This study is part of a doctoral dissertation of the first author, developed at the Escuela Doctoral Franco Peruana en Ciencias de la Vida, Universidad Peruana Cayetano Heredia and École Doctorale GAIA Biodiversité, Agriculture, Alimentation, Environnement, Terre, Eau, Université de Montpellier, France. We thank the directors and curators of the herbaria listed in the Material & Methods for loan of material, providing digital images, and for providing working space during our visits. This project was supported by a Franco-Peruvian doctoral fellowship from the National Council for Science, Technology and Technological Innovation (Consejo Nacional de Ciencia, Tecnología e Innovación Tecnológica, CONCYTEC) of the Peruvian Government, and the French Embassy in Peru. The Institut de Recherche pour le Developpement (IRD) also supported this project by sponsoring field work in Peru throug...