Is the Carboniferous †Adiphlebia lacoana really the oldest beetle? Critical reassessment and description of a new Permian beetle family

Bethoux recently identified the species dagger Adiphlebia lacoana Scudder from the Carboniferous of Mazon Creek, Ill., USA as the oldest beetle. The fossils bear coriaceous tegmina with pseudo-veins allegedly aligned with rows of cells as they occur in Permian beetles and extant Archostemata. The examination of four new specimens of dagger Adiphlebia lacoana from the same locality revealed that the cells are in fact clumps of clay inside a delicate meshwork, and no derived features shared with Coleoptera or Coleopterida (= Coleoptera + Strepsiptera) were found. Instead, dagger Adiphlebia lacoana bears veinal fusions and braces similar to extant Neuroptera. These features support a placement in dagger Strephocladidae, and are also similar to conditions found in dagger Tococladidae. These unplaced basal holometabolan families were erroneously re-analyzed as ancestral Mantodea and Orthoptera. Homologization of the wing pairs in neopteran lineages is updated and identification errors are corrected. A new Permian beetle family dagger Moravocoleidae [dagger Protocoleoptera (= Permian Coleoptera with pointed unpaired ovipositor; e.g., dagger Tshekardocoleidae)] is described.