Investigating the potential use of aerial line transect surveys for estimating polar bear abundance in sea ice habitats: A case study for the Chukchi Sea

The expense of traditional capture‐recapture methods, interest in less invasive survey methods, and the circumpolar decline of polar bear (Ursus maritimus) habitat require evaluation of alternative methods for monitoring polar bear populations. Aerial line transect distance sampling (DS) surveys are thought to be a promising monitoring tool. However, low densities and few observations during a survey can result in low precision, and logistical constraints such as heavy ice and fuel and safety limitations may restrict survey coverage. We used simulations to investigate the accuracy and precision of, DS for estimating polar bear abundance in sea ice habitats, using the Chukchi Sea subpopulation as an example. Simulation parameters were informed from a recent pilot survey. Predictions from a resource selection model were used for stratification, and we compared two ratio estimators to account for areas that cannot be sampled. The ratio estimator using predictions of resource selection by polar bears allowed for extrapolation beyond sampled areas and provided results with low bias and CVs ranging from 21% to 36% when abundance was >1,000. These techniques could be applied to other DS surveys to allocate effort and potentially extrapolate estimates to include portions of the landscape that are logistically impossible to survey.     

Sea ice resource selection models for polar bears in the Barents Sea subpopulation

The extent, thickness and age of Arctic sea ice has dramatically declined since the late 1990s, and these trends are predicted to continue. Exploring the habitat use of sea‐ice‐dependent species can help us understand which resources they use and how their distribution responds to a changing environment. The goal of this study was to develop predictive models of the habitat use of an Arctic apex predator. Polar bear Ursus maritimus habitat use in the Barents Sea subpopulation was modelled with seasonal resource selection functions (RSFs) using satellite‐linked telemetry data from 294 collars deployed on female polar bears between 1991 and 2015. Polar bears selected habitat in the Marginal Ice Zone, with a preference for intermediate sea ice concentrations (40–80%). They spent most time in areas with relatively short travel distances to 15 or 75% ice concentration, and during spring and autumn they exhibited a preference for sea ice areas over the continental shelf or over shallower bathymetry). Predictions of the distribution of polar bears in the Barents Sea area can be made for specific sea ice scenarios using these models. Two such predictive distribution maps based on the autumn seasonal model were made and validated against two independent polar bear survey datasets collected in August 2004 and August 2015. The distribution of optimal polar bear habitat has shifted strongly northwards in all seasons of the year during the 25 yr study period.     

East Greenland and Barents Sea polar bears (Ursus maritimus): adaptive variation between two populations using skull morphometrics as an indicator of environmental and genetic differences

A morphometric study was conducted on four skull traits of 37 male and 18 female adult East Greenland polar bears (Ursus maritimus) collected 1892-1968, and on 54 male and 44 female adult Barents Sea polar bears collected 1950-1969. The aim was to compare differences in size and shape of the bear skulls using a multivariate approach, characterizing the variation between the two populations using morphometric traits as an indicator of environmental and genetic differences. Mixture analysis testing for geographic differentiation within each population revealed three clusters for Barents Sea males and three clusters for Barents Sea females. East Greenland consisted of one female and one male cluster. A principal component analysis (PCA) conducted on the clusters defined by the mixture analysis, showed that East Greenland and Barents Sea polar bear populations overlapped to a large degree, especially with regards to females. Multivariate analyses of variance (MANOVA) showed no significant differences in morphometric means between the two populations, but differences were detected between clusters from each respective geographic locality. To estimate the importance of genetics and environment in the morphometric differences between the bears, a PCA was performed on the covariance matrix derived from the skull measurements. Skull trait size (PC1) explained approx. 80% of the morphometric variation, whereas shape (PC2) defined approx. 15%, indicating some genetic differentiation. Hence, both environmental and genetic factors seem to have contributed to the observed skull differences between the two populations. Overall, results indicate that many Barents Sea polar bears are morphometrically similar to the East Greenland ones, suggesting an exchange of individuals between the two populations. Furthermore, a subpopulation structure in the Barents Sea population was also indicated from the present analyses, which should be considered with regards to future management decisions.     

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.     

