Carbon and nitrogen assimilation in the Bering Sea clams Nuculana radiata and Macoma moesta

We analyzed bulk carbon and nitrogen stable isotope values (delta C-13 and delta N-15) of the benthic clams Nuculana radiata and Macoma moesta from the Bering Sea during controlled feeding experiments (spring of 2009 and 2010) using isotopically labeled sea ice algae. The aim was to determine the ability of these clam species to assimilate carbon and nitrogen from sea ice algae. Specimens were collected in the Bering Sea and placed into jars without sediment (2009, N. radiata only) or into natural sediment cores (2010, both species). The clams were offered isotopically enriched (both C and N) or non-enriched algal feeds for time periods of 42 (2009) and 18 d (2010). Isotopic assimilation rates for carbon and nitrogen were calculated using the change in the isotope ratios of the clams over the experimental time. N. radiata in the jar experiments had slow isotopic assimilation rates (0.01 to 0.23 parts per thousand d(-1)), with solvent-extractable organic matter/lipids taking up both of the isotope markers fastest and muscle tissue the slowest. Lipids may thus be particularly suitable to track the immediate ingestion of sea ice algal production in benthic consumers. M. moesta showed 30% higher isotopic assimilation compared to N. radiata in sediment cores, likely reflecting the different feeding behaviors of these two species. Based on our results, N. radiata is likely better able to utilize food sources buried in the sediment and may be more competitive over the sediment surface feeding M. moesta under conditions of reduced ice algal production in the northern Bering Sea. (C) 2012 Elsevier B.V. All rights reserved.This project was undertaken as a part of the National Science Foundation funded Bering Sea Ecosystem Study (BEST Project) #0732767awarded to R. Gradinger, K. Iken, and B. Bluhm at the University of Alaska Fairbanks. Additional research funding to J. Weems was provided by the UAF Center for Global Change Student Research Grant with funds from the Cooperative Institute for Alaska Research and the Harland and Genevieve Emerson Foundation of Adel, Iowa. Bering Sea field support and operations in 2009 and 2010 were provided by chief scientists L. Cooper, J. Grebmeier, and C. Ashjian, and the vessels and crews of the United States Coast Guard Cutters Healy and Polar Sea. Additionally, field and laboratory aid was provided by B. Bluhm, M. Kaufman, B. McConnell, M. Schuster, and J. von Meltzer. Services rendered by the Alaska Stable Isotope Facility and its staff, N. Haubenstock and T. Howe, were very much appreciated. Academic support for J. Weems was provided by the School of Fisheries and Ocean Sciences at the University of Alaska Fairbanks. We thank M. Castellini and two anonymous reviewers for comments on earlier versions of this manuscript. Support for the fatty acid analyses and partial support forWooller came from NSF grant ARC 0902177 awarded to (Gradinger, Iken and Wooller). [ST]Ye

Timing of Ice Algal Grazing by the Arctic Nearshore Benthic Amphipod Onisimus litoralis

Sea ice algae have been widely discussed as a potential food source for pelagic and benthic animals in ice-covered waters, specifically in the light of current substantial changes in the Arctic ice regime. Stomach and gut contents of the Arctic nearshore lysianassid amphipod Onisimus litoralissampled from February to May 2003 indicate that Arctic ice algae were dominant food no earlier than the onset of ice melt. Crustaceans, common prey in a previous study, were absent in stomachs and guts during the survey period. Our data support the concept that sea ice-derived organic carbon is of specific relevance for Arctic plankton and benthos during the period of ice melt.On a souvent discuté du fait que les algues de glace de mer pourraient être une source d'alimentation pour les animaux pélagiques et benthiques des eaux recouvertes de glace, surtout à la lumière des changements considérables qui caractérisent le régime de glaciation de l'Arctique. Le contenu de l'estomac et des intestins des amphipodes lysianassides Onisimus litoralisen proximité des rives de l'Arctique qui a été échantillonné de février à mai 2003 indique que les algues de glace de l'Arctique étaient dominantes pas plus tôt qu'au moment de la fonte des glaces. Les crustacés, qui étaient une proie courante dans le cadre d'une étude antérieure, ne se retrouvaient pas dans les estomacs et les intestins des animaux au cours de la période d'échantillonnage. Nos données viennent soutenir le concept selon lequel le carbone organique dérivé de la glace de mer revêt une importance particulière pour le plancton de l'Arctique et le benthos pendant la période de la fonte des glaces

