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.     

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.     

Biodiversity and exploitation of the main fish stocks in the Norwegian - Barents Sea ecosystem

Juvenile herring and capelin are the main stocks of plankton feeders in the Barents Sea, the cod is the dominant predator. Warm climate favours recruitment of herring and cod, but large stocks of juvenile herring hamper survival of the capelin fry. Since the early 1970s, the herring stock has been grossly overexploited, which could have led to an imbalance in the state of the predatorprey relationships in the Barents Sea. In the 1970s and early 80s, however, cod could feed on capelin which had excellent growth and recruitment conditions when the herring was absent. The consequences of the reduced herring stock were triggered in the mid 1980s, when excellent recruitment conditions for herring and cod occurred. Three abundant year classes of cod were recruited, but the herring stock was not sufficiently large to take full advantage of the favourable recruitment conditions. Given the lack of juvenile herring and a reduced capelin stock, the rapidly growing cod stock grazed down all other available prey species in the area, including its own progeny, and starved cod, seabirds and seals have in later years appeared on the north Norwegian coast. The capelin fishery collapsed, and the traditional coastal cod fisheries have been struck by the most serious crisis on record.     

Harp seal foraging behaviour during summer around Svalbard in the northern Barents Sea: diet composition and the selection of prey

The harp seal Pagophilus groenlandicus is a major high trophic level predator in the Barents Sea, and to better understand their function in the Barents Sea ecosystem, we need to understand their foraging behaviour during their most intensive feeding period. We analysed the diet composition and prey preference of 184 harp seals and 94 faeces samples, sampled in the northern Barents Sea (around Svalbard) during the period May–August in 1996, 1997, and 2004–2006. Concurrent with the sampling of seals, prey availability was assessed in one area in 1996 and 1997 and in two areas in 2006 using standard acoustic methods. Our study showed that harp seal diet composition varied significantly both in time (year) and space, and that their diets appeared to be size dependent. Both subadult (<150 cm) and adult seals were associated with pelagic crustaceans (particularly krill), whereas primarily adult seals were associated with fish (capelin, gadoids and flatfish). Krill was the most important prey group (63 %) followed by polar cod (16 %) and other fish species (10 %). The prey preference of harp seals varied in time and space; polar cod was often preferred by the seals whereas krill was commonly consumed in lower proportion than observed in the survey area. Gadoids and capelin had either been exploited in the same or less proportion as observed in the survey sea. This study emphasises the ecological significance of krill as prime food for harp seals during their intensive feeding period in summer.     

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.     

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.     

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.     

Resource‐driven colonization by cod in a high Arctic food web

1. Climate change is commonly associated with many species redistributions and the influence of other factors may be marginalized, especially in the rapidly warming Arctic.
2. The Barents Sea, a high latitude large marine ecosystem in the Northeast Atlantic has experienced above‐average temperatures since the mid‐2000s with divergent bottom temperature trends at subregional scales.
3. Concurrently, the Barents Sea stock of Atlantic cod Gadus morhua, one of the most important commercial fish stocks in the world, increased following a large reduction in fishing pressure and expanded north of 80°N.
4. We examined the influence of food availability and temperature on cod expansion using a comprehensive data set on cod stomach fullness stratified by subregions characterized by divergent temperature trends. We then tested whether food availability, as indexed by cod stomach fullness, played a role in cod expansion in subregions that were warming, cooling, or showed no trend.
5. The greatest increase in cod occupancy occurred in three northern subregions with contrasting temperature trends. Cod apparently benefited from initial high food availability in these regions that previously had few large‐bodied fish predators.
6. The stomach fullness in the northern subregions declined rapidly after a few years of high cod abundance, suggesting that the arrival of cod caused a top‐down effect on the prey base. Prolonged cod residency in the northern Barents Sea is, therefore, not a certainty.

     

Lack of spatial coherence of predators with prey: a bioenergetic explanation for Atlantic cod feeding on capelin

We tested two biologically based predictions that potentially influence scales of spatial association between Atlantic cod, Gadus morhua , and prey populations of capelin, Mallotus vilhsus . If cod aggregate in response to concentrations of prey, then spatial association (coherence) between capelin and cod was predicted to peak at the scale of maximum capelin spatial variance. If capelin-cod coherence did not match the scale of maximum prey spatial variability, then capelin-cod coherence was predicted to peak at the spatial scale that maximizes net energetic benefit to the predator. Contrary to predictions, we found no evidence of aggregative responses of cod to capelin over resolution scales of 20 m to 10 km. This result was observed consistently at the temporal scale of a single transect ( c . 1 h duration) and at the scale of averaged transects ( c . 2 weeks duration). Estimates of cod foraging energetics showed that they were not constrained by physiology to aggregate relative to capelin at any scale less than 10 km. A net energetic gain of 478 to 784 kJ would result if a 44 cm, 752 g cod consumed a ration of eight to 12 capelin over a period of 58 h. Energetic calculations included costs of egestion and excretion (317 to 476 kJ), maintenance (58 kJ), digestion (125 to 188 kJ), and continuous swimming during ration assimilation (79 kJ). During this period, a 44 cm cod could travel over 38 km swimming at 1 b.l. s⁻¹. Foraging cod are virtually certain to encounter capelin over this distance based on the abundance of pre-spawning capelin present in coastal bays during the spawning season. This study illustrates that aggregative responses of predators do no occur at all scales and possibly occur over a very limited range of scales.     

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.     

