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

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-1α 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-1α 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

Exaggeration and suppression of iridescence: the evolution of two-dimensional butterfly structural colours

Many butterfly species possess ‘structural’ colour, where colour is due to optical microstructures found in the wing scales. A number of such structures have been identified in butterfly scales, including three variations on a simple multi-layer structure. In this study, we optically characterize examples of all three types of multi-layer structure, as found in 10 species. The optical mechanism of the suppression and exaggeration of the angle-dependent optical properties (iridescence) of these structures is described. In addition, we consider the phylogeny of the butterflies, and are thus able to relate the optical properties of the structures to their evolutionary development. By applying two different types of analysis, the mechanism of adaptation is addressed. A simple parsimony analysis, in which all evolutionary changes are given an equal weighting, suggests convergent evolution of one structure. A Dollo parsimony analysis, in which the evolutionary ‘cost’ of losing a structure is less than that of gaining it, implies that ‘latent’ structures can be reused

Morfologia externa de Thyridia psidii cetoides (Rosenberg & Talbot) (Lepidoptera, Nymphalidae, Ithomiinae). III. Abdome e apêndices External morphology of Thyridia psidii cetoides (Rosenberg & Talbot). III. Abdomen and appendages (Lepidoptera, Nymphalidae, Ithomiinae)

<abstract language="eng">A detailed study of the abdominal external morphology of both sexes of Thyridia psidii cetoides (Rosenberg & Talbot, 1914) is presented. The material for this research was obtained at the city's plant nursery "Horto Florestal de Curitiba", Paraná, Brazil; mainly by rearing eggs and larvae collected on Cyphomandra betacea (Canavilles) Sendtner, 1845 (Solanaceae). When possible, the results obtained were compared with those already available in the literature concerning other Nymphalidae subfamilies morphology (Brassolinae, Morphinae and Danainae); the most striking feature being the asymmetrical valvae of the male and the length and faint sclerotinization of the third abdominal sternite in both sexes. A resume containing the main morphological differences to other nymphalid subfamillies, found throughout this research concerning head, thorax and abdome, is presented

Morfologia externa do adulto de Actinote melanisans (Nymphalidae: Heliconiinae: Acraeini)

A morfologia externa do adulto de A. melanisans Oberthür, 1917 é descrita e ilustrada. Os resultados obtidos foram comparados com outros já publicados para a morfologia externa de Nymphalidae (Danainae, Brassolinae/Brassolini sensu CASAGRANDE (2004), Morphinae/Morphini sensu LAMAS (2004a), Ithomiinae e Charaxinae). É apresentado um padrão para a caracterização morfológica do gênero, sendo esse o primeiro estudo detalhado de morfologia de uma espécie de Acraeini, agregando informações úteis para subsidiar a análise filogenética das espécies Neotropicais.<br>The external morphology of adult A. melanisans Oberthür, 1917 is described and illustrated. The results obtained were compared with other studies already published on the external morphology of Nymphalidae - Danainae: Brassolinae: Brassolini sensu CASAGRANDE (2004), Morphinae: Morphini sensu LAMAS (2004a), Ithomiinae and Charaxinae. A pattern for the morphological characterization of the genus is presented. It is the first detailed study of the morphology of a species of Acraeini, therefore contributing with information on characters useful for the phylogenetic analysis of the Neotropical species

Estudo comparado da morfologia externa de Zaretis itys itylus (Westwood) e Agrias claudina annetta (Gray) (Lepidoptera, Nymphalidae, Charaxinae) I. Cabeça, apêndices e região cervical Comparative study of the external morphology of Zaretis itys itylus (Westwood) and Agrias claudina annetta (Gray) (Lepidoptera, Nymphalidae, Charaxinae). I. Head, appendages and cervical region

Realizou-se um estudo detalhado e comparado da morfologia externa da cabeça, apêndices e região cervical, de duas espécies de Charaxinae, Zaretis itys itylus (Westwood, 1850) e Agrias claudina annetta (Gray, 1832). Os resultados obtidos foram comparados com outros já publicados e relacionados com morfologia externa de outras subfamílias de Nymphalidae (Brassolinae, Morphinae, Danainae e Ithomiinae).<br>Two species of Charaxinae, Zaretis itys itylus (Westwood, 1850) and Agrias claudina annetta (Gray, 1832) were subject of a detailed and comparative study of external morphology of the head, appendages and cervical region. The results obtained were compared with other studies published and related to the external morphology of other Nymphalidae subfamilies (Brassolinae, Morphinae, Danainae and Ithomiinae)