Eversion and withdrawal of an intromittent organ before sexual maturation prepares male beetles for copulation

Some species of criocerine beetles have a hyper-elongated part of the intromittent organ called a flagellum. In resting position, the flagellum is stored in a specialized internal sac in the intromittent organ. This specialized state of the flagellum and internal sac is indispensable during copulation for flagellar insertion into the female spermathecal duct for sperm transfer. However, the morphogenesis of the flagellum does not generate the active state of the flagellum; rather, the flagellum is generated in an inactive and completely coiled state. After eclosion, males of Lema coronata evert and withdraw the internal sac multiple times before sexual maturation, without mounting a female. This behaviour serves to uncoil the flagellum and guide it into the active state with the aid of surface structures on the internal sac. A closely related species, Lema dilecta, also has a long flagellum and undergoes the same behaviour to place the flagellum in the active position. However, some other species of criocerine beetles with much shorter flagella can attain the active state without exhibiting this behaviour. Based on a previously proposed phylogenetic tree, we discuss the evolutionary history of the hyper-elongation of the flagellum and associated behaviour.