Lipids and trophic interactions of ice fauna and pelagic zooplankton in the marginal ice zone of the Barents Sea

Gammarus wilkitzkii, Apherusa glacialis, Onismus nanseni, Onismus glacialis, Boreogadus saida, Parathemisto libellula and Calanus hyperboreus, collected in late June in the Barents Sea marginal ice zone, contained substantial levels (28–51% of the dry mass) of total lipid, the highest levels (51% and 41% respectively) being in  A. glacialis and  C. hyperboreus. Neutral lipids were present in greater amounts than polar lipids in all species. Triacylglycerols were major neutral lipids in A. glacialis, G. wilkitzkii and O. nanseni; triacylglycerols and wax esters were present in similar amounts in O. glacialis; higher levels of wax esters than triacylglycerols occurred in P. libellula; wax esters greatly exceeded triacylglycerols in C. hyperboreus, the opposite being true for B. saida. Diatom fatty acid markers were prominent in the triacylglycerols of G. wilkitzkii, O. nanseni, O. glacialis and, particularly, of  A. glacialis; 20:1(n-9) and 22:1(n-11) moieties were abundant in wax esters of G. wilkitzkii, O. nanseni, O. glacialis, P. libellula and  C. hyperboreus, and in triacylglycerols of B. saida. We deduce that  A. glacialis feeds mainly on ice algae and phytodetritus, G. wilkitzkii and the Onismus spp. feed on calanoid copepods as well as ice algae, whereas P. libellula and especially B. saida feed extensively on calanoid copepods. Accepted: 17 May 1998     

Vertical distributions of primary production and grazing by Calanus glacialis jaschnov and C. hyperboreus krøyer in Arctic waters (Barents Sea)

Summary Simultaneous measurements of phytoplankton primary production, and zooplankton grazing by Calanus glacialis and C. hyperboreus were carried out over a 24h period at two stations northeast of Spitzbergen in July and August 1985. Phytoplankton biomass was concentrated in a sub-surface maximum, and our results suggested that 65% to 90% of the carbon produced was grazed by these copepods. At both stations maximum grazing rates by these zooplankton were observed within and below the depth of maximum primary production. At least one third of regenerated primary production could be sustained by ammonia excreted by the large zooplankton species Calanus glacialis and C. hyperboreus.     

Intra-seasonal variation in zooplankton availability,chick diet and breeding performance of a high Arctic planktivorous seabird

Different phenological responses to climate changes by species representing preys and predators may lead to mismatch between functionally dependent components of an ecosystem, with important effects on its structure and functioning. Here, we investigate within-season variation in zooplankton availability, chick diet composition and breeding performance of a small planktivorous seabird, the little auk (Alle alle) in two large colonies in Hornsund and Magdalenefjorden, Spitsbergen, differing in synchrony of breeding (11-day vs. 22-day hatching period, respectively). Assuming similar zooplankton phenology and existing differences in duration of the little auk breeding period, we expected lower availability of the preferred food in the less synchronized colony in Magdalenefjorden and in consequence a negative effect on nestling body mass and survival. We found that in both colonies Calanus glacialis (copepodite stage CV) was the most important prey item in the chick diet making up 68–87 % of the biomass and energy of all prey items. The only exception was the end of the chick-rearing period in Magdalenefjorden, when contribution of this prey item was significantly lower (24–26 %). Thus, late breeders in Magdalenefjorden were apparently mismatched regarding C. glacialis CV availability. However, the hatching date did not affect birds fitness (reproductive output and chick pre-fledging mass) significantly. Results of our study indicate that little auks breeding on Spitsbergen can respond to a wide range of environmental conditions and prey availabilities through the plasticity of their foraging behaviour, which may help them to maintain their optimum fitness level in changing and unpredictable environments.     

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     

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.     

