The importance of turbulent ocean–sea ice nutrient exchanges for simulation of ice algal biomass and production with CICE6.1 and Icepack 1.2

Different sea ice models apply unique approaches in the computation of nutrient diffusion between the ocean and the ice bottom, which are generally decoupled from the calculation of turbulent heat flux. A simple molecular diffusion formulation is often used. We argue that nutrient transfer from the ocean to sea ice should be as consistent as possible with heat transfer, since all of these fluxes respond to varying forcing in a similar fashion. We hypothesize that biogeochemical models that do not consider such turbulent nutrient exchanges between the ocean and the sea ice, despite considering brine drainage and bulk exchanges through ice freezing and melting, may underestimate bottom-ice algal production. The Los Alamos Sea Ice Model (CICE + Icepack) was used to test this hypothesis by comparing simulations without and with diffusion of nutrients across the sea ice bottom that are dependent on velocity shear, implemented in a way that is consistent with turbulent heat exchanges. Simulation results support the hypothesis, showing a significant enhancement of ice algal production and biomass when nutrient limitation was relieved by bottom-ice turbulent exchange. Our results emphasize the potentially critical role of turbulent exchanges to sea ice algal blooms and thus the importance of properly representing them in biogeochemical models. The relevance of this becomes even more apparent considering ongoing trends in the Arctic Ocean, with a predictable shift from light-limited to nutrient-limited growth of ice algae earlier in the spring, as the sea ice becomes more fractured and thinner with a larger fraction of young ice with thin snow cover

Microalgal community structure and primary production in Arctic and Antarctic sea ice : A synthesis

Sea ice is one the largest biomes on earth, yet it is poorly described by biogeochemical and climate models. In this paper, published and unpublished data on sympagic (ice-associated) algal biodiversity and productivity have been compiled from more than 300 sea-ice cores and organized into a systematic framework. Significant patterns in microalgal community structure emerged from this framework. Autotrophic flagellates characterize surface communities, interior communities consist of mixed microalgal populations and pennate diatoms dominate bottom communities. There is overlap between landfast and pack-ice communities, which supports the hypothesis that sympagic microalgae originate from the pelagic environment. Distribution in the Arctic is sometimes quite different compared to the Antarctic. This difference may be related to the time of sampling or lack of dedicated studies. Seasonality has a significant impact on species distribution, with a potentially greater role for flagellates and centric diatoms in early spring. The role of sea-ice algae in seeding pelagic blooms remains uncertain. Photosynthesis in sea ice is mainly controlled by environmental factors on a small scale and therefore cannot be linked to specific ice types. Overall, sea-ice communities show a high capacity for photoacclimation but low maximum productivity compared to pelagic phytoplankton. Low carbon assimilation rates probably result from adaptation to extreme conditions of reduced light and temperature in winter. We hypothesize that in the near future, bottom communities will develop earlier in the season and develop more biomass over a shorter period of time as light penetration increases due to the thinning of sea ice. The Arctic is already witnessing changes. The shift forward in time of the algal bloom can result in a mismatch in trophic relations, but the biogeochemical consequences are still hard to predict. With this paper we provide a number of parameters required to improve the reliability of sea-ice biogeochemical models.Peer reviewe

Floating Ice-Algal Aggregates below Melting Arctic Sea Ice

During two consecutive cruises to the Eastern Central Arctic in late summer 2012, we observed floating algal aggregates in the melt-water layer below and between melting ice floes of first-year pack ice. The macroscopic (1-15 cm in diameter) aggregates had a mucous consistency and were dominated by typical ice-associated pennate diatoms embedded within the mucous matrix. Aggregates maintained buoyancy and accumulated just above a strong pycnocline that separated meltwater and seawater layers. We were able, for the first time, to obtain quantitative abundance and biomass estimates of these aggregates. Although their biomass and production on a square metre basis was small compared to ice-algal blooms, the floating ice-algal aggregates supported high levels of biological activity on the scale of the individual aggregate. In addition they constituted a food source for the ice-associated fauna as revealed by pigments indicative of zooplankton grazing, high abundance of naked ciliates, and ice amphipods associated with them. During the Arctic melt season, these floating aggregates likely play an important ecological role in an otherwise impoverished near-surface sea ice environment. Our findings provide important observations and measurements of a unique aggregate-based habitat during the 2012 record sea ice minimum yea

Dependency of Arctic zooplankton on pelagic food sources: New insights from fatty acid and stable isotope analyses

