The Phylogeny of Butterflies (Papolopnoidea and Hesperioidae)

A phylogenetic tree for butterflies and skippers is derived, using the common possession of derived traits to delimit monophyletic taxa. All available characters of larvae, pupae, adults, and behavior are used, including various new characters. The traits of the progenitor are deduced, and the exact character changes at each point of the tree are specified. The tree accepted is mostly similar to that of Ehrlich. However, within Nymphalidae, Apaturinae is elevated to a subfamily distinct from Charaxinae; and within Lycaenidae, Curetinae is elevated to a subfamily branching from the base of the Riodininae line, and Aphnaeini is elevated to tribal status; Megathyminae clearly belongs to the monocotyledon-feeding branch of Hesperiidae. Most characters support the origin of Pieridae from the Papilionid ancestral line, and the origin of Lycaenidae from the Nymphalid-Libytheid ancestral line. The few characters that seem to have been subject to state reversal, or evolved the same state independently repeatedly, are discussed.