The food supply of the Pacific salmon of the genus Oncorhynchus in the northwestern Pacific Ocean. 2. Comparative characterization and general state

The interannual variations and general state of the food supply of Pacific salmon (Oncorhynchus spp.) in the 2000s in the northwestern Pacific Ocean (including the Bering Sea and the Sea of Okhotsk) were analyzed based on indirect characteristics that indicate the variability of their forage base, feeding habits, growth, and biomass. A new index for the quantitative evaluation of food supply was suggested. The food supply of the Pacific salmon during the 2000s was found to be sufficient to maintain the normal functioning of populations. With high abundance of Pacific salmon, the food supply tended to decrease. However, this caused no negative consequences for the survival of major salmon stocks during the marine period of life and, as a rule, no marked decrease in the food consumption and growth rates of fish. A relative increase in food competition was compensated by adaptive changes in the diet and diel feeding rhythm of salmon. With the shortage of preferred food organisms (amphipods, euphausiids, and pteropods), Pacific salmon changed to consuming minor prey (copepods and chaetognaths), and numerous mesopelagic species of macroplankton and micronekton in the evening hours.     

Comparative characteristics of food supply of Pacific salmon (<Emphasis Type="Italic">Oncorhynchus</Emphasis> spp.) in the Bering Sea from 2002 to 2006

Based on complex epipelagic surveys in the western Bering Sea, a comparative analysis of food supply of Pacific salmon (Oncorhynchus spp.) was conducted in summer and fall from 2002 to 2006. Nine indirect indices of food supply used in the study were as follows: feeding similarity, width of the feeding spectrum, diet feeding ration, diet feeding rhythms, fraction of accessory food in the ration, growth rate of the fish, abundance of food resources, and abundance of salmon. The food supply of salmon is lower in summer 2003 and fall 2006 in comparison to the food supply in other years of the study. However, well expressed feeding selectivity, consumption of prey items of certain type, and small proportion of accessory food (copepods and chaetognaths) prevailed in plankton, suggests the presence of sufficient food resources for Pacific salmon in the western Bering Sea.     

Daily feeding rhythm and rations of Pacific salmon of the genus Oncorhynchus in the period of marine prespawning migrations

The daily feeding rhythm and rations of the humpback salmon Oncorhynchus gorbuscha, the sockeyed salmon O. nerka, and the chum salmon O. keta during marine prespawning migrations is investigated with consideration of materials collected at daily stations in waters off eastern Kamchatka in June–July 1999 and 2001 (from catches of drift nets). The bulk of humpback salmon and sockeyed salmon food consists of euphausiids, hyperiids, large copepods, pteropods, and fish juveniles. In the food of chum salmon, pteropods dominated. In a 24-h period, salmon manifest a pronounced evening peak of stomach fullness, while at night feeding discontinues. Samples collected in the morning consisted of fish who had just started feeding after the night pause. In addition to nightly period of rest, all species manifested a daytime decrease in foraging activity, though less pronounced than in the night. The daily rhythm of Pacific salmons’s feeding depends on the vertical migrations of their food items (representatives of sound-scattering layers). During the marine feeding period, the most intensive feeding is recorded in the humpback salmon and chum salmon. The daily ration of the humpback salmon is lower than that of the chum salmon but includes animals of higher food value. Due to a high digestion rate in combination with a large stomach volume, the chum salmon can consume a large quantity of low-calorific food in a short time. The daily ration of the sockeyed salmon is considerably lower than that of other salmon species.     

