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The determinants of the structure, functioning and resilience of pelagic ecosystems across most of the polar regions are not well known. Improved understanding is essential for assessing the value of biodiversity and predicting the effects of change (including in biodiversity) on these ecosystems and the services they maintain. Here we focus on the trophic interactions that underpin ecosystem structure, developing comparative analyses of how polar pelagic food webs vary in relation to the environment. We highlight that there is not a singular, generic Arctic or Antarctic pelagic food web, and, although there are characteristic pathways of energy flow dominated by a small number of species, alternative routes are important for maintaining energy transfer and resilience. These more complex routes cannot, however, provide the same rate of energy flow to highest trophic-level species. Food-web structure may be similar in different regions, but the individual species that dominate mid-trophic levels vary across polar regions. The characteristics (traits) of these species are also different and these differences influence a range of food-web processes. Low functional redundancy at key trophic levels makes these ecosystems particularly sensitive to change. To develop models for projecting responses of polar ecosystems to future environmental change, we propose a conceptual framework that links the life histories of pelagic species and the structure of polar food webs.

Traditionally, the advice provided to fishery managers has focused on the trade-offs between short- and long-term yields, and between future resource size and expected future catches. The harvest control rules that are used to provide management advice consequently relate catches to stock biomass levels expressed relative to reference biomass levels. There are, however, additional trade-offs. Ecosystem-based fisheries management (EBFM) aims to consider fish and fisheries in their ecological context, taking into account physical, biological, economic, and social factors. However, making EBFM operational remains challenging. It is generally recognized that end-to-end modeling should be a key part of implementing EBFM, along with harvest control rules that use information in addition to estimates of stock biomass to provide recommendations for management actions. Here we outline the process for selecting among alternative management strategies in an ecosystem context and summarize a Field-integrated End-To-End modeling program, or FETE, intended to implement this process as part of the Bering Sea Project. A key aspect of this project was that, from the start, the FETE included a management strategy evaluation component to compare management strategies. Effective use of end-to-end modeling requires that the models developed for a system are indeed integrated across climate drivers, lower trophic levels, fish population dynamics, and fisheries and their management. We summarize the steps taken by the program managers to promote integration of modeling efforts by multiple investigators and highlight the lessons learned during the project that can be used to guide future use and design of end-to-end models.

This paper compares and contrasts the ecosystems of the Barents and Chukchi Seas. Despite their similarity in a number of features, the Barents Sea supports a vast biomass of commercially important fish, but the Chukchi does not. Here we examine a number of aspects of these two seas to ascertain how they are similar and how they differ. We then indentify processes and mechanisms that may be responsible for their similarities and differences.Both the Barents and Chukchi Seas are high latitude, seasonally ice covered, Arctic shelf-seas. Both have strongly advective regimes, and receive water from the south. Water entering the Barents comes from the deep, ice-free and "warm" Norwegian Sea, and contains not only heat, but also a rich supply of zooplankton that supports larval fish in spring. In contrast, Bering Sea water entering the Chukchi in spring and early summer is cold. In spring, this Bering Sea water is depleted of large, lipid-rich zooplankton, thus likely resulting in a relatively low availability of zooplankton for fish. Although primary production on average is similar in the two seas, fish biomass density is an order of magnitude greater in the Barents than in the Chukchi Sea. The Barents Sea supports immense fisheries, whereas the Chukchi Sea does not. The density of cetaceans in the Barents Sea is about double that in the Chukchi Sea, as is the density of nesting seabirds, whereas, the density of pinnipeds in the Chukchi is about double that in the Barents Sea. In the Chukchi Sea, export of carbon to the benthos and benthic biomass may be greater. We hypothesize that the difference in fish abundance in the two seas is driven by differences in the heat and plankton advected into them, and the amount of primary production consumed in the upper water column. However, we suggest that the critical difference between the Chukchi and Barents Seas is the pre-cooled water entering the Chukchi Sea from the south. This cold water, and the winter mixing of the Chukchi Sea as it becomes ice covered, result in water temperatures below the physiological limits of the commercially valuable fish that thrive in the southeastern Bering Sea. If climate change warms the Barents Sea, thereby increasing the open water area via reducing ice cover, productivity at most trophic levels is likely to increase. In the Chukchi, warming should also reduce sea ice cover, permitting a longer production season. However, the shallow northern Bering and Chukchi Seas are expected to continue to be ice-covered in winter, so water there will continue to be cold in winter and spring, and is likely to continue to be a barrier to the movement of temperate fish into the Chukchi Sea. Thus, it is unlikely that large populations of boreal fish species will become established in this Arctic marginal sea. © 2012 Elsevier B.V.

Walleye pollock (Theragra chalcogramma) is an important component of the eastern Bering Sea ecosystem and subject to major fisheries. The Oscillating Control Hypothesis (OCH) predicted that recruitment of pollock year classes should be greatest in years with early ice retreat and late blooms in warm water, because more energy would flow into the pelagic (vs. benthic) community. The OCH further predicted that, with pollock population growth, there should be a shift from bottom-up to top-down regulation. New data support the predictions that in those years with early ice retreat, more primary production accrues to the pelagic compartment and that large numbers of age-0 pollock survive to summer. However, in these years, production of large crustacean zooplankton is reduced, depriving age-0 pollock of lipid-rich prey in summer and autumn. Consequently, age-0 pollock energy reserves (depot lipids) are low and predation on them is increased as fish switch to age-0 pollock from zooplankton. The result is weak recruitment of age-1 recruits the following year. A revised OCH indicates bottom-up constraints on pollock recruitment in very warm periods. Prolonged warm periods with decreased ice cover will likely cause diminished pollock recruitment and catches relative to recent values. © 2011 International Council for the Exploration of the Sea.

Ivory gulls (Pagophila eburnea, Phipps, 1774), one of the world's least-known species, have declined throughout their range in recent years. This study describes the patterns of ivory gull use of the North Water polynya, a large polynya that occurs every year near ivory gull breeding sites on Ellesmere Island, Nunavut, Canada. We conducted at-sea surveys from Canadian icebreakers during the summers of 1997-99. In 1998, stomach contents of five ivory gulls were analyzed. We measured stable isotope ratios (δ 13C, δ 15N) of liver, muscle, feather and bone to determine how ivory gull diets vary during the year. We observed a total of 275 individuals, most of which were seen on the western side of the polynya. Flying was the predominant behavior (76% of individuals); other behaviors included sitting on ice, sitting on water, and feeding. Four juveniles were seen in August and September. Birds collected had arctic cod (Boreogadus saida) in their stomachs. Other food items included an otolith from an unidentified species of Liparid fish and two bones from a mole, presumed to be a star-nosed mole (Condylura cristata). Stable isotope results indicated that ivory gull trophic levels (TLs) were high (around TL 4 based on δ 15N) but varied over the season. From the three individuals whose feathers were analyzed, we conclude that differences among individuals in their winter diets or foraging locations are possible. © The Arctic Institute of North America.

Each year, millions of seabirds migrate to the North Water Polynya, northern Baffin Bay, to feed in its productive waters during the 6 months that the polynya is free of ice. We evaluated seasonal shifts in diets of 3 species using the North Water: dovekie Alle alle, thick-billed murre Uria lomvia, and black-legged kittiwake Rissa tridactyla. Diets were assessed through a combination of stable isotope analysis of muscle tissue, fatty acid analysis of subcutaneous fat, and stomach content analysis. Dovekies had the lowest δ15N values and hence lowest trophic level in spring and summer, corresponding to their consumption of herbivorous copepods. In fall, their δ15N values increased as they switched to feeding at a higher trophic level on primary carnivores such as amphipods and fish. Throughout the study period, kittiwakes and murres had stomach contents and stable isotope values similar to dovekies in fall. Fatty acid signatures of black-legged kittiwakes and dovekies feeding in fall were similar, likely due to the reliance of both species on the pteropod Limacina limacina. Our study highlights the seasonal nature of prey availability and seabird diets in the polynya, as well as the utility of simultaneously using conventional stomach content analysis and stable isotope and fatty acid analyses to infer dietary patterns in marine food webs. © Inter-Research 2008.

