Phenological decoupling of mortality from wave forcing in kelp beds

Kelps often live in a harsh hydrodynamic environment where wave-driven dislodgement of individuals can alter the biodiversity and functioning of reef systems, and increase production in coastal ecosystems adjacent to reefs. The current paradigm is that winter storms tear kelps from reefs once hydrodynamic forces exceed attachment or tissue strength--a threshold response that implies a pulsed relationship between wave forces and dislodgement. Here, we challenge this understanding by showing how kelp phenology can decouple susceptibility to dislodgement from seasonal patterns in wave forces. We measured kelp dislodgement rates and hydrodynamic forces at nine subtidal reefs over two years (n = 4320 kelps tagged and monitored). Contrary to expectation, we found relatively low and constant dislodgement rates for all reefs (13% +/- 6% [mean per season +/- SD]) in spite of a strong temporal pattern in wave action and extreme water velocities (winter peaks up to 3-4 m/s). A biomechanical model, based on the balance between kelp attachment strength and hydrodynamic drag, demonstrated that severe reduction in individual kelp size toward winter (>50% decrease in biomass for all sites) minimized drag and made the kelps less susceptible to high water velocities, allowing individuals to survive storm velocities over 3-4 m/s. We conclude that the timing of reduced susceptibility to disturbance, through the seasonal reduction of individual kelp biomass that coincides with times of highest water velocities is critical to the dynamics of kelp dislodgement and survival. We propose that phenological processes maintain many kelp beds in a higher degree of population stability and equilibrium with hydrodynamic forces than previously believed.