Feeding and daily ration of the chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> in the western Bering Sea in the summer-fall of 2004

This study was based on data on the feeding of the chum salmon Oncorhynchus keta sampled at daily stations in areas of the Navarin Shelf and Aleutian Basin in the Bering Sea in August and September 2004. The food composition and daily dynamics of chum feeding were investigated. The rate of food evacuation from the stomach and the amount of the daily diet were estimated. The distinctions in the feeding patterns of chum sampled from different areas were considered. In the Navarin Shelf area the chumís prey were primarily nekton species (mostly Alaskan pollack underyearlings) and the amount of the daily diet for chum of various size groups was 1.8–3.8% of their body weight. In the deep-water Aleutian Basin, the daily dynamics of chum feeding depended on migration of euphausiids, amphipods and mesopelagic fishes to the surface water layers of epipelagics during the night. The daily ration for chum that fed mainly on plankton did not exceed 2.6–2.8% of their body weight in August and 1.2–2.5% in September.     

Feeding pattern of the great sculpin Myoxocephalus polyacanthocephalus (Cottidae) and its position in the trophic system of near-Kamchatka waters

Feeding patterns of the great sculpin Myoxocephalus polyacanthocephalus in near-Kamchatka waters of the Sea of Okhotsk, the Bering Sea and Pacific Ocean are considered using materials collected during 1978–2002. This species of Cottidae is characterized by wide feeding spectra (over 150 nutritive objects); nonetheless, fishes (on average 61.1%) and Decapoda (20.0%) have the highest significance in its diet. The fishes include Alaska pollock Theragra chalcogramma (32.7% of food mass), flatfish Pleuronectidae (12.2%), and sculpins Cottidae (10.5%); Decapoda include crabs from the family Majidae (18.5%). Seasonal, local, interannual, and age-related changes in food composition of the great sculpin are considered. This species is a facultative ambuscade predator; it is characterized by wide feeding spectra that permits him to use a considerable range of food components. In the trophic system of near-Kamchatka regions, units that form the biomass of the great sculpin are selected.     

Trophic ecology of Pacific salmon (<Emphasis Type="Italic">Oncorhynchus</Emphasis> spp.) in the ocean: a synthesis of stable isotope research

Increasing interest in the marine trophic dynamics of Pacific salmon has been motivated by the recognition of their sensitivity to changing climate and to the competitive effects of hatchery fish on wild stocks. It has become more common to use stable isotopes to supplement traditional diet studies of salmon in the ocean; however, there have been no integrated syntheses of these data to determine whether stable isotope analyses support the existing conventional wisdom of feeding strategies of the Pacific salmon. We performed a meta-analysis of stable isotope data to examine the extent of trophic partitioning among five species of Pacific salmon during their marine lives. Pink, sockeye, and chum salmon showed very high overlap in resource use and there was no consistent evidence for chum relying on alternative food webs dominated by gelatinous zooplankton. δ¹⁵N showed that Chinook and coho salmon fed at trophic levels higher than the other three species. In addition, these two species were distinctly enriched in ¹³C, suggesting more extensive use of coastal food webs compared to the more depleted (pelagic) signatures of pink, sockeye, and chum salmon. This paper presents the first synthesis of stable isotope work on Pacific salmon and provides δ¹⁵N and δ¹³C values applicable to research on the fate of the marine derived nutrients these organisms transport to freshwater and riparian ecosystems.     

On the Persistence of Stereotypes Concerning the Marine Ecology of Pacific Salmon (<Emphasis Type="Italic">Oncorhynchus</Emphasis> spp.)