Our objective was to understand how marine birds respond to oceanographic variability across the Southern Indian Ocean using data collected during an 16-day cruise (4-21 January 2003). We quantified concurrent water mass distributions, ocean productivity patterns, and seabird distributions across a heterogeneous pelagic ecosystem from subtropical to sub-Antarctic waters. We surveyed 5155 km and sighted 15,606 birds from 51 species, and used these data to investigate how seabirds respond to spatial variability in the structure and productivity of the ocean. We addressed two spatial scales: the structure of seabird communities across macro-mega scale (1000 s km) biogeographic domains, and their coarse-scale (10 s km) aggregation at hydrographic and bathymetric gradients. Both seabird density and species composition changed with latitudinal and onshore-offshore gradients in depth, water temperature, and chlorophyll-a concentration. The average seabird density increased across the subtropical convergence (STC) from 2.4 birds km-2 in subtropical waters to 23.8 birds km-2 in sub-Antarctic waters. The composition of the avifauna also differed across biogeographic domains. Prions (Pachyptila spp.) accounted for 57% of all sub-Antarctic birds, wedge-tailed shearwaters (Puffinus pacificus) accounted for 46% of all subtropical birds, and Indian Ocean yellow-nosed albatross (Thallasarche carteri) accounted for 32% of all birds in the STC. While surface feeders were the most abundant foraging guild across the study area, divers were disproportionately more numerous in the sub-Antarctic domain, and plungers were disproportionately more abundant in subtropical waters. Seabird densities were also higher within shallow shelf-slope regions, especially in sub-Antarctic waters, where large numbers of breeding seabirds concentrated. However, we did not find elevated seabird densities along the STC, suggesting that this broad frontal region is not a site of enhanced aggregation. © 2007 Elsevier Ltd. All rights reserved.

Declines of Steller sea lion (Eumetopias jubatus) populations in the Aleutian Islands and Gulf of Alaska could be a consequence of physical oceanographic changes associated with the 1976-77 climate regime shift. Changes in ocean climate are hypothesized to have affected the quantity, quality, and accessibility of prey, which in turn may have affected the rates of birth and death of sea lions. Recent studies of the spatial and temporal variations in the ocean climate system of the North Pacific support this hypothesis. Ocean climate changes appear to have created adaptive opportunities for various species that are preyed upon by Steller sea lions at mid-trophic levels. The east-west asymmetry of the oceanic response to climate forcing after 1976-77 is consistent with both the temporal aspect (populations decreased after the late 1970s) and the spatial aspect of the decline (western, but not eastern, sea lion populations decreased). These broad-scale climate variations appear to be modulated by regionally sensitive biogeographic structures along the Aleutian Islands and Gulf of Alaska, which include a transition point from coastal to open-ocean conditions at Samalga Pass westward along the Aleutian Islands. These transition points delineate distinct clusterings of different combinations of prey species, which are in turn correlated with differential population sizes and trajectories of Steller sea lions. Archaeological records spanning 4000 yr further indicate that sea lion populations have experienced major shifts in abundance in the past. Shifts in ocean climate are the most parsimonious underlying explanation for the broad suite of ecosystem changes that have been observed in the North Pacific Ocean in recent decades. © 2007 Blackwell Publishing Ltd.

We investigated seabird-eddy associations in subtropical waters of the southern Indian Ocean during a summer (January) cruise from Amsterdam Island to Western Australia. To determine habitat associations and to enable predictions about foraging strategies, we related the distributions of satellite altimetry data (sea surface height anomalies) to the 3 most abundant taxa in the study area: terns (sooty Sterna fuscata and bridled S. anaethetus) and shearwaters (wedge-tailed Puffinus pacificus) were significantly more common and numerous within convergence zones. However, a step-wise logistic regression model revealed that these seabirds were more strongly associated with other habitat variables, including the distance to breeding colonies and warm sea surface temperature conditions. Great-winged petrels Pterodroma macroptera were widely scattered and did not show associations with convergences or divergences. These contrasting results highlight the need to consider smaller-scale hydrographic features, such as frontal systems, when guantifying seabird associations with mesoscale eddies. Large-scale patterns of ocean circulation and mesoscale variability also seem to play a critical role in determining the extent of seabird-eddy associations. In contrast with previous studies, which have focused on dynamic frontal regions and energetic current systems, we surveyed a region of weak mesoscale variability north of the Subtropical Convergence zone. © Inter-Research 2006.

The short-tailed shearwater Puffinus tenuirostris is an apex predator in the southeastern Bering Sea ecosystem. During 1997 to 1999, a period of great variability in the Bering Sea, we used a multi-pronged approach to study transfer of carbon and nitrogen to short-tailed shearwaters through analysis of stomach contents of birds collected while foraging, and stable isotope and fatty acid composition of tissues from shearwaters and their prey. Two conclusions result from these 3 analyses of feeding history. First, short-tailed shearwaters demonstrated localized differences in diet with respect to sampling location and season, indicating that shearwaters feed in discrete locations long enough (several weeks) to reflect regional differences in prey availability. Second, elevated δ15N levels (∼1 to 2‰) in the liver of shearwaters in the fall of 1997 and 1998 were likely a response to nutritional stress and overturn of nutrients in the tissues of birds or elevated isotope levels in tissues of prey, rather than an increase in the trophic level of the diet. Over the 3 yr period, shearwater diets switched from adult euphausiids Thysanoessa raschii and T. inermis (usually females with spermatophores) taken in spring to an increase in the amount of fish (Pacific sandlance Ammodytes hexapterus), juvenile Gadidae fishes and larval walleye pollock Theragra chalcograma particularly in fall (1997 excepted). For each location and year, there was a consistent trend in isotopic values, with an increase of 3 to 5‰ for δ15N and 1 to 3‰ for δ13C between shearwaters and their primary prey. The resolution of our sampling techniques indicates little movement of shearwater flocks among sample locations for periods of up to several weeks. © Inter-Research 2006.

We examined the hypothesis that short-tailed shearwaters Puffin us tenuirostris aggregate to forage at the inner front of the SE Bering Sea because of enhanced production there. We tested this hypothesis by comparing primary production, the distribution of euphausiids and the distribution of shearwaters relative to the front during late spring and late summer/early fall of 1997, 1998 and 1999. We found enhanced primary production at the front and offshore of the front during summer but not during spring. Primary production varied between seasons and years. Major differences were related to anomalous conditions in 1997 and 1998. The density of euphausiids was higher at the front and offshore of the front during summer, but there were no differences among regions during spring. Foraging shearwaters aggregated in high densities at the front during summer, but foraged close to shore during spring. At the front, shearwaters foraged on euphausiids Thysanoessa raschii and T. inermis as expected, and on copepods that accumulated in the area. The proportion of zooplankton consumed at the front decreased from summer 1997 to summer 1999, while consumption of sandlance Ammodytes hexapterus at this feature increased. Our results show that, during summer, the inner front supports aggregations of euphausiids and their seabird predators. The means by which the frontal system supports enhanced production and the subseguent trophic transfers is dependent on the availability of nutrients at depth in the frontal region and the aggregation of small zooplankton organisms in this feature. © Inter-Research 2005.

Strong tidal currents crossing over the abrupt topography of the Aleutian Passes result in regions with high horizontal property gradients. These frontal regions vary with the tidal cycle and form the boundary between vertically mixed and stratified regions. Concentrations of seabirds were associated with convergence zones in the mixed water (MW) and with the front between North Pacific (NP) water and MW. Species that were foraging by picking at prey from the surface were associated with surface convergences that appeared to be associated with Langmuir circulation cells or tidal features (all fulmar aggregations) in the central passes (Samalga, Seguam). In contrast, sub-surface foraging puffins and small alcids were mostly observed in areas of turbulent, well-mixed water near the shallow regions of the passes. Short-tailed shearwater flocks that were plunge-diving for prey were associated with the front between the NP water and MW in the passes. On our transects, we observed no significant aggregations of seabirds associated with Bering Sea water or NP water away from the frontal zones. The interaction of strong currents with bathymetric features results in zones of vertical advection, mixing, and surface convergences that make island passes attractive foraging regions for seabirds. Deep passes lacking these features, such as many of the passes in the western Aleutian Archipelago, are not as likely to facilitate trophic transfer to top predators as shallow passes, such as those found in the eastern Aleutian Islands. © 2005 Blackwell Publishing Ltd.

