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.     

Schooling behaviour of arctic cod,Boreogadus saida,in relation to drifting pack ice

Synopsis Concentrations of arctic cod were detected with a hydroacoustic system in Resolute Bay, NWT during 2 weeks in August of 1986. Fish biomass within the bay was about 30 t. The fish were feeding primarily on amphipods, which were abundant. When the daily location of the schools was examined in relation to the extent and position of drifting pack ice, a pattern emerged suggesting that the distribution of the fish was influenced by the amount and location of ice cover. If the bay was relatively ice-free, the density of schooling cod was high and the size of the schools, as 2-dimensional surface area, was generally small. When ice covered the bay, density within the schools was lower and they occupied more area. Arctic cod were most dispersed after the bay had been filled with pack ice for several days. It is postulated that this behaviour is a response to potential predation by seabirds and marine mammals.     

Polar cod, Boreogadus saida (Gadidae), show an intermediate stress response between Antarctic and temperate fishes

To examine whether the attenuated stress response observed in Antarctic notothenioid fishes is a specialism for life in sub-zero waters, the polar cod, Boreogadus saida, and the temperate shorthorned sculpin, Myoxocephalus scorpius, were subjected to various stress treatments. Activity stress in both species had no effect on plasma catecholamine and cortisol levels, splenic mass, and on the haematological variables in B. saida. In contrast, heat stress caused a significant rise in circulating noradrenaline and adrenaline levels in B. saida, accompanied by a significant increase in haematocrit and haemoglobin concentrations, at constant plasma cortisol levels, red blood cell count and splenic mass. A concomitant rise in blood lactate concentrations indicated that heat-stressed B. saida were hypoxaemic. The capacity to synthesise catecholamines in B. saida was 38% of the value in M. scorpius, but similar to the values for Antarctic notothenioids. The lack of any adrenergic response to activity stress suggests that dominance of cholinergic control of the cardiovascular system may not be restricted to Antarctic notothenioids. Rather, the stress response in B. saida appears to be intermediate between Antarctic and temperate teleosts, in keeping with their relatively recent occupation of cold Arctic waters.     

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.     

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.     

Sourcing fatty acids to juvenile polar cod (Boreogadus saida) in the Beaufort Sea using compound-specific stable carbon isotope analyses

Juvenile polar cod (Boreogadus saida) are often found in close association with sea ice and represent an important trophic link in the Arctic food web. However, the proportional contribution of sea ice algal production via the sympagic food web to the diet of polar cod is unknown. To estimate the proportional contribution of fatty acids (FAs) from sea ice-derived particulate organic matter (i-POM) to the diet of juvenile polar cod, we used FA profiling and compound-specific stable carbon isotope analysis of individual FAs from juvenile polar cod collected from three regions in the Beaufort Sea. The δ¹³C values of the FAs 14:0, 16:4n-1, 18:0, 20:5n-3 and 22:6n-3 in the polar cod were found to most strongly resemble pelagic POM rather than i-POM. Results from isotope-mixing models using diatom FA markers indicated that the proportional contribution of FAs from i-POM to juvenile polar cod was ≤2 %, which suggests that juvenile polar cod had not sourced their FAs from i-POM. Thus, changes in sea ice coverage due to environmental change may not affect juvenile polar cod in regard to nutrients such as FAs but may still affect their populations by reducing critical shelter from predators.     

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.     

