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.     

Evidence that size-selective mortality affects growth of Atlantic cod (Gadus morhua L.) in the southern Gulf of St Lawrence

The effects of size-selective fishing mortality on the growth of Atlantic cod ( Gadus morhuu L.) in the southern Gulf of St Lawrence were investigated and compared between: (1) a period when fishing mortality was relatively high, growth was relatively rapid, and abundance low (1967–1972 year classes): and (2) a period when fishing mortality was lower, growth was slow, and density high (1977–1982 year classes). Cod first entered the fishery at age 3 during both periods. The 1967–1972 year classes (fast growing) were fully recruited to the fishery by age 5 or 6 and the fishery removed over twice as many fish from the lower than upper quartiles of length-at-age distributions for cod 4 to 10 years old (disproportionately high exploitation of slow-growing fish). In contrast. the 1977–1982 year classes (slow growing) did not fully recruit to the fishery until age 9 or 10 and the fishery removed four times as many fish from the upper than lower quartiles of the length-at-age distributions for 4- to 10-year-old cod (disproportionately high exploitation of fast-growing fish). The reduced mean lengths-at-age of the 1977–1982 year classes compared with the 1967–I972 year-classes is consistent with the different patterns of exploitation of fast- and slow-growing fish for the two periods.     

Early Environmental Influences on Growth of Arcto-Norwegian Cod, Gadus Morhua, From The 0-group To Adults

A high growth rate for Arcto-Norwegian cod, Gadus morhua, in the Barents Sea and adjacent areas from the larva period to the 0-group enhances survival and ultimately recruitment to the fishery. However, it appeared that high growth rates for a cohort through the 0-group were not continued as the cohort ages. Based on survey data, there was a significant negative correlation between the average length at the 0-group and its average length at ages 2 through 8. We provided evidence suggesting that this phenomenon was caused by the inter-annual variability in inflow of warm, prey-rich Atlantic water into the Barents Sea from the Norwegian Sea. Enhanced inflow provided favorable conditions for cod growth during the larva and juvenile pelagic intervals. However, this same strong inflow carried a proportion of the cohort farther to the east in the Barents Sea, where the bottom water is colder than in the west. The colder conditions experienced by such cohorts, as compared to cohorts that have a more westerly settlement, led to slower growth prior to age 2. Slow growth during this interval appeared to be the reason for these cohorts' relatively smaller mean length at older ages.     

Estimating the Barents Sea polar bear subpopulation size

A large-scale survey was conducted in August 2004 to estimate the size of the Barents Sea polar bear subpopulation. We combined helicopter line transect distance sampling (DS) surveys in most of the survey area with total counts in small areas not suitable for DS. Due to weather constraints we failed to survey some of the areas originally planned to be covered by DS. For those, abundance was estimated using a ratio estimator, in which the auxiliary variable was the number of satellite telemetry fixes (in previous years). We estimated that the Barents Sea subpopulation had approximately 2,650 (95% CI approximately 1,900–3,600) bears. Given current intense interest in polar bear management due to the potentially disastrous effects of climate change, it is surprising that many subpopulation sizes are still unknown. We show here that line transect sampling is a promising method for addressing the need for abundance estimates.     

Regional and local spatial genetic structure of Siberian primrose populations in Northern Europe

We have investigated the local and regional scale genetic structure of Siberian primrose (Primula nutans) populations in Northern Europe. The genetic diversity and structure of fifteen populations sampled from the Bothnian Bay in Finland, the Barents Sea in Norway and the White Sea in Russia were assessed using eleven microsatellite markers. We investigated the distribution of genetic variation within and between populations, and studied the local genetic structure using spatial autocorrelation analysis. We found very low genetic and allelic diversity in the Bothnian Bay and Barents Sea populations, and only slightly higher in the White Sea population. The level of genetic differentiation between the regions was very high, whereas differentiation between the populations within the regions was moderate. We found no spatial structuring of populations in any region suggesting efficient dispersal on a local scale. Clonal reproduction seemed to have no effect on genetic structure.     

