Inter-specific variations in distribution, abundance and possible life-cycle patterns of Themisto spp. (Amphipoda) in the Barents Sea

Amphipods from the central and northern areas of the Barents Sea were studied by a series of MOCNESS profiles between 1990 and 1997. Themisto abyssorum, though dominant in warmer Atlantic water, was also present in Arctic water. In contrast, T. libellula was a typical Arctic species and penetrated very little into the Atlantic water masses. T. compressa was seldom found in the studied area. Linear regression analysis showed a statistically significant negative relationship between abundance of T. libellula and the variability in the Atlantic inflow. The abundance of this species seems to be, to a large extent, determined by the amount of Arctic water in the Barents Sea. The sub-Arctic species, T. abyssorum, only has a 1-year life-cycle, with the peak in release of young (2-3 mm) occurring in May and June. A few individuals may survive to be older. The Arctic species, T. libellula, seems to release the young earlier and the length-frequency distributions seem to indicate a 2-year life-span. The spring phytoplankton, which usually blooms during April in the Barents Sea, followed by high abundance of Calanus spp. and krill species, are regarded as important factors that influence the release of the amphipods' young.     

Biomass changes and trophic amplification of plankton in a warmer ocean

Ocean warming can modify the ecophysiology and distribution of marine organisms, and relationships between species, with nonlinear interactions between ecosystem components potentially resulting in trophic amplification. Trophic amplification (or attenuation) describe the propagation of a hydroclimatic signal up the food web, causing magnification (or depression) of biomass values along one or more trophic pathways. We have employed 3‐D coupled physical‐biogeochemical models to explore ecosystem responses to climate change with a focus on trophic amplification. The response of phytoplankton and zooplankton to global climate‐change projections, carried out with the IPSL Earth System Model by the end of the century, is analysed at global and regional basis, including European seas (NE Atlantic, Barents Sea, Baltic Sea, Black Sea, Bay of Biscay, Adriatic Sea, Aegean Sea) and the Eastern Boundary Upwelling System (Benguela). Results indicate that globally and in Atlantic Margin and North Sea, increased ocean stratification causes primary production and zooplankton biomass to decrease in response to a warming climate, whilst in the Barents, Baltic and Black Seas, primary production and zooplankton biomass increase. Projected warming characterized by an increase in sea surface temperature of 2.29 ± 0.05 °C leads to a reduction in zooplankton and phytoplankton biomasses of 11% and 6%, respectively. This suggests negative amplification of climate driven modifications of trophic level biomass through bottom‐up control, leading to a reduced capacity of oceans to regulate climate through the biological carbon pump. Simulations suggest negative amplification is the dominant response across 47% of the ocean surface and prevails in the tropical oceans; whilst positive trophic amplification prevails in the Arctic and Antarctic oceans. Trophic attenuation is projected in temperate seas. Uncertainties in ocean plankton projections, associated to the use of single global and regional models, imply the need for caution when extending these considerations into higher trophic levels.     

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.     

Seasonal changes in zooplankton composition in the Barents Sea, with special attention to Calanus spp. (Copepoda)

The Barents Sea is an important area with respect to fisheriesresources (i.e. capelin and cod). In May, June and August 1981zooplankton biomass was measured along a transect at 30°E,from the ice border southwards. A maximum was recorded in Atlanticwater by the end of June (>100 g wet weight m^–2 InAugust the biomass values were relatively low south of the Polarfront and increased northwards into Arctic water (–50g m^–2 The species composition was influenced by the distributionof cold Arctic water and warmer Atlantic water. The zooplanktonwas dominated by the copepods Calanus finmarchicus and C. glacialis;the former is regarded as an Atlantic species and C. glacialisas an Arctic species.     

Climate and cyclic hydrobiological changes of the Barents Sea from the twentieth to twenty-first centuries

The Barents Sea is a transition zone between North Atlantic and Arctic waters, so its marine ecosystem is highly sensitive to climate dynamics. Understanding of marine biota response to climate changes is necessary to assess the environmental stability and the state of marketable biological resources. These processes are analyzed using a database from the Murmansk Marine Biological Institute which holds oceanographic and hydrobiological data sets collected for more than 100?years along the meridional Kola Transect in the Barents Sea. The data demonstrate high variability in thermal state of the upper layer of the Barents Sea, which is regulated by varying the inflow of Atlantic water and by regional climate. At irregular intervals, cold periods with extended seasonal ice cover are followed by warm periods. The most recent warm period started in the late 1980s and reached its maximum from 2001 to 2006. These cyclic changes in hydrologic regime across the twentieth century and first decade of the twenty-first century are reflected (with a specific lag of 1–5?years) by changes in species composition, as well as abundance and distribution of boreal and arctic groups of macrozoobenthos and fish fauna. For instance, cod and cod fisheries in the Barents Sea are closely linked to the marine climate. Furthermore, Kamchatka crab stock recruitment benefited from the warm climate of 1989 and 1990. In general, studies in this region have shown that climatic dynamics may be assessed using biological indices of abundance, biomass, and migration of marine organisms, including commercial species.     

