Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers

Abstract

Phylogenetic relationships among major clades of butterflies and skippers have long been controversial, with no general consensus even today. Such lack of resolution is a substantial impediment to using the otherwise well studied butterflies as a model group in biology. Here we report the results of a combined analysis of DNA sequences from three genes and a morphological data matrix for 57 taxa (3258 characters, 1290 parsimony informative) representing all major lineages from the three putative butterfly super-families (Hedyloidea, Hesperioidea and Papilionoidea), plus out-groups representing other ditrysian Lepidoptera families. Recently, the utility of morphological data as a source of phylogenetic evidence has been debated. We present the first well supported phylogenetic hypothesis for the butterflies and skippers based on a total-evidence analysis of both traditional morphological characters and new molecular characters from three gene regions (COI, EF-1alpha and wingless). All four data partitions show substantial hidden support for the deeper nodes, which emerges only in a combined analysis in which the addition of morphological data plays a crucial role. With the exception of Nymphalidae, the traditionally recognized families are found to be strongly supported monophyletic clades with the following relationships: (Hesperiidae+(Papilionidae+(Pieridae+(Nymphalidae+(Lycaenidae+Riodinidae))))). Nymphalidae is recovered as a monophyletic clade but this clade does not have strong support. Lycaenidae and Riodinidae are sister groups with strong support and we suggest that the latter be given family rank. The position of Pieridae as the sister taxon to nymphalids, lycaenids and riodinids is supported by morphology and the EF-1alpha data but conflicted by the COI and wingless data. Hedylidae are more likely to be related to butterflies and skippers than geometrid moths and appear to be the sister group to Papilionoidea+Hesperioidea.

Light on the moth-eye corneal nipple array of butterflies

“…The most likely evolutionary scenario for the corneal nipple arrays of butterflies is that the diurnal butterflies descended from nocturnal moths (Yack & Fullard 2000;Grimaldi & Engel 2005;Wahlberg et al 2005). Most nymphalids, considered to be the least evolved butterflies, thus have retained the full-grown nipples of the moths, but the highly developed papilionids have completely lost the nipple trait.…”

Beauty in the eye of the beholder: the two blue opsins of lycaenid butterflies and the opsin gene-driven evolution of sexually dimorphic eyes

“…Within the true butterflies (Papilionoidea) the current understanding of familial relationships is (Papilionidae+(Pieridae+(Nymphalidae+(Riodinidae+Lycaeni dae)))) (Wahlberg et al, 2005), where papilionid and pierid butterflies represent the most basal lineages, and riodinid and lycaenid the most derived. Because the two pairs of B opsin genes identified from pierid (Pieris rapae) (Arikawa et al, 2005) and lycaenid butterflies (this study) are from distantly related families, we were interested in testing the hypothesis that the blue opsin genes of pierids and lycaenids evolved independently.…”

From the Phylogeny of the Satyrinae Butterflies to the Systematics of Euptychiina (Lepidoptera: Nymphalidae): History, Progress and Prospects

“…These studies yielded similar results with morphological characters of adults and immatures (Freitas & Brown 2004) and with DNA sequences of mitochondrial (COI) and nuclear (EF-1α and wingless) (Brower 2000, Wahlberg et al 2003 genes, inding a "satyroid" clade made up of Charaxinae, Calinaginae, Satyrinae and Morphinae. Also, combining morphological characters of adults and molecular data provides further support for this clade (Wahlberg et al 2005), but its relationship to the rest of Nymphalidae remains unresolved.…”