The hydrodynamics of swimming at intermediate Reynolds numbers in the water boatman (Corixidae)

The fluid forces that govern propulsion determine the speed and energetic cost of swimming. These hydrodynamics are scale dependent and it is unclear what forces matter to the tremendous diversity of aquatic animals that are between a millimeter and a centimeter in length. Animals at this scale generally operate within the regime of intermediate Reynolds numbers, where both viscous and inertial fluid forces have the potential to play a role in propulsion. The present study aimed to resolve which forces create thrust and drag in the paddling of the water boatman (Corixidae), an animal that spans much of the intermediate regime (10<200). By measuring the force generated by tethered water boatmen, we found that thrust is generated primarily by drag on the paddling appendages, with a negligible contribution from the acceleration reaction force. Based on these findings, we developed a forward-dynamic model of propulsion in free swimming that accurately predicted changes in the body's center of mass over time. For both tethered and free swimming, we used non-linear optimization algorithms to determine the force coefficients that best matched our measurements. With this approach, the drag coefficients on the body and paddle were found to be up to three times greater than on static structures in fully developed flow at the same Reynolds numbers. This is likely a partial consequence of unsteady interactions between the paddles or between the paddles and the body. In addition, the maximum values for these coefficients were inversely related to the Reynolds number, which suggests that viscous forces additionally play an important role in the hydrodynamics of small water boatmen. This understanding for the major forces that operate at intermediate Reynolds numbers offers a basis for interpreting the mechanics, energetics and functional morphology of swimming in many small aquatic animals.