Hydrodynamic pumping by serial gill arrays in the mayfly nymph Centroptilum triangulifer

Aquatic nymphs of the mayfly ventilatory flow using a serial array of seven abdominal gill pairs that operates across a Reynolds numbers (Re) range from 2 to 22 during ontogeny. Net flow in small animals is directed ventrally and essentially parallel to the stroke plane (i.e. rowing), but net flow in large animals is directed dorsally and essentially transverse to the stroke plane (i.e. flapping). Detailed flow measurements based on Particle Image Velocimetry (PIV) ensemble-correlation analysis revealed that the phasing of the gills produces a time-dependent array of vortices associated with a net ventilatory current, a fluid kinematic pattern, here termed a 'phased vortex pump'. Absolute size of vortices does not change with increasing animal size or Re, and thus the vortex radius (R(v)) decreases relative to inter-gill distance (L(is)) during mayfly growth. Given that effective flapping in appendage-array animals requires organized flow between adjacent appendages, we hypothesize that rowing should be favored when L(is)/R(v)<1 and flapping should be favored when L(is)/R(v)>1. Significantly, the rowing-to-flapping transition in Centroptilum occurs at Re similar to 5, when the mean dynamic inter-gill distance equals the vortex radius. This result suggests that the Re-based rowing-flapping demarcation observed in appendage-array aquatic organisms may be determined by the relative size of the propulsive mechanism and its self-generated vortices.