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.     

Diel variation in feeding rate and prey composition of herring and mackerel in the southern Gulf of St Lawrence

Diel feeding patterns of herring Clupea harengus and mackerel Scomber scombrus in the southern Gulf of St Lawrence were examined based on samples obtained by midwater trawling between 19 and 26 June 2001. Within 3 h time periods, stomach contents tended to be more similar between fish from the same tow than between fish from different tows. Thus, in contrast to previous diet studies, which have used individual fish stomachs as independent observations, tow was used as the experimental unit in statistical analyses in this study. Diel patterns in stomach fullness were identified using generalized additive models. Two peaks in stomach fullness occurred for herring, one in the morning and the other in the evening. Mackerel showed an increase in feeding intensity throughout the day with a peak in mid‐afternoon. The diel changes in stomach contents suggested rapid gastric evacuation rates for both species, especially for herring. The estimate of the instantaneous evacuation rate for herring was twice that for mackerel. Calanus copepods (mainly C. hyperboreus ), fishes (mainly capelin Mallotus villosus ) and euphausiids were the main prey found in the stomachs of both species. Calanus copepods dominated the diet of herring regardless of time period. They also dominated the diet of mackerel during the late afternoon, evening and night while fishes and euphausiids were dominant during the morning and early afternoon. These diel patterns emphasize the need for sampling throughout the day and night in order to estimate ration and diet composition for bioenergetic and ecosystem models.     

The feeding strategy of dab Limanda limanda in the southern North Sea: linking stomach contents to prey availability in the environment

The feeding strategy of dab Limanda limanda was investigated within a predefined research area in the southern North Sea for 2 years. The aims were: 1) to establish the relationship between prey availability and dab abundance, distribution, body condition and prey selection, and 2) to determine the underlying functional processes of prey choice in dab. Temporal and spatial trends of prey availability in relation to dab abundances, distribution, condition and prey selection were analysed. Temporal trends in dab condition, percentage of empty stomachs and numbers of ingested prey were apparent. In particular in the winter period 2000–2001 dab were in poor condition and prey items were only rarely found in stomachs. Similarly prey availability decreased in the environment while the abundance of dab increased. Thus, temporal changes in the nutritive state were partly thought to be attributed to density‐dependent effects. The stomach composition of dab during the two consecutive summers was mainly dominated by Pariambus typicus and Ophiura albida while in the winter period only O. albida made a significant contribution to the stomach content. Analysis of prey characteristics indicated that only prey densities in the environment significantly influenced prey choice in dab, while other characteristics such as the position of the prey in the habitat, its palatability or mobility did not have significant effects. The feeding strategy of dab thus was highly opportunistic, although trends were apparent that suggested that buried living fauna was less likely to be ingested, which was also observed in previous studies.     

Pacific walrus (Odobenus rosmarus divergens):Differential prey digestion and diet

Stomach content data from 798 Pacific walruses ( Odobenus rosmarus divergens ) collected during 1952–1991 were analyzed using a method that evaluates the stage of digestion of prey remains. Non-molluscan prey taxa were not well represented in previous interpretations of walrus diet due to digestion biases. Stomach contents least affected by digestion (fresh stomachs) contained more prey taxa than stomachs of an unknown or more digested state. Bivalves, gastropods, and polychaete worms were the most frequent prey items in both the Bering and Chukchi seas, although bivalves occurred more frequently in stomachs from the Bering Sea and gastropods occurred more frequently in stomachs from the Chukchi Sea. Male and female walruses consumed essentially the same prey when in the same location. Using only fresh stomachs collected between 1975 and 1985, there was no significant difference between the proportion that contained mostly bivalves and the proportion that contained non-bivalve prey items. Earlier interpretations of a change in walrus diet in this period compared to the prior two decades may have been due to digestion as well as sampling biases. Current climatic changes may affect walrus's access to diverse, productive shallow water feeding areas.     