The recent population expansion of boarfish,Capros aper (Linnaeus, 1758): interactions of climate,growth and recruitment

The objectives of this study were to evaluate whether temperature changes in the Northeast Atlantic influence the growth and recruitment dynamics of boarfish, Capros aper. Two geographically separate areas were examined, ‘north’ at the northern distribution range west of Ireland and ‘south’ on the main fishing grounds south of Ireland. No significant differences in length‐at‐age were observed between the two areas. Interannual otolith growth patterns were similar between the two areas with distinct years of faster and slower growth. In the ‘north’, no significant relationship between adult growth and temperature was observed, while growth in the ‘south’ was positively related to temperature up to approximately 16°C growth rates were suppressed in the years with temperatures above that. Recruitment showed a positive correlation with adult growth the previous year for the Spanish recruitment index only, suggesting spatial connectivity between the Celtic Sea and the Bay of Biscay. The age distributions were similar in both areas and despite the boarfish's longevity of >30 years, are dominated by the age classes corresponding to the years with high recruitment, suggesting that increased recruitment is responsible for the observed stock expansion.     

Specific features of the spatial and temporal distribution of the White Sea population of harp seal

Analysis of the data obtained during aerial survey of sea mammals in the 1980s–1990s (White and Barents seas) has revealed that, during all the seasons of the year, harp seals of the White Sea population form two types of aggregations which differ in size and some ecological features. Large herds of seals (tens of thousands of individuals) are able to occupy areas extending tens and even hundreds of kilometers. Harp seals prefer to inhabit the northern part of the population range (Barents Sea), whereas the southern part (White Sea) is mainly used for reproduction and molting. Small herds (several hundred animals) can be scattered over vast territories, but they tend to dwell in the southern part of the area including the White Sea and southern areas of the Barents Sea. The opportunity to distinguish between these two types of aggregations makes it possible to study the biological features of each of them and to specify characteristics of the species biology.     

Long-term changes in Krill biomass and distribution in the Barents Sea: are the changes mainly related to capelin stock size and temperature conditions?

Krill plays a significant role in the Barents Sea ecosystem, providing energy transport between different trophic levels. The current paper presents the results of a long-term study (1980–2009) based on pelagic trawl catches from August to September. Our investigations show that the krill species were distributed widely in the Barents Sea and that the largest krill concentrations were restricted to the west-central and eastern parts of the Barents Sea. The current paper presents the relative biomass indices, and the estimates must be interpreted as minimum biomass. The mean annual krill biomass was estimated to be 22 million tonnes in wet weight, with the highest values being as much as 48 million tonnes. Capelin is the largest pelagic stock, and in some years, their biomass can amount to 4–7 million tonnes, which can impose high predation pressure on krill. When their biomass is high, capelin may consume close to 26 million tonnes annually. The predation from pelagic (herring and blue whiting) and bottom (cod and haddock) fish species was much lower, being 9 and 1 million tonnes, respectively. A negative relationship between krill biomass and capelin stock size above 74°N was observed during the study period. However, during the last decade, the krill biomass has increased despite heavy predation from capelin in some years. A positive significant linear relationship between the mean annual Kola temperature and the krill biomass seems to indicate that the recent warming conditions have favourable impacts on the krill populations in the Barents Sea.     

Late Pleistocene and Holocene distribution of Mytilus edulis in the Barents Sea region and its palaeoclimatic implications

Aim For decades, subfossil shells of the bivalve Mytilus edulis Linnaeus, 1758 in Svalbard have been taken as evidence of higher surface temperatures during the early Holocene because the modern northern occurrence of this mollusc was, until recently, in the southern Barents Sea. Here, we elucidate and discuss the spatial and temporal Late Pleistocene and Holocene distribution of the species within the entire Barents Sea region. Location The Barents Sea region. Methods Radiocarbon dates of Mytilus shells from the Barents Sea region and information about the present distribution of the species were compiled, including two new radiocarbon dates from north‐eastern Spitsbergen. The dataset was divided into time slices, each covering 1000 years, and compared with Holocene temperature variations, ocean current systems and present‐day temperature patterns. Results Maps show the Late Pleistocene and Holocene spatial and temporal distribution of Mytilus edulis in the Barents Sea region. M. edulis was already present in northern Norway about 14,000 cal. yr bp . It appeared at western Spitsbergen about 11,000 cal. yr bp , and slowly spread to the rest of the archipelago. The maximum distribution in the region was reached 10,000–7000 cal. yr bp , coinciding with the Holocene climatic optimum. The species gradually disappeared in the late Holocene and became absent from the northern and eastern parts of the region 3000–1000 cal. yr bp . Today, M. edulis lives in the southern part and has begun to recolonize the northern parts. Main conclusions The time slices illustrate strong connections between the ocean current regimes, the climate and the distribution of M. edulis. The species settled in the southern part of the Barents Sea region several thousand years before it spread to the northern part during the Holocene climatic optimum. It may even have been completely absent from the region for a short time during the late Holocene cold period. The Holocene distribution of Mytilus implies that the underlying pattern of coastal sea surface temperatures in the region was very stable.     

