Description: Understanding species' responses to upwelling may be especially important in light of ongoing environmental change. Upwelling frequency and intensity are expected to increase in the future, while ocean acidification and deoxygenation are expected to decrease the pH and dissolved oxygen of upwelled waters. However, the acute effects of a single upwelling event and the integrated effects of multiple upwelling events on marine organisms are poorly understood. Here, we use in situ measurements of pH, temperature, and dissolved oxygen to characterize the covariance of environmental conditions within upwelling-dominated kelp forest ecosystems. We then test the effects of acute (0-3 days) and chronic (1-3 month) upwelling on the performance of two species of kelp forest grazers, the echinoderm, Mesocentrotus franciscanus, and the gastropod, Promartynia pulligo. We exposed organisms to static conditions in a regression design to determine the shape of the relationship between upwelling and performance and provide insights into the potential effects in a variable environment. We found that respiration, grazing, growth, and net calcification decline linearly with increasing upwelling intensity for M. francicanus over both acute and chronic timescales. Promartynia pulligo exhibited decreased respiration, grazing, and net calcification with increased upwelling intensity after chronic exposure, but we did not detect an effect over acute timescales or on growth after chronic exposure. Given the highly correlated nature of pH, temperature, and dissolved oxygen in the California Current, our results suggest the relationship between upwelling intensity and growth in the 3-month trial could potentially be used to estimate growth integrated over long-term dynamic oceanographic conditions for M. franciscanus. Together, these results indicate current exposure to upwelling may reduce species performance and predicted future increases in upwelling frequency and intensity could affect ecosystem function by modifying the ecological roles of key species.

Description: Despite a growing interest in identifying tipping points in response to environmental change, our understanding of the ecological mechanisms underlying non-linear ecosystem dynamics is limited. Ecosystems governed by strong species interactions can provide important insight into how non-linear relationships between organisms and their environment propagate through ecosystems, and the potential for environmentally mediated species interactions to drive or protect against sudden ecosystem shifts. Here, we experimentally determine the functional relationships (i.e., the shapes of the relationships between predictor and response variables) of a seagrass assemblage with well-defined species interactions to ocean acidification (enrichment of CO2) in isolation and in combination with nutrient loading. We demonstrate that the effect of ocean acidification on grazer biomass (Phyllaplysia taylori and Idotea resecata) was quadratic, with the peak of grazer biomass at mid-pH levels. Algal grazing was negatively affected by nutrients, potentially due to low grazer affinity for macroalgae (Ulva intestinalis), as recruitment of both macroalgae and diatoms were favored in elevated nutrient conditions. This led to an exponential increase in macroalgal and epiphyte biomass with ocean acidification, regardless of nutrient concentration. When left unchecked algae can cause declines in seagrass productivity and persistence through shading and competition. Despite quadratic and exponential functional relationships to stressors that could cause a non-linear decrease in seagrass biomass, productivity of our model seagrass – the eelgrass (Zostera marina)- remained highly resilient to increasing acidification. These results suggest that important species interactions governing ecosystem dynamics may shift with environmental change, and ecosystem state may be decoupled from ecological responses at lower levels of organization.