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

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. Accepted: 3 January 2000     

Biogeographic responses of the copepod Calanus glacialis to a changing Arctic marine environment

Dramatic changes have occurred in the Arctic Ocean over the past few decades, especially in terms of sea ice loss and ocean warming. Those environmental changes may modify the planktonic ecosystem with changes from lower to upper trophic levels. This study aimed to understand how the biogeographic distribution of a crucial endemic copepod species, Calanus glacialis, may respond to both abiotic (ocean temperature) and biotic (phytoplankton prey) drivers. A copepod individual‐based model coupled to an ice‐ocean‐biogeochemical model was utilized to simulate temperature‐ and food‐dependent life cycle development of C. glacialis annually from 1980 to 2014. Over the 35‐year study period, the northern boundaries of modeled diapausing C. glacialis expanded poleward and the annual success rates of C. glacialis individuals attaining diapause in a circumpolar transition zone increased substantially. Those patterns could be explained by a lengthening growth season (during which time food is ample) and shortening critical development time (the period from the first feeding stage N3 to the diapausing stage C4). The biogeographic changes were further linked to large‐scale oceanic processes, particularly diminishing sea ice cover, upper ocean warming, and increasing and prolonging food availability, which could have potential consequences to the entire Arctic shelf/slope marine ecosystems.     

Records of the gadoid fish <Emphasis Type="Italic">Arctogadus glacialis</Emphasis> (Peters, 1874) in the European Arctic

The Arctogadus glacialis is endemic to the Arctic Ocean and its apparently disjunct circumpolar distribution range from the Siberian coast through the Chukchi Sea and the Canadian Arctic to the shelf off NE Greenland. Records of A. glacialis are scarce in the European Arctic and here we present all available and reliable records of the species in the area. Altogether, 296 specimens of A. glacialis are reported from 53 positions in the European Arctic during the period 1976–2008. The specimens were registered off Iceland and the Jan Mayen Island, northwest and northeast of Svalbard, northeast in the Barents Sea, and south and east off Franz Josef Land. The additional records show that A. glacialis display a circumpolar and more continuous distribution than described before. In the European Arctic, A. glacialis has been caught at 155–741 m depth with the highest abundance at 300–400 m. We therefore suggest that A. glacialis is more associated to the continental shelves surrounding the Arctic Ocean than previously thought. The length–weight relation of A. glacialis is similar across the European Arctic.     

The relationship between sea surface temperature and population change of Great Cormorants Phalacrocorax carbo breeding near Disko Bay,Greenland

Arctic seas have warmed and sea ice has retreated. This has resulted in range contraction and population declines in some species, but it could potentially be a boon for others. Great Cormorants Phalacrocorax carbo have a partially wettable plumage and seem poorly suited to foraging in Arctic waters. We show that rates of population change of Cormorant colonies around Disko Bay, Greenland, are positively correlated with sea surface temperature, suggesting that they may benefit from a warming Arctic. However, although Cormorant populations may increase in response to Arctic warming, the extent of expansion of their winter range may ultimately be limited by other factors, such as sensory constraints on foraging behaviour during long Arctic nights.     

Zooplankton boom and ice amphipod bust below melting sea ice in the Amundsen Gulf, Arctic Canada

Early summer in the Arctic with extensive ice melt and break-up represents a dramatic change for sympagic–pelagic fauna below seasonal sea ice. As part of the International Polar Year-Circumpolar Flaw Lead system study (IPY-CFL), this investigation quantified zooplankton in the meltwater layer below landfast ice and remaining ice fauna below melting ice during June (2008) in Franklin Bay and Darnley Bay, Amundsen Gulf, Canada. The ice was in a state of advanced melt, with fully developed melt ponds. Intense melting resulted in a 0.3- to 0.5-m-thick meltwater layer below the ice, with a strong halocline to the Arctic water below. Zooplankton under the ice, in and below the meltwater layer, was sampled by SCUBA divers. Dense concentrations (max. 1,400 ind. m⁻³) of Calanus glacialis were associated with the meltwater layer, with dominant copepodid stages CIV and CV and high abundance of nauplii. Less abundant species included Pseudocalanus spp., Oithona similis and C. hyperboreus. The copepods were likely feeding on phytoplankton (0.5–2.3 mg Chl-a m⁻³) in the meltwater layer. Ice amphipods were present at low abundance (<10 ind. m⁻²) and wet biomass (<0.2 g m⁻²). Onisimus glacialis and Apherusa glacialis made up 64 and 51% of the total ice faunal abundance in Darnley Bay and Franklin Bay, respectively. During early summer, the autochthonous ice fauna becomes gradually replaced by allochthonous zooplankton, with an abundance boom near the meltwater layer. The ice amphipod bust occurs during late stages of melting and break-up, when their sympagic habitat is diminished then lost.     

