High elevation temperate mountains have long been considered species poor owing to high extinction or low speciation rates during the Pleistocene. We performed a phylogenetic and population genomic investigation of an emblematic high-elevation plant clade (Androsace sect. Aretia, 31 currently recognized species), based on plant surveys conducted during alpinism expeditions. We inferred that this clade originated in the Miocene and continued diversifying through Pleistocene glaciations, and discovered three novel species of Androsace dwelling on different bedrock types on the rooftops of the Alps. This highlights that temperate high mountains have been cradles of plant diversity even during the Pleistocene, with in-situ speciation driven by the combined action of geography and geology. Our findings have an unexpected historical relevance: H.-B. de Saussure likely observed one of these species during his 1788 expedition to the Mont Blanc and we describe it here, over two hundred years after its first sighting.

Center for Theoretical Study Charles University and the Academy of Sciences of the Czech Republic 110 00 Prague Czech Republic

Department of Plant Biology University of Vermont Burlington VT 05405 USA

Parc National Des Écrins Domaine de Charance Gap France

Systematics and Evolution of Vascular Plants Associated Unit To CSIC Departament de Biologia Animal Biologia Vegetal 1 Ecologia Facultat de Biociències Universitat Autònoma de Barcelona 08193 Bellaterra Spain

Univ Grenoble Alpes CNRS Sciences Po Grenoble Pacte 38000 Grenoble France

Univ Grenoble Alpes Univ Savoie Mont Blanc CNRS LECA 38000 Grenoble France

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