Undulatory Locomotion in Freshwater Stingray Potamotrygon Orbignyi: Kinematics, Pectoral Fin Morphology, and Ground Effects on Rajiform Swimming

Abstract

Fishes are the most speciose group of living vertebrates, making up more than half of extant vertebrate diversity. They have evolved a wide array of swimming modes and body forms, including the batoid elasmobranchs, the dorsoventrally flattened skates and rays, which swim via oscillations or undulations of a broad pectoral fin disc. In this work I offer insights into locomotion by an undulatory batoid, freshwater stingray Potamotrygon orbignyi (Castelnau, 1855), combining studies of live animals, physical models, and preserved specimens. In Chapter 1, I quantify the three-dimensional kinematics of the P. orbignyi pectoral fin during undulatory locomotion, analyzing high-speed video to reconstruct three-dimensional pectoral fin motions. A relatively small portion (~25%) of the pectoral fin undulates with significant amplitude during swimming. To swim faster, stingrays increase the frequency, not the amplitude of propulsive motions, similar to the majority of studied fish species. Intermittently during swimming, a sharp, concave-down lateral curvature occurred at the fin margin; as the fin was cupped against the pressure of fluid flow this curvature is likely to be actively controlled. Chapter 2 employs a simple physical model of an undulating fin to examine the ground effects that stingrays may experience when swimming near a substrate. Previous research considering static air- and hydrofoils indicated that near-substrate locomotion offers a benefit to propulsion. Depending on small variations in swimming kinematics, undulating fins can swim faster near a solid boundary, but can also experience significant increases (~25%) in cost-of-transport. In Chapter 3, I determine how pectoral and pelvic fin locomotion are combined in P. orbignyi during augmented punting, a hybrid of pectoral and pelvic fin locomotion sometimes employed as stingrays move across a substrate. The timing of pectoral and pelvic fin motions is linked, indicating coordination of thrust production. Chapter 4 discusses pectoral fin structure and morphological variations within the fin, correlating morphology with the swimming kinematics observed in Chapter 1. Passive and active mechanisms may stiffen the anterior fin to create the stable leading edge seen during swimming; stingrays have converged on several structural features (fin ray segmentation and branching) shared by actinopterygian fishes.