To examine the marine habitat of the endangered western stock of the Steller's sea lion (Eumetopias jubatus), two interdisciplinary research cruises (June 2001 and May to June 2002) measured water properties in the eastern and central Aleutian Passes. Unimak, Akutan, Amukta, and Seguam Passes were sampled in both years, and three additional passes (Umnak, Samalga, and Tanaga) were sampled in 2002. In the North Pacific (and to a lesser extent in the Bering Sea), a strong front in water properties was observed near Samalga Pass in June of both years, with significantly warmer, fresher, and more nitrate-poor water east of Samalga Pass than west of the pass. These water properties reflect differences in source waters (Alaska Coastal Current versus Alaskan Stream), mixing depth, and Bering Sea influence. Strong cross-Aleutian gradients were also observed with warmer, fresher water on the North Pacific side of the archipelago. The nutrient content of the waters flowing through the passes, combined with the effects of mixing within the passes, influences the transport of nutrients into the Bering Sea. As water moves away from the strong mixing of the passes and becomes more stratified, phytoplankton can take advantage of the enhanced nutrient concentrations. Thus, the northern side of the Aleutian Islands (especially in the lee of the islands) appears to be more productive. Combined with evidence of coincident changes in many ecosystem parameters near Samalga Pass, it is hypothesized that Samalga Pass forms a physical and biogeographic boundary between the eastern and central Aleutian marine ecosystems. © 2005 Blackwell Publishing Ltd.

We examined the hypothesis that seabird distribution, abundance and diets differ among the eastern and central Aleutian Islands in response to distinct marine environments and energy pathways in each region. Research cruises were conducted in June 2001 and May-June 2002. We determined the distribution, abundance, diet and prey consumption of seabirds, and related these to zooplankton abundance and water masses that possess different physical properties. We found that distribution, abundance and diets of seabirds could be partitioned into two regions that correspond to marine environments determined by the extent of the Alaska Coastal Current along the eastern and central Aleutian Islands. Short-tailed shearwaters (Puffinus tenuirostris) were the most abundant seabird in the coastal waters of the eastern Aleutian Islands, and northern fulmars (Fulmarus glacialis) were the most abundant seabird in the oceanic waters of the central Aleutian Islands. Seabird communities in the central and eastern Aleutian Islands were likely associated with different food webs. In the central Aleutian Islands, short-tailed shearwaters and northern fulmars consumed shelf-break species of euphausiids (Thyssanoesa longipes) and oceanic copepods (Neocalanus cristatus), respectively; in the eastern Aleutian Islands, both short-tailed shearwaters and northern fulmars consumed shelf species of euphausiids (T. inermis). Carbon transport to seabirds was highest in Unimak and Akutan Passes where shearwaters removed large quantities of shelf euphausiids, followed by Samalga and Seguam Passes where northern fulmars removed large amounts of oceanic copepods. © 2005 Blackwell Publishing Ltd.

The distribution of nutrients (nitrate, phosphate, and silicic acid), chlorophyll and primary productivity were examined in the central and eastern Aleutian Archipelago. The data were collected from moorings (temperature, salinity, nitrate, and currents) and two hydrographic research cruises (June 2001 and May-June 2002). During the hydrographic cruises salinity, temperature, nutrients, chlorophyll and primary production were measured in and around the eastern and central Aleutian Passes. The net nutrient transport through the passes was northward, and it was relatively low in the eastern passes compared to the central passes. In the shallow eastern passes the source water from the Pacific was the nutrient-poor Alaska Coastal Current, while in the deeper central passes it was the nutrient-rich Alaskan Stream. Within the passes, vigorous tidal mixing resulted in enriched surface concentrations, especially in the central passes. Chlorophyll sections and satellite composites from summer indicate high chlorophyll east of Samalga Pass and very low chlorophyll between Samalga and Seguam Passes. Production was relatively low across the study area, except at the northern end of Seguam Pass and in the vicinity of the shelf edge in the southeastern Bering Sea. Production and chlorophyll concentrations were lowest in the deep passes. These results suggest that, within the passes, deep mixing inhibits new production, but substantial blooms may occur downstream of the passes subsequent to stratification. © 2005 Blackwell Publishing Ltd.

The eastern Bering Sea and the Barents Sea share a number of common biophysical characteristics. For example, both are seasonally ice-covered, high-latitude, shelf seas, dependent on advection for heat and for replenishment of nutrients on their shelves, and with ecosystems dominated by a single species of gadoid fish. At the same time, they differ in important respects. In the Barents Sea, advection of Atlantic Water is important for zooplankton vital to the Barents Sea productivity. Advection of zooplankton is not as important for the ecosystems of the southeastern Bering Sea, where high levels of diatom production can support production of small, neritic zooplankton. In the Barents Sea, cod are the dominant gadoid, and juvenile and older fish depend on capelin and other forage fish to repackage the energy available in copepods. In contrast, the dominant fish in the eastern Bering Sea is the walleye pollock, juveniles and adults of which consume zooplankton directly. The southeastern Bering Sea supports considerably larger fish stocks than the Barents. In part, this may reflect the greater depth of much of the Barents Sea compared with the shallow shelf of the southeastern Bering. However, walleye pollock is estimated to occupy a trophic level of 3.3 as compared to 4.3 for Barents Sea cod. This difference alone could have a major impact on the abilities of these seas to support a large biomass of gadoids. In both seas, climate-forced variability in advection and sea-ice cover can potentially have major effects on the productivity of these Subarctic seas. In the Bering Sea, the size and location of pools of cold bottom waters on the shelf may influence the likelihood of predation of juvenile pollock. © 2005 International Council for the Exploration of the Sea. Published by Elsevier Ltd. All rights reserved.

We investigated prey consumption by marine birds and their contribution to cross-shelf fluxes in the northern Gulf of Alaska. We utilized data from the North Pacific Pelagic Seabird Database for modeling energy demand and prey consumption. We found that prey consumption by marine birds was much greater over the continental shelf than it was over the basin. Over the shelf, subsurface-foraging marine birds dominated food consumption, whereas over the basin, surface-foraging birds took the most prey biomass. Daily consumption by marine birds during the non-breeding season ("winter") from September through April was greater than daily consumption during the breeding season, between May and August. Over the shelf, shearwaters, murres and, in winter, sea ducks, were the most important consumers. Over the basin, northern fulmars, gulls and kittiwakes predominated in winter and storm-petrels dominated in May to August. Our results suggest that marine birds contribute little to cross-shelf fluxes of energy or matter, but they do remove energy from the marine system through consumption, respiration and migration. © 2005 Elsevier Ltd. All rights reserved.

This compilation of new information and summaries of earlier work emphasizes variability within marine waters of the Aleutian Archipelago. From the Alaska Peninsula to Near Strait, net flow through the passes is northward, with four passes (Amukta, Amchitka, Buldir, and Near Strait) contributing most of the flow. East of Samalga Pass (169°W), waters derived from the Alaska Coastal Current predominate, whereas west of Samalga Pass, waters of the Alaskan Stream predominate. The pattern of storm tracks creates a climatological (interannual and long term) transition zone in weather features (e.g. surface air temperature) near 170°W. The marine ecosystem of the Aleutian Archipelago also has a strong discontinuity at Samalga Pass, where cold-water corals, zooplankton, fish, marine mammals and foraging seabirds show a step change in species composition. Diets of ground fish, Steller sea lions (Eumetopias jubatus) and some seabirds also change there. Lower growth rates of some fish species and stable isotope data indicate that productivity declines westward along the archipelago. The available data demonstrate considerable ecosystem variability over time scales of decades to millennia. Abrupt changes in composition of fish communities at several of the major passes suggest that Samalga Pass may mark only one of several ecological divisions of Aleutian waters. This spatial and temporal heterogeneity provides an important context within which to view recent declines in populations of Steller sea lions and other species, and has important implications for the management of regional marine resources. We conclude that the marine waters of the Aleutian Archipelago are divided into at least two different ecological regions, with potential for a concomitant separation of some fishery resources. © 2005 Blackwell Publishing Ltd.

