Morphological description, character conceptualization and the reconstruction of ancestral states exemplified by the evolution of arthropod hearts

Arthropods are the most species-rich taxon within Metazoa and have gone through major evolutionary changes with regard to body organization. Arthropod hearts and their associated vascular systems are thus morphologically highly disparate: while some arthropods exhibit very powerful hearts and complex vascular systems, other arthropods do not possess any kind of vascular system or heart at all. A comprehensive study investigating the structure of arthropods hearts has never been undertaken. In this study, we therefore investigate the hearts of 34 species from all major arthropod groups using various imaging techniques (confocal laser scanning microscopy, micro-computed tomography, histology) and describe them by addressing different aspects of heart morphology, e.g. the structure of the myocard or the composition of ostia. In a next step, we conceptualize 18 characters related to heart morphology and their respective character states and-using additional data from the literature-score a matrix for a total of 45 species from 38 supraspecific taxa. We map the characters onto prevailing phylogenetic hypotheses and perform parsimony-based ancestral state reconstruction to trace the evolutionary transformations undergone by arthropod hearts. An exploration of the character concepts (as explanatory hypotheses) reveals ontological peculiarities of character statements that clearly distinguish them in terms of ontological status from descriptive statements (i.e. descriptions of morphemes). The implications of these findings influence the interpretation of ground patterns as explanations. This first phylogenetic approach to heart morphology in the arthropod ground pattern reveals numerous new putative synapomorphies and leads to a reconsideration of the morphology of circulatory systems in early arthropods. Hypotheses on the evolution of hearts in (Pan-) Arthropoda are illustrated and discussed.