Brucella sp. antibodies in polar bears from Svalbard and the Barents Sea

A prevalence of 5.4% of anti-Brucella sp. antibodies was found in plasma samples from 297 polar bears (Ursus maritimus) from Svalbard and the Barents Sea. Plasma was tested by the classical brucellosis tests Slow Agglutination of Wright (SAW), EDTA modified SAW and Rose Bengal test, as well as by an indirect Protein A ELISA. Only samples classified as positive in all tests were regarded as containing anti-Brucella sp. antibodies. A significant west to east increase in the proportion of bears with anti-Brucella sp. antibodies was found, with 3.6% (n = 253) at Svalbard (Spitsbergen, Nordaustlandet, Edge?ya, Barents?ya and Hopen), and 15.9% (n = 44) in the central Barents Sea. Anti-Brucella sp. antibodies were previously found in ringed seals (Phoca hispida) and harp seals (Phoca groenlandica) from the same geographical areas. The ringed seal is an important prey species for the Svalbard polar bear population, and may thus be a source of brucellosis for the bears. There are no indications of reproductive disorders caused by Brucella sp. or other infectious agents in our study polar bear population. Potential impacts of Brucella sp. exposure on individuals or the population are unknown.     

Implications of the Circumpolar Genetic Structure of Polar Bears for Their Conservation in a Rapidly Warming Arctic

We provide an expansive analysis of polar bear (Ursus maritimus) circumpolar genetic variation during the last two decades of decline in their sea-ice habitat. We sought to evaluate whether their genetic diversity and structure have changed over this period of habitat decline, how their current genetic patterns compare with past patterns, and how genetic demography changed with ancient fluctuations in climate. Characterizing their circumpolar genetic structure using microsatellite data, we defined four clusters that largely correspond to current ecological and oceanographic factors: Eastern Polar Basin, Western Polar Basin, Canadian Archipelago and Southern Canada. We document evidence for recent (ca. last 1–3 generations) directional gene flow from Southern Canada and the Eastern Polar Basin towards the Canadian Archipelago, an area hypothesized to be a future refugium for polar bears as climate-induced habitat decline continues. Our data provide empirical evidence in support of this hypothesis. The direction of current gene flow differs from earlier patterns of gene flow in the Holocene. From analyses of mitochondrial DNA, the Canadian Archipelago cluster and the Barents Sea subpopulation within the Eastern Polar Basin cluster did not show signals of population expansion, suggesting these areas may have served also as past interglacial refugia. Mismatch analyses of mitochondrial DNA data from polar and the paraphyletic brown bear (U. arctos) uncovered offset signals in timing of population expansion between the two species, that are attributed to differential demographic responses to past climate cycling. Mitogenomic structure of polar bears was shallow and developed recently, in contrast to the multiple clades of brown bears. We found no genetic signatures of recent hybridization between the species in our large, circumpolar sample, suggesting that recently observed hybrids represent localized events. Documenting changes in subpopulation connectivity will allow polar nations to proactively adjust conservation actions to continuing decline in sea-ice habitat.     

Improving Aquatic Warbler Population Assessments by Accounting for Imperfect Detection

Monitoring programs designed to assess changes in population size over time need to account for imperfect detection and provide estimates of precision around annual abundance estimates. Especially for species dependent on conservation management, robust monitoring is essential to evaluate the effectiveness of management. Many bird species of temperate grasslands depend on specific conservation management to maintain suitable breeding habitat. One such species is the Aquatic Warbler (Acrocephalus paludicola), which breeds in open fen mires in Central Europe. Aquatic Warbler populations have so far been assessed using a complete survey that aims to enumerate all singing males over a large area. Because this approach provides no estimate of precision and does not account for observation error, detecting moderate population changes is challenging. From 2011 to 2013 we trialled a new line transect sampling monitoring design in the Biebrza valley, Poland, to estimate abundance of singing male Aquatic Warblers. We surveyed Aquatic Warblers repeatedly along 50 randomly placed 1-km transects, and used binomial mixture models to estimate abundances per transect. The repeated line transect sampling required 150 observer days, and thus less effort than the traditional ‘full count’ approach (175 observer days). Aquatic Warbler abundance was highest at intermediate water levels, and detection probability varied between years and was influenced by vegetation height. A power analysis indicated that our line transect sampling design had a power of 68% to detect a 20% population change over 10 years, whereas raw count data had a 9% power to detect the same trend. Thus, by accounting for imperfect detection we increased the power to detect population changes. We recommend to adopt the repeated line transect sampling approach for monitoring Aquatic Warblers in Poland and in other important breeding areas to monitor changes in population size and the effects of habitat management.     

