Pectoral fin locomotion in batoid fishes: undulation versus oscillation

This study explores the dichotomy between undulatory (passing multiple waves down the fin or body) and oscillatory (flapping) locomotion by comparing the kinematics of pectoral fin locomotion in eight species of batoids (Dasyatis americana, D. sabina, D. say, D. violacea, Gymnura micrura, Raja eglanteria, Rhinobatos lentiginosus and Rhinoptera bonasus) that differ in their swimming behavior, phylogenetic position and lifestyle. The goals of this study are to describe and compare the pectoral fin locomotor behavior of the eight batoid species, to clarify how fin movements change with swimming speed for each species and to analyze critically the undulation/oscillation continuum proposed by Breder using batoids as an example. Kinematic data were recorded for each species over a range of swimming velocities (1-3 disc lengths s(-1)). The eight species in this study vary greatly in their swimming modes. Rhinobatos lentiginosus uses a combination of axial-based and pectoral-fin-based undulation to move forward through the water, with primary thrust generated by the tail. The pectoral fins are activated in short undulatory bursts for increasing swimming speed and for maneuvering. Raja eglanteria uses a combination of pectoral and pelvic locomotion, although only pectoral locomotion is analyzed here. The other six species use pectoral locomotion exclusively to propel themselves through the water. Dasyatis sabina and D. say have the most undulatory fins with an average of 1.3 waves per fin length, whereas Rhinoptera bonasus has the most oscillatory fin behavior with 0.4 waves per fin length. The remaining species range between these two extremes in the degree of undulation present on their fins. There is an apparent trade-off between fin-beat frequency and amplitude. Rhinoptera bonasus has the lowest frequency and the highest fin amplitude, whereas Rhinobatos lentiginosus has the highest frequency and the lowest amplitude among the eight species examined. The kinematic variables that batoids modify to change swimming velocity vary among different species. Rhinobatos lentiginosus increases its tail-beat frequency to increase swimming speed. In contrast, the four Dasyatis species increase swimming speed by increasing frequency and wavespeed, although D. americana also changes wave number. Raja eglanteria modifies its swimming velocity by changing wavespeed and wave number. Rhinoptera bonasus increases wavespeed, Gymnura micrura decreases wave number, and both Rhinoptera bonasus and Gymnura micrura increase fin-tip velocity to increase swimming velocity. Batoid species fall onto a continuum between undulation and oscillation on the basis of the number of waves present on the fins.