From Carboniferous to Recent: wing venation enlightens evolution of thysanopteran lineage

Recent Thysanoptera are characterized by two pairs of slender wings fringed by long hairs and with reduced venation. Fossils presenting more complete venation have been used in earlier studies to link Thysanoptera to 'Zoropsocinae', which form with 'Lophioneurinae' the 'Lophioneuridae', an extinct family of the superorder Thripida. On the basis of one new Carboniferous fossil described herein, Westphalothripides oudardi sp. nov. (Westphalothripidesidae fam. nov.), as well as new Cretaceous fossils, we revise the current interpretation of venation in Thripida and propose new cladistic analyses, which divide this superorder into three clades: Panthysanoptera nov. (including 'Zoropsocidae' stat. rest. and Thysanoptera), Lophioneurida (including 'Lophioneurinae' and Moundthripidae), and Westphalothripidesidae. We confirm that the Thysanoptera belong to the superorder Thripida but show that 'Lophioneuridae' are paraphyletic. Thanks to the venation observed in fossils, we describe a new character of the wing in some Recent species which allows them to be integrated in phylogenetic studies. Results show two clades inside Thysanoptera: Aeolothripidae + Melanthripidae + Merothripidae, and Thripidae + Stenurothripidae + Fauriellidae. Phlaeothripidae could not be studied due to the absence of the diagnostic wing veins in this family. We discuss the appearance of Thripida within Paraneoptera and particularly the importance of one apomorphy, the reduction of the right mandible as a putative adaptation to spore and pollen piercing.