Upwelling and Other Environmental Influences on Growth of a Nearshore Benthic Fish

The role of upwelling in nearshore benthic systems is more uncertain compared to the relatively strong positive associations with pelagic production. To understand how upwelling and other environmental conditions influence nearshore benthic production, we developed an annual index of production from growth increments recorded in otoliths of kelp greenling (Hexagrammos decagrammus) at nine sites in the seasonally-upwelling California Current and downwelling Alaska Coastal currents. Kelp greenling are a benthic-feeding fish common in kelp forests with food webs sustained by both kelp and phytoplankton primary production. We explored the influence of basin- and local-scale conditions, including upwelling, across all seasons at lags up to two years taken to represent changes in the quantity and quality of prey. Upwelling strength was positively related to fish growth in both current systems, although relationships in the Alaska Coastal Current were indicative of faster growth with relaxed downwelling, rather than upwelling. Looking across a suite of basin- and local-scale environmental indicators, complex relationships emerged in the California Current, with faster growth related to within-year warm conditions and lagged-year cool conditions. In contrast, fish in the downwelling system grew faster both during and subsequent to warm conditions. The complex lag-dependent dynamics in the upwelling system may reflect differences in conditions that promote quantity versus quality of benthic invertebrate prey. Thus, we hypothesize that benthic production is maximized when cool and warm years alternate during periods of high frequency climate variability in the California Current. Such a pattern is consistent with previous findings suggesting that benthic invertebrate abundance (e.g., recruitment) is food-limited during warm years with reduced upwelling, while quality (e.g., energy content) is temperature-limited during cool years.