Functional importance of the mandibular skeleto-muscular system in the bivalved arthropod Heterocypris incongruens (Crustacea, Ostracoda, Cyprididae).

Arthropods with a pair of mandibles (Mandibulata) emerged by the end of the Cambrian period. The mandible is one of the apomorphic characteristics of this monophyletic clade, which is composed of Pancrustacea and Myriapoda. Acquisition of the mandible is one of the important events of the evolutionary pathway of arthropods because the powerful masticatory system provides benefits to individuals regarding food selection. Ancestral mandibulates are well known as so-called Cambrian bivalved arthropods, and a few of them provide a pair of valid mandibles with a broad molar process. However, extant bivalved arthropods can only be found in a few lineages of crustaceans, though all of them are equipped with mandibles. This study focuses on the neuroanatomy of the mandibular skeleto-muscular system of Heterocypris incongruens (Ostracoda, Crustacea), a millimeter-sized bivalved crustacean. Electron microscopy reveals that numerous mechanoreceptive sensilla are distributed inside the mandibular gnathal edges and that there are two types (heterodynal and monodynal) of sensilla, which differ in the number of dendrites and their probable function. This sensory nervous system in the gnathal edges contributes to the precise interdigitation of the right and left mandibles to allow for powerful omnivorous mastication, and the mandibular interdigitation plays a role as the fulcrum of triggering action to open the valves. Therefore, by reversing its fulcrum and load, the mandibular skeleto-muscular system in podocopid ostracods has two sub-systems with different functions, namely the "mandibular masticatory system" and the "valve opening system." Furthermore, this investigation provides significant information on the feeding mode of Cambrian bivalved arthropods, from the view of the functional morphology of the mandibular skeleto-muscular system.