Haloperoxidase activities in Arctic macroalgae

Haloperoxidase activities were assayed in 21 species of Arctic green, red and brown macroalgae collected in the Kongsfjord at Ny-Ålesund (Svalbard). The enzymes were specific for bromide and/or iodide, but not for chloride. Highest bromoperoxidase activities were found in the brown seaweads Laminaria saccharina and L. digitata, whereas highest iodoperoxidase activities were measured in the green species Acrosiphonia sonderi and Enteromorpha compressa. Optimum pH for bromination ranged between pH 4 and 7.     

Life strategy and diet of Calanus glacialis during the winter�Cspring transition in Amundsen Gulf, south-eastern Beaufort Sea

The copepod Calanus glacialis plays a key role in the lipid-based energy flux in Arctic shelf seas. By utilizing both ice algae and phytoplankton, this species is able to extend its growth season considerably in these seasonally ice-covered seas. This study investigated the impacts of the variability in timing and extent of the ice algal bloom on the reproduction and population success of C. glacialis. The vertical distribution, reproduction, amount of storage lipids, stable isotopes, fatty acid and fatty alcohol composition of C. glacialis were assessed during the Circumpolar Flaw Lead System Study. Data were collected in the Amundsen Gulf, south-eastern Beaufort Sea, from January to July 2008 with the core-sampling from March to April. The reduction in sea ice thickness and coverage observed in the Amundsen Gulf in 2007 and 2008 affected the life strategy and reproduction of C. glacialis. Developmental stages CIII and CIV dominated the overwintering population, which resulted in the presence of very few CV and females during spring 2008. Spawning began at the peak of the ice algal bloom that preceded the precocious May ice break-up. Although the main recruitment may have occurred later in the season, low abundance of females combined with a potential mismatch between egg production/development to the first feeding stage and phytoplankton bloom resulted in low recruitment of C. glacialis in the early summer of 2008.     

Synergistic effects of ocean acidification and warming on overwintering pteropods in the Arctic

Ocean acidification and warming will be most pronounced in the Arctic Ocean. Aragonite shell‐bearing pteropods in the Arctic are expected to be among the first species to suffer from ocean acidification. Carbonate undersaturation in the Arctic will first occur in winter and because this period is also characterized by low food availability, the overwintering stages of polar pteropods may develop into a bottleneck in their life cycle. The impacts of ocean acidification and warming on growth, shell degradation (dissolution), and mortality of two thecosome pteropods, the polar Limacina helicina and the boreal L. retroversa, were studied for the first time during the Arctic winter in the Kongsfjord (Svalbard). The abundance of L. helicina and L. retroversa varied from 23.5 to 120 ind m^?2 and 12 to 38 ind m^?2, and the mean shell size ranged from 920 to 981 μm and 810 to 823 μm, respectively. Seawater was aragonite‐undersaturated at the overwintering depths of pteropods on two out of ten days of our observations. A 7‐day experiment [temperature levels: 2 and 7 °C, pCO₂ levels: 350, 650 (only for L. helicina) and 880 μatm] revealed a significant pCO₂ effect on shell degradation in both species, and synergistic effects between temperature and pCO₂ for L. helicina. A comparison of live and dead specimens kept under the same experimental conditions indicated that both species were capable of actively reducing the impacts of acidification on shell dissolution. A higher vulnerability to increasing pCO₂ and temperature during the winter season is indicated compared with a similar study from fall 2009. Considering the species winter phenology and the seasonal changes in carbonate chemistry in Arctic waters, negative climate change effects on Arctic thecosomes are likely to show up first during winter, possibly well before ocean acidification effects become detectable during the summer season.     

