The aggregation of polar cod (Boreogadus saida) in the deep Atlantic layer of ice-covered Amundsen Gulf (Beaufort Sea) in winter

During the Circumpolar Flaw Lead System Study (CFL, 2007–2008), large aggregations of polar cod were detected in winter in the Amundsen Gulf (Western Canadian Arctic) using the EK60 echosounder of the CCGS Amundsen research icebreaker. Biomass estimated over 10 months reached a maximum of 0.732 kg m⁻² in February. Aggregations were encountered only in the presence of an ice cover from December to April. The vertical extent of the aggregations was dictated by temperature and zooplankton prey distribution. In winter, polar cod generally occupied the relatively warm deep Atlantic Layer (>0°C), but a fraction of the densest aggregations occasionally followed zooplankton prey up into the cold Pacific Halocline (−1.6 to 0°C). The diel vertical migration of polar cod was precisely synchronized with the seasonally increasing photoperiod. Throughout winter, polar cod aggregations migrated to progressively deeper regions (from 220 to 550 m bottom depths) in response to increasing light intensity, presumably to avoid predation by visual predators such as the ringed seal. Comparing Amundsen Gulf and Franklin Bay indicates that the entrapment of polar cod in embayments during winter is an important mechanism to provide marine mammal predators with dense concentrations of their main prey within their diving range.     

Characteristics of under-ice swarming of polar cod Boreogadus saida (Gadidae) in the Central Arctic Ocean

Phenological observations on under-ice swarming behavior of Polar cod Boreogadus saida were performed during the winter period 2009–2010 in the Canadian sector of the Arctic Ocean on the North Pole NP-37 drifting station. These observations and ichthyologic collections were the basis of the present study. The schools were mostly formed by the immature specimens (TL = 75–169 mm; 1+ to 4+ age), the dominating group was presented by fish of 2+ age (86–94% of total abundance). The driving factors of such large under-ice swarming of Polar cod B. saida in the Arctic are discussed, particularly, the interactions of Polar cod development and the ice drift in the studied region. This species inhabits the overcooled water (?1.8°C) under the ice, so the scheme of the antifreeze agent producing in the fish blood is considered. We make assumptions about under ice migrations of Polar cod swarming.     

Sub-lethal levels of waterborne petroleum may depress routine metabolism in polar cod Boreogadus saida (Lepechin, 1774)

Petroleum-related activities in Arctic waters are rapidly increasing parallel to the ongoing thinning of the Arctic sea ice. As part of a series of studies on petroleum-induced stress in polar cod Boreogadus saida, we tested the effects of acute (~60 min) and chronic (4 weeks) exposure to the water soluble fraction (WSF) of petroleum on whole body metabolism inferred from measurements of oxygen consumption rates. The exposure of polar cod to WSF leads to a statistically significant depression in routine metabolism in the order Control (0.260 mg O₂ g fish⁻¹ h⁻¹; N = 6) > Chronic (0.191 mg O₂ g fish⁻¹ h⁻¹; N = 6) > Acute (0.110 mg O₂ g fish⁻¹ h⁻¹; N = 2), decoupling of routine metabolism and body mass but possibly also to a partial metabolic compensation after 4 weeks of exposure. The results are reviewed in context with similar studies on Antarctic and non-polar fishes.     

Respiration of juvenile Arctic cod (Boreogadus saida): effects of acclimation,temperature, and food intake