We hypothesized that change in the annual population size of guano-producing seabirds (cormorant, Phalacrocorax bougainvillii; booby, Sula variegata; pelican, Pelecanus thagus) is a response to changes in primary and secondary production of the Peruvian upwelling system. We tested this hypothesis by modeling nitrate input through upwelling to the upper layers of the ocean off Peru between 6° and 14°S using data on wind stress and sea surface temperature. The model predicted the amount of carbon fixed by primary production each year from 1925 to 2000, which was then apportioned to the Peruvian anchovy (Engraulis ringens) biomass and ultimately to the seabird population and the anchovy fishery, the largest single-species fishery on Earth. The model predicted a marked increase in primary production as a consequence of increasing wind stress. It overestimated the anchovy biomass after the collapse of the fishery in 1972, but closely predicted the growth of seabird populations from 1925 to the mid-1960s, and their decline thereafter, explaining about 94% of the variation in seabird numbers from 1925 to 2000. The model indicates the seabirds consumed 14.4% of the available anchovies and, thus, that seabirds consumed 2.3% of the new production, before the development of the anchovy fishery, and only 2.2% of the available anchovies and 0.3% of the new production after the development of the fishery. The model results clarify the roles that environmental and anthropogenic factors may have had in regulating the guano-producing seabird populations. It indicates that the growth of seabird populations from 1925 to 1955 was likely a response to increased productivity of the Peruvian upwelling system and that the subsequent drastic decline in seabird abundance was likely due to competition for food with the fishery, which caught ∼85% of the anchovies, which otherwise would have been available for the seabirds. This model also shows that an increase in oceanic primary production promotes reproductive success and population growth in higher trophic level organisms.

The inner front of the southeastern Bering Sea shows marked spatial variability in frontal characteristics created by regional differences in forcing mechanisms. Differences in forcing mechanisms (sea ice advance/retreat and storm strength and timing) and early spring water properties result in strong interannual variability in biological, chemical, and physical features near the front. We have developed a simple model based on surface heat flux and water-column mixing to explain the existence of cold belts (Cont. Shelf Res. 19(14) (1999) 1833) associated with such fronts. Hydrography, fluorescence and nutrient observations show that pumping of nutrients into the euphotic zone occurs, and this can prolong primary production at the inner front. The effectiveness of this process depends on two factors: the existence of a reservoir of nutrients in the lower layer on the middle shelf and the occurrence of sufficient wind and tidal energy to mix the water column. © 2002 Elsevier Science Ltd. All rights reserved.

We modeled the energy demand of seabirds in the North Water, focusing on the planktivorous dovekie (Alle alle), the dominant species in the polynya. For the dovekie we provided an estimate of carbon flux that included aspects of spatial and temporal variability. We estimated the density, diet, and carbon consumption of dovekies throughout the polynya, from the time of their arrival in mid-May until they began to migrate south in September. Our model showed that this species is responsible for 92-96% of the energy demand and therefore carbon flux to seabirds. Dovekies consumed 73.7-147 × 103 mt C yr-1 in the North Water. Average flux rates to dovekies in the polynya were 0.74 g C m-2 yr-1, with a maximum estimated rate of 24 mg C m-2 d-1 in the eastern portion of the study area in May. However, when averaged over the entire polynya and period of occupancy, the proportion of pelagic primary production that went to dovekies was negligible (0.3-0.6%). Our observations of dovekie distribution, indicate that the major flux of carbon to seabirds occurred close inshore along the Greenland coast. There in May, carbon flux to dovekies was estimated to be 5-14% of the potential particulate export of phytoplankton. Our estimates of the spatial distribution of carbon flux to birds suggest the extraordinary importance of production along the west coast of Greenland. © 2002 Elsevier Science Ltd. All rights reserved.

There has been a dramatic increase in jellyfish biomass over the eastern Bering Sea shelf since the early 1990s, which was previously hypothesized to have been triggered by changing climate and ocean conditions. We examine the hypothesis that the presence of these large carnivores has affected fisheries resources, either through direct predation on larval stages, or through competition for zooplankton prey. In this paper, we explore the impact of this jellyfish increase on zooplankton and fish communities based on field data on the composition of the jellyfish community, and the abundance, size, stable isotopic signatures, and feeding habits of the principal scyphomedusae in the region. These data, together with those on zooplankton biomass, are used to estimate the ecosystem impacts of this increase. The center of jellyfish biomass has shifted from the SE Middle Shelf Domain in the early 1980s to the NW in the late 1990s. In recent years, the species composition of large medusae caught in trawls was dominated (>80% by number and >95% by weight) by the scyphozoan Chrysaora melanaster. Dense aggregations of this species occupied the water column in daytime between 10 and 40 m. Their food habits consisted mainly of pelagic crustaceans (euphausiids, copepods, amphipods), although other jellyfish and juvenile pollock were also consumed. Based on stable isotope ratios, the trophic level of this scyphozoan is equivalent to, or higher than, that of Age 0 pollock. Preliminary estimates showed that medusae have a moderate grazing impact on zooplankton in the area around the Pribilof Islands; C. melanaster was estimated on average to consume seasonally about one-third of the standing stock and 4.7% of the annual production of zooplankton in this region. Daily consumption of Age 0 pollock was estimated to be 2.8% of the standing stock around the Pribilof Islands during 1999. A hypothesis for the increase in jellyfishes observed in the eastern Bering Sea, based on release from competition from planktivorous forage fishes, is proposed.

We propose a new hypothesis, the Oscillating Control Hypothesis (OCH), which predicts that pelagic ecosystem function in the southeastern Bering Sea will alternate between primarily bottom-up control in cold regimes and primarily top-down control in warm regimes. The timing of spring primary production is determined predominately by the timing of ice retreat. Late ice retreat (late March or later) leads to an early, ice-associated bloom in cold water (e.g., 1995, 1997, 1999), whereas no ice, or early ice retreat before mid-March, leads to an open-water bloom in May or June in warm water (e.g., 1996, 1998, 2000). Zooplankton populations are not closely coupled to the spring bloom, but are sensitive to water temperature. In years when the spring bloom occurs in cold water, low temperatures limit the production of zooplankton, the survival of larval/juvenile fish, and their recruitment into the populations of species of large piscivorous fish, such as walleye pollock (Theragra chalcogramma), Pacific cod (Gadus macrocephalus) and arrowtooth flounder (Atheresthes stomias). When continued over decadal scales, this will lead to bottom-up limitation and a decreased biomass of piscivorous fish. Alternatively, in periods when the bloom occurs in warm water, zooplankton populations should grow rapidly, providing plentiful prey for larval and juvenile fish. Abundant zooplankton will support strong recruitment of fish and will lead to abundant predatory fish that control forage fish, including, in the case of pollock, their own juveniles. Piscivorous marine birds and pinnipeds may achieve higher production of young and survival in cold regimes, when there is less competition from large piscivorous fish for coldwater forage fish such as capelin (Mallotus villosus). Piscivorous seabirds and pinnipeds also may be expected to have high productivity in periods of transition from cold regimes to warm regimes, when young of large predatory species of fish are numerous enough to provide forage. The OCH predicts that the ability of large predatory fish populations to sustain fishing pressure will vary between warm and cold regimes. The OCH points to the importance of the timing of ice retreat and water temperatures during the spring bloom for the productivity of zooplankton, and the degree and direction of coupling between zooplankton and forage fish. Forage fish (e.g., juvenile pollock, capelin, Pacific herring [Clupea pallasii]) are key prey for adult pollock and other apex predators. In the southeastern Bering Sea, important changes in the biota since the mid-1970s include a marked increase in the biomass of large piscivorous fish and a concurrent decline in the biomass of forage fish, including age-1 walleye pollock, particularly over the southern portion of the shelf. Populations of northern fur seals (Callorhinus ursinus) and seabirds such as kittiwakes (Rissa spp.) at the Pribilof Islands have declined, most probably in response to a diminished prey base. The available evidence suggests that these changes are unlikely the result of a decrease in total annual new primary production, though the possibility of reduced post-bloom production during summer remains. An ecosystem approach to management of the Bering Sea and its fisheries is of great importance if all of the ecosystem components valued by society are to thrive. Cognizance of how climate regimes may alter relationships within this ecosystem will facilitate reaching that goal.