Regulation of gene expression is associated with tolerance of the Arctic copepod Calanus glacialis to CO2‐acidified sea water

Ocean acidification is the increase in seawater pCO₂ due to the uptake of atmospheric anthropogenic CO₂, with the largest changes predicted to occur in the Arctic seas. For some marine organisms, this change in pCO₂, and associated decrease in pH, represents a climate change‐related stressor. In this study, we investigated the gene expression patterns of nauplii of the Arctic copepod Calanus glacialis cultured at low pH levels. We have previously shown that organismal‐level performance (development, growth, respiration) of C. glacialis nauplii is unaffected by low pH. Here, we investigated the molecular‐level response to lowered pH in order to elucidate the physiological processes involved in this tolerance. Nauplii from wild‐caught C. glacialis were cultured at four pH levels (8.05, 7.9, 7.7, 7.5). At stage N6, mRNA was extracted and sequenced using RNA‐seq. The physiological functionality of the proteins identified was categorized using Gene Ontology and KEGG pathways. We found that the expression of 151 contigs varied significantly with pH on a continuous scale (93% downregulated with decreasing pH). Gene set enrichment analysis revealed that, of the processes downregulated, many were components of the universal cellular stress response, including DNA repair, redox regulation, protein folding, and proteolysis. Sodium:proton antiporters were among the processes significantly upregulated, indicating that these ion pumps were involved in maintaining cellular pH homeostasis. C. glacialis significantly alters its gene expression at low pH, although they maintain normal larval development. Understanding what confers tolerance to some species will support our ability to predict the effects of future ocean acidification on marine organisms.     

Lipid composition of copepodite stages and adult females of Calanus glacialis in Arctic waters of the Barents Sea

Summary The composition of lipid classes and their component fatty acid are described for copepodite stages III, IV, V, and adult females of Calanus glacialis sampled from Arctic waters of the Barents Sea during summer. Was esters were the principal component of the lipid in all copepodite stages examined, averaging 73% over all the stages. The proportion of triacylglycerols varied from 1.5% to 10.5% of total lipid among copepodite stages. The lipids of adult females contained lower levels of wax esters and higher levels of triacylglycerols than copepodite stages III, IV and V. Fatty alcohols of wax esters from copepods sampled in June and July were dominated by 20:1 (n-9) and 22:1 (n-11) alcohols with the proportion of 20:1 (n-9) increasing from stages III to adult female. 14:0 and 16:0 alcohols were the principal fatty alcohols of wax esters of a sample comprising mainly of copepodites stage III taken in August. 16:1 (n-7), 20:1 (n-9) and 20:5 (n-3) were the major fatty acids in the was esters of animals captured in June and July whereas 18:1 (n-9) predominated in the August sample. The polar lipids of the copepodite stage III from August also contained lower levels of polyunsaturated fatty acids than from all stages of copepods from June and July. The data are discussed in relation to life cycle strategies and trophic aspects of Calanus glacialis in the Arctic pelagic community of the Barents Sea.     

Relations between the thermocline meso and microzooplankton,chlorophyll a and primary production distributions in Lancaster Sound

Summary Primary productivity in Lancaster Sound was estimated from chlorophyll and light irradiance profiles and light saturation measurments. Mesozooplankton concentrations were measured using an in situ zooplankton counter mounted on a Batfish and with the BIONESS. Microzooplankton concentrations were estimated from samples taken with the BIONESS. The horizontal and vertical distribution of the two groups of zooplankton are described and related to the distribution of chlorophyll primary production and physical features of the Sound. A westward intrusion of Baffin Bay water along the north shore of the sound which mixed with Arctic water flowing easterly resulted in a complex physical environment in the sound. The admixing of these water masses resulted in a highly variable biological structure. The thermocline was the main physical feature that affected the depth of the chlorophyll layer and the levels of primary production, with a shallow thermocline resulting in higher primary production. The admixing of Baffin Bay water with Arctic water at different areas of the sound combined with the different times of appearance of open water resulted in the populations of Calanus hyperboreus and C. glacialis showing different age structures, the youngest being in the Arctic water at the western end of the Sound. Microzooplankton populations were concentrated at various depths depending on the species. Oithona juveniles preferred the upper 10 m when the thermocline was near the surface. Most macrozooplankton were found below the thermocline and the primary production maxium. The largest concentrations of microzooplankton were found at the western end of the Sound in the Arctic water believed to be still in the biological spring.     

