Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species dagger Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage 'Protorthoptera', is re-interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem-Mantodea, as are some other 'protorthopterous' taxa. This hypothesis implies that the total-group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous 'Protorthoptera' are stem-representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock-crawlers), suggesting a survivorship of several main Pterygota lineages at the end-Permian extinction event higher than previously expected. (C) 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 79-113.