The first demersal trawl survey of benthic fish and invertebrates in the Beaufort Sea since the late 1970s

This study represents the first demersal trawl survey of marine fishes and invertebrates in offshore waters of the Beaufort Sea since 1977. Species composition, distribution, and abundance of demersal fish and benthic invertebrates were assessed with standard methods and demersal trawl gear by the Alaska Fisheries Science Center. Fishes made up 6% of the total catch weight, and invertebrates made up the remaining 94% of the catch weight. A total of 32 species of fish were identified, two taxa were identified to genus and one to family, and 174 taxa of invertebrates were identified. The most abundant demersal fishes were polar cod (Boreogadus saida), eelpouts (Lycodes spp.), Bering flounder (Hippoglossoides robustus), and walleye pollock (Theragra chalcogramma). The most abundant invertebrates were notched brittle stars (Ophiura sarsi), snow crab (Chionoecetes opilio), mussels (Musculus spp.), and the mudstar (Ctenodiscus crispatus). We documented or confirmed extension to the known ranges of four species of fishes: walleye pollock, Pacific cod (Gadus macrocephalus), festive snailfish (Liparis marmoratus), and eyeshade sculpin (Nautichthys pribilovius). We also documented the presence of commercial-sized snow crab (Chionoecetes opilio), which has not previously been recorded in the North American Arctic.     

Feeding habits the Pacific cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> in oceanic waters of the Northern Kuril Islands and Southeast Kamchatka

This study examines the feeding habits of the Pacific cod Gadus macrocephalus in waters off the eastern coast of the northern Kuril Islands and southern Kamchatka. In November–December 1996, the cod primarily consumed fish, which made up 47.6% of the total food mass. The proportion of cephalopods, fishery offal discarded from fishing vessels, and decapods did not exceed 18.5, 17.4, and 12.2%, respectively. Among fishes, the main prey item of the cod was atka mackerel (15.4%); among cephalopods, octopus (16.8%); among fishery offal, heads of atka mackerel (14.2%); and among decapods, majid crabs (6.4%). The rather low percentage of walleye pollock (7.3%) in the cod diet was due to the decline of the east-Kamchatka walleye pollock stock.     

Isolation of twenty low stutter di- and tetranucleotide microsatellites for population analyses of walleye pollock and other gadoids

Fourteen tetra- and six dinucleotide microsatellites, which exhibit minimal stuttering following amplification via PCR were developed from walleye pollock Theragra chalcogramma . Most of these loci were isolated from a library enriched for tetranucleotide microsatellites by hybridization of genomic DNA to (gata)₇ oligonucleotides bound to streptavidin-coated paramagnetic beads. The average heterozygosity of these loci is ∼80%, and ranges from 53–95%. Mendelian inheritance was confirmed in five families, each consisting of a minimum of 10 or more offspring. Primer sets for all 20 loci were also evaluated in Arctic cod Boreogadus saida , Pacific cod Gadus macrocephalus , Pacific tomcod Microgadus proximus , sa.ron cod Eleginus gracilis , Pacific hake Merluccius productus , Atlantic cod Gadus morhua , haddock Melanogrammus aeglefinus , blue whiting Micromesistius poutassou , and European hake Merluccius merluccius . In each of these species, 3–19 primer sets amplified variable microsatellite loci. These loci, which exhibit little stutter and moderate to high variability, should be useful population markers in pollock and other gadoid fishes.     

Metabolic adjustments to varying protein intake in harbor seals (Phoca vitulina): evidence from serum free amino acids

Effects of varying dietary protein intake on serum free amino acid (FAA) concentrations were studied in harbor seals (Phoca vitulina) fed two different prey fish diets: either exclusively low-fat, high-protein walleye pollock (Theragra chalcogramma) or high-fat, relatively high-energy-density Pacific herring (Clupea pallasi). Significant differences in FAA concentrations and patterns were observed between the two diets. All essential amino acids (EAA), except methionine and phenylalanine, and two nonessential amino acids (NEAA), glycine and tyrosine, decreased when the diet was switched from herring to pollock and increased on switching back to herring. Both total EAA concentrations and EAA : NEAA ratios decreased with the elevated protein intake typical of a low-fat pollock diet, indicating an inverse correlation between EAA concentrations and dietary protein intake levels. We propose that differing dietary protein intake, caused by differences in macronutrient composition of the two prey fish species, induced a change in protein metabolism that was reflected in blood-circulating amino acids. These findings suggest that surveys of amino acid profiles may be useful to partially determine the protein metabolic status of harbor seals.     