Some of the views on the marine ecology of Pacific salmon (Oncorhynchus spp.) that were popular in the second half of the 20th century are discussed critically: the absolutization of the influence of sea surface temperature on distribution of salmon and strength of their year classes, as well as the conclusions on the shortage of food (particularly in winter) and the fierce competition for food, the “suppression” of other salmon species and own adjacent broodline by pink salmon, the limited carrying capacity of the pelagic zone of subarctic ocean waters for salmon, the distortion of the structure of epipelagic communities in ecosystems of the North Pacific due to the large-scale stock enhancement of chum salmon, etc. Most of these ideas have not been confirmed by the data of long-term monitoring conducted in the form of complex marine expeditions by the Pacific Research Fisheries Center (TINRO Center) in the Far-Eastern Seas and adjacent North Pacific waters since the 1980s. The data show that Pacific salmon are ecologically very flexible species with a wider temperature range of habitat than was previously believed. Salmon are able to make considerable vertical migrations, easily crossing zones of sharp temperature gradient and different water masses. Having the wide feeding spectra and being dispersed (as non-schooling fish) when feeding in the sea and ocean, they successfully satisfy their dietary needs in vast areas even with relatively low concentrations of prey organisms (macroplankton and small nekton). The total biomass of all the Pacific salmon species in the North Pacific is not greater than 4–5 million t (including 1.5–2.0 million t in Russian waters), whereas the biomass of other common species of nekton is a few hundreds of millions of tons. Salmon account for 1.0–5.0% of the total amount of food consumed by nekton in the epipelagic layer of the western Bering Sea, 0.5–1.0% in the Sea of Okhotsk, less than 1% in the ocean waters off the Kuril Islands, and 5.0–15.0% in the ocean waters off East Kamchatka. Thus, the role of Pacific salmon in the trophic webs of subarctic waters is rather moderate. Therefore, neither pink nor chum salmon can be considered as the species responsible for the large reorganization in ecosystems and the population fluctuations in other common nekton species.     

Long-term trends in food habits of Asiatic black bears in the Misaka Mountains on the Pacific coast of central Japan

I did the food habits of the Asiatic black bear Ursus thibetanus from 1013 fecal samples collected between 1999 and 2005 in the Misaka Mountains on the Pacific coast of central Japan. The food habits of the bears showed clear seasonal changes, and I classified the food resources of the bears into three types. Staple foods were green vegetation in spring, soft mast (Prunus spp.) and insects in summer, and hard mast (Quercus spp.) in autumn. Alternative foods were green vegetation and other soft mast (Rubus spp.) in summer and Japanese chestnuts Castanea crenata and vine fruits in autumn. Foods of opportunity were hard mast (Quercus spp.) that had been shed in the previous autumn and were found in spring and other fruits in autumn. Seasonal food habits showed yearly variations: bears used alternative foods and foods of opportunity in response to the yearly variation in staple food amount, but the magnitude of variability of food habits differed among seasons, with large variability in autumn and small variability in summer and spring. The primary influence on the yearly variation in food habit is presumably the fluctuation in fruit production among years. Summer is probably the most difficult season in terms of the bear's food supply, because the number of fruiting species is limited and staple foods such as new green vegetation and fruits are less available. Long-term studies of the availability of the main food items and food habits of bears will be critical for further understanding these animals’ feeding ecology and for determining the factors that influence their behavior.     

Evidence for competitive dominance of Pink salmon (<Emphasis Type="Italic">Oncorhynchus gorbuscha</Emphasis>) over other Salmonids in the North Pacific Ocean

Relatively little is known about fish species interactions in offshore areas of the world’s oceans because adequate experimental controls are typically unavailable in such vast areas. However, pink salmon (Oncorhynchus gorbuscha) are numerous and have an alternating-year pattern of abundance that provides a natural experimental control to test for interspecific competition in the North Pacific Ocean and Bering Sea. Since a number of studies have recently examined pink salmon interactions with other salmon, we reviewed them in an effort to describe patterns of interaction over broad regions of the ocean. Research consistently indicated that pink salmon significantly altered prey abundance of other salmon species (e.g., zooplankton, squid), leading to altered diet, reduced total prey consumption and growth, delayed maturation, and reduced survival, depending on species and locale. Reduced survival was observed in chum salmon (O. keta) and Chinook salmon (O. tshawytscha) originating from Puget Sound and in Bristol Bay sockeye salmon (O. nerka). Growth of pink salmon was not measurably affected by other salmon species, but their growth was sometimes inversely related to their own abundance. In all marine studies, pink salmon affected other species through exploitation of prey resources rather than interference. Interspecific competition was observed in nearshore and offshore waters of the North Pacific Ocean and Bering Sea, and one study documented competition between species originating from different continents. Climate change had variable effects on competition. In the North Pacific Ocean, competition was observed before and after the ocean regime shift in 1977 that significantly altered abundances of many marine species, whereas a study in the Pacific Northwest reported a shift from predation- to competition-based mortality in response to the 1982/1983 El Nino. Key traits of pink salmon that influenced competition with other salmonids included great abundance, high consumption rates and rapid growth, degree of diet overlap or consumption of lower trophic level prey, and early migration timing into the ocean. The consistent pattern of findings from multiple regions of the ocean provides evidence that interspecific competition can significantly influence salmon population dynamics and that pink salmon may be the dominant competitor among salmon in marine waters.     

