Abstract. Eastern boundary upwelling systems (EBUS) are among the most productive marine ecosystems on Earth. The production of organic material is fueled by upwelling of nutrient-rich deep waters and high incident light at the sea surface. However, biotic and abiotic factors can modify surface production and related biogeochemical processes. Determining these factors is important because EBUS are considered hotspots of climate change, and reliable predictions of their future functioning requires understanding of the mechanisms driving the biogeochemical cycles therein. In this field experiment, we used in situ mesocosms as tools to improve our mechanistic understanding of processes controlling organic matter cycling in the coastal Peruvian upwelling system. Eight mesocosms, each with a volume of ∼55 m3, were deployed for 50 d ∼6 km off Callao (12∘ S) during austral summer 2017, coinciding with a coastal El Niño phase. After mesocosm deployment, we collected subsurface waters at two different locations in the regional oxygen minimum zone (OMZ) and injected these into four mesocosms (mixing ratio ≈1.5 : 1 mesocosm: OMZ water). The focus of this paper is on temporal developments of organic matter production, export, and stoichiometry in the individual mesocosms. The mesocosm phytoplankton communities were initially dominated by diatoms but shifted towards a pronounced dominance of the mixotrophic dinoflagellate (Akashiwo sanguinea) when inorganic nitrogen was exhausted in surface layers. The community shift coincided with a short-term increase in production during the A. sanguinea bloom, which left a pronounced imprint on organic matter C : N : P stoichiometry. However, C, N, and P export fluxes did not increase because A. sanguinea persisted in the water column and did not sink out during the experiment. Accordingly, export fluxes during the study were decoupled from surface production and sustained by the remaining plankton community. Overall, biogeochemical pools and fluxes were surprisingly constant for most of the experiment. We explain this constancy by light limitation through self-shading by phytoplankton and by inorganic nitrogen limitation which constrained phytoplankton growth. Thus, gain and loss processes remained balanced and there were few opportunities for blooms, which represents an event where the system becomes unbalanced. Overall, our mesocosm study revealed some key links between ecological and biogeochemical processes for one of the most economically important regions in the oceans.

The subtropical oceans are home to one of the largest ecosystems on Earth, contributing to nearly one third of global oceanic primary production. Ocean warming leads to enhanced stratification in the oligotrophic ocean but also intensification in cross-shore wind gradients and thus in eddy kinetic energy across eastern boundary regions of the subtropical gyres. Phytoplankton thriving in a future warmer oligotrophic subtropical ocean with enhanced CO 2 levels could therefore be patchily fertilized by increased mesoscale and submesoscale variability inducing nutrient pumping into the surface ocean. Under this premise, we have tested the response of three size classes (0.2-2, 2-20, and >20 µm) of subtropical phytoplankton communities in terms of primary production, chlorophyll and cell biomass, to increasing CO 2 concentrations and nutrient fertilization during an in situ mesocosm experiment in oligotrophic waters off of the island of Gran Canaria. We found no significant CO 2 -related effect on primary production and biomass under oligotrophic conditions (phase I). In contrast, primary production, chlorophyll and biomass displayed a significant and pronounced increase under elevated CO 2 conditions in all groups after nutrient fertilization, both during the bloom (phase II) and post-bloom (phase III) conditions. Although the relative increase of primary production in picophytoplankton (250%) was 2.5 higher than in microphytoplankton (100%) after nutrient fertilization, comparing the high and low CO 2 treatments, microphytoplankton dominated in terms of biomass, contributing >57% to the total. These results contrast with similar studies conducted in temperate and cold waters, where consistently small phytoplankton benefitted after nutrient additions at high CO 2 , pointing to different CO 2 -sensitivities across plankton communities and ecosystem types in the ocean.