What we do in the dark: Prevalence of omnivorous feeding activity in Arctic zooplankton during polar night

During the productive polar day, zooplankton and sea-ice amphipods fulfill a critical role in energy transfer from primary producers to higher trophic-level species in Arctic marine ecosystems. Recent polar night studies on zooplankton and sea-ice amphipods suggest higher levels of biological activity than previously assumed. However, it is unknown if these invertebrates maintain polar night activity on stored lipids, opportunistic feeding, or a combination of both. To assess how zooplankton (copepods, amphipods, and krill) and sea-ice amphipods support themselves on seasonally varying resources, we studied their lipid classes, fatty acid compositions, and compound-specific stable isotopes of trophic biomarker fatty acids during polar day (June/July) and polar night (January). Lipid storage and fatty acid results confirm previously described dietary sources in all species during polar day. We found evidence of polar night feeding in all species, including shifts from herbivory to omnivory. Sympagic-, pelagic-, and Calanus spp.-derived carbon sources supported zooplankton and sea-ice amphipods in both seasons. We provide a first indication of polar night feeding of sea-ice amphipods in the pelagic realm

Ice-Associated Amphipods in a Pan-Arctic Scenario of Declining Sea Ice

Sea-ice macrofauna includes ice amphipods and benthic amphipods, as well as mysids. Amphipods are important components of the sympagic food web, which is fuelled by the production of ice algae. Data on the diversity of sea-ice biota have been collected as a part of scientific expeditions over decades, and here we present a pan-Arctic analysis of data on ice-associated amphipods and mysids assimilated over 35 years (1977–2012). The composition of species differed among the 13 locations around the Arctic, with main differences between basins and shelves and also between communities in drift ice and landfast sea ice. The sea ice has been dramatically reduced in extent and thickness during the recorded period, which has resulted in reduced abundance of ice amphipods as well as benthic amphipods in sea ice from the 1980’s to the 2010’s. The decline mainly involved Gammarus wilkitzkii coinciding with the disappearance of much of the multiyear sea ice, which is an important habitat for this long-lived species. Benthic amphipods were most diverse, and also showed a decline over the time-span. They had higher abundance closer to land where they are associated with landfast ice. However, they also occurred in the Central Arctic Ocean, which is likely related to the origin of sea ice over shallow water and subsequent transport in the transpolar ice drift. Recent sampling in the waters east and north of Svalbard has found continued presence of Apherusa glacialis, but almost no G. wilkitzkii. Monitoring by standardized methods is needed to detect further changes in community composition of ice amphipods related to reductions in sea-ice cover and ice type

Terrestrial Inputs Shape Coastal Bacterial and Archaeal Communities in a High Arctic Fjord (Isfjorden, Svalbard)

The Arctic is experiencing dramatic changes including increases in precipitation, glacial melt, and permafrost thaw, resulting in increasing freshwater runoff to coastal waters. During the melt season, terrestrial runoff delivers carbon- and nutrient-rich freshwater to Arctic coastal waters, with unknown consequences for the microbial communities that play a key role in determining the cycling and fate of terrestrial matter at the land-ocean interface. To determine the impacts of runoff on coastal microbial (bacteria and archaea) communities, we investigated changes in pelagic microbial community structure between the early (June) and late (August) melt season in 2018 in the Isfjorden system (Svalbard). Amplicon sequences of the 16S rRNA gene were generated from water column, river and sediment samples collected in Isfjorden along fjord transects from shallow river estuaries and glacier fronts to the outer fjord. Community shifts were investigated in relation to environmental gradients, and compared to river and marine sediment microbial communities. We identified strong temporal and spatial reorganizations in the structure and composition of microbial communities during the summer months in relation to environmental conditions. Microbial diversity patterns highlighted a reorganization from rich communities in June toward more even and less rich communities in August. In June, waters enriched in dissolved organic carbon (DOC) provided a niche for copiotrophic taxa including Sulfitobacter and Octadecabacter. In August, lower DOC concentrations and Atlantic water inflow coincided with a shift toward more cosmopolitan taxa usually associated with summer stratified periods (e.g., SAR11 Clade Ia), and prevalent oligotrophic marine clades (OM60, SAR92). Higher riverine inputs of dissolved inorganic nutrients and suspended particulate matter also contributed to spatial reorganizations of communities in August. Sentinel taxa of this late summer fjord environment included taxa from the class Verrucomicrobiae (Roseibacillus, Luteolibacter), potentially indicative of a higher fraction of particle-attached bacteria. This study highlights the ecological relevance of terrestrial runoff for Arctic coastal microbial communities and how its impacts on biogeochemical conditions may make these communities susceptible to climate change