Microzooplankton Growth Rates Examined across a Temperature Gradient in the Barents Sea

Growth rates (µ) of abundant microzooplankton species were examined in field experiments conducted at ambient sea temperatures (−1.8–9.0°C) in the Barents Sea and adjacent waters (70–78.5°N). The maximum species-specific µ of ciliates and athecate dinoflagellates (0.33–1.67 d⁻¹ and 0.52–1.14 d⁻¹, respectively) occurred at temperatures below 5°C and exceeded the µ_max predicted by previously published, laboratory culture-derived equations. The opposite trend was found for thecate dinoflagellates, which grew faster in the warmer Atlantic Ocean water. Mixotrophic ciliates and dinoflagellates grew faster than their heterotrophic counterparts. At sub-zero temperatures, microzooplankton µ_max matched those predicted for phytoplankton by temperature-dependent growth equations. These results indicate that microzooplankton protists may be as adapted to extreme Arctic conditions as their algal prey.     

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.     

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.     

Divergent trends in anadromous salmonid populations in Norwegian and Scottish rivers

The Atlantic salmon (Salmo salar) is a charismatic anadromous fish of high conservation and economic value. Concerns have been expressed regarding the long-term viability of fisheries throughout the species''s distributional range because of abundance variations that cannot currently be explained or predicted. Here, we analyse long-term catch data obtained over a wide geographical range and across a range of spatial subscales to understand more fully the factors that drive population abundance. We use rod catch data from 84 Norwegian rivers over 125 years (1876–2000) and 48 Scottish rivers over 51 years (1952–2002). The temporal correlation in catches is very long-term, with trends persisting over several decades. The spatial correlation is relatively short-range, indicating strong local-scale effects on catch. Furthermore, Scottish salmon populations exhibit recent negative trends in contrast to some more positive trends in Norway—especially in the north.     

Spatial variance of mobile aquatic organisms: capelin and cod in Newfoundland coastal waters

Spatial variance in the distribution of aquatic mobile organisms differs from that of passive tracers such as phytoplankton or water temperature. On average, spatial variance of phytoplankton scales with sample unit as $L^2$ or equivalently with frequency as $f^{-2}$. Limited evidence suggests that spatial variance in the distribution of mobile organisms is concentrated at relatively small scales, with little increase over larger scales: spatial variance scales as $f^{-1}$ or less. We investigated whether spatial variance in distributions of a mobile predator, Atlantic cod (Gadus morhua), and a schooling prey, capelin (Mallotus villosus), also scale with frequency as $f^{-1}$. Acoustic surveys showed that at short time scales spatial variance in cod and capelin densities, as measured by spectral density, peaked at various scales ranging from 20 m to 10 km. At longer time scales, spatial variance of cod scaled as $f^{-1.08}$ at resolutions finer than 90 m, while scaling as $f^{-0.18}$ at coarser scales. Spatial variance of capelin scaled as $f^{-1.1}$ at resolutions finer than 400 m, while scaling as $f^{-0.21}$ at coarser scales. Spatial variance plots of krill and marine birds showed similar transitions from shallow to steep scaling. Shoaling, schooling and the aggregative response by predators to concentrations of prey were three processes hypothesized to influence spatial variance in distributions of mobile organisms. Numerical experiments showed that shoaling injects variance at large to intermediate scales, resulting in scalings flatter than $f^{-1}$. Additional experiments showed that schooling produces a transition from shallow to steep scaling as frequency increases. Spatial variance patterns in cod density were not due to aggregative responses by the predator to concentrations of capelin: there was no association, on average, at resolution scales from 20 m to 10 km. Exponent values for aquatic or terrestrial mobile organisms are predicted to be approximately two at the scale of an individual organism, 0.2 at scales that contain aggregations, and two at scales larger than that of populations. These findings suggest that relations between mobile organisms and large scale habitat variables will be difficult to detect, that stratified survey designs used to estimate commercial population sizes will be inefficient, and that rates of interaction between predator and prey will be underestimated if local observations are averaged over the spatial scale of the population.     

Climate-Driven Ichthyoplankton Drift Model Predicts Growth of Top Predator Young

Climate variability influences seabird population dynamics in several ways including access to prey near colonies during the critical chick-rearing period. This study addresses breeding success in a Barents Sea colony of common guillemots Uria aalge where trophic conditions vary according to changes in the northward transport of warm Atlantic Water. A drift model was used to simulate interannual variations in transport of cod Gadus morhua larvae along the Norwegian coast towards their nursery grounds in the Barents Sea. The results showed that the arrival of cod larvae from southern spawning grounds had a major effect on the size of common guillemot chicks at fledging. Furthermore, the fraction of larvae from the south was positively correlated to the inflow of Atlantic Water into the Barents Sea thus clearly demonstrating the mechanisms by which climate-driven bottom-up processes influence interannual variations in reproductive success in a marine top predator.     

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.     

Ecology of Arctic cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> (Gadiformes,Gadidae) and its fishery potential in Kara Sea

According to the bottom trawl-survey data, 97% of the ichthyomass in the southwestern region of the Kara Sea are composed of the Arctic cod Boreogadus saida; its stock is significantly higher than the previously registered resources. The Arctic cod is most unevenly distributed across the water area and capable to form the high-density aggregations, which can be caught by the targeted trawls. A wide range of the age composition (0+?6+), the size-age composition, and the growth rates of the Arctic cod in the trawl catches in the Kara Sea, which are different from those in the fish in the adjacent Barents Sea, can indicate their assignment of the Arctic cod in these seas to different populations.     

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.