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Microalgae growing within and attached to the bottom of Arctic sea ice (sympagic algae) can serve as a nutritious food resource for animals inhabiting the sea-ice water interface (under-ice fauna), particularly during the bottom ice-algal bloom in spring. As a consequence, under-ice fauna is likely impacted by sea-ice decline and changes in ice-algal primary production. To investigate this, samples of pelagic (=PPOM) and ice-associated particulate organic matter (=IPOM) and the ice-associated amphipods Apherusa glacialis and Eusirus holmii, and polar cod (Boreogadus saida), collected below ridged sea ice at two locations with pronounced differences in productivity in the northern Barents Sea during May 2021, were assessed for their trophic marker content. Specifically, we investigated the composition of diatom- and dinoflagellate-produced fatty acids (FAs), pelagic and sympagic highly branched isoprenoid (HBI) lipids as well as sterols to determine the animals’ dietary preferences and trophic association to the sea-ice habitat during spring. Relative proportions of FAs differed strongly between PPOM and IPOM, indicating differences in species composition and degradation state between pelagic and sympagic habitats, respectively. FA signatures and sterol content of the consumers largely resembled known diet compositions with a strong reliance on diatom-derived carbon in A. glacialis, a higher degree of carnivory in E. holmii and evidence of Calanus-feeding in polar cod. Sympagic HBIs were detected at either low concentrations or not at all, in both producers and consumers, likely as a result of the very low abundance of their source diatoms. Pronounced trophic marker variability in A. glacialis collected at the highly productive shelf slope station versus the less productive central Arctic Basin station suggests a surprisingly high flexibility in carbon-source composition with a stronger reliance on pelagic food when available versus a higher importance of ice algal carbon when pelagic production is low. Nevertheless and despite the general lack (below detection limit) of sympagic HBIs in our dataset, high ice-algal biomass and elevated proportions of polyunsaturated FAs in IPOM compared to other seasons indicate that ice algae constitute a valuable nutritional carbon source as alternative to pelagic carbon during spring.

Over the past decades, two key grazers in the Southern Ocean (SO), krill and salps, have experienced drastic changes in their distribution and abundance, leading to increasing overlap of their habitats. Both species occupy different ecological niches and long-term shifts in their distributions are expected to have cascading effects on the SO ecosystem. However, studies directly comparing krill and salps are lacking. Here, we provide a direct comparison of the diet and fecal pellet composition of krill and salps using 18S metabarcoding and fatty acid markers. Neither species’ diet reflected the composition of the plankton community, suggesting that in contrast to the accepted paradigm, not only krill but also salps are selective feeders. Moreover, we found that krill and salps had broadly similar diets, potentially enhancing the competition between both species. This could be augmented by salps’ ability to rapidly reproduce in favorable conditions, posing further risks to krill populations.

The thecosome pteropods Limacina helicina and L. retroversa are important contributors to the zooplankton community in high-latitude environments but little is known about their distribution and life cycle under polar conditions. We collected the early life stages (< 1 mm) of the thecosome population in 2012 and 2013 at a bi-weekly to monthly resolution in fjord highly influenced by Arctic waters as well as Atlantic inflows (Adventfjorden, Svalbard, 78°N), together with environmental parameters. L. retroversa only occurred episodically, in association with the inflow of Atlantic water, with low numbers and random size distributions. This suggests that this boreal species does not fulfill its life cycle in Adventfjorden. In contrast, young specimens of L. helicina were present during the entire study. Veligers hatched in late summer/autumn and measured 0.14 mm on average. They grew with rates of 0.0006 mm day− 1 over the 10–11 months of development. Only thereafter, growth accelerated by one order of magnitude and maximal rates were reached in autumn (0.0077 mm day− 1). Our results indicate that L. helicina reaches a size of 1 mm after approximately 1.5 years in Adventfjorden. We therefore suggest that L. helicina overwinters the first year as a small juvenile and that it needs at least 2 years to reach an adult size of 5 mm in Adventfjorden. This reveals an complex and delicate aspect of the life-cycle of L. helicina and further research is needed to determine if it makes the population especially vulnerable towards climate changes.

Biogeochemical markers in combination with bacterial community composition were studied at two contrasting stations at the Río Negro (RN) estuary to assess the outwelling hypothesis in the Argentinian Patagonia. Inorganic nutrients and dissolved organic matter were exported clearly during the last hours of the ebb at the station Wetland. Moreover, a considerable outwelling of polyunsaturated fatty acids (PUFA), particulates and microalgae was inferred by this combined approach. The exported 22:6(n-3) and 20:5(n-3) contributed very likely to sustain higher trophic levels in the coasts of the Southwest Atlantic. The stable isotopes did not evidence clearly the outwelling; nevertheless, the combination of δ

Sea ice continues to decline across many regions of the Arctic, with remaining ice becoming increasingly younger and more dynamic. These changes alter the habitats of microbial life that live within the sea ice, which support healthy functioning of the marine ecosystem and provision of resources for human-consumption, in addition to influencing biogeochemical cycles (e.g. air–sea CO2 exchange). With the susceptibility of sea ice ecosystems to climate change, there is a pressing need to fill knowledge gaps surrounding sea ice habitats and their microbial communities. Of fundamental importance to this goal is the development of new methodologies that permit effective study of them. Based on outcomes from the DiatomARCTIC project, this paper integrates existing knowledge with case studies to provide insight on how to best document sea ice microbial communities, which contributes to the sustainable use and protection of Arctic marine and coastal ecosystems in a time of environmental change. Supplementary Information The online version contains supplementary material available at 10.1007/s13280-021-01658-z.

