The abundance and distribution of euphausiids and zero-age pollock on the inner shelf of the southeast Bering Sea near the Inner Front in 1997–1999

Acoustic data and net samples were collected during late spring and early fall 1997–1999 to assess zooplankton and micronekton abundance and distribution relative to the Inner Front at three sampling grids (Port Moller, Cape Newenham and Nunivak Island) on the inner shelf of the southeast Bering Sea. Epibenthic scattering layers were observed during May–June and August–September in all three years. Acoustic data were scaled to euphausiid biomass using target strength models. Mean euphausiid biomass determined acoustically for each transect line was 0.7–21 g m⁻², with most values below 5 g m⁻². There was no consistent relationship between the distribution and biomass of euphausiids and the location of the Inner Front. Zero age pollock were observed on the inner shelf in August–September during all years, but were confined primarily to the stratified side of the Inner Front and to the frontal regime. The acoustic data for pollock were scaled to biomass using laboratory measurements of gas bladder dimensions and target strength models. Acoustic determinations of mean transect biomass for euphausiids did not differ from literature values for the inner shelf of the southeast Bering Sea, and pollock biomass on the inner shelf did not differ from that around the Pribilof Islands. Despite recent anomalies in climate and oceanographic conditions on the inner shelf, and high mortality of shorttail shearwaters during 1997, we found no evidence of significant interannual differences in the biomass of euphausiids or zero-age pollock on the inner shelf of the southeast Bering Sea.     

Diets of short-tailed shearwaters in the southeastern Bering Sea

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.     

Interannual and decadal variability in zooplankton communities of the southeast Bering Sea shelf

The southeastern Bering Sea shelf ecosystem is an important fishing ground for fin- and shellfish, and is the summer foraging grounds for many planktivorous seabirds and marine mammals. In 1997 and 1998, Northern Hemisphere climate anomalies affected the physical and biological environment of the southeastern Bering Sea shelf. The resulting anomalous conditions provided a valuable opportunity to examine how longer-term climate change might affect this productive ecosystem. We compared historical and recent zooplankton biomass and species composition data for the southeastern Bering Sea shelf to examine whether or not there was a response to the atmosphere–ocean–ice anomalies of 1997 and 1998. Summer zooplankton biomass (1954–1994) over the southeastern shelf did not exhibit a decline as previously reported for oceanic stations. In addition, zooplankton biomass in 1997 and 1998 was not appreciably different from other years in the time series. Spring concentrations of numerically abundant copepods (Acartia spp., Calanus marshallae, and Pseudocalanus spp.), however, were significantly higher during 1994–1998 than 1980–1981; spring concentrations of Metridia pacifica and Neocalanus spp. were not consistently different between the two time periods. Neocalanus spp. was the only taxon to have consistent differences in stage composition between the two time periods—CV copepodites were much more prevalent in May of the 1990s than early 1980s. Since relatively high zooplankton concentrations were observed prior to 1997, we do not attribute the high concentrations observed in the summers of 1997 and 1998 directly to the acute climate anomalies. With the present data it is not possible to distinguish between increased production (control from below) and decreased predation (control from above) to explain the recent increase in concentrations of the species examined.     

Characteristics and variability of the inner front of the southeastern Bering Sea

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.     

Ecological characteristics of Walleye pollock eggs and larvae in the southeastern Bering Sea during the late 1970s

Walleye pollock (Theragra chalcogramma) is an ecologically and economically important groundfish in the eastern Bering Sea. Its population size fluctuates widely, driving and being driven by changes in other components of the ecosystem. It is becoming apparent that dramatic shifts in climate occur on a decadal scale, and these “regime shifts” strongly affect the biota. This paper examines quantitative collections of planktonic eggs and larvae of pollock from the southeastern Bering Sea during 1976–1979. Mortality, advection, and growth rates were estimated, and compared among the years encompassing the 1970s’ regime shift. These data indicate that pollock spawning starts in late February over the basin north of Bogoslof Island. Over the shelf, most spawning occurs north of Unimak Island near the 100 m isobath in early or mid April. Pollock eggs are advected to the northwest from the main spawning area at 5–10 cm/sec. Larvae are found over the basin north of Bogoslof Island in April, and over the shelf between Unimak Island and the Priblof Islands in May. Compared to 1977, the spawning period appeared to be later in 1976 (a cold year) and earlier in 1978 (a warm year) in the study area. At the lower temperatures in 1976, egg duration would be longer and thus egg mortality would operate over a longer period than in the other years. Mean larval growth appeared to be lower in 1976 than in 1977 and 1979. Estimated egg mortality rate in 1977 was 0.6 in April and 0.3 in early May.     

