Slawomir Kwasniewski

Lipids and life strategies of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in late autumn, Kongsfjorden, Svalbard

Abstract Stage IV and V copepodites were the dominant forms of Calanus finmarchicus, C. glacialis and C. hyperboreus in Kongsfjorden in late September 1997. Stage IV and V copepodites of C. glacialis and C. hyperboreus were rich in lipid, largely wax esters, and were well fitted to overwinter. Stage IV copepodites of C. finmarchicus were also rich in wax esters, but stage V copepodites of C. finmarchicus were less wax ester-rich. Large size increments between stage IV and V copepodites and between stage V copepodites and females were noted in C. finmarchicus. A very large increment between stage IV and V copepodites was noted for C. glacialis but the size difference between stage V copepodites and females was very small in this species. Particularly large increments were noted between stage IV and V copepodites of C. hyperboreus and also between stage V copepodites and females of this species. The very large, wax ester-rich C. hyperboreus is well adapted to survive the most extreme variations in the Arctic, in Arctic basin waters, whereas the smaller, wax ester-rich C. glacialis is adapted to survive less extreme Arctic variations, as in Arctic shelf waters. The smallest of the three, C. finmarchicus, is best adapted to survive the more predictable waters of the North Atlantic and the Barents Sea.

Distribution, reproduction and development of Calanus species in the Northeast water in relation to environmental conditions

Abstract The vertical and horizontal distribution of mesozooplankton biomass and its composition, together with the reproduction and development of the three dominant herbivorous copepods Calanus finmarchicus, C. glacialis and C. hyperboreus , were studied in the Northeast Water between the end of May and end of July, 1993. Biomass varied between 0.15 and 10.6 g m −2 and was lower on the shallow banks. Highest biomass was found on the shelf slope and in the entrances to Belgica and Westwind Troughs. There was no seasonal trend during the study period. Among the zooplankton taxa, copepods were dominant, with 84% biomass of all other taxa, followed by chaetognaths with 14%. The large Calanus species made up 91% of copepod biomass. The most abundant species, C. glacialis , inhabited areas of low current speeds on Belgica and Ob Bank, C. hyperboreus dominated shelf slope and trough stations, while C. finmarchicus was most abundant in the Return Atlantic Current along the shelf slope and also eastern Belgica Trough. C. glacialis was the only Calanus species spawning during this period, but young copepodites of the other species were also abundant. Egg production of C. glacialis was at a maximum by our arrival and continued to at least mid August. According to the results from starvation experiments, its egg production was fuelled by food uptake, but was decoupled from phytoplankton chlorophyll until July, indicating ice-algae and microzooplankton as an alternative food source. Only when the polynya approached its maximum extent was a close relationship to phytoplankton established. Due to both spatial and temporal heterogeneity of the occurrence of young copepodite stages it was impossible to follow the growth of cohorts of developmental stages. Herbivorous carbon requirements estimated from egg production rates were mostly less than one third of the phytoplankton stock. From egg production and the distribution of young stages, the outer Westwing Trough seems to be the centre of biological activities. This may be related to the formation of young ice in winter in this area, which was found to carry a large mass of under-ice algae. High biomass but low production in Belgica Trough indicate this as an advective and expatriated community of C. finmarchicus and C. hyperboreus , where grazing is negligible, at least during early summer.

Leads in Arctic pack ice enable early phytoplankton blooms below snow-covered sea ice

The Arctic icescape is rapidly transforming from a thicker multiyear ice cover to a thinner and largely seasonal first-year ice cover with significant consequences for Arctic primary production. One critical challenge is to understand how productivity will change within the next decades. Recent studies have reported extensive phytoplankton blooms beneath ponded sea ice during summer, indicating that satellite-based Arctic annual primary production estimates may be significantly underestimated. Here we present a unique time-series of a phytoplankton spring bloom observed beneath snow-covered Arctic pack ice. The bloom, dominated by the haptophyte algae Phaeocystis pouchetii, caused near depletion of the surface nitrate inventory and a decline in dissolved inorganic carbon by 16 ± 6 g C m−2. Ocean circulation characteristics in the area indicated that the bloom developed in situ despite the snow-covered sea ice. Leads in the dynamic ice cover provided added sunlight necessary to initiate and sustain the bloom. Phytoplankton blooms beneath snow-covered ice might become more common and widespread in the future Arctic Ocean with frequent lead formation due to thinner and more dynamic sea ice despite projected increases in high-Arctic snowfall. This could alter productivity, marine food webs and carbon sequestration in the Arctic Ocean.

