Evolutionary history of Neotropical butterflies of the infratribe Neosatyriti based on target enrichment (Lepidoptera: Nymphalidae, Satyrinae, Satyrini, Pronophilina)

Abstract

The infratribe Neosatyriti is a section of the entirely Neotropical subtribe Pronophilina, accountingfor 57 species distributed from southern Patagonia to the Andes of northern Peru, and along theAtlantic coast, with the highest diversity in central Chile. They are found in two main types ofhabitats, i.e. puna, pampas and subantarctic grasslands, and Valdivian forests and Chilean matorral,from sea level to nearly 5000 m above sea level. We propose a phylogenetic hypothesis of theinfratribe based on molecular data obtained by target enrichment (TE) of 621 nuclear loci, totalling248,373 base pairs, from 53 species of this infratribe and 12 outgroups. Our analysis confirms thatNeosatyriti is monophyletic with full support. Based on these results, we propose eight newcombinations and two status reinstatements. Molecular data are congruent with morphologicalcharacters except for Homoeonympha which appears to be paraphyletic, thus almost all the generadescribed originally by L. Herrera, K. Hayward and W. Heimlich in the Twentieth century based onlyon morphological characters are confirmed as valid, except for Haywardella that is synonymisedwith Pampasatyrus. Neosatyriti diverged from other Pronophilina some 23 Ma and split into twomajor lineages some 20 Ma. Our reconstructions do not indicate a single area of origin, rather amultisource origin, but they suggest the Neosatyriti originated in the lowlands, and that theirancestral plants were non-bambusoid grasses. Major divergence of the infratribe started some12–11 Ma when it split into the ‘Neomaenas clade’ (9 genera), the ‘Pampasatyrus clade’ (4 genera) and the ‘Neosatyrus clade’ (5 genera). The next main radiation took place some 7–5 Ma with the switch to bamboo host plants and the colonisation of Valdivian forests. The final dispersal of the tribe was associated with the colonisation of Mata Atlantica some 6–5 Ma, and high Andean puna in Peru in the Early Pleistocene.