The COI phylogeny indicated that the basal Verrucarum series lineage probably diverged before the Townsendi and Serrana series. This is the first molecular evidence of the ancestry and ranking within species of the Verrucarum group. The phylogenetic positions of three series (Verrucarum, Serrana, and Townsendi: sensu Young and Duncan1) in the Verrucarum group have been positioned unambiguously and support the taxonomic reorganization by Galati4 based on morphology.