Letter | Published:

A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings

Nature volume 521, pages 7073 (07 May 2015) | Download Citation



The wings of birds and their closest theropod relatives share a uniform fundamental architecture, with pinnate flight feathers as the key component1,2,3. Here we report a new scansoriopterygid theropod, Yi qi gen. et sp. nov., based on a new specimen from the Middle–Upper Jurassic period Tiaojishan Formation of Hebei Province, China4. Yi is nested phylogenetically among winged theropods but has large stiff filamentous feathers of an unusual type on both the forelimb and hindlimb. However, the filamentous feathers of Yi resemble pinnate feathers in bearing morphologically diverse melanosomes5. Most surprisingly, Yi has a long rod-like bone extending from each wrist, and patches of membranous tissue preserved between the rod-like bones and the manual digits. Analogous features are unknown in any dinosaur but occur in various flying and gliding tetrapods6,7,8,9,10, suggesting the intriguing possibility that Yi had membranous aerodynamic surfaces totally different from the archetypal feathered wings of birds and their closest relatives. Documentation of the unique forelimbs of Yi greatly increases the morphological disparity known to exist among dinosaurs, and highlights the extraordinary breadth and richness of the evolutionary experimentation that took place close to the origin of birds.

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  1. 1.

    , , & Two feathered dinosaurs from northeastern China. Nature 393, 753–761 (1998)

  2. 2.

    et al. Four-winged dinosaurs from China. Nature 421, 335–340 (2003)

  3. 3.

    , , & A pre-Archaeopteryx troodontid from China with long feathers on the metatarsus. Nature 461, 640–643 (2009)

  4. 4.

    et al. Jurassic System in the North of China (VII): The Stratigraphic Region of Northeast China (Petroleum Industry Press, 2003)

  5. 5.

    et al. Melanosome evolution indicates a key physiological shift within feathered dinosaurs. Nature 507, 350–353 (2014)

  6. 6.

    & Morphology and homology of the chiropteran calcar, with comments on the phylogenetic relationships of Archaeopteropus. J. Mamm. Evol. 5, 1–32 (1998)

  7. 7.

    & Wing tip anatomy and aerodynamics in flying squirrels. J. Mamm. 79, 245–250 (1998)

  8. 8.

    Articulation and function of the pteroid bone of pterosaurs. J. Vertebr. Paleontol. 27, 881–891 (2007)

  9. 9.

    , & Oldest fossil record of gliding in rodents. Nature 379, 439–441 (1996)

  10. 10.

    , , & Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation. Nature 451, 818–821 (2008)

  11. 11.

    et al. The vertebrates of the Jurassic Daohugou Biota of northeastern China. J. Vertebr. Paleontol. 34, 243–280 (2014)

  12. 12.

    Axial skeletal ontogeny in the Parasuchia (Archosauria: Pseudosuchia) and its implications for ontogenetic determination in Archosaurs. J. Vertebr. Paleontol. 27, 350–361 (2007)

  13. 13.

    & Mass prediction in theropod dinosaurs. Hist. Biol. 16, 85–92 (2004)

  14. 14.

    , , , & A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455, 1105–1108 (2008)

  15. 15.

    & Reinterpretation of the Early Cretaceous maniraptoran (Dinosauria: Theropoda) Zhongornis haonae as a scansoriopterygid-like non-avian, and morphological resemblances between scansoriopterygids and basal oviraptorosaurs. Vertebr. Palasiat. 52, 3–30 (2014)

  16. 16.

    , , & A juvenile coelurosaurian theropod from China indicates arboreal habits. Naturwissenschaften 89, 394–398 (2002)

  17. 17.

    & in Feathered Dinosaurs and the Origin of Flight (ed. Czerkas, S. J.) 63–95 (The Dinosaur Museum, 2002)

  18. 18.

    Myology of the gliding membranes of some Petauristine rodents (genera: Glaucomys, Pteromys, Petinomys, and Petaurista). J. Mamm. 58, 374–384 (1977)

  19. 19.

    Glide angle in the genus Petaurus and a review of gliding mammals. Mammal Rev. 30, 9–30 (2000)

  20. 20.

    Bird origins anew. Auk 130, 1–12 (2013)

  21. 21.

    et al. Melanosomes or microbes: testing an alternative hypothesis for the origin of microbodies in fossil feathers. Sci. Rep. 4, 4233 (2014)

  22. 22.

    & Jurassic archosaur is a non-dinosaurian bird. J. Ornithol. 155, 841–851 (2014)

  23. 23.

    , & New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature 511, 79–82 (2014)

  24. 24.

    in Functional Morphology in Vertebrate Paleontology (ed. Thomason, J. J.) 19–33 (Cambridge Univ. Press, 1995)

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We thank Z. Zhou, R. Dudley, J. Clarke and T. Stidham for discussion, X. Ding for specimen preparation, Y. Liu and Y. Han for providing illustrations, Z. Zhang for assistance with the EDS analysis, and Y. Hou for CT scanning the specimen. This research was funded by the National Natural Science Foundation of China (41372014, 41472023, 41120124002 and 41125008) and Major Basic Research Projects of the Ministry of Science and Technology, China (2012CB821900).

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    • Xing Xu
    •  & Xiaoting Zheng

    These authors contributed equally to this work.


  1. Institute of Geology and Paleontology, Linyi University, Linyi City, Shandong 276005, China

    • Xing Xu
    • , Xiaoting Zheng
    • , Xiaoli Wang
    •  & Yan Wang
  2. Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China

    • Xing Xu
    • , Corwin Sullivan
    • , Jingmai K. O’Connor
    •  & Fucheng Zhang
  3. Shandong Tianyu Museum of Nature, Pingyi, Shandong 273300, China

    • Xiaoting Zheng
    •  & Xiaomei Zhang
  4. School of the Earth Sciences and Resources, China University of Geosciences, Beijing 100083, China

    • Lida Xing
  5. Key Laboratory of Economic Stratigraphy and Palaeogeography of Chinese Academy of Sciences, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China

    • Yanhong Pan


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X.X., X. Zheng and C.S. designed the project, X.X., X. Zheng, C.S., X.W., Y.W., L.X., X. Zhang, J.K.O’C., F.Z. and Y.P. performed the research, and X.X. and C.S. wrote the manuscript.

Competing interests

The authors declare no competing financial interests.

Corresponding authors

Correspondence to Xing Xu or Xiaoting Zheng.

Data have been deposited in ZooBank under Life Science Identifier (LSID) http://zoobank.org/urn:lsid:zoobank.org:pub:5B765FD5-A534-45D5-AE14-62C25EADDD2D. The raw CT data have been deposited in the Dryad Digital Repository (http://dx.doi.org/10.5061/dryad.fp060).

Extended data

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  1. 1.

    Supplementary Information

    This file contains Supplementary Text and Data, Supplementary Tables 1-2 and Supplementary References.


  1. 1.

    CT data for holotype of Yi qi

    This data archive contains CT slice images in .dcm format for the slab containing the holotype specimen (STM, Shandong Tianyu Museum, 31-2) of Yi qi, a scansoriopterygid dinosaur from the Middle-Upper Jurassic of Mutoudeng, Hebei Province, China. The CT data reveal the 3D structure of the specimen, and of the slab itself.

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