Trophic ecology and food consumption of fishes in a hypersaline tropical lagoon

This study evaluated the trophic ecology (diet composition, trophic strategy, similarities and overlap between species, feeding period and food consumption) of six benthivorous fish species in Araruama Lagoon, the largest hypersaline tropical lagoon on the east coast of South America, with an area of 210 km² and an average salinity of 52. The burrfish Chilomycterus spinosus fed on Anomalocardia flexuosa shell deposits, ingesting associated fauna. The caitipa mojarra Diapterus rhombeus differed from all other species, having not only the highest proportions of algae and Nematoda, but also feeding on polychaete tentacles. The two mojarras Eucinostomus spp. showed similar trophic strategies, feeding mostly on Polychaeta. The corocoro grunt Orthopristis ruber also fed mainly on Polychaeta, but differed from Eucinostomus spp. in secondary items. The whitemouth croacker Micropogonias furnieri fed mainly on small Crustacea at night, showing a high number of secondary prey items with low frequencies and high prey‐specific abundance. The daily food consumption (g food g^?1 fish mass) for Eucinostomus argenteus was 0·012 and was 0·031 and 0·027 for M. furnieri in two different sampling events. The diet similarities between Araruama Lagoon and other brackish and marine environments indicate that hypersalinity is not a predominant factor shaping the trophic ecology of fishes in this lagoon. The stability of hypersaline conditions, without a pronounced gradient, may explain the presence of several euryhaline fishes and invertebrates well adapted to this condition, resulting in a complex food web.     

Diel feeding rhythms and daily food consumption of juvenile Atlantic salmon in the River Alta, northern Norway

Atlantic salmon fry (0+) sampled from the River Alta exhibited only minor differences in stomach content weights and feeding rates throughout diel periods, but feeding rates were generally lowest at night. In contrast, salmon parr (1+ to 3+) had large diel fluctuations in stomach content weight, with the largest weights usually being recorded during the night and early morning. Accordingly, their feeding rates were highest at night. This nocturnal feeding pattern was consistent throughout all sampling occasions, and appeared to reflect a persistent feeding periodicity in the salmon parr. The daily food consumption rates of both fry and parr were highest during midsummer and decreased towards autumn.     

Spatial distribution and features of biology of Pacific sleeper shark Somniosus pacificus in the North Pacific

The results of the investigations of spatial and vertical distribution of Pacific sleeper shark Somniosus pacificus in the North Pacific Ocean conducted for many years are presented. In addition, the size distribution and features of biology of the species are studied. The largest abundance of the species is registered in the Bering Sea, western Gulf of Alaska, eastern Aleutian Islands, and Pacific waters of northern Kuril Islands and southeastern Kamchatka. The species is the most abundant near the bottom at the depth from 200 to 700 m and in the pelagic waters at a depth of 100–200 m. The average depths of the catches of Pacific sleeper shark substantially change over the year reaching minimum values in June and maximum values in December. Vertical daily migrations (to the water column at night and to the bottom during the day) are registered. The catches are represented by fish 26–352 cm in length, and sharks 100–200 cm in length prevail. The males are noticeably smaller than the females. In general, condition of the fishes decreases and feeding intensity increases with growth. Food composition substantially changes with the increase of body length: consumption of squids decreases and consumption of crustaceans, fishes, and fishery wastes increases. The food composition is slightly different in the females and males.     

