The evolution of Strepsiptera (Hexapoda)

An evolutionary scenario for the enigmatic group Strepsiptera is provided, based on the results of a comprehensive cladistic analysis of characters of all life stages. A recently described fossil--+Protoxenos janzeni--the most archaic strepsipteran, sheds new light on the early evolution of the group and reduces the "morphological gap" between Strepsiptera and other insects. It weakens both current hypotheses--Coleoptera+Strepsiptera and Diptera+Strepsiptera (="Halteria"). The splitting into +Protoxenos (Protoxenidae) and the remaining Strepsiptera was linked with a distinct size reduction and many morphological changes. Unlike males of extant strepsipteran species, +Protoxenos was still able to process food. Mengeidae (+Mengea), with two small species, is the sister group of extant Strepsiptera. A unique characteristic of extant males (Strepsiptera s. str.) is the mouthfield sclerite. It is part of an air uptake apparatus which belongs to an extremely modified air-filled "balloon gut". Besides this, male strepsipterans possess specialised antennae and compound eyes, a strongly developed flight apparatus, large testes, and a sperm pump, whereas other organ systems are strongly reduced (e.g., fat body, malpighian tubules). Males are designed to find females within a few hours and to copulate. A dramatic change is linked with the split into Mengenillidae and Stylopidia. The change to pterygote hosts and the permanent endoparasitism of the females are evolutionary novelties acquired by the latter clade, and linked with far-reaching morphological transformations, e.g. the presence of unique brood organs. Hairy tarsal adhesive devices are present in males and guarantee efficient attachment to the host during copulation. A well-founded clade within Stylopidia is Stylopiformia, which are characterised by a unique fissure-shaped birth opening. The evolutionary development towards the most specialised and successful forms (parasites of aculeate Hymenoptera [e.g., Xenidae+Stylopidae], ca. 46% of the species) is a stepwise process. The presented evolutionary scenario comprises a complex network of functionally correlated morphological changes in primary larvae, secondary larvae, females and males.