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



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.

Access options

Rent or Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Figure 1: Overview photograph of the 11th skeletal specimen of Archaeopteryx.
Figure 2: Details of the plumage of the 11th specimen of Archaeopteryx.
Figure 3: Evolution of pennaceous feathers in maniraptoran theropods.


  1. 1

    Wellnhofer, P. Archaeopteryx. Der Urvogel von Solnhofen (Dr Friedrich Pfeil, 2008)

    Google Scholar 

  2. 2

    Ji, Q., Currie, P. J., Norell, M. A. & Ji, S. Two feathered dinosaurs from northeastern China. Nature 393, 753–761 (1998)

    ADS  Article  Google Scholar 

  3. 3

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

    ADS  CAS  Article  Google Scholar 

  4. 4

    Hu, D., Hou, L., Zhang, L. & Xu, X. A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus. Nature 461, 640–643 (2009)

    ADS  CAS  Article  Google Scholar 

  5. 5

    Xu, X., You, H., Du, K. & Han, F. An Archaeopteryx-like theropod from China and the origin of Avialae. Nature 475, 465–470 (2011)

    CAS  Article  Google Scholar 

  6. 6

    Godefroit, P. et al. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498, 359–362 (2013)

    ADS  CAS  Article  Google Scholar 

  7. 7

    Prum, R. O. & Brush, A. H. The evolutionary origin and diversification of feathers. Q. Rev. Biol. 77, 261–295 (2002)

    Article  Google Scholar 

  8. 8

    Nudds, R. L. & Dyke, G. J. Narrow primary feathers rachises in Confuciusornis and Archaeopteryx suggest poor flight ability. Science 328, 887–889 (2010)

    ADS  CAS  Article  Google Scholar 

  9. 9

    Longrich, N. R., Vinther, J., Meng, Q., Li, Q. & Russell, A. P. Primitive wing feather arrangement in Archaeopteryx lithographica and Anchiornis huxleyi. Curr. Biol. 22, 1–6 (2012)

    Article  Google Scholar 

  10. 10

    Xu, X. & Guo, Y. The origin and early evolution of feathers: insights from recent paleontological and neontological data. Vertebrata PalAsiatica 47, 311–329 (2009)

    Google Scholar 

  11. 11

    Li, Q. et al. Plumage color patterns of an extinct dinosaur. Science 327, 1369–1372 (2010)

    ADS  CAS  Article  Google Scholar 

  12. 12

    Li, Q. et al. Reconstruction of Microraptor and the evolution of iridescent plumage. Science 335, 1215–1219 (2012)

    ADS  CAS  Article  Google Scholar 

  13. 13

    Longrich, N. R. Structure and function of hindlimb feathers in Archaeopteryx lithographica. Paleobiology 32, 417–431 (2006)

    Article  Google Scholar 

  14. 14

    Turner, A. H., Makovicky, P. J. & Norell, M. A. A review of dromaeosaurid systematics and paravian phylogeny. Bull. Am. Mus. Nat. Hist. 371, 1–206 (2012)

    Article  Google Scholar 

  15. 15

    Zheng, X. et al. Hind wings in basal birds and the evolution of leg feathers. Science 339, 1309–1312 (2013)

    ADS  CAS  Article  Google Scholar 

  16. 16

    Lucas, A. M. & Stettenheim, P. R. Avian Anatomy. Integument, Parts I and II 197–233 (U.S. Government Printing Office, 1972)

    Google Scholar 

  17. 17

    Gatesy, S. M. & Dial, K. P. From frond to fan: Archaeopteryx and the evolution of short-tailed birds. Evolution 50, 2037–2048 (1996)

    Article  Google Scholar 

  18. 18

    Meseguer, J. et al. Lift devices in the flight of Archaeopteryx. Span. J. Palaeontol. 27, 125–130 (2012)

    Google Scholar 

  19. 19

    Foth, C. On the identification of feather structures in stem-line representatives of birds: evidence from fossils and actuopalaeontology. Paläontol. Z. 86, 91–102 (2012)

    Article  Google Scholar 

  20. 20

    Xu, X., Zheng, X. & You, H. Exceptional dinosaur fossils show ontogenetic development of early feathers. Nature 464, 1338–1341 (2010)

    ADS  CAS  Article  Google Scholar 

  21. 21

    Nachtigall, W. & Kempf, B. Vergleichende Untersuchungen zur flugbiologischen Funktion des Daumenfittichs (Alula spuria) bei Vögeln. Z. Vgl. Physiol. 71, 326–341 (1971)

    Article  Google Scholar 

  22. 22

    Chiappe, L. M. & Dyke, G. J. The Mesozoic radiation of birds. Annu. Rev. Ecol. Syst. 33, 91–124 (2002)

    Article  Google Scholar 

  23. 23

    Ji, Q., Norell, M. A., Gao, K., Ji, S. & Ren, D. The distribution of integumentary structures in a feathered dinosaur. Nature 410, 1084–1088 (2001)

    ADS  CAS  Article  Google Scholar 

  24. 24

    Xu, X. & Zhang, F. A new maniraptoran dinosaur from China with long feathers on the metatarsus. Naturwissenschaften 92, 173–177 (2005)