The hydrography of the Arctic Seas is being altered by ongoing climate change, with knock-on effects to nutrient dynamics and primary production. As the major pathway of exchange between the Arctic and the Atlantic, the Fram Strait hosts two distinct water masses in the upper water column, northward flowing warm and saline Atlantic Waters in the east, and southward flowing cold and fresh Polar Surface Water in the west. Here, we assess how physical processes control nutrient dynamics in the Fram Strait using nitrogen isotope data collected during 2016 and 2018. In Atlantic Waters, a weakly stratified water column and a shallow nitracline reduce nitrogen limitation. To the west, in Polar Surface Water, nitrogen limitation is greater because stronger stratification inhibits nutrient resupply from deeper water and lateral nitrate supply from central Arctic waters is low. A historical hindcast simulation of ocean biogeochemistry from 1970 to 2019 corroborates these findings and highlights a strong link between nitrate supply to Atlantic Waters and the depth of winter mixing, which shoaled during the simulation in response to a local reduction in sea-ice formation. Overall, we find that while the eastern Fram Strait currently experiences seasonal nutrient replenishment and high primary production, the loss of winter sea ice and continued atmospheric warming has the potential to inhibit deep winter mixing and limit primary production in the future.

Submesoscale eddies and fronts are important components of oceanic mixing and energy fluxes. These phenomena occur in the surface ocean for a period of several days, on scales between a few hundred meters and few tens of kilometers. Remote sensing and modeling suggest that eddies and fronts may influence marine ecosystem dynamics, but their limited temporal and spatial scales make them challenging for observation and in situ sampling. Here, the study of a submesoscale filament in summerly Arctic waters (depth 0–400 m) revealed enhanced mixing of Polar and Atlantic water masses, resulting in a ca. 4 km wide and ca. 50 km long filament with distinct physical and biogeochemical characteristics. Compared to the surrounding waters, the filament was characterized by a distinct phytoplankton bloom, associated with depleted inorganic nutrients, elevated chlorophyll a concentrations, as well as twofold higher phyto- and bacterioplankton cell abundances. High-throughput 16S rRNA gene sequencing of bacterioplankton communities revealed enrichment of typical phytoplankton bloom-associated taxonomic groups (e.g., Flavobacteriales) inside the filament. Furthermore, linked to the strong water subduction, the vertical export of organic matter to 400 m depth inside the filament was twofold higher compared to the surrounding waters. Altogether, our results show that physical submesoscale mixing can shape distinct biogeochemical conditions and microbial communities within a few kilometers of the ocean. Hence, the role of submesoscale features in polar waters for surface ocean biodiversity and biogeochemical processes need further investigation, especially with regard to the fate of sea ice in the warming Arctic Ocean.

Mitochondrial respiration is a multi-step pathway that involves matrix and membrane-associated enzymes and plays a key role in acclimation to variable environmental conditions, but until now it has not been clear which of these steps would be most important in acclimation to changing temperatures and CO2 levels. Considering scenarios of ocean warming and acidification we assessed the role and limitation to phenotypic plasticity in the hearts of two Gadoid species adapted to different thermal ranges: the polar cod (Boreogadus saida), an Arctic stenotherm, and the Northeast Arctic population of Atlantic cod (NEAC, Gadus morhua), a cold eurytherm. We analysed the capacity of single enzymes involved in mitochondrial respiration [citrate synthase (CS), succinate dehydrogenase (SDH), cytochrome c oxidase (CCO)], the capacity of the electron transport system and the lipid class composition of the cellular membranes. Juveniles of the two species were held for four months at four temperatures (0, 3, 6, 8 °C for polar cod and 3, 8, 12, 16 °C for NEAC), at both ambient and elevated $$P_{{{\text{CO}}_{2} }}$$ (400 µatm and 1170 µatm, respectively). Polar cod showed no changes in mitochondrial enzyme capacities and in the relative lipid class composition in response to altered temperature or elevated $$P_{{{\text{CO}}_{2} }} .$$ The lack of cardiac cellular plasticity together with evidence at the whole-animal level coming from other studies is indicative of little or no ability to overcome stenothermy, in particular during acclimation to 8 °C. In contrast, eurythermal NEAC exhibited modifications of membrane composition towards a more rigid structure and altered enzyme capacities to preserve functionality at higher temperatures. Furthermore, in NEAC, the capacities of SDH, CCO and CS were increased by high levels of CO2 if combined with high temperatures (12 and 16 °C), suggesting the compensation of an inhibitory effect. These results indicate that the cold eurythermal species (NEAC) is able to alter its mitochondrial function to a far greater extent than the Arctic stenotherm (polar cod), indicating greater resilience to variable environmental conditions. This difference in plasticity may underpin differences in the resilience to climate change and affect future species distributions and, eventually, survival.

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ13C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ13C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ13C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ13C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs.

International audience

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.