Lipids in copepodite stages of <Emphasis Type="Italic">Calanus glacialis</Emphasis>

Calanus glacialis is a key herbivore in Arctic shelf seas. It feeds on primary producers and accumulates large energy reserves, primarily as wax esters. Lipid classes, fatty acids (FAs) and fatty alcohols (FAlcs) from copepodite stage II (CII) to adult females (AF) from Kongsfjorden, Svalbard, were studied in May 2004. Wax esters were the dominating lipid class in all stages, ranging from 34% of total lipids in CII to 60% in CIII–CV. Triacylglycerols increased from 8% of total lipids in CII to 23% in AF. In the earlier stages, 16:1n7 and 16:0 FAs and FAlcs were the major components of the neutral lipids, whereas the later stages were mainly characterized by the long-chained FAs and FAlcs 20:1n9 and 22:1n11. C. glacialis utilizes the short spring bloom to build up lipid reserves, mainly as wax esters, and it also incorporates effectively essential polyunsaturated FAs such as 20:5n3 and 22:6n3 in its polar lipids.     

Seasonal dynamics of mesozooplankton in the Arctic Kongsfjord (Svalbard) during year-round observations from August 1998 to July 1999

We conducted a year-round mesozooplankton study in the Arctic Kongsfjord from August 1998 until July 1999 to investigate seasonal abundance and vertical as well as stage distributions of the prevalent taxa. It is the first investigation in Kongsfjord that covers the Arctic winter season and provides reasonable estimates also of small-sized copepod species. Abundant smaller copepods comprised Oithona similis, Pseudocalanus minutus, Microcalanus spp., Triconia borealis and Acartia longiremis. Among the larger copepods, Calanus finmarchicus, C. glacialis, C. hyperboreus and Metridia longa dominated. The thecosome pteropod Limacina helicina was also an important component. Abundance maxima occurred in November (988,669 ind. m^?2) with one to two orders of magnitude higher numbers as compared to all other months (39,832–200,067 ind. m^?2). The summers of 1998 and 1999 were characterized by intrusions of Atlantic water, but the community was not entirely dominated by advected boreal species. During winter, the majority of the mesozooplankton occurred below 100 m. Advection is the most likely reason for the accumulation of zooplankton at depth in winter, but local production may also contribute to high overwintering numbers. Much lower abundances of most species in spring suggest high winter mortality and emphasize the importance of sufficient reproductive success during the previous summer to ensure enough winter survivors as seed stock for the coming reproductive season. This study was conducted prior to the recent warming trend in the Arctic. Therefore, it provides valuable baseline data and allows comparing present and future states of the zooplankton community in Kongsfjord.     

Ovary development in Greenland halibut Reinhardtius hippoglossoides in west Greenland waters

Maturity in adult female Greenland halibut Reinhardtius hippoglossoides was studied in three areas in west Greenland waters: the inshore area in Disko Bay and two offshore areas, Baffin Bay and Davis Strait. The aim was to monitor maturity changes in the inshore fjords of Disko Bay over an extended period from winter to autumn and compare these findings with specimens from Baffin Bay and the presumed spawning area in Davis Strait. A significant difference in maturity level was observed in and between the three areas. In Disko Bay maturity indices increased significantly in August and September both with respect to the gonado‐somatic index ( I _G) and the size in the leading oocyte cohort. In the period February to May no significant changes were observed. Mature ovaries were only observed among fish >80 cm total length and only among a fraction of these large fish. Offshore areas of Baffin Bay, even though poorly sampled, showed similar signs in the maturity indices as in Disko Bay. Relative to Disko Bay and Baffin Bay, female fish in Davis Strait had more progressed maturity indices. Furthermore, almost all fish in Davis Strait showed signs of progressed maturity contrary to Disko and Baffin Bay. A large proportion of the Greenland halibut in Disko and Baffin Bay apparently did not begin the maturation cycle until very late in their life history or were repeat spawners with a multi‐year maturation cycle. These observations could thus support the hypothesis that Greenland halibut have a prolonged adolescent phase. Atresia was highest in the early phases of maturation in Greenland halibut but relatively high levels of atresia were also observed in fish in more advanced maturity phase. The first was ascribed to fecundity regulation while the latter could be linked to the fish's fitness condition but it was not possible to show this with the available condition index.     

