New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers

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Discoveries of bird-like theropod dinosaurs and basal avialans in recent decades have helped to put the iconic ‘Urvogel’ Archaeopteryx1 into context2,3,4,5,6 and have yielded important new data on the origin and early evolution of feathers7. However, the biological context under which pennaceous feathers evolved is still debated. Here we describe a new specimen of Archaeopteryx with extensive feather preservation, not only on the wings and tail, but also on the body and legs. The new specimen shows that the entire body was covered in pennaceous feathers, and that the hindlimbs had long, symmetrical feathers along the tibiotarsus but short feathers on the tarsometatarsus. Furthermore, the wing plumage demonstrates that several recent interpretations8,9 are problematic. An analysis of the phylogenetic distribution of pennaceous feathers on the tail, hindlimb and arms of advanced maniraptorans and basal avialans strongly indicates that these structures evolved in a functional context other than flight, most probably in relation to display, as suggested by some previous studies10,11,12. Pennaceous feathers thus represented an exaptation and were later, in several lineages and following different patterns, recruited for aerodynamic functions. This indicates that the origin of flight in avialans was more complex than previously thought and might have involved several convergent achievements of aerial abilities.

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We thank B. Pohl for making the specimen available for study, C. Keilmann and R. Albersdörfer for assistance, Xu X. for access to material, and R. Carney, G. Mayr and A. López-Arbarello for discussions. This study was supported by the Volkswagen Foundation under grant I/84 640 (to O.W.M.R.).

Author information


  1. Staatliche Naturwissenschaftliche Sammlungen Bayerns, Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Straße 10, 80333 Munich, Germany

    • Christian Foth
    •  & Oliver W. M. Rauhut
  2. Department of Earth and Environmental Sciences and GeoBioCenter, Ludwig-Maximilians-Universität München, Richard-Wagner-Straße 10, 80333 Munich, Germany

    • Christian Foth
    •  & Oliver W. M. Rauhut
  3. Tannenweg 16, 85134 Stammham, Germany

    • Helmut Tischlinger


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C.F. and O.W.M.R. designed the study, collected and analysed the data and wrote the paper; H.T. did all the photography and helped with discussions.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Oliver W. M. Rauhut.

The data matrix for the phylogenetic analysis has been deposited in MorphoBank (http://www.morphobank.org) as project 1126.

Extended data

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains Supplementary Text 1-6, Supplementary Tables 1-2, a list of Characters 1-356 and additional References.


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