Essential omega-3 fatty acids are depleted in sea ice and pelagic algae of the Central Arctic Ocean.

Microalgae are the main source of the omega-3 fatty acids eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), essential for the healthy development of most marine and terrestrial fauna including humans. Inverse correlations of algal EPA and DHA proportions (% of total fatty acids) with temperature have led to suggestions of a warming-induced decline in the global production of these biomolecules and an enhanced importance of high latitude organisms for their provision. The cold Arctic Ocean is a potential hotspot of EPA and DHA production, but consequences of global warming are unknown. Here, we combine a full-seasonal EPA and DHA dataset from the Central Arctic Ocean (CAO), with results from 13 previous field studies and 32 cultured algal strains to examine five potential climate change effects; ice algae loss, community shifts, increase in light, nutrients, and temperature. The algal EPA and DHA proportions were lower in the ice-covered CAO than in warmer peripheral shelf seas, which indicates that the paradigm of an inverse correlation of EPA and DHA proportions with temperature may not hold in the Arctic. We found no systematic differences in the summed EPA and DHA proportions of sea ice versus pelagic algae, and in diatoms versus non-diatoms. Overall, the algal EPA and DHA proportions varied up to four-fold seasonally and 10-fold regionally, pointing to strong light and nutrient limitations in the CAO. Where these limitations ease in a warming Arctic, EPA and DHA proportions are likely to increase alongside increasing primary production, with nutritional benefits for a non-ice-associated food web.

Myctobase, a circumpolar database of mesopelagic fishes for new insights into deep pelagic prey fields.

The global importance of mesopelagic fish is increasingly recognised, but they remain poorly studied. This is particularly true in the Southern Ocean, where mesopelagic fishes are both key predators and prey, but where the remote environment makes sampling challenging. Despite this, multiple national Antarctic research programs have undertaken regional sampling of mesopelagic fish over several decades. However, data are dispersed, and sampling methodologies often differ precluding comparisons and limiting synthetic analyses. We identified potential data holders by compiling a metadata catalogue of existing survey data for Southern Ocean mesopelagic fishes. Data holders contributed 17,491 occurrence and 11,190 abundance records from 4780 net hauls from 72 different research cruises. Data span across 37 years from 1991 to 2019 and include trait-based information (length, weight, maturity). The final dataset underwent quality control processes and detailed metadata was provided for each sampling event. This dataset can be accessed through Zenodo. Myctobase will enhance research capacity by providing the broadscale baseline data necessary for observing and modelling mesopelagic fishes.

Allometric relationships of ecologically important Antarctic and Arctic zooplankton and fish species.

Allometric relationships between body properties of animals are useful for a wide variety of purposes, such as estimation of biomass, growth, population structure, bioenergetic modelling and carbon flux studies. This study summarizes allometric relationships of zooplankton and nekton species that play major roles in polar marine food webs. Measurements were performed on 639 individuals of 15 species sampled during three expeditions in the Southern Ocean (winter and summer) and 2374 individuals of 14 species sampled during three expeditions in the Arctic Ocean (spring and summer). The information provided by this study fills current knowledge gaps on relationships between length and wet/dry mass of understudied animals, such as various gelatinous zooplankton, and of animals from understudied seasons and maturity stages, for example, for the krill Thysanoessa macrura and larval Euphausia superba caught in winter. Comparisons show that there is intra-specific variation in length-mass relationships of several species depending on season, e.g. for the amphipod Themisto libellula. To investigate the potential use of generalized regression models, comparisons between sexes, maturity stages or age classes were performed and are discussed, such as for the several krill species and T. libellula. Regression model comparisons on age classes of the fish E. antarctica were inconclusive about their general use. Other allometric measurements performed on carapaces, eyes, heads, telsons, tails and otoliths provided models that proved to be useful for estimating length or mass in, e.g. diet studies. In some cases, the suitability of these models may depend on species or developmental stages. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s00300-021-02984-4.

Unexpected fish and squid in the central Arctic deep scattering layer.

The retreating ice cover of the Central Arctic Ocean (CAO) fuels speculations on future fisheries. However, very little is known about the existence of harvestable fish stocks in this 3.3 million-square kilometer ecosystem around the North Pole. Crossing the Eurasian Basin, we documented an uninterrupted 3170-kilometer-long deep scattering layer (DSL) with zooplankton and small fish in the Atlantic water layer at 100- to 500-meter depth. Diel vertical migration of this central Arctic DSL was lacking most of the year when daily light variation was absent. Unexpectedly, the DSL also contained low abundances of Atlantic cod, along with lanternfish, armhook squid, and Arctic endemic ice cod. The Atlantic cod originated from Norwegian spawning grounds and had lived in Arctic water temperature for up to 6 years. The potential fish abundance was far below commercially sustainable levels and is expected to remain so because of the low productivity of the CAO.

