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Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing

We undertook a comprehensive morphological and molecular phylogenetic analysis of dragonfly phylogeny, examining both extant and fossil lineages in simultaneous analyses. The legitimacy of higher‐level family groups and the phylogenetic relationship between families were tested. Thirteen families were supported as monophyletic (Aeshnidae, Calopterygidae, Chlorocyphidae, Euphaeidae, Gomphidae, Isostictidae, Lestidae, Libellulidae, Petaluridae, Platystictidae, Polythoridae, Pseudostigmatidae and Synthemistidae) and eight as non‐monophyletic (Amphipterygidae, Coenagrionidae, Corduliidae, Megapodagrionidae, Protoneuridae and Synlestidae), although Perilestidae and Platycnemididae were recovered as monophyletic under Bayesian analyses. Nine families were represented by one species, thus monophyly was not tested (Epiophlebiidae, Austropetaliidae, Chlorogomphidae, Cordulegastridae, Macromiidae, Chorismagrionidae, Diphlebiidae, Lestoideidae and Pseudolestidae). Epiprocta and Zygoptera were recovered as monophyletic. Ditaxinerua is supported as the sister lineage to Odonata, Epiophlebiidae and the lestid‐like damselflies are sister to the Epiprocta and Zygoptera, respectively. Austropetaliidae + Aeshnidae is the sister lineage to the remaining Anisoptera. Tarsophlebia's placement as sister to Epiprocta or as sister to Epiprocta + Zygoptera was not resolved. Refinements are made to the current classification. Fossil taxa did not seem to provide signals crucial to recovering a robust phylogeny, but were critical to understanding the evolution of key morphological features associated with flight. Characters associated with wing structure were optimized revealing two wing character complexes: the pterostigma–nodal brace complex and the costal wing base & costal–ScP junction complex. In turn, these two complexes appear to be associated; the pterostigma–nodal brace complex allowing for further modification of the wing characters comprised within the costal wing base & costal–ScP junction complex leading the modern odonate wing.