Feeding of Polar cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> in the Kara Sea

In the summer–autumn period of 2007, 2010, and 2013, the peculiarities of Polar cod feeding Boreogadus saida in the Kara Sea were considered. In these years, the intensity of feeding of adult individuals was quite similar. Spatial and interannual differences in the composition of the food have been observed. Copepoda, Euphausiacea, and tunicates (Oikopleura sp.) dominated in the food of young fish. The main preys of adult individuals are Hyperiidea, Copepoda, Euphausiacea, and fish. As the length of Polar cod increases, the weight percentage of copepods decreases and the portion of fish increases.     

Dietary characteristics of co-occurring polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) and capelin (<Emphasis Type="Italic">Mallotus villosus</Emphasis>) in the Canadian Arctic,Darnley Bay

Reduction in sea ice due to climate change is expected to have a negative impact on habitat availability for Arctic marine fishes and induce range expansion of species from southern environments. Such an effect will likely be observed in the abundance of polar cod, Boreogadus saida (Lepechin, 1774), as well as interspecific interactions of this intermediate-level trophic taxon, particularly in more southerly fringing seas in the Arctic. Polar cod and capelin, Mallotus villosus (Müller, 1776), are pelagic, planktivorous forage fishes, which occupy similar dietary niches and are the primary prey of marine predators. Co-occurring polar cod and capelin were collected at seven stations in Darnley Bay, NT, during August 2013. Standard length (SL), used as a proxy for age, suggested that polar cod (mean ± 1 SD: 71.1 ± 10.3 mm) were predominantly age 1+ and capelin (96.2 ± 13.4 mm) were mostly age 2+. Stomach content analyses indicated that both species feed extensively on calanoid copepods (Calanus hyperboreus, C. glacialis, Metridia longa) and amphipods (Themisto libellula). There was high dietary overlap between capelin and polar cod, evidenced by Schoener’s index (0.80). Additionally the quantity of dietary items, biomass and energetic content consumed differed among size classes in both capelin (SL, 70.5–132.0 mm) and polar cod (SL, 42.1–114.4 mm). This study illustrates that the diets of these sympatric forage fishes in an Arctic ecosystem are very similar, indicating a high potential for interspecific competition as the sub-Arctic capelin expands its range into Arctic regions with climate change.     

Temperature-dependent growth and behavior of juvenile Arctic cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) and co-occurring North Pacific gadids

The thermal sensitivity of Arctic fish species is poorly understood, yet such data are a critical component of forecasting and understanding ecosystem impacts of climate change. In this study, we experimentally measured temperature-dependent growth and routine swim activity in the juvenile stage of two Arctic gadids (Arctic cod, Boreogadus saida and saffron cod, Eleginus gracilis) and two North Pacific gadids (walleye pollock, Gadus chalcogrammus and Pacific cod, Gadus macrocephalus) over a 6-week growth period across five temperatures (0, 5, 9, 16 and 20 °C). Arctic cod demonstrated a cold-water, stenothermic response in that there was relatively high growth at 0 °C (0.73 % day^?1), near-maximal growth at 5 °C (1.35 % day^?1) and negative impacts on activity, growth and survival at 16 °C. In contrast, saffron cod demonstrated a warmer-water, eurythermic response, and temperature had a positive effect on growth and condition beyond 16 °C. However, despite these distinct thermal responses, walleye pollock and Pacific cod grew 2–3 times faster than Arctic gadids across a relatively broad temperature range above 5 °C. These results, coupled with possible northward expansion by both Pacific cod and walleye pollock, suggest Arctic cod are highly vulnerable to continued climate change in the Arctic, especially in coastal areas of the Beaufort and Chukchi Seas where temperatures already exceed 14 °C in the summer growth period.     

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.     

Variation in the diet of Arctic Cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) in the Pacific Arctic and Bering Sea

Arctic cod, Boreogadus saida (Lepechin, 1774), is a nodal species in Arctic marine foodwebs as an important prey of many birds, marine mammals, and other fishes, as well as an abundant predator of zooplankton and epibenthic fauna. We examined the summer diet of Arctic cod across a latitudinal gradient extending from the southern limit of their distribution in the eastern Bering Sea to the northern margins of the eastern Chukchi Sea (ECS) continental shelf. Specimens were collected from demersal and pelagic trawls conducted between 1999 and 2012, and across a range of predator sizes (3–26 cm). Arctic cod diets vary with body size and between regions within the study area, and appear to vary between years in the eastern Bering Sea, indicating opportunistic feeding habits. Constrained Analysis of Principal Coordinates was conducted on ECS demersal samples and revealed consumption of fish and decapod crustacea were positively correlated with Arctic cod length while consumption of euphausiids and copepods had the opposite relationship. The demersal Arctic cod diet in the northern latitudes of the ECS was dominated by copepod consumption (47% by weight, %W), but copepods were less important (12–26%W) in the central and southern latitudes of the ECS and in the northern and eastern Bering Sea—areas where diets were more varied in their composition. High levels of variation in the diet of Arctic cod highlights the need to monitor Arctic cod diets to identify consistently dominant prey types and potential future changes to trophic relationships related to climate change or increasing anthropogenic activity.     

