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.     

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.     

Endurance of simulated winter conditions by age-0 walleye pollock: effects of body size, water temperature and energy stores

Survival of age-0 walleye pollock Theragra chalcogramma in the absence of food followed simple bioenergetic models, with large body size, high initial condition, and cold temperatures all increasing survival rates. High survival after >200 days at cold temperatures (<3·0° C) indicated extended tolerance of extreme cold, as long as sufficient body size and condition are attained during the summer growth period. Analysis of body constituents demonstrated a substantial increase in tissue water and depletion of lipid during starvation. Survivors had significantly higher lipid stores than mortalities, and larger fish had higher levels of lipid than smaller fish among experimental survivors, laboratory fish that were never starved, and wild fish. Fish returned to warm temperatures and high rations following 205 days of food deprivation displayed nearly complete recovery, with rapid increases in length, weight, and condition and minimal mortality (6·8%) during the subsequent 3 months. Age-0 walleye pollock collected in September in the Bering Sea were substantially smaller and generally had lower lipid levels than fish used in laboratory starvation experiments, suggesting they are susceptible to size- and condition-dependent mortality during the winter. The results are interpreted with respect to field distributions of age-0 walleye pollock, overwinter survival, and synergistic effects of food and temperature under varying models of climate change.     

Otolith size and location in digestive tracts of northern fur seals (Callorhinus ursinus): Implications for dietary interpretations

Walleye pollock (Theragra chalcogramma) otoliths (n= 2,706) recovered from stomachs, small intestines, and colons of 43 northern fur seals (Callorhinus ursinus) were evaluated for size and wear by location in the digestive tract. Pollock fork length was regressed on otolith length after correction for erosion, and age was estimated from the calculated body size. Age‐1+ pollock otoliths (≥6.3‐mm length) were concentrated in stomachs while age‐0 otoliths (≤6.2‐mm length) were concentrated in colons. Less than 10% of otoliths were found in the small intestines. Pollock age decreased with progression along seal gastrointestinal tracts. Otolith quality increased along gastrointestinal tracts in numbers ≥20, which was typical of age‐0 otoliths recovered from colons. Otolith distribution by age and quality along gastrointestinal tracts suggests that small (≤12 cm) schooling prey are consumed in large volume and passed as a bolus rapidly through the digestive tract before significant erosion of bony remains occurs; while larger prey are eaten in smaller volume and subjected to otolith erosion due to longer retention in the stomach. Our results illustrate the importance of multiple sampling strategies to comprehensively represent prey size in pinniped diet.     

Spring cannibalism on 1 year walleye pollock in the Doto area,northern Japan: is it density dependent?

Cannibalism in walleye pollock off the eastern coast of the Hokkaido Island, Japan was important only during spring (April to June), and its importance increased from 0% in dry mass for <200 mm L _S fish to 48·9% for >400 mm L _S fish. Most of the prey was represented by age 1 year fish, showing a unimodal body size distribution with a mode at 121–130 mm. Although cannibal body size was larger in deeper (>150 m) water, there was no difference in prey size by depth, suggesting impingement of the predators inhabiting deeper water into the shallow areas to cannibalize 1 year fish. The minimum ratio cannibal: prey size was 1·74. There was a positive but non-significant correlation between the contribution of cannibalism to a potential predator's (>300 mm) diet and an estimate of the previous year's recruitment. This was due to an extremely high contribution of cannibalism during 1992, when a distinctly larger size of predators seemed to bias the contribution. When the 1992 data were removed from the analysis, a significant correlation was obtained ( r ²=0·77, P <0·01), showing that Pollock cannibalism is rather density dependent. Based on the results, it is hypothesized that the'overflow' of 1 year fish from the shelf waters due to their high abundance and the weak stratification in the spring water column results in increased co-occurrence with adult fish and consequent cannibalism.     

Predation of Japanese dace, Tribolodon hakonensis, by largemouth bass, Micropterus salmoides, in experimental aquaria

 To test the size range of prey fish that largemouth bass, Micropterus salmoides, can successfully consume, live Japanese dace, Tribolodon hakonensis, were given as prey fish to individual largemouth bass in aquaria. The ratio of maximum standard length (SL) of the Japanese dace consumed by largemouth bass was 46–69% of bass SL. The maximum length of Japanese dace consumed did not differ significantly between largemouth bass and smallmouth bass (M. dolomieu) previously studied, although largemouth bass have relatively larger mouth sizes than smallmouth bass. Largemouth bass occasionally injured and killed Japanese dace larger than the limit that could be consumed.     

