The systematics of Mantodea revisited: an updated classification incorporating multiple data sources (Insecta: Dictyoptera)

After a brief synopsis of the history of mantodean classification, a re-organized systematic arrangement of extant praying mantids is provided. To overcome past homoplasy problems, a phylogenetic framework based on male genital structure was used, supplemented by published morphological, chromosomal and molecular data. As already noticed by previous authors, external morphology is highly homoplastic and does not provide useful systematic tools above subfamily level. In contrast, the morphology of male external genitalia is largely congruent with the results of recent molecular phylogenies, but contradicts the most widely used past systems. Additionally, some genital structures widely used for taxonomic purposes could be shown to be not homologous, most notably the distal process. Evolutionary transitions of the distal process and the phalloid apophysis across the mantodean phylogenetic tree are identified and named. The phalloid apophysis of many derived mantodeans shows a tendency towards bifurcation into an anterior and a posterior lobe. This and other observed genital traits are hypothesized to be an adaptation of males towards a stable copulatory grasp in groups exhibiting increased sexual dimorphism, associated with an increased risk for the male to be cannibalized during copulation. Genital characters allowed most genera to be unambiguously assigned to the major clades (superfamilies) recovered by our genital and previous molecular data. The few exceptions concern genera with secondarily simplified genitalia lacking diagnostic structures. Taxonomic literature is very heterogeneous, and several subfamilies yet lacking any modern revisionary treatment will need further refinement. To account for phylogenetic constraints, i.e. correct for past polyphyletic groupings, the number of families was elevated to 29, and the number of subfamilies to 60. We establish the new family Leptomantellidae, the new subfamilies Brancsikiinae and Deiphobinae, the new tribes Leptomiopterygini, Hagiomantini, Gonypetellini, Bolbellini, Epsomantini, Neomantini, Amantini, Armenini, Danuriellini, Deiphobini, Cotigaonopsini, Didymocoryphini, Oxyelaeini, Heterochaetulini, Rhodomantini and Pseudoxyopsidini, and the new subtribes Amphecostephanina, Bolbina, Tricondylomimina, Gonypetyllina, Antistiina, Toxomantina and Tarachomantina. New morphological diagnoses are provided for the currently recognized families. Despite a few yet to be solved problems, this work offers the urgently needed working base for future studies in Mantodean systematics, life history and ecology.