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.     

Benthic-pelagic trophic interactions within the fish assemblage in the western Bering Sea shelf area according to stomach content analysis and ratios of C and N stable isotopes

The composition, abundance, diet and trophic status of zooplankton, bottom invertebrates, fish and nekton were analyzed based on the data collected by the staff of the TINRO-Center during complex bottom trawl catches on the Bering Sea shelf in the fall of 2004. The stomach contents of mass fish species were analyzed and the nitrogen and carbon isotopic composition of 36 mass species of plankton, benthos, nekton and nektobenthos, which together make up the basis of pelagic and bottom communities, was determined. It was found that zooplankton noticeably differ from benthic invertebrates in carbon isotopic composition: δ¹³C values in zooplankton varied from −20.3‰ to −17.9‰; in benthos—from −17.5‰to −13.0‰; and in fish—from −19.2‰ (juvenile walleye pollock) to −15.3‰ (saffron cod). The levels of ¹³C isotope in the tissues of fish depended mostly on the share of pelagic or benthic animals in their diet. δ¹⁵N values in the studied species ranged from 8.6‰ (in sea urchins) to 17.2‰ (in large Pacific cods), which corresponds to a trophic level of 2.8. Obviously the δ¹⁵N values reflect the degree of predation and generally show the ratio of primary, secondary and tertiary consumers in a fish’s diet. Trophic interactions manifest a high degree of interdependence between benthic and pelagic communities (even without taking into account such lower components of the food web as phytoplankton, bacteria, and protozoa) occurring in most nektonic species that depend on both bottom and pelagic food.     

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.     

Using a baited camera to assess relative abundance of juvenile Pacific cod: Field and laboratory trials

Juvenile fishes are sometimes difficult to survey because they occur in a variety of structurally complex habitats that are not readily sampled with traditional trawl or seine gear. Laboratory experiments and field sampling were conducted to determine whether a baited-camera system could be used effectively to survey age-0 gadids. Pacific cod (Gadus macrocephalus Tilesius), saffron cod (Eleginus gracilis Tilesius), and walleye pollock (Theragra chalcogramma Pallas) all responded to bait bags presented in the laboratory, with increased general activity and approaches to the bait. Pacific cod responded most strongly, and remained in close proximity to the bait bag. In nearshore habitats of Kodiak, Alaska, age-0 gadids (mostly Pacific cod) were quickly attracted to a baited-camera system. Arrival rates stabilized within 15-min, and several metrics for Pacific cod abundance taken from the video records were closely correlated with the numbers collected in seine hauls. Highest correlations occurred with the total number of fish arriving in camera view during 15-min sets (NFA₁₅) and with the overall maximum number of fish observed at one time during the sets (MaxN_O). The time for first arrival of fish in view (TFA) was not correlated significantly with the numbers of fish in seine collections. NFA₁₅ was also useful for age-1+ saffron cod. Occasionally, the presence of large fish had a negative effect on assessments of age-0 fish, but the impact was important only when the large fish were present continuously. The baited-camera system performed equally well in seagrass, kelp and open habitats, and can be used to rapidly assess the relative abundance of Pacific cod and selected other species in a wide range of habitats in shallow and deep water.     

Mitochondrial genomics of gadine fishes: implications for taxonomy and biogeographic origins from whole-genome data sets.

Phylogenetic analysis of 13 substantially complete mitochondrial DNA genome sequences (14,036 bp) from 10 taxa of gadine codfishes and pollock provides highly corroborated resolution of outstanding questions on their biogeographic evolution. Of 6 resolvable nodes among species, 4 were supported by >95% of bootstrap replications in parsimony, distance, likelihood, and similarly high posterior probabilities in bayesian analyses, one by 85%-95% according to the method of analysis, and one by 99% by one method and a majority of the other two. The endemic Pacific species, walleye pollock (Theragra chalcogramma), is more closely related to the endemic Atlantic species, Atlantic cod (Gadus macrocephalus), than either is to a second Pacific endemic, Pacific cod (Gadus macrocephalus). The walleye pollock should thus be referred to the genus Gadus as originally described (Gadus chalcogrammus Pallas 1811). Arcto-Atlantic Greenland cod, previously regarded as a distinct species (G. ogac), are a genomically distinguishable subspecies within pan-Pacific G. macrocephalus. Of the 2 endemic Arctic Ocean genera, Polar cod (Boreogadus) as the outgroup to Arctic cod (Arctogadus) and Gadus sensu lato is more strongly supported than a pairing of Boreogadus and Arctogadus as sister taxa. Taking into consideration historical patterns of hydrogeography, we outline a hypothesis of the origin of the 2 endemic Pacific species as independent but simultaneous invasions through the Bering Strait from an Arcto-Atlantic ancestral lineage. In contrast to the genome data, the complete proteome sequence (3830 amino acids) resolved only 3 nodes with >95% confidence, and placed Alaska pollock outside the Gadus clade owing to reversal mutations in the ND5 locus that restore ancestral, non-Gadus, amino acid residues in that species.     