Oxygen consumption (VO₂) of juvenile Arctic cod (Boreogadus saida) was investigated at low tempera tures (six temperatures; range -0.5 to 2.7°C). Small (mean wt. 6–8 g) and large (mean wt. 14 g) fish were acclimated, or adjusted to a constant temperature (0.4°C), for 5 months and then tested for metabolic cold adaptation (elevated metabolic rates in polar fishes). Short-term (2 weeks) acclimated fish showed elevated VO₂ similar to previously established values for polar fishes, but there was no such evidence after longterm acclimation. Long-term acclimation caused VO₂ values to drop significantly (from 86.0 to 46.5 mg O₂·kg^–1·h^–1, at 0.4°C), which showed that metabolic cold adaptation was a phenomenon caused by insufficien: acclimation time for fish in respiration experiments. We also measured the effects of temperature and feeding on VO₂. A temperature increase of 2.3°C resulted in relatively large increases in VO₂ for both longand short-term acclimated fish (Q₁₀ = 6.7 and 7.1, respectively), which suggests that metabolic processes are strongly influenced by temperature when it is close to zero. Feeding individuals to satiation caused significant increases in VO₂ above pre-fed values (34–60% within 1–2 days after feeding). Respiration budgets of starved and fed Arctic cod at ambient temperatures in Resolute Bay N.W.T., Canada, were used to model annual respiration costs and potential weight loss. Low respiration costs for Arctic cod at ambient temperatures result in high growth efficiency during periods of feeding and low weight loss during periods of starvation.     

Size,age and diet of polar cod,Boreogadus saida (Lepechin 1773), in ice covered waters

Summary Polar cod (Boreogadus saida) associated with drifting sea-ice were collected in the western Barents sea and north of Svalbard with dip-nets while SCUBA-diving in 1986 and 1987. Length-frequency measurements and otolith-readings suggested that the specimens were either one or two years old. The diet of fish from the western Barents sea (first-year ice) consisted mainly of copepods (Calanus finmarchicus, Calanus glacialis) and the hyperiid amphipod Parathemisto libellula. Fish collected north of the Svalbard archipelago (multi-year ice) had a more diverse diet, in which P. libellula and the sympagic amphipod Apherusa glacialis contributed more to the total diet biomass than copepods.     

From polar night to midnight sun: photoperiod, seal predation, and the diel vertical migrations of polar cod (Boreogadus saida) under landfast ice in the Arctic Ocean

The winter/spring vertical distributions of polar cod, copepods, and ringed seal were monitored at a 230-m station in ice-covered Franklin Bay. In daytime, polar cod of all sizes (7–95 g) formed a dense aggregation in the deep inverse thermocline (160–230 m, −1.0 to 0°C). From December (polar night) to April (18-h daylight), small polar cod <25 g migrated into the isothermal cold intermediate layer (90–150 m, −1.4°C) at night to avoid visual predation by shallow-diving immature seals. By contrast, large polar cod (25–95 g), with large livers, remained below 180 m at all times, presumably to minimize predation by deep-diving mature seals. The diel vertical migration (DVM) of small polar cod was precisely synchronized with the light/dark cycle and its duration tracked the seasonal lengthening of the photoperiod. The DVM stopped in May coincident with the midnight sun and increased schooling and feeding. We propose that foraging interference and a limited prey supply in the deep aggregation drove the upward re-distribution of small polar cod at night. The bioluminescent copepod Metridia longa could have provided the light needed by polar cod to feed on copepods in the deep aphotic layers.     

Summer feeding ecology of harp seals (Phoca groenlandica) in relation to arctic cod (Boreogadus saida) in the Canadian high arctic

Summary The diet and feeding behaviour of harp seals, Phoca groenlandica, was examined in two high arctic locations. Fish otoliths were used to evaluate dietary composition and aspects of the population dynamics of the major prey species, arctic cod, Boreogadus saida. Harp seals, primarily adults, arrive in the high arctic in mid to late June and depart by early October. Their migration is undertaken specifically for feeding. Harp seals feed intensively on arctic cod, often occurring in dense multispecies aggregations in late summer. The average weight of harp seal stomach contents was high; glutted individuals contained as much as 6% of their body weight in food. Although arctic cod declined in abundance between years, size of cod ingested was similar between areas and years, and overlapped completely with cod taken by other marine mammals. Age/size segregation of arctic cod may account for poor representation of fish <3 years old in the seal diet. Widespread reproductive failure of arctic cod could have a profound influence on the energy balance of adult harp seals since there does not appear to be an alternate food source of equivalent energy value and abundance in arctic waters. Increasing harp seal populations will likely result in increased competition with a host of arctic cod predators, particularly ringed seals.     

