Phylogenetic position and taxonomic rearrangement of Davidina (Lepidoptera: Nymphalidae), an enigmatic butterfly genus new for Europe and America

Davidina, an enigmatic butterfly genus described from China in the 19th century, has for a long time been considered a member of the family Pieridae due to its pierid-like wing pattern. In the 20th century, it was transferred to Satyridae (now subfamily Satyrinae of Nymphalidae) based on analysis of the structure of genitalia and placed next to the species-rich genus Oeneis (subtribe Satyrina), being separated from the latter by supposed differences in wing venation. We have conducted a phylogenetic and taxonomic study of the subtribe Satyrina using analysis of molecular and morphological characters. We show that the genus Oeneis is not monophyletic, and consists of two genetically diverged and morphologically differentiated groups that are not sister-groups (Oeneis s.a and Protoeneis). We also demonstrate that Davidina is closely related to Protoeneis, but not to Oeneis s.s. To resolve this newly discovered non-monophyly and morphological heterogeneity, several species should be removed from Oeneis and transferred to the genus Davidina. As a consequence, we synonymize the name Protoeneis Gorbunov, 2001 with Davidina Oberthur, 1879. We conclude that Davidina is not a monotypic Chinese endemic genus, as has been previously supposed, but is composed of nine species that have a broad distribution area across the Holarctic region, extending to Europe and America.