We investigated diets of 24 Barents Sea zooplankton taxa to understand pelagic food-web processes during late summer, including the importance of sea ice algae-produced carbon. This was achieved by combining insights derived from multiple and complementary trophic marker approaches to construct individual aspects of feeding. Specifically, we determined proportions of algal-produced fatty acids (FAs) to reflect the reliance on diatom- versus dinoflagellate-derived carbon, highly branched isoprenoid (HBI) lipids that distinguish between ice-associated and pelagic carbon sources, and sterols to indicate the degree of carnivory. Copepods had the strongest diatom signal based on FAs, while a lack of sea ice algae-associated HBIs (IP25, IPSO25) suggested that they fed on pelagic rather than ice-associated diatoms. The amphipodThemisto libellulaand the ctenophoresBeroë cucumisandMertensia ovumhad a higher contribution of dinoflagellate-produced FAs. There was a high degree of carnivory in this food web, as indicated by the FA carnivory index 18:1(n−9)/18:1(n−7) (mean value < 1 only in the pteropodClione limacina), the presence of copepod-associated FAs in most of the taxa, and the absence of algal-produced HBIs in small copepod taxa, such asOithona similisandPseudocalanusspp. The coherence between concentrations of HBIs and phytosterols within individuals suggested that phytosterols provide a good additional indication for algal ingestion. Sea ice algae-associated HBIs were detected in six zooplankton species (occurring in krill, amphipods, pteropods, and appendicularians), indicating an overall low to moderate contribution of ice-associated carbon from late-summer sea ice to pelagic consumption. The unexpected occurrence of ice-derived HBIs in pteropods and appendicularians, however, suggests an importance of sedimenting ice-derived material at least for filter feeders within the water column at this time of year.

Submesoscale eddies and fronts are recognized as important components of oceanic mixing and energy fluxes. These submesoscale phenomena occur in the surface ocean for a period of a few days on scales between several hundred meters and a few tens of kilometers. Remote sensing and modeling suggest that they may influence marine ecosystem dynamics, but their limited temporal and spatial scales make them challenging for observation and in situ sampling. Here, the study of a submesoscale filament in summerly Arctic waters (depth 0 - 400 m) revealed enhanced vertical mixing of Polar and Atlantic water masses, resulting in a ca. 4 km wide and ca. 50 km long filament with distinct physical and biogeochemical conditions. Compared to the surrounding waters the filament was characterized by a distinct phytoplankton bloom dominated by diatoms and two-fold higher bacterioplankton cell densities. High-throughput 16S rRNA gene sequencing of both bacterioplankton communities revealed 3-4 orders of magnitude higher sequence abundance of Synechococcus inside the filament, as well as tenfold higher sequence abundance of taxonomic groups typically found during summertime in aging phytoplankton blooms (e.g., Flavobacteriales). In contrast, the surrounding waters contained severalfold higher sequence abundance of winterly taxonomic groups that are also associated with polar water masses (e.g., SAR202 clade). Altogether, our results show that physical submesoscale processes in the ocean can shape distinct biogeochemical conditions and microbial communities within a few kilometers. Furthermore, our results underline the importance of such submesoscale features for our understanding of surface ocean diversity and biogeochemical processes.

Publisher: Elsevier B.V.; International audience; Meiofauna are known to have an important role on many ecological processes, although, their role in food web dynamics is often poorly understood, partially as they have been an overlooked and under sampled organism group. Here, we used quantitative food web modeling to evaluate the trophic relationship between meiofauna and their food sources and how meiofauna can mediate the carbon flow to higher trophic levels in five contrasting soft-bottom intertidal habitats (including seagrass beds, mudflats and sandflats). Carbon flow networks were constructed using the linear inverse model-Markov chain Monte Carlo technique, with increased resolution of the meiofauna compartments (i.e. biomass and feeding ecology of the different trophic groups of meiofauna) compared to most previous modeling studies. These models highlighted that the flows between the highly productive microphytobenthos and the meiofauna compartments play an important role in transferring carbon to the higher trophic levels, typically more efficiently so than macrofauna. The pathway from microphytobenthos to meiofauna represented the largest flow in all habitats and resulted in high production of meiofauna independent of habitat. All trophic groups of meiofauna, except for selective deposit feeders, had a very high dependency on microphytobenthos. Selective deposit feeders relied instead on a wider range of food sources, with varying contributions of bacteria, microphytobenthos and sediment organic matter. Ecological network analyses (e.g. cycling, throughput and ascendency) of the modeled systems highlighted the close positive relationship between the food web efficiency and the assimilation of high-quality food sources by primary consumers, e.g. meiofauna and macrofauna. Large proportions of these flows can be attributed to trophic groups of meiofauna. The sensitivity of the network properties to the representation of meiofauna in the models leads to recommending a greater attention in ecological data monitoring and integrating meiofauna into food web models. © 2020 Elsevier B.V.

The study revealed species- and stage-specific differences in lipid accumulation of the dominant Antarctic copepods, the primarily herbivorous Calanoides acutus (copepodite stage V (CV), females) and the more omnivorous Calanus propinquus (females) storing wax esters and triacylglycerols, respectively, which were collected in summer (end of December). Feeding carbon-labelled diatoms to these copepods, 13C elucidated assimilation and turnover rates of copepod total lipids as well as specific fatty acids and alcohols. The 13C incorporation was monitored by compound-specific stable isotope analysis (CSIA). CV stages of C. acutus exhibited an intense total lipid turnover and 55% of total lipids were labelled after 9 days of feeding. By contrast, total lipid assimilation of female C. acutus and C. propinquus was lower with 29% and 32%, respectively. The major dietary fatty acids 16:0, 16:1(n − 7) and 20:5(n − 3) had high turnover rates in all specimens. In C. acutus CV, the high rates of the de novo synthesized long-chain monounsaturated fatty acids and alcohols 20:1(n − 9) and 22:1(n − 11) indicate intense lipid deposition, whereas these rates were low in females. The differences in lipid assimilation and turnover clearly show that the copepod species exhibit a high variability and plasticity to adapt their lipid production to their various life phases. This article is part of the theme issue ‘The next horizons for lipids as ‘trophic biomarkers': evidence and significance of consumer modification of dietary fatty acids'.