Global warming causes dramatic environmental change to Arctic ecosystems. While pelagic primary production is initiated earlier and its intensity can be increased due to earlier ice melt and extended open-water periods, sea-ice primary production is progressively confined on a spatio-temporal scale, leading to unknown consequences for the ice-associated (sympagic) food web. Understanding ecological responses to changes in the availability and composition of pelagic and sympagic food sources is crucial to determine potential changes of food-web structure and functioning in Arctic marine communities under increasingly ice-free conditions. Focus was placed on the importance of suspended particulate organic matter vs. sympagic organic matter for 12 zooplankton species with different feeding modes covering five taxonomic groups (copepods, krill, amphipods, chaetognaths, and appendicularians) at two ice-covered, but environmentally different, stations in the north-western Barents Sea in August 2019. Contributions of diatom- and flagellate-associated fatty acids (FAs) to total lipid content and carbon stable isotopic compositions of these FAs were used to discriminate food sources and trace flows of organic matter in marine food webs. Combination of proportional contributions of FA markers with FA isotopic composition indicated that consumers mostly relied, directly (herbivorous species), or indirectly (omnivorous and carnivorous species), on pelagic diatoms and flagellates, independently of environmental conditions at the sampling locations, trophic position, and feeding mode. Differences were nevertheless observed between species. Contrary to other studies demonstrating a high importance of sympagic organic matter for food-web processes, our results highlight the complexity and variability of trophic structures and dependencies in different Arctic food webs

Nitrate isotope investigations reveal future impacts of climate change on nitrogen inputs and cycling in Arctic fjords: Kongsfjorden and Rijpfjorden (Svalbard)

Ongoing climate change in the Arctic has caused tidewater glaciers to retreat while increasing the discharge of freshwater and terrestrial material into fjords. This can affect both nutrient inputs and cycling within the fjord systems. In particular, tidewater glaciers and the presence of associated subglacial meltwater plumes can have a large impact on fjord circulation and biogeochemistry. In this study, we assess the influence of tidewater glaciers on nitrogen inputs and cycling in two fjords in Svalbard during the summer using stable isotopic analyses of dissolved nitrate (δ15N and δ18O) in combination with nutrient and hydrographic data. Kongsfjorden receives inputs from tidewater glaciers, whereas Rijpfjorden mainly receives surface inputs from land-terminating glaciers. Results showed that both fjords are enriched in nutrients from terrestrial inputs. Nutrient ratios indicate excess Si and P relative to N. In both fjords, terrestrial nitrate from snowpack and glacier melting are identified as the dominant sources based on high δ18O-NO3- and low δ15N-NO3- of dissolved nitrate. In Kongsfjorden, mixed-layer nitrate is completely consumed within the fjord system, which we attribute to vigorous circulation at the glacial front influenced by the subglacial plume and longer residence time in the fjord. This is in contrast to Rijpfjorden where nutrients are only partially consumed perhaps due to surface river discharge and light limitation. In Kongsfjorden, we estimate terrestrial and marine N contributions to the nitrate pool from nitrogen isotopic values (δ15N-NO3-), and this suggests that nearly half the nitrate in the subglacial plume (50 ± 3 %) and the water column (44 ± 3 %) originates from terrestrial sources. We show that terrestrial N contributes significantly to the regenerated N pool (63 %–88 %) within this fjord suggesting its importance in sustaining productivity here. Given this importance of terrestrial nutrient sources within the fjords, increase in these inputs due to climate change can enhance the fjord nutrient inventory, productivity and nutrient export offshore. Specifically, increasing Atlantification and warmer Atlantic Water will encourage tidewater glacier retreat and in turn increase surface discharge. In fjords akin to Rijpfjorden this is expected to foster more light limitation and less dynamic circulation, ultimately aiding the export of nutrients offshore contributing to coastal productivity. Climate change scenarios postulated for fjords such as Kongsfjorden include more terrestrial N-fuelled productivity and N cycling within the fjord, less vigorous circulation due to the retreat of tidewater glaciers, and the expansion of oxygen-depleted deep waters isolated by the sill.</p

Tidewater glaciers as “climate refugia” for zooplankton-dependent food web in Kongsfjorden, Svalbard

