Copepod dynamics across warm and cold periods in the eastern Bering Sea: Implications for walleye pollock (Gadus chalcogrammus) and the Oscillating Control Hypothesis

Differences in zooplankton populations in relation to climate have been explored extensively on the southeastern Bering Sea shelf, specifically in relation to recruitment of the commercially important species walleye pollock (Gadus chalcogrammus). We addressed two research questions in this study: (i) Does the relative abundance of individual copepod species life history stages differ across warm and cold periods and (ii) Do estimated secondary production rates for copepods differ across warm and cold periods? For most copepod species, warmer conditions resulted in increased abundances in May, the opposite was observed in colder conditions. Abundances of smaller‐sized copepod species did not differ significantly between the warm and cold periods, whereas abundances of larger‐sized Calanus spp. increased during the cold period during July and September. Estimated secondary production rates in the warm period were highest in May for smaller‐sized copepods; production in the cold period was dominated by the larger‐sized Calanus spp. in July and September. We hypothesize that these observed patterns are a function of temperature‐driven changes in phenology combined with shifts in size‐based trophic relationships with primary producers. Based on this hypothesis, we present a conceptual model that builds upon the Oscillating Control Hypothesis to explain how variability in copepod production links to pollock variability. Specifically, fluctuations in spring sea‐ice drive regime‐dependent copepod production over the southeastern Bering Sea, but greatest impacts to upper trophic levels are driven by cascading July/September differences in copepod production.     

Multispecies biomass dynamics models reveal effects of ocean temperature on predation of juvenile pollock in the eastern Bering Sea

Walleye pollock (Gadus chalcogrammus) supports one of the largest commercial fisheries in the world. Juvenile pollock are important forage fish in the eastern Bering Sea (EBS) ecosystem, often representing the largest fraction in the diets of major Bering Sea piscivores. Large variability in the EBS pollock stock biomass in recent years has been attributed primarily to fluctuations in recruitment. It has been hypothesized that predation rates on forage fishes increase when the cold pool (a body of cold water < 2°C) is extensive and covers much of the middle continental shelf, which tends to concentrate larger predatory fishes in the outer shelf and slope regions. In contrast, young pollock appear to tolerate colder temperatures than older fish and can stay in the cold pool, thereby reducing predation. We used a multispecies modeling approach to examine the effects of the cold pool size on predation of juvenile pollock. We found that predation on age‐1 pollock by age‐3+ pollock decreased, and predation on age‐1 and age‐2 pollock by arrowtooth flounder increased with increasing bottom temperature, which was used as a proxy for the cold pool size. These results suggest that the cold pool creates spatial separation between juvenile pollock and arrowtooth flounder, but not between adult and juvenile pollock. The model developed in this study could be used to examine the effects of other covariates on interspecific interactions, help explain observed changes in fish communities, and understand implications of climate change on ecosystems and their productivity.     

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.     

Eastern Bering Sea pollock recruitment,abundance, distribution and approach to fishery management

Eastern Bering Sea pollock have two distinctly different stable spawning grounds—along the shelf and in the eastern and central Aleutian Islands between 400 and 500 m water columns. Pollock spawning behavior supports the hypothesis that the shelf and deepwater “basin” spawning pollock are completely independent reproductive stocks. Deepwater pollock inhabit the shelf and, once mature at age 5–6 years, migrate from the shelf onto the continental slope into the Zhemchug, Pribilof, and Bering canyons by the end of winter. Bering Sea pollock recruitment and year class abundance have high annual variability, but there are no clear relationships between pollock year class strength and water temperature, ice distribution or survival on early ontogenesis stages (eggs and larvae). Young-of-the-year fish survival varies dramatically during winter supporting the hypothesis that the Bering Sea pollock recruitment and strength of year class have high annual variability depending on young-of-the-year fish survival during winter. The annual change of physical oceanography condition, productivity and species composition of zooplankton community are associated with great differences in pollock seasonal migrations and distribution, reproduction, survival of recruits at early stages of development and finally with abundance of year classes and total biomass. Implementation of ecosystem-based fishery management most important for application of pollock research both of Russian national program and on base of International Agreements.     

Spatial and temporal patterns of walleye pollock (Theragra chalcogramma) spawning in the eastern Bering Sea inferred from egg and larval distributions

Walleye pollock Theragra chalcogramma (pollock hereafter) is a key ecological and economic species in the eastern Bering Sea, yet detailed synthesis of the spatial and temporal patterns of pollock ichthyoplankton in this important region is lacking. This knowledge gap is particularly severe considering that egg and larval distribution are essential to reconstructing spawning locations and early life stages drift pathways. We used 19 yr of ichthyoplankton collections to determine the spatial and temporal patterns of egg and larval distribution. Generalized additive models (GAMs) identified two primary temporal pulses of pollock eggs, the first occurring from 20 February to 31 March and the second from 20 April to 20 May; larvae showed similar, but slightly lagged, pulses. Based on generalized cross‐validation and information theory, a GAM model that allowed for different seasonal patterns in egg density within three unique areas outperformed a GAM that assumed a single fixed seasonal pattern across the entire eastern Bering Sea. This ‘area‐dependent’ GAM predicted the highest densities of eggs (i.e., potential spawning locations) in three major areas of the eastern Bering Sea: near Bogoslof Island (February–April), north of Unimak Island and the Alaska Peninsula (March–April), and around the Pribilof Islands (April–August). Unique temporal patterns of egg density were observed for each area, suggesting that pollock spawning may be more spatially and temporally complex than previously assumed. Moreover, this work provides a valuable baseline of pollock spawning to which future changes, such as those resulting from climate variability, may be compared.     

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.     