Variability in the summer diets of juvenile polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the northeastern Chukchi and western Beaufort Seas

Polar cod (Boreogadus saida) is an important link between top predators and lower trophic levels in high-latitude marine ecosystems. Previous findings describe differences in its diet throughout the western Arctic; however, the causes of this variation are not well known. This study examined the diets of juvenile polar cod collected via demersal trawling methods over three summers in the northeastern Chukchi Sea (2010–2012) and one summer in the western Beaufort Sea (2011) to determine the amount of variability explained by biological, spatial, and interannual factors. Prey were identified, measured for length, and aggregated by percent mean weight into taxonomically coarse prey categories for analysis. Within seas, variation in juvenile polar cod diet composition was significantly related to body size, latitude, longitude, depth, and interannual (Chukchi Sea only) factors. Canonical correspondence analysis indicated body size was the most important factor contributing to the total variance in juvenile polar cod diet in the Chukchi and Beaufort Seas. Body size-based diet differences between the Chukchi and Beaufort Seas were evaluated using non-metric multidimensional scaling. This method revealed that similar-sized polar cod consumed similar-sized prey in both seas, but their diets were more benthically influenced in the Chukchi Sea and more pelagically influenced in the Beaufort Sea. Juvenile polar cod diet compositions vary by body size and region of inhabitance throughout their distribution. Here, we show that body size was the primary factor explaining variation in the summer diet of juvenile polar cod within the Chukchi and Beaufort Seas.     

Distribution of polar cod and age-0 fish in the U.S. Beaufort Sea

Little is known about species composition, distribution, and abundance of pelagic fish in the U.S. portion of the Beaufort Sea continental shelf and slope. To inventory the community and describe pelagic fish distributions relative to water characteristics, a systematic survey was conducted in August 2008. Acoustics (38 kHz), midwater trawling, and CTD casts were used to sample water depths from 20 to 500 m. Age-1+ polar cod (Boreogadus saida) was the dominant fish species, with peak densities of 155,000 # ha⁻¹ at bottom depths of 100–350 m. Age-0 fish (polar cod, sculpin (Cottidae family), and eelblenny (Lumpenus sp.)), dominated the pelagic biomass at bottom depths between 20 and 75 m, with peak densities of 160,000 # ha⁻¹, but were also found in surface waters at bottom depths >75 m. Age-1+ polar cod were associated with cold, saline waters likely of Chukchi Sea origin and mirrored published foraging distributions for beluga whales (Delphinapterus leucas). Conversely, age-0 fish were found in warm, fresher water, likely of ice melt and/or riverine origin, throughout the study area. This study provides a necessary baseline for the development of Arctic assessment surveys and management plans for polar cod.     

Trophic variability of Arctic fishes in the Canadian Beaufort Sea: a fatty acids and stable isotopes approach

Trophic ecology of most demersal Arctic fishes remains one of the major knowledge gaps for understanding food web dynamics and connectivity among ecosystems. In this study, fatty acids (FA) and stable isotopes (SI) were used to study the feeding ecology of seven species (n = 106) of the most abundant benthic fishes (eelpouts, sculpins and agonids) in the Canadian Beaufort Sea from shallow (20–75 m), slope (200–350 m) and deep (500–1000 m) habitats. Both FA and SI results revealed among- and within-species variability in diet composition. Correspondence analysis of FA signatures identified high within-species variability in diet, resulting in high overlap among species. Calanus-derived FA were present in all species (Calanus markers up to 13 % of total FA) and were particularly important in Ribbed Sculpin, Adolf’s and Longear Eelpout collected in deep habitats, suggesting a strong contribution of pelagic-derived FA to benthic fish communities. Incorporation of this signal in the benthos may result from either direct consumption of deep overwintering copepods (i.e., off-bottom feeding) or through detrital accumulation in benthic invertebrate prey. Mean SI values differed among species and indicated that a large range of trophic positions (δ¹⁵N varied from 14.09 to 17.71 ‰ for Canadian Eelpout and Adolf’s Eelpout, respectively) and carbon dietary sources are preyed upon (δ¹³C range from ?21.13 to ?23.85 ‰ for Longear Eelpout and Ribbed Sculpin, respectively). SI analyses suggested that most species examined were low- to mid-trophic generalist benthic carnivores, with the exception of Ribbed Sculpin, which was a low-trophic pelagic predator.     

