Flow field-induced drag forces and swimming behavior of three benthic fish species

Modern ethohydraulics is the study of the behavioral responses of swimming fish to flow fields. However, the exact drag forces experienced by fish remain poorly studied; this information is required to obtain a better understanding of the behavioral responses of fish and their current resistance strategies. We measured near-ground frontal drag forces on preserved individuals of three benthic fish species, round goby (Neogobius melanstomus), gudgeon (Gobio gobio) and bullhead (Cottus gobio), in a flow channel. The forces were compared to acoustic Doppler velocity (ADV) measurements and fish tracking data based on video observations of live fish in the flow channel. Overall, we observed drag coefficients (CD) of ∼10$^{-3}$ at Reynolds numbers ∼10$^{5}$. The frontal drag forces acting on preserved fish with non-spread fins ranged from -1.96 mN*g$^{-1}$ (force per fish wet weight, velocity 0.55 m*s$^{-1}$) to 11.01 mN*g$^{-1}$ (velocity 0.85 m*s$^{-1}$). Spreading the fins strongly increased the drag forces for bullhead and round goby. In contrast, the drag forces were similar for gudgeon with spread fins and all fish with non-spread fins. Video tracking revealed no clear relationship between the position of the fish in the flow field and the forces experienced by the preserved fish at these positions. Collectively, these results suggest that i) the differences in frontal drag forces between species are small in homogenous flow, ii) individuals chose their position in the flow field based on factors other than the drag forces experienced, and iii) whether fins are spread or non-spread is an essential quality that modulates species-specific differences. The methodology and results of this study will enable integration of flow measurements, fish behavior and force measurements and inform ethohydraulics research. More advanced force measurements will lead to a detailed understanding of the current resistance strategies of benthic fish and improve the design of fish passes