Diversification of Morpho butterflies (Lepidoptera, Nymphalidae): a re-evaluation of morphological characters and new insight from DNA sequence data

This study compiles previously published morphological, colour and behavioural characters and includes new DNA sequence data for eight markers (one mitochondrial and seven nuclear) to re-evaluate phylogenetic relationships and estimate times of divergence for Morpho butterflies using parsimony and Bayesian methods. We note an effect of missing data on phylogenetic inference and calculations of Partitioned Bremer Support. Morphology and DNA trees were moderately congruent, and the combined analyses of all data included elements of both sources. Both morphology and DNA support the monophyly of Morpho and the early separation of the sister pair M. marcus plus M. eugenia, but trees from different data sources are congruent mostly at derived nodes, and differ at several internal nodes. The analyses of combined data indicate that Morpho is composed of four clades each of which include one or more previously proposed subgenera. The subgenera Pessonia and Morpho were not monophyletic, and to address this issue we propose that Pessonia, syn.nov. be subsumed within Morpho. The ancestor of Morpho probably arose during the Oligocene, and most diversification seems to have occurred during the late Miocene. S-DIVA analysis suggests eastern Andean region as the ancestral area for Morpho, and that the South American Atlantic Forest was colonized multiple times.