Long‐term changes in the feeding of Pollachius virens on the Scotian Shelf: responses to a dynamic ecosystem

The diet and feeding ecology of pollock Pollachius virens from the Scotian Shelf and Bay of Fundy in the north‐west Atlantic changed over the last few decades, which was associated with decreases in euphausiid abundance. Stomach contents data for 2078 pollock collected during the 1958–1967 period and for 1230 pollock collected during the 1996–2002 period indicated that pollock diet contained fewer euphausiids and feeding activity decreased. During the early period, euphausiids were present in 65% of the pollock stomachs that contained food and only 9% of these stomachs in the recent period. The decrease of euphausiids was not wholly offset by an increase in piscivory, since there was little increase in the frequency of fish prey in the diet or in the fullness index for this prey type. Empty stomachs were significantly more common in the recent period during both the winter and summer. The decreased occurrence of euphausiids in stomach samples coincided with a significantly decreased abundance of this prey, suggesting that the near‐absence of euphausiids in recently collected pollock stomachs reflected prey abundance. Concurrent with changes in diet and feeding intensity, the condition or 'plumpness' of pollock significantly declined from the early to the late sampling periods.     

The potential for intracohort cannibalism in age-0 walleye pollock,Theragra chalcogramma,as determined under laboratory conditions

Synopsis In laboratory experiments, we tested the capability of larger age-0 walleye pollock to consume smaller members of their cohort. In separate aquaria, 81 pairs of juveniles covering a wide range of size differences (total lengths differing by 12 to 61 mm) were held and monitored over a 4 day period. Complete consumption, in which a smaller fish was swallowed whole by a larger fish, occurred 11% of the time. In 36% of the pairs, attacks by the larger fish resulted in mortality of the smaller fish. The mouth width:body depth ratio between the larger and smaller fish of a pair differed significantly depending on whether the smaller fish survived, was killed but not consumed, or was ingested whole by the larger fish. Cannibalistic individuals could consume fish close to the maximum size physically possible under gape limitation; at this size the length of the cannibal was approximately 1.7 times the length of the prey. Length-frequency distributions of age-0 pollock in field concentrations suggested that, at least in some geographical areas, potential cannibals and prey commonly co-occur. Unsuccessful predatory attacks by larger individuals may have additional detrimental effects on smaller pollock in natural populations.     

Intertidal recruitment and feeding in relation to optimal utilization of nursery areas by juvenile English sole (Parophrys vetulus: Pleuronectidae)

Synopsis Juvenile Parophrys vetulus 19–102 mm were collected on an intertidal flat in Humboldt Bay during 1976–1977. Recently metamorphosed fish that fed almost exclusively on harpacticoid copepods and other epibenthic crustaceans were found in large numbers during spring and early summer. A feeding transition occurred among fish 50–65 mm in length, and infaunal polychaetes were the dominant prey of fish greater than 65 mm. Increases in the feeding niche width and average size of prey items, as well as a decrease in the number of prey items per stomach, accompanied growth.The disappearance of English sole from intertidal areas in early fall at an average size of 82 mm and their subsequent residence in subtidal channels until a size of 140 mm suggest a size-depth segregation within the nursery ground. The advantages of such a distribution are discussed in terms of optimal foraging and reduction of intra- and interspecific competition.     

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.     

Investigation of foraging habits and prey selection by humpback whales (Megaptera novaeangliae) using acoustic tags and concurrent fish surveys

Tags containing acoustic time-depth transmitters (ATDT) were attached to four humpback whales near Kodiak, Alaska. Tags allowed for whale dive depths to be recorded in real time. Acoustic and mid-water trawl surveys were conducted concurrent with tagging efforts within the study area to quantify available fish resources and describe potential prey selection by humpback whales. Recorded dives were grouped through visual assessment and t -tests. Dives that indicated likely foraging occurred at a mean maximum depth of 106.2 m with 62% of dives occurring between 92 m and 120 m. Acoustic backscatter from fish surveys was attributed to potential humpback prey based on known target strength values and 10 net tows. Capelin comprised 84% of the total potential prey abundance in the region followed by age 0 (12%) and juvenile pollock (2%), and eulachon (<1%). Although horizontally segregated in the region, both capelin and age 0 pollock were distributed at depths exceeding 92 m with maximum abundance between 107 m and 120 m. The four-tagged humpbacks were found to forage in areas with greatest capelin densities but bypassed areas of high age 0 pollock abundance. The location and diving behavior of tagged whales suggested that whales were favoring capelin over pollock as a prey source.     

