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Feathered dinosaur

dinosaurs possessing feathers
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A feathered dinosaur is a dinosaur with feathers. It is now thought that all coelurosaurs, and perhaps all theropods, had feathers.

Archaeopteryx feather, showing the off-centre rachis of a typical flight feather. Solnhofen 1860

The possibility that dinosaurs are closely related to birds raised the possibility of feathers. Fossils of Archaeopteryx include well-preserved feathers, but it was not until the early 1990s that clearly non-bird dinosaur fossils were discovered with preserved feathers. About 40 genera of theropods are now known to be feathered.

Most of these fossils come from the Yixian formation in China. The fossil feathers of one specimen, Shuvuuia deserti, have been tested positive for beta-keratin in immunological tests.[1] This is the main protein in bird feathers.

Fossil evidence

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Sinosauropteryx fossil, first fossil of a definitively non-avialan dinosaur with feathers

After a century without clear evidence, well-preserved fossils of feathered dinosaurs were discovered during the 1990s, and more continue to be found. The fossils were preserved in a Lagerstätte — a sedimentary deposit exhibiting remarkable richness and completeness in its fossils — in Liaoning, China.

The area had repeatedly been smothered in volcanic ash by eruptions in Inner Mongolia 124 million years ago, during the Lower Cretaceous. The fine-grained ash preserved the living organisms that it buried in fine detail. The area was teeming with life, with millions of leaves, angiosperms (the oldest known), insects, fish, frogs, salamanders, mammals, turtles, lizards and crocodilians discovered to date.

The most important discoveries at Liaoning have been a host of feathered dinosaur fossils, with a steady stream of new finds filling in the picture of the dinosaur-bird connection and adding more to theories of the evolutionary development of feathers and flight. Quill knobs were found an ulna of Velociraptor mongoliensis. These are associated with large, well-developed secondary feathers.[2]

 
A nesting Citipati osmolskae specimen, at the AMNH.

Behavioural evidence, in the form of an oviraptorosaur on its nest, showed another link with birds. Its forearms were folded, like those of a bird.[3] Although no feathers were preserved, it is likely that these would have been present to keep eggs and juveniles warm.[4]

Evidence in amber

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The feathered tail of a non-avialan (non-bird) theropod has been found preserved in mid-Cretaceous (∼99 mya) amber from Kachin State, Myanmar (Burma).[5] The BBC describe it as "perfectly preserved".[6] Lida Xing found it at an amber market in Myitkina, Myanmar. The amber was already polished for jewellery: the seller thought it was plant material. However, it was the tail of a feathered dinosaur about the size of a sparrow. The feathers were not flight feathers. Dr Paul Barrett, from London's Natural History Museum, called the specimen a "beautiful fossil", describing it as a "really rare occurrence of vertebrate material in amber".[6]

Cretaceous period

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In 2011, samples of amber were discovered to contain preserved feathers from the Cretaceous period, with evidence that they were from both dinosaurs and birds. Initial analysis suggests that some of the feathers were used for insulation, and not flight.[7][8]

Current knowledge

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List of dinosaurs with evidence of feathers

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Fossil of Sinornithosaurus, the first evidence of feathers in dromaeosaurids
 
Cast of a Caudipteryx fossil with feather impressions and stomach content
 
Fossil of Microraptor gui impressions of feathered wings
 
Fossil cast of NGMC 91, an unnamed species
 
Jinfengopteryx elegans fossil

Many non-avialan dinosaurs are now known to have been feathered. Direct evidence of feathers exists for the following genera, listed in the order of when the evidence was first published.

The evidence consists of feather impressions, or convincing skeletal or chemical evidence. Skeletal evidence would be the presence of quill knobs (the anchor points for wing feathers on the forelimb) or a pygostyle (the fused vertebrae at the tail tip which often supports large feathers).

For comparison: Archaeopteryx (1861; avialan)[9][10] definitely had working flight feathers with off-centre rachis (stem). Otherwise, apart from the front limbs, it was a typical small carnivorous dinosaur.

