaurines, may have been too fragile to be used for active defense, and so they were likely used for sexual or aggressive displays, though little is known about dinosaur mating and territorialism. Most dinosaurs seem to have relied on land-based locomotion. A good understanding of how dinosaurs moved on the ground is key to models of dinosaur behavior; the science of biomechanics, in particular, has provided significant insight in this area. For example, studies of the forces exerted by muscles and gravity on dinosaurs' skeletal structure have investigated how fast dinosaurs could run, whether diplodocids could create sonic booms via whip-like tail snapping, and whether sauropods could float.Ganzhousaurus
Ganzhousaurus is an extinct genus of oviraptorine oviraptorid dinosaur known from the Late Cretaceous Nanxiong Formation of Nankang County, Ganzhou City of Jiangxi Province, southern China. It was found in a Maastrichtian deposit and contains a single species, Ganzhousaurus nankangensis. It is distinguished by a combination of primitive and derived features.Heyuannia
Heyuannia ("from Heyuan") is a genus of oviraptorid dinosaur that lived during the Late Cretaceous Period in China. It was the first oviraptorid found in that country; most others were found in neighbouring Mongolia. Two species are known: H. huangi, named by Lü Junchang in 2002; and H. yanshini, originally named as a separate genus Ingenia by Rinchen Barsbold in 1981 (and renamed to Ajancingenia in 2013 due to the preoccupation of Ingenia).Huanansaurus
Huanansaurus is an extinct genus of oviraptorid dinosaur that lived approximately 72 million years ago, between the Campanian and Maastrichtian, during the latter part of the Cretaceous period in what is now China, in the Nanxiong Formation.Jiangxisaurus
Jiangxisaurus is an extinct genus of oviraptorid theropod dinosaur from Late Cretaceous of China.
It was similar to Heyuannia, but with more strongly curved anterior claws and a thinner, frailer mandible.
This find is paleontologically significant because it contributes to current knowledge about the paleogeographical distribution of oviraptorids in southern China.Khaan
Khaan (; Mongol [χaːŋ] 'lord') was an oviraptorid dinosaur that was found in the Djadochta Formation of Mongolia and lived in the Late Cretaceous Period (Campanian), 75 million years ago.Machairasaurus
Machairasaurus is a genus of oviraptorid dinosaur which was found in the Bayan Mandahu Formation, China dating to the late Cretaceous period.During the Sino-Canadian expeditions of 1988 and 1990 some skeletons of unknown oviraptorosaurians were discovered by Philip J. Currie in Inner Mongolia. Based on two of these a new genus was named and described by Nicholas R. Longrich, Currie and Dong Zhiming in 2010 with as type species Machairasaurus leptonychus. The generic name is derived from Greek μάχαιρα (makhaira), "short scimitar". The specific name is derived from Greek λεπτός (leptos), "slender", and ὄνυξ (onyx), "claw". The species name as a whole refers to the sabre-like claws of the hand.The holotype, IVPP V15979, was found in layers of the Bayan Mandahu dating from the late Campanian. It mainly consists of a left frontlimb, including the lower end of the lower arm, two carpal bones and a complete hand. Also some fragmentary foot elements were present. The other find is the paratype, IVPP V15980, consisting of a very fragmentary skeleton including tail vertebrae, chevrons, ribs, phalanges of the hands, fragments of the second and fourth metatarsals and pedal phalanges.Five oviraptorid specimens associated with a nest, the female having been found brooding near the eggs, may belong to Machairasaurus.Machairasaurus was a small bipedal theropod. The describers established a single autapomorphy, unique derived trait: the hand claws are very elongated and blade-like in side view, with a length four times that of the joint height. The long claws would be proof that basal oviraptorids used their hands to pull down branches; the more curved claws of more derived forms would have served to dig up roots.Machairasaurus was in 2010 assigned to the Oviraptoridae, more precisely to the Ingeniinae. It formed a smaller clade with "Ingenia" yanshini, Heyuannia huangi, Conchoraptor gracilis, and Nemegtomaia barsboldi.Nankangia
Nankangia is an extinct genus of caenagnathoid oviraptorosaurian dinosaur known from the Late Cretaceous Nanxiong Formation of Nankang County, Ganzhou City of Jiangxi Province, southeastern China. It contains a single species, Nankangia jiangxiensis. N. jiangxiensis coexisted with at least four other caenagnathoids, including an unnamed oviraptorid, Banji long, Ganzhousaurus nankangensis and Jiangxisaurus ganzhouensis. The relatively short dentary and non-downturned mandibular symphysis of Nankangia suggest that it may have been more herbivorous than carnivorous.Nomingia
Nomingia is a genus of oviraptorid theropod dinosaur hailing from the Late Cretaceous Bugin Tsav Beds of Mongolia.Oviraptor
Oviraptor is a genus of small Mongolian theropod dinosaurs, first discovered by technician George Olsen in an expedition led by Roy Chapman Andrews, and first described by Henry Fairfield Osborn, in 1924. Its name is Latin for 'egg taker' or "egg seizer", referring to the fact that the first fossil specimen was discovered atop a pile of what were thought to be Protoceratops eggs, and the specific name philoceratops means "lover of ceratopsians", also given as a result of this find. In his 1924 paper, Osborn explained that the name was given due to the close proximity of the skull of Oviraptor to the nest (it was separated from the eggs by only 4 inches or 10 centimetres of sand). However, Osborn also suggested that the name Oviraptor "may entirely mislead us as to its feeding habits and belie its character". In the 1990s, the discovery of nesting oviraptorids like Citipati proved that Osborn was correct in his caution regarding the name. These finds showed that the eggs in question probably belonged to Oviraptor itself, and that the specimen was actually brooding its eggs, when it died at the nest.
