Evidence for a substantial increase in gelatinous zooplankton in the Bering Sea, with possible links to climate change

We examined quantitative catches of large medusae from summer bottom trawl surveys that sampled virtually the same grid station on the eastern Bering Sea shelf using the same methodology every year from 1979 to 1997. This series shows a gradual increase in biomass of medusae from 1979 to 1989, followed by a dramatic increase in the 1990s. The median biomass increased tenfold between the 1982–1989 and 1990–1997 periods. Most of this biomass was found within the Middle Shelf Domain (50 <  z  < 100 m). The greatest rate of increase occurred in the north-west portion of this domain. Whether this dramatic increase in biomass of gelatinous zooplankton has resulted from some anthropogenic perturbation of the Bering Sea environment or is a manifestation of natural ecosystem variability is unclear. However, several large-scale winter/spring atmospheric and oceanographic variables in the Bering Sea exhibited concomitant changes beginning around 1990, indicating that a possible regime change occurred at this time.     

North Pacific Atmospheric and SST Anomalies in 1997: Links to ENSO?

In the summer of 1997, positive sea surface temperature anomalies (SSTA) extended across the Gulf of Alaska (GOA) and into the eastern Bering Sea (EBS). The SSTA in the EBS are at least in part due to atmospheric causes. Anomalously high 925 mb temperatures and 700 mb geopotential heights and low 925 mb relative humidities, and hence decreased low cloud cover, occurred over the region during April to August. This resulted in enhanced warming of the GOA and EBS owing to increased insolation. The anomalous solar heating was particularly great in the EBS from mid-May to mid-July. The pattern of positive 700 mb height anomalies for April to August 1997 is similar to its counterpart formed by compositing the April to August anomalies that occurred during previous El Niños. The positive equatorial SSTA for 1997 was one of the strongest on record for summer months. The existence of an equatorial/high-latitude connection and the strength of the summer equatorial SSTA in 1997 suggest an El Niño/Southern Oscillation (ENSO) influence in the GOA and EBS. The warming in the Bering Sea and North Pacific during summer 1997 appears to be due in part to the confluence of three meteorological factors which favoured clear skies. There was not only an El Niño influence, but also a decadal trend toward higher 700 mb geopotential heights and a particularly strong blocking ridge weather pattern over the Gulf of Alaska in May.     

The importance of episodic weather events to the ecosystem of the Bering Sea shelf

Climate variability on decadal time scales is generally recognized to influence high‐latitude marine populations. Our recent work in studying air–sea interactions in the Bering Sea suggests that interannual to decadal climate variability is important through its modulation of the frequencies and magnitudes of weather events on intraseasonal time scales. We hypothesize that it is these weather events that directly impact the marine ecosystem of the Bering Sea shelf. The linkages between the event‐scale weather and the ecosystem are illustrated with three examples: walleye pollock (Theragra chalcogramma), Tanner crabs (Chionoecetes bairdi), and coccolithophorid phytoplankton (Emiliania huxleyi). We hypothesize that the strong recruitment of walleye pollock that occurred in 1978, 1982, and 1996 can be attributed in part due to the seasonably strong storms that occurred in the early summer of those years. These storms caused greater than normal mixing of nutrients into the euphotic zone which presumably led to sustained primary productivity after the spring bloom and, possibly, enhanced prey concentrations for pollock larvae and their competitors. Recruitment of Tanner crab was particularly strong for the 1981 and 1984 year‐classes. These years had periods of prominent east wind anomalies along the Alaska Peninsula during the previous winter. Such winds promote flow through Unimak Pass, and hence an enhanced flux of nutrient‐rich water onto the shelf. This mechanism may have ultimately resulted in favorable feeding conditions for Tanner crab larvae. Finally, an unprecedented coccolithophorid bloom occurred over the Bering Sea shelf in the summer of 1997. This summer featured lighter winds and greater insolation than usual after a spring that included a very strong May storm. This combination brought about a warm, nutrient‐poor upper mixed layer by mid‐summer. This provided a competitive advantage for coccolithophorid phytoplankton in 1997 and to a lesser extent in 1998. Unusually high concentrations of coccolithophores persisted for the following two years although physical environmental conditions did not remain favorable. While slow variations in the overall aspects of the physical environment may be important for setting the stage, we propose that the significant multi‐year adjustments in the marine ecosystem of the Bering Sea shelf are more directly caused by major air–sea interaction events on intraseasonal time scales.     

Anomalous conditions in the south-eastern Bering Sea, 1997: nutrients, phytoplankton and zooplankton

Anomalies in the regional weather over the south-eastern 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.     

Mass mortality of short-tailed shearwaters in the south-eastern Bering Sea during summer 1997

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.     

On the temporal variability of the physical environment over the south-eastern Bering Sea

During 1997 and 1998, unusual physical conditions occurred in the Bering Sea: strong May storms and calm conditions in July; record high sea surface temperature; a shallow wind mixed layer; a fresher-than-normal water column; and abnormal cross-shelf currents. Accompanying these conditions were changes in the dominant phytoplankton, a die-off of seabirds, increased sightings of large whales and diminished returns of salmon. Changes to the physical environment during 1997 and 1998 are placed in context of historical meteorological and oceanographic data sets. Although 1997 had the warmest sea surface temperature ever observed on the south-east Bering Sea shelf, the heat content of the water column was cooler than average. In contrast, during 1998, the sea surface temperature was cooler than in 1997 but the water column had significantly higher heat content. During recent years, the water column has freshened over the middle shelf because of increased sea ice and reduction of on-shelf transport of the saline, high-nutrient water from the slope. The timing of the spring bloom is directly related to the presence of ice. When ice is advected over the south-east shelf during March/April an early, sharp phytoplankton bloom occurs. The absence of ice during this critical time is associated with a May/June bloom.     