Reduction of anthropogenic CO2 emissions alone will not sufficiently restrict global warming and enable the 1.5°C goal of the Paris agreement to be met. To effectively counteract climate change, measures to actively remove carbon dioxide from the atmosphere are required. Artificial upwelling has been proposed as one such carbon dioxide removal technique. By fueling primary productivity in the surface ocean with nutrient-rich deep water, it could potentially enhance downward fluxes of particulate organic carbon (POC) and carbon sequestration. In this study we investigated the effect of different intensities of artificial upwelling combined with two upwelling modes (recurring additions vs. one singular addition) on POC export, sinking matter stoichiometry and remineralization depth. We carried out a 39 day-long mesocosm experiment in the subtropical North Atlantic, where we fertilized oligotrophic surface waters with different amounts of deep water. The total nutrient inputs ranged from 1.6 to 11.0 μmol NO3– L–1. We found that on the one hand POC export under artificial upwelling more than doubled, and the molar C:N ratios of sinking organic matter increased from values around Redfield (6.6) to ∼8–13, which is beneficial for potential carbon dioxide removal. On the other hand, sinking matter was remineralized at faster rates and showed lower sinking velocities, which led to shallower remineralization depths. Particle properties were more favorable for deep carbon export in the recurring upwelling mode, while in the singular mode the C:N increase of sinking matter was more pronounced. In both upwelling modes roughly half of the produced organic carbon was retained in the water column until the end of the experiment. This suggests that the plankton communities were still in the process of adjustment, possibly due to the different response times of producers and consumers. There is thus a need for studies with longer experimental durations to quantify the responses of fully adjusted communities. Finally, our results revealed that artificial upwelling affects a variety of sinking particle properties, and that the intensity and mode with which it is applied control the strength of the effects.

Diatoms are silicifying phytoplankton contributing about one quarter to primary production on Earth. Ocean acidification (OA) could alter the competitiveness of diatoms relative to other taxa and/or lead to shifts among diatom species. In spring 2016, we set up a plankton community experiment at the coast of Gran Canaria (Canary Islands, Spain) to investigate the response of subtropical diatom assemblages to elevated seawater pCO 2. Therefore, natural plankton communities were enclosed for 32 days in in situ mesocosms (∼8 m 3 volume) with a pCO 2 gradient ranging from 380 to 1140 µatm. Halfway through the study we added nutrients to all mesocosms (N, P, Si) to simulate injections through eddy-induced upwelling which frequently occurs in the region. We found that the total diatom biomass remained unaffected during oligotrophic conditions but was significantly positively affected by high CO 2 after nutrient enrichment. The average cell volume and carbon content of the diatom community increased with CO 2. CO 2 effects on diatom biomass and species composition were weak during oligotrophic conditions but became quite strong above ∼620 µatm after the nutrient enrichment. We hypothesize that the proliferation of diatoms under high CO 2 may have been caused by a fertilization effect on photosynthesis in combination with reduced grazing pressure. Our results suggest that OA in the subtropics may strengthen the competitiveness of (large) diatoms and cause changes in diatom community composition, mostly under conditions when nutrients are injected into oligotrophic systems.

Cyclonic and anticyclonic eddies are common mesoscale features in the flow past the Canary Islands throughout the year. While drifting southward, eddy pairs interact among them but also with upwelling filaments and eddies generated at the coastal jet of the nearby African upwelling system. These interactions force the generation of frontal zones where ageostrophic secondary circulation (ASC) may occur. With the aim of contributing to understand how meso-submesoscale interactions modulate plankton distribution, we carried out an interdisciplinary cruise across a mesoscale eddy field. The sampled region was characterized by the presence of a cyclonic eddy interacting with two anticyclonic eddies and an upwelling filament. High-resolution sampling allowed us to assess the upwelling/downwelling processes associated with eddy pumping and ASC, the injection of nutrients into the euphotic zone, and the subduction of particles related to these processes. The planktonic community, which included heterotrophic bacteria, cyanobacteria-like Prochlorococcus and Synechococcus, pico and nanoautotrophic eukaryotes, and heterotrophic nanoflagellates, showed a heterogeneous distribution in response to meso-submesoscale processes. Redundancy analysis and plankton distribution suggest that while the distribution of small organisms (picoplankton) is modulated by a combination of physical and biogeochemical drivers, the distribution of larger autotrophic and heterotrophic nanoflagellates is modulated by nutrient inputs and grazing, respectively. These observational results provide new insights in the study of the impact of mesoscale structures in the dynamics of nutrients, chlorophyll and planktonic communities, and valuable to validate theoretical and modeling studies.