We examine how coupling between physical and biological processes influences the production and transfer of energy to upper trophic-level species in the southeastern Bering Sea. We review time series that illustrate changes in the marine climate of the southeastern Bering Sea since the mid-1970s, particularly variability in the persistence of sea ice and the timing of its retreat. Time series (1995 - 2001) from a biophysical mooring in the middle domain of the southeastern shelf support the hypothesis that retreat of the winter sea ice before mid-March (or the failure of ice to be advected into a region) results in an open water bloom in May or June in relatively warm water (≥3°C). Conversely, when ice retreat is delayed until mid-March or later, an ice-associated bloom occurs in cold (≤0°C) water in early spring. These variations are important because the growth and production of zooplankton and the growth and survival of larval and juvenile fish are sensitive to water temperature. The Oscillating Control Hypothesis (OCH) recently proposed by Hunt et al. (2002), predicts that control of the abundance of forage fish, and in the case of walleye pollock (Theragra chalcogramma), recruitment of large piscivorous fish, will switch from bottom-up limitation in extended periods with late ice retreat to top-down in warmer periods when ice retreat occurs before mid-March. In support of this hypothesis, we review recent data from the southeastern Bering Sea that show 2- to 13-fold changes in copepod abundance with changes in spring water temperatures of 3 to 5°C. We also provide indirect evidence that the abundance of adult pollock on the eastern Bering Sea shelf negatively affects the abundance forage fishes (including juvenile pollock) available to top predators. Although there is evidence that pollock year-class strength is positively related to temperature, we lack the time series of pollock populations in extended periods (8 - 10 years) of cold-water blooms necessary to test the OCH. © 2002 Elsevier Science Ltd. All rights reserved.

In the late 1990s, the southeastern Bering Sea exhibited a number of anomalous conditions, including a major die-off of short-tailed shearwaters (Puffinus tenuirostris), a trans-equatorial migrant that constitutes a major portion of the marine bird biomass in the southeastern Bering Sea. As part of a larger study of the ecological role of the inner or structural front over the southeastern Bering Sea shelf, in 1997-1999, we collected short-tailed shearwaters to determine diet composition. In spring 1997, we found that short-tailed shearwaters were consuming predominately the euphausiid Thysanoessa raschii, a diet expected on the basis of past studies. However, in subsequent years, short-tailed shearwater diets in spring contained increasingly larger proportions of fish, in particular, sandlance (Ammodytes hexapterus), as well as other species of euphausiids (T. inermis in 1999). In summer and fall collections, short-tailed shearwater diets were more varied than in spring, and included both fish (age-0 gadids, 21-35% by weight) and a wider variety of euphausiid species (T. inermis and T. spinifera). In summer and fall, crab zoea (August 1998) and copepods (August 1999) were eaten by shearwaters collected while feeding within the inner front. Diets in 1997-1999 were broader than those found in previous studies of short-tailed shearwaters over the inner shelf and Bristol Bay, which had documented diets composed almost solely of T. raschii. Our data are consistent with the hypothesis that euphausiids were less available to short-tailed shearwaters foraging over the middle and coastal domains of the southeastern Bering Sea in 1997-1999 than has previously been true. Our results are also consistent with hypothesis that the inner front can affect the availability of prey to shearwaters. © 2002 Elsevier Science Ltd. All rights reserved.

Two Bering Sea marine research programs collaborated during the final years of the 1990s to forge advances in understanding the southeastern Bering Sea pelagic ecosystem. Southeast Bering Sea Carrying Capacity, sponsored by NOAA Coastal Ocean Program, investigated processes on the middle and outer shelf and the continental slope. The Inner Front Program, sponsored by NSF, investigated processes of the inner domain and the front between the inner and middle domains. The purposes of these programs were to (1) increase understanding of the southeastern Bering Sea ecosystem, including the roles of juvenile walleye pollock, (2) investigate the hypothesis that elevated primary production at the inner front provides a summer-long energy source for the food web, and (3) develop and test annual indices of pre-recruit pollock abundance. The observations occurred during a period of unusually large variability in the marine climate, including a possible regime shift. Sea-ice cover ranged from near zero to one of the heaviest ice years in recent decades. Sea-surface temperatures reached record highs during summer 1997, whereas 1999 was noted for its low Bering Sea temperatures. Moreover, the first recorded observations of coccolithophore blooms on the shelf were realized in 1997, and these blooms now appear to be persistent. The programs' results include an archive of physical and biological time series that emphasize large year-to-year regional variability, and an Oscillating Control Hypothesis that relates marine productivity to climate forcing. Further investigations are needed of the confluences of interannual and even intra-seasonal variability with low-frequency climate variability as potential producers of major, abrupt changes in the southeastern Bering Sea ecosystem.

During summer 1997, hundreds of thousands of emaciated short-tailed shearwaters (Puffinus tenuirostris) died in the south-eastern Bering Sea. Using strip transect methodology, we documented the distribution and abundance of short-tailed shearwaters during cruises conducted prior to, during, and after the die-off, as well as the distributions and abundances of floating carcasses. The distributions and abundances of short-tailed shearwaters in 1997 were similar to those found during the 1970s and early 1980s. In August-September 1997, we observed 163 floating shearwater carcasses, most of which were between St Paul Island and Nunivak Island. We estimated ∼ 190 000 carcasses were afloat in the study area, about 11% of the surveyed population. Between spring (June) and autumn (August/September), mean net body mass of shearwaters decreased by 19%, mean pectoral muscle mass decreased by 14%, and mean percentage body lipid content decreased by 46%, from 15.6% in spring to 8.4% in autumn. Compared with spring, short-tailed shearwater diets broadened in autumn 1997, to include, in addition to adult euphausiids Thysanoessa raschii, juveniles of T. inermis, T. raschii and T. spinifera, crab megalops, fish and squid. We discuss how the ecosystem anomalies in the south-eastern Bering Sea during spring and summer 1997 relate to the mortality event and suggest possible implications of long-term climate change for populations of apex predators in the south-eastern Bering Sea.

Short-tailed shearwaters Puffinus tenuirostris migrate annually from breeding areas in southeast Australia and Tasmania to the Bering Sea to feed on abundant prey aggregations, mainly euphausiids. Occasionally thousands of shearwaters die of starvation en route, within, or on return from the Bering Sea. Collection of live and dead shearwaters in the southeastern Bering Sea in 1997, 1998, and 1999 allowed us to measure seasonal changes in energy reserves during a major mortality event. As birds lost body mass, lipid mass initially decreased faster than that of pectoralis muscle, but loss of pectoralis mass increased markedly at a body mass around 500 g when lipids were almost depleted (∼33 g remaining). Death occurred as body mass approached 426 g. Individuals near this body mass had lipid values permitting estimated flight ranges of 140 to 400 km, a range less than that potentially covered in 1 d by shearwaters searching for prey (440 to 1124 km d-1). Seasonal differences in body composition were most striking among body and bone marrow lipid contents, with the lowest values occurring during the die-off in fall 1997 and in fall 1998. The lack of shearwater mortality in fall 1998 may have resulted from more consistent winds that decreased flight costs and from greater availability of alternative fish prey. Our data allow estimates of usable energy stores and flight ranges based on lipid reserves in short-tailed shearwaters. Estimated flight ranges suggest that if feeding conditions are poor near Japan or near other termination points of the transequatorial migration routes shearwaters may have few reserves available to support foraging for food and starvation events may occur. Our findings suggest how their energetic strategies and migration are shaped by seasonal and annual variability of prey during transglobal movements of short-tailed shearwaters between oceanic regions.