Gastric evacuation rates and daily rations of Arctic cod (Boreogadus saida) at low temperatures

Gastric evacuation rates were determined for different sizes of Arctic cod (Boreogadus saida) at sub-zero temperatures (−1.4 and −0.5°C). These temperatures represent ambient conditions for Arctic cod in the Canadian high Arctic. Evacuation half-times, the time required for half of the content of the stomach to be evacuated, were longer (36–70 h; mean=51 h) than those reported in studies carried out on other fish species. Gastric evacuation rates at low temperatures were equal to, or below, those predicted by extrapolation from experiments conducted at higher temperatures. There were no significant differences in evacuation rates among fish size-groups or diets, but evacuation rates were slower for fish that had been starved prior to experiments. Estimated daily rations for Arctic cod in Resolute Bay, N.W.T., were 0.51% body weight for small fish (4.5 g) and 1.13% body weight for large fish (51 g). Slow stomach evacuation rates at low temperatures may limit daily food intake when food is seasonally abundant. This may contribute to slow growth rates and limited maximum size of Arctic cod in Canadian high Arctic waters. Received: 14 July 1997 / Accepted: 15 November 1997     

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.     

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.     

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).     

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.     

Aspects of reproductive biology of wild-caught polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) from Svalbard waters

Polar cod (Boreogadus saida) is considered a key species in the Arctic marine ecosystems. Yet detailed or even basic knowledge regarding its biology and adaptations, especially during the polar night, is in many cases poor. Data are presently unavailable in Western literature on the gonad development of polar cod and its reproductive biology in wild specimens. Accordingly, gonad development of wild-caught polar cod from fjords of the Svalbard archipelago was studied across seasons (April, August, September, November and January). Histological analyses of polar cod showed strong indication of a group-synchronous oocyte development with determinate fecundity and iteroparity. Females started gonadal development prior to April and had not yet reached the final stage of maturation in January. Testes matured more rapidly, with males ready to spawn in January. Furthermore, our data show that polar cod were able to reach sexual maturity at age 1+. Based on our data and previous reports, we hypothesise that polar cod is a total spawner.     

Population Structure and Growth of Polar Cod Boreogadus saida in the Sea of Laptev

Journal of Ichthyology - The population structure and growth rate of polar cod Boreogadus saida in the Laptev Sea are analyzed based on the materials collected in the summer–autumn period of...     

Environmental confirmation of multiple ice age refugia for Pacific cod, Gadus macrocephalus

The concept of species surviving through quaternary climatic extremes by retreating to glacial refugia, and then evolving genetically during re-population movements of the following interglacial, has been in the literature for over 40 years. Recently, advances in genetic analysis have enabled this concept to be validated and theories regarding population expansions and contractions to be built. For the major Northern Hemisphere species of cod, Gadus morhua (Atlantic cod) and Gadus macrocephalus (Pacific cod), genetic analysis has suggested retreat to separate refugia on both sides of their respective ocean basins during the last glacial period. Ecosystem niche modelling has previously confirmed that environmental conditions during the last glacial were compatible with the existence of these separate refugia for Atlantic cod. Here it is shown that such modelling also confirms a reduced core glacial distribution for G. macrocephalus, but probable refugia on either side of the Pacific. Existing mitochondrial DNA analyses suggest two separate glacial populations in the northwest Pacific, which modelling confirms, with predicted separate marine refugia in the land-locked Sea of Japan basin and the Sea of Okhotsk. Existing mitochondrial DNA for the northeast Pacific populations is less conclusive regarding whether there were one or two separate refugia off this coast, and their location. Using environmental niche models this study shows the glacial NE Pacific environment could support two marine refugia, one centred in the Aleutians/Gulf of Alaska and the other off British Columbia. The intervening Cordilleran Ice Sheet, and the glacially sub-aerial and ice-free Queen Charlotte Islands shelf, is hypothesised to have constrained exchange between glacial stocks either side of the Islands. The postulated southern marine refugium is off-shore from an established terrestrial refugium, suggesting greater dependence of species and ecosystems during environmental change. An earth system approach to evolutionary change could enhance understanding of past and future ecosystems.     

