Assessing hydrodynamic space use of brown trout, Salmo trutta, in a complex flow environment: a return to first principles.

It is commonly assumed that stream-dwelling fish should select positions where they can reduce energetic costs relative to benefits gained and enhance fitness. However, the selection of appropriate hydrodynamic metrics that predict space use is the subject of recent debate and a cause of controversy. This is for three reasons: (1) flow characteristics are often oversimplified, (2) confounding variables are not always controlled and (3) there is limited understanding of the explanatory mechanisms that underpin the biophysical interactions between fish and their hydrodynamic environment. This study investigated the space use of brown trout, Salmo trutta, in a complex hydrodynamic flow field created using an array of different sized vertically oriented cylinders in a large open-channel flume in which confounding variables were controlled. A hydrodynamic drag function (D) based on single-point time-averaged velocity statistics that incorporates the influence of turbulent fluctuations was used to infer the energetic cost of steady swimming. Novel hydrodynamic preference curves were developed and used to assess the appropriateness of D as a descriptor of space use compared with other commonly used metrics. Zones in which performance-enhancing swimming behaviours (e.g. Kármán gaiting, entraining and bow riding) that enable fish to hold position while reducing energetic costs (termed 'specialised behaviours') were identified and occupancy was recorded. We demonstrate that energy conservation strategies play a key role in space use in an energetically taxing environment with the majority of trout groups choosing to frequently occupy areas in which specialised behaviours may be adopted or by selecting low-drag regions.