Diet of Atlantic cod in the southern Gulf of St Lawrence as an index of ecosystem change, 1959–2000

Major temporal changes in the importance of euphausiids and Atlantic herring Clupea harengus in the diet of Atlantic cod Gadus morhua , 10–75 cm L _F, occurred in shallow waters (<100 m depths) of the southern Gulf of St Lawrence between 1959 and 2000. Euphausiids represented 6–70% of prey mass for Atlantic cod 31–60 cm L _F from 1959 to 1987 but only trace amounts were detected in stomachs collected from 1990 to 2000. Mysids and gammarid amphipods (for Atlantic cod ≤45 cm L _F), and Atlantic herring (for Atlantic cod >45 cm L _F) largely replaced euphausiids in the Atlantic cod diets from 1990 to 2000. This diet change suggested there has been a major perturbation of the food web of the southern Gulf of St Lawrence. The importance of fishes (mostly Atlantic herring) in the diet of Atlantic cod >45 cm L _F increased significantly between the periods 1959–1980 and 1987–2000. Atlantic herring comprised 0–4% (mean 1.3%) of prey mass of Atlantic cod 46–60 cm L _F from 1959 to 1980 and increased to 2–42% (mean 19.6%) of the diet from 1987 to 2000. Atlantic herring comprised 0–25% (mean 9.4%) of the prey mass of Atlantic cod 61–75 cm L _F from 1959 to 1980 and increased to 42–81% (mean 54.2%) of prey mass from 1987 to 2000. This increased consumption of Atlantic herring was consistent with observed changes in abundance of Atlantic herring in the ecosystem between the late 1970s and 2000. The large changes in consumption of euphausiids and Atlantic herring represent highly significant changes and would need to be included in the development and interpretation of ecosystem-based management models for this ecosystem.     

Food habits of the sandpaper skate, Bathyraja kincaidii (Garman, 1908) off central California: seasonal variation in diet linked to oceanographic conditions

The stomachs of 130 sandpaper skates, Bathyraja kincaidii (Garman, 1908), were sampled from off central California to determine their diet composition. The overall diet was dominated by euphausiids, but shrimps, polychaetes and squids were also important secondary prey. A three-factor MANOVA demonstrated significant differences in the diet by sex, maturity status and oceanographic season using numeric and gravimetric measures of importance for the major prey categories. These three main factors explained more variation in diet than interactions between the factors, and season explained the most variance overall. A detailed analysis of the seasonal variation among the prey categories indicated that abundance changes in the most important prey, euphausiids, were coupled with seasonal changes in the importance of other prey. When upwelling occurred and productivity was great (Upwelling and Oceanic seasons), euphausiids were likely highly abundant in the study area and were the most important prey for B. kincaidii. As productivity declined (Davidson Current season), euphausiids appeared to decrease in abundance and B. kincaidii switched to secondary prey. At that time, gammarid amphipods and shrimps became the most important prey items and polychaetes, mysids and euphausiids were secondary.     

DIETS OF FIN, SEI, AND SPERM WHALES IN BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS, 1963–1967

Diets of fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), and sperm whales ( Physeter macrocephalus ) were estimated from the stomach contents of individuals killed along the British Columbia coast from 1963 to 1967. The dominant prey types of fin whales were euphausiids, with minor contributions from copepods and fish. Sei whale stomachs contained primarily copepods in three years, whereas euphausiids or a variety of fish dominated the diet in the other two years. Sperm whales consumed primarily North Pacific giant squid ( Moroteuthis robusta ), but secondary prey differed between males and females. Female sperm whales frequently consumed ragfish ( Icosteus spp.) and other fish, whereas the male diet also contained rockfish ( Sebastes spp.). The high abundance of euphausiids along the British Columbia coast likely contributed to the presence of a summer resident population of fin whales. The high abundance of large copepods farther north probably influenced the migration of sei whales through the offshore waters of British Columbia. Sperm whale stomach contents differed by sex reflecting location and possibly breeding behaviors.     

