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.).
The authors declare no competing financial interests.
The data matrix for the phylogenetic analysis has been deposited in MorphoBank (http://www.morphobank.org) as project 1126.
Extended data figures and tables
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.
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.
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.
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.
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.
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.
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Foth, C., Tischlinger, H. & Rauhut, O. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature 511, 79–82 (2014). https://doi.org/10.1038/nature13467
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