Archaeopteryx is an iconic fossil taxon with feathered wings from the Late Jurassic of Germany that occupies a crucial position for understanding the early evolution of avian flight. After over 150 years of study, its mosaic anatomy unifying characters of both non-flying dinosaurs and flying birds has remained challenging to interpret in a locomotory context. Here, we compare new data from three Archaeopteryx specimens obtained through phase-contrast synchrotron microtomography to a representative sample of archosaurs employing a diverse array of locomotory strategies. Our analyses reveal that the architecture of Archaeopteryx's wing bones consistently exhibits a combination of cross-sectional geometric properties uniquely shared with volant birds, particularly those occasionally utilising short-distance flapping. We therefore interpret that Archaeopteryx actively employed wing flapping to take to the air through a more anterodorsally posteroventrally oriented flight stroke than used by modern birds. This unexpected outcome implies that avian powered flight must have originated before the latest Jurassic.

Bayerische Staatssammlung für Paläontologie und Geologie Richard Wagner Str 10 D 80333 München Germany

Bürgermeister Müller Museum Bahnhofstrasse 8 91807 Solnhofen Germany

CNRS Laboratoire d'éthologie animale et humaine Université de Rennes 1 Université de Caen Normandie 263 Avenue du Général Leclerc 35042 Rennes France

Department of Zoology and Laboratory of Ornithology Palacký University 17 listopadu 50 771 46 Olomouc Czech Republic

European Synchrotron Radiation Facility 71 Avenue des Martyrs CS 40220 38043 Grenoble Cedex France

Science for Life Laboratory and Uppsala University Subdepartment of Evolution and Development Department of Organismal Biology Evolutionary Biology Centre Norbyvägen 18A 752 36 Uppsala Sweden

Sorbonne Université CNRS INSU Institut des Sciences de la Terre Paris ISTeP UMR 7193 F 75005 Paris France

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