Key fish species, northern fur seals, Callorhinus ursinus, and fisheries interactions involving walleye pollock, Theragra chalcogramma, in the eastern Bering Sea

The recent decline of the northern fur seal population breeding on the Pribilof Islands has not yet been explained. This study estimates the amounts and sizes of walleye pollock (the fur seal's main prey) removed by predatory fish, fur seals and commercial fisheries in the main fur seal feeding area in the eastern Bering Sea during summer 1985. Fur seals relied primarily on age-1 pollock during 1985, while predatory fish consumed pollock mainly of ages 0-1. The fishery took pollock older than age-3. This indicated no direct effect of fishery removal on fur seal prey abundance, at least during the study period. In the short term, the 1985 pollock fishery may have indirectly affected fur seal prey abundance in a positive manner by removing adult pollock that compete with fur seals for age-1 pollock.     

Seasonal, diel and ontogenetic variation in feeding patterns of small yellow croaker in the central Yellow Sea

Stomach contents of 1603 small yellow croaker Pseudosciaena polyactis , sampled from seasonal bottom trawl surveys in the central Yellow Sea between March 2001 and January 2002, were examined. The results showed that small yellow croaker was a carnivorous predator and >30 prey species were identified from stomach contents analysis. Crustaceans (mainly euphausiids and decapods) were the most important prey, occurring in 93·1% of the stomachs containing food, and accounting for 77·6% of the total food by mass. Feeding activity was highest in autumn and lowest in spring and winter. Decapods were more important in summer, whereas euphausiids were more important during other seasons. Ontogenetic differences were found in the diet composition and feeding activity within the range of size (standard length, L _S) studied. The importance of fishes and decapods increased with L _S, whereas euphausiids, copepods and amphipods decreased in importance with L _S. Dietary breadth increased markedly for adults. A positive relationship was found between L _S and prey size. In each season the maximum diel feeding activity occurred at 0800 and 2400 hours, indicating that there was crepuscular and nocturnal feeding by small yellow croaker.     

Variation in the diet of Genypterus blacodes(Ophidiidae) around the Falkland Islands

The diet change with size, season and area was investigated using the stomachs of 496 kingclip Genypterus blacodes collected around the Falkland Islands (south‐west Atlantic) between August 2001 and September 2002. The key prey species were rockcod Patagonotothen spp., benthic isopods and Patagonian grenadier Macruronus magellanicus . Kingclip <50 cm total length ( L _T) fed mainly on crustaceans and small fishes. With size the diet shifted away from crustaceans towards Patagonotothen spp. in kingclip 50–100 cm L _T, and finally towards larger fishes such as M. magellanicus and Micromesistius australis australis in kingclip >100 cm L _T. The niche breadth was highest in fish >100 cm L _T and the lowest in fish <50 cm L _T. The larger kingclip generally selected larger individuals of the same prey species, with the exception of the Patagonian squid Loligo gahi , where all ingested squid were of similar size, regardless of the predator length. The importance of the main prey species varied substantially between five consequent seasons studied, and appeared to follow the seasonal abundance and availability of prey. The spatial variability in the diet was found in kingclip caught in regions occupied by transformed temperate and sub‐Antarctic waters. The rockcod, which is available throughout the year around the Falkland Islands, was the most important prey in the kingclip diet. Kingclip takes advantage of other seasonally abundant prey species during their seasonal migrations ( e.g . L. gahi ) and also scavenge on discards from fishing vessels when available.     