Seasonal changes in the diel vertical migration of walleye pollock (<Emphasis Type="Italic">Theragra chalcogramma</Emphasis>) in the northern Gulf of Alaska

Walleye pollock (Theragra chalcogramma) perform diel vertical migration (DVM) as juveniles, but have an increasing tendency to be associated with the bottom with age. We studied the DVM of a local population of adult pollock in the northern Gulf of Alaska in August and November 2003. There was no relationship between the depth of pollock and the isolume (line of equal light intensity) necessary for visual foraging in August. Pollock passed through the thermocline at this time. In November there was a significant relationship between pollock biomass above/below the 200 m isobath and the isolume necessary for visual foraging. It is hypothesized that in August pollock ignore the isolume and thermocline, simply tracking the movements of their prey (euphausiids) to feed upon them near the surface at night. In November, relatively denser pollock shoals migrate up and down with the isolume necessary for visual foraging to feed on decapods.     

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.     

Morphology and diet are decoupled in nearshore notothenoids from King George Island,West Antarctica

Antarctic notothenioid fishes show wide adaptive morphological radiation, linked to habitat preferences and food composition. However, direct comparisons of phenotypic variability and feeding habits are still lacking, particularly in stages inhabiting nearshore areas. To assess these relationships, we collected juveniles and adults of the most common benthic species inhabiting shallow waters off the South Shetland Islands within a similar size range, the plunderfish Harpagifer antarcticus, the black rockcod Notothenia coriiceps, and the marbled rockcod Notothenia rossii. Individual size ranges varied from 44.0 to 98.9 mm standard length (LS) (H. antarcticus), from 95.8 to 109.3 mm LS (N. coriiceps), and from 63.0 to 113.0 mm LS (N. rossii). Notothenioid fish showed different morphospace variability, being larger for H. antarcticus than the other Notothenia species and associated with the position of the posterior end of the operculum, along with the location and relative size of the eye. The evolutionary allometry was low, but the static allometry was much higher, especially for H. antarcticus and N. rossii. The diet was mainly carnivorous, consisting of amphipods and euphausiids. Macroalgae were scarce or totally absent in the gut contents of all species. Only H. antarcticus showed an increase in the prey number and ingested prey volume with fish size. Finally, there was a significant covariation between shape changes and LS in all species (allometric effects), however, not with prey composition, probably due to the small size range or ontogenetic stage and the relative similarity (or lack of contrast) in the benthic environment that they utilized.     

Changing handling times during feeding and consequences for prey size selection of 0+ zooplanktivorous fish

The interrelationship of fish size, prey size and handling time within a 15-min feeding period was studied in three size groups of 0 + roach, Rutilus rutilus, and bleak, Alburnus alburnus. Four size classes of cladoceran prey were used to measure changes in feeding rate and handling time from initial rapid feeding to sustained feeding. Observed differences in increase of handling time between prey size classes led to a change in the prey profitability ranking of those size classes within the first 2 min of the experiments. A 1-min feeding period is interpreted as reflecting an intermediate motivational status between extreme hunger and satiation. The use of average handling times for this period revealed a substantial change in prey profitability estimates compared to previous studies which used handling times based on short-term (a few seconds up to 1 min) feeding. It is not the largest prey items a fish can handle and swallow that are most profitable, but prey of intermediate size. By this approach a closer fit between expectations derived from optimal foraging theory and empirical data on prey size selection of 0 + zooplanktivorous fish is qualitatively achieved. Optimal prey size was found to be close the mouth gape width in small fish of 15 mm standard length, decreasing to 50% of mouth gape width in fish of 40 mm standard length.     

