Climate and Demography Dictate the Strength of Predator-Prey Overlap in a Subarctic Marine Ecosystem

There is growing evidence that climate and anthropogenic influences on marine ecosystems are largely manifested by changes in species spatial dynamics. However, less is known about how shifts in species distributions might alter predator-prey overlap and the dynamics of prey populations. We developed a general approach to quantify species spatial overlap and identify the biotic and abiotic variables that dictate the strength of overlap. We used this method to test the hypothesis that population abundance and temperature have a synergistic effect on the spatial overlap of arrowtooth flounder (predator) and juvenile Alaska walleye pollock (prey, age-1) in the eastern Bering Sea. Our analyses indicate that (1) flounder abundance and temperature are key variables dictating the strength of flounder and pollock overlap, (2) changes in the magnitude of overlap may be largely driven by density-dependent habitat selection of flounder, and (3) species overlap is negatively correlated to juvenile pollock recruitment when flounder biomass is high. Overall, our findings suggest that continued increases in flounder abundance coupled with the predicted long-term warming of ocean temperatures could have important implications for the predator-prey dynamics of arrowtooth flounder and juvenile pollock. The approach used in this study is valuable for identifying potential consequences of climate variability and exploitation on species spatial dynamics and interactions in many marine ecosystems.     

Interannual differences in growth and hatch date distributions of age-0 year walleye pollock Theragra chalcogramma (Pallas) sampled from the Shumagin Islands region of the Gulf of Alaska, 1985 – 2001

Daily increments of age-0 year walleye pollock Theragra chalcogramma otoliths from 1985 to 2001 were examined for interannual variability in growth and hatch dates. Fish were collected in summer and autumn surveys near the Shumagin Islands in the western Gulf of Alaska (GOA). Hatch date distributions of these fish were compared with hatch dates of larvae from spawning aggregations located in Shelikof Strait and the Shumagin Islands based on their spawning times. The hatch date distributions of age-0 year fish captured from the Shumagin Islands area were similar to those observed for larvae from the Shelikof Strait spawning group. Age-0 year fish whose hatch dates corresponded to the Shumagin Islands spawning, which occurred earlier in January and February, were not found. Sea surface temperature was associated with variability in hatch date distribution and growth.     

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.     

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.     

A CRITICAL REVIEW OF THE REGIME SHIFT-"JUNK FOOD"-NUTRITIONAL STRESS HYPOTHESIS FOR THE DECLINE OF THE WESTERN STOCK OF STELLER SEA LION

Steller sea lions ( Eumetopias jubatus ) in the central and western Gulf of Alaska, Aleutian Islands, and Bering Sea have declined by 80% in the last 30 yr. One hypothesis for the decline in this western Steller sea lion population is that a climate regime shift in 1976–1977 changed the species composition of the fish community and reduced the nutritional quality (energy density) of the sea lion prey field. This in turn led to nutritional stress and reduced individual fitness, survival, and reproduction of sea lions. Implications of this regime shift-"junk food" hypothesis are that (1) the recruitment and abundance of supposed high quality species ( e.g. , Pacific herring, Clupea pallasi ) decreased while those of supposed low quality ( e.g. , species in the family Gadidae) increased following the regime shift, (2) Steller sea lion diets shifted in response to this change in fish community structure, and (3) a diet composed principally of gadids ( e.g. , walleye pollock, Theragra chalcogramma ) is detrimental to sea lion fitness and survival. We examine data relating to each of these implications and find little support for the hypothesis that increases in the availability and consumption of gadids following the regime shift are primarily responsible for the decline of the western population of Steller sea lion.     

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.     