The extreme disjunction between Beringia and Europe in Ranunculus glacialis s. l. (Ranunculaceae) does not coincide with the deepest genetic split – a story of the importance of temperate mountain ranges in arctic–alpine phylogeography

The arctic–alpine Ranunculus glacialis s. l. is distributed in high‐mountain ranges of temperate Europe and in the North, where it displays an extreme disjunction between the North Atlantic Arctic and Beringia. Based on comprehensive sampling and employing plastid and nuclear marker systems, we (i) test whether the European/Beringian disjunction correlates with the main evolutionary diversification, (ii) reconstruct the phylogeographic history in the Arctic and in temperate mountains and (iii) assess the susceptibility of arctic and mountain populations to climate change. Both data sets revealed several well‐defined lineages, mostly with a coherent geographic distribution. The deepest evolutionary split did not coincide with the European/Beringian disjunction but occurred within the Alps. The Beringian lineage and North Atlantic Arctic populations, which reached their current distribution via rapid postglacial colonization, show connections to two divergent pools of Central European populations. Thus, immigration into the Arctic probably occurred at least twice. The presence of a rare cpDNA lineage related to Beringia in the Carpathians supports the role of these mountains as a stepping stone between temperate Europe and the non‐European Arctic, and as an important area of high‐mountain biodiversity. The temperate and arctic ranges presented contrasting phylogeographic histories: a largely static distribution in the former and rapid latitudinal spread in the latter. The persistence of ancient lineages with a strictly regional distribution suggests that the ability of R. glacialis to survive repeated climatic changes within southern mountain ranges is greater than what recently was predicted for alpine plants from climatic envelope modelling.     

Lipid composition and trophic relationships of krill species in a high Arctic fjord

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.     

Dynamics of coexisting Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus populations in a high-Arctic fjord

We studied the population dynamics of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Billefjorden, Svalbard (78°40N). All three species reproduced in the fjord with different timing. The maximum abundance of Calanus spp. copepodite stages peaked on the 11th of July (29,000 ind m^–2). C. glacialis was the dominant species accounting for 60–80% of the total Calanus abundance. C. finmarchicus appear to thrive in the fjord despite the low temperatures (–1.86°C to 5°C) and accounted for 20–30% of the total population. C. hyperboreus contributed less to the total abundance (5–20%). A 1-year life cycle is suggested for C. finmarchicus and C. hyperboreus in the fjord, C. glacialis has a 1- to 2-year life cycle. Highest mortality rates were observed for copepodite stage CV in C. finmarchicus and C. glacialis (0.09 and 0.075 d^–1, respectively) and for females in C. hyperboreus (0.149 d^–1). Mortality of copepodite stages was substantially lower in C. glacialis than in the other species. This is particularly obvious in the early and numerous copepodite stages (CI + CII) during the period of recruitment to these stages. This suggests that differences in secondary production in Arctic pelagic ecosystems are controlled partly by population loss rates.     

Size,age and diet of polar cod,Boreogadus saida (Lepechin 1773), in ice covered waters

Summary Polar cod (Boreogadus saida) associated with drifting sea-ice were collected in the western Barents sea and north of Svalbard with dip-nets while SCUBA-diving in 1986 and 1987. Length-frequency measurements and otolith-readings suggested that the specimens were either one or two years old. The diet of fish from the western Barents sea (first-year ice) consisted mainly of copepods (Calanus finmarchicus, Calanus glacialis) and the hyperiid amphipod Parathemisto libellula. Fish collected north of the Svalbard archipelago (multi-year ice) had a more diverse diet, in which P. libellula and the sympagic amphipod Apherusa glacialis contributed more to the total diet biomass than copepods.     

Geographic variation in cranial morphology of narwhals (Monodon monoceros) from Greenland and the eastern Canadian Arctic

We analyzed variation in nine non-metric and eight metric variables in the skulls of 132 narwhals (Monodon monoceros) from five localities in Greenland (Inglefield Bredning, Melville Bay, Uummannaq, Disko Bay, and Scoresby Sound) and one in the eastern Canadian Arctic (Eclipse Sound). Metric variables were used to compare the combined Disko Bay and Uummannaq samples with the samples from Inglefield Bredning and Scoresby Sound using three different multivariate techniques for each sex. None of the results were significant. Seven of the non-metric variables were independent of age and sex and were used in comparing samples from the six localities. No differences were found among the four localities in West Greenland, but differences were found in two of the non-metric variables between the combined West Greenland sample and the one from Scoresby Sound. A major shortcoming of the analysis based on metric data was the small sample size from several of the areas, which resulted in low statistical power. Genetic as well as environmental factors could explain the differences detected here between narwhals living along the west and the east coasts of Greenland.