Summer diet of beluga whales inferred by fatty acid analysis of the eastern Beaufort Sea food web

Beluga whales (Delphinapterus leucas) are the most abundant odontocetes in Arctic waters and are thus thought to influence food web structure and function. The diet of the Beaufort Sea beluga population is not well known, partly due to the inherent difficulty of observing feeding behaviour in Arctic marine cetaceans. To determine which prey items are critical to the Beaufort Sea beluga diet we first examine and describe the Mackenzie Delta and Beaufort Sea food web using fatty acid analyses. Fatty acid profiles effectively partitioned prey items into groups associated with their habitat and feeding ecology. Next, the relative contribution of various prey items to beluga diet was investigated using fatty acids. Finally, beluga diet variability was examined as a function of body size, a known correlate of habitat use. Beluga appeared to feed predominantly on Arctic cod (Boreogadus saida) collected from near shore and offshore regions. Size related dietary differences suggested larger sized beluga preferred offshore Arctic cod given the shared high levels of long chain monounsaturates, whereas smaller sized beluga appeared to feed on prey in their near shore habitats that included near shore Arctic cod. The presence of Arctic cod groups in shallow near shore and deep offshore habitats may facilitate the behavioural segregation of beluga habitat use as it relates to their size and resource requirements. Given Arctic cod are a sea ice associated fish combined with the accelerated sea ice loss in this region, beluga whales may need to adapt to new dietary regimes.     

A radiographic method for feeding studies on fish using metallic iron powder as a marker

A novel method for studying trophic dynamics of fish is described which involves X-raying animals which have been fed a diet labelled with metallic iron powder. Results of feeding studies on juvenile Atlantic salmon, Salmo salar L., are presented which demonstrate that food consumption and gastric evacuation rates can be measured without incurring the major disadvantages offeree feeding, starvation or autopsy which are associated with most of the established methods.     

Underyearling pink and chum salmon in the western Bering Sea during the fall season (September and October 2013): Distribution,feeding habits,and growth patterns

The ontogeny of a year class of pink and chum salmon is described for the period after the redistribution of underyearling individuals from coastal waters to deep-sea areas of the western Bering Sea in September and October, 2013. The intensity of their feeding was high; their diet included hyperiids, pteropods, and juvenile euphausiids. The metabolic costs of growth reached only 20% of the consumed food, which indicates significant energy costs for locomotion; moreover, as the body size increases, the level of metabolic functions rises at a decreasing rate, which causes the body growth to slow down and food consumption to decrease. The main items in the diet of underyearling salmon are characterized by a low content of dry matter, low lipid content, and, consequently, a low calorie content, i.e., underyearlings mainly consume protein-rich food with a low fat content. The chemical composition of the tissues almost did not differ between underyearling pink and chum salmon. Both species typically had a low fat content in their muscles. Thus, fat is not accumulated at this stage of ontogeny; all energy that is obtained with food, after being used for locomotion and metabolism, is spent for linear growth.     

Feeding ecology and trophic impact of the hydroid Obelia dichotoma in the Kongsfjorden (Spitsbergen, Arctic)

Obelia dichotoma is a thecate hydroid with a worldwide distribution, occurring mainly on shallow water hard substrates. Since the trophic ecology of hydroids in polar waters is badly understood, the aim of the present work was to study qualitatively and quantitatively the diet of these organisms in an Arctic environment and to determine their trophic significance. For this purpose, the density of the hydroid population was documented, and simultaneously, zooplankton was sampled in two different years (1997 and 1998). Prey capture rates were estimated by analysing the gastrovascular content of the polyps in a diurnal cycle. Additionally, the digestion time of O. dichotoma was measured by laboratory feeding experiments using diatoms as food items. The analyses of the gastrovascular cavities of the polyps sampled during the diurnal cycles showed that O. dichotoma fed mainly on faecal pellets, organic matter and microalgae. Zooplankton prey was also observed, but gastrovascular contents and zooplankton abundance did not show any correlation in both years. The consumption rates of the hydroid populations differed between the 2 years. It was almost double (8.9 mg Carbon m^?2) in 1998 compared to 1997 (5.5 mg Carbon m^?2). The significance of the environmental variability in the feeding ecology and population dynamics of hydroids under Arctic conditions is discussed.     