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.     

Interannual fluctuations of zooplankton in the Kola Section (Barents Sea) in relation to environmental factors

An analysis of interannual variations of zooplankton composition and biomass in the Kola Section (Barents Sea) in summer was conducted based on the data of 2003–2010. Maximum values of the mean water temperature and temperature anomaly were found in 2006 and in 2007. Variations in the zooplankton composition and relative biomass of common species were studied in relation to climatic factors. It is discussed which parameters may be used as indicators of climatic changes in the southern Barents Sea.     

Cod, Gadus morhua L., populations identified by mitochondrial DNA

Isozyme and haemoglobin analysis has shown that the cod, Gadus morhua L., in the NE Atlantic can be regarded as two populations: Arctic and coastal cod. One hundred and one individual cod from nine different locations were sampled and restriction fragment analysis uncovered 14 different mtDNA clones. Calculation of sequence divergence between localities displayed large divergence between samples from coastal and Arctic areas (l.77–5.62%), as opposed to the low intrapopulation divergence (∼0.10%) and the divergence between localities along the coast (∼0.17%) and in the Barents Sea (∼ 1.00%).     

Feeding of Greenland halibut <Emphasis Type="Italic">Reinhardtius hippoglossoides</Emphasis> (Pleuronectidae) in the Kara Sea

Feeding of Greenland halibut Reinhardtius hippoglossoides in the northern Kara Sea was studied based on data collected in summer–autumn 2007–2013. The main food of all size groups of halibut were fish—up to 98% of weight of the food bolus. Larger individuals had lower intensity of feeding as compared to juveniles, which was probably owing to the lack of suitable food for large fish and, along with gonad maturation process, could be one of the reasons of their migration to the Barents Sea. The northern part of the Kara Sea, as well as the adjacent areas of Barents Sea, can be considered as an important area of habitation of juvenile Greenland halibut of the Norwegian–Barents Sea population.     

Egg Size and Formation of the Geographical Range in <Emphasis Type="Italic">Thysanoessa inermis</Emphasis> (Kroyer, 1846) (Crustacea: Euphausiacea) in the Norwegian and Barents Seas

Using materials on the euphausiacean Thysanoessa inermis collected in the Norwegian and Barents seas (from 60°N to 73°N) we studied the variability of size characteristics in the eggs (diameter of embryos and egg capsules and the width of the perivitelline space) and their correlations with water temperature and salinity. The size of embryos showed almost no variability irrespective of the location of the sampling site and the water temperature and salinity; the perivitelline space performs a protective function, and its width showed a tendency to decrease with increasing water temperature and salinity. The similarities in the size of embryos in T. inermis populations thousands of kilometers away from each other might be explained, first, by the relatively small age of the populations (the age of the population from the Barents Sea did not exceed 6000–7000 years) and second, by movement of the crustaceans with currents from the Norwegian Sea to the Barents Sea.     

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).     

Food‐web structure varies along environmental gradients in a high‐latitude marine ecosystem

Large‐scale patterns in species diversity and community composition are associated with environmental gradients, but the implications of these patterns for food‐web structure are still unclear. Here, we investigated how spatial patterns in food‐web structure are associated with environmental gradients in the Barents Sea, a highly productive shelf sea of the Arctic Ocean. We compared food webs from 25 subregions in the Barents Sea and examined spatial correlations among food‐web metrics, and between metrics and spatial variability in seawater temperature, bottom depth and number of days with ice cover. Several food‐web metrics were positively associated with seawater temperature: connectance, level of omnivory, clustering, cannibalism, and high variability in generalism, while other food‐web metrics such as modularity and vulnerability were positively associated with sea ice and negatively with temperature. Food‐web metrics positively associated with habitat heterogeneity were: number of species, link density, omnivory, path length, and trophic level. This finding suggests that habitat heterogeneity promotes food‐web complexity in terms of number of species and link density. Our analyses reveal that spatial variation in food‐web structure along the environmental gradients is partly related to species turnover. However, the higher interaction turnover compared to species turnover along these gradients indicates a consistent modification of food‐web structure, implying that interacting species may co‐vary in space. In conclusion, our study shows how environmental heterogeneity, via environmental filtering, influences not only turnover in species composition, but also the structure of food webs over large spatial scales.     