Evaluation of the productivity of walleye pollock generations in the northern Sea of Okhotsk

Based on data on the abundances of various age groups in the northern Sea of Okhotsk in 1984–2008, the productivity of 33 generations of walleye pollock born between 1975 and 2007 was evaluated in two ways. The first approach allowed a retrospective estimation by abundance of the first nine one-year age groups, and the second one was a perspective estimation using the abundance index of the nearest recruitment—3- and 4-year-olds—that enabled us to evaluate the productivity of a generation before it enters into spawning and commercial stocks. These productivity estimates were compared to the data of annual catches and the dynamics of spawning stock for many years. A high value of walleye pollock stock was shown to be supported by several productive and highly productive generations, a medium value—by one productive and one moderately productive or by two moderately productive generations, and a low value—by one productive, or moderately productive, and low productive generations. In addition, overall mortality rates were calculated for generations with various productivity levels. The greatest variability of mortality rates was observed in immature walleye pollock; the maximum mortality rate was typical for the recruitment of highly abundant generations, and the minimum, for the least abundant ones. The lowest mortality was recorded at the age of mass maturation. Mature individuals of generations with various productivity levels showed insignificant differences in overall mortality rates.     

Evidence of predatory control of yellow perch (Perca flavescens) recruitment in Lake Erie, U.S.A.

Predation can be a major factor in recruitment success of yellow perch ( Perca flavescens Mitchitl). Trawl catches of age-0 yellow perch in western Lake Erie declined from 870·3 per trawling h in June to 3·3 per trawling h in late July 1988. Coincident with the decline in relative abundance of age-0 yellow perch we found large numbers of age-0 yellow perch in the stomachs of small walleyes ( Stizostedion vitreum Mitchill). From this evidence we hypothesized that predation by walleyes may have caused the demise of the 1988 year-class of yellow perch. We used a population and bioenergetics modelling approach to estimate the impact of walleye predation on the abundance of age-0 yellow perch. Modelling showed that 6·8 × 10⁹ age-0 yellow perch that had attained 18 mm total length ( t.l. ), were eaten by small (age-2 and younger) walleye from June through July 1988. We estimated that walleye ate 28·4–89·7% of the yellow perch reaching 18 mm t.l. during 1988. The majority of this predation (77% of total) was by the abundant age-2 cohort of walleyes. We concluded that, even in a large system such as Lake Erie, predation can play a major role in structuring year-class strength of yellow perch and, thus, management of percid fisheries should be conducted on a fish-community basis.     

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.     

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.     

Projected shifts in fish species dominance in Wisconsin lakes under climate change

Temperate lakes may contain both coolwater fish species such as walleye (Sander vitreus) and warmwater fish species such as largemouth bass (Micropterus salmoides). Recent declining walleye and increasing largemouth bass populations have raised questions regarding the future trajectories and management actions for these species. We developed a thermodynamic model of water temperatures driven by downscaled climate data and lake‐specific characteristics to estimate daily water temperature profiles for 2148 lakes in Wisconsin, US, under contemporary (1989–2014) and future (2040–2064 and 2065–2089) conditions. We correlated contemporary walleye recruitment and largemouth bass relative abundance to modeled water temperature, lake morphometry, and lake productivity, and projected lake‐specific changes in each species under future climate conditions. Walleye recruitment success was negatively related and largemouth bass abundance was positively related to water temperature degree days. Both species exhibited a threshold response at the same degree day value, albeit in opposite directions. Degree days were predicted to increase in the future, although the magnitude of increase varied among lakes, time periods, and global circulation models (GCMs). Under future conditions, we predicted a loss of walleye recruitment in 33–75% of lakes where recruitment is currently supported and a 27–60% increase in the number of lakes suitable for high largemouth bass abundance. The percentage of lakes capable of supporting abundant largemouth bass but failed walleye recruitment was predicted to increase from 58% in contemporary conditions to 86% by mid‐century and to 91% of lakes by late century, based on median projections across GCMs. Conversely, the percentage of lakes with successful walleye recruitment and low largemouth bass abundance was predicted to decline from 9% of lakes in contemporary conditions to only 1% of lakes in both future periods. Importantly, we identify up to 85 resilient lakes predicted to continue to support natural walleye recruitment. Management resources could target preserving these resilient walleye populations.     