Lipids and fatty acids in the copepod Jaschnovia brevis (Jaschnov) and in particulates from Arctic waters

Abstract. The small, sub-ice copepod Jaschnovia brevis is rich in triacylglycerols, suggesting a feeding behaviour not constrained to the seasonal phytoplankton bloom. The copepods triacylglycerol reserves contain: the diatom biomarkers 16:1n-7 (23.9%), 20:5n-3 (8.5%) and C16 PUFA (1.3%), the flagellate biomarkers 18:4n-3 (3.7%) and 22:6n-3 (3.3%), and the Calanus copepod biomarkers 20:1n-9 (7.7%) and 22:1n-11 (6.2%). Total lipid from particulates in the water column contained polar lipid (45.0%), wax esters (24.9%) and triacylglycerols (11.2%) as major components. The total lipids in the particulates were rich in 18:1n-9 (31.5%) and 16:0 (21.2%), and relatively rich in 18:0 (7.8%) and 18:2n-6 (9.2%). The triacylglycerols in the particulates contained 16:1n-7 (20.7%), C16 PUFA (4.1%), 18:4n-3 (1.9%), 20:5n-3 (3.6%), 22:6n-3 (1.9%), 20:1n-9 (5.2%) and 22:1n-11 (3.9%). The polar lipids in the particulates contained 16:1n-7 (17.3%), C16 PUFA (7.8%), 18:4n-3 (3.3%), 20:5n-3 (14.5%) and 22:6n-3 (9.6%). The fatty alcohols in the wax esters of the particulates were mainly 16:0 (11.3%), 20:1n-9 (21.1%) and 22:1n-11 (30.6%). The nature of the particulates, their possible origin in living and non-living material, and their role in the nutrition of J. brevis are discussed.

Zooplankton in Svalbard fjords on the Atlantic–Arctic boundary

Zooplankton abundance and community structures were studied in three west Spitsbergen fjords at the beginning of the warm phase, which seem to have entered in 2006. Sampling was conducted in summer 2007 at stations distributed along transects in Hornsund, Isfjorden and Kongsfjorden. Variations in zooplankton standing stocks and community structures (assessing taxonomic diversity and zoogeographical affiliations) were analysed in relation to the environmental variables using multivariate techniques. The hydrographic conditions in Hornsund were influenced by the cold Arctic Water, whereas those in Isfjorden and especially in Kongsfjorden were, to a greater extent, under the influence of the warm Atlantic Water. High abundances of both meroplankton and holoplankton organisms were observed in Kongsfjorden, with high contributions of boreal and ubiquitous species (Calanus finmarchicus and Oithona similis, respectively). In Hornsund at the same time, the zooplankton consisted mainly of boreo-Arctic and Arctic species, the abundances of which were comparable along the West Spitsbergen Shelf. Our results indicate that the difference in hydrography had measurable effects on the zooplankton community in the study area. Furthermore, by comparing regions of contrasting oceanographic conditions, we present evidence as to how the zooplankton structure will change in the Arctic ecosystems if the warming trends continue to operate with the same dynamics. The advection of Atlantic waters to the Arctic seas may lead to changes in zooplankton structure, with increased abundance and contributions of boreal and small ubiquitous species. The ‘warmer Arctic fjords’ scenarios may also induce more rapid development of both holoplankton and meroplankton populations and, consequently, modify the trophic interactions in plankton communities.

Genetics redraws pelagic biogeography of Calanus

Planktonic copepods of the genus Calanus play a central role in North Atlantic/Arctic marine food webs. Here, using molecular markers, we redrew the distributional ranges of Calanus species inhabiting the North Atlantic and Arctic Oceans and revealed much wider and more broadly overlapping distributions than previously described. The Arctic shelf species, C. glacialis, dominated the zooplankton assemblage of many Norwegian fjords, where only C. finmarchicus has been reported previously. In these fjords, high occurrences of the Arctic species C. hyperboreus were also found. Molecular markers revealed that the most common method of species identification, prosome length, cannot reliably discriminate the species in Norwegian fjords. Differences in degree of genetic differentiation among fjord populations of the two species suggested that C. glacialis is a more permanent resident of the fjords than C. finmarchicus. We found no evidence of hybridization between the species. Our results indicate a critical need for the wider use of molecular markers to reliably identify and discriminate these morphologically similar copepod species, which serve as important indicators of climate responses.

Variations in the structural and functional diversity of zooplankton over vertical and horizontal environmental gradients en route to the Arctic Ocean through the Fram Strait

A multi-scale approach was used to evaluate which spatial gradient of environmental variability is the most important in structuring zooplankton diversity in the West Spitsbergen Current (WSC). The WSC is the main conveyor of warm and biologically rich Atlantic water to the Arctic Ocean through the Fram Strait. The data set included 85 stratified vertical zooplankton samples (obtained from depths up to 1000 metres) covering two latitudinal sections (76°30’N and 79°N) located across the multi-path WSC system. The results indicate that the most important environmental variables shaping the zooplankton structural and functional diversity and standing stock variability are those associated with depth, whereas variables acting in the horizontal dimension are of lesser importance. Multivariate analysis of the zooplankton assemblages, together with different univariate descriptors of zooplankton diversity, clearly illustrated the segregation of zooplankton taxa in the vertical plane. The epipelagic zone (upper 200 m) hosted plentiful, Oithona similis-dominated assemblages with a high proportion of filter-feeding zooplankton. Although total zooplankton abundance declined in the mesopelagic zone (200–1000 m), zooplankton assemblages in that zone were more diverse and more evenly distributed, with high contributions from both herbivorous and carnivorous taxa. The vertical distribution of integrated biomass (mg DW m-2) indicated that the total zooplankton biomass in the epipelagic and mesopelagic zones was comparable. Environmental gradients acting in the horizontal plane, such as the ones associated with different ice cover and timing of the spring bloom, were reflected in the latitudinal variability in protist community structure and probably caused differences in succession in the zooplankton community. High abundances of Calanus finmarchicus in the WSC core branch suggest the existence of mechanisms advantageous for higher productivity or/and responsible for physical concentration of zooplankton. Our results indicate that regional hydrography plays a primary role in shaping zooplankton variability in the WSC on the way to the Arctic Ocean, with additional effects caused by biological factors related to seasonality in pelagic ecosystem development, resulting in regional differences in food availability or biological production between the continental slope and the deep ocean regions.

