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Timeline of oviraptorosaur research

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Restored profiles of variousoviraptorids

Thistimeline of oviraptorosaur researchis a chronological listing of events in thehistoryof paleontology focused on theoviraptorosaurs,a group ofbeaked,bird-liketheropoddinosaurs.The earlyhistoryof oviraptorosaur paleontology is characterized bytaxonomicconfusion due to the unusual characteristics of these dinosaurs. When initially described in1924Oviraptoritself was thought to be a member of theOrnithomimidae,popularly known as the "ostrich"dinosaurs, because both taxa share toothless beaks.[1]Earlycaenagnathidoviraptorosaur discoveries likeCaenagnathusitself were also incorrectly classified at the time, having been misidentified as birds.[1]

Thehypothesisthat caenagnathids were birds was questioned as early as1956byRomer,but not corrected untilOsmolskaformally reclassified them as dinosaurs in1976.Meanwhile, the classification ofOviraptoras an ornithomimid persisted unquestioned by researchers like Romer andSteel[disambiguation needed]until the early1970swhenDale Russellargued against the idea in1972.In 1976 when Osmolska recognizedOviraptor's relationship with the Caenagnathids, she also recognized that it was not an ornithomimid and reclassified it as a member of the former family.[1]However, that same yearRinchen Barsboldargued thatOviraptorbelonged to a distinct family he named theOviraptoridae[1]and he also formally named the Oviraptorosauria later in the same year.[2]

Like their classification, thepaleobiologyof oviraptorosaurs has been subject to controversy and reinterpretation. The first scientifically documentedOviraptorskeleton was found lying on a nest ofeggs.Because its powerfulparrot-like beak appeared well-adapted to crushing hard food items and the eggs were thought to belonged to theneoceratopsianProtoceratops,oviraptorosaurs were thought to be nest-raiders that preyed on the eggs of other dinosaurs. In the1980s,Barsbold proposed that oviraptorosaurs used their beaks to crackmolluskshells as well. In1993,Currieand colleagues hypothesized that smallvertebrateprey may have also been part of the oviraptorosaur diet. Not long after,fossil embryonic remainscast doubt on the popular reconstruction of oviraptorosaurs as egg thieves when it was discovered that the "Protoceratops"eggs thatOviraptorwas thought to be "stealing" actually belonged toOviraptoritself. The discovery of additionalOviraptorpreserved on top of nests in lifelikebroodingposture firmly established that oviraptorosaurs had been "framed" as egg thieves and were actually caring parentsincubatingtheir own nests.[3]

20th century

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Drawing of the left arm and both hands of theOviraptortype specimen AMNH 6517

1920s

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1923

  • A specimen of the species that would come to be namedOviraptor philoceratopswas found preserved on top of a nest of eggs.[3]

1924

  • Osborn described the new genus and speciesOviraptor philoceratops.[2]He classified it as an ornithomimid because it didn't have any teeth in its jaws[1]and interpreted the genus as being adapted to a diet of eggs.[3]Since a specimen was found preserved on top of a nest of eggs presumed to belong to Protoceratops, Osborn thought that it was smothered by a sandstorm while in the act of raiding the nest.[3]
Skeletal reconstruction ofChirostenotes

1930s

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1932

1933

1940s

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Drawing of the skull ofOviraptor,type specimen AMNH 6517

1940

1950s

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1956

  • Romer followed Osborn's original classification ofOviraptoras an ornithomimid. He also observed that caenagnathids had reptilian characteristics and may have been coelurosaurs.[1]

1960s

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1960

  • Wetmore questioned the hypothesis that caenagnathids were birds because their remains exhibit some reptilian traits.[1]

1966

  • Romer continued to follow Osborn's original classification ofOviraptoras an ornithomimid.[1]

1970s

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Skeletal reconstruction ofMicrovenator

1970

  • Ostrom described the new genus and speciesMicrovenator celer.[4]
  • Steel followed Osborn's original classification ofOviraptoras an ornithomimid.[1]He also questioned the avian status of caenagnathids and proposed that they might actually be coelurosaurs instead.[1]
Artist's restoration ofMicrovenator

1971

  • Cracroft named the new speciesCaenagnathus sternbergi.[2]He thought it was a bird.[1]

1972

  • Russell argued against the classification ofOviraptoras an ornithomimid.[1]

