Diatoms are one of the major primary producers in the ocean, responsible annually for ~20% of photosynthetically fixed CO2 on Earth. In oceanic models, they are typically represented as large (>20 µm) microphytoplankton. However, many diatoms belong to the nanophytoplankton (2–20 µm) and a few species even overlap with the picoplanktonic size-class (<2 µm). Due to their minute size and difficulty of detection they are poorly characterized. Here we describe a massive spring bloom of the smallest known diatom (Minidiscus) in the northwestern Mediterranean Sea. Analysis of Tara Oceans data, together with literature review, reveal a general oversight of the significance of these small diatoms at the global scale. We further evidence that they can reach the seafloor at high sinking rates, implying the need to revise our classical binary vision of pico- and nanoplanktonic cells fueling the microbial loop, while only microphytoplankton sustain secondary trophic levels and carbon export.

International audienceThe biological composition of the material exported to a moored sediment trap located under the winter mixed layer of the naturally fertilized Kerguelen Plateau in the Southern Ocean was studied over an annual cycle. Despite iron availability in spring, the annual particulate organic carbon (POC) export (98.2 mmol m−2) at 289 m was low, but annual biogenic silica export was significant (114 mmol m−2). This feature was related to the abundance of empty diatom cells and the ratio of full to empty cells exerted a first-order control in BSi : POC export stoichiometry of the biological pump. Chaetoceros Hyalochaete spp. and Thalassiosira antarctica resting spores were responsible for more than 60% of the annual POC flux that occurred during two very short export events of < 14 days in spring–summer. Relatively low diatom fluxes were observed over the remainder of the year. Faecal pellet contribution to annual carbon flux was lower (34%) and reached its seasonal maximum in autumn and winter (> 80%). The seasonal progression of faecal pellet types revealed a clear transition from small spherical shapes (small copepods) in spring, to larger cylindrical and ellipsoid shapes in summer (euphausiids and large copepods) and finally to large tabular shapes (salps) in autumn and winter. We propose in this high-biomass, low-export (HBLE) environment that small but highly silicified and fast-sinking resting spores are able to bypass the intense grazing pressure and efficient carbon transfer to higher trophic levels that are responsible for the low fluxes observed the during the remainder of the year. More generally our study also provides a statistical framework linking the ecological succession of diatom and zooplankton communities to the seasonality of carbon and silicon export within an iron-fertilized bloom region in the Southern Ocean

The Southern Ocean (SO) hosts plankton communities that impact the biogeochemical cycles of the global ocean. However, weather conditions in the SO restrict mainly in situ observations of plankton communities to spring and summer, preventing the description of biological successions at an annual scale. Here, we use shipboard observations collected in the Indian sector of the SO to develop a multivariate relationship between physical and bio‐optical data, and, the composition and carbon content of the plankton community. Then we apply this multivariate relationship to five biogeochemical Argo (BGC‐Argo) floats deployed within the same bio‐geographical zone as the ship‐board observations to describe spatial and seasonal changes in plankton assemblage. The floats reveal a high contribution of bacteria below the mixed layer, an overall low abundance of picoplankton and a seasonal succession from nano‐ to microplankton during the spring bloom. Both naturally iron‐fertilized waters downstream of the Crozet and Kerguelen Plateaus show elevated phytoplankton biomass in spring and summer but they differ by a nano‐ or microplankton dominance at Crozet and Kerguelen, respectively. The estimated plankton group successions appear consistent with independent estimations of particle diameter based on the optical signals. Furthermore, the comparison of the plankton community composition in the surface layer with the presence of large mesopelagic particles diagnosed by spikes of optical signals provides insight into the nature and temporal changes of ecological vectors that drive particle export. This study emphasizes the power of BGC‐Argo floats for investigating important biogeochemical processes at high temporal and spatial resolution.

Hydrothermal activity is significant in regulating the dynamics of trace elements in the ocean. Biogeochemical models suggest that hydrothermal iron might play an important role in the iron-depleted Southern Ocean by enhancing the biological pump. However, the ability of this mechanism to affect large-scale biogeochemistry and the pathways by which hydrothermal iron reach the surface layer have not been observationally constrained. Here we present the first observational evidence of upwelled hydrothermally influenced deep waters stimulating massive phytoplankton blooms in the Southern Ocean. Captured by profiling floats, two blooms were observed in the vicinity of the Antarctic Circumpolar Current, downstream of active hydrothermal vents along the Southwest Indian Ridge. These hotspots of biological activity are supported by mixing of hydrothermally sourced iron stimulated by flow-topography interactions. Such findings reveal the important role of hydrothermal vents on surface biogeochemistry, potentially fueling local hotspot sinks for atmospheric CO 2 by enhancing the biological pump.