Climate change and control of the southeastern Bering Sea pelagic ecosystem

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 cold-water 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.     

Modelling phytoplankton succession on the Bering Sea shelf: role of climate influences and trophic interactions in generating Emiliania huxleyi blooms 1997–2000

Several years of continuous physical and biological anomalies have been affecting the Bering Sea shelf ecosystem starting from 1997. Such anomalies reached their peak in a striking visual phenomenon: the first appearance in the area of bright waters caused by massive blooms of the coccolithophore Emiliania huxleyi (E. huxleyi). This study is intended to provide an insight into the mechanisms of phytoplankton succession in the south-eastern part of the shelf during such years and addresses the causes of E. huxleyi success by means of a 2-layer ecosystem model, field data and satellite-derived information. A number of potential hypotheses are delineated based on observations conducted in the area and on previous knowledge of E. huxleyi general ecology. Some of these hypotheses are then considered as causative factors and explored with the model. The unusual climatic conditions of 1997 resulted most notably in a particularly shallow mixed layer depth and high sea surface temperature (about 4 °C above climatological mean). Despite the fact that the model could not reproduce for E. huxleyi a clear non-bloom to bloom transition (pre- vs. post-1997), several tests suggest that this species was favoured by the shallow mixed layer depth in conjunction with a lack of photoinhibition. A top-down control by microzooplankton selectively grazing phytoplankton other than E. huxleyi appears to be responsible for the long persistence of the blooms. Interestingly, observations reveal that the high N:P ratio hypothesis, regarded as crucial in the formation of blooms of this species in previous studies, does not hold on the Bering Sea shelf.     

On the recent warming of the southeastern Bering Sea shelf

During the last decade, the southeastern Bering Sea shelf has undergone a warming of 3 °C that is closely associated with a marked decrease of sea ice over the area. This shift in the physical environment of the shelf can be attributed to a combination of mechanisms, including the presence over the eastern Bering Sea shelf of a relatively mild air mass during the winter, especially from 2000 to 2005; a shorter ice season caused by a later fall transition and/or an earlier spring transition; increased flow through Unimak Pass during winter, which introduces warm Gulf of Alaska water onto the southeastern shelf; and the feedback mechanism whereby warmer ocean temperatures during the summer delay the southward advection of sea ice during winter. While the relative importance of these four mechanisms is difficult to quantify, it is evident that for sea ice to form, cold arctic winds must cool the water column. Sea ice is then formed in the polynyas during periods of cold north winds, and this ice is advected southward over the eastern shelf. The other three mechanisms can modify ice formation and melt, and hence its extent. In combination, these four mechanisms have served to temporally and spatially limit ice during the 5-year period (2001–2005). Warming of the eastern Bering Sea shelf could have profound influences on the ecosystem of the Bering Sea—from modification of the timing of the spring phytoplankton bloom to the northward advance of subarctic species and the northward retreat of arctic species.     

Variations in age-0 pollock distribution among eastern Bering Sea nursery areas: A comparative study through acoustic indices

Cohort abundance of walleye pollock (Theragra chalcogramma) is subject to strong interannual variation in the eastern Bering Sea, and this variation is known to be determined largely at the age-0 stage. We estimated the spatial distributions and densities of age-0 walleye pollock in five nursery areas around the eastern Bering shelf in three successive years (1997–1999) from acoustic survey data. Concurrently, we calculated estimates of the spatial distribution of euphausiids, a major prey of age-0 walleye pollock, and estimates of spatial overlap of groundfish predators with the age-0 walleye pollock. The analyses showed that all nursery areas had low densities of age-0 walleye pollock in 1997, which ultimately produced the weakest adult year-class. In the intermediate year of 1998, age-0 densities were low to medium, and in 1999, which produced the strongest of the three adult year-classes, all nursery areas had medium to high age-0 walleye pollock densities. Euphausiid distributions had a consistently positive spatial relationship with age-0 walleye pollock. Groundfish predator density ratios were positively related to age-0 walleye pollock density when age-0 walleye pollock were displaced relatively northward. Our results suggest that abundance of age-0 walleye pollock, and hence of adult cohorts in the eastern Bering Sea, can be predictable from a concise set of indicators: the densities of age-0 walleye pollock at nursery areas in mid- to late-summer, their spatial relationship to euphausiids and groundfish predators, and the latitudinal trend of their distributions. The 3 years 1997–1999 had significant differences of physical conditions in the eastern Bering Sea, and represent an advantageous framework for testing these hypotheses.     

