Winter mixing and coastal upwelling drive long-term changes in zooplankton in the Bay of Biscay (1993-2010)

Long-term studies are necessary to gain insight into the influence of external climate forcing on zooplankton dynamics. Using monthly time series (1993-2010) collected in the Cantabrian Sea (southern Bay of Biscay), we studied long-term trends and seasonal cycles of zooplankton abundance and biomass along a coastal-ocean gradient. We also analysed changes in environmental variables such as sea surface temperature (SST) and chlorophyll a concentration. Zooplankton abundance and biomass increased during the study period, with greater increases in biomass towards the coast and for larger size classes. These results contrast with the decrease in biomass and size expected under global warming, which is shown in the region by the rise in summer SST, suggesting the operation of other processes. Indeed, winter mixing and coastal upwelling were key drivers of zooplankton dynamics in spring and autumn, respectively. In these seasons, zooplankton inter-annual increases were stronger. Including these hydrographic processes in future models of zooplankton dynamics for the Bay of Biscay and other temperate areas will improve their accuracy. Our results highlight the importance of disentangling the main mechanisms behind long-term changes in zooplankton, especially within the context of climate change.