  1. Avimimus portentosus (inferred 1987: quill knobs)[11][12]
  2. Sinosauropteryx prima (1996)[13]
  3. Protarchaeopteryx robusta (1997)[14]
  4. GMV 2124 (1997).[15]
  5. Caudipteryx zoui (1998)[16]
  6. Shuvuuia deserti (1999)[1]
  7. Sinornithosaurus millenii (1999)[17]
  8. Beipiaosaurus inexpectus (1999)[18]
  9. Caudipteryx dongi (2000)[19]
  10. Caudipteryx sp. (2000)[20]
  11. Microraptor zhaoianus (2000)[21]
  12. Nomingia gobiensis (inferred 2000: pygostyle)[22]
  13. Psittacosaurus sp.? (2002)[23]
  14. Scansoriopteryx heilmanni (2002)[24]
  15. Yixianosaurus longimanus (2003)[25]
  16. Dilong paradoxus (2004)[26]
  17. Jinfengopteryx elegans (2005)[27][28]
  18. Juravenator starki (2006)[29][30]
  19. Sinocalliopteryx gigas (2007)[31]
  20. Velociraptor mongoliensis (inferred 2007: quill knobs)[2]
  21. Epidexipteryx hui (2008)[32]
  22. Similicaudipteryx yixianensis (inferred 2008: pygostyle; confirmed 2010)[33][34]
  23. Zhongornis haoae (2008)[35][36]
  24. Tianyulong confuciusi? (2009)[37]
  25. Concavenator corcovatus? (inferred 2010: quill knobs?)[38]
  26. Yutyrannus huali (2012)[39]
  27. Microraptor hanqingi (2012)[40]
  28. Ornithomimus edmontonicus (2012)[41]
  29. Ningyuansaurus wangi (2012)[42]
  30. Eosinopteryxbrevipenna (2013)[43]
  31. Citipati osmolskae (inferred 2013: pygostyle)[44]
  32. Conchoraptor gracilis (inferred 2013: pygostyle)[44]
  33. Jianchangosaurus yixianensis (2013)[45]
  34. Aurornis xui (2013; possibly avialan)[46]
  35. Changyuraptor yangi (2014)[47]
  36. Kulindadromeus zabaikalicus? (2014)[48]
  37. Citipati osmolskae (inferred 2014: pygostyle)[44]
  38. Conchoraptor gracilis (inferred 2014: pygostyle)[44]
  39. Deinocheirus mirificus (inferred 2014: pygostyle)[49]
  40. Yi qi (2015)[50]
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References

