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UNLABELLED: 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. ispartof: POLAR BIOLOGY vol:45 issue:2 pages:203-224 ispartof: location:Germany status: published

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.

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.

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.

Ice algae are critical components to the lipid-driven Arctic marine food web, particularly early in the spring. As little is known about these communities in multiyear ice (MYI), we aimed to provide a baseline of fatty acid (FA) and stable isotope signatures of sea-ice communities in MYI from the Lincoln Sea and compare these biomarkers to first-year ice (FYI). Significant differences in the relative proportions of approximately 25% of the identified FAs and significantly higher nitrogen stable isotope values (δ15N) in bottom-ice samples of FYI (δ15N = 6.4 ± 0.7%) compared to MYI (δ15N = 5.0 ± 0.4%) reflect different community compositions in the two ice types. Yet, the relative proportion of diatom- and dinoflagellate-associated FAs, as well as their bulk and most of the FA-specific carbon stable isotope compositions (δ13C) were not significantly different between bottom FYI (bulk δ13C: –28.4% to –26.7%, FA average δ13C: –34.4% to –31.7%) and MYI (bulk δ13C: –27.6% to –27.2%, FA average δ13C: –33.6% to –31.9%), suggesting at least partly overlapping community structures and similar biochemical processes within the ice. Diatom-associated FAs contributed, on average, 28% and 25% to the total FA content of bottom FYI and MYI, respectively, indicating that diatoms play a central role in structuring sea-ice communities in the Lincoln Sea. The differences in FA signatures of FYI and MYI support the view that different ice types harbor different inhabitants and that the loss of Arctic MYI will impact complex food web interactions with ice-associated ecosystems. Comparable nutritional quality of FAs, however, as indicated by similar average levels of polyunsaturated FAs in bottom FYI (33%) and MYI (28%), could help to ensure growth and reproduction of ice-associated grazers despite the shift from a MYI to FYI-dominated sea-ice cover with ongoing climate warming.

In the Arctic Ocean, sea-ice decline will significantly change the structure of biological communities. At the same time, changing nutrient dynamics can have similarly strong and potentially interacting effects. To investigate the response of the taxonomic and trophic structure of planktonic and ice-associated communities to varying sea-ice properties and nutrient concentrations, we analysed four different communities sampled in the Eurasian Basin in summer 2012: (1) protists and (2) metazoans from the under-ice habitat, and (3) protists and (4) metazoans from the epipelagic habitat. The taxonomic composition of protist communities was characterised with 18S meta-barcoding. The taxonomic composition of metazoan communities was determined based on morphology. The analysis of environmental parameters identified (i) a ‘shelf-influenced’ regime with melting sea ice, high-silicate concentrations and low NOx (nitrate + nitrite) concentrations; (ii) a ‘Polar’ regime with low silicate concentrations and low NOx concentrations; and (iii) an ‘Atlantic’ regime with low silicate concentrations and high NOx concentrations. Multivariate analyses of combined bio-environmental datasets showed that taxonomic community structure primarily responded to the variability of sea-ice properties and hydrography across all four communities. Trophic community structure, however, responded significantly to NOx concentrations. In three of the four communities, the most heterotrophic trophic group significantly dominated in the NOx-poor shelf-influenced and Polar regimes compared to the NOx-rich Atlantic regime. The more heterotrophic, NOx-poor regimes were associated with lower productivity and carbon export than the NOx-rich Atlantic regime. For modelling future Arctic ecosystems, it is important to consider that taxonomic diversity can respond to different drivers than trophic diversity.

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.

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.

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.

Atlantic mackerel (Scomber scombrus L.) is a widely distributed commercially important pelagic species. Little is known about the stock structure of this species, but it is thought to be undergoing a range extension due to environmental changes. Knowledge of the stock structure under these changing conditions is fundamental for effective management. In this paper, 30 highly polymorphic microsatellite loci developed with next generation sequencing are described. The number of alleles per locus ranged from 4 to 39 in two geographically distant populations, observed and expected heterozygosities ranged between 0. 370-0. 978 and 0. 426-0. 962, respectively. These loci are an important resource that will allow assessment of the current population genetic structure of this species, and enable monitoring of climate related changes in the species range and distribution. © 2012 Springer Science+Business Media Dordrecht.