Empirical orthogonal function regression: Linking population biology to spatial varying environmental conditions using climate projections

Ecologists and oceanographers inform population and ecosystem management by identifying the physical drivers of ecological dynamics. However, different research communities use different analytical tools where, for example, physical oceanographers often apply rank‐reduction techniques (a.k.a. empirical orthogonal functions [EOF]) to identify indicators that represent dominant modes of physical variability, whereas population ecologists use dynamical models that incorporate physical indicators as covariates. Simultaneously modeling physical and biological processes would have several benefits, including improved communication across sub‐fields; more efficient use of limited data; and the ability to compare importance of physical and biological drivers for population dynamics. Here, we develop a new statistical technique, EOF regression, which jointly models population‐scale dynamics and spatially distributed physical dynamics. EOF regression is fitted using maximum‐likelihood techniques and applies a generalized EOF analysis to environmental measurements, estimates one or more time series representing modes of environmental variability, and simultaneously estimates the association of this time series with biological measurements. By doing so, it identifies a spatial map of environmental conditions that are best correlated with annual variability in the biological process. We demonstrate this method using a linear (Ricker) model for early‐life survival (“recruitment”) of three groundfish species in the eastern Bering Sea from 1982 to 2016, combined with measurements and end‐of‐century projections for bottom and sea surface temperature. Results suggest that (a) we can forecast biological dynamics while applying delta‐correction and statistical downscaling to calibrate measurements and projected physical variables, (b) physical drivers are statistically significant for Pacific cod and walleye pollock recruitment, (c) separately analyzing physical and biological variables fails to identify the significant association for walleye pollock, and (d) cod and pollock will likely have reduced recruitment given forecasted temperatures over future decades.     

Springtime microprotozoan abundance and biomass in the southeastern Bering Sea and Shelik of Strait, Alaska

We surveyed springtime biomass and abundance of the >20 µmmicroprotozoa in surface waters of the SE Bering Sea and Shelikof Strait, Alaska. This study was part of the Fisheries OceanographyCoordinated Investigations (FOCI) program examining processeswhich affect the recruitment variability of walleye pollock(Theragra chalcogramma). Microprotozoa are a potential preyresource for larval pollock which has not been previously examined.In both areas, the >20 µm microprotozoa were predominantlydinoflagellates and ciliates. At the time of sampling (May 1990in Shelikof Strait and April 1992 in the SE Bering Sea), thespring diatom bloom was under way in Shelik of Strait, but notin the SE Bering Sea. Heterotrophic dinoflagellates dominatedthe microprotozoan assemblage in Shelik of Strait, but not inthe SE Bering Sea. In the SE Bering Sea. total microprotozoanabundances ranged from 300 to 6233 organisms 1^–1 and biomassfrom 0.58 to 9.73 µg C 1^–1. In Shelik of Strait,abundance and biomass were higher, ranging from 850 to 14 960organisms 1^–1 and from 1.29 to 70.73 µg C 1^–1,respectively. These biomass levels are comparable to those reportedfrom other coastal and oceanic regions. Microprotozoan biomasslevels were sufficient to support the estimated metabolic needsof first-feeding larval walleye pollock. It remains to be shownwhether larval pollock use this resource.     

Mitochondrial DNA Variation in Northwestern Bering Sea Walleye Pollock, Theragra chalcogramma (Pallas)

Genetic variability of the highly valuable gadid species, walleye pollock, Theragra chalcogramma (Pallas 1811), from five spatially separated northwestern Bering Sea areas (Ozernoi Bay, Olutorskyi Bay, Koryak shelf, Navarin region, and Anadyr Gulf) was investigated. Haplotype diversity within the samples ranged from 0.8788±0.0393 to 0.9436±0.0162. Nucleotide diversity within the samples ranged from 0.0108 to 0.0127. Nucleotide diversity among the regional collections ranged from 0% to 0.18%. Walleye pollock from the Anadyr Gulf appeared genetically separate from the other four samples in a clustering of genetic distances. Total heterogeneity among all five samples was significant, while there was no heterogeneity among the four samples excluding that from the Anadyr Gulf. Pair-wise comparisons using tests for heterogeneity and F_st supported the dendrogram of genetic distances in that only the collection from the Anadyr Gulf was significantly different from the others. The observed pattern of genetic differentiation among walleye pollock from the northwestern Bering Sea presumably emerged as a result of a population of the species subdividing in the northernmost part of its geographic range, though further analysis is needed to verify this supposition.     