Light and freshwater discharge drive the biogeochemistry and microbial ecology in a sub-Arctic fjord over the Polar night

The polar night has recently received increased attention as a surprisingly active biological season. Yet, polar night microbial ecology is a vastly understudied field. To identify the physical and biogeochemical parameters driving microbial activity over the dark season, we studied a sub-Arctic fjord system in northern Norway from autumn to early spring with detailed monthly sampling. We focused on the impact of mixing, terrestrial organic matter input and light on microbial ecosystem dynamics. Our study highlights strong differences in the key drivers between spring, autumn, and winter. The spring bloom started in March in a fully mixed water column, opposing the traditional critical depth hypothesis. Incident solar radiation was the key driver maximum Chlorophyll was reached in April. The onset of the autumn phytoplankton bloom was controlled by vertical mixing, causing nutrient upwelling and dilution of zooplankton grazers, which had their highest biomass during this time. According to the dilution-recoupling hypothesis grazer dilution reduced grazing stress and allowed the fall bloom formation. Mixing at that time was initiated by strong winds and reduced stratification as a consequence of freezing temperatures and lower freshwater runoff. During the light-limited polar night, the primary production was extremely low but bacteria continued growing on decaying algae, their exudates and also allochthonous organic matter. A melting event in January could have increased input of organic matter from land, supporting a mid-winter bacterial bloom. In conclusion, polar night biogeochemistry and microbial ecology was not only driven by light availability, but strongly affected by variability in reshwater discharge and allochthonous carbon input. With climate change freshwater discharge will increase in the Arctic, which will likely increase importance of the dynamics described in this study

Light and freshwater discharge drive the biogeochemistry and microbial ecology in a sub-Arctic fjord over the Polar night

The polar night has recently received increased attention as a surprisingly active biological season. Yet, polar night microbial ecology is a vastly understudied field. To identify the physical and biogeochemical parameters driving microbial activity over the dark season, we studied a sub-Arctic fjord system in northern Norway from autumn to early spring with detailed monthly sampling. We focused on the impact of mixing, terrestrial organic matter input and light on microbial ecosystem dynamics. Our study highlights strong differences in the key drivers between spring, autumn, and winter. The spring bloom started in March in a fully mixed water column, opposing the traditional critical depth hypothesis. Incident solar radiation was the key driver maximum Chlorophyll was reached in April. The onset of the autumn phytoplankton bloom was controlled by vertical mixing, causing nutrient upwelling and dilution of zooplankton grazers, which had their highest biomass during this time. According to the dilution-recoupling hypothesis grazer dilution reduced grazing stress and allowed the fall bloom formation. Mixing at that time was initiated by strong winds and reduced stratification as a consequence of freezing temperatures and lower freshwater runoff. During the light-limited polar night, the primary production was extremely low but bacteria continued growing on decaying algae, their exudates and also allochthonous organic matter. A melting event in January could have increased input of organic matter from land, supporting a mid-winter bacterial bloom. In conclusion, polar night biogeochemistry and microbial ecology was not only driven by light availability, but strongly affected by variability in reshwater discharge and allochthonous carbon input. With climate change freshwater discharge will increase in the Arctic, which will likely increase importance of the dynamics described in this study

Early spring subglacial discharge plumes fuel under-ice primary production at a Svalbard tidewater glacier

