Optimal Environmental Conditions and Anomalous Ecosystem Responses: Constraining Bottom-up Controls of Phytoplankton Biomass in the California Current System

In Eastern Boundary Current systems, wind-driven upwelling drives nutrient-rich water to the ocean surface, making these regions among the most productive on Earth. Regulation of productivity by changing wind and/or nutrient conditions can dramatically impact ecosystem functioning, though the mechanisms are not well understood beyond broad-scale relationships. Here, we explore bottom-up controls during the California Current System (CCS) upwelling season by quantifying the dependence of phytoplankton biomass (as indicated by satellite chlorophyll estimates) on two key environmental parameters: subsurface nitrate concentration and surface wind stress. In general, moderate winds and high nitrate concentrations yield maximal biomass near shore, while offshore biomass is positively correlated with subsurface nitrate concentration. However, due to nonlinear interactions between the influences of wind and nitrate, bottom-up control of phytoplankton cannot be described by either one alone, nor by a combined metric such as nitrate flux. We quantify optimal environmental conditions for phytoplankton, defined as the wind/nitrate space that maximizes chlorophyll concentration, and present a framework for evaluating ecosystem change relative to environmental drivers. The utility of this framework is demonstrated by (i) elucidating anomalous CCS responses in 1998-1999, 2002, and 2005, and (ii) providing a basis for assessing potential biological impacts of projected climate change.