Physical–biological coupling in spore dispersal of kelp forest macroalgae

The physical–biological linkages controlling the dispersal of spores produced by macroalgae that reside in kelp forests are complicated and laced with feedbacks. Here we discuss the fundamental elements of these interactions. Biological considerations include spore swimming and sinking speeds, their periods of viability in the plankton, and the height of spore release above the seafloor, which together determine the durations over which spores can be swept by horizontal currents before they contact the seafloor. Morphologies and material properties of canopy forming kelps may also influence the drag exerted on passing waters by the kelps, the plants' ability to persist in the face of rapid flows, and thereby the degree to which impinging currents are redirected around, or slowed within, kelp forests. Macroalgal life histories, and the size of spore sources as controlled by the dimensions of kelp forests and the density and fecundity of individuals within them, influence effective dispersal distances as well. Physical considerations encompass the mean speed, direction, and timescales of variability of currents relative to spore suspension times, the interaction of surface gravity waves with currents in producing turbulence in the benthic boundary layer, wind-driven surface mixing, water stratification, and shoreline bathymetry and substratum roughness, all of which can affect the interplay of vertical and horizontal transport of macroalgal spores. Intricate within-forest processes may induce attenuation of current speeds and consequent reductions in seabed shear, along with simultaneous production of small-scale turbulence in kelp wakes. Slower mean currents and smaller eddy scales in turn may attenuate vertical mixing within forests, thus extending spore suspension times. Further complexities likely arise due to changes in the relative rates of horizontal and vertical dispersion, modifications to the overall profiles of vertical mixing, and the creation of fine-scale secondary flows around kelp individuals and substratum features. Under conditions of more rapid currents, submergence of the surface canopy and the establishment of skimming flows at the canopy–fluid interface may introduce additional coherent flow structures that alter rates of fluid exchange to and from the forest. Many of these coupled physical–biological processes are just beginning to be examined in a rigorous fashion in kelp forests, but their potential importance is clear.