Scaling Coastal Ecosystems to Oceanographic and Climatic Drivers: Making Sense of Community Variation on Rocky Shores Using the Comparative-Experimental Approach in Upwelling and Downwelling Systems

Ecology focuses increasingly on the issue of matching spatial and temporal scales responsible for ecosystem pattern and dynamics. Benthic coastal communities traditionally were studied at local scales using mostly short-term research, while environmental (oceanographic, climatic) drivers were investigated at large scales (e.g., regional to oceanic, mostly offshore) using combined snapshot and monitoring (time series) research. The comparative-experimental approach combines local-scale studies at multiple sites spanning large-scale environmental gradients in combination with monitoring of inner shelf oceanographic conditions including upwelling/downwelling wind forcing and their consequences (e.g., temperature), and inputs of subsidies (larvae, phytoplankton, detritus). Temporal scale varies depending on the questions, but can extend from years to decades. We discuss two examples of rocky intertidal ecosystem dynamics, one at a regional scale (California Current System, CCS) and one at an interhemispheric scale. In the upwelling-dominated CCS, 52% and 32% of the variance in local community structure (functional group abundances at 13 sites across 725 km) was explained by external factors (ecological subsidies, oceanographic conditions, geographic location), and species interactions, respectively. The interhemispheric study tested the intermittent upwelling hypothesis (IUH), which predicts that key ecological processes will vary unimodally along a persistent downwelling to persistent upwelling gradient. Using 14-22 sites, unimodal relationships between ecological subsidies (phytoplankton, prey recruitment), prey responses (barnacle colonization, mussel growth) and species interactions (competition rate, predation rate and effect) and the Bakun upwelling index calculated at each site accounted for ~50% of the variance. Hence, external factors can account for about half of locally-expressed community structure and dynamics.