Anomalies in the regional weather over the southeastern Bering Sea during spring and summer of 1997 resulted in significant differences in nutrient availability, phytoplankton species composition, and zooplankton abundance over the continental shelf as compared with measurements in the 1980s. Calm winds and the reduction of cloud cover in spring and summer produced a very shallow mixed layer in which nitrate and silicate were depleted after an April diatom bloom. High submarine light levels allowed subsequent phytoplankton growth below the pycnocline and eventual depletion of nitrate from the water column to depths of 70 m or more. Thus, total new production during 1997 may have exceeded that of previous years when nitrate was not depleted below the pycnocline. A bloom of the coccolithophorid, Emiliania huxleyi, was observed in early July in the warm, nutrient-depleted waters over the middle and inner shelf. Emiliania huxleyi concentrations reached 4.5 × 106 cells L-1 by September, and the bloom persisted through the autumn. There was evidence for increased abundance of some species of copepods in 1997 as compared with data from the middle domain in June 1981. The abundance of adult and juvenile euphausiids in 1997 was statistically similar to values measured in 1980 and 1981. However, near-surface swarms were rarely observed on the inner shelf in August-September 1997. Lack of euphausiid availability in the upper water column may partially explain the August-September mass mortality of planktivorous short-tailed shearwaters (Puffinus tenuirostris) observed on the inner shelf.

Short-tailed Shearwaters (Puffinus tenuirostris) migrate between breeding areas in Australia and wintering areas in the Bering Sea. These extreme movements allow them to feed on swarms of euphausiids (krill) that occur seasonally in different regions, but they occasionally experience die-offs when availability of euphausiids or other prey is inadequate. During a coccolithophore bloom in the Bering Sea in 1997, hundreds of thousands of Short-tailed Shearwaters starved to death. One proposed explanation was that the calcareous shells of phytoplanktonic coccolithophores reduced light transmission, thus impairing visual foraging underwater. This hypothesis assumes that shearwaters feed entirely by vision (bite-feeding), but their unique bill and tongue morphology might allow nonvisual filter-feeding within euphausiid swarms. To investigate these issues, we developed simulation models of Short-tailed Shearwaters bite-feeding and filter-feeding underwater on the euphausiid Thysanoessa raschii. The visual (bite-feeding) model considered profiles of diffuse and beam attenuation of light in the Bering Sea among seasons, sites, and years with varying influence by diatom and coccolithophore blooms. The visual model indicated that over the huge range of densities in euphausiid swarms (tens to tens of thousands per cubic meter), neither light level nor prey density had appreciable effects on intake rate; instead, intake was severely limited by capture time and capture probability after prey were detected. Thus, for shearwaters there are strong advantages of feeding on dense swarms near the surface, where dive costs are low relative to fixed intake rate, and intake might be increased by filter-feeding. With minimal effects of light conditions, starvation of shearwaters during the coccolithophore bloom probably did not result from reduced visibility underwater after prey patches were found. Alternatively; turbidity from coccolithophores might have hindered detection of euphausiid swarms from the air.

In 1997, the Bering Sea ecosystem, a productive, high-latitude marginal sea, demonstrated that it responds on very short time scales to atmospheric anomalies. That year, a combination of atmospheric mechanisms produced notable summer weather anomalies over the eastern Bering Sea. Calm winds, clear skies, and warm air temperatures resulted in a larger-than-normal transfer of heat to surface waters and the establishment of a shallow mixed layer. In spring, significant new production occurred below the shallow pycnocline over the Middle Shelf, depleting the subpycnocline nutrient reservoir that normally exists during summer. Following the depletion of nitrate and silicate from the system, a sustained (≥ 4 months) bloom of coccolithophores (Emiliania huxleyi) was observed - a phenomenon not previously documented in this region. Summer Middle Shelf Domain copepod concentrations were higher for some species in 1997 than in the early 1980s. Warmer surface water and lack of wind mixing also changed the basic distribution of hydrographic regimes on the south-eastern shelf and altered the strength and position of fronts or transition zones where apex predators seek elevated food concentrations. The Inner Front was well inshore of its normal position, and adult euphausiids (the primary prey of short-tailed shearwaters, Puffinus tenuirostris) were unavailable at, and shoreward of, the front in autumn. High shearwater mortality rates followed the period of low euphausiid availability. Some, but not all, of these anomalous conditions re-occurred in 1998. These observations are another demonstration that the structure and function of marine ecosystems are intimately tied to forcing from the atmosphere. Alteration of climatological forcing functions, expressed as weather, can be expected to have large impacts on this ecosystem and its natural resources.

In the vicinity of the Pribilof Islands in the Bering Sea, abundance of food available to surface-foraging seabirds was greater during the chick-rearing period in 1988 than in 1987, whereas abundance of food available to pursuit-diving seabirds was greater in 1987. Here we examine how breeding success and resource allocation of surface-foraging black-legged kittiwakes Rissa tridacty1a (BLKI) and pursuit-diving thick-billed murres Uria lomvia (TBMU) varied with the fluctuations in their food supply. We also examine a difference in resource allocation among parents raising chicks at the large colony on St. George Island and those at the nearby small colony on St. Paul Island. We studied breeding success (BS), field metabolic rates (FMR, assessed by using doubly labeled water), foraging distribution, and nest attendance of parents and growth rate (GR) of chicks. The BS of BLKIs was lower in 1987 (a season of less abundant food for kittiwakes) than in 1988 (a season of more abundant food), and parents had higher FMRs in 1987 than in 1988. At-sea distributions and nest attendance suggested that in 1987 BLKIs foraged farther from the colonies, which could have resulted in the higher FMR of the parents. GR of BLKI chicks did not vary between 1987 and 1988. The BS of TBMUs was not significantly different between 1987 (a season of more abundant food for TBMUs) and 1988 (a season of less abundant food). Parent TBMUs had similar FMRs between the seasons. Densities of foraging TBMUs were higher within 20 km around colonies in 1987 than in 1988. Although the total time parent TBMUs spent foraging did not vary inter-seasonally, they performed more foraging trips of a shorter duration in 1987 than in 1988, and the GR of TBMU chicks was higher in 1987 than in 1988. Inter-colony comparisons do not suggest that parents reproducing at the large colony work harder to raise their young compared to parents breeding at the small colony. In 1987 parent BLKIs failed in raising young at the large colony, whereas one-third of BLKIs fledged their chicks at the small colony. In 1988, however, RS and FMRs of parent BLKIs were not significantly different between the colonies. Also, TBMUs at the large colony had higher BS than those at the small colony in both 1987 and 1988. Furthermore, in both years parent TBMUs feeding young at the small colony foraged farther from the colony and had significantly higher FMRs than at the large colony. These results suggest that fluctuations in food supply affect resource allocation in seabirds. However, a decrease in food abundance is likely to cause an increase in energy expenditures of parent BLKIs, whereas growth rates of their chicks are less affected. For the TBMUs, food shortages are likely to cause a decrease in growth of the chicks, but not an increase in energy expenditures of the parents.