High gene flow in polar cod (Boreogadus saida) from West-Svalbard and the Eurasian Basin.

The current and projected environmental change of the Arctic Ocean contrasts sharply with the limited knowledge of its genetic biodiversity. Polar cod Boreogadus saida (Lepechin, 1774) is an abundant circumpolar marine fish and ecological key species. The central role of polar cod in the Arctic marine food web warrants a better understanding of its population structure and connectivity. In this study, the genetic population structure of 171 juveniles, collected from several fjords off West-Svalbard (Billefjorden, Hornsund and Kongsfjorden), the northern Sophia Basin and the Eurasian Basin of the Arctic Ocean, was analysed using nine DNA microsatellite loci. Genetic analyses indicated moderate to high genetic diversity, but absence of spatial population structure and isolation-by-distance, suggesting ongoing gene flow between the studied sampling regions. High levels of connectivity may be key for polar cod to maintain populations across wide spatial scales. The adaptive capacity of the species will be increasingly important to face challenges such as habitat fragmentation, ocean warming and changes in prey composition. In view of a limited understanding of the population dynamics and evolution of polar cod, a valuable next step to predict future developments should be an integrated biological evaluation, including population genomics, a life-history approach, and habitat and biophysical dispersal modelling.

Review: the energetic value of zooplankton and nekton species of the Southern Ocean.

Understanding the energy flux through food webs is important for estimating the capacity of marine ecosystems to support stocks of living resources. The energy density of species involved in trophic energy transfer has been measured in a large number of small studies, scattered over a 40-year publication record. Here, we reviewed energy density records of Southern Ocean zooplankton, nekton and several benthic taxa, including previously unpublished data. Comparing measured taxa, energy densities were highest in myctophid fishes (ranging from 17.1 to 39.3 kJ g-1 DW), intermediate in crustaceans (7.1 to 25.3 kJ g-1 DW), squid (16.2 to 24.0 kJ g-1 DW) and other fish families (14.8 to 29.9 kJ g-1 DW), and lowest in jelly fish (10.8 to 18.0 kJ g-1 DW), polychaetes (9.2 to 14.2 kJ g-1 DW) and chaetognaths (5.0-11.7 kJ g-1 DW). Data reveals differences in energy density within and between species related to size, age and other life cycle parameters. Important taxa in Antarctic food webs, such as copepods, squid and small euphausiids, remain under-sampled. The variability in energy density of Electrona antarctica was likely regional rather than seasonal, although for many species with limited data it remains difficult to disentangle regional and seasonal variability. Models are provided to estimate energy density more quickly using a species' physical parameters. It will become increasingly important to close knowledge gaps to improve the ability of bioenergetic and food web models to predict changes in the capacity of Antarctic ecosystems to support marine life.

Dependency of Antarctic zooplankton species on ice algae-produced carbon suggests a sea ice-driven pelagic ecosystem during winter.

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.

Plastic ingestion by juvenile polar cod (Boreogadus saida) in the Arctic Ocean.

One of the recently recognised stressors in Arctic ecosystems concerns plastic litter. In this study, juvenile polar cod (Boreogadus saida) were investigated for the presence of plastics in their stomachs. Polar cod is considered a key species in the Arctic ecosystem. The fish were collected both directly from underneath the sea ice in the Eurasian Basin and in open waters around Svalbard. We analysed the stomachs of 72 individuals under a stereo microscope. Two stomachs contained non-fibrous microplastic particles. According to µFTIR analysis, the particles consisted of epoxy resin and a mix of Kaolin with polymethylmethacrylate (PMMA). Fibrous objects were excluded from this analysis to avoid bias due to contamination with airborne micro-fibres. A systematic investigation of the risk for secondary micro-fibre contamination during analytical procedures showed that precautionary measures in all procedural steps are critical. Based on the two non-fibrous objects found in polar cod stomachs, our results show that ingestion of microplastic particles by this ecologically important fish species is possible. With increasing human activity, plastic ingestion may act as an increasing stressor on polar cod in combination with ocean warming and sea-ice decline in peripheral regions of the Arctic Ocean. To fully assess the significance of this stressor and its spatial and temporal variability, future studies must apply a rigorous approach to avoid secondary pollution.

Pan-Arctic sea ice-algal chl a biomass and suitable habitat are largely underestimated for multiyear ice.