Contamination of food by crude oil affects food selection and growth performance,but not appetite,in an Arctic fish,the polar cod (Boreogadus saida)

The polar cod (Boreogadus saida) is recognized as a key species in Arctic marine food webs and it may, therefore, be important for the transfer of xenobiotics from lower trophic levels to its main predators, birds and sea mammals. The present work examines the effects of foods contaminated with 200 or 400 ppm crude oil on food selection patterns and appetite-growth relationships in polar cod using X-radiography. It is shown that sexually mature polar cod consumed mixtures of uncontaminated and oil-contaminated foods, and did not show a reduced overall appetite as compared with fish provided with uncontaminated food only. Food selection was, however, influenced by both sex and individual appetite. Male fish selected uncontaminated food when appetite was low, whereas females ingested contaminated and uncontaminated foods equally, irrespective of appetite level. The ingestion of oil-contaminated food led to a significant depression in growth performance in both male and female fish. Food contaminated with oil at a concentration of 500 ppm was completely rejected by both sexes.     

Vertical segregation of age-0 and age-1+ polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) over the annual cycle in the Canadian Beaufort Sea

The offshore marine ecosystem of the Canadian Beaufort Sea faces the double pressure of climate change and industrialization. Polar cod (Boreogadus saida) is a pivotal forage species in this ecosystem, accounting for 95 % of the pelagic fish assemblage. Its vertical distribution over the annual cycle remains poorly documented. Hydroacoustic records from 2006 to 2012 were analysed to test the hypothesis that age-0 polar cod segregate vertically from larger congeners. Trawls and ichthyoplankton nets validated the acoustic signal. Fish length, weight, and biomass were estimated from new regressions of target strength and weight on standard length. Polar cod were vertically segregated by size in all months, with small age-0 juveniles in the epipelagic (<100 m) layer and larger age-1+ deeper in the water column. From December to March, the biomass of age-1+ peaked in a mesopelagic layer between 200 and 400 m. With increasing irradiance from April to July, the mesopelagic layer deepened and extended to 600 m. Starting in July, age-0 polar cod formed an epipelagic scattering layer that persisted until November. From September onward, age-0 left the epipelagic layer to join small age-1+ in the upper mesopelagic layer. Low biomass in the mesopelagic layer from February to September likely resulted from large polar cod settling on the seafloor to avoid diving marine mammals. Longer ice-free seasons, warmer sea-surface temperatures, or an oil spill at the surface would likely impact epipelagic age-0, while mesopelagic age-1+ would be vulnerable to an eventual oil plume spreading over and above the seafloor.     

Diets and body condition of polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the northern Bering Sea and Chukchi Sea

To understand trophic responses of polar cod Boreogadus saida (a key species in Arctic food webs) to changes in zooplankton and benthic invertebrate communities (prey), we compared its stomach contents and body condition between three regions with different environments: the northern Bering Sea (NB), southern Chukchi Sea (SC), and central Chukchi Sea (CC). Polar cod were sampled using a bottom trawl, and their potential prey species in the environment were sampled using a plankton net and a surface sediment sampler. Polar cod fed mainly on appendicularians in the NB and SC where copepods were the most abundant in the environment, while they fed on copepods, euphausiids, and gammarids in the CC where barnacle larvae were the most abundant species in plankton samples on average. The stomach fullness index of polar cod was higher in the NB and SC than CC, while their body condition index did not differ between these regions. The lower lipid content of appendicularians compared to other prey species is the most plausible explanation for this inconsistency.     