SUMMER FEEDING HABITS OF DALL'S PORPOISE, PHOCOENOIDES DALLI, IN THE SOUTHERN SEA OF OKHOTSK

Stomach contents were analyzed from 85 Dall's porpoises, Phocoenoides dalli , taken in the southern Sea of Okhotsk during the summer of 1988. Thirteen species of fish and five species of cephalopods were identified. Fishes comprising 9 families were predominant and made up 79.9% of the total number of prey, with an overall occurrence in the stomachs of 100%. Three families of cephalopods made up 20.1% of the total number of prey ingested, with an overall occurrence of 74%. The Japanese pilchard, Sardinops sagax melanostictus , was the primary prey and represented 72.0% of the total prey in 97.3% of the stomachs examined. The gonatid squid, Berryteuthis magister , ranked second and made up 16.9% of the total prey in 61.6% of the stomachs. Walleye pollock, Theragra chalcogramma , was the third most abundant prey item consumed and represented 6.0% of the total, with an occurrence of 53.4%. Composition of the sample by the calculated total mass and caloric intake supports the numeric findings on the relative importance of the three major prey species to the DaIl's porpoise diet. The Japanese pilchard was clearly dominant using these indices and constituted 65.4% of the total mass and 80.7% of the caloric intake. Berryteuthis magister made up 26.8% of the mass and 15.5% of the caloric intake. Walleye pollock contributed the least to the diet with 7.9% of the total mass and 3.8% of the caloric intake. Volumetric data indicate that porpoises are feeding heavily on Japanese pilchard throughout the daylight hours. Berryteuthis magister was probably fed upon in the late night or early morning hours. Occurrence of both Dall's porpoise and Japanese pilchard in the coastal waters of the southern Sea of Okhotsk is seasonal. The summer movements of Dall's porpoise into this region may be related to the seasonal migration and abundance of the Japanese pilchard.     

Feeding habits of ocellate spot skate,Okamejei kenojei (Müller & Henle, 1841), in coastal waters of Taean,Korea

The feeding habits of Okamejei kenojei were studied using 592 specimens collected in the coastal waters of Taean, Korea from April 2008 to March 2009. O. kenojei is a bottom‐feeding carnivore that consumes mainly shrimp, fishes, and crabs. Its diet also includes small quantities of amphipods, mysids, cephalopods, euphausiids, copepods, isopods, and polychaetes. The total length (TL) of individuals in this study ranged from 8.2 to 49.0 cm. Cluster analysis based on %IRI (index of relative importance) identified three size classes. Group A (< 20 cm TL) ate primarily caridean shrimp and amphipods; group B (20–30 cm TL) ate exclusively shrimp; and group C (> 30 cm TL) ate penaeoidean shrimp, fishes, and crabs. O. kenojei showed ontogenetic changes in feeding habits. Although shrimps were the primary food consumed by all size groups, the proportion of shrimp in the total diet decreased and the consumption of fishes and crabs gradually increased with the body size of O. kenojei. Size of the prey organisms also increased. Smaller individuals fed mainly on small prey, such as amphipods, mysids, and small shrimp, whereas larger individuals preferred larger prey, such as larger shrimp, fishes, and crabs. The size‐related diet breadth and the percentage of empty stomachs were significant; the diet breadth gradually increased with body size, whereas the percentage of empty stomachs decreased. Seasonal changes in the O. kenojei diet were not significant, but shrimp constituted 97.3% of the summer diet by %IRI. Seasonal changes in diet breadth and the percentage of empty stomachs were not significant.     

Stomach content analysis of minke whales Balaenoptera acutorostrata from the Lofoten and Vesterålen areas, Norway

Stomach contents from 15 minke whales Balaenoptera acutorostrata collected in the Lofoten and Vesterålen areas of Norway, August 1988, were analysed. All stomachs contained identifiable remains and constituted of fish only. Herring Clupea harengus was the dominant prey species found in all whale stomachs and constituted 92% of the total prey volume. Cod Gadus morhua was the second most abundant prey. In addition pollock Pollachius virens , haddock Melanogrammus aeglefinus , and great silver smelt Argentina silus were found in single whale stomachs. All fish species found in the whale stomachs in our investigations are of commercial importance, and the sizes of the herrings and cods in minke whale stomachs are of interest for commercial exploitation.     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua , hake Merluccius merluccius , megrim Lepidorhombus whiffiagonis , whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey ( e.g . Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey ( e.g . mackerel Scomber scombrus ), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes ( Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