1976

  • Osmolska recognized that caenagnathids were theropods and classifiedOviraptoras a member of the family.[1]
  • Barsbold classifiedOviraptoras a member of the new taxaOviraptorinaeandOviraptoridae.[1]
  • Barsbold named the Oviraptorosauria.[2]

1980s

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Artist's restoration ofAvimimus

1981

Known skeletal material ofAjancingenia,formerly "Ingenia"
  • Barsbold described the new genus and speciesIngenia yanshini.[4]He also named the subfamily Ingeniinae[4]and classified the family Caenagnathidae in the Oviraptorosauria.[1]
  • Osmolska described the new genus and speciesElmisaurus rarus.[4]

1983

  • Barsbold proposed that oviraptorosaurs used their powerful beaks to feed on shelled mollusks.[3]

1986

1988

  • Currie and Russell observed that caenagnathids had arctometatarsalian feet.[3]

1990s

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FossilizedOviraptornest, specimen AMNH FR 6508

1991

  • Jerzykiewicz and Russell observed that oviraptorosaurs seem to have been most common during the Djadokhta stage.[3]
  • Sabath described an oviraptorosaur nest mound.[3]

1992

  • Smith interpreted the biomechanics of oviraptorosaur jaws to imply that these dinosaurs were actually herbivorous.[3]

1993

  • Jerzykiewicz and others observed that oviraptorosaurs seem to have been most common in central Asia.[3]
  • Currie, Godfrey, and Nessov described the new genus and speciesCaenagnathasia martinsoni.[2]
  • Currie and others built a case for interpreting oviraptorosaurs as egg-eaters who supplemented their diet with small prey. They noted supporting traits like the animals' ability to give a "powerful nipping bite" with the front of its beak. Oviraptorosaurs also had tooth-like projections from the roof of the mouth, which resemble similar adaptations in modern egg-eating mammals.
    NestingCitipatispecimen nicknamed "Big Auntie"
    The orientation of the throat on the underside of the jaw is also consistent with this reconstruction of oviraptorosaur paleoecology.[3]
  • Russell and Dong argued that the Maniraptora (which includes Oviraptorosauria) was a polyphyletic assemblage of unrelated groups. Instead they classified the traditional oviraptorosaurs with the ornithomimids, therizinosauroids, and troodontids in a new, greatly expanded Oviraptorosauria.[5]

1994

  • Norell and others reported the discovery of a tiny theropod skeleton in an oviraptorid nest. They suggested this find was evidence that oviraptorosaur hunted tiny game. They also noted that the supposedProtoceratopseggs of the Central Asiatic Expeditions actually preserved the embryonic remains of oviraptorosaurs.[3]

1996

  • Currie reported the presence of nests of large eggs more than 40 centimetres (16 in) long in China. Oviraptorosaurs would come to be considered possible candidates for the egg layers.[3]
    FossilizedCitipatiegg with preserved embryo
  • Dong and Currie reported the discovery of an oviraptorid from the Djadokhta Formation of northern China preserved on top of a nest of eggs. This overturned more than 60 years of interpreting theOviraptorof the Central Asiatic Expeditions as a rapacious egg-thief in favor of it likely being a faithful mother at her nest. The researchers reconstructed the way mother oviraptorosaurs built their nests. The standing mother would lay a pair of eggs and bury them by hand. Then she would turn and repeat the process until she had made a ring of egg-pairs completely around herself. Since by then the area where she was standing would be higher than the eggs, she would repeat the process with another ring of egg-pairs as a second layer. This process would gradually build an egg mound containing as many as 30 eggs in up to three layers.[3]

1997

  • Sues published a critical review of earlier interpretations of the oviraptorosaurs' evolutionary relationships and formulated the clade's firstsynapomorphy-based diagnosis. He also performed a cladistic analysis and found oviraptorosaurs to be the sister group of the therizinosaurs.[6]
  • Sereno regarded oviraptorosaurs as maniraptorans and found them to be the sister group of Paraves (which includes deinonychosaurs and birds) in a cladistic analysis.[6]
  • Padian and others published a cladistic definition for Oviraptorsauris for the first time; all taxa closer toOviraptorthan to birds.[6]
Cast of aCaudipteryx zouispecimen,Houston Museum of Natural Science
  • Barsbold described the new genusRincheniafor the speciesOviraptor mongoliensis.[2]Barsbold credited Currie and Padian for defining Oviraptorosauria as the Oviraptoridae and all taxa closer toOviraptor.[6]