Fatty acid (FA) compositions provide insights about storage and feeding modes of marine organisms, characterizing trophic relationships in the marine food web. Such compositional data, which are normalized to sum to 1, have values—and thus derived statistics as well—that depend on the particular mix of components that constitute the composition. In FA studies, if the set of FAs under investigation is different in two separate studies, all the summary statistics and relationships between the FAs that are common to the two studies are artificially changed due to the normalization, and thus incomparable. Ratios of FAs, however, are invariant to the particular choice of FAs under consideration—they are said to be subcompositionally coherent. Here, we document the collaboration between a biochemist (M.G.) and a statistician (M.J.G.) to determine a suitable small set of FA ratios that effectively replaces the original data set for the purposes of univariate and multivariate analysis. This strategy is applied to two FA data sets, on copepods and amphipods, respectively, and is widely applicable in other contexts. The selection of ratios is performed in such a way as to satisfy substantive requirements in the context of the respective data set, namely to explain phenomena of interest relevant to the particular species, as well as the statistical requirement to explain as much variance in the FA data set as possible. Benefits of this new approach are (1) univariate statistics that can be validly compared between different studies, and (2) a simplified multivariate analysis of the reduced set of ratios, giving practically the same results as the analysis of the full FA data set.

Assessing the relative importance of sea ice algal-based production is often vital for studies about climate change impacts on Arctic marine ecosystems. Several types of lipid biomarkers and stable isotope ratios are widely used for tracing sea ic-associated (sympagic) vs. pelagic particulate organic matter (POM) in marine food webs. However, there has been limited understanding about the plasticity of these compounds in space and time, which constrains the robustness of some of those approaches. Furthermore, some of the markers are compromised by not being unambiguously specific for sea ice algae, whereas others might only be produced by a small sub-group of species. We analyzed fatty acids, highly branched isoprenoids (HBIs), stable isotope ratios of particulate organic carbon (POC) (&delta, 13C), as well as &delta, 13C of selected fatty acid markers during an Arctic sea ice algal bloom, focusing on spatial and temporal variability. We found remarkable differences between these approaches and show that inferences about bloom characteristics might even be contradictory between markers. The impact of environmental factors as causes of this considerable variability is highlighted and explained. We emphasize that awareness and, in some cases, caution is required when using lipid and stable isotope markers as tracers in food web studies and offer recommendations for the proper application of these valuable approaches.

Antarctic krill Euphausia superba is an ecological key species in the Southern Ocean and a major fisheries resource. The winter survival of age class 0 (AC0) krill is susceptible to changes in the sea-ice environment due to their association with sea ice and their need to feed during their first winter. However, our understanding of their overwintering diet and its variability is limited. We studied the spatio-temporal variability of the diet in 4 cohorts of AC0 krill in the Northern Weddell Sea during late winter 2013 using stomach contents, fatty acids (FAs) and bulk stable isotope analysis (BSIA). Stomach contents were dominated by diatoms in numbers and occasionally contained large volumes of copepods. Many of the prey species found in the stomachs were sea ice-associated. Our results show that the diet of overwintering AC0 krill varies significantly in space and time. Variability in stomach content composition was related to environmental factors, including chlorophyll a concentration, copepod abundance and sea-ice cover. In contrast, FA composition mainly varied between cohorts, indicating variation in the long-term diet. The condition of the AC0 krill was reflected in FA and BSIA analysis, suggesting that the availability of sea ice-derived food sources over a long period may impact the condition of developing AC0 krill significantly. The spatio-temporal availability of sea-ice resources is a potentially important factor for AC0 krill winter survival.

The rhodophytes Mastocarpus stellatus and Chondrus crispus occupy the lower intertidal zone of rocky shores along North Atlantic coastlines, with C. crispus generally occurring slightly deeper. Consequently, M. stellatus is exposed to more variable environmental conditions, related to a generally higher stress tolerance of this species. In order to extend our understanding of seasonal modulation of stress tolerance, we subjected local populations of M. stellatus and C. crispus from Helgoland, North Sea, to short-term high-light stress experiments over the course of a year (October 2011, March, May and August 2012). Biochemical analyses (pigments, antioxidants, total lipids, fatty acid compositions) allowed to reveal mechanisms behind modulated high-light tolerances. Overall, C. crispus was particularly more susceptible to high-light at higher water temperatures (October 2011 and August 2012). Furthermore, species-specific differences in antioxidants, total lipid levels and the shorter-chain/longer-chain fatty acid ratio (C14 + C16/C18 + C20) were detected, which may enhance the tolerance to high-light and other abiotic stress factors in M. stellatus, so that this species is more competitive in the highly variable upper intertidal zone compared to C. crispus. Since the high-light tolerance in C. crispus seemed to be affected by water temperature, interactions between both species may be impacted in the future by rising mean annual sea surface temperature around the island of Helgoland.