Supplementary diet components of little auk chicks in two contrasting regions on the West Spitsbergen coast

The complete diet composition structure of the most numerous planktivorous sea bird, little auk (Alle alle), in the European Arctic, is still not fully recognized. Although regular constituents of little auk chick diets, the copepods, Calanus glacialis and C. finmarchicus have been previously relatively well described, more taxa were frequent ingredients of the bird’s meals. Therefore, the role of the little auks supplementary diet components (SDCs) at two colonies in the Svalbard Archipelago, Hornsund and Magdalenefjorden, in 2007–2009, is a main subject of this comparative study. Because the SDCs often consisted of scarce but large zooplankters, this investigation was focused on biomass as a proxy of the SDCs’ energy input. Although the total biomass of the food delivered to chicks in both colonies was similar, in Magdalenefjorden, the proportion of SDCs was twice that found in Hornsund. The main SDCs in Hornsund were Decapoda larvae (with predominating Pagurus pubescens) and Thysanoessa inermis, whereas the main SDCs in Magdalenefjorden were C. hyperboreus and Apherusa glacialis. Previous investigations, which indicated lipid richness of SDCs, together with our ecological results from the colonies, suggest that this category might play a compensatory role in little auk chick diets. The ability to forage on diverse taxa may help the birds to adapt to ongoing Arctic ecosystem changes.

Interannual zooplankton variability in the main pathways of the Atlantic water flow into the Arctic Ocean (Fram Strait and Barents Sea branches)

Original Article Interannual zooplankton variability in the main pathways of the Atlantic water flow into the Arctic Ocean (Fram Strait and Barents Sea branches) Marta Gluchowska*, Padmini Dalpadado, Agnieszka Beszczynska-Möller, Anna Olszewska, Randi B. Ingvaldsen, and Slawomir Kwasniewski Institute of Oceanology, Polish Academy of Sciences (IO PAN), Powstancow Warszawy 55, Sopot 81-712, Poland Institute of Marine Research (IMR), P.O. Box 1870 Nordnes, Bergen 5817, Norway *Corresponding author: tel: þ48 58 7311786; fax: þ48 58 551 21 30; e-mail: mgluchowska@iopan.gda.pl

Seasonal Variation in Transport of Zooplankton Into the Arctic Basin Through the Atlantic Gateway, Fram Strait

The largest contribution of oceanic heat to the Arctic Ocean is the warm Atlantic Water (AW) inflow through the deep Fram Strait. The AW current also carries Atlantic plankton into the Arctic Basin and this inflow of zooplankton biomass through the Atlantic-Arctic gateway far exceeds the inflow through the shallow Pacific-Arctic gateway. However, because this transport has not yet been adequately quantified based on observational data, so that the present contribution is poorly defined, and future changes in Arctic zooplankton communities are difficult to project and observe. Our objective was to quantify the inflow of zooplankton biomass through the Fram Strait during different seasons, including winter. We collected data with high spatial resolution covering hydrography (CTD), currents (ADCP and LADCP) and zooplankton distributions (LOPC and MultiNet) from surface to 1000 m depth along two transects crossing the AW inflow during three cruises in January, May and August 2014. Long-term variations (1997-2016) in the AW inflow were analyzed based on moored current meters. Water transport across the inflow region was of the same order of magnitude during all months (January 2.2 Sv, May 1.9 Sv, August 1.7 Sv). We found a higher variability in zooplankton transport between the months (January 51 kg C s-1, May 34 kg C s-1, August 50 kg C s-1), related to seasonal changes in the vertical distribution of zooplankton. However, high abundances of carbon-rich copepods were observed in the AW inflow during all months. Surface patches with high abundances of C. finmarchicus, Microcalanus spp., Pseudocalanus spp. and Oithona similis clearly contributed to the advected biomass, also in winter. The data reveal that the phenology of species is important for the amount of advected biomass, and that the advective input of zooplankton carbon into the Arctic Basin is important during all seasons. The advective zooplankton input might be especially important for mesopelagic planktivorous predators that were recently observed in the region, particularly during winter. The inflow of C. finmarchicus with AW was estimated to be in the order of 500 000 metric tons C y-1, which compares well to modeled estimates.