With climate warming, many tidewater glaciers are retreating. Fresh, sediment-rich sub-glacial meltwater is discharged at the glacier grounding line, where it mixes with deep marine water resulting in an upwelling of a plume visible in front of the glacial wall. Zooplankton may suffer increased mortality within the plume due to osmotic shock when brought in contact with the rising meltwater. The constant replenishment of zooplankton and juvenile fish to the surface areas attracts surface-foraging seabirds. Because access to other feeding areas, such as the marginal ice zone, has become energetically costly due to reduced sea-ice extent, glacial plumes may become increasingly important as “climate refugia” providing enhanced prey availability. Here, we investigated zooplankton concentrations within the plume and adjacent waters of four tidewater glaciers in Kongsfjorden, Svalbard, in early August 2016 and late July 2017. Our aim was to compare the zooplankton composition, abundance, and isotopic signatures within the plumes to those in adjacent fjord and shelf waters. Our hypothesis was that the plumes resulted in increased zooplankton mortality through osmotic shock and increased prey availability to predators. The mortality due to osmotic shock in the glacial plume was low (<5% dead organisms in samples), although slightly higher than in surrounding waters. This indicates that plumes are inefficient “death traps” for zooplankton. However, the high abundance and biomass of zooplankton within plume areas suggest that the “elevator effect” of rising glacial water supplies zooplankton to the sea surface, thereby enhancing prey availability for surface-feeding seabirds. Thus, our study provides evidence that glacial plumes are important as “climate refugia” for foraging seabirds. Stable isotope signatures showed that the glacial bay zooplankton and fish community represent a distinct isotopic niche. Additionally, zooplankton mortality associated with the plume estimated over 100-days of melt season supports a flux of 12.8 tonnes of organic carbon to benthic communities in the glacial bays. Benthic scavengers, such as Onisimus caricus and Anonyx nugax, were abundant in the glacial bay, where they feed on sinking organic matter.publishedVersio

A Winter-to-Summer Transition of Bacterial and Archaeal Communities in Arctic Sea Ice

The Arctic is warming 2–3 times faster than the global average, leading to a decrease in Arctic sea ice extent, thickness, and associated changes in sea ice structure. These changes impact sea ice habitat properties and the ice-associated ecosystems. Sea-ice algal blooms provide various algal-derived carbon sources for the bacterial and archaeal communities within the sea ice. Here, we detail the transition of these communities from winter through spring to early summer during the Norwegian young sea ICE (N-ICE2015) expedition. The winter community was dominated by the archaeon Candidatus Nitrosopumilus and bacteria belonging to the Gammaproteobacteria (Colwellia, Kangiellaceae, and Nitrinocolaceae), indicating that nitrogen-based metabolisms, particularly ammonia oxidation to nitrite by Cand. Nitrosopumilus was prevalent. At the onset of the vernal sea-ice algae bloom, the community shifted to the dominance of Gammaproteobacteria (Kangiellaceae, Nitrinocolaceae) and Bacteroidia (Polaribacter), while Cand. Nitrosopumilus almost disappeared. The bioinformatically predicted carbohydrate-active enzymes increased during spring and summer, indicating that sea-ice algae-derived carbon sources are a strong driver of bacterial and archaeal community succession in Arctic sea ice during the change of seasons. This implies a succession from a nitrogen metabolism-based winter community to an algal-derived carbon metabolism-based spring/ summer community

Carbon export in the seasonal sea ice zone north of Svalbard from winter to late summer

Phytoplankton blooms in the Arctic Ocean's seasonal sea ice zone are expected to start earlier and occur further north with retreating and thinning sea ice cover. The current study is the first compilation of phytoplankton bloom development and fate in the seasonally variable sea ice zone north of Svalbard from winter to late summer, using short-term sediment trap deployments. Clear seasonal patterns were discovered, with low winter and pre-bloom phytoplankton standing stocks and export fluxes, a short and intense productive season in May and June, and low Chl a standing stocks but moderate carbon export fluxes in the autumn post-bloom conditions. We observed intense phytoplankton blooms with Chl a standing stocks of >350 mg m−2 below consolidated sea ice cover, dominated by the prymnesiophyte Phaeocystis pouchetii. The largest vertical organic carbon export fluxes to 100 m, of up to 513 mg C m−2 day−1, were recorded at stations dominated by diatoms, while those dominated by P. pouchetii recorded carbon export fluxes up to 310 mg C m−2 day−1. Fecal pellets from krill and copepods contributed a substantial fraction to carbon export in certain areas, especially where blooms of P. pouchetii dominated and Atlantic water advection was prominent. The interplay between the taxonomic composition of protist assemblages, large grazers, distance to open water, and Atlantic water advection was found to be crucial in determining the fate of the blooms and the magnitude of organic carbon exported out of the surface water column. Previously, the marginal ice zone was considered the most productive region in the area, but our study reveals intense blooms and high export events in ice-covered waters. This is the first comprehensive study on carbon export fluxes for under-ice phytoplankton blooms, a phenomenon suggested to have increased in importance under the new Arctic sea ice regime