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.     

Feeding habits of megrim, Lepidorhombus whiffiagonis (Walbaum, 1792), from the central Adriatic Sea

The feeding habits of the megrim, Lepidorhombus whiffiagonis , from the central Adriatic Sea were investigated with respect to fish size and seasons. Stomach contents of 510 specimens, 9.0–28.9 cm total length (TL), collected by bottom-trawl from 2004 to 2006 were analyzed. The percentage of 97 empty stomachs varied significantly with season (maximum during the winter, 32.1%; minimum during summer, 7.4%). The food composition suggests that the megrim is a carnivorous fish. Prey items identified in the stomachs belong to four major groups: Teleostei, Decapoda, Mysidacea and Cephalopoda. Teleosts were the most important (%IRI = 82.2), especially in fish larger than 14 cm TL. Mysidacea constituted the main prey for fish less than 14 cm TL. At the species level, the teleost Antonogadus megalokynodon (%IRI = 1.5) and decapod Processa canaliculata (%IRI = 0.9) were the most frequent prey. The mean weight of stomach contents increased significantly for fish larger than 14 cm TL, while the mean number of prey items did not differ significantly among size classes. Diet composition showed little seasonal variation; teleosts were the most important prey in all seasons, especially during spring. The lowest intensity of feeding recorded in winter could be related to the lower sea temperature and to the spawning period.     

Do abiotic and ontogenetic factors influence the diet of a generalist predator? Feeding ecology of the Pacific spiny dogfish (<Emphasis Type="Italic">Squalus suckleyi</Emphasis>) in the northeast Pacific Ocean

Stomach contents from 1221 Pacific spiny dogfish (Squalus suckleyi) were collected from the Gulf of Alaska over 2004–2006 and analyzed to determine predominant prey species. Pacific spiny dogfish in the Gulf of Alaska have a variable diet suggesting that they are generalist feeders. Of the 68.30% of stomachs with prey contents, the most important prey groups based on the percent prey-specific index of relative importance (%PSIRI) were shrimp (27.06%), cephalopods (17.16%), and forage fish (17.11%). Remaining components of the diet were inconsistent. Commercially valuable species, such as salmon, Oncorhynchus spp., and rockfish, Sebastes spp. constituted only 2.37% and 1.51% of the diet of Pacific spiny dogfish, respectively. Diet diversity was not significantly influenced by spatial, temporal or ontogenetic factors; however, trophic level of prey and average prey item weight varied both interannually and with ontogeny. With increasing size, Pacific spiny dogfish incorporate larger prey items into their diet, and diet composition may be driven more by seasonal availability and prey size than any other factors.     

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.     

Diet and feeding habits of Economidichthys pygmaeus (Perciformes: Gobiidae) in Lake Pamvotis,NW Greece

The purpose of the study was to define the diet composition, feeding preferences, seasonal, size and sex related changes in diet, feeding strategy and diel cycle of Economidichthys pygmaeus. Important features of the overall feeding patterns of the goby include (i) feeding activity, (ii) bottom habits and (iii) consumption of food items to identify the dietary breadth. Stomach contents of 533 specimens, 13.42–48.65 mm total length (TL), collected by hand net (2 mm mesh size) from October 2006 to September 2007 were analyzed. The percentage of 76 empty stomachs (14.26%) did not vary significantly with season. The food composition suggested that the goby is a carnivorous fish. Prey items identified in the stomachs belonged to four major groups: Crustaceans, Insects, Mollusca and Plants. Copepods were the most important (%IRI = 57.51), especially in fish larger than 23 mm TL. Chironomids (%IRI = 43.23) constituted the main prey for fish <23 mm TL. Diet composition showed little seasonal variation. Copepoda dominated the diet in autumn and winter (43.4%; 82.4%) and were replaced by Chironomidae larvae in spring (39.7%) and summer (47.7%). During the reproductive season, large males show a narrow food spectrum as opposed to large females, reflecting the different activity pattern exhibited by males and females during the breeding season, since the former supports parental care. According to the modified Costello graphical method, specialist individuals form the goby population that feeds on two preferential prey types (e.g. Copepods, Chironomids). Nevertheless, they consume some occasional prey (e.g. Cladocera, Insects, Gastropods). The species foraged chiefly at night and early morning, during lower light intensity. The highest feeding activity recorded at night (02.00 hours) and during early morning (08.00 hours) could be related to the cryptic behaviour displayed by the species.     