Effects of climate and demography on reproductive phenology of a harvested marine fish population

Shifts in phenology are a well‐documented ecological response to changes in climate, which may or may not be adaptive for a species depending on the climate sensitivity of other ecosystem processes. Furthermore, phenology may be affected by factors in addition to climate, which may accentuate or dampen climate‐driven phenological responses. In this study, we investigate how climate and population demographic structure jointly affect spawning phenology of a fish species of major commercial importance: walleye pollock (Gadus chalcogrammus). We use 32 years of data from ichthyoplankton surveys to reconstruct timing of pollock reproduction in the Gulf of Alaska and find that the mean date of spawning has varied by over 3 weeks throughout the last >3 decades. Climate clearly drives variation in spawn timing, with warmer temperatures leading to an earlier and more protracted spawning period, consistent with expectations of advanced spring phenology under warming. However, the effects of temperature were nonlinear, such that additional warming above a threshold value had no additional effect on phenology. Population demographics were equally as important as temperature: An older and more age‐diverse spawning stock tended to spawn earlier and over a longer duration than a younger stock. Our models suggest that demographic shifts associated with sustainable harvest rates could shift the mean spawning date 7 days later and shorten the spawning season by 9 days relative to an unfished population, independent of thermal conditions. Projections under climate change suggest that spawn timing will become more stable for walleye pollock in the future, but it is unknown what the consequences of this stabilization will be for the synchrony of first‐feeding larvae with production of zooplankton prey in spring. With ongoing warming in the world’s oceans, knowledge of the mechanisms underlying reproductive phenology can improve our ability to monitor and manage species under changing climate conditions.     

The influence of light on egg buoyancy and hatching rate of the walleye pollock, Theragra chalcogramma

Changes in buoyancy in fertilized bathypelagic eggs of the walleye pollock, Theragra chalcogramma , collected from Shelikof Strait in the Gulf of Alaska were measured under controlled laboratory conditions in density gradient columns from 90 h post–fertilization through hatching. Eggs were incubated at 6° C and exposed to either diel light or constant dark. Eggs held under diel light conditions became more dense than eggs under constant dark beginning <10 h after exposure to light and remained so until 12 h before hatching. Eggs held under constant dark then became more dense than those under diel light. Hatching of eggs under both conditions began at the same time but eggs under diel light showed a delayed hatching rate. Light–induced changes in egg density indicate the ability of walleye pollock eggs to respond to external stimuli and thereby alter their position in the water column in an ecologically meaningful way.     

Temperature and prey quality effects on growth of juvenile walleye pollock Theragra chalcogramma (Pallas): a spatially explicit bioenergetics approach

A bioenergetics model for juvenile age‐0 year walleye pollock Theragra chalcogramma was applied to a spatially distinct grid of samples in the western Gulf of Alaska to investigate the influence of temperature and prey quality on size‐specific growth. Daily growth estimates for 50, 70 and 90 mm standard length (L_S) walleye pollock during September 2000 were generated using the bioenergetics model with a fixed ration size. Similarities in independent estimates of prey consumption generated from the bioenergetics model and a gastric evacuation model corroborated the performance of the bioenergetics model, concordance correlation (r_c) = 0·945, lower 95% CL (transformed) (L₁) = 0·834, upper 95% CL (transformed) (L₂) = 0·982, P < 0·001. A mean squared error analysis (M_SE) was also used to partition the sources of error between both model estimates of consumption into a mean component (M_C), slope component (S_C), and random component (R_C). Differences between estimates of daily consumption were largely due to differences in the means of estimates (M_C= 0·45) and random sources (R_C= 0·49) of error, and not differences in slopes (S_C= 0·06). Similarly, daily growth estimates of 0·031–0·167 g day^?1 generated from the bioenergetics model was within the range of growth estimates of 0·026–0·190 g day^?1 obtained from otolith analysis of juvenile walleye pollock. Temperature and prey quality alone accounted for 66% of the observed variation between bioenergetics and otolith growth estimates across all sizes of juvenile walleye pollock. These results suggest that the bioenergetics model for juvenile walleye pollock is a useful tool for evaluating the influence of spatially variable habitat conditions on the growth potential of juvenile walleye pollock.     

