Head structures of males of Strepsiptera (Hexapoda) with emphasis on basal splitting events within the order

Internal and external head structures of males of Strepsiptera were examined and the head of a species of Mengenilla is described in detail. The results suggest a reinterpretation of some structures. The head of basal extant strepsipterans is subprognathous, whereas it is strictly orthognathous in the groundplan of Strepsiptera s.l. The labrum and hypopharynx are not part of the mouthfield sclerite. The labial palps are absent in all strepsipterans. A very slightly modified mandibular articulation is preserved in Eoxenos, whereas it is distinctly reduced in other extant groups. A salivary duct, salivary glands, and a cephalic aorta are absent. The cladistic analysis of 44 characters of the head results in the following branching pattern: (Protoxenos + (Mengea + (Eoxenos + (Mengenilla Austr.] + Mengenilla) + (Elenchus + Dundoxenos + Xenos + Stylops)))). Most apomorphies of males are associated with the necessity of finding females within a short time span and with a reduced necessity to consume food: large "raspberry" eyes, flabellate antennae with numerous dome-shaped chemoreceptors, Hofeneder's organ, an ovoid sensillum of the maxillary palp, and the simplified condition of the maxilla and the labium. Strepsiptera excl. Protoxenos are supported by the dorsomedian frontal impression, the dorsally shifted antennal insertions, a reduced number of antennal segments, absence of the galea, and probably by the presence of the mouthfield sclerite, which is a unique apomorphic feature. The balloon-gut combined with an unusual air-uptake apparatus is another possible autapomorphy of this clade. It is likely that the last common ancestor of Strepsiptera excl. Protoxenos did not process food. Strepsiptera s.str. are characterized by the strongly reduced condition of the labrum and the absence of the epistomal suture. Eoxenos is the sister group of the remaining Strepsiptera s.str. Synapomorphies of Mengenilla + Stylopidia are the advanced reduction of the mandibular articulation and the secondary absence of the ovoid sensillum. The monophyly of Mengenilla is confirmed, even though a small free labrum is present in Australian species. Derived features of Stylopidia are the absence of the coronal suture and the reduced condition of the frontal suture. Apomorphies that have evolved within Stylopidia are the membranization of parts of the head, the fusion of antennal segments, the increase or decrease of the number of flabellate flagellomeres, reductions and modifications of the mandibles, and modifications of the mouthfield sclerite. The monophyly of Stylopiformia is not unambiguously supported. A position of the mandibles posterior to the mouthfield sclerite (when adducted) is a possible synapomorphy shared by Xenos, Stylops, and other "higher Stylopidia." The blade-like distal part of the mandibles suggests a closer relationship of Elenchus with these taxa.