Morphological, functional, and phylogenetic aspects of the head capsule of the cockroach Ergaula capucina (Insecta/Blattodea)

Background: Cockroaches are usually typical omnivorous detritivores and their cephalic morphology is considered to be ancestral in various aspects. Thus, several studies addressed the morphology and function of the blattodean head, and the cockroach usually serves as a model for standard mouthparts in text books. However, so far only two of the three major lineages of Blattodea have been studied and no detailed information for the head of any Corydioidea was available. The present study closes this gap by providing a detailed morphological description of the head of Ergaula capucina, studying some important functional parameters of the mandible and discussing it in a phylogenetic framework. Methods: The cephalic morphology of Ergaula studied in detail using a broad set of different techniques including digital microscopy, µ-computed tomography, and 3-dimensional reconstructions. Concerning the functional morphology of the mandible, we compared the volume and effective cross sections of the eight compartments of the primary mandibular adductor muscle for Ergaula, Blattella germanica, and Salganea rossi and measured the mechanical advantage, i.e., the force transmission ratio for all teeth of the mandible of Ergaula. Results: The head capsule of Ergaula is characterized by a strong sexual dimorphism and typical orthopteran mouthparts. It resembles the head capsule of other roaches in several respects and confirms oesotendons, the reduction of the mesal occelus, and bipartite M. verticopharyngealis and M. hypopharyngosalivaris as blattodean apomorphies. But it also shows some unique adaptations. It is the first described cockroach that lacks the dorsal tentorial arms which has various consequences for the cephalic musculature. On the maxillary lacinia, Ergaula is the first described blattodean to show strong and blunt setae instead of a lacinula, which might be homologues to the dentisetae of dragonflies and mayflies. Like other corydiid roaches that inhabit xeric areas, Ergaula has an atmospheric water-vapor absorption mechanism that includes a gland and a ductus on the epipharnyx and bladders on the hypopharynx. The mandibular adductor is in cockroaches asymmetric, a pattern not found in termites, mantids, or other closely related insects.