Scavenging on discarded saury by demersal fishes off Sendai Bay, northern Japan

Pacific saury Cololabis saira were found in 24 out of 1314 stomachs of demersal fishes collected by bottom trawling at 245–393 m depths. The same area was occupied by the stick-held dipnet fishery for saury, in which a landing limitation was enforced to avoid overfishing. The fish species containing saury were: Pacific cod Gadus macrocephalus , walleye pollock Theragra chalcogramma , and oilfish Ruvettus pretiosus . These demersal species would not encounter living saury naturally, which normally are distributed above thermoclines. The ingested saury were smaller than market size which suggests that the saury from fish stomachs were discarded by fishermen because of small sizes to maximize profits under the landing limitation.
The scavenged saury made up 41.5, 24.6 and 77.7% of the diets of large-sized (>30cm)Pacific cod, walleye pollock and oilfish, respectively in terms of DW composition. The extent to which discarded saury contributed to the total diet in the demersal fish assemblage, calculated by considering species composition and diets of bottom fishes, was 21.8% of the total diet. The discarded saury seemed to compensate the less productive feeding environment during autumn for the bottom fishes.     

Temporal variation in chum salmon, Oncorhynchus keta, diets in the central Bering Sea in summer and early autumn

Seasonal, ontogenetic, and diel variations in the diets of chum salmon, Oncorhynchus keta, were examined by analyzing the stomach contents of 1398 fish (300–755 mm fork length) collected in the Bering Sea during summer and early autumn of 2002. Whereas mesozooplankton, including euphausiids, hyperiids, and gastropods, constituted the greatest portion of the stomach contents during the summer, forage fishes (Stenobrachius leucopsarus and Atka mackerel, Pleurogrammus monopterygius) were the most important items during early autumn. Although no apparent diel trend was found in feeding intensity, distinct diel differences in prey composition were observed. Chum salmon caught in the morning contained Stenobrachius leucopsarus, whereas those caught in the afternoon had mainly fed on euphausiids. Thus, chum salmon diets change temporally because of changes in prey availability that result from differences in the annual life cycles and diurnal vertical migrations of prey species.     

Feeding and prey-selection of wild Atlantic salmon post-smolts

The diet of post-smolt Atlantic salmon Salmo salar caught in the Trondheimsfjord and Frohavet in central Norway, based on stomach contents analysis, showed a gradual change during migration from the river to the estuary, fjord and coastal areas. Post-smolts caught in the estuary had eaten intertidal gammarid amphipods, while post-smolts caught further seawards preyed upon available marine prey such as Calanus spp., adult euphausiids and fish larvae. The frequency of adult insects was high in all post-smolt stomachs. The gradual change in diet suggested that feeding conditions in the early marine phase were important for post-smolt survival and growth. With the exception of the copepods, there was no overall similarity between species composition of the plankton samples and the stomach contents. Although the hypothesis that the post-smolts are opportunistic feeders cannot be rejected, the composition of the stomach contents suggests a possible selectivity of advantageous prey.     

WINTER FEEDING INTENSITY OF NARWHALS (MONODON MONOCEROS)

Stomach contents from 121 narwhals ( Monondon monoceros ) harvested in the eastern Canadian High Arctic and West Greenland were used to quantify seasonal changes in feeding activity and prey selection. Stomachs collected from summer harvests were mostly empty with little evidence of recent feeding. Stomachs collected in late fall and winter harvests had considerable amounts of undigested material with evidence of recent feeding. In summer, Arctic cod ( Arctogadus glacialis ), polar cod ( Boreogadus saida ), and Gonatus squid spp. constituted the narwhal diet. In fall, Gonatus fabricii was the only prey item observed. In late fall and winter, Greenland halibut ( Reinhardtius hippoglossoides ) and G. fabricii were the dominant prey items, observed in 51% and 73% of stomachs collected, respectively. Greenland halibut taken by narwhals were on average 39 cm (SD 8) and 556 g (306) and G. fabricii were on average 23 g (15) with mean mantle lengths of 85 mm (24). The low diversiry of prey species indicates narwhals have a restricted diet across all seasons. This study presents the first information on the winter diet of the narwhal and suggests Baffin Bay and Davis Strait are heavily utilized for feeding, in contrast to limited food intake during the summer period.