Linking predators to seasonality of upwelling: Using food web indicators and path analysis to infer trophic connections

Upwelling in eastern boundary current systems is a primary driver of ecosystem productivity. Typically, peak upwelling occurs during spring and summer, but winter upwelling may also be important to ecosystem functions. In this study, we investigated the hypothesis that winter and spring/summer upwelling, operating through indirect trophic interactions, are important to a suite of top predators in the California Current. To test this hypothesis, we collated information on upwelling, chlorophyll-a concentrations, zooplankton and forage fish, and related these to predator responses including rockfish growth, salmon abundance, seabird productivity and phenology (timing of egg-laying), and whale abundance. Seabird diets served in part as food web indicators. We modeled pathways of response using path analysis and tested for significance of the dominant paths with multiple regression. We found support for the hypothesis that relationships between upwelling and top predator variables were mediated primarily by intermediate trophic levels. Both winter and summer upwelling were important in path models, as were intermediate lower and mid trophic level functional groups represented by chlorophyll-a, zooplankton, and forage fish. Significant pathways of response explained from 50% to 80% of the variation of seabird (Cassin's auklet (Ptychoramphus aleuticus) and common murre (Uria aalge)), humpback whale (Megaptera novaeangliae) and Chinook salmon (Oncorhynchus tshawytscha) dependent variables, whereas splitnose rockfish (Sebastes diploproa) showed no significant response pathways. Upwelling and trophic responses for salmon were established for both the year of ocean entry and the year of return, with zooplankton important in the year of ocean entry and forage fish important in the year of return. This study provides one of the first comparative investigations between upwelling and predators, from fish to marine mammals and birds within a geographically restricted area, demonstrates often difficult to establish "bottom-up" trophic interactions, and establishes the importance of seasonality of upwelling to various trophic connections and predator demographic traits. Understanding change in the seasonality of upwelling is therefore required to assess dynamics of commercially and recreationally important upper trophic level species in eastern boundary current ecosystems. (C) 2012 Elsevier Ltd. All rights reserved.