There is mounting evidence that multiyear ice (MYI) is a unique component of the Arctic Ocean and may play a more important ecological role than previously assumed. This study improves our understanding of the potential of MYI as a suitable habitat for sea ice algae on a pan-Arctic scale. We sampled sea ice cores from MYI and first-year sea ice (FYI) within the Lincoln Sea during four consecutive spring seasons. This included four MYI hummocks with a mean chl a biomass of 2.0 mg/m2 , a value significantly higher than FYI and MYI refrozen ponds. Our results support the hypothesis that MYI hummocks can host substantial ice-algal biomass and represent a reliable ice-algal habitat due to the (quasi-) permanent low-snow surface of these features. We identified an ice-algal habitat threshold value for calculated light transmittance of 0.014%. Ice classes and coverage of suitable ice-algal habitat were determined from snow and ice surveys. These ice classes and associated coverage of suitable habitat were applied to pan-Arctic CryoSat-2 snow and ice thickness data products. This habitat classification accounted for the variability of the snow and ice properties and showed an areal coverage of suitable ice-algal habitat within the MYI-covered region of 0.54 million km2 (8.5% of total ice area). This is 27 times greater than the areal coverage of 0.02 million km2 (0.3% of total ice area) determined using the conventional block-model classification, which assigns single-parameter values to each grid cell and does not account for subgrid cell variability. This emphasizes the importance of accounting for variable snow and ice conditions in all sea ice studies. Furthermore, our results indicate the loss of MYI will also mean the loss of reliable ice-algal habitat during spring when food is sparse and many organisms depend on ice-algae.

Comparing springtime ice-algal chlorophyll a and physical properties of multi-year and first-year sea ice from the Lincoln Sea.

With near-complete replacement of Arctic multi-year ice (MYI) by first-year ice (FYI) predicted to occur within this century, it remains uncertain how the loss of MYI will impact the abundance and distribution of sea ice associated algae. In this study we compare the chlorophyll a (chl a) concentrations and physical properties of MYI and FYI from the Lincoln Sea during 3 spring seasons (2010-2012). Cores were analysed for texture, salinity, and chl a. We identified annual growth layers for 7 of 11 MYI cores and found no significant differences in chl a concentration between the bottom first-year-ice portions of MYI, upper old-ice portions of MYI, and FYI cores. Overall, the maximum chl a concentrations were observed at the bottom of young FYI. However, there were no significant differences in chl a concentrations between MYI and FYI. This suggests little or no change in algal biomass with a shift from MYI to FYI and that the spatial extent and regional variability of refrozen leads and younger FYI will likely be key factors governing future changes in Arctic sea ice algal biomass. Bottom-integrated chl a concentrations showed negative logistic relationships with snow depth and bulk (snow plus ice) integrated extinction coefficients; indicating a strong influence of snow cover in controlling bottom ice algal biomass. The maximum bottom MYI chl a concentration was observed in a hummock, representing the thickest ice with lowest snow depth of this study. Hence, in this and other studies MYI chl a biomass may be under-estimated due to an under-representation of thick MYI (e.g., hummocks), which typically have a relatively thin snowpack allowing for increased light transmission. Therefore, we suggest the on-going loss of MYI in the Arctic Ocean may have a larger impact on ice-associated production than generally assumed.

The association of Antarctic krill Euphausia superba with the under-ice habitat.

The association of Antarctic krill Euphausia superba with the under-ice habitat was investigated in the Lazarev Sea (Southern Ocean) during austral summer, autumn and winter. Data were obtained using novel Surface and Under Ice Trawls (SUIT), which sampled the 0-2 m surface layer both under sea ice and in open water. Average surface layer densities ranged between 0.8 individuals m(-2) in summer and autumn, and 2.7 individuals m(-2) in winter. In summer, under-ice densities of Antarctic krill were significantly higher than in open waters. In autumn, the opposite pattern was observed. Under winter sea ice, densities were often low, but repeatedly far exceeded summer and autumn maxima. Statistical models showed that during summer high densities of Antarctic krill in the 0-2 m layer were associated with high ice coverage and shallow mixed layer depths, among other factors. In autumn and winter, density was related to hydrographical parameters. Average under-ice densities from the 0-2 m layer were higher than corresponding values from the 0-200 m layer collected with Rectangular Midwater Trawls (RMT) in summer. In winter, under-ice densities far surpassed maximum 0-200 m densities on several occasions. This indicates that the importance of the ice-water interface layer may be under-estimated by the pelagic nets and sonars commonly used to estimate the population size of Antarctic krill for management purposes, due to their limited ability to sample this habitat. Our results provide evidence for an almost year-round association of Antarctic krill with the under-ice habitat, hundreds of kilometres into the ice-covered area of the Lazarev Sea. Local concentrations of postlarval Antarctic krill under winter sea ice suggest that sea ice biota are important for their winter survival. These findings emphasise the susceptibility of an ecological key species to changing sea ice habitats, suggesting potential ramifications on Antarctic ecosystems induced by climate change.