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  1. ↑ 1.0 1.1 Schweitzer, M.H.; et al. (1999). "Beta-keratin specific immunological reactivity in feather-like structures of the Cretaceous Alvarezsaurid, Shuvuuia deserti". Journal of Experimental Zoology. 285 (2): 146–57. doi:10.1002/(SICI)1097-010X(19990815)285:2<146::AID-JEZ7>3.0.CO;2-A. PMID 10440726.
  2. ↑ 2.0 2.1 Turner, Alan H.; Makovicky, Peter J.; Norell, Mark A. (2007). "Feather quill knobs in the dinosaur Velociraptor" (PDF). Science. 317 (5845): 1721. Bibcode:2007Sci...317.1721T. doi:10.1126/science.1145076. PMID 17885130. S2CID 11610649.
  3. ↑ Norell M.A.; et al. (1995). "A nesting dinosaur". Nature. 378 (6559): 774–776. Bibcode:1995Natur.378..774N. doi:10.1038/378774a0. S2CID 4245228.
  4. ↑ Hopp, Thomas J. & Orsen, Mark J. 2004. In Philip J. Currie, Eva B. Koppelhus and Martin A. Shugar (eds) Feathered Dragons: studies on the transition from dinosaurs to birds. Chapter 11: Dinosaur brooding behavior and the origin of flight feathers. Indiana University Press. Bloomington IN. USA.
  5. ↑ Lida Xing et al 2016. A feathered dinosaur tail with primitive plumage trapped in mid-Cretaceous amber. Current Biology. [1]
  6. ↑ 6.0 6.1 Rincon, Paul 2016. 'Beautiful' dinosaur tail found preserved in amber. BBC News Science & Environment. [2]
  7. ↑ CBC News: Dinosaur feathers found in Alberta amber
  8. ↑ BBC News: Dinosaur feather evolution trapped in Canadian amber
  9. ↑ von Meyer H. 1861. Archaeopteryx litographica (Vogel-Feder) und Pterodactylus von Solenhofen. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde. 1861: 678–679, plate V [Article in German] Fulltext at Google Books.
  10. ↑ Xu, Xing; You, Hailu; Du, Kai; Han, Fenglu (July 2011). "An Archaeopteryx-like theropod from China and the origin of Avialae". Nature. 475 (7357): 465–470. doi:10.1038/nature10288. ISSN 0028-0836. PMID 21796204. S2CID 205225790.
  11. ↑ 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]
  12. ↑ Chiappe L.M. and Witmer L.M. 2002. Mesozoic Birds: above the heads of dinosaurs. Berkeley: University of California Press. ISBN 0-520-20094-2
  13. ↑ Ji Q., Ji S. (1996). "On discovery of the earliest bird fossil in China and the origin of birds". Chinese Geology. 10 (233): 30–33.
  14. ↑ Ji, Q. and Ji, S. 1997. A Chinese archaeopterygian, Protarchaeopteryx gen. nov." Geological Science and Technology (Di Zhi Ke Ji), 238: 38-41. Translated By Will Downs Bilby Research Center Northern Arizona University January 2001
  15. ↑ Ji, Q.; Ji, S. (1997). "Advances in Sinosauropteryx research". Chinese Geology. 7: 30–32.
  16. ↑ Currie, Philip J.; et al. (1998). "Two feathered dinosaurs from northeastern China" (PDF). Nature. 393 (6687): 753–761. Bibcode:1998Natur.393..753Q. doi:10.1038/31635. S2CID 205001388.
  17. ↑ Wu, Xiao-Chun; et al. (1999). "A dromaeosaurid dinosaur with a filamentous integument from the Yixian Formation of China". Nature. 401 (6750): 262–266. Bibcode:1999Natur.401..262X. doi:10.1038/45769. S2CID 4430574.
  18. ↑ Xu, Xing; et al. (1999). "A therizinosauroid dinosaur with integumentary structures from China". Nature. 399 (6734): 350–354. Bibcode:1999Natur.399..350X. doi:10.1038/20670. S2CID 204993327.
  19. ↑ Zhou, Z.; et al. (2000). "A new species of Caudipteryx from the Yixian Formation of Liaoning, northeast China" (PDF). Vertebrata PalAsiatica. 38 (2): 113–130. Archived from the original (PDF) on 2007-10-07. Retrieved 2014-04-19.