Development and application of a bioenergetics model for juvenile walleye Pollock

A bioenergetics model was parameterized for age-0 walleye pollock, Theragra chalcogramma , based on a synthesis of literature data. The sensitivity of the new parameters was tested by individual parameter perturbation (IPP) analysis. The model was applied to estimate individual and total cohort food consumption of age-0 pollock in two areas of high pollock density in the Gulf of Alaska during the summer of 1990. Total cohort consumption was compared with zooplankton biomass and production estimates for the same areas and times of the year. The model was also used to examine the bioenergetic implications of age-0 pollock diel vertical migration through a thermal gradient. During a 1-month bioenergetics simulation, individual daily consumption decreased from 16·0 to 6·0% of wet body weight. Daily ration estimates corresponded well with independent field estimates of daily ration for the same areas and time of the year. Comparison of total cohort consumption with prey availability (production and biomass) indicated minimal potential for food limitation. Bioenergetic optimization of growth can be a potential benefit of diel vertical migration to age-0 pollock, however more information on prey density and distribution is needed to test this hypothesis thoroughly.     

Diet composition of starry smooth-hound Mustelus asterias and methodological considerations for assessing the trophic level of predatory fish

The stomach contents of 640 starry smooth-hound Mustelus asterias from the north-east Atlantic were examined. The diet was dominated by crustaceans (98.8% percentage of index of relative importance, %IRI), with the two main prey species being hermit crab Pagurus bernhardus (34% IRI) and flying crab Liocarcinus holsatus (15% IRI). Ontogenetic dietary preferences showed that smaller individuals [20–69 cm total length (L_T) n = 283] had a significantly lower diversity of prey than larger individuals (70–124 cm L_T, n = 348); however, 18 prey species were found exclusively in smaller individuals and eight prey taxa were found exclusively in larger individuals. Larger commercially important brachyurans such as edible crab Cancer pagurus and velvet swimming crab Necora puber were more prevalent in the diet of larger individuals. Specimens from the North Sea ecoregion had a lower diversity of prey types for a given sample size than fish from the Celtic Seas ecoregion. Whilst cumulative prey curves did not reach an asymptote, this was primarily due to the high taxonomic resolution utilized and 95% of the diet was described by just seven crustacean taxa. The trophic level (TL) was calculated as 4.34 when species-level prey categories were used. This fine-scale taxonomic resolution resulted in a TL estimate close to a whole level above that estimated using wider taxonomic groupings. This large bias has important methodological implications for TL studies based on categorized prey data, particularly those of predatory fish.     

Modelling Size-Dependent Cannibalism in Barramundi Lates calcarifer: Cannibalistic Polyphenism and Its Implication to Aquaculture

This study quantified size-dependent cannibalism in barramundi Lates calcarifer through coupling a range of prey-predator pairs in a different range of fish sizes. Predictive models were developed using morphological traits with the alterative assumption of cannibalistic polyphenism. Predictive models were validated with the data from trials where cannibals were challenged with progressing increments of prey sizes. The experimental observations showed that cannibals of 25–131 mm total length could ingest the conspecific prey of 78–72% cannibal length. In the validation test, all predictive models underestimate the maximum ingestible prey size for cannibals of a similar size range. However, the model based on the maximal mouth width at opening closely matched the empirical observations, suggesting a certain degree of phenotypic plasticity of mouth size among cannibalistic individuals. Mouth size showed allometric growth comparing with body depth, resulting in a decreasing trend on the maximum size of ingestible prey as cannibals grow larger, which in parts explains why cannibalism in barramundi is frequently observed in the early developmental stage. Any barramundi has the potential to become a cannibal when the initial prey size was <50% of the cannibal body length, but fish could never become a cannibal when prey were >58% of their size, suggesting that 50% of size difference can be the threshold to initiate intracohort cannibalism in a barramundi population. Cannibalistic polyphenism was likely to occur in barramundi that had a cannibalistic history. An experienced cannibal would have a greater ability to stretch its mouth size to capture a much larger prey than the models predict. The awareness of cannibalistic polyphenism has important application in fish farming management to reduce cannibalism.     

Morphological limitations, prey size selectivity, and growth response of juvenile atlantic salmon, Salmo salar

The head and jaw movements involved in capture, buccal manipulation, ingestion and rejection of prey were investigated using sequential photography of juvenile Atlantic salmon feeding in a simulated stream environment. The results are described and discussed and mouth breadth and gill raker spacing are proposed as morphometric limitations to the range of prey sizes available which remains constant at 0·06 · fish fork length ( PFR ).
A recirculatory flume tank was used to study prey size selectivity behavior. Simplified downstream-drifting prey items elicited a variety of responses depending on their physical size. One hundred percent of offered prey of PFR 0·025 were ingested, while 90 % of prey at PFR 0·051 and 100% of prey at PFR 0·105 were rejected. It is demonstrated that fish show negative selection for prey sizes smaller than PFR 0·025 and that prey of this size elicits maximum growth response.
The validity of the proposed morphometric limitations on the available prey sizes is demonstrated by reference to selectivity behaviour and prey size related differential growth.     

