The homology of cephalic muscles and endoskeletal elements between Diplura and Ectognatha (Insecta)

Diplura (two-pronged bristletails) are key to our understanding of hexapod head evolution. A sister group relationship with Ectognatha (=Insecta), comprising bristletails, silverfish and winged insects, is advocated in most modern studies, however, homologization of head muscles and endoskeletal elements between Diplura and Ectognatha is still lacking. Here, we present the first homologization of a number of head muscles and endoskeletal structures between Diplura and Ectognatha. A homologization of these structures is possible if a range of species, both from Japygidae and Campodeidae, are studied in order to reconstruct the potential groundplan characteristics and account for inner anatomy variations within Diplura. Japygidae and Campodeidae show differences in the origin, insertion, and presence of mandibular and maxillary muscles as well as the shape of the maxillary cardo. Taking into account recent embryological studies on the formation of the endoskeleton in Protura, Collembola and Diplura, we furthermore reconstruct the potential evolution of the endoskeleton in early Hexapoda. The tentorium is a defining feature of dicondylic insects (including Archaeognatha) while anterior and posterior cephalic invaginations (the later tentorial pits of dicondylic insects) are groundplan features of Hexapoda. Additionally, we clarify the composition of the gnathal pouches (i.e. the type of entognathy) in Diplura and Collembola. The pouches in Diplura are posteriorly separated, similar to the state encountered in Collembola. This contrasts to former studies emphasizing the differences in the ellipuran and dipluran type of entognathy