Oviraptorosauria

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Oviraptorosaurs
Temporal range:
Ma
Oviraptorosauria diversity. Clockwise from top left: GIN 100/42 (which may represent Citipati or a different taxon), "Ronaldoraptor" (undescribed), Avimimus, Anzu and Gigantoraptor
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Pennaraptora
Clade: Oviraptorosauria
Barsbold, 1976
Subgroups
Synonyms

Oviraptorosaurs ("egg thief lizards") are a group of feathered

Maryanska et al (2002) and Osmólska et al. (2004) have proposed that they may represent primitive flightless birds.[5][6] The most complete oviraptorosaur specimens have been found in Asia.[7] The North American oviraptorosaur record is sparse.[7]

The earliest and most basal ("primitive") known oviraptorosaurs are

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).[8]

Description

Oviraptorosaurs to scale

Oviraptorosaurians have shortened rostrums, massive, beaklike mandibles, and long parietal bones. With the exception of the 8-meter long Gigantoraptor, they are generally medium-sized and rarely exceeded 2 meters in length. The most primitive members have four pairs of teeth in the premaxillae, such as in Caudipteryx[9] and in Incisivosaurus they are enlarged and form bizarrely prominent bucktoothed incisors. The more advanced members have no teeth in the jaws.

biceps tubercle.[10]

Anzu wyliei skeleton cast in the Rocky Mountain Dinosaur Resource Center in Woodland Park, Colorado

Oviraptorosaurians are different from most other maniraptorans in the form of their skulls. They have shortened snouts, beak-like jaws with few or no teeth, and a large opening in the lower jaw bone. Some have bony crests atop the skull. The most primitive members have a few teeth in the front of the mouth; in Incisivosaurus, they are enlarged and form bizarrely prominent "bucktoothed" incisors. The arms and hands are generally long (though very reduced in some advanced species) and the shoulder girdle is large and massive, with flexed coracoid bones and prominent attachments for strong arm muscles.

Their tails are very short compared to other maniraptorans. In Nomingia and Similicaudipteryx, the tail ends in four fused vertebrae which Osmólska, He, and others have referred to as a "pygostyle", but which Witmer found was anatomically different and evolved separately from the pygostyle of birds (a bone which serves as the attachment point for a fan of tail feathers).[2][10]

Feathers

Evidence for feathered oviraptorosaurs exists in several forms. Most directly, four species of primitive oviraptorosaurs (in the genera

Citipati, and Conchoraptor) preserved tails ending in something like a pygostyle, a bony structure at the end of the tail that, in modern birds, is used to support a fan of feathers.[2][11] Similarly, quill knobs (anchor points for wing feathers on the ulna) have been reported in the oviraptorosaurian species Avimimus portentosus.[12] Additionally, a number of oviraptorid specimens have famously been discovered in a nesting position similar to that of modern birds. The arms of these specimens are positioned in such a way that they could perfectly cover their eggs if they had small wings and a substantial covering of feathers.[13]

Notably, a study on flight feathers has concluded that Caudipteryx was secondarily flightless, implying an ancestral volant ancestor for oviraptorosaurs.[14]

Paleobiology

Diet

The eating habits of these animals are not fully known: they have been suggested to have been either carnivorous, herbivorous, mollusk-eating or egg-eating (the evidence that originally supported the latter is no longer considered valid); these options are not necessarily incompatible.

Some ate small

gastroliths preserved with Caudipteryx
. There are also arguments for the inclusion of mollusks in their diet.

Originally these animals were thought to be egg raiders, based on a Mongolian find showing Oviraptor on top of a nest. Recent studies have shown that the animal was actually on top of its own nest.[15]

A 2022 study of the bite forces of oviraptorosaurs such as Incisivosaurus, Khaan, Citipati, and Conchoraptor suggests that most if not all oviraptorosaurs had a very strong bite force. The moderate jaw gape seen in oviraptorosaurs is indicative of herbivory in the majority of the group, but it is clear they were likely feeding on much tougher or more types of vegetation than other herbivorous theropods in their environment, such as ornithomimosaurs and therizinosaurs were able to. The examinations suggest oviraptorosaurs may have been powerful-biting generalists or specialists that partook of niche partitioning both in body size and cranial function. One particular group, the Caenagnathidae, may have also been more omnivorous or even carnivorous than other oviraptorosaurs.[16]

Reproduction

Hatchling specimen known as "baby Louie"

Several oviraptorosaurian nests are known, with several oviraptorid specimens preserved in a brooding position over large clutches of up to a dozen or more eggs. The eggs are usually arranged in pairs, and forming a circular pattern within the nest. One oviraptorosaurian specimen from China has been found with 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 were shaped like highly elongated ovals — the greatest egg elongation among diapsids — with the more pointed end pointing backward from the cloaca and oriented toward the center of the nest.[17][18] Geochemical analysis also revealed that oviraptorosaurs incubated their eggs in the 35–40 °C (95–104 °F) range, as many modern bird species do today, based on the oxygen isotope ratios in the bones of the fossil embryos of various species during development.[19]

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.[17]

Relationship to birds

Oviraptorosaurs, like

deinonychosaurs, are so bird-like that several scientists consider them to be true birds, more advanced than Archaeopteryx. Gregory S. Paul has written extensively on this possibility, and Teresa Maryańska and colleagues published a technical paper detailing this idea in 2002.[5][20][21] Michael Benton
, in his widely respected text Vertebrate Paleontology, wrote placement of oviraptorosaurs among birds is highly controversial .