1998

  • Sereno regarded oviraptorosaurs as maniraptorans and found them to be the sister group of Paraves (which includes deinonychosaurs and birds) in a cladistic analysis.[6]He defined oviraptorosaurs as all maniraptorans closer toOviraptorthan to Neornithes.[6]
  • Currie, Norell, and Ji described the new genus and speciesCaudipteryx zoui.[2]
  • Ji and others reported the presence of gastroliths inCaudipteryx.These are evidence for an herbivorous diet.[3]
  • Makovicky and Sues considered oviraptorosaurs to be the sister group of the therizinosaurs.[6]

1999

  • Sereno foundCaudipteryxto be a basal oviraptorosaur. He also erected the clade Caenagnathoidea for the caenagnathids and oviraptorids.[6]
  • Elzanowski performed a cladistic analysis and found a group consisting of oviraptorosaurs, ornithomimosaurs and therizinosaurs were more closely related to birds than deinonychosaurs. No other cladistic study in the history of dinosaur research had come up with this result.[6]
  • Padian and others changed the definition of Oviraptorosauria from a stem-based clade to a node-based one. They defined the oviraptorosaurs as "OviraptorandChirostenotes(=Caenagnathus) and all the descendants of their most recent common ancestor. "[6]
  • Clark and others observed that oviraptorosaurs are among the most common dinosaurs found at Ukhaa Tolgod in Mongolia.[3]They reported further specimens preserved on nests in brooding position. They suggested contrary to Dong and Currie's 1996 reconstruction of oviraptorosaur nest-building behavior that the animals may have constructed their nests by maneuvering the eggs into position by hand. However, this explanation is less parsimonious and has less evidentiary support, so it never gained favor among paleontologists.[3]

21st century

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Artist's restoration ofNomingia

2000s

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2000

2001

Citipati osmolskaeand an unnamedCitipatispecies
  • Clark, Norell, and Barsbold described the new genus and speciesCitipati osmolskae.[4]
  • Clark, Norell, and Barsbold described the new genus and speciesKhaan mckennai.[4]
  • David J. Varrichioreported the first occurrences ofoviraptorosaursfromMontana.[7]The first find was anarticularregion from the lower jaw ofCaenagnathus sternbergiofCampanianage from theTwo Medicine Formation.[7]This species had previously only been known from the Canadian province ofAlberta.[7]Another new Montanan oviraptorosaur specimen, a foot found in theHell Creek Formation,was assigned toLeptorhynchus elegans(asElmisaurus elegans).[7]
  • Kevin Padian,Ji QiangandJi Shu-anpublished a review of knownfeathered dinosaursand their implications for theorigin of flight.[8]The authors observe that many aspects of the distribution of feather homologues meet the expectations of earlier phylogenetic hypotheses, including a gradual transition from primitive filaments inSinosauropteryxto the shared filaments and "rudimentary" true feathers inCaudipteryxandProtarchaeopteryx,to flight feathers inArchaeopteryx.[9]The team speculates that the plumulaceous feathers inCauditeryxandProtarchaeopteryxmay have originated as tufts ofSinosauropteryx-style filaments, the shafts of which possibly formed by the consolidation of individual filaments.[10]The parallel nature of the barbs inCaudipteryxandProtarcheopteryxsuggest the existence ofbarbules.[11]This suggests that barbules, which are necessary for flight-worthy wings, evolved prior to flight.[11]
The skull ofIncisivosaurus

2000

  • 2002?Teresa Maryańskaand others confirmed Sereno's finding thatCaudipteryxwas an oviraptorosaur. They also foundAvimimusto be an oviraptorosaur as well.[6]

2002

  • Xu and others described the new genus and speciesIncisivosaurus gauthieri.[2]Their research supported recent findings thatCaudipteryxandAvimimuswere oviraptorosaurs.[6]
  • Maryanska and others performed a cladistic analysis that found oviraptorosaurs to be avialans.[6]
  • Zelenitsky and others studied the shape and shell histology of the large fossil eggs reported from China by Currie in 1996 and concluded that they may have been laid by oviraptorosaurs. Given their large size, this implied that giant oviraptorosaurs remained to be discovered.[3]

2003

Skeletal mount ofGigantoraptor

2004

2005

2007

2008

2009

2010s

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Artist's restoration ofMachairasaurus