Comparison of atmospheric forcing in four sub-arctic seas

A comparative analysis was conducted on climate variability in four sub-arctic seas: the Sea of Okhotsk, the Bering Sea shelf, the Labrador Sea, and the Barents Sea. Based on data from the NCEP/NCAR reanalysis, the focus was on air–sea interactions, which influence ice cover, ocean currents, mixing, and stratification on sub-seasonal to decadal time scales. The seasonal cycles of the area-weighted averages of sea-level pressure (SLP), surface air temperature (SAT) and heat fluxes show remarkable similarity among the four sub-arctic seas. With respect to variation in climate, all four seas experience changes of comparable magnitude on interannual to interdecadal time scales, but with different timing. Since 2000 warm SAT anomalies were found during most of the year in three of the four sub-arctic seas, with the exception of the Sea of Okhotsk. A seesaw (out of phase) pattern in winter SAT anomalies between the Labrador and the Barents Sea in the Atlantic sector is observed during the past 50 years before 2000; a similar type of co-variability between the Sea of Okhotsk and the Bering Sea shelf in the Pacific is only evident since 1970s. Recent positive anomalies of net heat flux are more prominent in winter and spring in the Pacific sectors, and in summer in the Atlantic sectors. There is a reduced magnitude in wind mixing in the Sea of Okhotsk since 1980, in the Barents Sea since 2000, and in early spring/late winter in the Bering Sea shelf since 1995. Reduced sea-ice areas are seen over three out of four (except the Sea of Okhotsk) sub-arctic seas in recent decades, particularly after 2000 based on combined in situ and satellite observations (HadISST). This analysis provides context for the pan-regional synthesis of the linkages between climate and marine ecosystems.     

Polychaete assemblage as surrogate for prey availability in assessing southeastern Bering Sea flatfish habitat

The flatfish yellowfin sole (Limanda aspera), northern rock sole (Lepidopsetta polyxystra), and Alaska plaice (Pleuronectes quadrituberculatus) in the southeastern Bering Sea prey mainly on infauna. Spatial correspondence between their stomach contents and infauna assemblages across habitat types was examined to identify indices of prey availability for flatfish habitat characterization and quality assessment. Benthic samples and flatfish stomachs were collected in 2009 near the Alaska Peninsula in the southeastern Bering Sea. Polychaetes and bivalves were the most dominant infauna groups, each comprising 35–60% by weight in each infauna sample. These two were also the only prey groups that frequently averaged > 50% of stomach content by weight. Bivalves dominated the infauna biomass on the relatively sandy inner shelf (0–50 m depth). The muddier middle shelf (50–100 m) had the highest infauna biomass, which was dominated by polychaetes. Diet compositions of the flatfish varied spatially in correspondence with the infauna assemblage. Polychaetes were prevalent in all flatfish diets on the middle shelf, even yellowfin sole whose typical primary prey are amphipods and bivalves. Polychaete-rich habitats are potentially prime for flatfish as polychaetes are readily utilized where available and generally have high nutritional value. Flatfish did not select for specific polychaete taxa, so an index of habitat quality could be based on the biomass of aggregate polychaetes or on dominant polychaete families of the region. Under normal environmental conditions, the three flatfish have slightly-offset spatial distributions, enabling each to utilize different infauna assemblages across the shelf. However, during cold phases in the Bering Sea ecosystem, as when this study was conducted, a cold pool of < 2 °C bottom water from the spring ice melt extends over the middle shelf in summer. This physiological barrier displaces all three flatfish to the inner shelf, intensifying competition for prey resources.     

The Bering Sea—A dynamic food web perspective

The Bering Sea is a high-latitude, semi-enclosed sea that supports extensive fish, seabird, marine mammal, and invertebrate populations and some of the world's most productive fisheries. The region consists of several distinct biomes that have undergone wide-scale population variation, in part due to fisheries, but also in part due to the effects of interannual and decadal-scale climatic variation. While recent decades of ocean observation have highlighted possible links between climate and species fluctuations, mechanisms linking climate and population fluctuations are only beginning to be understood. Here, we examine the food webs of Bering Sea ecosystems with particular reference to some key shifts in widely distributed, abundant fish populations and their links with climate variation. Both climate variability and fisheries have substantially altered the Bering Sea ecosystem in the past, but their relative importance in shaping the current ecosystem state remains uncertain.     

