The locomotor apparatus of one of the smallest beetles - The thoracic skeletomuscular system of Nephanes titan (Coleoptera, Ptiliidae).

Pterothoracic structures of the minute ptiliid Nephanes titan were examined and described in detail. Effects of miniaturization and the phylogenetic and functional background are discussed. Apomorphies shared with Hydraenidae are the large metascutal shield, the fringe of setae along the posterior edge of the wings, and the fusion of the mesoventrite with the mesanepisternum. Autapomorphies of Ptiliidae are the highly modified feather-like wings, the strongly elongated alacristae, the loss of the mesotrochantin, the enlarged metathoracic pleural wing joint, and the simplification of the direct flight musculature. Most phylogenetically relevant features are linked with miniaturization, especially the modifications of the wings and skeleton, for instance the reduced wing membrane or the simplified metanotum without distinct notal wing processes. Ptiliids show flight characteristics that distinctly differ from what is known in other insects, such as an unusually high amplitude of beats of the elytra, and a unique trajectory of wing return movements, as well as the presence of upper and lower claps. A comparison of cross-sections of ptiliid principal flight muscles with those of larger staphyliniform beetles suggests that muscles of the tiniest beetles are relatively smaller. This may reflect the higher aerodynamic efficiency of ptiliid feather wings compared to the common membranous wings found in other groups.