Contrasting the early life histories of sympatric Arctic gadids <Emphasis Type="Italic">Boreogadus saida</Emphasis> and <Emphasis Type="Italic">Arctogadus glacialis</Emphasis> in the Canadian Beaufort Sea

The early life stages of Boreogadus saida and Arctogadus glacialis are morphologically similar, making it difficult to assess differences in their ecological niche. The present study documented for the first time the early life stage ecology of A. glacialis, compared it to that of B. saida, and identified the factors separating the niches of the two sympatric species. The 10,565 larval gadids collected in the Beaufort Sea from April to August of 2004 and 2008 were identified to species either directly by genetics and/or otolith nucleus size, or indirectly with a redistribution procedure. Between 8.0 and 8.7 % of all gadids were assigned to A. glacialis. Larvae of A. glacialis were longer at hatch and experienced lower mortality rates than those of B. saida. The two species shared similar spatiotemporal and vertical distributions, hatching season, and growth rate. Under the ice, feeding incidence of B. saida was low (14 %) relative to A. glacialis (88 %). At lengths <15 mm, both species specialized on different prey. The diet of fish >15 mm overlapped (Schoener’s index = 0.7), with Calanus glacialis and C. hyperboreus providing >50 % of the carbon intake of both species. The higher mortality in B. saida may be explained by the smaller size at age from hatching to metamorphosis and a lower under-ice feeding incidence. The early larval stage appears to be the key period of niche divergence between the two species.     

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.     

<Emphasis Type="Italic">Boreogadus saida</Emphasis> (Lepechin) (Gadidae): a review of its metazoan parasite fauna from Greenland,eastern Canada,Alaska and the Russian Arctic

The metazoan parasite fauna of 50 specimens of Boreogadus saida (Lepechin) (Gadidae) from eastern Greenland is very similar to those of previous studies of the parasite fauna of B. saida from Greenland, eastern Canada, Alaska and the Russian Arctic. The digeneans Hemiurus levinseni, Derogenes varicus and Lecithaster gibbosus and cestode larvae were found at most stations. Single specimens of the nematode larvae Anisakis simplex were found at four stations. A comparison of the distribution of the larvae of Contracaecum sp. and Hysterothylacium sp. is difficult due to a possible confusion of the two genera. Most of the metazoan parasites of B. saida are generalist species found in several fish families. Boreogadus saida acquires most of its endoparasites by eating pelagic crustaceans, mainly copepods and amphipods. It plays an important role in the arctic ecosystem and its parasites are transferred to predatory fish, birds and mammals through the food web.     

On genetic differentiation of the Pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> tilesius, 1810 (Gadiformes: Gadidae)

The variability of the Gmo3, Gmo34, Gmo35, and Pgmo32 DNA microsatellite loci in Pacific cod Gadus macrocephalus samples from different areas of the North Pacific was analyzed. The data obtained show that Pacific cod from the southern Kuril Islands significantly differs from the populations of the Bering Sea, the Sea of Okhotsk, and the coastal waters of Canada (the microsatellite loci Gmo3 and Pgmo32 bear the highest differentiating capacity). Despite the significant geographical remoteness of these areas from one another, the above three Pacific cod populations exhibit a high degree of similarity (I = 0.997–0.999).     

New encounters in Arctic waters: a comparison of metabolism and performance of polar cod (<Emphasis Type="Italic">Boreogadus saida</Emphasis>) and Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) under ocean acidification and warming

