Aesthete Pattern Diversity in Chiton Clades (Mollusca: Polyplacophora): Balancing Sensory Structures and Strength in Valve Architecture.

Chitons possess the most elaborate system of shell pores found in any hard-shelled invertebrate. Although chitons possess some anteriorly located sense organs, they lack true cephalization, as their major sensory systems are not concentrated in a distinct head region. Instead, the aesthete system within their shells forms a dense sensory network that overcomes the barrier of their hard dorsal armour. The basic arrangement of neural structures embedded within a solid, opaque matrix, has confounded understanding of the overall network. In this study, we use synchrotron X-ray μCT to visualise the aesthete canal networks inside chiton valves. We selected representatives from all three major chiton clades: Lepidopleurida, the basal branching clade, and Callochitonida and Chitonida, which both have more complex shell morphology, to compare internal structure. Lepidopleurida aesthete canals are oriented vertically and pass directly through the shell to connect with the body. By contrast, aesthetes canals in Callochitonida and Chitonida have complex internal structures with extended horizontal passages, coalescing at the shell diagonal that corresponds to the valve insertion slits. This represents a stepwise evolution of chiton shell form, where thicker and more complex valves require a diverting and rewiring of the entire sensory network. Aspects of the aesthete system, such as the microscopic arrangement of surface pores, have long been used in chiton taxonomy for species diagnoses; insertion slits should also be understood as a secondary feature of the aesthete system. Chiton shell structures that are used for morphological systematics are driven by sensory adaptations.