Winter–spring feeding and metabolism of Arctic copepods: insights from faecal pellet production and respiration measurements in the southeastern Beaufort Sea

Faecal pellet production (FPP) and respiration rates of Calanus glacialis, C. hyperboreus and Metridia longa were measured under land-fast ice in the southeastern Beaufort Sea during the winter–spring transition (March–May 2004) prior to the phytoplankton spring bloom. Despite different overwintering and life cycle strategies and remaining low concentrations of suspended chlorophyll a and particulate organic matter, all species showed increasing FPP rates in spring. A corresponding increase in respiration was only observed in C. glacialis, while respiration remained constant in C. hyperboreus and M. longa. In C. glacialis and C. hyperboreus calculated ingestion covered respiratory expenditures. The constancy of the oil sac volume in M. longa suggests that the animals fed during winter-spring. Pre-bloom grazing as shown here seems to acclimate the copepod populations physiologically for the upcoming high feeding season, so that they are able to resume maximum grazing and reproduction as soon as the phytoplankton bloom is initiated.     

Distribution of dominant calanoid copepod species in the Greenland sea during late fall

Summary Between 6 November and 12 December 1988, vertical distributions of Calanus finmarchicus, C. hyperboreus, C. glacialis and Metridia longa were studied at three stations in the Arctic water of the Greenland Sea Gyre (GSG) and compared with two stations in the Atlantic water (AW) of the Westspitsbergen Current. Nine depth strata down to 3,000 m were sampled. C.finmarchicus was most abundant in AW, C. hyperboreus in GSG, M. longa showed no preference and C. glacialis was rare everywhere. Stage composition differed with species and water mass. Vertical distribution varied also with water mass in all species but C. hyperboreus, which was always centered between 1,000 and 1,500 m. The other species were concentrated in the upper 300 m in AW and between 1,000 and 1,500 m in GSG, although not all stages followed this trend. Ontogenetic vertical migration leads to significant dislocations of organic matter from the euphotic zone to great depth in the Greenland Sea. In egg production experiments, C. hyperboreus spawned up to 149 eggs female^–1 day^–1. Implications of stage composition, sex ratio, and gonad maturation on copepod life cycles are discussed.     

Inter‐ and intraspecific variation in body‐ and genome size in calanoid copepods from temperate and arctic waters

The tendency of ectotherms to get larger in the cold (Bergmann clines) has potentially great implications for individual performance and food web dynamics. The mechanistic drivers of this trend are not well understood, however. One fundamental question is to which extent variation in body size is attributed to variation in cell size, which again is related to genome size. In this study, we analyzed body and genome size in four species of marine calanoid copepods, Calanus finmarchicus, C. glacialis, C. hyperboreus and Paraeuchaeta norvegica, with populations from both south Norwegian fjords and the High Arctic. The Calanus species showed typical interspecific Bergmann clines, and we assessed whether they also displayed similar intraspecific variations—and if correlation between genome size and body size differed between species. There were considerable inter‐ as well as intraspecific variations in body size and genome size, with the northernmost populations having the largest values of both variables within each species. Positive intraspecific relationships suggest a functional link between body and genome size, although its adaptiveness has not been settled. Impact of additional drivers like phylogeny or specific adaptations, however, was suggested by striking divergences in body size – genome size ratios among species. Thus, C. glacialis and C. hyperboreus, had fairly similar genome size despite very different body size, while P. norvegica, of similar body size as C. hyperboreus, had the largest genome sizes ever recorded from copepods. The inter‐ and intraspecific latitudinal body size clines suggest that climate change may have major impact on body size composition of keystone species in marine planktonic food webs.     

Differences in food delivered to chicks by males and females of little auks ( Alle alle ) on South Spitsbergen

Sex differences in the quality and quantity of food loads brought to little auk (Alle alle) chicks were investigated in a large colony in Hornsund (South Spitsbergen). Adults returning to the colony were caught in mist-nets and food loads were taken from their gular pouch. The sex of each bird was determined by means of molecular methods. Females brought significantly more food per single load than males in terms of wet weight (30% more on average), number of prey items (46%) and energy contents (39%). However, there was no difference between the sexes in the size of prey taken. Energy-rich Calanus glacialis, originating from cold Arctic waters, was the most frequent prey item and made up the majority of food loads brought by both males and females (75 and 72%, respectively). This indicates that both sexes forage mainly in Arctic waters. However, differences in the proportion of cold water Calanus species (C. glacialis and C. hyperboreus), warm water Calanus species (C. finmarchicus), as well as other taxa, between males and females may suggest different ways of exploiting the feeding area.     