The distribution of chlorophyll a and its influencing factors in different regions of the Bering Sea

The distribution of chlorophyll a(Chl a) and its relationships with physical and chemical parameters in different regions of the Bering Sea were discussed in July 2010. The results showed the seawater column Chl a concentrations were 13.41–553.89 mg/m2 and the average value was 118.15 mg/m2 in the study areas. The horizontal distribution of Chl a varied remarkably from basin to shelf in the Bering Sea. The regional order of Chl a concentrations from low to high was basin, slope, outer shelf, inner shelf, and middle shelf. The vertical distribution of Chl a was grouped mainly from single-peak type in basin, slope, outer shelf, and middle shelf, where the deep Chl a maxima(DCM) layer was observed at 25–50 m, 30–35 m, 36–44 m, and 37–47 m, respectively. The vertical distribution of Chl a mainly had three basic patterns: standard single-peak type, surface maximum type, and bottom maximum type in the inner shelf. The analysis also showed that the transportation of ocean currents may control the distribution of Chl a, and the effects were not simple in the basin of the Bering Sea. There was a positive correlation between Chl a and temperature, but no significant correlation between Chl a and nutrients. The Bering Sea slope was an area deeply influenced by slope current. Silicate was the factor that controlled the distribution of Chl a within parts of the water in the slope. Light intensity was an important environmental factor in controlling seawater column Chl a in the shelf, where Chl a was limited by nitrate rather than phosphate within the upper water. Meanwhile, there was a positive relationship between Chl a and salinity. Algal blooms broke out at Sta. B6 of the southwestern St. Lawrence Island and Stas F6 and F11 in the middle of the Bering Strait.     

Low-temperature incubation of walleye pollock (Theragra chalcogramma) eggs from the southeast Bering Sea shelf and Shelikof Strait, Gulf of Alaska

To determine the effect of low water temperature on development, walleye pollock (Theragra chalcogramma) eggs from the Bering Sea were reared at −0.6°C, 0.4°C, 2.0°C, and 3.8°C. One group of eggs was reared at 3.9°C under a diel light cycle (14 h light, 10 h dark) to observe the effect of light on development and hatching. Development was normal for all temperatures except −0.6°C; abnormal development of the tail and lack of development of eyes occurred in some embryos. Time to 50% hatch was 820, 620, and 424 h at 0.4°C, 2.0°C, and 3.8°C. Eggs incubated in diel light at 3.9°C developed at the same rate as eggs incubated in constant dark at 3.8°C, but required an additional 72 h to reach 50% hatch. A piece-wise regression model was generated to predict egg age for incubation temperatures of −0.6°C to 3.8°C. For temperatures recorded in the southeastern Bering Sea 1995–1998, the model predicted incubation periods for walleye pollock eggs that varied by 13 days between the warmest and coldest years.Walleye pollock eggs from Shelikof Strait, Alaska, were incubated at 0.2°C, 1.8°C, and 2.8°C. Development was normal for all temperatures. A piece-wise regression model (as above) was generated for incubation temperatures 0.2–2.8°C. When the regression models were compared, Bering Sea eggs (1.4–1.7 mm in diameter), required more time for development prior to hatch than Shelikof Strait eggs (1.2–1.3 mm in diameter) at 1.8°C and 2.8°C. However, for temperatures 0.2–2.0°C, Bering Sea walleye pollock began hatching earlier and at a developmentally younger age than Shelikof Strait walleye pollock.     

Size-dependent, Spatial, and Temporal Variability of Juvenile Walleye Pollock (Theragra chalcogramma) Feeding at a Structural Front in the Southeast Bering Sea

Abstract. The waters surrounding the Pribilof Islands are an important nursery ground for juvenile walleye pollock (Theragra chalcogramma), an important forage fish in the pelagic food web of the productive Bering Sea shelf region. The diet of juvenile pollock was studied in two consecutive years along a transect line crossing from a well‐mixed coastal domain, through a frontal region to stratified water farther offshore. Variability in stomach fullness was high and evidence for increased feeding intensity in the front was weak. Prey diversity and prey size generally increased with increasing fish size, shifting from predominantly small copepods to larger, more evasive prey items such as euphausiids, crab megalopae and fish. The diet of the fish reflected changes in the relative abundance of copepods and euphausiids in the prey fields between years. Juvenile pollock showed increased feeding rates at dusk, and stomach fullness as well as prey condition were generally lowest just before sunrise; however, the proportion of euphausiids increased in the diet of pollock caught at night, suggesting that some food was also ingested during darkness. Juvenile pollock and their euphausiid prey both vertically migrated above the thermocline at night, although each had a different daytime depth.     