Adaptive line transect sampling

Adaptive line transect sampling offers the potential of improved population density estimation efficiency over conventional line transect sampling when populations are spatially clustered. In adaptive sampling, survey effort is increased when areas of high animal density are located, thereby increasing the number of observations. Its disadvantage is that the survey effort required is not known in advance. We develop an adaptive line transect methodology that, by varying the degree of adaptation, allows total effort to be fixed at the design stage. Relative to conventional line transect surveys, it also provides better survey coverage in the event of disruption in survey effort, e.g., due to poor weather. In analysis, sightings from the adaptive sections are downweighted in proportion to the increase in effort. We evaluate the methodology by simulation and report on surveys of harbor porpoise in the Gulf of Maine, in which the approach was compared with conventional line transect sampling.     

Pooling robustness in distance sampling: Avoiding bias when there is unmodelled heterogeneity

The pooling robustness property of distance sampling results in unbiased abundance estimation even when sources of variation in detection probability are not modeled. However, this property cannot be relied upon to produce unbiased subpopulation abundance estimates when using a single pooled detection function that ignores subpopulations. We investigate by simulation the effect of differences in subpopulation detectability upon bias in subpopulation abundance estimates. We contrast subpopulation abundance estimates using a pooled detection function with estimates derived using a detection function model employing a subpopulation covariate. Using point transect survey data from a multispecies songbird study, species-specific abundance estimates are compared using pooled detection functions with and without a small number of adjustment terms, and a detection function with species as a covariate. With simulation, we demonstrate the bias of subpopulation abundance estimates when a pooled detection function is employed. The magnitude of the bias is positively related to the magnitude of disparity between the subpopulation detection functions. However, the abundance estimate for the entire population remains unbiased except when there is extreme heterogeneity in detection functions. Inclusion of a detection function model with a subpopulation covariate essentially removes the bias of the subpopulation abundance estimates. The analysis of the songbird point count surveys shows some bias in species-specific abundance estimates when a pooled detection function is used. Pooling robustness is a unique property of distance sampling, producing unbiased abundance estimates at the level of the study area even in the presence of large differences in detectability between subpopulations. In situations where subpopulation abundance estimates are required for data-poor subpopulations and where the subpopulations can be identified, we recommend the use of subpopulation as a covariate to reduce bias induced in subpopulation abundance estimates.     

Demographic and traditional knowledge perspectives on the current status of Canadian polar bear subpopulations

Subpopulation growth rates and the probability of decline at current harvest levels were determined for 13 subpopulations of polar bears (Ursus maritimus) that are within or shared with Canada based on mark–recapture estimates of population numbers and vital rates, and harvest statistics using population viability analyses (PVA). Aboriginal traditional ecological knowledge (TEK) on subpopulation trend agreed with the seven stable/increasing results and one of the declining results, but disagreed with PVA status of five other declining subpopulations. The decline in the Baffin Bay subpopulation appeared to be due to over‐reporting of harvested numbers from outside Canada. The remaining four disputed subpopulations (Southern Beaufort Sea, Northern Beaufort Sea, Southern Hudson Bay, and Western Hudson Bay) were all incompletely mark–recapture (M‐R) sampled, which may have biased their survival and subpopulation estimates. Three of the four incompletely sampled subpopulations were PVA identified as nonviable (i.e., declining even with zero harvest mortality). TEK disagreement was nonrandom with respect to M‐R sampling protocols. Cluster analysis also grouped subpopulations with ambiguous demographic and harvest rate estimates separately from those with apparently reliable demographic estimates based on PVA probability of decline and unharvested subpopulation growth rate criteria. We suggest that the correspondence between TEK and scientific results can be used to improve the reliability of information on natural systems and thus improve resource management. Considering both TEK and scientific information, we suggest that the current status of Canadian polar bear subpopulations in 2013 was 12 stable/increasing and one declining (Kane Basin). We do not find support for the perspective that polar bears within or shared with Canada are currently in any sort of climate crisis. We suggest that monitoring the impacts of climate change (including sea ice decline) on polar bear subpopulations should be continued and enhanced and that adaptive management practices are warranted.     

Using distance sampling techniques to estimate bottlenose dolphin (Tursiops truncatus) abundance at Turneffe Atoll,Belize

Reliable abundance estimates are critical for management and conservation of coastal small cetaceans. This is particularly important in developing countries where coastal human populations are increasing, the impacts of anthropogenic activities are often unknown, and the resources necessary to assess coastal cetaceans are limited. We adapted ship‐based line transect methods to small‐boat surveys to estimate the abundance of bottlenose dolphins (Tursiops truncatus) at Turneffe Atoll, Belize. Using a systematic survey design with random start and uniform coverage, 34 dolphin clusters were sighted during small‐boat line transect surveys conducted in 2005–2006. Distance sampling methods estimated abundance at 216 individuals (CV = 27.7%, 95% CI = 126–370). Due to species rarity in the Atoll, small sample size, and potential violations in line transect assumptions, the estimate should be considered preliminary. Nevertheless, it provides up‐to‐date information on the status of a regional population in an area under increasing threat of habitat loss and prey depletion via uncontrolled development and unsustainable fishing. This information will be useful as Belize develops a new conservation initiative to create a comprehensive and resilient marine protected area system. Our study illustrates the application of distance sampling methods to small‐boat surveys to obtain abundance estimates of coastal cetaceans in a region lacking resources.     