The Arctic represents an extreme habitat for phototrophic algae due to long periods of darkness caused by the polar night (~4 months darkness). Benthic diatoms, which dominate microphytobenthic communities in shallow water regions, can survive this dark period, but the underlying physiological and biochemical mechanisms are not well understood. One of the potential mechanisms for long-term dark survival is the utilisation of stored energy products in combination with a reduced basic metabolism. In recent years, water temperatures in the Arctic increased due to an ongoing global warming. Higher temperatures could enhance the cellular energy requirements for the maintenance metabolism during darkness and, therefore, accelerate the consumption of lipid reserves. In this study, we investigated the macromolecular ratios and the lipid content and composition of Navicula cf. perminuta Grunow, an Arctic benthic diatom isolated from the microphytobenthos of Adventfjorden (Svalbard, Norway), over a dark period of 8 weeks at two different temperatures (0 and 7 °C). The results demonstrate that N. perminuta uses the stored lipid compound triacylglycerol (TAG) during prolonged dark periods, but also the pool of free fatty acids (FFA). Under the enhanced temperature of 7 °C, the lipid resources were used significantly faster than at 0 °C, which could consequently lead to a depletion of this energy reserves before the end of the polar night. On the other hand, the membrane building phospho- and glycolipids remained unchanged during the 8 weeks darkness, indicating still intact thylakoid membranes. These results explain the shorter survival times of polar diatoms with increasing water temperatures during prolonged dark periods.

This study aimed at a better understanding of the fatty acid (FA) turnover in Arctic pteropods. Thecosome pteropods, i.e. Limacina helicina (juveniles and adults) and L. retroversa (adults), were collected in summer/autumn in Kongsfjorden and Isfjorden (Svalbard, 78° N) and, for the first time, successfully fed with 13 C-labeled algae for 6 d. The gymnosome pteropod Clione limacina was sampled in summer in northern Svalbard and fed with 13 C-labeled L. retroversa for 23 d. FA compositions were determined by gas chromatography, and 13 C enrichment of FAs was analyzed by compound-specific isotope analysis. Among the thecosomes, maximum lipid turnover occurred in L. retroversa adults (1.3% d −1 ). L. helicina adults and juveniles showed lower lipid turnover rates (0.1 and 0.2% d −1 , respectively). The thecosomes exhibited the ability to assimilate omega-3 FAs (up to 8.0% d −1 ). The lipid turnover rate of C. limacina averaged at only 0.07% d −1 . However, C. limacina clearly showed the unusual capacity of de novo synthesis of odd-chain FAs (up to 1.2% d −1 ). Lipid turnover rates of pteropods were lower than those reported for Arctic copepods. However, pteropods may play a substantial role in the transfer of lipids to higher trophic levels, especially in autumn, when copepods have descended from the upper layers of the water column. The pteropods also showed the capacity to channel particular compounds such as omega-3 and odd-chain FAs, and therefore could be important for the functional diversity of the Arctic zooplankton community.

International audience; Meiofauna can play an important role in the carbon fluxes of soft-bottom coastal habitats. Investigation of their feeding behavior and trophic position remains challenging due to their small size. In this study, we determine and compare the food sources used by nematodes and benthic copepods by using stable isotope compositions, fatty acid profiles and compound specific isotope analyses of fatty acids in the mudflats, seagrass beds and a sandflat of the Marennes-Oléron Bay, France, and the Sylt-Rømø Bight, Germany. Suspended particulate organic matter was much more 13C-depleted than other food sources and meiofauna, highlighting its poor role in the different studied habitats. The very low proportions of vascular plant fatty acid markers in meiofauna demonstrated that these consumers did not rely on this food source, either fresh or detrital, even in seagrass beds. The combined use of stable isotopes and fatty acids emphasized microphytobenthos and benthic bacteria as the major food sources of nematodes and benthic copepods. Compound specific analyses of a bacteria marker confirmed that bacteria mostly used microphytobenthos as a substrate.

Calanus is one of the best studied genera of Arctic zooplankton, but still we know very little about the males since they are short-lived and mainly present in winter. Their short life-span compared to females is assumed to be a combination of high mating activity, no feeding and consequential depletion of lipid stores. In this study we tested 1) if the life span of male Calanus glacialis is limited by their lipid storage reserves and 2) if males are capable of feeding and utilize food if present. We ran two separate experiments from January to March; one on starvation and one on feeding. In the 39-days long starvation experiment we followed the lipid sac size of individually incubated males until their time of death. On average the total lipid (TL) content decreased by 2.6 to 4.5 μg day−1, but despite this males had substantial amounts of lipids left (131.4 μg, SD 44.0) when they died. This strongly suggests that the depletion of lipid reserves is not the main reason for males' short life span which in this study was measured to be up to 73 days. In the feeding experiment, we fed both C. glacialis males and females ad libitum with 13C labelled microalgae. Both males and females were capable of feeding and assimilate the diatom monoculture, but females responded faster to the sudden favourable food conditions, and produced more and larger fecal pellets than the males. Assimilation of 13C labelled 20:5(n-3), an essential polyunsaturated fatty acid (PUFA), from the diatom diet was traceable in both males and females on day 21, and then with a higher enrichment in females than males. Morphological investigations of the feeding appendages showed some differences between sexes, suggesting males to be more omnivorous than females. In conclusion, lipid storage depletion is not the cause of death for male C. glacialis, and males may even compensate for some of the mating energy costs by feeding. In future, we recommend further studies on the role of essential fatty acids (FA) for sperm formation and aging as determining factors for males' relatively short life span.

