Locomotion on the water surface: hydrodynamic constraints on rowing velocity require a gait change

Fishing spiders, Dolomedes triton (Araneae, Pisauridae), propel themselves across the water surface using two gaits: they row with four legs at sustained velocities below 0.2 m s(-)(1) and they gallop with six legs at sustained velocities above 0.3 m s(-)(1). Because, during rowing, most of the horizontal thrust is provided by the drag of the leg and its associated dimple as both move across the water surface, the integrity of the dimple is crucial. We used a balance, incorporating a biaxial clinometer as the transducer, to measure the horizontal thrust forces on a leg segment subjected to water moving past it in non-turbulent flow. Changes in the horizontal forces reflected changes in the status of the dimple and showed that a stable dimple could exist only under conditions that combined low flow velocity, shallow leg-segment depth and a long perimeter of the interface between the leg segment and the water. Once the dimple disintegrated, leaving the leg segment submerged, less drag was generated. Therefore, the disintegration of the dimple imposes a limit on the efficacy of rowing with four legs. The limited degrees of freedom in the leg joints (the patellar joints move freely in the vertical plane but allow only limited flexion in other planes) impose a further constraint on rowing by restricting the maximum leg-tip velocity (to approximately 33 % of that attained by the same legs during galloping). This confines leg-tip velocities to a range at which maintenance of the dimple is particularly important. The weight of the spider also imposes constraints on the efficacy of rowing: because the drag encountered by the leg-cum-dimple is proportional to the depth of the dimple and because dimple depth is proportional to the supported weight, only spiders with a mass exceeding 0.48 g can have access to the full range of hydrodynamically possible dimple depths during rowing. Finally, the maximum velocity attainable during rowing is constrained by the substantial drag experienced by the spider during the glide interval between power strokes, drag that is negligible for a galloping spider because, for most of each inter-stroke interval, the spider is airborne. We conclude that both hydrodynamic and anatomical constraints confine rowing spiders to sustained velocities lower than 0.3 m s(-)(1), and that galloping allows spiders to move considerably faster because galloping is free of these constraints.