2010

2011

Artist's restoration of twoAjancingenia

2012

Skull and neck ofAnzu

2013

2014

2015

2016

2017

2018

2019

See also

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Footnotes

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  1. ^abcdefghijklmnopqOsmolska, Currie, and Barsbold (2004);"Systematics and Evolution", page 178.
  2. ^abcdefghijklmnopqOsmolska, Currie, and Barsbold (2004);"Table 8.1: Oviraptorosauria", page 166.
  3. ^abcdefghijklmnopqrOsmolska, Currie, and Barsbold (2004);"Paleoecology", page 183.
  4. ^abcdefghOsmolska, Currie, and Barsbold (2004);"Table 8.1: Oviraptorosauria", page 167.
  5. ^Osmolska, Currie, and Barsbold (2004);"Systematics and Evolution", pages 178-179.
  6. ^abcdefghijklmOsmolska, Currie, and Barsbold (2004);"Systematics and Evolution", page 179.
  7. ^abcdVarricchio (2001);"Abstract," page 42.
  8. ^Padian, Ji, and Ji (2001);"Abstract," page 117.
  9. ^Padian, Ji, and Ji (2001);"Conclusions," pages 131-132.
  10. ^Padian, Ji, and Ji (2001);"Possible Evolutionary Sequence from Protofeathers to True Feathers," page 126.
  11. ^abPadian, Ji, and Ji (2001);"Possible Evolutionary Sequence from Protofeathers to True Feathers," page 127.
  12. ^Lü et al. (2004);"Abstract," page 95.
  13. ^Zanno and Sampson (2005);"Abstract," page 897.
  14. ^Lü et al. (2005);"Abstract," page 51.
  15. ^Lü and Zhang (2005);"Abstract," page 412.
  16. ^Xu et al. (2007);"Abstract," page 844.
  17. ^He, Wang, and Zhou (2008);"Abstract," page 178.
  18. ^Lü et al. (2009);"Abstract," page 43.
  19. ^Xu and Han (2010);"Abstract," page 11.
  20. ^Longrich, Currie, and Dong (2010);"Abstract," page 23.
  21. ^abSullivan, Jasinski and Van Tomme (2011);"Abstract," page 418.
  22. ^Ji et al. (2012);"Abstract," page 2102.
  23. ^Easter (2013);"Abstract," page 184.
  24. ^Wang et al. (2013);"Abstract," page 242.
  25. ^Wei et al. (2013);"Abstract," page 11.
  26. ^Longrich et al. (2013);"Abstract," page 899.
  27. ^Lü et al. (2013b);"Abstract," page 1.
  28. ^Xu et al. (2013);"Abstract," page 85.
  29. ^Lü et al. (2013a);"Abstract," page 165.
  30. ^Lamanna et al. (2014);"Abstract," page 1.
  31. ^Lü et al. (2015);in passim.
  32. ^Funston and Currie (2016);in passim.
  33. ^Lü et al. (2016);in passim.
  34. ^Yilun Yu; Kebai Wang; Shuqing Chen; Corwin Sullivan; Shuo Wang; Peiye Wang; Xing Xu (2018)."A new caenagnathid dinosaur from the Upper Cretaceous Wangshi Group of Shandong, China, with comments on size variation among oviraptorosaurs".Scientific Reports.8(1): Article number 5030.Bibcode:2018NatSR...8.5030Y.doi:10.1038/s41598-018-23252-2.PMC5864915.PMID29567954.
  35. ^G.F. Funston; S.E. Mendonca; P.J. Currie; R. Barsbold (2018). "Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin".Palaeogeography, Palaeoclimatology, Palaeoecology.494:101–120.Bibcode:2018PPP...494..101F.doi:10.1016/j.palaeo.2017.10.023.
  36. ^Sungjin Lee; Yuong-Nam Lee; Anusuya Chinsamy; Junchang Lü; Rinchen Barsbold; Khishigjav Tsogtbaatar (2019)."A new baby oviraptorid dinosaur (Dinosauria: Theropoda) from the Upper Cretaceous Nemegt Formation of Mongolia".PLOS ONE.14(2): e0210867.Bibcode:2019PLoSO..1410867L.doi:10.1371/journal.pone.0210867.PMC6364893.PMID30726228.
  37. ^Rui Qiu; Xiaolin Wang; Qiang Wang; Ning Li; Jialiang Zhang; Yiyun Ma (2019)."A new caudipterid from the Lower Cretaceous of China with information on the evolution of the manus of Oviraptorosauria".Scientific Reports.9(1): Article number 6431.Bibcode:2019NatSR...9.6431Q.doi:10.1038/s41598-019-42547-6.PMC6483983.PMID31024012.

References

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