A regional tidal/subtidal circulation model of the southeastern Bering Sea: development, sensitivity analyses and hindcasting

A regional eddy-resolving primitive equation circulation model was used to simulate circulation on the southeastern Bering Sea (SEBS) shelf and basin. This model resolves the dominant observed mean currents, eddies and meanders in the region, and simultaneously includes both tidal and subtidal dynamics. Circulation, temperature, and salinity fields for years 1995 and 1997 were hindcast, using daily wind and buoyancy flux estimates, and tidal forcing derived from a global model. This paper describes the development of the regional model, a comparison of model results with available Eulerian and Lagrangian data, a comparison of results between the two hindcast years, and a sensitivity analysis. Based on these hindcasts and sensitivity analyses, we suggest the following: (1) The Bering Slope Current is a primary source of large (100 km diameter) eddies in the SEBS basin. Smaller meanders are also formed along the 100 m isobath on the southeastern shelf, and along the 200-m isobath near the shelf break. (2) There is substantial interannual variability in the statistics of eddies within the basin, driven by variability in the strength of the ANSC. (3) The mean flow on the shelf is not strongly sensitive to changes in the imposed strength of the ANSC; rather, it is strongly sensitive to the local wind forcing. (4) Vertical mixing in the SEBS is strongly affected by both tidal and subtidal dynamics. Strongest mixing in the SEBS may in fact occur between the 100- and 400-m isobaths, near the Pribilof Islands, and in Unimak Pass.     

Combining field observations and modeling approaches to examine Greenland halibut (Reinhardtius hippoglossoides) early life ecology in the southeastern Bering Sea

Spawning in Greenland halibut (Reinhardtius hippoglossoides) occurs along the continental slope and in submarine canyons in the eastern Bering Sea. It is assumed that these bathymetric features and their associated circulation patterns deliver eggs and larvae to suitable nursery habitats over the continental shelf. However, there have been no directed field studies examining spawning areas or transport of Greenland halibut early life stages in the Bering Sea, nor is it known how large-scale oceanographic forcing modulates specific physical mechanisms of delivery. The present study was undertaken to: better define spawning areas of Greenland halibut, examine development and distribution of larvae, and understand the influence of climate variations on interannual patterns of transport, distribution and abundance. Eggs were found in Bering and Pribilof Canyons and over the adjacent slope in February and early March, confirming that spawning occurs in these regions. Larvae were present over the slope, outer shelf and middle shelf in winter and spring, and settled juveniles were collected over the shelf in September. Oceanographic modeling approaches that simulate larval advection from spawning to nursery habitats indicate that depth-discrete variations in transport pathways from submarine canyons to the adjacent shelf contribute to interannual variability in transport trajectories. Overall, our results highlight specific physical mechanisms of delivery that are modulated by large-scale atmospheric and oceanographic forcing, potentially varying the degree of slope–shelf connectivity for Greenland halibut and other slope-spawning species.     

Stable C and N isotopic composition of sinking particles and zooplankton over the southeastern Bering Sea shelf

Stable carbon and nitrogen isotopic composition of zooplankton, suspended particulate organic matter (SPOM), and sinking particles collected using sediment traps were measured for samples obtained from the southeastern Bering Sea middle and outer shelf during 1997–1999. The quantity of material collected by the middle shelf sediment trap was greater in both spring and late summer and fall than in early and mid-summer. The δ¹⁵N of SPOM, sinking material and zooplankton showed greater inter-annual variability at the middle shelf site (M2) than at the outer shelf site (M3). Zooplankton and sinking organic matter collected by M2 sediment traps became more depleted in ¹⁵N from 1997 through 1999, associated with a change from unusually warm to unusually cold conditions. Suspended and sinking organic matter and zooplankton collected from M3 decreased only slightly in δ¹⁵N from 1998 to 1999. SPOM, zooplankton, and sediment trap samples collected at M2 were usually enriched in δ¹⁵N and δ¹³C over those from M3. However, in 1999 sediment trap samples from the middle shelf were enriched in ¹³C over M3 material, but the δ¹⁵N of samples from the two sites was similar. The geographic pattern could be explained greater productivity over the middle shelf, associated with either isotopically heavy nitrogen being regenerated from sediments, or with utilization of a greater fraction of the available inorganic nitrogen pool during most years.     

