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| Oviraptorids | |
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| Montage of four oviraptorids. Clockwise from top left:Citipati? sp.,Nemegtomaia,Huanansaurus andConchoraptor | |
Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Reptilia |
| Clade: | Dinosauria |
| Clade: | Saurischia |
| Clade: | Theropoda |
| Superfamily: | †Caenagnathoidea |
| Family: | †Oviraptoridae Barsbold, 1976 |
| Type species | |
| †Oviraptor philoceratops Osborn, 1924 | |
| Subgroups | |
Oviraptoridae is a group of bird-like, herbivorous and omnivorousmaniraptorandinosaurs. Oviraptorids are characterized by their toothless, parrot-like beaks and, in some cases, elaboratecrests. 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 theGobi Desert ofMongolia and northwesternChina.
The most characteristic feature of this group is the skull structure. Oviraptorids had short snouts and very deepmandibles. Some taxa (such asCitipati,Corythoraptor,Rinchenia) had a midline crest on top of the skull, resembling that of acassowary. Other distinguishing characteristics include a bony spike intruding on themandibular fenestra, nostrils placed very high and far back on the snout, an extremely thin bony bar beneath the eye, and highlypneumatized skull bones. Like their relatives thecaenagnathids, 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 be more closely related to the crested oviraptorids. Because of this, and the fact that at least one study found thatCaenagnathus 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,[1] 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 subfamilyHeyuanniinae and the larger, crested, long-armedOviraptorinae (Oviraptor +Citipati). Some phylogenetic studies have shown thatOviraptor is the most primitive known oviraptorid, thus makingCitipati a closer relative of the "ingeniines" and this traditional division into crestless and crested forms artificial.[2]
Other possible oviraptorids includeNomingia gobiensis,Gigantoraptor erlianensis,Jiangxisaurus ganzhouensis andShixinggia oblita. All four have been suggested to be oviraptorids, caenagnathids, or more primitive than either group.
In 2020, during their description ofOksoko, the cladogram recovered by Funstonet al. is shown below.[3] BecauseOviraptor did not clade withCitipati and the other "oviraptorines", they named the latter's clade Citipatiinae, although they did not provide a formal definition. However, according to Mickey Mortimer, the clade can be considered valid because its describers explicitly name it as new, which satisfies ICZN Article 16.1.[4]
| Oviraptoridae | |
The diet of oviraptorids is not fully understood. Though some appear to have been at least partially carnivorous, they were probably primarily herbivorous or omnivorous.[5]
Originally, oviraptorids were thought to be specialized egg raiders, based on a Mongolian find showingOviraptor on top of a nest erroneously attributed to theceratopsian dinosaurProtoceratops. However, discoveries in the 1990s, includingCitipati specimens clearly brooding (rather than preying on) the same types of nests, and aCitipatiembryo inside the same type of egg preserved in these nests, showed that the "specialized egg thief" idea was incorrect.[6] 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 herbivorousdicynodontsynapsids, which are usually considered herbivorous.(This leads to the possibility that these animals are omnivorous).[5]
Their beaks also share similarities with the beaks of herbivorousparrots andtortoises.[7]
Evidence of partial carnivory among some oviraptorines comes from a lizard skeleton preserved in the body cavity of thetype specimen ofOviraptor[8] and two hatchlingByronosaurus skulls found in aCitipati nest.[9] Some scientists have also suggested that some oviraptorids (especially the small-handed, weak-clawed "ingeniines") fed mainly on plant material.[5]
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Although fossilizeddinosaur eggs are generally rare, oviraptorid eggs are relatively well known. Several oviraptorid nests, eggs, and embryos are known, mostly uncovered in theGobi Desert. Some specimens ofOviraptor philoceratops,Citipati osmolskae,Nemegtomaia andcf.Machairasaurus have been found inbrooding positions in association with nests.[8][10][11] 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.[10]
Oviraptorid eggs are shaped like elongated ovals (elongatoolithid) and resemble the eggs ofratite birds (such asostriches) 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.[12] The eggs ofCitipati are the largest known definitive oviraptorid eggs, at 18 cm. In contrast, eggs associated withOviraptor are only up to 14 cm long.[10]
The first oviraptorid eggs (of the genusOviraptor, which mean "Egg thief") were found in close proximity to the remains of theceratopsian dinosaurProtoceratops and it was assumed that the oviraptorids were preying upon the eggs of the ceratopsians.[13] It was not until 1993, when aCitipati embryo was discovered inside an egg of the type assigned toProtoceratops, that the error was corrected.[6] Norellet al., who recognized the embryo as oviraptorid, assigned it to the genusCitipati. The egg containing the embryo was smaller than most knownCitipati eggs at only 12 cm, though it was partially eroded and broken into three pieces, making an accurate estimate of its original size difficult.[10] 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.[6]
An oviraptorosaurian specimen fromChina described in 2005 was found to have two unlaid eggs within the pelvic canal. This suggests that, unlike moderncrocodilians, 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.[14]
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 twooviducts. However, crocodilians produce multiple shelled eggs per oviduct at a time, whereas oviraptorosaurs, like birds, produced only one egg per oviduct at a time.[14]
In 2017, paleontologists discovered colored pigments in some fossilized oviraptorid embryos of the egg-shell genusMacroolithus (which may represent eggs ofHeyuannia). Examinations of eggs attributed toHeyuannia by Jasmina Wiemann and Tzu-Ruei Yanget al revealed the eggs preserved the blue-green pigmentbiliverdin and the reddish-brown pigmentprotoporphyrin, 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.[15]
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 likeBrachiosaurus, which had body temperatures of up to 100 °F (38 °C) and were fully endothermic.[16]
Oviraptorids were probablyfeathered, since some close relatives were found with feathers preserved (including species ofCaudipteryx,Protarchaeopteryx andSimilicaudipteryx).[17][18] Another finding pointing to this is the discovery inNomingia of apygostyle, 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.[19] Finally, the arm position of the broodingCitipati would have been far more effective if feathers were present to cover the eggs.[20]
The brooding oviraptorid specimenIGM 100/979 showed acallus and possiblelongitudinal groove left over from a healed fracture of the rightulna. Otheroviraptorids have had pathological features reported in their phalanges but these have not been described in detail in the scientific literature.[21]
Almost all oviraptorids come fromdesert deposits of theGobi 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 toankylosaurs andprotoceratopsids. 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 theBarun Goyot andDjadochta Formations) than in wetter ecosystems where large dinosaurs are common (such as theNemegt 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.[5]
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