Observations on larval fish transport and retention on the Scotian Shelf in relation to geostrophic circulation

We draw inferences about the dynamic processes responsible for the dispersal of ichthyoplankton on small marine banks using physical and biological data derived from static point-estimates of water-mass characteristics and ichthyoplankton collected concomitantly on the central Scotian Shelf. Where the density field evolves slowly and ageostrophic forcing is weak, the near-surface geostrophic flow can be derived from hydrographic data using the dynamic height method modified for shallow seas. We assess our interpretations of larval distributions using simple particle tracking. The hydrography of the Scotian Shelf during November of 1997 was typical of late autumn, when density is determined by surface variation in salinity. Surface isopycnals generally paralleled isobaths, and there was no evidence of strong surface fronts. Sizes of larvae of pelagic origin (e.g. cod and hake) on Western Bank (sole spawning source) increased and became skewed towards larger animals (cod, 3–10 mm; hake, 5–15 mm) in water-mass (Temperature and Salinity) space along isopycnals, consistent with gradual mixing and limited transport in the geostrophic flow (i.e. retention). Conversely, larvae of benthic origin (e.g. herring, 6–25 mm) were distributed across water-mass space, consistent with multiple origins and substantial transport. Our results indicate that dispersal from small, low-energy marine banks results from the interaction of spawning location, geostrophic currents and bathymetric steering, and requires neither convergence nor larval behaviour.     

Pacific herring, Clupea pallasi, recruitment in the Bering Sea and north-east Pacific Ocean, II: relationships to environmental variables and implications for forecasting

Previous studies have shown that Pacific herring populations in the Bering Sea and north-east Pacific Ocean can be grouped based on similar recruitment time series. The scale of these groups suggests large-scale influence on recruitment fluctuations from the environment. Recruitment time series from 14 populations were analysed to determine links to various environmental variables and to develop recruitment forecasting models using a Ricker-type environmentally dependent spawner–recruit model. The environmental variables used for this investigation included monthly time series of the following: southern oscillation index, North Pacific pressure index, sea surface temperatures, air temperatures, coastal upwelling indices, Bering Sea wind, Bering Sea ice cover, and Bering Sea bottom temperatures. Exploratory correlation analysis was used for focusing the time period examined for each environmental variable. Candidate models for forecasting herring recruitment were selected by the ordinary and recent cross-validation prediction errors. Results indicated that forecasting models using air and sea surface temperature data lagged to the year of spawning generally produced the best forecasting models. Multiple environmental variables showed marked improvements in prediction over single-environmental-variable models.     

Early- to late-summer population growth and prey consumption by age-0 pollock (Theragra chalcogramma), in two years of contrasting pollock abundance near the Pribilof Islands, Bering Sea

Acoustic survey data were used to estimate the abundance and distribution of age-0 walleye pollock and zooplankton near the Pribilof Islands, Bering Sea, nursery area at two time periods in two consecutive years: the beginning of August, and mid-September, of 1996 and 1997. The 1996 pollock year class ultimately produced a large adult cohort in the eastern Bering Sea, while the 1997 year class produced a below-average adult cohort. Acoustic densities of age-0 pollock were significantly lower in August – and declined more strongly from August to September – in 1997 than in 1996, indicating that the trend to adult cohort strength was already set by August. Diet composition analyses revealed that age-0 pollock ate a much higher proportion of euphausiids in 1997 than in 1996, despite lower acoustic abundance of euphausiids in 1997. We infer that in 1996, age-0 pollock experienced greater feeding success by August, with high concentrations of copepods available for smaller fish to consume, and high concentrations of euphausiids available for larger individuals. In 1997, age-0 pollock had lower body condition in August and may have been limited by the availability of small (<2 mm) copepods. Bioenergetic modeling of prey consumption did not indicate a likelihood that age-0 pollock would begin to deplete euphausiids until late August in 1996, and not at all between August and mid-September in 1997.     

Distribution and drift pathways of Greenland halibut (Reinhardtius hippoglossoides) during early life stages in the eastern Bering Sea and Aleutian Islands

We describe the spatial distribution and dispersal pathways of Greenland halibut (Reinhardtius hippoglossoides) early life stages based on historical field data from the eastern Bering Sea and adjacent water along the eastern Aleutian Islands. Our results indicate that Greenland halibut from preflexion larvae to newly settled juveniles have a long pelagic duration and are subject to extended drift pathways. Hatching may occur in deep water, below 530 m, and larvae rise in the water column as they grow. Flexion/postflexion larvae are mostly found around the Pribilof Islands over the middle shelf (50–100‐m isobaths) in July, and settling occurs during late summer on the middle shelf near St. Matthew Island. However, given that age‐1 individuals were primarily found on the outer shelf, it appears that Greenland halibut actively move to deeper water with age (or size). The mechanisms of slope–shelf connectivity in preflexion larvae may be related to the Bering Slope Current in the vicinity of both Bering and Pribilof Canyons. This study shows that Greenland halibut early life stages have extensive horizontal ontogenetic migrations in the Bering Sea, and utilize a range of geographic areas over the basin and slope along the Aleutian Islands and in the eastern Bering Sea. Based on these results, it is hypothesized that settlement success and recruitment of Greenland halibut may be influenced by variability in currents and flows of the Bering Sea slope and shelf during their transport.