Comparison of productivity and phytoplankton in a warm (Kongsfjorden) and a cold (Hornsund) Spitsbergen fjord in mid-summer 2002

Kongsfjorden and Hornsund are two glacial fjords without sills on the West Spitsbergen coast. Both sites are under the influence of relatively warm Atlantic-derived water, although Hornsund is more influenced by cold water from the Barents Sea. In this study, we compared the impacts of cold Arctic and warmer Atlantic waters on the pelagic ecosystems of Kongsfjorden and Hornsund. Both fjords were strongly influenced by Atlantic-derived waters during summer (2002). Diatoms were the most substantial contributors to phytoplankton biomass, especially in outer basins of both fjords, whereas the second most important contributors were autotrophic dinoflagellates in Kongsfjorden and nanoflagellates in Hornsund. Total phytoplankton biomass was highest in Hornsund. Primary production rates were an order of magnitude lower in Kongsfjorden than in Hornsund, and increased from inner to outer fjord (from 2.47 to 4.48 mg C m⁻² h⁻¹ in Kongsfjorden and from 14.00 to 86.65 mg C m⁻² h⁻¹ in Hornsund). Chlorophyll-a concentration was also substantially lower in Kongsfjorden. Zooplankton was dominated by omnivorous species in Kongsfjorden and herbivorous in Hornsund. Observed differences between the fjords may originate from (1) advection of different waters into the fjords; (2) differences in freshwater runoff and turbidity, and (3) timing of the phytoplankton bloom. Climate warming will likely increase the Atlantic water influence, and result in reduced production of diatoms and increase in flagellates.     

Distribution and diet of 0-group cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) and haddock (<Emphasis Type="Italic">Melanogrammus aeglefinus</Emphasis>) in the Barents Sea in relation to food availability and temperature

Distribution of 0-group cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) in August–September 2005 and 2006 was mainly restricted to the Atlantic waters of the western and central areas of the Barents Sea. The main distribution of 0-group fish overlapped largely with areas of high biomass (>7 gm⁻² dry weight) of zooplankton. The copepod Calanus finmarchicus and krill Thysanoessa inermis, which are dominant zooplankton species in both Atlantic and boreal waters of the Barents Sea, were the main prey of 0-group cod and haddock. The main distribution, feeding areas and prey of 0-group cod and haddock overlapped, implying that competition for food may occur between the two species. However, though their diet coincided to a certain degree, haddock seems to prefer smaller and less mobile prey, such as Limacina and appendicularians. As 0-group fish increased in size, there seems to be a shift in diet, from small copepods and towards larger prey such as krill and fish. Overall, a largely pelagic feeding behaviour of 0-group cod and haddock was evident from this study.     

Estimated copepod production rate and structure of mesozooplankton communities in the coastal Barents Sea during summer–autumn 2007

The southern Barents Sea is considered to be the most productive area in the Arctic Ocean; however, there are no assessments of daily production rates in the coastal waters. During the summer and autumn of 2007, we investigated the variation of mesozooplankton community structure relative to environmental conditions at 12 coastal stations. Copepods dominated the total zooplankton biomass and abundance during both periods. Diversity indices and the total biomass of zooplankton communities differed significantly between the two seasons. Cluster analyses revealed two distinct groups of stations which were associated with Ura Bay and the adjacent open sea, respectively. Daily production rates of the copepod species examined were calculated using three methods based on: (1) a temperature-dependent equation and (2) two multiple regressions that consider temperature, body weight, and chlorophyll a concentration. Significant seasonal differences for daily production rates were found using all three model equations (p?<?0.05): 358?±?188–1,775?±?791 versus 198?±?85–1,584?±?559?μg?dry?mass?m^?3?day^?1. Results of principal components analyses demonstrated that the abundance and biomass of herbivorous species were related to variation in chlorophyll a concentration while the abundance and biomass of other species (omnivorous copepods and Ctenophora) were related mainly with water temperature and salinity. Mesozooplankton biomass was higher during this study relative to previous studies. Computed copepod production rates were higher compared with other Arctic seas confirming a high productive potential of the coastal southern Barents Sea.     