Subglacial upwelling of nutrient-rich bottom water is known to sustain elevated summer primary production in tidewater-glacier-influenced fjord systems. However, the importance of subglacial upwelling during the early spring season has not been considered yet. We hypothesized that subglacial discharge under sea ice is present in early spring and that its flux is sufficient to increase phytoplankton primary productivity. We evaluated the effects of the submarine discharge on primary production in a seasonally fast-ice covered Svalbard fjord (Billefjorden) influenced by a tidewater outlet glacier in April and May 2019. We found clear evidence for subglacial discharge and upwelling. Although the estimated bottom-water entrainment factor (1.6) and total fluxes were lower than in summer studies, we still observed substantial impact on the fjord ecosystem and primary production at this time of the year. The subglacial discharge leads to a salinity-stratified surface water layer and sea ice formation with low bulk salinity and permeability. The combination of the stratified surface layer, a 2-fold higher under-ice irradiance due to thinner snow cover, and higher N and Si concentrations at the glacier front supported phytoplankton primary production 2 orders of magnitude higher (42.6 mg C m−2 d−1) compared to a marine reference site at the fast-ice edge. Reciprocal transplant experiments showed that nutrient supply increased phytoplankton primary production by approximately 30 %. The brackish-water sea ice at the glacier front with its low bulk salinity contained a reduced brine volume, limiting the inhabitable brine channel space and nutrient exchange with the underlying seawater compared to full marine sea ice. Microbial and algal communities were substantially different in subglacial-influenced water and sea ice compared to the marine reference site, sharing taxa with the subglacial outflow water. We suggest that with climate change, the retreat of tidewater glaciers in early spring could lead to decreased under-ice phytoplankton primary production. In contrast, sea ice algae production and biomass may become increasingly important, unless sea ice disappears first, in which case spring phytoplankton primary production may increase

Ice-algal carbon supports harp and ringed seal diets in the European Arctic: evidence from fatty acid and stable isotope markers

Sea-ice declines in the European Arctic have led to substantial changes in marine food webs. To better understand the biological implications of these changes, we quantified the contributions of ice-associated and pelagic carbon sources to the diets of Arctic harp and ringed seals using compound-specific stable isotope ratios of fatty acids in specific primary producer biomarkers derived from sea-ice algae and phytoplankton. Comparison of fatty acid patterns between these 2 seal species indicated clear dietary separation, while the compound-specific stable isotope ratios of the same fatty acids showed partial overlap. These findings suggest that harp and ringed seals target different prey sources, yet their prey rely on ice and pelagic primary production in similar ways. From Bayesian stable isotope mixing models, we estimated that relative contributions of sympagic and pelagic carbon in seal blubber was an average of 69% and 31% for harp seals, and 72% and 28% for ringed seals, respectively. The similarity in the Bayesian estimations also indicates overlapping carbon sourcing by these 2 species. Our findings demonstrate that the seasonal ice-associated carbon pathway contributes substantially to the diets of both harp and ringed seals

Seasonal Variability in the Zooplankton Community Structure in a Sub-Arctic Fjord as Revealed by Morphological and Molecular Approaches

Phyto- and zooplankton in Arctic and sub-Arctic seas show very strong seasonal changes in diversity and biomass. Here we document the seasonal variability in the mesozooplankton community structure in a sub-Arctic fjord in Northern Norway based on monthly sampling between November 2018 and February 2020. We combined traditional morphological zooplankton identification with DNA metabarcoding of a 313 base pair fragment of the COI gene. This approach allowed us to provide the most detailed mesozooplankton species list known for this region across an entire year, including both holo- and meroplankton. The zooplankton community was dominated by small copepods throughout the sampling period both in terms of abundance and relative sequence counts. However, meroplankton was the most diverse group, especially within the phylum polychaeta. We identified four distinct periods based on the seasonal analysis of the zooplankton community composition. The pre-spring bloom period (February–March) was characterized by low abundance and biomass of zooplankton. The spring bloom (April) was characterized by the presence of Calanus young stages, cirripedia and krill eggs. The spring-summer period (May–August) was characterized by a succession of meroplankton and a relatively high abundance of copepods of the genus Calanus spp. Finally, the autumn-winter period (September–December) was characterized by a high copepod diversity and a peak in abundance of small copepods (e.g., Oithona similis, Acartia longiremis, Pseudocalanus acuspes, Pseudocalanus elongatus, Pseudocalanus moultoni, Pseudocalanus minutus). During this period, we also observed an influx of boreal warm-water species which were notably absent during the rest of the year. Both the traditional community analysis and metabarcoding were highly complementary and with a few exceptions showed similar trends in the seasonal changes of the zooplankton community structure