The diets of polar cod (Boreogadus saida) from August 2008 in the US Beaufort Sea

Polar cod (Boreogadus saida) play an integral part in the Arctic ecosystems linking the upper and lower trophic levels. Though their estimated biomass is considerable, recent knowledge of their diets in the US Beaufort Sea is sparse. Collections of polar cod from the US Beaufort Sea were made during August 2008 using demersal and pelagic trawls. Polar cod diet composition was quantified as percent prey weight, percent prey count, and frequency of occurrence of prey. The diet composition between the demersal- and pelagic-captured cod showed differences in all these categories. Polar cod captured in the demersal nets primarily fed on fish (by weight), and pelagic cod primarily fed on copepods (frequency of occurrence) and euphausiids (by weight). In general, these dominant preys are different than what has been reported in other studies describing polar cod diets.     

Quantifying food web interactions in the North Pacific – a data-based approach

As part of its annual bottom-trawl survey program, the Alaska Fisheries Science Center (AFSC) has been collecting and analyzing the stomach contents of groundfish predators since 1981. Between 1981 and 2011, a total of 233,451 fish stomachs were collected and analyzed from the eastern Bering Sea, the Gulf of Alaska, and the Aleutian Islands large marine ecosystems; these data are now available online as AFSC’s Groundfish Trophic Interactions Database. Here, we discuss features of the survey and data to aid in the interpretation and use of this extensive dataset for the Alaska region. The primary fish sampled include walleye pollock (Gadus chalcogrammus), Pacific cod (Gadus macrocephalus), Pacific halibut (Hippoglossus stenolepis), and arrowtooth flounder (Atheresthes stomias), although 159 predator species have been included in the stomach content analysis. Prey length measurements are included for important commercial prey and can identify age or size classes of prey prior to their recruitment into fisheries and most other surveys. With these data, one can track time trends in growth, mortality, and prey composition as ecosystem indicators, and include food web interactions in fish stock assessments for ecosystem-based fisheries management.     

Spatial Match-Mismatch between Juvenile Fish and Prey Provides a Mechanism for Recruitment Variability across Contrasting Climate Conditions in the Eastern Bering Sea

Understanding mechanisms behind variability in early life survival of marine fishes through modeling efforts can improve predictive capabilities for recruitment success under changing climate conditions. Walleye pollock (Theragra chalcogramma) support the largest single-species commercial fishery in the United States and represent an ecologically important component of the Bering Sea ecosystem. Variability in walleye pollock growth and survival is structured in part by climate-driven bottom-up control of zooplankton composition. We used two modeling approaches, informed by observations, to understand the roles of prey quality, prey composition, and water temperature on juvenile walleye pollock growth: (1) a bioenergetics model that included local predator and prey energy densities, and (2) an individual-based model that included a mechanistic feeding component dependent on larval development and behavior, local prey densities and size, and physical oceanographic conditions. Prey composition in late-summer shifted from predominantly smaller copepod species in the warmer 2005 season to larger species in the cooler 2010 season, reflecting differences in zooplankton composition between years. In 2010, the main prey of juvenile walleye pollock were more abundant, had greater biomass, and higher mean energy density, resulting in better growth conditions. Moreover, spatial patterns in prey composition and water temperature lead to areas of enhanced growth, or growth ‘hot spots’, for juvenile walleye pollock and survival may be enhanced when fish overlap with these areas. This study provides evidence that a spatial mismatch between juvenile walleye pollock and growth ‘hot spots’ in 2005 contributed to poor recruitment while a higher degree of overlap in 2010 resulted in improved recruitment. Our results indicate that climate-driven changes in prey quality and composition can impact growth of juvenile walleye pollock, potentially severely affecting recruitment variability.     