Distribution and seasonal dynamics of <Emphasis Type="Italic">Boroecia maxima</Emphasis> (Ostracoda: Halocypridinae) in the Arctic basin and adjacent Atlantic waters

Based on materials obtained during Soviet and Russian expeditions to the Arctic basin and adjacent Atlantic waters during the period from 1929–1993, as well as data from the literature, we examined the distribution range of the common deep-sea ostracod Boroecia maxima. This species occurs throughout the pelagic water column (down to 3000 m depth and below) and prefers low water temperatures (surface Arctic and intermediate water masses). During the year, at all depths, the ratio between age groups of B. maxima remains virtually unchanged. Breeding apparently continues year-round. During the polar night, B. maxima makes no migration into warm Atlantic water, where wintering predatory zooplankton occur in large numbers, and thus avoids grazing pressure.     

Dynamics of biomass of Calanus finmarchicus and zooplankton in the coastal zone of the Barents Sea under various thermal regimes

The dynamics of zooplankton biomass and the biomass of the key copepod species Calanus finmarchicus were studied in the southern Barents Sea. The effect of climatic factors, i.e., water temperature and the atmospheric indices, on the zooplankton was assessed. It was found that the biomass of the zooplankton correlated positively both with the water temperature and winter NAO index of the previous year. The phenomenon and its reasons are discussed.     

Qualitative assessment of the diet of European eel larvae in the Sargasso Sea resolved by DNA barcoding

European eels (Anguilla anguilla) undertake spawning migrations of more than 5000 km from continental Europe and North Africa to frontal zones in the Sargasso Sea. Subsequently, the larval offspring are advected by large-scale eastward ocean currents towards continental waters. However, the Sargasso Sea is oligotrophic, with generally low plankton biomass, and the feeding biology of eel larvae has so far remained a mystery, hampering understanding of this peculiar life history. DNA barcoding of gut contents of 61 genetically identified A. anguilla larvae caught in the Sargasso Sea showed that even the smallest larvae feed on a striking variety of plankton organisms, and that gelatinous zooplankton is of fundamental dietary importance. Hence, the specific plankton composition seems essential for eel larval feeding and growth, suggesting a linkage between eel survival and regional plankton productivity. These novel insights into the prey of Atlantic eels may furthermore facilitate eel larval rearing in aquaculture, which ultimately may replace the unsustainable use of wild-caught glass eels.     

Structure and distribution of pelagic ostracods (Ostracoda: Myodocopa) in the Arctic Ocean

The study of the pelagic ostracod fauna of the Arctic Ocean based on materials collected by numerous Russian expeditions (1929–1993) and data from the literature showed the extreme poorness of the Arctic pelagic ostracod fauna, its mainly North Atlantic genesis and complete isolation from the Pacific fauna. Maximum ostracod abundance was observed in the epipelagic zone, and the greatest species diversity occurred in the relatively warm deep Atlantic layer throughout the year. To the north, east, and west of Franz Josef Land and Spitsbergen, the number of species and abundance indices of pelagic ostracods were decreased. In superficial water layers of the Central Arctic, maximum ostracod density and biomass were recorded in June and September. The best bioindicator of warm Atlantic water in the Arctic basin is Obtusoecia obtusata; and of cold polar water in the North Atlantic, Boroecia maxima.     

Depth profiles of plankton,particulate organic matter and microbial activity in the eastern Canadian Arctic during summer

Summary Deep profiles of particulate organic matter, microplankton (phytoplankton and bacteria), zooplankton and their metabolic activities were investigated during two summer voyages to the eastern Canadian Arctic. Magnitudes and depth distributions were similar in many respects to observations from temperate and tropical waters. Strong gradients in most properties were observed in the upper 50–100 m and subsurface maxima were generally associated with the upper mixed-layer (>50 m). In addition to the general vertical decreases in plankton biomass and metabolic activity there was evidence for both rapid transport (sinking) of organic matter and for enhanced (above background) levels of microbial metabolic activity in deep waters (>500 m). Zooplankton depth distributions differed from the pattern generally observed at lower latitudes; in the Arctic, zooplankton abundance decreased to a lesser degree with depth than particulate organics and microplankton. The overwintering behavior of high-latitude zooplankton appeared to be the best explanation for their relatively high abundance at depth. Despite this, however, zooplankton apparently contributed little to the total column community metabolism.     