黄海中部高眼鲽的摄食及随体长的变化

对2004年1月-2005年10月在黄海中部采集的622尾高眼鲽样品的胃含物进行分析.结果表明：高眼鲽摄食的食物种类包括11类38种,其中甲壳类和瓣鳃类为优势饵料类群,共占食物相对重要性指数百分比的99%;食物中的优势饵料种类为太平洋磷虾、脊腹褐虾和樱蛤.与20世纪80年代相比,黄海高眼鲽的摄食强度下降,食物组成发生了较大的变化;鱼类不再是优势饵料类群,鳀鱼在食物中所占的比例明显下降.通过聚类分析和列联表检验等方法对高眼鲽摄食随体长的变化进行研究,结果表明：体长在51～370 mm范围内的高眼鲽的摄食强度没有显著的体长变化,但不同体长组摄食的食物种类有一定差异.＜119 mm的主要以磷虾类和底层虾类为食;体长在120～199 mm的除以磷虾类和底层虾类为主要食物外,还摄食相当比例的瓣鳃类和鱼类;＞200 mm的则主要摄食底层虾类和鱼类,磷虾类所占比例减小.     

Electron microprobe analysis of juvenile walleye pollock,Theragra chalcogramma,otoliths from Alaska: a pilot stock separation study

Synopsis The incorporation of dissolved oceanic constituents in the otoliths of fish has potential as a chemical tracer for reconstructing the early life history of marine fish. Wavelength dispersive spectrometers on an electron microprobe were used to measure Na, Mg, P, S, Cl, K, Ca, and Sr concentrations on the outer margins of 57 juvenile walleye pollock, Theragra chalcogramma, otoliths from five locations in the Gulf of Alaska and Bering Sea. Discriminant analyses that used various combinations of Na, P, K, Sr, and fish standard length and/or age showed that 60–80% of the samples could be assigned to the correct capture locality. While the concentrations of some of the measured elements correlated with standard length or age of the fish, there are measurable differences among localities when concentrations are length or age corrected, mainly due to differences in Na and K concentrations. Elemental composition of otoliths potentially could be used to assign fish from a mixed stock fishery to original stocks, information that is greatly needed for the effective management of fish stocks.     

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.     

The food of barramundi, Lates calcarifer (Bloch), in coastal and inland waters of Van Diemen Gulf and the Gulf of Carpentaria, Australia

Examination of the diet of Lates calcarifer , based on 3684 specimens of 4–1200 mm total length (T.L.), from two localities in north Australia showed it is an opportunistic predator with an ontogenetic progression in its diet from microcrustacea to macrocrustacea to fish. The increasing importance of fish in the diet of L. calcarifer in salt water (up to 87% of food consumed) involved a dietary shift from feeding on bottom-dwelling forms to pelagic forms. There were few differences in the diet of L. calcarifer between the two localities. Cannibalism was rarely encountered in the Gulf of Carpentaria although it was significant (11–4% of the diet of L. calcarifer of 1001–1200 mm T.L.) in Van Diemen Gulf. A significant correlation between predator and prey length was found for Ariidae, Dorosomidae, Engraulidae, L. calcarifer , Mugilidae and Polynemidae. The maximum prey/predator length ratio observed was 0–61 for the dorosomid, Nematolosa erebi.