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Responses of marine primary production to a changing climate are determined by a concert of multiple environmental changes, for example in temperature, light, pCO 2 , nutrients, and grazing. To make robust projections of future global marine primary production, it is crucial to understand multiple driver effects on phytoplankton. This meta-analysis quantifies individual and interactive effects of dual driver combinations on marine phytoplankton growth rates. Almost 50% of the single-species laboratory studies were excluded because central data and metadata (growth rates, carbonate system, experimental treatments) were insufficiently reported. The remaining data (42 studies) allowed for the analysis of interactions of pCO 2 with temperature, light, and nutrients, respectively. Growth rates mostly respond non-additively, whereby the interaction with increased pCO 2 profusely dampens growth-enhancing effects of high temperature and high light. Multiple and single driver effects on coccolithophores differ from other phytoplankton groups, especially in their high sensitivity to increasing pCO 2. Polar species decrease their growth rate in response to high pCO 2 , while temperate and tropical species benefit under these conditions. Based on the observed interactions and projected changes, we anticipate primary productivity to: (a) first increase but eventually decrease in the Arctic Ocean once nutrient limitation outweighs the benefits of higher light availability; (b) decrease in the tropics and mid-latitudes due to intensifying nutrient limitation, possibly amplified by elevated pCO 2 ; and (c) increase in the Southern Ocean in view of higher nutrient availability and synergistic interaction with increasing pCO 2. Growth-enhancing effect of high light and warming to coccolithophores, mainly Emiliania huxleyi, might increase their relative abundance as long as not offset by acidification. Dinoflagellates are expected to increase their relative abundance due to their positive growth response to increasing pCO 2 and light levels. Our analysis reveals gaps in the knowledge on multiple driver responses and provides recommendations for future work on phytoplankton.

Atmospheric and oceanic CO2 concentrations are rising at an unprecedented rate. Laboratory studies indicate a positive effect of rising CO2 on phytoplankton growth until an optimum is reached, after which the negative impact of accompanying acidification dominates. Here, we implemented carbonate system sensitivities of phytoplankton growth into our global biogeochemical model FESOM-REcoM and accounted explicitly for coccolithophores as the group most sensitive to CO2. In idealized simulations in which solely the atmospheric CO2 mixing ratio was modified, changes in competitive fitness and biomass are not only caused by the direct effects of CO2, but also by indirect effects via nutrient and light limitation as well as grazing. These cascading effects can both amplify or dampen phytoplankton responses to changing ocean pCO2 levels. For example, coccolithophore growth is negatively affected both directly by future pCO2 and indirectly by changes in light limitation, but these effects are compensated by a weakened nutrient limitation resulting from the decrease in small-phytoplankton biomass. In the Southern Ocean, future pCO2 decreases small-phytoplankton biomass and hereby the preferred prey of zooplankton, which reduces the grazing pressure on diatoms and allows them to proliferate more strongly. In simulations that encompass CO2-driven warming and acidification, our model reveals that recent observed changes in North Atlantic coccolithophore biomass are driven primarily by warming and not by CO2. Our results highlight that CO2 can change the effects of other environmental drivers on phytoplankton growth, and that cascading effects may play an important role in projections of future net primary production.

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The collection of zooplankton swimmers and sinkers in time-series sediment traps provides unique insight into year-round and interannual trends in zooplankton population dynamics. These samples are particularly valuable in remote and difficult to access areas such as the Arctic Ocean, where samples from the ice-covered season are rare. In the present study, we investigated zooplankton composition based on swimmers and sinkers collected by sediment traps at water depths of 180-280, 800-1320, and 2320-2550 m, over a period of 16 yr (2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007)(2008)(2009)(2010)(2011)(2012)(2013)(2014)(2015)(2016) at the Long-Term Ecological Research observatory HAUSGARTEN located in the eastern Fram Strait (79 N, 4 E). The time-series data showed seasonal and interannual trends within the dominant zooplankton groups including copepoda, foraminifera, ostracoda, amphipoda, pteropoda, and chaetognatha. Amphipoda and copepoda dominated the abundance of swimmers while pteropoda and foraminifera were the most important sinkers. Although the seasonal occurrence of these groups was relatively consistent between years, there were notable interannual variations in abundance, suggesting the influence of various environmental conditions such as sea-ice dynamic and lateral advection of water masses, for example, meltwater and Atlantic water. Statistical analyses revealed a correlation between the Arctic dipole climatic index and sea-ice dynamics (i.e., ice coverage and concentration), as well as the importance of the distance from the ice edge on swimmer composition patterns and carbon export.