We studied hydrographic structure, zooplankton distributions, and foraging by planktivorous seabirds in the Anadyr Strait, northern Bering Sea, during 4 summer cruises (1984-1986, 1993). The western portion of the strait was occupied by cold, dense Anadyr water that was mixed from top to bottom. This mixed water was separated from the stratified Bering Shelf water on the eastern side of the strait by a sharp surface front (the 'Anadyr Front'). Net sampling indicated that calanoid copepods were the numerically dominant component of the zooplankton, and that densities of several species were elevated in the frontal zone, apparently due to mechanical accumulation resulting from surface convergence. Hydroacoustic surveys showed that overall zooplankton biomass was concentrated along the thermocline and at the front. Although the location of the Anadyr Front was highly variable over time scales as short as 1 d, large numbers of least auklets Aethia pusilla often flew 25 to 50 km from their breeding colonies to feed at the front. Diet samples indicated that the copepod Neocalanus plumchrus was the principal prey taken by least auklets both at the front and away from it, indicating that heavy use of the distant frontal habitat was due to the higher densities of their preferred prey (i.e. rather than absence of suitable prey species closer to shore). Whenever aggregations of least auklets were found away from the front, there was evidence that they were exploiting near-surface high-density patches of zooplankton, though the exact mechanisms responsible for the formation of such patches are unclear. In contrast to least auklets, crested auklets Aethia cristatella were usually found away from the front. In several cases, compact aggregations of crested auklets were located over acoustically observed epibenthic layers of zooplankton. Hydrographic data suggested that intense subsurface jets and/or upwelling along the eastern side of the strait might have increased the availability of the crested auklets' preferred euphausiid prey. Thus, spatial segregation of the 2 principal planktivores in Anadyr Strait likely arises because different physical mechanisms cause concentrations of preferred prey originating at different depths.

We studied hydrographic structure, zooplankton distributions, and foraging by planktivorous seabirds in the Anadyr Strait, northern Bering Sea, during 4 summer cruises (1984-1986, 1993). The western portion of the strait was occupied by cold, dense Anadyr water that was mixed from top to bottom. This mixed water was separated from the stratified Bering Shelf water on the eastern side of the strait by a sharp surface front (the 'Anadyr Front'). Net sampling indicated that calanoid copepods were the numerically dominant component of the zooplankton, and that densities of several species were elevated in the frontal zone, apparently due to mechanical accumulation resulting from surface convergence. Hydroacoustic surveys showed that overall zooplankton biomass was concentrated along the thermocline and at the front. Although the location of the Anadyr Front was highly variable over time scales as short as 1 d, large numbers of least auklets Aethia pusilla often flew 25 to 50 km from their breeding colonies to feed at the front. Diet samples indicated that the copepod Neocalanus plumchrus was the principal prey taken by least auklets both at the front and away from it, indicating that heavy use of the distant frontal habitat was due to the higher densities of their preferred prey (i.e. rather than absence of suitable prey species closer to shore). Whenever aggregations of least auklets were found away from the front, there was evidence that they were exploiting near-surface high-density patches of zooplankton, though the exact mechanisms responsible for the formation of such patches are unclear. In contrast to least auklets, crested auklets Aethia cristatella were usually found away from the front. In several cases, compact aggregations of crested auklets were located over acoustically observed epibenthic layers of zooplankton. Hydrographic data suggested that intense subsurface jets and/or upwelling along the eastern side of the strait might have increased the availability of the crested auklets' preferred euphausiid prey. Thus, spatial segregation of the 2 principal planktivores in Anadyr Strait likely arises because different physical mechanisms cause concentrations of preferred prey originating at different depths.

1. The foraging ecology of Brunnich's guillemots Uria lomvia was studied during the breeding season in south-eastern Svalbard. In the region of Storfjorden there are two large breeding colonies comprising a total of about 540 000 individuals. These birds forage in the western part of Storfjorden and further to the south. Their main prey: are polar cod Boreogadus saida, pelagic amphipods Parathemisto spp. and euphausiids Thysanoessa inermis. 2. A ship-based transect survey was used to record bird abundance and the acoustically determined biomass of presumed prey. The five transects were divided into 33 segments, each 8-11 km in length. The resolution of the survey was 150 m, and analyses of correlations between predators and prey were performed at length scales: from 150 m to 9 km. We differentiated acoustic signals into aggregated and dispersed categories according to the estimated horizontal distribution of presumed prey. 3. Foraging guillemots were consistently more strongly correlated with the aggregated prey than with dispersed prey over scales ranging from 150 m to 9 km. Correlations were weak at small scales (150 m - 1 km) and increased and stabilized at scales of 23 km. The spatial scale at which we obtained a shift from weak to strong correlations between guillemots and their prey was similar to the scale at which the spatial variances in both guillemot and prey abundance were high. 4. Guillemots showed low correlations with prey at low prey densities. Similarly, correlations between guillemots and prey were low at low bird densities. The data support the hypothesis that the birds associate with prey patches with densities above a certain threshold, and that 'regional' prey abundances affect local use of patches. 5. The numerical aggregative response curves between guillemot and prey density were classified as being neither hyperbolic (type II) or sigmoidal (type III) within the range of prey densities observed in this study. The aggregative response curves were sensitive to spatial scales, which suggest that studies of response curves should be conducted at a range of spatial scales.

The at-sea distribution of thick-billed murres (Uria lomvia) in southeastern Svalbard waters was studied during the summers of 1992, 1993, and 1996. The Storfjordrenna region south of Svalbard was confirmed as an important foraging area for thick-billed murres; murre aggregations were located at distances of 85 to 126 km from the closest breeding colonies. Fish, mainly polar cod (Boreogadus saida), but also capelin (Mallotus villosus), were the main prey found in 16 murres and 3 black-legged kittiwakes (Rissa tridactyla) collected from these aggregations. Murres were seen flying with fish in their beaks at four locations 78 to 102 km away from the colonies. Murre aggregations were associated with frontal zones between cold Arctic waters and warmer Atlantic water, and in areas with strong stratification in salinity at 15-30 m. A positive association was found between the abundance of murres and the occurrence of cetaceans. Murres and other marine birds were often seen near surfacing cetaceans. The most common cetaceans were minke whales (Balaenoptera acutorostrata) and white-beaked dolphins (Lagenorhynchus albirostris).

We tested the hypothesis that the spatial distributions of foraging least, crested and parakeet auklets (Aethia pusilla, A. cristatella and A. psittacula, respectively) in the shallow passes of the Aleutian Islands would be determined by physical mechanisms that control near-surface prey concentrations. We recorded currents using an Acoustic Doppler Current Profiler, volume scattering using 200 and 420 kHz scientific echosounders, and the numbers of foraging birds. Zooplankton were sampled using a multiple opening/closing net and environmental sampling system (MOCNESS). Prey choice of birds was ascertained by collecting foraging birds and examining their stomach contents. Most sampling occurred between 8 July and 6 August 1993, when we conducted 50 passages along a transect that crossed a sill between Unalga and Kavalga Islands, western Aleutian Islands, thereby sampling the foraging activity of auklets at a variety of times of day and tidal phases. We found that the abundance of foraging individuals of each of the 3 auklet species was a function of tidal speed. Auklet species were selective about the species of prey taken. Regardless of tidal direction, crested auklets foraged on euphausiids upwelled on the upstream side of the pass, whereas least auklets consumed copepods concentrated in near-surface convergences on the downstream side. Parakeet auklets foraged over the top of the pass and took fish and invertebrates. Tidal speed and direction influenced the distance between the peak numbers of some, but not all, species of auklets. Auklet prey preferences dictated where they foraged in the pass and the physical mechanisms exploited for successful foraging. Thus, in this instance, resource partitioning by these closely related planktivores was enhanced by a spatial segregation forced by the physical processes that enhanced the availability of prey. Our findings emphasize the important role of physical processes in the structuring of marine communities.