Organisms in shallow waters at high latitudes are under pressure due to climate change. These areas are typically inhabited by microphytobenthos (MPB) communities, composed mainly of diatoms. Only sparse information is available on the ecophysiology and acclimation processes within MPBs from Arctic regions. The physico-chemical environment and the ecology and ecophysiology of benthic diatoms in Kongsfjorden (Svalbard, Norway) are addressed in this review. MPB biofilms cover extensive areas of sediment. They show high rates of primary production, stabilise sediment surfaces against erosion under hydrodynamic forces, and affect the exchange of oxygen and nutrients across the sediment-water interface. Additionally, this phototrophic community represents a key component in the functioning of the Kongsfjorden trophic web, particularly as a major food source for benthic suspension- or deposit-feeders.

Life cycle and reproduction of Calanus hyperboreus were studied during a year of record low ice cover in the southeastern Beaufort Sea. Stages CIV, adult females and CV dominated the overwintering population, suggesting a 2- to 3-year life cycle. Within two spring-summer months in the upper water column females filled their energy reserves before initiating their downward seasonal migration. From February to March, vigorous reproduction (20–65 eggs f−1 d−1) delivered numerous eggs (29 000 eggs m−2) at depth and nauplii N1-N3 (17 000 ind. m−2) in the water column. However, CI copepodite recruitment in May, coincident with the phytoplankton bloom, was modest in Amundsen Gulf compared to sites outside the gulf. Consequently, C. hyperboreus abundance and biomass stagnated throughout summer in Amundsen Gulf. As a mismatch between the first-feeding stages and food was unlikely under the favourable feeding conditions of April-May 2008, predation on the egg and larval stages in late winter presumably limited subsequent recruitment and population growth. Particularly abundant in Amundsen Gulf, the copepods Metridia longa and C. glacialis were likely important consumers of C. hyperboreus eggs and nauplii. With the ongoing climate-driven lengthening of the ice-free season, intensification of top-down control of C. hyperboreus recruitment by thriving populations of mesopelagic omnivores and carnivores like M. longa may counteract the potential benefits of increased primary production over the Arctic shelves margins for this key prey of pelagic fish, seabirds and the bowhead whale.

To better predict ecological consequences of changing Arctic sea ice environments, we aimed to quantify the contribution of ice algae-produced carbon (αIce) to pelagic food webs in the central Arctic Ocean. Eight abundant under-ice fauna species were submitted to fatty acid (FA) analysis, bulk stable isotope analysis (BSIA) of nitrogen (δ15N) and carbon (δ13C) isotopic ratios, and compound-specific stable isotope analysis (CSIA) of δ13C in trophic marker FAs. A high mean contribution αIce was found in Apherusa glacialis and other sympagic (ice-associated) amphipods (BSIA: 87% to 91%, CSIA: 58% to 92%). The pelagic copepods Calanus glacialis and C. hyperboreus, and the pelagic amphipod Themisto libellula showed substantial, but varying αIce values (BSIA: 39% to 55%, CSIA: 23% to 48%). Lowest αIce mean values were found in the pteropod Clione limacina (BSIA: 30%, CSIA: 14% to 18%). Intra-specific differences in FA compositions related to two different environmental regimes were more pronounced in pelagic than in sympagic species. A comparison of mixing models using different isotopic approaches indicated that a model using δ13C signatures from both diatom-specific and dinoflagellate-specific marker FAs provided the most conservative estimate of αIce. Our results imply that ecological key species of the central Arctic Ocean thrive significantly on carbon synthesized by ice algae. Due to the close connectivity between sea ice and the pelagic food web, changes in sea ice coverage and ice algal production will likely have important consequences for food web functioning and carbon dynamics of the pelagic system.

The study revealed species- and stage-specific differences in lipid accumulation of the dominant Antarctic copepods, the primarily herbivorous Calanoides acutus (copepodite stage V (CV), females) and the more omnivorous Calanus propinquus (females) storing wax esters and triacylglycerols, respectively, which were collected in summer (end of December). Feeding carbon-labelled diatoms to these copepods, 13 C elucidated assimilation and turnover rates of copepod total lipids as well as specific fatty acids and alcohols. The 13 C incorporation was monitored by compound-specific stable isotope analysis (CSIA). CV stages of C. acutus exhibited an intense total lipid turnover and 55% of total lipids were labelled after 9 days of feeding. By contrast, total lipid assimilation of female C. acutus and C. propinquus was lower with 29% and 32%, respectively. The major dietary fatty acids 16:0, 16:1(n − 7) and 20:5(n − 3) had high turnover rates in all specimens. In C. acutus CV, the high rates of the de novo synthesized long-chain monounsaturated fatty acids and alcohols 20:1(n − 9) and 22:1(n − 11) indicate intense lipid deposition, whereas these rates were low in females. The differences in lipid assimilation and turnover clearly show that the copepod species exhibit a high variability and plasticity to adapt their lipid production to their various life phases. This article is part of the theme issue ‘The next horizons for lipids as ‘trophic biomarkers': evidence and significance of consumer modification of dietary fatty acids'.