Variability of the Characteristics of the First Scale Annulus and Length of Year-Old Fish in the Bering Sea Walleye Pollock Theragra chalcogramma

Geographic and interannual variability of the number of annuli and the radius of the first scale annulus, as well as the retrospective length of year-old walleye pollock from the western, northern, and eastern parts of the Bering Sea, was examined using data for the years 1995, 1996, 1998, and 1999. The characteristics of the first scale annulus were varying. By these parameters, walleye pollock from the western Bering Sea significantly differs statistically from walleye pollock of the eastern Bering Sea. With respect to number of annuli, the radius of the first ring, and the retrospective length of year-old specimens, the walleye pollock from the Navarinskii region occupies an intermediate position between fish from the western and eastern parts of the Bering Sea. Interannual variability of the three parameters was found for walleye pollock from the Navarinskii region.     

The first annulus of otoliths: a tool for studying intra-annual growth of walleye pollock (Theragra chalcogramma)

We quantified the growing season of yearling walleye pollock (Theragra chalcogramma) and related it to annual cycles of water temperature and day length. The study was restricted to members of the 2000 year class and thereby controlled for inter-annual variability. Juveniles were sampled from the year class during 10 cruises in the western Gulf of Alaska (GOA). Fifty percent of juveniles exhibited an annulus on 16 March 2001 (± 11 days 95% confidence interval). No regional difference was detected in the timing of annulus formation or in post-annulus growth trajectories. A model, derived from growth trajectories, estimated that the growing season lasted 204 days (22 March to 13 October 2001) and that growth rate peaked at 0.59 mm day⁻¹ on 2 July 2001. Growth rate increased with day length and water temperature during spring and decreased in late summer possibly due to thermal stress. Secondarily, we explored the utility of otolith size at the first annulus as a natural tag to identify nursery area, but this potential was curtailed by overlap in length among regions. Our results indicate that the first annulus can be used to advance our understanding of climate forcing on marine fish growth by providing fine temporal resolution of the growing season.     

Inter-island variation in the diet of female northern fur seals (Callorhinus ursinus) in the Bering Sea

The diet of adult female northern fur seals ( Callorhinus ursinus ) is examined through the analysis of faecal material collected during the summer breeding season at three breeding locations in the Bering Sea: St. Paul Island (1988, 1990) and St. George Island (1988, 1990) of the Pribilof Islands Group (USA), and Medny Island (1990) of the Commander Islands Group (Russia). Prey consumption varies annually and accordingly with the physical and biological environment surrounding each island. Juvenile walleye pollock ( Theragra chalcogramma ) is the most common prey of northern fur seals from St. Paul Island; the island is surrounded by a broad neritic environment with widely separated frontal zones and is the greatest distance from the continental shelf-edge. Gonatid squid ( Gonatopsis borealis/Berryteuthis magister and Gonatus madokail Gonatus middendorffi ) were the most common prey of northern fur seals from Medny Island; the island is surrounded by a compressed neritic environment and is adjacent to the continental shelf-edge and the oceanic marine environment. A combination of walleye pollock and gonatid squid is consumed by northern fur seals from St. George Island; the island has a surrounding oceanographic environment intermediate between the other two islands.
Variability in predation on walleye pollock is consistent with fishery information concerning the relative abundance and availability of walleye pollock around St. George and St. Paul Islands. The abundance and availability of these prey resources during the summer breeding season are key factors which influence the health and growth of the northern fur seal populations in the Bering Sea.