The survival of Pacific salmon in the North Pacific in the winter and spring

The effects of various factors (water temperature, food availability, predation, and the size of juveniles) on the survival of Pacific salmons during overwintering in open ocean waters are analyzed based on the data collected by expeditions of the Pacific Research Fisheries Center to the northwestern Pacific Ocean in the winter and spring seasons of 1986–1992 and 2009–2011, as well as in the summer seasons of 2004–2011. The temperature factor is unlikely to be a direct cause of the high salmon mortality in the ocean during the winter, as there is no clear evidence that it affects food availability for salmon. The biomass of forage zooplankton in the Subarctic Front zone in February and March is lower than that in April and June–July, but it does not decrease substantially in the winter months. Taking the fact into account that the total abundance of planktivorous nekton is also low in this area during the winter, food availability cannot be considered a crucial factor that has a serious influence on salmon mortality in this period. The difference in feeding intensity between salmon species and between their size groups in the winter and spring is determined by their life strategies. The observed variations in feeding intensity and lipid accumulation from autumn to spring are caused by cyclic seasonal changes in physiological processes in salmon rather than by the amount and availability of food resources. The low abundance of predators in subarctic waters and in the Subarctic Front zone in the winter also cannot reduce salmon abundance substantially. The probable relationship between the critical size of juveniles and their survival in the winter is considered using the example of a Sea of Okhotsk stock of pink salmon. The conclusion is that the size of juvenile pink salmon cannot always be used as a predictor of the values of its subsequent returns, because survival of salmon during the ocean period of life depends both on the initial conditions during downstream migration and on the ocean conditions that form in the winter. Thus, none of the factors above can be considered as strictly limiting the abundance of Pacific salmon in the winter. It is more probable that the survival of salmon in the ocean is influenced, to a lesser or greater extent, by the combined effects of abiotic and biotic factors.     

Diel diet shift of roach and its implications for the estimation of daily rations

In a small, 12 ha, mesotropic lake, roach Rutilus rutilus performed diel habitat shifts that clearly influenced the composition of their diet. During daytime, roach stayed in the littoral zone and concentrated on littoral prey. At night they were found in the pelagic zone, and pelagic prey items such as Daphnia spp. or Chaoborus flaricans dominated their food. On a seasonal scale, there were shifts in the importance of different food items and in the diel pattern of feeding intensity. Bioenergetics modelling in combination with an evacuation rate method for estimating daily rations allowed for changes in feeding modes to be taken into account, and so food item specific daily rations over the season could be determined. With the evacuation rate method applied on selected days, diel changes in diet compositions and feeding intensities could be quantitatively accounted for. When the 24 h integrated diet proportions were then used as an input parameter for bioenergetics modelling, food item specific consumption could be determined over the entire sampling season. The consideration of the diel diet shifts proved to be essential for the model output. If only the daytime or the night-time diet composition (derived from one single daily sampling) was taken into account for bioenergetics modelling, severe under- or overestimations of daily rations for specific food items resulted.     

A population model for limited food competition

In this paper a deterministic differential equation system is proposed to model the population dynamics of a biological community in which two species on the same trophic level compete for a common food, taken to be in limited supply. Food limitation is assumed to be the only inhibition of the growth of the populations and food quantity is assumed to be only affected by consumption. The model is thus designed to mimic a closed experimental situation rather than a natural community.Analytical properties of the solution of the differential equation system are developed and corresponding biological interpretations suggested.Cited laboratory data on the experimental batch community consisting of the marine ciliates Euplotes vannus and Uronema marinum feeding on bacteria motivated the model and supported its analytic properties.     

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.