A combined approach merging stable isotopes and fatty acids was applied to study anthropogenic pollution in the Río Negro estuary. Fatty acid markers of vegetal detritus indicated considerable allochthonous inputs at freshwater sites. Correlative evidence of diatom fatty acids, δ13C, chlorophyll and particulate organic matter suggested the importance of diatoms for the autochthonous organic matter production at the river mouth. Low δ15N values (~0‰) and high fatty acid 18:1(n-7) concentrations in the suspended particulate matter, in combination with the peaks of coliforms and ammonium, indicated a strong impact of untreated sewage discharge. The 15N depletion was related to oxygen-limited ammonification processes and incorporation of 15N depleted ammonium to microorganisms. This work demonstrates that the combined use of lipid and isotopic markers can greatly increase our understanding of biogeochemical factors and pollutants influencing estuaries, and our findings highlight the urgent need for water management actions to reduce eutrophication. Fil: Kopprio, Germán Adolfo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Bahía Blanca. Instituto Argentino de Oceanografía. Universidad Nacional del Sur. Instituto Argentino de Oceanografía; Argentina. Leibniz Centre for Tropical Marine Research; Alemania Fil: Dutto, María Sofía. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Bahía Blanca. Instituto Argentino de Oceanografía. Universidad Nacional del Sur. Instituto Argentino de Oceanografía; Argentina Fil: Garzon Cardona, John Edison. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Bahía Blanca. Instituto Argentino de Oceanografía. Universidad Nacional del Sur. Instituto Argentino de Oceanografía; Argentina Fil: Gärdes, A.. Leibniz Centre for Tropical Marine Research; Alemania Fil: Lara, Ruben Jose. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Bahía Blanca. Instituto Argentino de Oceanografía. Universidad Nacional del Sur. Instituto Argentino de Oceanografía; Argentina Fil: Graeve, Martin. Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research; Alemania

Maternal investment (MI), the energy allocated by mothers to offspring, has important effects on the life-history traits of marine organisms. Variation in such traits shows strong correlation with latitude for several marine taxa (Thorson’s rule). Large-scale latitudinal variation in MI within a single species suggests population genetic divergence, while temporal changes in MI, rather, reflect plasticity. At higher latitudes (i.e., colder waters), traits associated with MI (brood weight, fecundity, egg volume, and energy content) increase. To identify phenotypic plasticity along a latitudinal gradient in MI traits (brood weight, egg volume, density number, and egg lipid composition), five populations of the kelp crab Taliepus dentatus along the coast of Chile (30°S–42°S) were investigated during the summer (December–February) and winter months (June–August) of 2015–2016. Despite this wide latitudinal range, the sea surface temperature (SST) difference between the northernmost and the southernmost sites was only approximately 2.0 °C in winter and 5.5 °C in summer. In summer, when latitudinal variation in SST was highest, brood weight, egg density, fecundity, and egg lipids increased with latitude, while egg volume decreased. No trends in MI were observed in winter when the SST gradient was almost non-existent. These results suggest that the relationship between MI and latitude is shaped by temperature rather than being site-specific. The seasonality of latitudinal MI traits also suggests a trade-off between the costs of female maintenance and/or brooding behaviours and MI. When investigating latitudinal and temporal variation in marine brooder MI, the effect of temperature on life-history traits and the associated costs of female brooding should be quantified.

How the abundant pelagic life of the Southern Ocean survives winter darkness, when the sea is covered by pack ice and phytoplankton production is nearly zero, is poorly understood. Ice-associated ("sympagic") microalgae could serve as a high-quality carbon source during winter, but their significance in the food web is so far unquantified. To better understand the importance of ice algae-produced carbon for the overwintering of Antarctic organisms, we investigated fatty acid (FA) and stable isotope compositions of 10 zooplankton species, and their potential sympagic and pelagic carbon sources. FA-specific carbon stable isotope compositions were used in stable isotope mixing models to quantify the contribution of ice algae-produced carbon (αIce ) to the body carbon of each species. Mean αIce estimates ranged from 4% to 67%, with large variations between species and depending on the FA used for the modelling. Integrating the αIce estimates from all models, the sympagic amphipod Eusirus laticarpus was the most dependent on ice algal carbon (αIce : 54%-67%), and the salp Salpa thompsoni showed the least dependency on ice algal carbon (αIce : 8%-40%). Differences in αIce estimates between FAs associated with short-term vs. long-term lipid pools suggested an increasing importance of ice algal carbon for many species as the winter season progressed. In the abundant winter-active copepod Calanus propinquus, mean αIce reached more than 50% in late winter. The trophic carbon flux from ice algae into this copepod was between 3 and 5 mg C m-2 day-1 . This indicates that copepods and other ice-dependent zooplankton species transfer significant amounts of carbon from ice algae into the pelagic system, where it fuels the food web, the biological carbon pump and elemental cycling. Understanding the role of ice algae-produced carbon in these processes will be the key to predictions of the impact of future sea ice decline on Antarctic ecosystem functioning.