Cooling in the western Bering Sea in 1999: quick propagation of La Niña signal or compensatory processes effect?

In 1999, synoptic and hydrological conditions in the western Bering Sea were characterized by negative SST and air temperature anomalies, extensive ice coverage and late melting. Biological processes were also delayed. In 1999, the average zooplankton biomass was 1.76 g/m³, approximately half the average 3.07 g/m³ in 1998. Pacific salmon migrated to the northeastern Kamchatka streams two weeks later. This contrasts with 1997 (spring and summer) and 1998 (summer) when positive SST anomalies were widely distributed throughout the northwestern Bering Sea shelf. Since the second half of the 1990s, seasonal atmospheric processes developed over the western Bering Sea that were similar to those of the cold decades of the 1960–1970s. A meridional atmospheric circulation pattern began to replace zonal transport. Colder Arctic air masses have shifted over the Bering Sea region and shelf water temperatures have cooled considerably with the weakening of zonal atmospheric circulation. Temperature decreased in the cold intermediate layer during its renewal in winter. Besides, oceanic water inflow intensified into the Bering Sea in intermediate layers. Water temperature warmed to 4°C and a double temperature maximum existed in the warm intermediate layer in late summer in both 1997 and 1998. Opposing trends of cold water temperature and a warm intermediate layer led to an increase of vertical gradients in the main thermocline and progressing frontogenesis. It accelerates frontal transport and can be regarded as a chief cause of increased water exchange with the Pacific Ocean.     

Plankton distribution associated with frontal zones in the vicinity of the Pribilof Islands

We studied the effect of four types of fronts, the coastal front, the middle front, the shelf partition front and the shelf break front on the quantitative distribution and the composition of plankton communities in the Pribilof area of the eastern Bering Sea shelf in late spring and summer of 1993 and 1994. The coastal fronts near St. Paul and St. George Islands and the coastal domains encircled by the fronts featured specific taxonomic composition of planktonic algae, high abundance and production of phytoplankton, as well as large numbers of heterotrophic nanoplankton. The coastal fronts also were characterized by high values of total mesozooplankton biomass, high concentrations of Calanus marshallae, as well as relatively high abundances of Parasagitta setosa and Euphausiacea compared to surrounding shelf waters. We hypothesize that wind-induced erosion of a weak thermocline in the inner part of the coastal front as well as transfrontal water exchange in subthermocline layers result in nutrient enrichment of the euphotic layer in the coastal fronts and coastal domains in summer time. This leads to prolonged high primary production and high phytoplankton biomass. In this paper a new type of front—the shelf partition front located 45–55 km to the north-east off St. Paul Island—is described, which is assumed to be formed by the flux of oceanic domain waters onto the shelf. This front features a high abundance of phytoplankton and a high level of primary production compared to the adjacent middle shelf. Near the southwestern periphery of the front a mesozooplankton peak occurred, composed of C. marshallae, with biomass in the subthermocline layer, reaching values typical for the shelf break front and the highest for the area. High abundance of phyto- and zooplankton as well as heterotrophic nanoplankton and elevated primary production were most often observed in the area adjacent to the shelf break front at its oceanic side. The phyto- and mesozooplankton peaks here were formed by oceanic community species. The summer levels of phytoplankton numbers, biomass and primary production in the shelf break frontal area were similar to those reported for the outer and middle shelf during the spring bloom and the coastal domains and coastal fronts in summer. In the environment with a narrow shelf to the south of St. George Island, the mesozooplankton peak was observed at the inner side of the shelf break front as close as 20 km from the island shore and was comprised of a “mixed” community of shelf and oceanic species. The biomass in the peak reached the highest values for the Pribilof area at 2.5 g mean wet weight m⁻³ in the 0–100 m layer. Details of the taxonomic composition and the numbers and production of phytoplankton hint at the similarity of processes that affect the phytoplankton summer community in the coastal domains of the islands, at the coastal fronts, and at the oceanic side of the shelf break front. The middle front was the only one that had no effect on plankton composition or its quantitative characteristics in June and July. Location of a variety of frontal productive areas within 100 km of the Pribilof Islands creates favorable foraging habitat for higher trophic level organisms, including sea birds and marine mammals, populating the islands.