Regional, interannual and size-related variation of age 0 year walleye pollock whole body energy content around the Pribilof Islands, Bering Sea

Mean whole energy content ( E _wb) of age 0 year walleye pollock Theragra chalcogramma was 19.928 KJ g⁻¹ dry mass in 3943 fish collected from different habitats around the Pribilof Islands frontal structure, south-east Bering Sea, during September 1994–1996 and 1999. It varied, however, with habitat type. Fish residing offshore had higher E _wb than fish residing inshore of the frontal regions. Age 0 year walleye Pollock E _wb changed in a non-linear fashion with fish size, with larger juveniles typically having higher E _wb. Size thresholds were identified at which the relationship between age 0 year walleye pollock E _wb and L _S changed. One such threshold was found at 46 mm where E _wb reached a local minimum. Another threshold was found at 80 mm beyond which E _wb tended to remain constant with size. Overall mass-length and E _wb-length residuals were highly correlated with each other ( r =0.73, P ≪0.0001). The slope of the regression, however, was higher for smaller fish. Possible mechanisms are proposed to explain the observed ontogenetic variation in nutritional status and the role of age 0 year walleye pollock late summer E _wb on survival over their first critical winter of life.     

Response of predators to prey abundance: separating the effects of prey density and patch size

We tested the relative and combined effects of prey density and patch size on the functional response (number of attacks per unit time and duration of attacks) of a predatory reef fish (Cheilodactylus nigripes (Richardson)) to their invertebrate prey. Fish attacked prey at a greater rate and for longer time in large than small patches of prey, but large patches had naturally greater densities of prey. We isolated the effects of patch size and prey density by reducing the density of prey in larger patches to equal that of small patches; thereby controlling for prey density. We found that the intensity at which fish attacked prey (combination of attack rate and duration) was primarily a response to prey density rather than the size of patch they occupied. However, there was evidence that fish spent more time foraging in larger than smaller patches independent of prey density; presumably because of the greater total number of prey available. These experimental observations suggest that fish can distinguish between different notions of prey abundance in ways that enhance their rate of consumption. Although fish may feed in a density dependent manner, a critical issue is whether their rate of consumption outstrips the rate of increase in prey abundance to cause density dependent mortality of prey.     

Feeding on Profitable and Unprofitable Prey: Comparing Behaviour of Growth-Enhanced Transgenic and Normal Coho Salmon (Oncorhynchus kisutch)

We compared the performance of normal and growth hormone‐transgenic coho salmon feeding on surface drifting edible and inedible novel prey items in various social environments. With an inherently higher appetite, we predicted that transgenic fish would be more willing to feed on novel prey, and that visual company with another fish would enhance this difference further. Transgenic and normal fish, of similar size and age, were equally willing to attack both the edible (live insects) and inedible (artificial angling lure flies) prey, but transgenic fish did so faster and were more likely to make repeated attacks. Transgenic fish managed to seize and consume the edible prey after fewer attacks than did normal fish. However, swallowing of prey took longer than for normal fish. More transgenic individuals interacted with the inedible prey compared with normal salmon, and initially, transgenic fish in visual company with another fish also interacted more with the prey than single transgenic or any constellation of normal focal fish. With repeated exposures, the number of individuals attacking and the number of interactions with the prey decreased. These responses were stronger in transgenic fish, partly explained by the initially low response in normal fish. The observed differences are most likely the consequences of elevated levels of growth hormone in transgenic fish generating enhanced feeding motivation and reinforcement capacity. In a natural environment, the performance of a growth hormone‐transgenic fish may therefore depend on the relative abundance of profitable vs. unprofitable prey, as well as the presence of other transgenic individuals.     

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.     

Prey orientation in piscivorous brown trout

Piscivorous brown trout Salmo trutta change their feeding behaviour depending on prey species, prey size and number of prey eaten. In trout which had eaten fish recently, most had one fish in their stomach, but up to 16 prey fish were found. Individuals of the small-sized minnow Phoxinus phoxinus were swallowed chiefly tail first, whereas individuals of the larger Arctic charr Salvelinus alpinus were taken both head and tail first. The largest charr were swallowed head first. In stomachs containing more than one fish prey, prey orientation was likely to be mixed. For all three types of prey orientation (i.e. tail first, head first and mixed), significant and positive correlations existed between prey length and predator length. The maximum prey size eaten tail first or in mixed orientation was about 70–85% of the size of prey eaten head first, indicating morphological advantages in eating the prey head first.     