Biology of the sand-smelt (Atherina presbyter Valenciennes) around Fawley power station

Past studies of fish impingement at Fawley power station have shown that sand-smelts, Atherina presbyter Valenciennes, are susceptible to impingement and may be a useful species for examining effects of impingement mortalities on population size and structure. A 21 month study of populations in the vicinity of Fawley power station was carried out to obtain data on the biology of the species for future population dynamics modelling studies. Aspects included in the study were growth, sex ratio, reproduction, survival and mortality rates and diet. Differences in apparent and true rates of growth indicated some size-selective mortality towards younger age groups, possibly caused by the power station. A seasonallyoscillating von Bertalanffy growth equation is given to predict mean length-at-age. Most female sand-smelts (73%) matured at the end of their first year of life, fecundities ranging on average from 1394 eggs per female for 1+ group fish to 6872 for III+ fish. The sex ratio was even (1 : 1). An annual survival rate of 10–8% was estimated for mature fish, and the consequent age structure of the population indicates that the bulk of egg production lies with the smaller (1+ group) fish. Since smaller fish are generally more vulnerable to impingement this factor may render populations sensitive to impingement mortalities. Gut content analysis indicated that sand-smelts probably actively select zooplankters from the general plankton when young and, once larger than 104 mm in length, increasingly feed upon natant macrofaunal species.     

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.     

Relationships between smolt size, postsmolt growth and sea age at maturity in Atlantic salmon ranched in the Baltic Sea

Tagging data were used to examine the relationships between smolt size, post-smolt growth and sea age at first maturity for the short-migrating Neva strain of Atlantic salmon ( Salmo salar L.) ranched in the Bothnian Sea and the Gulf of Finland. The results provided evidence that post-smolt growth was influenced by both relative and absolute smolt size. For both sea-areas, 2-year smolts of small relative size within a release group grew more rapidly in the sea than did smolts of higher relative, but equivalent absolute size. The negative influence of increasing relative smolt size on marine growth was, however, outweighed by the stronger positive influence of increasing absolute smolt size. A 160-mm increase in smolt size (140–300 mm) resulted in an overall growth advantage of about 1 year. In the Bothnian Sea, the predicted mean length after 1 year in the sea was 288 ± 25 mm for 140-mm smolts and 560 ± 16 mm for 300-mm smolts. Under the more favourable conditions of the Gulf of Finland, the respective mean lengths were 369 ± 15 mm and 613 ± 12 mm. The sea age at first maturity was inversely related to both freshwater and marine growth rates. For both sea areas, large smolts yielded proportionately more grilse than did small ones. Smolt years with good post-smolt growth rates yielded more grilse than did years with poor growth rates. The overall level of grilsing was higher in the Gulf of Finland than in the Bothnian Sea. These results suggest that the relationships between smolt size, post-smolt growth and age at first maturity in the sea are influenced by the environmental conditions of the respective sea area. A framework explaining the links between smolt size, marine growth, survival and sea age at maturity in Neva salmon is presented for the Gulf of Finland and the Bothnian Sea.     

The distribution and biology of the cestode Eubothrium parvum in capelin, Mallotus villosus, (Pallas) in the Barents Sea, and its use as a biological tag

Samples of Eubothrium parvum were obtained from capelin Mallotus villosus at 55 stations throughout the Barents Sea and from Balsfjord, North Norway. The parasite is distributed widely throughout the Barents Sea, but both incidence and intensity of infection are higher in the regions off Murmansk and the Kola peninsula, and Spitsbergen. E. parvum exhibits a seasonal peak in maturation and probably also in acquisition of new infections. The incidence of infection is greatest in 1 + fish, whereas the intensity is more independent of host age. It is suggested that the parasite requires only a single intermediate host, a plank-tonic copepod, and its distribution in relation to age of host is a reflection of the dietary preference shown by young capelin for copepods. The frequency distribution of E. parvum in capelin was over-dispersed in Balsfjord, where infection levels of between 1 and 28 parasites per fish were encountered in all samples, but under-dispersed in the Barents Sea, where infections of more than four parasites per fish were never found and even infections with three and four parasites were very local. It is suggested that the underdispersion is due to a very low probability of infection in the open waters of the sea. Although the presence of E. parvum cannot be used as a biological tag for capelin, its abundance and frequency distribution can. The difference in frequency distribution and the failure to find any heavily infected fish in the Barents Sea confirm the suggestion that the capelin of Balsfjord form a local isolated population, which does not migrate into the Barents Sea. The differences in infection levels within the Barents Sea suggest the further possibility that there are at least two stocks of capelin there, but this requires further investigation and confirmation.