Gender Specific Reproductive Strategies of an Arctic Key Species (Boreogadus saida) and Implications of Climate Change

The Arctic climate is changing at an unprecedented rate. What consequences this may have on the Arctic marine ecosystem depends to a large degree on how its species will respond both directly to elevated temperatures and more indirectly through ecological interactions. But despite an alarming recent warming of the Arctic with accompanying sea ice loss, reports evaluating ecological impacts of climate change in the Arctic remain sparse. Here, based upon a large-scale field study, we present basic new knowledge regarding the life history traits for one of the most important species in the entire Arctic, the polar cod (Boreogadus saida). Furthermore, by comparing regions of contrasting climatic influence (domains), we present evidence as to how its growth and reproductive success is impaired in the warmer of the two domains. As the future Arctic is predicted to resemble today''s Atlantic domains, we forecast changes in growth and life history characteristics of polar cod that will lead to alteration of its role as an Arctic keystone species. This will in turn affect community dynamics and energy transfer in the entire Arctic food chain.     

Size, growth and reproductive biology of the giant mudskipper, Periophthalmodon schlosseri (Pallas, 1770), in Malaysian waters

The size, growth and reproductive biology of the giant mudskipper, Periophthalmodon schlosseri, was studied in the inter‐tidal mudflat of a mangrove swamp along a 0.4–0.6 × 5 km stretch of the Sepang coastal waters, Selangor Malaysia. The size ranges of the giant mudskipper specimens caught in the study area were between 10–27.5 cm TL (20.6 ± 3.1 cm) for males and 14.5–28.5 cm TL (21.9 ± 2.3 cm) for females. Male to female sex ratio during the study was 1 : 1.1. The estimated von Bertalanffy growth parameters were L_∞ = 29 cm and K = 1.4 year^?1, with a growth performance index of Ø′ = 3.1. Otolith microstructure analysis revealed a strong correlation (Female: r = 0.920, n = 35; Male: r = 0.943, n = 41, P < 0.05) between the number of daily rings (number of days old) over the respective size ranges of male (8.5–19.5 cm SL) and female fish (11.8–22.3 cm SL). Four maturity stages were described to follow gonadal development based on external features; these stages were further validated through histological examination of the ovary and oocyte diameter measurements. The gonadal development cycle of P. schlosseri was asynchronous, as marked by the random mixture of early stages oocytes (diameter: 70.7 ± 14.9 μm) to matured stages (356.9 ± 20.2 μm), and had a prolonged spawning season extending from June to October. Monthly fluctuation of the gonadosomatic and hepatosomatic indices for this species was highly correlated (r = 0.126, P < 0.05).     

Oceanic micronektonic/macrozooplanktonic community structure and feeding in ice covered Antarctic waters during the winter (AMERIEZ 1988)

Summary Fifty-seven species of oceanic micronekton and macrozooplankton were collected under pack ice during the winter in the vicinity of the Weddell-Scotia Confluence with a modified opening-closing Tucker trawl. The majority of the 57 species did not vertically migrate and lived deeper during the winter than during the spring or fall. However, despite the short day length, several of the most common mesopelagic fish and crustaceans did migrate. Fish moved into shallower depths at night but apparently most did not continue into the near-freezing upper mixed layer, leaving that zone to the migratory crustaceans. In the upper 1000 m, the dominant species were, in order of decreasing biomass, Euphausia superba, the cnidarian Atolla wyvillei, the ctenophore Beroe sp., and the mesopelagic fish Electrona antarctica, Bathylagus antarcticus and Gymnoscopelus braueri. Thysanoessa macrura and Salpa thompsoni were biomass subdominants. The majority of the dominant species showed little seasonal differences in biomass. However, the biomass of gelatinous species varied considerably with A. wyvillei and Beroe sp. being most abundant and S. thompsoni least abundant during the winter. Incidence of food in the stomachs in several important species was low, suggesting a low impact on their Zooplankton prey. Specimens of S. thompsoni had high quantities of food in their guts but this species was uncommon so its net impact would also have been low. Euphausia superba and the three common mesopelagic fish had significantly lower stomach fullness ratings during the winter than during the fall, suggesting an overall decrease in feeding activity of dominant species during the winter.