Ecological redundancy between coral reef sharks and predatory teleosts

Reef sharks may be ecologically redundant, such that other mesopredatory fishes compensate for their functions when they decline in number, preventing trophic cascades. Oral jaw gape, hereafter referred to as gape, determines maximum prey size in many piscivores and therefore affects the size structure of prey assemblages. Here, we examine whether gape and maximum prey size differ between five species of reef shark and 21 species of teleost (n?=?754) using data collected from 38 reefs in the Indo-Pacific. Sharks displayed relatively small gape dimensions compared to most teleost species and, at smaller sizes, the giant trevally Caranx ignobilis and other teleosts may be able to consume larger prey than similar-sized sharks. However, ecological redundancy between reef sharks and teleosts appears to decline at larger sizes, such that the grey reef shark Carcharhinus amblyrhynchos, for example, may be capable of consuming larger prey than any other reef predator at its largest sizes, regardless of prey body shape. Moreover, sharks may be able to consume proportionally larger prey as they grow, in contrast to reef teleosts, which may largely be limited by their gapes to ever-smaller prey as a proportion of their body size. Our results also suggest that reef sharks may be unable to swallow whole prey that are >?36% of their length, consistent with gut-content studies. Conservation of reef ecological function may therefore depend not only on the protection of sharks but also particular size classes and key components of the mesopredatory guild.

     

Ontogenetic variations in the type and size of prey consumed by juvenile coho,Oncorhynchus kisutch,and chinook,O. tshawytscha,salmon

Synopsis Stomach contents of juvenile coho,Oncorhynchus kisutch, and chinook,O. tshawytscha, salmon collected in purse seines off the coast of Washington and Oregon were examined for variations related to predator size. There was a general trend toward increasing consumption of fish with increasing body size, due mainly to the increase in northern anchovy biomass consumed by the larger salmon. Most of the major prey taxa showed significant differences among the size classes examined for both salmon species. There was a direct relationship between predator and prey size for both coho and chinook, but considerable variation was found in prey length consumed within each size class. Prey width did not provide as good a fit as prey length for either species. In general, coho consumed larger fish prey in relation to their body length than chinook but there were substantial differences by month or year of collection.     

Prey size selection and cannibalistic behaviour of juvenile barramundi Lates calcarifer

This study assessed the cannibalistic behaviour of juvenile barramundi Lates calcarifer and examined the relationship between prey size selection and energy gain of cannibals. Prey handling time and capture success by cannibals were used to estimate the ratio of energy gain to energy cost in prey selection. Cannibals selected smaller prey despite its capability of ingesting larger prey individuals. In behavioural analysis, prey handling time significantly increased with prey size, but it was not significantly affected by cannibal size. Conversely, capture success significantly decreased with the increase of both prey and cannibal sizes. The profitability indices showed that the smaller prey provides the most energy return for cannibals of all size classes. These results indicate that L. calcarifer cannibals select smaller prey for more profitable return. The behavioural analysis, however, indicates that L. calcarifer cannibals attack prey of all size at a similar rate but ingest smaller prey more often, suggesting that prey size selection is passively orientated rather than at the predator's choice. The increase of prey escape ability and morphological constraint contribute to the reduction of intracohort cannibalism as fish grow larger. This study contributes to the understanding of intracohort cannibalism and development of strategies to reduce fish cannibalistic mortalities.     

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.     

Prey size,prey abundance,and temperature as correlates of growth in stream populations of cutthroat trout

Growth and maximum size of stream fishes can be highly variable across populations. For salmonid fishes in streams, individuals from populations confined to headwater streams often exhibit small size at maturity in comparison to populations with access to main-stem rivers. Differences in prey size, prey availability, and metabolic constraints based on temperature may explain patterns of maximum size and growth. In this study, cutthroat trout from headwater stream populations that were isolated above a waterfall were compared to individuals from populations in similar sized streams without a movement barrier and from large main-stem rivers. Cutthroat trout from smaller streams with or without a movement barrier were significantly smaller at a given age than fish from main-stem rivers, where individuals were able to achieve a much larger maximum size. Comparisons of invertebrate drift abundance and size in the three types of streams revealed that drift size did not differ between stream categories, but was highest per volume of water in large main-stem rivers. Across all stream types, prey abundance declined from summer to fall. Temperature declined over the course of the season in a similar manner across all stream types, but remained relatively high later in the season in main-stem river habitats. Prey availability and temperature conditions in main-stem rivers may provide more optimal growing conditions for fish as individuals increase in size and become constrained by prey availability and temperature conditions in small streams. Maintaining connectivity between small spawning and rearing tributary streams and main-stem river habitats may be critical in maintaining large-bodied populations of stream salmonids.