Internal wave intensity and angle of propagation modulate small-scale settlement patterns of intertidal barnacles during peak recruitment

The settlement of intertidal barnacles Chthamalus spp. was measured for an entire recruitment season at 3 sites separated by 100 m in Baja California, Mexico. During a 10 d pulse that accounted for nearly 30% of yearly settlement, coastal wind speed and direction, nearshore water column temperature, and current velocities were also measured. During this period, strong internal tidal forcing was observed, with short episodes (1-1.5 h) of rapid fluctuations in water column temperature, stratification, and currents. Chthamalus spp. settlement was significantly and positively correlated with cumulative high-frequency fluctuations in temperature, thermal stratification, and surface current flows, but not with onshore winds. Furthermore, the spatial pattern in the number of settlers was correlated with the angle of propagation estimated for onshore-moving internal waves, potentially modulated by nearshore bathymetry. This relationship between Chthamalus spp. settlement and high-frequency changes in water column temperature was also shown for a site in La Jolla, California, USA, but not for a similar barnacle, Balanus glandula, suggesting that interspecific differences in larval behavior and nearshore depth distribution may translate into differences in transport. Our results suggest that both the number of settlers at shore and their spatial pattern can be modulated by the intensity and direction of internal wave events, at least during peak settlement periods. Future research should consider both the intensity and geometry of internal-wave-induced variability in the nearshore, as well the small-scale features of coastal geomorphology and bathymetry.