Fish eggs and larvae in the diet of herring Clupea harengus membras Linnaeus, 1758 and the sprat Sprattus sprattus balticus (Schneider, 1904) (Clupeidae) in the Southeastern Baltic Sea

The qualitative and quantitative composition of fish eggs and larvae in the stomach contents of Baltic herring and sprat was examined. The occurrence of fish eggs and larvae in the diet of herring was markedly higher, compared to sprat. Eggs of four and larvae of nine fish species were found in the herring stomachs. The prey species composition of the herring and sprat diets was related to the ichthyoplankton composition and to the degree of overlapping of the prey and predator distributions. Sprat eggs were the predominant prey in the herring diet in May and sprat larvae prevailed in August. The daily ration estimates indicate that the sprat larvae consumption by herring in August exceeded the sprat egg consumption in May by many times. Cod eggs in the herring stomachs were only found at the optimal oxygen content and salinity for the cod eggs in the near-bottom water layer.     

Prey selection by age-0 walleye pollock, Theragra chalcogramma, in nearshore waters of the Gulf of Alaska

Juvenile walleye pollock, Theragra chalcogramma, is the dominant forage fish on the continental shelf of the Gulf of Alaska, yet little is known about the feeding habits of this important interval of pollock life history. The taxonomic composition and size of prey found in the stomachs of age-0 juveniles collected at three nearshore locations in the Gulf of Alaska in September 1990 were compared to the composition and size of zooplankton collected in concurrent plankton tows. The maximum length of prey consumed increased dramatically over the length range of pollock examined (58–110 mm) from approximately 7 mm to 30 mm, due mainly to the consumption of large euphausiids and chaetognaths by the bigger individuals. The maximum width of prey changed little over this size range although there was a general increase in prey width with increasing predator size. The minimum prey length and width did not change with increasing fish size. Juvenile pollock generally selected the larger prey sizes relative to what was available. Juvenile pollock showed a marked preference for adult euphausiids and decapod larvae and an avoidance of copepods and chaetognaths relative to the numbers collected in net tows. These results are discussed relative to the feeding ecology of these juvenile fishes. This revised version was published online in July 2006 with corrections to the Cover Date.     

Oxygen and salinity characteristics of predator–prey distributional overlaps shown by predatory Baltic cod during spawning

In the distributional overlap volume of Baltic cod Gadus morhua and its prey, studied in the Bornholm Basin in the southern Baltic Sea, only a fraction of the sprat Sprattus sprattus population vertically overlapped with the Baltic cod population. Sprat occurred in the intermediate water, in the halocline and in the bottom water, while herring Clupea harengus and Baltic cod occurred exclusively in the halocline and in the bottom water. Only parts of the sprat population were hence accessible for Baltic cod, and only a fraction of the sprat had access to the Baltic cod eggs below the halocline. Baltic cod–clupeid overlap volumes appeared to be determined by salinity stratification and oxygenation of the bottom water. Hydrography time series were used to estimate average habitat volumes and overlap from July to September in 1958–1999. In the 1999 survey spawning Baltic cod had greater ratios of empty stomachs and lower average rations than non-spawning Baltic cod. The average ration for Baltic cod caught within 11· 4 m from the bottom (demersal) did not differ from the average ration of Baltic cod caught in shallower waters (pelagic), because spawning and non-spawning Baltic cod in both strata were caught at equal rates. The diet of the Baltic cod caught demersally contained more benthic invertebrates, especially Saduria entomon, but Baltic cod caught pelagically also had fresh benthic food in their stomachs, indicating vertical migration of individual fish.     

Isolation of the North American strain of viral hemorrhagic septicemia virus (VHSV) associated with epizootic mortality in two new host species of Alaskan marine fish

Thousands of dead Pacific herring Clupea pallasi, Pacific hake Merluccius productus and walleye pollock Theragra chalcogramma were reported in Lisianski Inlet near Pelican, Alaska, USA, on August 1, 1998. The Pacific hake and pollock continued to die through the end of September. Virological examinations of dead fish identified the North American strain of viral hemorrhagic septicemia virus (VHSV) from all 3 species of fish as well as associated high virus titers and possible histopathological lesions. No other primary fish pathogens were detected and there were no apparent environmental causes for fish mortality. This is the first report of VHSV in 2 new Alaskan fish host species and of a natural epizootic associated with VHSV in which progressive mass mortality was observed simultaneously in herring and 2 other species of free-ranging marine fish.     

