Integrative taxonomy, phylogeny and distribution patterns of the diverse Neotropical cloud forest butterfly genus Perisama Doubleday (Nymphalidae, Biblidinae, Callicorini)

Phylogenetic relationships of the predominantly Andean species-rich nymphalid genus Perisama are explored based on five genetic markers and a total of 3719 bps, and morphological characters of the adults with an emphasis on male genitalia and wing colour patterns. Its monophyly is recovered by including the genera Orophila, which turns out polyphyletic, and Mesotaenia. Two main, highly supported clades are identified, one comprising P. bomplandii and P. oppelii, the other all the remaining species. It is estimated that Perisama and its sister genus Diaethria diverged in Miocene, whereas its main radiation occurred in later Pliocene and throughout Pleistocene and into Holocene, simultaneously with the final Andean uplift. Alpha-taxonomic revision of Perisama was made. Based on comprehensive taxon sampling it is shown that colour patterns are less stable on a geographic scale than previously considered, thus several subspecies were synonymized. An updated checklist is presented. Male genitalia confirm as a more reliable taxonomic character for specific identification of Perisama.