Condamine, Fabien, Allio, Rémi, Reboud, Eliette, Dupuis, Julian, Toussaint, Emmanuel, Mazet, Nathan, Hu, Shao-Ji, Lewis, Delano, Kunte, Krushnamegh, Cotton, Adam, Sperling, Felix, Institut des Sciences de l'Evolution de Montpellier (UMR ISEM), Centre de Coopération Internationale en Recherche Agronomique pour le Développement (Cirad)-École Pratique des Hautes Études (EPHE), Université Paris sciences et lettres (PSL)-Université Paris sciences et lettres (PSL)-Institut de recherche pour le développement [IRD] : UR226-Centre National de la Recherche Scientifique (CNRS)-Université de Montpellier (UM), Centre de Biologie pour la Gestion des Populations (UMR CBGP), Centre de Coopération Internationale en Recherche Agronomique pour le Développement (Cirad)-Institut de Recherche pour le Développement (IRD [France-Sud])-Institut National de Recherche pour l’Agriculture, l’Alimentation et l’Environnement (INRAE)-Institut Agro Montpellier, Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro)-Institut national d'enseignement supérieur pour l'agriculture, l'alimentation et l'environnement (Institut Agro)-Université de Montpellier (UM), University of Kentucky (UK), Museum d'Histoire Naturelle [Genève] (MHN), Yunnan Agricultural University, Burman University, Tata Institute of Fundamental Research [Bangalore], National Centre for Biological Sciences [TIFR] (NCBS), Tata Institute for Fundamental Research (TIFR), Chercheur indépendant, University of Alberta, This project has received funding from the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (project GAIA, agreement no. 851188), and an ‘Investissement d’Avenir’ grant from the Agence Nationale de la Recherche (project CASMA, CEBA, ref. ANR-10-LABX-0025) to F.L.C., a Ramanujan Fellowship from the Dept. of Science and Technology, Govt. of India, to K.K., and support of the Dept. of Atomic Energy, Govt. of India, under project nos. 12-RandD-TFR-5.04-0800 and 12-RandD-TFR-5.04-0900 to TIFR/NCBS, a Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant to F.A.H.S. Sequencing of most Indian Papilio was done at NCBS under research and voucher specimen collection permits issued to K.K. by the state forest departments in Kerala (permit no. WL 10-3781/2012 dated 18/12/2012, and GO (RT) No. 376/2012/FandWLD dated 26/07/2012) and Nagaland (permit no. CWL/GEN/240/522-39 dated 14 August 2012). Alberto Fracasso (Italy) and Laurie Wills (New Zealand) are thanked for providing samples of some hard to obtain taxa., ANR-10-LABX-0025,CEBA,CEnter of the study of Biodiversity in Amazonia(2010), and European Project: 851188,GAIA
International audience; Highlights:• A dense and robust phylogeny of Papilio is reconstructed with seven DNA markers.• A taxonomic working list is proposed, including 235 extant species• Papilio originated in the Oligocene in a northern region centered on the Bering Land Bridge• The role of Paleotropics is revealed within the radiation of Old World Papilio• Synchronous southward dispersals explained the current tropical distribution of PapilioAbstract:The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide, and has broad morphological habits and ecological niches. Because of its elevated species richness, it has been historically difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with highly supported relationships within subgenera, although a few nodes in the early history of the Old World Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus, P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago (Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate further studies to address questions on their ecology and evolutionary biology using this model clade.