Hydrographic, acoustic and zooplankton data were collected during June 1992 and June 1995 along transects to the north and south of Kiska and Buldir Islands in the western Aleutian Islands to look for mesoscale relationships between zooplankton community composition, biomass, and associated frontal systems and pycnoclines. Hydrographic data revealed a front dividing the Bering Sea water mass from the mixed regime in the passes and Alaska Stream water south of the islands. Chlorophyll concentration and ADB (acoustically determined biomass) were consistently higher on the Bering Sea side of the front in June 1992. Elevated ADB in the Bering Sea was due to aggregates of Neocalanus cristatus copepodid V in the pycnocline separating the upper mixed layer from the underlying cold intermediate layer. N. cristatus abundance was significantly higher (p < 0.05) in Bering Sea water in June 1992 than in Pacific water in June 1992 and Bering Sea water in June 1995. N. fleminger abundance was significantly higher in Bering Sea water than in Pacific water in both 1992 and 1995. N. plumchrus and Eucalanus bungil stages III and IV were more abundant in Pacific water than in Bering Sea water. The results clearly demonstrate the utility of interfacing acoustics, net samples and hydrographic data to elucidate mesoscale associations of zooplankton to fronts, water masses and pycnoclines.

We investigated the relationship between surface-foraging prions Pachyptila spp. and their zooplankton prey by comparing counts of foraging birds with near-surface concentrations of zooplankton from net samples. Zooplankton abundance was assessed by employing a new design of sampler that combined a frame net with a pump at the cod-end and which was deployed at the sea surface. Six transects using the sampler with concurrent bird observations were made over a short stretch of Stewart Strait near Bird Island, South Georgia. Concentrations of a variety of zooplankton species occurred patchily along each transect. Peak numbers of prions were recorded consistently towards the southern end of each transect. These peaks coincided with peaks in zooplankton numbers, but with different prey species predominant on different occasions. Comparison of the distribution of foraging and sitting birds with the presence or absence of zooplankton peaks, after pooling zooplankton species into 2 broad classes, showed that birds were consistently observed over copepod maxima, but were not found to be associated with aggregations of larger zooplankton more often than expected by chance.

In the eastern Bering Sea, the overall distribution of least auklets (Aethia pusilia) reflects the distribution of different water masses and their associated plankton communities. Within these water masses, the foraging distribution of least auklets is influenced by physical processes that result in predictable aggregations of prey, most often because of property or flow gradients with which the plankton interact. Examples include the accumulation of zooplankton at, or above, the pycnocline and near-surface patches of plankton concentrated in downwellings formed at convergences. These small-scale processes influence avian use of the larger-scale features. Breeding colonies of least auklets are located within commuting distance of physical features in the ocean at which appropriate prey are concentrated; where large copepods or physical features to concentrate them are absent, auklets do not breed. Thus, control of the distribution of foraging seabirds and their breeding colonies in the eastern Bering Sea is a multi-scale process.

In the SE Bering Sea, sea surface temperatures increased from the late 1970s to the mid-1980s, then decreased. Over the same period, there were inter-annual variations in the water masses surrounding the Pribilof Islands. Subsequent to the mid-1970s, there were changes in the abundance of capelin Mallotus villosus and 1-group walleye pollock Theragra chalcogramma. Use of capelin by both black-legged kittiwakes Rissa tridactyla and red-legged kittiwakes R. brevirostris decreased at the Pribilof Islands subsequent to 1978, as did the use of 1-group pollock in the late 1970s. Availability of fatty fishes such as myctophids, capelin and sand lance decreased after the late 1970s. Beginning in the late 1970s, there was a decrease in the number of chicks produced per nest for both black-legged and red-legged kittiwakes nesting on the Pribilof Islands. Inter-annual variation in the availability of fatty fish was at least in part responsible for variations in the production of chicks by red-legged and possibly by black-legged kittiwakes.

We studied short-tailed shearwaters Puffinus tenuirostris foraging near the Pribilof Islands, Alaska, USA, during the summers of 1987, 1988, and 1989. Their foods were almost exclusively the euphausiid Thysanoessa raschii, which they obtained both from near-surface swarms and from epibenthic layers. Near-surface mating swarms of euphausiids occurred in areas of elevated phytoplankton standing stocks near inshore tidal fronts. Many of these euphausiids had attached spermatophores. Shearwaters also obtained euphausiids over shallow reefs and inshore of the fronts where euphausiids were trapped in water shallower than 40 m by irregularities in bottom topography ('bathymetric traps'). We hypothesize that the largely inshore distribution of shearwaters in the southeastern Bering Sea described by previous workers is the result of attraction of shearwaters to frontal areas where euphausiids may forage on phytoplankton stocks throughout the summer. These areas, when shallower than 40 m, would also permit shearwaters to access epibenthic aggregations of euphausiids during daylight, when euphausiids not engaged in mating swarms usually migrate to depth.

We investigated the foraging distribution of 2 species of murres (Uria lomvia and U. aalge) in relation to Acoustically Determined Biomass (ADB) and hydrographic structure surrounding the Pribilof Islands, Bering Sea, Alaska, USA. The distribution and abundance of both murre species were similar; therefore, we combined data for thick-billed and common murres in the analysis. We found that murre densities were higher in frontal regions than in non frontal regions. Maximum ADB was also found to be elevated in the frontal zone. Within frontal regions, high densities of murres were associated with regions containing relatively high levels of ADB. In contrast, we did not find significant correlations between murre density and ADB in the study area as a whole. We postulate that murres restrict their foraging to the frontal region because prey can be located more predictably within this zone than in other areas in the vicinity of the Pribilof Islands.

We recorded the distribution and abundance of marine birds in the northern Barents Sea from 27 February to 8 March 1987 and from 20 to 31 May 1988. Birds were more abundant in waters associated with pack ice than in open water away from pack ice. Within the pack ice, thick-billed murres (Uria lomvia) were the most commonly encountered birds in both periods. Murre densities in the pack ice north of the zone proximate to the ice edge were positively correlated with distance from the ice edge. Large leads were more frequently occupied by murres than small leads, and had larger numbers of birds present. In spring, we found more birds along a well-defined ice edge than were present either in open water or in leads in the pack ice within 5 nautical miles of the ice edge. Transects along the ice edge revealed little correlation in abundance between species, or within species when coverage was repeated during the same day. We conclude that the birds showed considerable specificity of habitat choice within the habitat divisions that we recognized and that avian patches were of short duration. We need information on the distribution, abundance and movements of prey patches if we are to understand the changing distribution patterns of the birds.

We studied the influence of the distribution of prey and hydrographic fronts on the spatial distribution of foraging Brunnich's guillemots (Uria lomvia) in Storfjorden, southeastern Svalbard in late July 1992. Two large breeding colonies, comprising a total of 540,000 individuals, were located adjacent to the study area, and large numbers of Brunnich's gullemots from these colonies foraged within the area, as well as to the south, outside of Storfjorden. Within the study area, most guillemots foraged on the west side of the fjord, coincident with a weak subsurface front between warm Atlantic water, which penetrated Storfjorden from the south, and cold Arctic water. Food samples from the guillemots collected in the study area contained primarily crustaceans (Parathemisto spp. and Thysanoessa inermis) and polar cod Boreogadus saida. Acoustic observations of prey were differentiated into two classes of signals, which we interpreted as originating from aggregated and dispersed organisms. The numbers of foraging guillemots were strongly correlated with the strength of echoes of the aggregated type, whereas correlations with dispersed echoes were consistently weaker. The distribution of foraging guillemots showed no significant correlations with either horizontal or vertical gradients of physical properties of the water column. Our finding that guillemots respond differently to aggregated and dispersed prey has important implications both for the interpretation of past work on the foraging ecology of marine birds, and for the management of fisheries.

The authors studied the relationship between the foraging radius and energy economy of A. pusilla breeding in colonies on three islands in the Bering Sea (St Lawrence, St Matthew, and St George Islands). The distance to which auklets commuted on foraging trips varied by more than an order of magnitude c5-56 km), but mean field metabolic rate did not vary significantly among birds from the three islands. The authors modeled the partitioning of energy to various activities and hypothesized that the added cost of commuting incurred by auklets from St Lawrence Island (foraging radius, 56 km) was offset by reduced energy costs while foraging at sea. Data indicated that auklets from St Lawrence Island fed on larger, more energy-rich copepods than auklets from St Matthew Island (foraging radius, 5 km). -from Authors