Revival of Palaeoptera—head characters support a monophyletic origin of Odonata and Ephemeroptera (Insecta)

The earliest branching event in winged insects, one of the core problems regarding early insect evolution, was addressed using characters of the head. The head is arguably one of the most complex body regions in insects and the phylogenetic information content of its features has been demonstrated. In contrast, the wings and other body parts related to the flight apparatus and sperm transmission are not useful in the context of this problem, as the outgroups (silverfish and bristletails) are wingless and transmit spermatophores externally. Therefore, they show profound differences in the organization of the postcephalic body, and assessment of homology and subsequent phylogenetic analysis of features of these body regions is extremely difficult. The core of this study is the investigation of head structures of representatives of the major clades of dragonflies. A detailed description of the head of Lestes virens is presented and was used as a starting point for the compilation of a character set and a character state matrix for the entire Dicondylia (winged insects + silverfish), with a main focus on the placement of dragonflies and consequently the basal branching event within winged insects. Our results indicate a sister‐group relationship between a clade Palaeoptera (dragonflies + mayflies) and the megadiverse monophyletic lineage Neoptera. We show that despite considerable structural similarity between the odonate and neopteran mandible, the muscle equipment in dragonflies is more plesiomorphic with respect to Dicondylia than previously known. Odonata and Ephemeroptera also share presumably derived features of the antenna, maxilla, and labial musculature. Parsimony analyses of the head data unambiguously support a clade Palaeoptera. © The Willi Hennig Society 2012.