Oceans are experiencing increasing acidification in parallel to a distinct warming trend in consequence of ongoing climate change. Rising seawater temperatures are mediating a northward shift in distribution of Atlantic cod (Gadus morhua), into the habitat of polar cod (Boreogadus saida), that is associated with retreating cold water masses. This study investigates the competitive strength of the co-occurring gadoids under ocean acidification and warming (OAW) scenarios. Therefore, we incubated specimens of both species in individual tanks for 4 months, under different control and projected temperatures (polar cod: 0, 3, 6, 8 °C, Atlantic cod: 3, 8, 12, 16 °C) and PCO₂ conditions (390 and 1170 µatm) and monitored growth, feed consumption and standard metabolic rate. Our results revealed distinct temperature effects on both species. While hypercapnia by itself had no effect, combined drivers caused nonsignificant trends. The feed conversion efficiency of normocapnic polar cod was highest at 0 °C, while optimum growth performance was attained at 6 °C; the long-term upper thermal tolerance limit was reached at 8 °C. OAW caused only slight impairments in growth performance. Under normocapnic conditions, Atlantic cod consumed progressively increasing amounts of feed than individuals under hypercapnia despite maintaining similar growth rates during warming. The low feed conversion efficiency at 3 °C may relate to the lower thermal limit of Atlantic cod. In conclusion, Atlantic cod displayed increased performance in the warming Arctic such that the competitive strength of polar cod is expected to decrease under future OAW conditions.     

Population genetic analysis of Euro-Arctic polar cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> suggests fjord and oceanic structuring

Polar cod, Boreogadus saida, is a key species in Arctic marine ecosystems; however, its genetic population structure is largely undescribed. The population genetic structure of 472 B. saida specimens among nine locations in the north-east Atlantic was revealed using 12 microsatellite loci. Pairwise F _ST comparisons showed significant population differentiation between B. saida sampled inside fjords in Svalbard and north-east Greenland, as compared to B. saida from the shelf. The observed genetic variation was not a function of isolation by distance, and it is speculated that B. saida populations inhabiting fjords may have become reproductively isolated from shelf-dwelling B. saida during the last post-glacial recolonization.     

A simple molecular approach to distinguish between two Arctic gadoid fishes <Emphasis Type="Italic">Arctogadus glacialis</Emphasis> (Peters, 1874) and <Emphasis Type="Italic">Boreogadus saida</Emphasis> (Lepechin, 1774)

Correct phenotypical identification of and discrimination between Boreogadus saida and Arctogadus glacialis is challenging, especially for larvae and young fish. We propose the use of a single microsatellite genetic marker, Gmo8, to distinguish between the two gadoid fish species. Amplified allele frequencies differed considerably, with B. saida (n = 97) being almost exclusively monomorphic at this locus, whereas A. glacialis (n = 136) is highly polymorphic. There was a clear separation between the amplified allele ranges for the two species. The species specific properties of Gmo8 enables the use of this marker to distinguish between B. saida and A. glacialis.     

Atlantic cod <Emphasis Type="Italic">Gadus morhua</Emphasis> (Gadiformes: Gadidae) from the Gulf of Ura: Morphobiological characteristic

Analysis of variation of biological and morphometric characteristics of cod Gadus morhua from the Gulf of Ura-a water area in the composition of Motovskii Bay of the Barents Sea was performed. The material was collected from 1999 to 2006 during spawning of the Atlantic cod. Parameters such as length, weight, sex composition, maturity stage, age, otolith structure, and variation of plastic characters were analyzed. It was shown that the Gulf of Ura, parallel to individuals of junior age groups, is inhabited by cod in the spawning state aged 5–6 months and older with coastal and Atlantic types of otolith.     

Embryonic development of the pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> (Gadidae)

Incubation of pacific cod eggs was divided into eight series, in which temperatures were set at −0.04°C to +4.03°C and warm and cold conditions alternated. The morphological changes that took place during the embryogenesis were described in detail using the results of the incubation. Twenty-two morphological characters that could be identified easily and that characterized the morphogenesis were defined in the course of development. The results of the incubation and data from the literature showed that the duration of the embryonic period in the Pacific cod’s lifecycle grew exponentially as water temperature decreased. It was found during the experiment that developing cod eggs survived low water temperatures up to freezing, as well as abrupt warming or cooling (over 3°C). According to the widely accepted Rass scale, the first stage of the Pacific cod embryogenesis takes 21% of its total duration, the second stage 23%, the third, 17%, and the fourth, 39%. However, at a temperature below 0°C, the relative duration of the stages of cleavage and embryonic shield was slightly shortened, whereas the mature embryo stage extended to almost half of the embryogenesis period. A more comprehensive analysis of temperature effects on embryogenesis revealed that the reduction of the rate of embryogenesis upon a temperature decrease occurred mostly at later stages of embryo growth. Modeling of development using defined morphological characters showed that the duration of embryogenesis grew linearly as the incubation temperature dropped in the first half of the embryogenesis and exponentially in the second half. A function was selected that described the obtained results most satisfactorily and that could be used for estimating the duration of the entire embryogenesis or any its stages within the range of water temperatures typical for Pacific cod.     