  20. ↑ Zhou Z.; et al. (2000). "Important features of Caudipteryx - Evidence from two nearly complete new specimens" (PDF). Vertebrata PalAsiatica. 38 (4): 241–254.[permanent dead link]
  21. ↑ Xu, Xing; Zhou, Zhonghe; Wang, Xiaolin (2000). "The smallest known non-avian theropod dinosaur" (PDF). Nature. 408 (6813): 705–708. doi:10.1038/35047056. PMID 11130069. S2CID 4411157. Archived from the original (PDF) on 2008-12-17. Retrieved 2011-09-16.
  22. ↑ Barsbold R.; et al. (2000). "New Oviraptorosaur (Dinosauria, Theropoda) from Mongolia: the first dinosaur with a pygostyle" (PDF). Acta Palaeontologica Polonica. 45 (2): 97–106.
  23. ↑ Mayr, Gerald; et al. (2002). "Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus". Naturwissenschaften. 89 (8): 361–365. Bibcode:2002NW.....89..361M. doi:10.1007/s00114-002-0339-6. PMID 12435037. S2CID 17781405.
  24. ↑ Czerkas S.A. and Yuan C. 2002. An arboreal maniraptoran from northeast China. In Czerkas S.J. (ed) Feathered dinosaurs and the origin of flight, 63-95. The Dinosaur Museum Journal 1. The Dinosaur Museum, Blanding, U.S.A. PDF abridged version
  25. ↑ Xu X., Wang X.-L. (2003). "A new maniraptoran from the early Cretaceous Yixian Formation of western Liaoning". Vertebrata PalAsiatica. 41 (3): 195–202.
  26. ↑ Xu X.; et al. (2004). "Basal tyrannosauroids from China and evidence for protofeathers in tyrannosauroids". Nature. 431 (7009): 680–684. Bibcode:2004Natur.431..680X. doi:10.1038/nature02855. PMID 15470426. S2CID 4381777.
  27. ↑ Ji Q.; et al. (2005). "First avialan bird from China (Jinfengopteryx elegans gen. et sp. nov.)". Geological Bulletin of China. 24 (3): 197–205.
  28. ↑ Turner, Alan H.; et al. (2007). "A basal dromaeosaurid and size evolution preceding avian flight" (PDF). Science. 317 (5843): 1378–1381. Bibcode:2007Sci...317.1378T. doi:10.1126/science.1144066. PMID 17823350. S2CID 2519726.
  29. ↑ Goehlich U.B; Tischlinger H; Chiappe L.M (2006). "Juraventaor starki (Reptilia, Theropoda) ein nuer Raubdinosaurier aus dem Oberjura der Suedlichen Frankenalb (Sueddeutschland): Skelettanatomie und Wiechteilbefunde". Archaeopteryx. 24: 1–26.
  30. ↑ Chiappe, Luis M.; Göhlich, Ursula B. (2010). "Anatomy of Juravenator starki (Theropoda: Coelurosauria) from the Late Jurassic of Germany" (PDF). Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 258 (3): 257–296. doi:10.1127/0077-7749/2010/0125. S2CID 130486197.
  31. ↑ Ji S.; et al. (2007). "A new giant compsognathid dinosaur with long filamentous integuments from Lower Cretaceous of Northeastern China". Acta Geologica Sinica. 81 (1): 8–15.
  32. ↑ Zhang, Fucheng; et al. (2008). "A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers". Nature. 455 (7216): 1105–1108. doi:10.1038/nature07447. PMID 18948955. S2CID 4362560.
  33. ↑ 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.
  34. ↑ Xu, Xing; Zheng, Xiaoting; You, Hailu (2010). "Exceptional dinosaur fossils show ontogenetic development of early feathers". Nature. 464 (7293): 1338–1341. Bibcode:2010Natur.464.1338X. doi:10.1038/nature08965. PMID 20428169. S2CID 205220207.
  35. ↑ Gao, Cunling et al (2008) A new basal lineage Of early Cretaceous birds from China and its implications on the evolution of the avian tail. Palaeontology 51, Part 4, pp. 775-791. doi:10.1111/j.1475-4983.2008.00793.x
  36. ↑ Jingmai K. O'Connor and Corwin Sullivan (2014). "Reinterpretation of the early Cretaceous maniraptoran (Dinosauria: Theropoda) Zhongornis haoae as a scansoriopterygid-like non-avian, and morphological resemblances between scansoriopterygids and basal oviraptorosaurs" (PDF). Vertebrata PalAsiatica. 52 (1): 3–30.