In-situ observations on the distribution and behavior of amphipods and Arctic cod (Boreogadus saida) under the sea ice of the High Arctic Canada Basin

The occurrence and behavior of sympagic amphipods and Arctic cod (Boreogadus saida) were studied in the High Arctic Canada Basin by diving under the ice at seven stations in summer 2002. Still images of video-transects were used to obtain animal abundances and information on the structure of the ice environment. Mean amphipod abundances for the stations varied between 1 and 23 individuals m^–2, with an increase towards the western part of the basin. The standard deviation within the 31–51 images analyzed per station was small (<1 individual m^–2). Gammarus wilkitzkii was found in low abundances, often hiding in small ice gaps. Small amphipods (Onisimus spp., Apherusa glacialis, and juveniles of all species) tended to move freely along the bottom of the floes. B. saida occurred in narrow wedges of seawater along the edges of melting ice floes at three stations in water depths of 10–50 cm and was never found under the ice. The fish occurred in schools of 1–28 per wedge. Fish were inactive and did not escape the approaching diver. Resting in the wedges may be a strategy to reduce energetic requirements and avoid predators.We dedicate this publication to Professor Dr. J. Lenz (Kiel University).     

Differentiation of western and eastern Baltic Sea cod stocks (Gadus morhua) by means of stable isotope ratios in muscles and otoliths

Stable isotope analysis of Baltic Sea cod from the Bornholm Sea (ICES‐SD 25) and western Baltic (Belt Sea, ICES‐SD 22) revealed significant differences in the δ¹⁵N and δ¹³C values of the dorsal muscle, as well as in the δ¹⁸O values of otoliths. The method pledges to become especially appropriate to the Baltic due to the high variability in oxygen isotope ratios associated with its estuarine nature.     

Fasting physiology of polar bears in relation to environmental change and breeding behavior in the Beaufort Sea

We examined the use of the ratio of serum urea to serum creatinine as a physiological biomarker of fasting to monitor temporal patterns in the feeding ecology of polar bears (Ursus maritimus). Blood was collected from 436 polar bears in the eastern Beaufort Sea during April and May of 1985–1986 and 2005–2006. The proportions of polar bears fasting were 9.6% in 1985, 10.5% in 1986, 21.4% in 2005, and 29.3% in 2006. We used stepwise logistic regression analysis to evaluate factors that could influence the binary response variable of fasting or not fasting. Significant predictor variables of fasting were: the 2005 and 2006 capture years, solitary adult male bears, and adult male bears that were accompanying an estrous female. The increased number of polar bears in a physiological fasting state from all sex, age, and reproductive classes in 2005 and 2006 corresponded with broad scale changes in Arctic sea ice composition, which may have affected prey availability. The higher proportion of adult males fasting from all years was attributed to spring breeding behavior.     

Unravelling complexities in benthic food webs using a dual stable isotope (hydrogen and carbon) approach

1. Stable carbon isotope studies have been an essential component of research regarding the contribution of methane (CH₄)-derived carbon to freshwater food webs and results have suggested that benthic macroinvertebrates in billabongs, streams and lakes may be partially, and in some instances, significantly 'fuelled' by methanotrophic biomass. However, the singular use of carbon isotopes can lead to ambiguous interpretations concerning the origin of carbon, especially in systems where phototrophs are likely to be using carbon respired sources and hence show more ¹³C-depleted values.
2. These uncertainties can be further resolved by the inclusion of additional isotopic data. Stable hydrogen isotopes are being increasingly used in food web studies with a marked advantage that sources may be isotopically distinct by one or two orders of magnitude greater than stable carbon or nitrogen, the isotopes most commonly used to delineate trophic interactions. By using hydrogen as a second biogeochemical tracer we provide further supportive evidence for the assimilation of methanotrophic microbial biomass by chironomid larvae.
3. Moreover, the hydrogen and carbon isotope values we found in chironomid tissues appear to reflect the original substrate used during methanogenesis; either acetate fermentation or carbonate reduction. Use of the former tends to result in relatively heavy carbon and light hydrogen isotope values due to kinetic isotope effects, whereas use of the latter results in relatively lighter carbon and heavier hydrogen isotope values.
4. We provide preliminary evidence to suggest that hydrogen and carbon isotope values in macroinvertebrates may be used to distinguish between CH₄ formation pathways and help to explain inter-depth and inter-specific differences between co-existing chironomid species found in the same lake.