Occurrence of a gelatinous predator (<Emphasis Type="Italic">Cyanea capillata</Emphasis>) may affect the distribution of <Emphasis Type="Italic">Boreogadus saida</Emphasis>, a key Arctic prey fish species

Although the Arctic cod (Boreogadus saida) has a pan-Arctic distribution, little is known about its occurrence in near-shore waters where this species is the principal prey for seabirds, marine mammals and other fish. Published research describes the scyphomedusa Cyanea capillata as an Arctic cod predator, and this paper presents observations from long-term investigations using active hydroacoustics that suggest the Arctic cod avoided C. capillata in two small bays of Cornwallis Island (Canadian High Arctic archipelago). Distribution patterns in echograms suggested that features such as boundary layer fronts restricted jellyfish movements and Arctic cod were often abundant on the side of fronts where C. capillata were absent. Thus, habitat partitioning allowed Arctic cod to share habitat with its predator, albeit exceptions to this sharing occurred when jellyfish abundance was high and Arctic cod were displaced. Thus, if a warmer Arctic triggers an increase in C. capillata abundance, it is possible that small-scale aspects of Arctic cod distribution could be affected. This in turn could have significant ripple effects within the Arctic food web, an additional and previously unrecognized consequence of climate change.     

Under-ice distribution of polar cod <Emphasis Type="Italic">Boreogadus saida</Emphasis> in the central Arctic Ocean and their association with sea-ice habitat properties

In the Arctic Ocean, sea-ice habitats are undergoing rapid environmental change. Polar cod (Boreogadus saida) is the most abundant fish known to reside under the pack-ice. The under-ice distribution, association with sea-ice habitat properties and origins of polar cod in the central Arctic Ocean, however, are largely unknown. During the RV Polarstern expedition ARK XXVII/3 in the Eurasian Basin in 2012, we used for the first time in Arctic waters a Surface and Under Ice Trawl with an integrated bio-environmental sensor array. Polar cod was ubiquitous throughout the Eurasian Basin with a median abundance of 5000 ind. km^?2. The under-ice population consisted of young specimens with a total length between 52 and 140 mm, dominated by 1-year-old fish. Higher fish abundance was associated with thicker ice, higher ice coverage and lower surface salinity, or with higher densities of the ice-amphipod Apherusa glacialis. The fish were in good condition and well fed according to various indices. Back-tracking of the sea-ice indicated that sea-ice sampled in the Amundsen Basin originated from the Laptev Sea coast, while sea-ice sampled in the Nansen Basin originated from the Kara Sea. Assuming that fish were following the ice drift, this suggests that under-ice polar cod distribution in the Eurasian Basin is dependent on the coastal populations where the sea-ice originates. The omnipresence of polar cod in the Eurasian Basin, in a good body condition, suggests that the central Arctic under-ice habitats may constitute a favourable environment for this species survival, a potential vector of genetic exchange and a recruitment source for coastal populations around the Arctic Ocean.     

Complete mitochondrial genome sequences of the Arctic Ocean codfishes <Emphasis Type="Italic">Arctogadus glacialis</Emphasis> and <Emphasis Type="Italic">Boreogadus saida</Emphasis> reveal oriL and tRNA gene duplications

We have determined the complete mitochondrial genome sequences of the codfishes Arctogadus glacialis and Boreogadus saida (Order Gadiformes, Family Gadidae). The 16,644 bp and 16,745 bp mtDNAs, respectively, contain the same set of 37 structural genes found in all vertebrates analyzed so far. The gene organization is conserved compared to other Gadidae species, but with one notable exception. B. saida contains heteroplasmic rearrangement-mediated duplications that include the origin of light-strand replication and nearby tRNA genes. Examination of the mitochondrial control region of A. glacialis, B. saida, and four additional representative Gadidae genera identified a highly variable domain containing tandem repeat motifs in A. glacialis. Mitogenomic phylogeny based on the complete mitochondrial genome sequence, the concatenated protein-coding genes, and the derived protein sequences strongly supports a sister taxa affiliation of A. glacialis and B. saida.     

Embryonic and early larval development of White Sea cod <Emphasis Type="Italic">Gadus morhua marisalbi</Emphasis> (Gadidae)

Embryonic and early larval development of White Sea cod Gadus morhua marisalbi are described. The study is conducted on live material obtained after spawning of fishes in the tanks and based on artificial insemination of eggs. Dynamics of contact and mobility properties of cells in vivo and in vitro (in cell culture) is assessed. Discrete chronological parameters of the morphogenetic movements are determined. Early ontogeny of the White Sea cod is very similar to that in G. morhua of the Northeast Atlantic. However, the former subspecies differs in larger egg diameter (on average, 1.50 mm), larger buoyancy (at a salinity up to 24.22‰), and a resistance to negative temperature (?1.8°C).