Dietary habits of polar bears in Foxe Basin,Canada: possible evidence of a trophic regime shift mediated by a new top predator

Polar bear (Ursus maritimus) subpopulations in several areas with seasonal sea ice regimes have shown declines in body condition, reproductive rates, or abundance as a result of declining sea ice habitat. In the Foxe Basin region of Nunavut, Canada, the size of the polar bear subpopulation has remained largely stable over the past 20 years, despite concurrent declines in sea ice habitat. We used fatty acid analysis to examine polar bear feeding habits in Foxe Basin and thus potentially identify ecological factors contributing to population stability. Adipose tissue samples were collected from 103 polar bears harvested during 2010–2012. Polar bear diet composition varied spatially within the region with ringed seal (Pusa hispida) comprising the primary prey in northern and southern Foxe Basin, whereas polar bears in Hudson Strait consumed equal proportions of ringed seal and harp seal (Pagophilus groenlandicus). Walrus (Odobenus rosmarus) consumption was highest in northern Foxe Basin, a trend driven by the ability of adult male bears to capture large‐bodied prey. Importantly, bowhead whale (Balaena mysticetus) contributed to polar bear diets in all areas and all age and sex classes. Bowhead carcasses resulting from killer whale (Orcinus orca) predation and subsistence harvest potentially provide an important supplementary food source for polar bears during the ice‐free period. Our results suggest that the increasing abundance of killer whales and bowhead whales in the region could be indirectly contributing to improved polar bear foraging success despite declining sea ice habitat. However, this indirect interaction between top predators may be temporary if continued sea ice declines eventually severely limit on‐ice feeding opportunities for polar bears.     

Double-Observer Line Transect Methods: Levels of Independence

Summary .  Double-observer line transect methods are becoming increasingly widespread, especially for the estimation of marine mammal abundance from aerial and shipboard surveys when detection of animals on the line is uncertain. The resulting data supplement conventional distance sampling data with two-sample mark–recapture data. Like conventional mark–recapture data, these have inherent problems for estimating abundance in the presence of heterogeneity. Unlike conventional mark–recapture methods, line transect methods use knowledge of the distribution of a covariate, which affects detection probability (namely, distance from the transect line) in inference. This knowledge can be used to diagnose unmodeled heterogeneity in the mark–recapture component of the data. By modeling the covariance in detection probabilities with distance, we show how the estimation problem can be formulated in terms of different levels of independence. At one extreme, full independence is assumed, as in the Petersen estimator (which does not use distance data); at the other extreme, independence only occurs in the limit as detection probability tends to one. Between the two extremes, there is a range of models, including those currently in common use, which have intermediate levels of independence. We show how this framework can be used to provide more reliable analysis of double-observer line transect data. We test the methods by simulation, and by analysis of a dataset for which true abundance is known. We illustrate the approach through analysis of minke whale sightings data from the North Sea and adjacent waters.     

Status,number, and monitoring of the common eider (<Emphasis Type="Italic">Somateria mollissima</Emphasis>) population in the barents sea and the White Sea

The breeding, molting, and wintering ranges of the common eider (Somateria mollissima) were mapped and described in the southern part of the Barents Sea region, including the White Sea population in the western White Sea and the Murmansk population in the northwestern White Sea and the southeastern Barents Sea. The present-day abundance was determined for both populations: since the 1950s, it was the highest in the 2000s for the entire period of ornithological monitoring in the region. Aerial survey from a helicopter is considered to be the only reliable method for estimation of the total population number. The best season for surveying the White Sea population is March–early April (wintering); the most favorable period for surveying the Murmansk population is August (postbreeding/molting). The population status of the common eider in the northern and eastern parts of the Barents Sea has not yet been evaluated. On Franz-Josef Land, the common eider is distributed sporadically, and its number was assessed using expert estimates. The current abundance of the common eider on Novaya Zemlya is unknown. Helicopter-based aerial survey performed in late summer during the postbreeding season is the only relevant method to obtain reliable estimates for the population of the common eider inhabiting Franz-Josef Land and Novaya Zemlya.     