  37. ↑ 37.0 37.1 Zheng, Xiao-Ting; et al. (2009). "An early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures". Nature. 458 (7236): 333–336. Bibcode:2009Natur.458..333Z. doi:10.1038/nature07856. PMID 19295609. S2CID 4423110.
  38. ↑ Ortega, Francisco; Escaso, Fernando; Sanz, José L. (2010). "A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain". Nature. 467 (7312): 203–206. Bibcode:2010Natur.467..203O. doi:10.1038/nature09181. PMID 20829793. S2CID 4395795.
  39. ↑ Xu, Xing; et al. (2012). "A gigantic feathered dinosaur from the Lower Cretaceous of China". Nature. 484 (7392): 92–95. Bibcode:2012Natur.484...92X. doi:10.1038/nature10906. PMID 22481363. S2CID 29689629.
  40. ↑ En-Pu Gong; et al. (2012). "A new species of Microraptor from the Jehol Biota of northeastern China". Palaeoworld. 21 (2): 81–91. doi:10.1016/j.palwor.2012.05.003.
  41. ↑ Darla K. Zelenitsky; et al. (2012). "Feathered non-avian dinosaurs from North America provide insight into wing origins". Science. 338 (6106): 510–514. Bibcode:2012Sci...338..510Z. doi:10.1126/science.1225376. PMID 23112330. S2CID 2057698.
  42. ↑ Ji Qiang; et al. (2012). "A new oviraptorosaur from the Yixian Formation of Jianchang, Western Liaoning Province, China". Regional Geology of China (12): 2102–2107.
  43. ↑ Pascal Godefroit; et al. (2013). "Reduced plumage and flight ability of a new Jurassic paravian theropod from China". Nature Communications. 4: Article number 1394. Bibcode:2013NatCo...4.1394G. doi:10.1038/ncomms2389. PMID 23340434. S2CID 205315142.
  44. ↑ 44.0 44.1 44.2 44.3 W. Scott Persons, IV; Philip J. Currie; Mark A. Norell (2013). "Oviraptorosaur tail forms and functions". Acta Palaeontologica Polonica. doi:10.4202/app.2012.0093. S2CID 54935594.
  45. ↑ Hanyong Pu; et al. (2013). "An unusual basal therizinosaur dinosaur with an ornithischian dental arrangement from northeastern China". PLOS ONE. 8 (5): Article number e63423. Bibcode:2013PLoSO...863423P. doi:10.1371/journal.pone.0063423. PMC 3667168. PMID 23734177.
  46. ↑ Pascal Godefroit; et al. (2013). "A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds". Nature. 498 (7454): 359–362. doi:10.1038/nature12168. PMID 23719374. S2CID 4364892.
  47. ↑ Gang Han; Chiappe; Ji; Habib; Turner; Chinsamy; Liu; Han; et al. (2014). "A new raptorial dinosaur with exceptionally long feathering provides insights into dromaeosaurid flight performance". Nature Communications. 5: Article number 4382. doi:10.1038/ncomms5382. PMID 25025742. S2CID 205327789.
  48. ↑ Pascal Godefroit; Sinitsa; Dhouailly; Bolotsky; Sizov; McNamara; Benton; Spagna; et al. (2014). "A Jurassic ornithischian dinosaur from Siberia with both feathers and scales". Science. 25 (345): 451–455. doi:10.1126/science.1253351. hdl:1983/a7ae6dfb-55bf-4ca4-bd8b-a5ea5f323103. PMID 25061209. S2CID 206556907.
  49. ↑ Yuong-Nam Lee; et al. (2014). "Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus". Nature. 515 (7526): 257–260. doi:10.1038/nature13874. PMID 25337880. S2CID 2986017.
  50. ↑ Xu, X. et al 2015. A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings. Nature. [3]
  51. ↑ Czerkas S.A., and Ji Q. 2002. A new rhamphorhynchoid with a headcrest and complex integumentary structures. Pp. 15-41 in: Czerkas, S.J. (Ed.). Feathered dinosaurs and the origin of flight. Blanding, Utah: The Dinosaur Museum. ISBN 1-932075-01-1.
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