A mitogenomic approach to the taxonomy of pollocks: <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> and <Emphasis Type="Italic">T. finnmarchica</Emphasis> represent one single species

Background  

The walleye pollock (Theragra chalcogramma) and Norwegian pollock (T. finnmarchica) are confined to the North Pacific and North Atlantic Oceans, respectively, and considered as distinct species within the family Gadidae. We have determined the complete mtDNA nucleotide sequence of two specimens of Norwegian pollock and compared the sequences to that of 10 specimens of walleye pollock representing stocks from the Sea of Japan and the Bering Sea, 2 specimens of Atlantic cod (Gadus morhua), and 2 specimens of haddock (Melanogrammus aeglefinus).     

Description of five new Loma (Microsporidia) species in pacific fishes with redesignation of the type species Loma morhua Morrison & Sprague, 1981, based on morphological and molecular species-boundaries tests

Five new species of Loma were described from five Pacific fishes using light-microscopic and ultrastructural features along with phylogenetic analysis of the gene sequences of ribosomal RNA (rRNA) and elongation factor 1-alpha. Morphological data revealed both qualitative and quantitative differences in developmental stages and timing, vesicles, xenoma features, and spore sizes with statistical support that differentiated Loma pacificodae n. sp. in Pacific cod, Loma wallae n. sp. in walleye pollock, Loma kenti n. sp. in Pacific tomcod, Loma lingcodae n. sp. in lingcod, and Loma richardi n. sp. in sablefish from each other and other species in the genus. Phylogenetic analyses combined with monophyly tests supported species designations, but with low resolution in two cases perhaps due to rRNA paralogs or recent speciation. Loma branchialis in haddock was shown to be separate from Loma morhua in Atlantic cod, thereby making L. morhua, and not L. branchialis, the type species. A species from brook trout was shown to be a separate species from Loma salmonae, not a variant strain selected in the laboratory. By comparison with gadid host phylogeny, these Loma species appear to have coevolved with their hosts, first colonizing the Pacific basin about 12 million years ago.     

Resource seasonality and fish diets in an Illinois stream

Synopsis The purpose of this study was to evaluate the intensity of competition for food among 9 species of stream fishes that primarily eat aquatic invertebrates. The taxonomic and size composition, and numerical abundance of aquatic invertebrates were monitored for one year using drift and benthic samples. Diet data were obtained from stomachs of fishes captured at the same time and place that invertebrate sampling was done. Diet characteristics examined included taxonomic and size composition, number of prey per fish, and diet breadth. Drifting invertebrates were more abundant early in the year (March–June) than later (July–January). The summer-early fall scarcity of invertebrates was especially notable among those>3.6 mm long, which comprised the bulk of prey found in fish stomachs. Average prey size eaten by a fish species was positively correlated with fish mouth size, but interspecific overlap in prey size was extensive. Cyprinids as a group (5 species) ate proportionally fewer small (< 3.6 mm long) prey from July to January than did the centrarchids and stonecat. Taxonomic compositions of available invertebrates and fish diets varied markedly among sampling dates, but the use of prey taxa by fishes was not correlated with the availability of those taxa. Use of aquatic prey taxa was generally similar among fish species, but cyprinids as a group ate proportionally more terrestrial prey from July to January than did the centrarchids and stonecat. Diet breadths for all species increased as food levels declined, indicating that these fishes experienced resource depression. Food scarcity was evidently more severe for cyprinids since their stomachs contained few prey through the summer and fall relative to the centrarchids and stonecat. Though the fish species studied probably compete for food in the summer and fall, this competition did not account for the community structure observed.     

Incidence of parasitic worms in stomachs of pelagic and demersal fish of the Rockall Trough, northeastern Atlantic Ocean

The incidence of gyrocotyllid worms in chimaeroids and cestode larvae and nematodes in the stomachs of some 13 600 demersal and pelagic fish from the Rockall Trough, northeastern Atlantic is determined. The observations were made in conjunction with a detailed study of the diets of the fish which belonged to 117 species. No dietary sources of the infections could be identified with certainty. Pelagically caught fish, from between the surface and 2500 m depth, had no helminth infestation of their stomachs. Infestations of demersal fish were more pronounced between 1500 and 2900 m than between 500 and 1250 m depth.