Body sizes,development rates,and genome sizes among Calanus species

Data on embryonic and larval development times (D) of Calanus species are analysed using Blehrádek's temperature (T) function, D = a (T – )^b, with b = – 2.05 as in previous studies. Among these species, for embryonic duration varies directly with temperatures in their geographical ranges and a is related to egg diameter. Using and b from embryonic durations, the fitted values of a for older stages are related to body sizes. Roughly estimated nucleus numbers in single adult females of C. finmarchicus, glacialis and hyperboreus were similar at 72 000, 85 000, and 96 000 respectively. Genome sizes (2C) of adult females are ca. 13 pg DNA in C. finmarchicus and pacificus, ca. 17 pg in C. sinicus, ca. 21 pg in C. helgolandicus and marshallae, and ca. 25 pg in C. glacialis and hyperboreus. These correspond roughly to body sizes and temperature-corrected development rates, quite precisely so in the sibling pair C. finmarchicus and C. glacialis, suggesting that, given similar nucleus numbers, there is nucleotypic control of whole-organism characteristics.     

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.     

Arctic seabird food chains explored by fatty acid composition and stable isotopes in Kongsfjorden, Svalbard

Marine birds are important predators in the marine ecosystem, and dietary studies can give useful information about their feeding ecology, food webs and oceanographic variability. The aim of this study was to increase our understanding of the diet and trophic level of the seabirds breeding in Kongsfjorden, Svalbard. We have used fatty acids and stable isotopes, both of which integrate diet information over space and time, to determine trophic relationships in marine food webs. Fatty acid compositions of muscle from Little auk (Alle alle), Brünnich’s guillemot (Uria lomvia), Black-legged kittiwake (Rissa tridactyla), Northern fulmar (Fulmarus glacialis) and Glaucous gull (Larus hyperboreus) were determined and compared with their prey species. Canonical analysis (CA) showed that fatty acid composition differed among the five seabird species. Little auk, Black-legged kittiwake and Northern fulmar had high levels of the Calanus markers 20:1n9 and 22:1, indicating that these seabirds are a part of the Calanus food chain. Brünnich’s guillemot differed from the other species with much lower levels of 20:1n9 and 22:1. Brünnich’s guillemot is a pursuit diver feeding on fish and amphipods deeper in the water column, below 30 m. Glaucous gull also differed from the other seabird species, with a larger variation in the fatty acid composition indicating a more diverse diet. Trophic level analysis placed Little auk at the lowest trophic level, Brünnich’s guillemot and Black-legged kittiwake at intermediate levels and Glaucous gull and Northern fulmar at the highest trophic level.     

The role of copepods in the planktonic ecosystems of mixed and stratified waters of the European shelf seas

The European shelf seas can be divided into regions which have tidally mixed waters and thermally stratified waters. The tidally mixed near shore environments support zooplankton communities dominated by smaller copepods and having large meroplankton contributions. These small copepods (Centropages spp., Temora spp., Acartia spp., Paral Pseudo/Microcalanus spp.) together with the microzooplankton component form a different and more complex food web than the larger copepod/diatom link associated with thermally stratified waters. The copepods Calanus finmarchicus and C. helgolandicus account for over 90% of the copepod dry weight biomass in stratified waters. Although occurring in lower numbers in mixed waters they can still make significant contributions to the biomass. A 31 year time series from the European shelf shows the inter- and intea-annual variability of these species. The basic biology and food web that these two systems support, and the transfer of energy, can result in marked differences in quantity and quality of particulates available as food for fish larvae. Calanus dominated systems allow the primary production to be directed straight through the trophic food chain (diatoms/Calanus/fish larvae) while the near shore communities of smaller copepods limit the amount of energy being transferred to the higher trophic levels. Eighty-two Longhurst Hardy Plankton Recorder hauls were used as the data base for this study. In all cases the zooplankton was dominated by copepods both in numbers and biomass accounting for > 80% of total zooplankton dry weight in the Irish Sea, Celtic Sea, shelf edge of the Celtic Sea and the northern and southern North Sea in Spring.     