    ADS  CAS  Article  Google Scholar 

  25. 25

    O’Connor, J. et al. Unique caudal plumage of Jeholornis and complex tail evolution in early birds. Proc. Natl Acad. Sci. USA 110, 17404–17408 (2013)

    ADS  Article  Google Scholar 

  26. 26

    Ji, Q. et al. First avialian bird from China. Geol. Bull. China 24, 197–210 (2005)

    Google Scholar 

  27. 27

    Godefroit, P. et al. Reduced plumage and flight ability of a new Jurassic paravian theropod from China. Nature Commun. 4, 1–6 (2013)

    Article  Google Scholar 

  28. 28

    Chiappe, L. M., Marugán-Lobón, J., Ji, S. & Zhou, Z. Life history of a basal bird: morphometrics of the Early Cretaceous Confuciusornis. Biol. Lett. 4, 719–723 (2008)

    Article  Google Scholar 

  29. 29

    O’Connor, J. K., Chiappe, L. M., Chuong, C., Bottjer, D. J. & You, H. Homology and potential cellular and molecular mechanisms for the development of unique feather morphologies in early birds. Geosciences 2, 157–177 (2012)

    ADS  Article  Google Scholar 

  30. 30

    Chiappe, L. M. et al. A new specimen of the Early Cretaceous bird Hongshanornis longicresta: insights into the aerodynamics and diet of a basal ornithuromorph. PeerJ 2, e234 (2014)

    Article  Google Scholar 

Download references


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




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.

Corresponding author

Correspondence to Oliver W. M. Rauhut.

Ethics declarations

Competing interests

The authors declare no competing financial interests.

Additional information

The data matrix for the phylogenetic analysis has been deposited in MorphoBank ( as project 1126.

Extended data figures and tables

Extended Data Figure 1 The 11th specimen of Archaeopteryx under ultraviolet light.

a, Overview photograph of the 11th skeletal specimen of Archaeopteryx. Scale bar, 5 cm. b, Presacral vertebral column. Scale bar, 2 cm. c, Right forelimb. Scale bar, 5 cm.

Extended Data Figure 2 Anatomical details of the 11th skeletal specimen of Archaeopteryx.

a, Preserved cranial remains under ultraviolet light. Scale bar, 1 cm. b, Enlargement of the tooth row of the left premaxilla in medial view under normal light. c, Pelvis under normal light. Arrow points to the impression of the (broken) ‘intermediate process’ of the ischium. Scale bar, 1 cm. Abbreviations: ar, articular; il, ilium; in, incision in posterior end of jugal; is, ischium; ld, left dentary; lf, left femur; lj, left jugal; lpm, left premaxilla; pu, pubis; qjp, quadratojugal process of the jugal; rd, right dentary; rpm, right premaxilla; sa, surangular; sp, splenial.

Extended Data Figure 3 Overview the plumage in the 11th specimen of Archaeopteryx.

Colour code: yellow, body feathers from different body regions, which cannot be assigned to a certain body region; light blue, heckle feathers; sky blue, body feathers; dark blue, tibial feathers; red, femoral feathers; black, metatarsal feathers; light green, rectrices; green, remiges. Scale bar, 5 cm.

Extended Data Figure 4 Details of the plumage of the 11th specimen of Archaeopteryx.

a, Primary wing feathers under ultraviolet light. b, First and second primary and first secondary wing feathers under ultraviolet light. c, Close up of the primary feathers showing the rhachides and the barbs. d, Close up of the metatarsal feathers. Scale bar, 1 cm.

Extended Data Figure 5 Wing morphology of the 11th specimen of Archaeopteryx in dorsal view.

a, Rhachides of the primaries shown as yellow dashed lines. b, Imprints of the dorsal coverts marked by yellow arrows. Scale bars, 1 cm.

Extended Data Figure 6 The relative rhachis diameter of the primaries of Archaeopteryx and modern birds in relation to body mass.

a, Measurements of primaries of the 11th specimen of Archaeopteryx. b, Plot of the ratio of rhachis diameter to feather length against body weight (both log transformed). Yellow dots represent feathers of the 11th specimen of Archaeopteryx; black dots represent data of modern birds; red rectangles represent data of Confuciusornithidae. See Supplementary Information for details.

Extended Data Figure 7 Phylogenetic hypothesis used for tracing plumage characters.

a, Strict consensus tree of 3,020 most parsimonious trees (tree length = 2,612; consistency index = 0.275; retention index = 0.730). b, Reduced consensus tree of the pruned data matrix after the exclusion of Shenzhousaurus, Segnosaurus, Erliansaurus, Albinykus, Saurornitholestes, Harygryphus, Tianyuraptor, Hesperonychus, Pyroraptor, Lithornis, Liaoningornis and Limneavis.

Extended Data Table 1 Measurements of the 11th specimen of Archaeopteryx

Supplementary information

Supplementary Information

This file contains Supplementary Text 1-6, Supplementary Tables 1-2, a list of Characters 1-356 and additional References. (PDF 1909 kb)

PowerPoint slides

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Foth, C., Tischlinger, H. & Rauhut, O. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature 511, 79–82 (2014).

Download citation

Further reading


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Sign up for the Nature Briefing newsletter for a daily update on COVID-19 science.
Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing