Temporal Dynamics of Top Predators Interactions in the Barents Sea

The Barents Sea system is often depicted as a simple food web in terms of number of dominant feeding links. The most conspicuous feeding link is between the Northeast Arctic cod Gadus morhua, the world''s largest cod stock which is presently at a historical high level, and capelin Mallotus villosus. The system also holds diverse seabird and marine mammal communities. Previous diet studies may suggest that these top predators (cod, bird and sea mammals) compete for food particularly with respect to pelagic fish such as capelin and juvenile herring (Clupea harengus), and krill. In this paper we explored the diet of some Barents Sea top predators (cod, Black-legged kittiwake Rissa tridactyla, Common guillemot Uria aalge, and Minke whale Balaenoptera acutorostrata). We developed a GAM modelling approach to analyse the temporal variation diet composition within and between predators, to explore intra- and inter-specific interactions. The GAM models demonstrated that the seabird diet is temperature dependent while the diet of Minke whale and cod is prey dependent; Minke whale and cod diets depend on the abundance of herring and capelin, respectively. There was significant diet overlap between cod and Minke whale, and between kittiwake and guillemot. In general, the diet overlap between predators increased with changes in herring and krill abundances. The diet overlap models developed in this study may help to identify inter-specific interactions and their dynamics that potentially affect the stocks targeted by fisheries.     

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.     

Seasonal distribution and dive behaviour of harp seals (<Emphasis Type="Italic">Pagophilus groenlandicus</Emphasis>) of the White Sea–Barents Sea stock

Eight adult female harp seals (Pagophilus groenlandicus) of the White Sea–Barents Sea stock were tagged with satellite-linked dive recorders during the nursing period and followed from breeding in late February 1995 until moulting in late April 1995. Another ten adult harp seals of both sexes were tagged and followed from moult in early May 1996 until breeding in late February the following year. Between breeding and moult the seals were distributed along the coasts of Kola of Russia and eastern Finnmark of Norway, coinciding in time and space with the spawning capelin (Mallotus villosus). Between moulting and breeding they encircled the entire Barents Sea, mostly in open water, using the water column from 20 to 300 m, and in so doing by and large reflecting the annual migrations of the capelin. Capelin is therefore assumed to be the main source of prey for the White Sea–Barents Sea stock of harp seals, to be substituted, in part, by amphipods (e.g. Themisto libellula) in mid-summer and polar cod (Boreogadus saida) and herring (Clupea pallasii) in late autumn and winter. These data provide a baseline for the evaluation of the effects of future climatic change in the rich Barents Sea ecosystem.     

Fisheries management in practice: review of 13 commercially important fish stocks

This paper comparatively reviews several commercially important fish stocks, their state and their management in various regions of the world including Japanese anchovy, Bay of Biscay anchovy, North Sea sandeel, North Sea herring, Icelandic cod, Barents Sea cod, South African cape hakes, sockeye salmon, chinook salmon, southern bluefin tuna, Pacific halibut, Greenland halibut and Patagonian toothfish. The reviewed fish stocks are systemized in three categories: (1) stock properties and status; (2) management structure and objectives; and (3) management advice. We gather evidence to outline qualities of management regimes that are recommended and highlight those that most often fail. Robust management, biological limits (reference points), implementation and consensus are critical points that separate successful and unsuccessful management regimes. We evaluate each fish stock’s management performance relative to its management objectives and current conservation issues. Furthermore, we point out the importance of stakeholder involvement in fisheries management as well as the problems that international fisheries commissions face through examples from the case studies. Management successes tended to be single-nation and single-stock fisheries with capacity control and clear stakeholder involvement. Fisheries with fleet overcapacity, unclear objectives and illegal activity characterized the case studies with management problems.     

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.     

Food web dynamics affect Northeast Arctic cod recruitment

Proper management of ecosystems requires an understanding of both the species interactions as well as the effect of climate variation. However, a common problem is that the available time-series are of different lengths. Here, we present a general approach for studying the dynamic structure of such interactions. Specifically, we analyse the recruitment of the world's largest cod stock, the Northeast Arctic cod. Studies based on data starting in the 1970-1980s indicate that this stock is affected by temperature through a variety of pathways. However, the value of such studies is somewhat limited by the fact that they are based on a quite specific ecological and climatic situation. Recently, this stock has consisted of fairly young fish and the spawning stock has consisted of relatively few age groups. In this study, we develop a model for the effect of capelin (the cod's main prey) and herring on cod recruitment since 1973. Based on this model, we analyse data on cod, herring and temperature going back to 1921 and find that food-web effects explain a significant part of the cod recruitment variation back to around 1950.     

Stock collapse and its effect on species interactions: Cod and herring in the Norwegian‐Barents Seas system as an example

Both the Norwegian Spring Spawning herring (Clupea harengus) and the Northeast Arctic (NEA) cod (Gadus morhua) are examples of strong stock reduction and decline of the associated fisheries due to overfishing followed by a recovery. Cod and herring are both part of the Barents Sea ecosystem, which has experienced major warming events in the early (1920–1940) and late 20th century. While the collapse or near collapse of these stocks seems to be linked to an instability created by overfishing and climate, the difference of population dynamics before and after is not fully understood. In particular, it is unclear how the changes in population dynamics before and after the collapses are associated with biotic interactions. The combination of the availability of unique long‐term time series for herring and cod makes it a well‐suited study system to investigate the effects of collapse. We examine how species interactions may differently affect the herring and cod population dynamic before and after a collapse. Particularly we explore, using a GAM modeling approach, how herring could affect cod and vice versa. We found that the effect of cod biomass on herring that was generally positive (i.e., covariation) but the effect became negative after the collapse (i.e., predation or competition). Likewise a change occurred for the cod, the juvenile herring biomass that had no effect before the collapse had a negative effect after. Our results indicate that the population collapses may alter the inter‐specific interactions and response to abiotic environmental changes. While the stocks are at similar abundance levels before and after the collapses, the system is potentially different in its functioning and may require different management action.     

Characteristics of the Norwegian Coastal Current during Years with High Recruitment of Norwegian Spring Spawning Herring (Clupea harengus L.)

Norwegian Spring Spawning herring (NSSH) Clupea harengus L. spawn on coastal banks along the west coast of Norway. The larvae are generally transported northward in the Norwegian Coastal Current (NCC) with many individuals utilizing nursery grounds in the Barents Sea. The recruitment to this stock is highly variable with a few years having exceptionally good recruitment. The principal causes of recruitment variability of this herring population have been elusive. Here we undertake an event analysis using data between 1948 and 2010 to gain insight into the physical conditions in the NCC that coincide with years of high recruitment. In contrast to a typical year when northerly upwelling winds are prominent during spring, the years with high recruitment coincide with predominantly southwesterly winds and weak upwelling in spring and summer, which lead to an enhanced northward coastal current during the larval drift period. Also in most peak recruitment years, low-salinity anomalies are observed to propagate northward during the spring and summer. It is suggested that consistent southwesterly (downwelling) winds and propagating low-salinity anomalies, both leading to an enhanced northward transport of larvae, are important factors for elevated recruitment. At the same time, these conditions stabilize the coastal waters, possibly leading to enhanced production and improved feeding potential along the drift route to Barents Sea. Further studies on the drivers of early life history mortality can now be undertaken with a better understanding of the physical conditions that prevail during years when elevated recruitment occurs in this herring stock.     

Evidence for Two Highly Differentiated Herring Groups at Goose Bank in the Barents Sea and the Genetic Relationship to Pacific Herring, Clupea pallasi

Genetic studies on Atlantic herring, Clupea harengus, have generally revealed a low level of genetic variation over large geographic areas. Genetically distinct herring populations in some of the Norwegian fjords are exceptions, and juvenile herring from the large oceanic herring, Norwegian Spring Spawners (NSS), are often found in mixture with local fjord populations as well as widely distributed in the Barents Sea. Research surveys in the eastern Barents Sea (Goose Bank) in 1993, 1994 and 2001 included collection of herring samples for allozyme analyses. As expected the results identified juveniles from NSS stock, but an additional unique group of herring (low vertebrae number), being almost fixed for alternative alleles at several allozyme loci, was detected. In some cases, the two groups of herring were taken in the same trawl catches as documented by highly significant departure from Hardy—Weinberg expectation with large excess of homozygotes providing evidence for population mixing. Large genetic differences (Nei's genetic distance = 1.53; F_ST = 0.754) were detected in pairwise comparisons based on five allozyme loci. The two herring groups were also compared with reference samples of Pacific herring, Clupea pallasi, including one sample from Japan Sea and three Alaskan samples. UPGMA dendrogram based on five allozyme loci revealed a close genetic relationship between the low vertebrae herring in the Barents Sea and the group of samples of Pacific herring. Although significant different in allele frequencies, one of the herring samples clustered together with the reference sample from Bering Sea with genetic distance of 0.008 and F_ST value of 0.032. The close genetic relationship found in this paper, suggest a re-evaluation of the taxonomic status of the Barents Sea herring populations investigated.     

Structure,biomass distribution,and energetics of the pelagic ecosystem in the Barents Sea: A synopsis

Biomass distribution and energetics of trophic levels in the pelagic ecosystem of the Barents Sea are presented as averages over several years for the whole Barents Sea using data from the research programme Pro Mare in 1984–1989 and mathematical ecosystem models. Average biomasses range from more than 3 tonnes carbon km^–2 (zooplankton) to 0.1 kg C km^–2 (polar bears) and P/B ratios from 300 (bacteria) to 0.035 (minke whales). However, the Barents Sea ecosystem is in a far from steady state with, for instance, capelin stocks ranging from 30–700 kg C km^–2 between years and cod stocks from 150–700 kg C km^–2. As a general rule, the various fish stocks grow adequately, albeit at different rates, in warm years characterized by large influxes of Atlantic water and high zooplankton productivity. The skewed populations distribution which arises in warm years may lead to grave imbalances in cold years and even to the collapses of stocks, such as of capelin in the eighties. The food requirements of average-sized stocks of cod, seabirds and marine mammals correspond to more than twice the average productivity of capelin. Thus other species of pelagic fish (herring, polar cod) and zooplankton obviously play major roles as prey for these animals.     

Diet of juvenile cod (age 0–2) in the Barents Sea in relation to food availability and cod growth

Diet investigations were carried out on 0-, 1- and 2-year-old Northeast Arctic cod (Gadus morhua) sampled in the Barents Sea during 1984–2002. Stomach-content analyses showed that the 0 and 1 group cod fed mainly on crustaceans, with krill and amphipods composing up to 70% of their diet. Krill (Thysanoessa spp. and Meganyctiphanes norvegica) and amphipods (Themisto spp.) were mainly found in cod stomachs sampled in the central and close to the Polar Front region in the Barents Sea where these prey organisms are reported to be abundant in summer. A shift in the main diet from crustaceans to fish was observed from age 1 to age 2. The diet of 2-year-old cod mainly comprised capelin (Mallotus villosus) and other fish, and to a lesser degree, krill and amphipods. Shrimp (mainly Pandalus spp.) was also an important prey in both age 1 and 2 cod. A statistically significant positive relationship was obtained between capelin stock size and the amount of capelin in the diet of 2-year-old cod. Results from this study also show that the larger age-2 cod preyed more on capelin in winter and that larger cod (>22 cm) prefer larger capelin (>12 cm). During periods of low capelin abundance, the 2-year-old cod shift their diet more to crustaceans, such as krill and amphipods. A positive significant relationship was also obtained between Total Fullness Index (TFI) and the amount of capelin in the diet and between TFI and the growth of 2-year-old cod, indicating that the growth of age-2 cod is to a large extent dependent on the amount of capelin consumed. Growth of age-1 cod was also positively correlated to TFI.     

Forecasting future recruitment success for Atlantic cod in the warming and acidifying Barents Sea

Productivity of marine fish stocks is known to be affected by environmental and ecological drivers, and global climate change is anticipated to alter recruitment success of many stocks. While the direct effects of environmental drivers on fish early life stage survival can be quantified experimentally, indirect effects in marine ecosystems and the role of adaptation are still highly uncertain. We developed an integrative model for the effects of ocean warming and acidification on the early life stages of Atlantic cod in the Barents Sea, termed SCREI (Simulator of Cod Recruitment under Environmental Influences). Experimental results on temperature and CO₂ effects on egg fertilization, egg and larval survival and development times are incorporated. Calibration using empirical time series of egg production, temperature, food and predator abundance reproduces age‐0 recruitment over three decades. We project trajectories of recruitment success under different scenarios and quantify confidence limits based on variation in experiments. A publicly accessible web version of the SCREI model can be run under www.oceanchange.uni-bremen.de/ ;SCREI. Severe reductions in average age‐0 recruitment success of Barents Sea cod are projected under uncompensated warming and acidification toward the middle to end of this century. Although high population stochasticity was found, considerable rates of evolutionary adaptation to acidification and shifts in organismal thermal windows would be needed to buffer impacts on recruitment. While increases in food availability may mitigate short‐term impacts, an increase in egg production achieved by stock management could provide more long‐term safety for cod recruitment success. The SCREI model provides a novel integration of multiple driver effects in different life stages and enables an estimation of uncertainty associated with interindividual and ecological variation. The model thus helps to advance toward an improved empirical foundation for quantifying climate change impacts on marine fish recruitment, relevant for ecosystem‐based assessments of marine systems under climate change.     

The dangers of ignoring stock complexity in fishery management: the case of the North Sea cod

The plight of the marine fisheries is attracting increasing attention as unsustainably high exploitation levels, exacerbated by more extreme climatic conditions, are driving stocks to the point of collapse. The North Atlantic cod (Gadus morhua), a species which until recently formed a major component of the demersal fisheries, has undergone significant declines across its range. The North Sea stock is typical of many, with a spawning stock biomass that has remained below the safe biological limit since 2000 and recruitment levels near the lowest on record. Cod within the North Sea are currently managed as a single stock, and yet mounting empirical evidence supports the existence of a metapopulation of regionally variable, genetically distinct, sub-stocks. Applying the same management strategies to multiple stocks that differ in their resilience to exploitation inevitably results in the overfishing and likely collapse of the weaker components. Indeed, recent studies have identified two North Sea spawning stocks that have undergone disproportionally large collapses with very substantial reductions in egg production. Similarly affected cod stocks in the northwest Atlantic have shown little evidence of recovery, despite fishery closures. The possible implications of ignoring sub-structuring within management units for biocomplexity, local adaptation and ecosystem stability are considered.     

Towards better Understanding of Successful Management of the Gulf of Riga Herring Stock

The Baltic herring is one of the key elements of the pelagic ecosystem of the Baltic Sea, being of the most important commercial species of the sea. The structure and dynamics of herring populations (stocks) have been assessed and managed on international level since 1970s. Since 1990, all local herring populations have been combined and assessed as one stock (Central Baltic Herring in Sub-divisions 25–29 and 32). However, the continuously decreasing trend in stock biomass throughout almost 30-years period of observations indicates the failure of the stock management implementation of the scientific advice. The separate assessments of different stocks and results of hydro-acoustic surveys of different sub-units (herring in Sub-divisions 25–27, Sub-divisions 28, 29 and 32 and the Gulf of Riga herring) have revealed rather different situation within combined assessment unit: e.g. fishing mortality of the Gulf of Riga herring has decreased while a sharp increase in mortality is observed in the herring stocks of the North-eastern Baltic (Sub-divisions 28, 29 and 32). At the same time stock abundance and biomass of the Gulf of Riga herring have increased to record high in early 2000s allowing also higher catches. It is concluded that only separate analytical assessment of local populations combined with regular acoustical surveys and following different protection measures have proved to be a successful way in managing the herring stocks. Joint assessment and management of several populations with different structure and dynamics as one combined stock do not allow revealing the real situation and trends in its parts.     

Feeding habits of harp seals (Phoca groenlandica) during early summer and autumn in the northern Barents Sea

The feeding habits of harp seals (Phoca groenlandica) in the Barents Sea were examined in studies conducted during June 1991, September 1990 and 1991, and October 1992. Analyses of stomach and intestinal contents were carried out and concurrent estimates of prey abundance were made using trawl gear. Harp seals appeared to feed at low intensity in the pack ice belt during the first half of June. There was little potential prey in the water column, but prawns (Pandalus borealis), capelin (Mallotus villosus) and polar cod (Boreogadus saida) were abundant close to the bottom. In September, the seals sampled in the northern pack ice areas of the Barents Sea fed on the pelagic amphipod Parathemisto libellula, krill (Thysanoessa spp.), prawns and, to a lesser extent, on fish species such as polar cod, sculpins (Cottidae) and snailfish (Liparidae). Trawling revealed that large quantities of Parathemisto libellala were present in the upper layers of the water column. Fish, mainly capelin and polar cod, were less abundant and occurred in deeper waters. In mid-October, the diet of seals in the northern Barents Sea consisted mainly of amphipods (Parathemisto sp.). Later in October, when increasing pack ice cover forced the harp seals to move south, the diet seemed to change from amphipods to fish prey, predominantly capelin and polar cod.     

Direct and indirect climate forcing in a multi-species marine system

Interactions within and between species complicate quantification of climate effects, by causing indirect, often delayed, effects of climate fluctuations and compensation of mortality. Here we identify direct and indirect climate effects by analysing unique Russian time-series data from the Norwegian Sea–Barents Sea ecosystem on the first life stages of cod, capelin, herring and haddock, their predators, competitors and zooplanktonic prey. By analysing growth and survival from one life stage to the next (eggs–larvae–juveniles–recruits), we find evidence for both bottom-up, direct and top-down effects of climate. Ambient zooplankton biomass predicts survival of all species, whereas ambient temperature mainly affects survival through effects on growth. In warm years, all species experienced improved growth and feeding conditions. Cohorts born following a warm year will, however, experience increased predation and competition because of increased densities of subadult cod and herring, leading to delayed climate effects. While climate thus affects early growth and survival through several mechanisms, only some of the identified mechanisms were found to be significant predictors of population growth. In particular, our findings exemplify that climate impacts are barely propagated to later life stages when density dependence is strong.     

Pantophysin (Pan I) locus divergence between inshore v. offshore and northern v. southern populations of Atlantic cod in the north‐east Atlantic

More than 6000 cod Gadus morhua , sampled in coastal and offshore waters stretching from the Barents Sea down to the North Sea, were analysed for frequencies of alleles at the scnDNA pantophysin locus ( Pan I)[formerly called synaptophysin ( Syp I)]. The significant allele frequency difference between the two major stocks of cod in Norway, north‐east Arctic cod (NEAC) and Norwegian coastal cod (NCC), was upheld in all years of the investigation (1993 to 2001), and applied both to larval cod and post‐juveniles of various ages. On a north‐south axis, the appearance of a latitudinal cline of post‐juvenile (≥1 year) allele frequencies was exposed. The intermediate allele frequencies in coastal areas of northern Norway, seem to a large extent to be caused by intermingling of the two stocks, although the existence of populations of coastal cod with alternative Pan I frequencies could not be ruled out. The role of selection is yet unresolved. Depth of the sampling location seemed to have an effect on the allele frequencies and their temporal stability, while there was no indication of seasonal variation in the frequencies. Breeding structure was the most likely cause for upholding the extreme divergence in Pan I frequencies between NEAC and NCC.     

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.     

Recent trends in the abundance of plaice Pleuronectes platessa and cod Gadus morhua in shallow coastal waters of the Northeastern Atlantic continental shelf – a review

Shallow, near-shore water habitats on the continental shelf of the Northeast Atlantic have been productive fishing areas in the past. Here, we review the present knowledge about (i) recent trends in the abundance of plaice and cod in these habitats and (ii) hypotheses regarding the factors responsible for any trends. At present, only a few studies exist on the trends of abundance of plaice or cod, namely from the Bay of Biscay, the North Sea and the Skagerrak/Kattegat. They suggest a declining abundance in coastal, shallow areas and – at least for plaice – a latitudinal gradient with an erosion of the southern distribution boundary in the Bay of Biscay and deepening of stocks in the North Sea. In contrast, no trend in shallow water abundance of plaice similar to a decline in deep-water stocks during the 1970s and their slow recovery during the 2000s is apparent in the Skagerrak/Kattegat. Although shallow habitats fundamentally differ from deeper areas by the prevalence of juvenile stages, the declining trends coincide with decreasing abundance/landings and spatial stock relocations in the deeper areas. Whether this indicates a common trend pointing at connectivity between shallow and deep water remains open. Fundamental differences exist in the suggested causes of the trends in different geographical areas. High fishing pressure together with low local recruitment apparently prevents the recovery of overexploited plaice and cod stocks in the Skagerrak/Kattegat. In contrast, the responses of juveniles and adult fish to increasing seawater temperature are the main hypotheses for changes in distribution and abundance of both fish species in the North Sea/Bay of Biscay. However, temperature alone cannot explain the observed decline of fish in coastal areas, and the causes may be more complex, involving nutrient loading, primary productivity or food availability, although at present, knowledge of these factors is insufficient.