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AbstractPowered flight evolved independently in vertebrates in the pterosaurs, birds and bats, each of which has a different configuration of the bony elements and epidermal structures that form the wings1,2. Whereas the early fossil records of pterosaurs and bats are sparse, mounting evidence (primarily from China) of feathered non-avian dinosaurs and stemward avians that derive primarily from the Middle–Upper Jurassic and Lower Cretaceous periods has enabled the slow piecing together of the origins of avian flight3,4. These fossils demonstrate that, close to the origin of flight, dinosaurs closely related to birds were experimenting with a diversity of wing structures3,5. One of the most surprising of these is that of the scansoriopterygid (Theropoda, Maniraptora) Yi qi, which has membranous wings—a flight apparatus that was previously unknown among theropods but that is used by both the pterosaur and bat lineages6. This observation was not universally accepted7. Here we describe a newly identified scansoriopterygid—which we name Ambopteryx longibrachium, gen. et sp. nov.—from the Upper Jurassic period. This specimen provides support for the widespread existence of membranous wings and the styliform element in the Scansoriopterygidae, as well as evidence for the diet of this enigmatic theropod clade. Our analyses show that marked changes in wing architecture evolved near the split between the Scansoriopterygidae and the avian lineage, as the two clades travelled along very different paths to becoming volant. The membranous wings supported by elongate forelimbs that are present in scansoriopterygids probably represent a short-lived experimentation with volant behaviour, and feathered wings were ultimately favoured during the later evolution of Paraves. Access optionsAccess options Subscribe to JournalGet full journal access for 1 year227,39 €only 4,46 € per issueAll prices include VAT for Germany.Rent or Buy articleGet time limited or full article access on ReadCube.from$8.99All prices are NET prices. Data availabilityAll data—including the measurements, source data for morphometric analysis and phylogenetic data matrix—that support the findings of this research are included as Supplementary Information. The specimen (IVPP V24192) described in this study is archived and available on request from the IVPP. A Life Science Identifier for the newly described species has been registered at ZooBank (http://zoobank.org/): urn:lsid:zoobank.org:act:0A2DE2F0-CE78-4149-B0BD-A0DE91FC1328. 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Author informationAffiliationsKey Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, ChinaMin Wang, Jingmai K. O’Connor, Xing Xu & Zhonghe ZhouCenter for Excellence in Life and Paleoenvironment, Chinese Academy of Science, Beijing, ChinaMin Wang, Jingmai K. O’Connor, Xing Xu & Zhonghe ZhouAuthorsSearch for Min Wang in:Search for Jingmai K. O’Connor in:Search for Xing Xu in:Search for Zhonghe Zhou in:ContributionsZ.Z. and M.W. designed the research project; Z.Z. and M.W. conducted the fieldwork; M.W. performed the phylogenetic, histological and phylogenetic principal component analyses; and M.W., J.K.O., X.X. and Z.Z. wrote the manuscript. Competing interests The authors declare no competing interests. Corresponding authorCorrespondence to Min Wang.Extended data figures and tablesExtended Data Fig. 1 Additional photographs of Ambopteryx, IVPP V24192.a, Counter slab. b, Skeletal reconstruction based on preserved bones. c, Skull. d, Gastroliths and the unidentified bony stomach content. Abbreviations as in Fig. 1, except for: fe, feather associated with the neck; lil, left ilium;; lti, left tibia; pd, pedal digits; and ub, unidentified bony element. The white box indicates the position from which the sample was taken for histological analysis. Scale bars, 10 mm (a, c, d), 20 mm (b).Extended Data Fig. 2 Bone histology of Ambopteryx.a–d, Thin cross-section of the left humerus (a, b) and the unidentified bony stomach content (c, d). The arrowheads indicate the osteocyte lacunae. Scale bars, 100 μm (a–c), 200 μm (d).Extended Data Fig. 3 Anatomy of Ambopteryx, IVPP V24192.a–d, Interpretative drawings of the neck and pectoral girdle (a), caudal vertebrae and pygostyle (b), left forelimb (c) and pelvis and left hindlimb (d). Abbreviations as in Figs. 1, 2, except for: ca, caudal vertebrae; cm, calcaneum; de, deltopectoral crest of humerus; do, dorsal vertebrae; I–III, metacarpals I–III; ip, iliac peduncle of ilium; mtII–IV, metatarsals II–IV; p1–4, pedal digits I to IV; and st, styliform element. Scale bars, 10 mm (a–d).Extended Data Fig. 4 Forelimb comparisons between Ambopteryx and Yi.a, Left forelimb of Ambopteryx (IVPP V24192). b–d, Left forelimb (b), right humerus (c) and the right styliform element (d) of Yi (STM 31-2). The proximal margins of the humeri are marked in white dashed lines to show the differences between these two taxa. Abbreviations as in Figs. 1, 2, except for: st, styliform element. Scale bars, 10 mm (a), 20 mm (b–d).Extended Data Fig. 5 Comparisons of hand morphology among Scansoriopterygidae.a–c, Line drawings of the hands of Ambopteryx (a), Epidendrosaurus (b) and Yi (c). Scale bars, 10 mm (a, b), 20 mm (c).Extended Data Fig. 6 Scanning electron microscopy photographs of the soft tissues that are preserved in Ambopteryx.a–d, Feather samples associated with the neck. e–f, Samples of membranous tissues taken from the area between the left femur and left manual digits. Arrows denote the positions of the samples. Scale bars, 2 μm.Extended Data Fig. 7 Time-scaled recovered strict-consensus tree of Mesozoic coelurosaurians.Bremer and bootstrap values are labelled near the corresponding node in bold italic and upright non-bold font, respectively.Extended Data Fig. 8 Compiled super-tree of the sampled Mesozoic coelurosaurians used in morphometric analyses.Complete tree for the coelurosaurians used in generating the PPCA morphospaces shown in Fig. 3b, c.Supplementary informationSupplementary InformationThis file contains Supplementary Text Sections 1–5; which include additional anatomical description, stomach contents and diet of Ambopteryx longibrachium, Supplementary Tables 1, 2, 4 and 5, and the data used in the phylogenetic analysis.Reporting SummarySupplementary Table 3This file contains appendicular limb bone measurements of Mesozoic coelurosaurians used in the phylogenetic principal components analysis.Rights and permissionsTo obtain permission to re-use content from this article visit RightsLink. 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