The Functional Morphology of Shark Control Surfaces: A Comparative Analysis

Sharks are an objectively diverse group of animals; ranging in maximum size from 2,000cm (whale shark) to 17cm (dwarf lantern shark); occupying habitats that are periodically terrestrial (epaulette shark) to the deepest parts of the ocean (frilled shark); relying on a diversity of diets from plankton to marine mammals; with vast amounts of morphology diversity such as the laterally expanded heads of hammerhead species, the elongate caudal fins of thresher species, and the tooth embedded rostrum of saw shark species representing some of the anatomical extremes. Yet despite these obvious differences in morphology, physiology, and ecology, the challenges associated with studying hard to access, large bodied, pelagic animals have limited our comparative understanding of form and function as it relates to swimming within this group. The majority of shark swimming studies examine species that succeed in captivity, which are usually benthic associated sharks that spend time resting on the substrate. These studies have also been limited by the use of flumes, in which the unidirectional flow and small working area precludes the analysis of larger animals, volitional swimming, and maneuvering. The few existing volitional kinematics studies on sharks quantify two-dimensional kinematics which are unable to capture movements not observable in the plane of reference. With this study, we quantified the volitional swimming kinematics of sharks in relation to morphological, physiological, and ecological variation among species. We developed a technique to analyze three-dimensional (3D) kinematics in a semi-natural, large volume environment, which, to our knowledge, provides the first3D analysis of volitional maneuvering in sharks. We demonstrated that Pacific spiny dogfish and bonnethead sharks rotate the pectoral fins substantially during yaw (horizontal) maneuvering and is correlated with turning performance. We proposed that ecomorphological differences correlate with the varied maneuvering strategies we observed between the two species. We also found that there is some mechanical constraint on shark pectoral fin shape that is explained by phylogenetic relationships but describe a continuum of morphological variables within that range. We propose standardized terminology and methodology for the future assessment of shark pectoral fin morphology and function. As with previous studies, the ease of access to species was a challenge in this study and future studies should continue to assess the functional ecomorphology of shark pectoral fins among species.