Oviraptor lived in the late Cretaceous period, during the late Campanian stage about 75 million years ago; only one definitive specimen is known (with associated eggs), from the Djadokhta Formation of Mongolia, though a possible second specimen (also with eggs) comes from the northeast region of Inner Mongolia, China, in an area called Bayan Mandahu.Oviraptorosauria
Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or without bony crests atop the head. They ranged in size from Caudipteryx, which was the size of a turkey, to the 8 metre long, 1.4 ton Gigantoraptor. The group (along with all maniraptoran dinosaurs) is close to the ancestry of birds. Analyses like those of Maryanska et al (2002) and Osmólska et al. (2004) suggest that they may represent primitive flightless birds. The most complete oviraptorosaur specimens have been found in Asia. The North American oviraptorosaur record is sparse.The earliest and most basal ("primitive") known oviraptorosaurs are Ningyuansaurus wangi, Protarchaeopteryx robusta and Incisivosaurus gauthieri, both from the lower Yixian Formation of China, dating to about 125 million years ago during the Aptian age of the early Cretaceous period. A tiny neck vertebra reported from the Wadhurst Clay Formation of England shares some features in common with oviraptorosaurs, and may represent an earlier occurrence of this group (at about 140 million years ago).Timeline of oviraptorosaur research
This timeline of oviraptorosaur research is a chronological listing of events in the history of paleontology focused on the oviraptorosaurs, a group of beaked, bird-like theropod dinosaurs. The early history of oviraptorosaur paleontology is characterized by taxonomic confusion due to the unusual characteristics of these dinosaurs. When initially described in 1924 Oviraptor itself was thought to be a member of the Ornithomimidae, popularly known as the "ostrich" dinosaurs, because both taxa share toothless beaks. Early caenagnathid oviraptorosaur discoveries like Caenagnathus itself were also incorrectly classified at the time, having been misidentified as birds.The hypothesis that caenagnathids were birds was questioned as early as 1956 by Romer, but not corrected until Osmolska formally reclassified them as dinosaurs in 1976. Meanwhile, the classification of Oviraptor as an ornithomimid persisted unquestioned by researchers like Romer and Steel until the early 1970s when Dale Russell argued against the idea in 1972. In 1976 when Osmolska recognized Oviraptor's relationship with the Caenagnathids, she also recognized that it was not an ornithomimid and reclassified it as a member of the former family. However, that same year Rinchen Barsbold argued that Oviraptor belonged to a distinct family he named the Oviraptoridae and he also formally named the Oviraptorosauria later in the same year.Like their classification, the paleobiology of oviraptorosaurs has been subject to controversy and reinterpretation. The first scientifically documented Oviraptor skeleton was found lying on a nest of eggs. Because its powerful parrot-like beak appeared well-adapted to crushing hard food items and the eggs were thought to belonged to the neoceratopsian Protoceratops, oviraptorosaurs were thought to be nest-raiders that preyed on the eggs of other dinosaurs. In the 1980s, Barsbold proposed that oviraptorosaurs used their beaks to crack mollusk shells as well. In 1993, Currie and colleagues hypothesized that small vertebrate prey may have also been part of the oviraptorosaur diet. Not long after, fossil embryonic remains cast doubt on the popular reconstruction of oviraptorosaurs as egg thieves when it was discovered that the "Protoceratops" eggs that Oviraptor was thought to be "stealing" actually belonged to Oviraptor itself. The discovery of additional Oviraptor preserved on top of nests in lifelike brooding posture firmly established that oviraptorosaurs had been "framed" as egg thieves and were actually caring parents incubating their own nests.Tongtianlong
Tongtianlong (meaning "Tongtianyan dragon") is a genus of oviraptorid theropod dinosaurs that lived in the late Maastrichtian epoch of the late Cretaceous period. It contains one species, T. limosus.Wulatelong
Wulatelong is an extinct genus of basal oviraptorid dinosaur known from the Late Cretaceous Wulansuhai Formation (Campanian stage) of Bayan Mandahu, Linhe District of Inner Mongolia, northern China. It contains a single species, Wulatelong gobiensis.