Despite evidence of internal waves in the NW Iberian upwelling region, their action and role on nutrient supply dynamics and phytoplankton community structure remain unexplored. A multidisciplinary approach, combining analysis of Synthetic Aperture Radar (SAR) images acquired during the summer months of 2008-2011, together with high-frequency samplings carried out in the R ıa de Vigo in August 2013 during spring (CHAOS1) and neap tides (CHAOS2), was used to characterize: (1) the internal wave activity, (2) its influence on mixing and nutrient supply, and (3) its role on phytoplankton community. SAR images revealed that internal waves were more energetic during spring tides. Turbulent mixing was higher during CHAOS1-springs (Kz 51.3 [1.0-2.0, 95% confidence interval] 3 10 23 m 2 s 21 ) compared to CHAOS2-neaps (Kz 5 0.7 [0.5-1.0] 3 10 23 m 2 s 21 ), and as a result nitrate diffusive fluxes were approximately fourfold higher (35 [17-73] mmol m 22 d 21 ) during CHAOS1-springs. The sampling covered a transition from relaxation-stratification (CHAOS1springs) to intensifying upwelling (CHAOS2-neaps) conditions, resulting in nitrate supply (including both diffusive and advective fluxes) being about 50% higher during CHAOS2-neaps. The phytoplankton community, which was overwhelmingly dominated by diatoms in both cruises, exhibited a shift in species composition, with an increase in the abundance of large Chaetoceros spp. during CHAOS2-neaps. About 50% of the primary production in the ecosystem during periods of upwelling relaxation-stratification could be sustained by enhanced nitrate diffusive fluxes during spring tides. Therefore, even in coastal upwelling regions, turbulent mixing driven by internal waves could play an important role in controlling phytoplankton productivity and community structure.

Abstract. The effect of inorganic nutrients on planktonic assemblages has traditionally relied on concentrations rather than estimates of nutrient supply. We combined a novel dataset of hydrographic properties, turbulent mixing, nutrient concentration, and picoplankton community composition with the aims of (i) quantifying the role of temperature, light, and nitrate fluxes as factors controlling the distribution of autotrophic and heterotrophic picoplankton subgroups, as determined by flow cytometry, and (ii) describing the ecological niches of the various components of the picoplankton community. Data were collected at 97 stations in the Atlantic Ocean, including tropical and subtropical open-ocean waters, the northwestern Mediterranean Sea, and the Galician coastal upwelling system of the northwest Iberian Peninsula. A generalized additive model (GAM) approach was used to predict depth-integrated biomass of each picoplankton subgroup based on three niche predictors: sea surface temperature, averaged daily surface irradiance, and the transport of nitrate into the euphotic zone, through both diffusion and advection. In addition, niche overlap among different picoplankton subgroups was computed using nonparametric kernel density functions. Temperature and nitrate supply were more relevant than light in predicting the biomass of most picoplankton subgroups, except for Prochlorococcus and low-nucleic-acid (LNA) prokaryotes, for which irradiance also played a significant role. Nitrate supply was the only factor that allowed the distinction among the ecological niches of all autotrophic and heterotrophic picoplankton subgroups. Prochlorococcus and LNA prokaryotes were more abundant in warmer waters (>20 ∘C) where the nitrate fluxes were low, whereas Synechococcus and high-nucleic-acid (HNA) prokaryotes prevailed mainly in cooler environments characterized by intermediate or high levels of nitrate supply. Finally, the niche of picoeukaryotes was defined by low temperatures and high nitrate supply. These results support the key role of nitrate supply, as it not only promotes the growth of large phytoplankton, but it also controls the structure of marine picoplankton communities.

We investigate the role of mixing, through its effect on nutrient and light availability, as a driver of phytoplankton community composition in the context of Margalef's mandala. Data on microstructure turbulence, irradiance, new nitrogen supply and phytoplankton composition were collected at 102 stations in three contrasting marine environments: the Galician coastal upwelling system of the northwest Iberian Peninsula, the northwestern Mediterranean, and the tropical and subtropical Atlantic, Pacific and Indian oceans. Photosynthetic pigments concentration and microscopic analysis allowed us to investigate the contribution of diatoms, dinoflagellates, pico-and nanoeukaryotes, and cyanobacteria to the phytoplankton community. Simple linear regression was used to assess the role of environmental factors on community composition, and environmental overlap among different phytoplankton groups was computed using nonparametric kernel density functions. Mixing and new nitrogen supply played an important role in controlling the phytoplankton community structure. At lower values of mixing and new nitrogen supply cyanobacteria dominated, pico-and nanoeukaryotes were dominant across a wide range of environmental conditions, and finally enhanced new nitrogen supply was favourable for diatoms and dinoflagellates. Dinoflagellates were prevalent at intermediate mixing levels, whereas diatoms spread across a wider range of mixing conditions. Occasional instances of enhanced diatom biomass were found under low mixing, associated with the high abundance of Hemiaulus hauckii co-occurring with high N 2 fixation in subtropical regions, and with the formation of thin layers in the Galician coastal upwelling. Our results verify the Margalef's mandala for the whole phytoplankton community, emphasizing the need to consider nutrient supply, rather than nutrient concentration, as an indicator of nutrient availability.

With two 25‐hour series of turbulent microstructure and currents observations carried out in August 2013, during spring (CHAOS 1) and neap tides (CHAOS 2), we investigated the semidiurnal cycle of turbulent dissipation in an embayment affected by coastal upwelling (Ría de Vigo, NW Iberia). At the time of sampling, the bay hosted a net, wind‐driven bi‐directional positive exchange flow and thermal stratification. Turbulent kinetic energy (TKE) dissipation (ɛ) at the interface between upwelled and surface waters was enhanced by two orders of magnitude during the ebbs ( ∼10−6 W kg− 1) with respect to the floods ( ∼10−8 W kg− 1). This pattern was caused by the constructive interference of the shear associated with the upwelling and tidal currents. The vertical structure of the tidal currents was consistent with a deformation of tidal ellipses by stratification, which was tightly coupled to the intensity of upwelling. This two‐pronged interaction resulted in a modulation of the semidiurnal cycle of turbulent dissipation by coastal upwelling. Thus, as a result of the upwelling relaxation conditions experienced during CHAOS 1, depth‐integrated interior TKE dissipation rates were higher, by a factor of ∼2, compared to CHAOS 2. By using a simple model, we determined that observed variations in turbulent mixing had a limited influence on the tidal variations of stratification, which were dominated by straining and advection. The mixing mechanism described here is potentially relevant for the ecology of upwelling bays, as it can stimulate the transport of nutrients from deep‐upwelled waters to the sun‐lit surface layers where primary production takes place.

Difficulties to quantify ocean turbulence have limited our knowledge about the magnitude and variability of nitrate turbulent diffusion, which constitutes one of the main processes responsible for the supply of nitrogen to phytoplankton inhabiting the euphotic zone. We use an extensive dataset of microturbulence observations collected in contrasting oceanic regions, to build a model for nitrate diffusion into the euphotic zone, and obtain the first global map for the distribution of this process. A model including two predictors (surface temperature and nitrate vertical gradient) explained 50% of the variance in the nitrate diffusive flux. This model was applied to climatological data to predict nitrate diffusion in oligotrophic mid and low latitude regions. Mean nitrate diffusion (~ 20 Tmol N y−1) was comparable to nitrate entrainment due to seasonal mixed-layer deepening between 40°N–40ºS, and to the sum of global estimates of nitrogen fixation, fluvial fluxes and atmospheric deposition. These results indicate that nitrate diffusion represents one of the major sources of new nitrogen into the surface ocean in these regions.