Phylogeny and systematics of the "Pronophila clade," with 2 new genera to resolve the formerly polyphyletic genus Pseudomaniola (Lepidoptera: Nymphalidae: Satyrinae)

Analysis of a target enrichment molecular dataset confirms the monophyly of the Neotropical montane butterfly group known as the Pronophila Westwood clade, 1 of 2 major lineages of the satyrine subtribe Pronophilina. The Pronophila clade comprises 18-20 recognized genera and some 125 species. Within this group, the genus Pseudomaniola Rober appears as paraphyletic, and is split here into 3 genera, Pseudomaniola sensu novum with 6 species, including 4 previously considered as subspecies of P. phaselis (Hewitson), the monobasic Fahraeusia Pyrcz n. gen. for Catargynnis asuba Thieme, n. comb., and Boyeriana Pyrcz, Espeland & Willmott n. gen., with 9 species. The adults of all 3 genera can be recognized by their wing color patterns, but the strongest synapomorphies are found in the genitalia, especially those of the male, supporting the above systematic decisions. Notable differences are also found in scale organization and morphology. A divergence time analysis suggests that Fahraeusia diverged from Pseudomaniola + Boyeriana in the mid-Miocene, around 12 Mya, and the subsequent separation of the last 2 genera occurred at the start of the Pliocene at around 5 Mya.