Abundance estimation of long-diving animals using line transect methods

Line transect sampling is one of the most widely used methods for estimating the size of wild animal populations. An assumption in standard line transect sampling is that all the animals on the trackline are detected without fail. This assumption tends to be violated for marine mammals with surfacing/diving behaviors. The detection probability on the trackline is estimated using duplicate sightings from double-platform line transect methods. The double-platform methods, however, are insufficient to estimate the abundance of long-diving animals because these animals can be completely missed while the observers pass. We developed a more flexible hazard probability model that incorporates information on surfacing/diving patterns obtained from telemetry data. The model is based on a stochastic point process and is statistically tractable. A simulation study showed that the new model provides near-unbiased abundance estimates, whereas the traditional hazard rate and hazard probability models produce considerably biased estimates. As an illustration, we applied the model to data on the Baird's beaked whale (Berardius bairdii) in the western North Pacific.     

A hierarchical modeling framework for multiple observer transect surveys

Ecologists often use multiple observer transect surveys to census animal populations. In addition to animal counts, these surveys produce sequences of detections and non-detections for each observer. When combined with additional data (i.e. covariates such as distance from the transect line), these sequences provide the additional information to estimate absolute abundance when detectability on the transect line is less than one. Although existing analysis approaches for such data have proven extremely useful, they have some limitations. For instance, it is difficult to extrapolate from observed areas to unobserved areas unless a rigorous sampling design is adhered to; it is also difficult to share information across spatial and temporal domains or to accommodate habitat-abundance relationships. In this paper, we introduce a hierarchical modeling framework for multiple observer line transects that removes these limitations. In particular, abundance intensities can be modeled as a function of habitat covariates, making it easier to extrapolate to unsampled areas. Our approach relies on a complete data representation of the state space, where unobserved animals and their covariates are modeled using a reversible jump Markov chain Monte Carlo algorithm. Observer detections are modeled via a bivariate normal distribution on the probit scale, with dependence induced by a distance-dependent correlation parameter. We illustrate performance of our approach with simulated data and on a known population of golf tees. In both cases, we show that our hierarchical modeling approach yields accurate inference about abundance and related parameters. In addition, we obtain accurate inference about population-level covariates (e.g. group size). We recommend that ecologists consider using hierarchical models when analyzing multiple-observer transect data, especially when it is difficult to rigorously follow pre-specified sampling designs. We provide a new R package, hierarchicalDS, to facilitate the building and fitting of these models.     

Diet composition of polar bears in Svalbard and the western Barents Sea

We estimated both the numerical and biomass composition of the prey of polar bears (Ursus maritimus) from 135 opportunistic observations of kills in Svalbard and the western Barents Sea collected from March to October 1984-2001. By number, the prey composition was dominated by ringed seals (Phoca hispida) (63%), followed by bearded seals (Erignathus barbatus) (13%), harp seals (P. groenlandica) (8%) and unknown species (16%). However, when known prey were converted to biomass, the composition was dominated by bearded seals (55%), followed by ringed seals (30%) and harp seals (15%). Results indicated that bearded seals are an important dietary item for polar bears in the western Barents Sea. We believe that different patterns of space use by different bears may result in geographic variation of diet within the same population.     

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.     

The Effect of Animal Movement on Line Transect Estimates of Abundance

Line transect sampling is a distance sampling method for estimating the abundance of wild animal populations. One key assumption of this method is that all animals are detected at their initial location. Animal movement independent of the transect and observer can thus cause substantial bias. We present an analytic expression for this bias when detection within the transect is certain (strip transect sampling) and use simulation to quantify bias when detection falls off with distance from the line (line transect sampling). We also explore the non-linear relationship between bias, detection, and animal movement by varying detectability and movement type. We consider animals that move in randomly orientated straight lines, which provides an upper bound on bias, and animals that are constrained to a home range of random radius. We find that bias is reduced when animal movement is constrained, and bias is considerably smaller in line transect sampling than strip transect sampling provided that mean animal speed is less than observer speed. By contrast, when mean animal speed exceeds observer speed the bias in line transect sampling becomes comparable with, and may exceed, that of strip transect sampling. Bias from independent animal movement is reduced by the observer searching further perpendicular to the transect, searching a shorter distance ahead and by ignoring animals that may overtake the observer from behind. However, when animals move in response to the observer, the standard practice of searching further ahead should continue as the bias from responsive movement is often greater than that from independent movement.     

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.