Reassessment of butterfly family relationships using independent genes and morphology

Evolutionary relationships of true butterflies (Papilionoidea) are addressed using sequence data from nuclear, 28S ribosomal RNA, and the mitochondrial, protein-coding gene, ND1. We include a reanalysis of morphological characters from the literature to obtain an independent estimate of phylogeny. Neither morphology nor sequence data produce a single tree, and their results disagree on the relationships of Pieridae, Lycaenidae, and Riodinidae. The total evidence analysis corroborates Kristensen's hypothesis of butterfly family evolution Lycaenids and riodinids form a clade and are placed as the sister to Nymphalidae. Pieridae is placed as the sister taxon to this apical clade and not as the sister to Papilionidae. Nymphalid subfamily relationships remain enigmatic, particularly the placement of Ithomiinae and Libytheinae.