Protein polymorphisms in six species of the genus Calanus

Isoelectric focusing was used to study total protein patterns and allozyme variations of six species of the genus Calanus. Each species could be characterized by total protein patterns. The results of the allozyme study indicated, in agreement with previous morphological studies, that the six Calanus species belong to three different groups: the C. finmarchicus group C. finmarchicus, C. glacialis and C. marshallae the C. helgolandicus group (C. helgolandicus, C. pacificus), and C. hyperboreus, which stands apart. There is no indication that there are more loci coding for the proteins studied in species with larger genome sizes. Nor is the degree of enzyme polymorphism related to genome size in these species.     

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.     

Seasonal changes in sedimentation of particulate matter and lipid content of zooplankton collected by sediment trap in the Arctic Ocean off Axel Heiberg Island

Summary Sedimentation of particulate matter measured at a depth of 100 m varied from 0.2 to 6.0 mg dry weight m^-2 d^-1 over approximately monthly intervals between September 1986 and June 1987 on the continental shelf of the western Arctic Ocean north of the Canadian Arctic Islands. Detritus deposited during December and January, the period of maximum sedimentation, contained relatively low amounts of organic carbon and nitrogen (4.0% and 0.5% of dry weight, respectively) with a carbon: nitrogen ratio > 7. Maximum rates of organic carbon and nitrogen sedimentation (0.60 and 0.12 mg m^-2 d^-1, respectively) during February were associated with debris enriched with organic matter (organic carbon and nitrogen content of 17.0% and 3.4% of dry weight, C:N=5). Dry weight of macrozooplankton, predominantly Calanus hyperboreus and Metridia longa, separated from preserved trap contents, equalled or exceeded by up to an order-of-magnitude the weight of sedimented debris. Lipid content of CV and adult female stages of C. hyperboreus decreased from 46%–64% of dry weight in September to 20%–30% in January. Females of C. hyperboreus with lower lipid levels (7% of dry weight) between April and June were probably in a post-reproductive condition.     

Zooplankton communities of the Arctic’s Canada Basin: the contribution by smaller taxa

Zooplankton was sampled at ten stations in the Canada Basin during August 2002 using both 53- and 236-m mesh nets to examine the contribution by smaller and less studied species. Copepod nauplii, the copepods Oithona similis, Oncaea borealis and Microcalanus pygmaeus, and the larvacean Fritillaria borealis typica dominated the upper 100 m of the water column numerically, while biomass was dominated by the copepods Calanus hyperboreus, Calanus glacialis and Paraeuchaeta glacialis, and the chaetognath Eukrohnia hamata. Zooplankton biomass ranged from 3.7 to 14.5 mg AFDW m^–3, with a mean of 9.6 mg AFDW m^–3 . While the three microcopepods contributed less than 5% of the biomass, estimates of their potential growth rates suggest they might contribute upwards of 25% of the metazoan zooplankton production. The true rates of growth and development of these microcopepods in the Arctic need to be determined to conclusively ascertain their importance.     

Feeding, respiration and life history of the hyperiid amphipod Themisto libellula in the Arctic marginal ice zone of the Greenland Sea