Fil: Kopprio, German Adolfo. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Centro Cientifico Tecnologico Conicet - Bahia Blanca. Instituto Argentino de Oceanografia. Universidad Nacional del Sur. Instituto Argentino de Oceanografia; Argentina

Lessonia nigrescens used to be an abundant kelp species along the Chilean coast, but recent molecular studies revealed the existence of a L. nigrescens species complex, which includes the two cryptic species Lessonia berteroana and Lessonia spicata. Since these species have different distributions (16°S–30°S for L. berteroana and 29°S–42°S for L. spicata), they experience differences in environmental conditions, such as solar irradiance, seawater temperature and air exposure during low tide. This study tested to what extent the genetic distinctness of each of the two species [identified by a mitochondrial marker (atp8/trnS)] is reflected by ecophysiological traits (total lipids, fatty acid composition, phlorotannins, pigments and variable chlorophyll a fluorescence of PSII) in response to the respective environmental conditions, prevailing along the latitudinal gradient. We studied algal individuals from eight populations (27°S–32°S, including the species overlapping zone). Phlorotannins, pigments and Chl a fluorescence of PSII were most crucial for species-specific adaptations at the respective growth sites, whereas changes in total lipids and fatty acid compositions were negligible. Hence, species differentiation within the L. nigrescens complex is also manifested at the ecophysiological level. These findings may help to predict kelp responses towards future environmental changes.

The polar cod ( Boreogadus saida ) is considered an ecological key species, because it reaches high stock biomasses and constitutes an important carbon source for seabirds and marine mammals in high-Arctic ecosystems. Young polar cod (1–2 years) are often associated with the underside of sea ice. To evaluate the impact of changing Arctic sea ice habitats on polar cod, we examined the diet composition and quantified the contribution of ice algae-produced carbon ( α Ice ) to the carbon budget of polar cod. Young polar cod were sampled in the ice-water interface layer in the central Arctic Ocean during late summer 2012. Diets and carbon sources of these fish were examined using 4 approaches: (1) stomach content analysis, (2) fatty acid (FA) analysis, (3) bulk nitrogen and carbon stable isotope analysis (BSIA) and (4) compound-specific stable isotope analysis (CSIA) of FAs. The ice-associated (sympagic) amphipod Apherusa glacialis dominated the stomach contents by mass, indicating a high importance of sympagic fauna in young polar cod diets. The biomass of food measured in stomachs implied constant feeding at daily rates of ∼1.2% body mass per fish, indicating the potential for positive growth. FA profiles of polar cod indicated that diatoms were the primary carbon source, indirectly obtained via amphipods and copepods. The α Ice using bulk isotope data from muscle was estimated to be >90%. In comparison, α Ice based on CSIA ranged from 34 to 65%, with the highest estimates from muscle and the lowest from liver tissue. Overall, our results indicate a strong dependency of polar cod on ice-algae produced carbon. This suggests that young polar cod may be particularly vulnerable to changes in the distribution and structure of sea ice habitats. Due to the ecological key role of polar cod, changes at the base of the sea ice-associated food web are likely to affect the higher trophic levels of high-Arctic ecosystems.

Antarctic krill Euphausia superba (‘krill’) constitute a fundamental food source for Antarctic seabirds and mammals, and a globally important fisheries resource. The future resilience of krill to climate change depends critically on the winter survival of young krill. To survive periods of extremely low production by pelagic algae during winter, krill are assumed to rely partly on carbon produced by ice algae. The true dependency on ice algae-produced carbon, however, is so far unquantified. This confounds predictions on the future resilience of krill stocks to sea ice decline. Fatty acid (FA) analysis, bulk stable isotope analysis (BSIA) and compound-specific stable isotope analysis (CSIA) of diatom- and dinoflagellate-associated marker FAs were applied to quantify the dependency of overwintering larval, juvenile and adult krill on ice algae-produced carbon (αIce) during winter 2013 in the Weddell-Scotia Confluence Zone. Our results demonstrate that the majority of the carbon uptake of the overwintering larval and juvenile krill originated from ice algae (up to 88% of the carbon budget), and that the dependency on ice algal carbon decreased with ontogeny, reaching less than 56% of the carbon budget in adults. Spatio-temporal variability in the utilization of ice algal carbon was more pronounced in larvae and juvenile krill than in adults. Differences between αIce estimates derived from short- versus long-term FA-specific isotopic compositions suggested that ice algae-produced carbon gained importance as the winter progressed, and might become critical at the late winter-spring transition, before the phytoplankton bloom commences. Where the sea ice season shortens, reduced availability of ice algae might possibly not be compensated by surplus phytoplankton production during wintertime. Hence, sea ice decline could seriously endanger the winter survival of recruits, and subsequently overall biomass of krill.

Our study deals with the lipid biochemistry of the krill community in the ecosystem of the high Arctic Kongsfjord (Svalbard). During the last decades, Kongsfjord experienced a change in krill species composition due to recent increased advection of Atlantic water masses carrying characteristic boreal as well as subtropical-boreal euphausiids into the ecosystem. The lipid biochemistry and trophic relationships of the species recently inhabiting the Arctic water masses are scarcely known, although a change in a krill population may have a significant impact on the ecosystem. A comparison of nutrition and energy storage strategies, stable isotopes, lipid profiles and fatty acid compositions showed remarkable differences between the krill species. These reflected the diverse feeding behaviours and specific adaptations to the environments of their origin: the boreal Meganyctiphanes norvegica and subtropical Nematoscelis megalops appear more carnivorous and have significantly lower mean lipid contents (29 and 10 %, respectively) and a different energy storage pattern (triacylglycerols and polar lipids, respectively) than the arcto-boreal Thysanoessa inermis, which consists of up to 54 % of lipids mainly stored as wax esters (>40 %). These differences may have significant implications for the rapidly changing marine food web of Kongsfjord—especially for higher trophic levels relying on the nutritional input of animal lipids.