Classification

The internal classification of the oviraptorosaurs has also been controversial. Most studies divide them into two primary sub-groups, the Caenagnathidae and the Oviraptoridae. The Oviraptoridae is further divided into the small, short-armed, and crestless subfamily Ingeniinae, and the larger, crested, long-armed Oviraptorinae. However, some phylogenetic studies have suggested that many traditional members of the Caenagnathidae were more closely related to the crested oviraptorids.

Phylogeny

The 2007 cladistic analysis of Turner and colleagues recovered the Oviraptorosauria as a maniraptoran clade (natural grouping) of maniraptorans more primitive than true birds. They found that the oviraptorosaurs are the sister group to the Therizinosauria and that the two, together, are more basal than any member of Paraves.[23] However, a more recent study by Zanno and colleagues challenged that finding, showing therizinosaurs to be more primitive and not closely related to oviraptorosaurs.[24]

The following cladogram was found by an analysis published with the description of the caenagnathid Anzu.[25]

Oviraptorosauria

Incisivosaurus gauthieri

Caudipteridae

Similicaudipteryx yixianensis

Caudipteryx zoui

Caudipteryx dongi

Avimimus portentosus

Caenagnathoidea
Caenagnathidae

Microvenator celer

Gigantoraptor erlianensis

Caenagnathasia martinsoni

Ojoraptorsaurus boerei

Alberta dentary morph 3

Epichirostenotes curriei

Elmisaurus rarus

Hagryphus giganteus

Chirostenotes pergracilis

Leptorhynchos gaddisi

Leptorhynchos elegans

"Caenagnathus" sternbergi

Anzu wyliei

Caenagnathus collinsi

Oviraptoridae

Nankangia jiangxiensis

Yulong mini

Nomingia gobiensis

Oviraptor philoceratops

Rinchenia mongoliensis

Zamyn Khondt oviraptorid

Huanansaurus ganzhouensis

Citipati osmolskae

Citipati sp.

Wulatelong gobiensis

Banji long

Shixinggia oblita

Jiangxisaurus ganzhouensis

Ganzhousaurus nankangensis

Nemegtomaia barsboldi

Machairasaurus leptonychus

Conchoraptor gracilis

Khaan mckennai

Ajancingenia yanshini

Heyuannia huangi

See also

References

  1. ^ Ji Qiang; Lü Jun-Chang; Wei Xue-Fang; Wang Xu-Ri (2012). "A new oviraptorosaur from the Yixian Formation of Jianchang, Western Liaoning Province, China". Geological Bulletin of China. 31 (12): 2102–2107.
  2. ^ a b c He, T.; Wang, X.-L.; Zhou, Z.-H. (2008). "A new genus and species of caudipterid dinosaur from the Lower Cretaceous Jiufotang Formation of western Liaoning, China". Vertebrata PalAsiatica. 46 (3): 178–189.
  3. PMID 33204472
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  4. .
  5. ^ a b Maryanska, T., Osmólska, H., & Wolsam, M. (2002). "Avialian status for Oviraptorosauria". Acta Palaeontologica Polonica. 47 (1): 97–116.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  6. ^ a b Osmólska, Halszka, Currie, Philip J., Brasbold, Rinchen (2004) "The Dinosauria" Weishampel, Dodson, Osmólska. "Chapter 8 Oviraptorosauria" University of California Press.
  7. ^ a b Varricchio, D. J. 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from Montana. pp. 42–57 in D. H. Tanke and K. Carpenter (eds.), Mesozoic Vertebrate Life. Indiana University Press, Indianapolis, Indiana.
  8. .
  9. .
  10. ^
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  12. ^ Kurzanov, S.M. (1987). "Avimimidae and the problem of the origin of birds." Transactions of the Joint Soviet-Mongolian Paleontological Expedition, 31: 5-92. [in Russian]
  13. ^ Hopp, Thomas J., Orsen, Mark J. (2004) "Feathered Dragons: Studies on the Transition from Dinosaurs to Birds. Chapter 11. Dinosaur Brooding Behavior and the Origin of Flight Feathers" Currie, Koppelhaus, Shugar, Wright. Indiana University Press. Bloomington, IN. USA.
  14. ^ https://www.pnas.org/doi/10.1073/pnas.2306639121
  15. S2CID 4245228
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  16. .
  17. ^ .
  18. ^ Eggs of Earliest Dinosaurs Were Leathery, Fossils Show
  19. .
  20. ^ Paul, G.S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press.
  21. ^ Paul, G.S. (1988). Predatory Dinosaurs of the World. New York: Simon & Schuster.
  22. ^ Benton, M. J. (2004). Vertebrate Paleontology, 3rd ed. Blackwell Science Ltd.
  23. ^
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  • Barsbold, R. (1983). "Carnivorous dinosaurs from the Cretaceous of Mongolia". Transactions of the Joint Soviet-Mongolian Paleontological Expedition. 8: 39–44.