Daily ingestion rates of the pelagic hyperiid amphipod Themisto libellula were studied in the marginal ice zone of the Arctic Fram Strait by feeding experiments, respiration measurements and an allometric approach based on body mass. Amphipods were collected by stratified multiple opening/closing net hauls and Rectangular Midwater Trawl (RMT 8) in August 2000 during the expedition ARK XVI/2 of R/V “Polarstern”. T. libellula occurred with abundances of 0.043 and 0.015 ind. m⁻³ in the upper 30 m of the water column at two RMT 8 stations. Based on respiration data, the daily ingestion necessary to cover metabolic energy demands measured 1.9±0.6% of body carbon per day. Actual prey consumption during feeding experiments with Calanus copepodids as prey was very similar and accounted for 1.9±1.5% day⁻¹, indicating that feeding on Calanus can meet the energy demands of T. libellula. In general, experimental results were slightly lower than the maximum potential ingestion (2% day⁻¹ for an individual of median body dry mass of 32 mg) estimated by an allometric equation based on body mass, but feeding experiments showed a strong variability. Reduced metabolism and low ingestion rates of T. libellula are consistent with low ambient temperature, large body size, slow growth and long life span of this polar species. The effect of the active pelagic life style of T. libellula on metabolism and ingestion rate is discussed in comparison to the sympagic (i.e. ice-associated) amphipod Gammarus wilkitzkii of similar body size living in the same environment. In relation to the mesozooplankton biomass in the investigation area, the predation impact by T. libellula was low. However, high-Arctic conditions also limit the secondary production of principal prey species, such as Calanus glacialis and Calanus hyperboreus, so that even low predation rates may affect the growth of prey populations.     

Vertical distribution of zooplankton biomass and species in northeastern Baffin Bay related to temperature and salinity

Summary Vertical distributions of various species and stages of zooplankton at different times of the day were determined by stratified sampling with the BIONESS in northeastern Baffin Bay during early August. The water column was divided into an upper subarctic zone (>0° C, salinity < 32), a lower subarctic zone (< 0° C, salinity 32 to 34) and a deep zone of Atlantic water (>0° C, salinity 34). The upper subarctic zone was dominated by two species of pteropod molluscs; the lower subarctic water was dominated by the copepods, Calanus finmarchicus, C. glacialis and C. hyperboreus whose copepodite stages showed depth distributions that were different from one another, with the copepodite stage 5 and adult females generally shallower than the younger stages. All stages of all Calanus species were in the zone of primary production (10 to 50 m) while the copepods Pseudocalanus, Metridia and Oithona were generally found below this zone. Only C. finmarchicus and C. glacialis showed evidence of diurnal migration, migrating to the surface waters when the sun was at its lowest position on the horizon (i.e. at 0100 h).     

Food and growth of arctic char, Salvelinus alpinus (L.), in the Cumberland Sound area of Baffin Island

Arctic char, Salvelinus alpinus (L.), captured during the summer, 1972, in Cumberland Sound, Baffin Island, fed mainly on planktonic amphipods (Parathemisto libellula, Pseudalibrotus glacialis), copepods (Calanus hyperboreus), and fish (Boreogadus saida, Myoxocephalus sp.), but failed to utilize coelenterates, planktonic gastropods, and epi- and infauna. The considerable variation in the species composition of the diet of char of different lengths was due primarily to size selection. The average length of all individuals in stomach contents and of representatives of most food species increased with the length of fish. The minimum and maximum length of frequently ingested organisms increased four and 90 times, respectively, as char increased from 4 to 85 cm. Char less than 10 cm in length captured in rivers tributary to the Sound fed mainly on larval Chironomidae (Eukiefferiella bavaria) during the summer, whereas those longer than 10 cm fed predominantly on other char. During the winter the stomachs of the smaller individuals were always empty while the diet of the large char was restricted to other fish. Most species available to the char were consumed in proportion to their relative abundance in the rivers. The dry weight of stomach contents, when expressed on a unit weight basis, decreased with the wet weight of char in both fresh and salt water. Thus, fish 10 g in weight contained approximately 1.5 times more food in their stomachs than those weighing 1000 g. The stomach contents of char captured in salt water weighed approximately 11 times more than those of char of comparable size captured simultaneously in fresh water. The fish fed at random intervals during the day and ceased feeding at night. Arctic char, at all sampling areas, had reached a length of approximately 9.6 cm after four years. Upon migrating to salt water, their growth rate increased sharply with the result that after eight years they were 26.5 cm in length, reflecting food availability. The growth rate gradually decreased in fish older than nine years so that 20 year olds were approximately 70.0 cm in length.