The Southern Ocean remains one of the least explored marine environments. The investigation of temporal microbial dynamics has thus far been hampered by the limited access to this remote ocean. We present here high-resolution seasonal observations of the prokaryotic community composition during phytoplankton blooms induced by natural iron fertilization. A total of 18 seawater samples were collected by a moored remote autonomous sampler over 4 months at 5-11 day intervals in offshore surface waters (central Kerguelen Plateau). Illumina sequencing of the 16S rRNA gene revealed that among the most abundant amplicon sequence variants, SAR92 and Aurantivirga were the first bloom responders, Pseudomonadaceae, Nitrincolaceae and Polaribacter had successive peaks during the spring bloom decline, and Amylibacter increased in relative abundance later in the season. SAR11 and SUP05 were abundant prior to and after the blooms. Using network analysis, we identified two groups of diatoms representative of the spring and summer bloom that had opposite correlation patterns with prokaryotic taxa. Our study provides the first seasonal picture of microbial community dynamics in the open Southern Ocean and thereby offers biological insights to the cycling of carbon and iron, and to an important puzzling issue that is the modest nitrate decrease associated to iron fertilization.

Abstract.A sediment trap moored in the naturally ironfertilized Kerguelen Plateau in the Southern Ocean provided an annual record of particulate organic carbon and nitrogen fluxes at 289 m. At the trap deployment depth, current speeds were typically low ( ∼ 10 cm s −1 ) and primarily tidaldriven (M2 tidal component). Although advection was weak, the sediment trap may have been subject to hydrodynamical and biological (swimmer feeding on trap funnel) biases. Particulate organic carbon (POC) flux was generally low (< 0.5 mmol m −2 d −1 ), although two episodic export events (< 14 days) of 1.5 mmol m −2 d −1 were recorded. These increases in flux occurred with a 1-month time lag from peaks in surface chlorophyll and together accounted for approximately 40 % of the annual flux budget. The annual POC flux of 98.2 ± 4.4 mmol m −2 yr −1 was low considering the shallow deployment depth but comparable to independent estimates made at similar depths (∼ 300 m) over the plateau, and to deep-ocean (> 2 km) fluxes measured from similarly productive iron-fertilized blooms. Although undertrapping cannot be excluded in shallow moored sediment trap deployment, we hypothesize that grazing pressure, including mesozooplankton and mesopelagic fishes, may be responsible for the low POC flux beneath the base of the winter mixed layer. The importance of plankton community structure in controlling the temporal variability of export fluxes is addressed in a companion paper.

a b s t r a c tBiogeochemical and diatom export fluxes are presented from two bathypelagic sediment trap deployments in the Antarctic Zone of the Southern Ocean. One of the sediment traps was deployed in very productive, naturally iron-fertilized waters downstream of South Georgia (P3, 2000 m) and compared to a deployment in moderately productive waters upstream of the island system (P2, 1500 m). At both sites significant diatom export events occurred in spring (November) and contained mostly empty cells that were associated with low particulate organic carbon (POC) fluxes. A summer export pulse occurred one month later at P2 (end February/March) compared to P3 (end January). Diatom fluxes at P3 were one order of magnitude higher than at P2, a difference mainly attributed to the short and intense export of resting spores from Chaetoceros Hyalochaete and Thalassiosira antarctica species. Aside from these resting spores, diatom export assemblages at both sites were dominated by empty Fragilariopsis kerguelensis frustules. The fraction of diatoms exported as empty frustules was considerably lower at P3 (52%) than P2 (91%). This difference was related to the flux of intact diatom resting spores at P3 and may partially explain the lower Si:C export stoichiometry observed at P3 (1.1) compared to P2 (1.5). Through the enumeration of full diatom frustules and subsequent biomass calculations we estimate that diatom resting spores account for 42% of annual POC flux in the productive waters downstream of South Georgia. At both sites the contribution of diatom vegetative stages to POC fluxes was considerably lower (o 5%). From these analyses we conclude that resting spore export contributes towards the slightly higher bathypelagic (POC) flux at P3 (40.6 mmol m À 2 y À 1 ) compared to P2 (26.4 mmol m À 2 y À 1 ). We compared our sediment trap records with previously published diatom assemblage data from the mixed layer and surface sediments (3760 m) around South Georgia. The relative proportion of diatom resting spores within diatom assemblages increases as a function of depth and is explained by selective preservation of their robust frustules. Our study highlights the significance of diatom resting spore export as a carbon vector out of the mixed layer. Furthermore, the contribution or resting spores to POC flux in the bathypelagic ocean and sediments suggests they play a particularly important role in sequestering biologically fixed CO 2 over climatically relevant timescales.

This study was a part of the DeWEX project (Deep Water formation Experiment), designed to better understand the impact of dense water formation on the marine biogeochemical cycles. Here, nutrient and phytoplankton vertical and horizontal distributions were investigated during a deep open‐ocean convection event and during the following spring bloom in the Northwestern Mediterranean Sea (NWM). In February 2013, the deep convection event established a surface nutrient gradient from the center of the deep convection patch to the surrounding mixed and stratified areas. In the center of the convection area, a slight but significant difference of nitrate, phosphate and silicate concentrations was observed possibly due to the different volume of deep waters included in the mixing or to the sediment resuspension occurring where the mixing reached the bottom. One of this process, or a combination of both, enriched the water column in silicate and phosphate, and altered significantly the stoichiometry in the center of the deep convection area. This alteration favored the local development of microphytoplankton in spring, while nanophytoplankton dominated neighboring locations where the convection reached the deep layer but not the bottom. This study shows that the convection process influences both winter nutrients distribution and spring phytoplankton distribution and community structure. Modifications of the convection's spatial scale and intensity (i.e., convective mixing depth) are likely to have strong consequences on phytoplankton community structure and distribution in the NWM, and thus on the marine food web.