Structure and mechanical implications of the pectoral fin skeleton in the Longnose Skate (Chondrichthyes, Batoidea)

Animal propulsion systems are believed to show high energy and mechanical efficiency in assisting movement compared to artificial designs. As an example, batoid fishes have very light cartilaginous skeletons that facilitate their elegant swimming via enlarged wing-like pectoral fins. The aim of this work is to illustrate the hierarchical structure of the pectoral fin of a representative batoid, the Longnose Skate (Raja rhina), and explain the mechanical implications of its structural design. At the macro level, the pectoral fins are comprised of radially oriented fin rays, formed by staggered mineralized skeletal elements stacked end-to-end. At the micro level, the midsection of each radial element is composed of three mineralized components, which consist of discrete segments (tesserae) that are mineralized cartilage and embedded in unmineralized cartilage. The radial elements are wrapped with aligned, unmineralized collagen fibers. This is the first report of the detailed structure of the ray elements, including the observation of a 3-chain mineralized tesserae. Structural analyses demonstrate that this configuration enhances stiffness in multiple directions. A two-dimensional numerical model based on the morphological analysis demonstrated that the tessera structure helps distributing shear, tensile and compressive stress more ideally, which can better support both lift and thrust forces when swimming without losing flexibility.