Meiobenthic stocks and benthic activity on the NE-Svalbard shelf and in the Nansen Basin

Summary High Arctic meiofaunal distribution, standing stock, sediment chemistry and benthic respiratory activity (determined by sediment oxygen consumption using a shipboard technique) were studied in summer 1980 on the NE Svalbard shelf (northern Barents Sea) and along a transect into the Nansen Basin, over a depth range of 240–3920 m. Particulate sediment proteins, carbohydrates and adenylates were measured as additional measures of benthic biomass. To estimate the sedimentation potential of primary organic matter, sediment bound chloroplastic pigments (chlorophylls, pheopigments) were assayed. Pigment concentrations were found comparable to values in sediments from the boreal and temperate N-Atlantic. Meiofauna, which was abundant on the shelf, decreased in numbers and biomasses with increasing depth, as did sediment proteins, carbohydrates, adenylates and sediment oxygen consumption. Meiofaunal abundances and biomasses within the Nansen Basin were comparable with those observed in abyssal sediments of the North Atlantic. Nematodes clearly dominated in metazoan meiofauna. Protozoans were abundant in shelf sediments. Probably in response to the sedimentation of the plankton bloom, meiofauna abundance and biomass as well as sediment proteins, carbohydrates and adenylates were significantly correlated to the amount of sediment bound chloroplastic pigments, stressing the importance of food quantity to determine benthic stocks. Ninety-four percent of the variance in sediment oxygen consumption were caused by chloroplastic pigments. Benthic respiration, calculated per unit biomass, was 3–10 times lower than in the East Atlantic, suggesting low turnover rates in combination with a high standing stocks for the high Arctic benthos.     

Ecological significance of 0-group fish in the Barents Sea ecosystem

Investigations into the 0-group fish in the Barents Sea have been carried out since 1965, with the goal of estimating the abundance of 0-group fish. 0-group abundance indices have been used in the assessment of the recruitment level and in recruitment variability studies. However, the ecological importance of the 0-group fish in the Barents Sea has been less studied. Although 0-group capelin, herring, cod and haddock are widely distributed in the Barents Sea, the central area seems to be the most important, accounting for approximately 50–80% of the annual biomass. The total biomass of the four most abundant 0-group fish species can be up to 3.3 million tonnes, with an average of 1.3 million tonnes (1993–2009). Wide distribution and high biomass of pelagically distributed 0-group fish make these fishes an important element in the energy transport between different trophic levels and different geographical areas, having a critical impact on the entire Barents Sea ecosystem. In recent years, capelin have shown a pronounced northward shift in biomass distribution, and several successive strong year classes occurred during warm temperature conditions. Cod biomasses were unexpectedly low during warm years and were positively correlated with spawning stock biomass, while the correlation with temperature was not significant. Haddock and herring show, as expected, increasing biomass with increased temperature when the spawning stock is at a sufficiently high level.     

Is the mass emergence of Themisto libellula in the northern Bering Sea an invasion or a bloom?

The mass occurrence of the large hyperiid Themisto libellula was recorded in both the western and the eastern Bering Sea within 2007–2011. Those were the years of a relatively long 6-year period of cold, which was caused mainly by the inflow of cold waters from the north; this is confirmed by the distribution of bottom and surface temperatures and also by the ice-cover values. This hyperiid became dominant in the diet of salmon, walleye pollock, herring, and several other nekton fish species. T. libellula periodically spreads southward with cold northern waters, finding favorable conditions in “new” areas. Being a rapidly growing species with a short life cycle, within 1 or 2 years it reaches a high abundance, which then gradually declines and remains at a mean or low level, as usually occurs with species that were introduced into a new habitat. After the environmental conditions deteriorate, as a “warm” period arrives with changes in the general circulation and a growing inflow of warmed Pacific waters, the southern boundary of the species range moves back far northward and it completely disappears in the areas where it prevailed in the plankton and was a main forage item in the diet of many fish species. Taking into account the durations of warm and cold periods from 1980 until 2010, an event like this in the Bering Sea can be expected within 1 or 2 years. In the eastern Bering Sea, the abundance and dominance of a number of zooplankton species may vary simultaneously. This effect is more pronounced in T. libellula and for this reason the species is considered as a biological indicator of the described climatic changes in the Bering Sea.     

Population status of Greenland halibut <Emphasis Type="Italic">Reinhardtius hippoglossoides</Emphasis> (Walbaum, 1793) of the Laptev Sea

This is the first study to perform a comparative genetic analysis of Greenland halibut in the samples from the Atlantic (waters of west and east of Greenland), Arctic (Laptev Sea), and Pacific (the western part of the Bering Sea) ocean basins using seven microsatellite loci. The obtained data clearly demonstrate that the Greenland halibut population in the Laptev Sea belongs to the groups of the Atlantic Ocean basin. Apparently, the Greenland halibut of the Laptev Sea is represented by a dependent population, which is replenished due to the drift of immatures from the spawning grounds in the Barents Sea with the transformed Atlantic water flow along the continental slope. In addition, the Arctic population can be partially replenished due to the breeding of the halibut in local spawning grounds.     

Demersal fish assemblages and spatial diversity patterns in the Arctic-Atlantic transition zone in the Barents Sea

Direct and indirect effects of global warming are expected to be pronounced and fast in the Arctic, impacting terrestrial, freshwater and marine ecosystems. The Barents Sea is a high latitude shelf Sea and a boundary area between arctic and boreal faunas. These faunas are likely to respond differently to changes in climate. In addition, the Barents Sea is highly impacted by fisheries and other human activities. This strong human presence places great demands on scientific investigation and advisory capacity. In order to identify basic community structures against which future climate related or other human induced changes could be evaluated, we analyzed species composition and diversity of demersal fish in the Barents Sea. We found six main assemblages that were separated along depth and temperature gradients. There are indications that climate driven changes have already taken place, since boreal species were found in large parts of the Barents Sea shelf, including also the northern Arctic area. When modelling diversity as a function of depth and temperature, we found that two of the assemblages in the eastern Barents Sea showed lower diversity than expected from their depth and temperature. This is probably caused by low habitat complexity and the distance to the pool of boreal species in the western Barents Sea. In contrast coastal assemblages in south western Barents Sea and along Novaya Zemlya archipelago in the Eastern Barents Sea can be described as diversity "hotspots"; the South-western area had high density of species, abundance and biomass, and here some species have their northern distribution limit, whereas the Novaya Zemlya area has unique fauna of Arctic, coastal demersal fish. (see Information S1 for abstract in Russian).     

Biomass of scyphozoan jellyfish, and its spatial association with 0-group fish in the Barents Sea

An 0-group fish survey is conducted annually in the Barents Sea in order to estimate fish population abundance. Data on jellyfish by-catch have been recorded since 1980, although this dataset has never been analysed. In recent years, however, the ecological importance of jellyfish medusae has become widely recognized. In this paper the biomass of jellyfish (medusae) in 0–60 m depths is calculated for the period 1980–2010. During this period the climate changed from cold to warm, and changes in zooplankton and fish distribution and abundance were observed. This paper discusses the less well known ecosystem component; jellyfish medusae within the Phylum Cnidaria, and their spatial and temporal variation. The long term average was ca. 9×10⁸ kg, with some years showing biomasses in excess of 5×10⁹ kg. The biomasses were low during 1980s, increased during 1990s, and were highest in early 2000s with a subsequent decline. The bulk of the jellyfish were observed in the central parts of the Barents Sea, which is a core area for most 0-group fishes. Jellyfish were associated with haddock in the western area, with haddock and herring in the central and coastal area, and with capelin in the northern area of the Barents Sea. The jellyfish were present in the temperature interval 1°C<T<10°C, with peak densities at ca. 5.5°C, and the greatest proportion of the jellyfish occurring between 4.0–7.0°C. It seems that the ongoing warming trend may be favourable for Barents Sea jellyfish medusae; however their biomass has showed a recent moderate decline during years with record high temperatures in the Barents Sea. Jellyfish are undoubtedly an important component of the Barents Sea ecosystem, and the data presented here represent the best summary of jellyfish biomass and distribution yet published for the region.     

Productivity in the Barents Sea - Response to Recent Climate Variability

The temporal and spatial dynamics of primary and secondary biomass/production in the Barents Sea since the late 1990s are examined using remote sensing data, observations and a coupled physical-biological model. Field observations of mesozooplankton biomass, and chlorophyll a data from transects (different seasons) and large-scale surveys (autumn) were used for validation of the remote sensing products and modeling results. The validation showed that satellite data are well suited to study temporal and spatial dynamics of chlorophyll a in the Barents Sea and that the model is an essential tool for secondary production estimates. Temperature, open water area, chlorophyll a, and zooplankton biomass show large interannual variations in the Barents Sea. The climatic variability is strongest in the northern and eastern parts. The moderate increase in net primary production evident in this study is likely an ecosystem response to changes in climate during the same period. Increased open water area and duration of open water season, which are related to elevated temperatures, appear to be the key drivers of the changes in annual net primary production that has occurred in the northern and eastern areas of this ecosystem. The temporal and spatial variability in zooplankton biomass appears to be controlled largely by predation pressure. In the southeastern Barents Sea, statistically significant linkages were observed between chlorophyll a and zooplankton biomass, as well as between net primary production and fish biomass, indicating bottom-up trophic interactions in this region.