Use of juvenile salmon growth and temperature change indices to predict groundfish post age‐0 yr class strengths in the Gulf of Alaska and eastern Bering Sea

Juvenile marine growth (SW1) of salmon and a new temperature change (TC) index were evaluated as ecosystem indicators and predictors for the post age‐0 year class strength (YCS) of groundfish in the Gulf of Alaska (GOA) and eastern Bering Sea (EBS). Our hypothesis was that SW1, as measured on the scales of adult Pacific salmon (Oncorhynchus spp.), is a proxy for ocean productivity on the continental shelf, a rearing area for young salmon and groundfish. Less negative TC index values are the result of a cool late summer followed by a warm spring, conditions favorable for groundfish YCS. In the GOA, SW1 was a positive predictor of age‐1 pollock (Theragra chalcogramma), but not age‐2 sablefish (Anoplopoma fimbria) YCS, indicating that the growth of the Karluk River sockeye salmon that enter Shelikof Strait is a proxy for ocean conditions experienced by age‐0 pollock. Contrary to our hypotheses, the TC index was a negative predictor of GOA pollock YCS; and the SW1 a negative predictor of EBS pollock and cod YCS since the 1980s. Recent fisheries oceanography survey results provide insight into possible mechanisms to support the inverse SW1 and YCS relationship. For the EBS, the TC index was a significant positive predictor for pollock and cod YCS, supporting the hypothesis that a cool late summer followed by a warm spring maximizes the over‐wintering survival of pollock and cod (Gadus macrocephalus), especially since the 1980s. The TC and SW1 index showed value for the assessment of pollock and cod, but not sablefish.     

Spatial and temporal patterns in summer ichthyoplankton assemblages on the eastern Bering Sea shelf 1996–2007

Larval and early juvenile fishes were sampled from the eastern Bering Sea (EBS) shelf from 2001 to 2005, and in 2007. Data from these collections were used to examine spatial and temporal patterns in species assemblage structure and abundance. The years 2001–2005 were unusual because the EBS water temperature was ‘warm’ compared with the long‐term mean temperature. In contrast, 2007 was a ‘cold’ year. The abundance of the five most numerous taxa at 12 stations common to all years sampled (1996–2005, 2007) were significantly different among years. Larval and early juvenile stage Theragra chalcogramma (walleye pollock), a commercially important gadid, were by far the most abundant fish in all years. Bottom depth alone best explained assemblage structure in most years, but in others, bottom depth and water column temperature combined and percent sea‐ice coverage were most important. Abundance of T. chalcogramma larvae increases with water column temperature until 5°C and then becomes level. Higher abundances of Gadus macrocephalus (Pacific cod) larvae occur in years with the greatest percent sea‐ice cover as indicated by GAM analysis. Larvae of Lepidopsetta polyxystra (northern rock sole) increase in abundance with increasing maximum wind speed, but decrease at a later date during the last winter storm. The data are consistent with the hypothesis that oceanographic conditions, specifically water temperature and sea‐ice coverage, affect the spatial and temporal pattern of larval abundances. In general, ichthyoplankton species assemblages can be important early indicators of environmental change in the Bering Sea and potentially other subarctic seas as well.     

Ichthyophonus‐infected walleye pollock Theragra chalcogramma (Pallas) in the eastern Bering Sea: a potential reservoir of infections in the North Pacific

In 2003, the Alaska walleye pollock industry reported product quality issues attributed to an unspecified parasite in fish muscle. Using molecular and histological methods, we identified the parasite in Bering Sea pollock as Ichthyophonus. Infected pollock were identified throughout the study area, and prevalence was greater in adults than in juveniles. This study not only provides the first documented report of Ichthyophonus in any fish species captured in the Bering Sea, but also reveals that the parasite has been present in this region for nearly 20 years and is not a recent introduction. Sequence analysis of 18S rDNA from Ichthyophonus in pollock revealed that consensus sequences were identical to published parasite sequences from Pacific herring and Yukon River Chinook salmon. Results from this study suggest potential for Ichthyophonus exposures from infected pollock via two trophic pathways; feeding on whole fish as prey and scavenging on industry‐discharged offal. Considering the notable Ichthyophonus levels in pollock, the low host specificity of the parasite and the role of this host as a central prey item in the Bering Sea, pollock likely serve as a key Ichthyophonus reservoir for other susceptible hosts in the North Pacific.     

Walleye pollock: global overview

Walleye pollock is the second most extensively fished species in the world. The major fishing grounds include the Bering and Okhotsk Seas. Large-scale fishing started in the 1960s and continues to date with average annual landings over this 50-year period of 1.5 million tons. Yet over this period catches were characterized by considerable volatility. This volatility makes rational management of stock and planning of annual fishing activities difficult. The changes in annual catches correlate with the changes in the biomass of walleye pollock. Existing data suggest a close link between climate change in the northern Pacific and biomass, which allows quantitative estimates of future trends in the biomass, and consequently annual catch, of walleye pollock. Cooling of the northern Pacific is expected to increase the biomass in the Sea of Japan and decrease it in the Bering Sea and Sea of Okhotsk. The opposite is predicted to occur if the northern Pacific experiences warming.     

Ontogeny matters: Climate variability and effects on fish distribution in the eastern Bering Sea

Analyses of climate effects often ignore differences in life history for individual species. We analyzed a 34‐year time series of eastern Bering Sea fish surveys to evaluate changes in distribution by length and between cold and warm shelf‐wide average water temperatures for 20 species over inhabited depth, temperature, and location. All species showed evidence of ontogenetic migration. Differences in distribution between years with warm and years with cold shelf‐wide water temperatures varied among species and within species at different lengths. For species where shelf‐wide temperature effects were detected, the mid‐sized fish were most active in changing spatial distribution. For aquatic organisms ontogenetic migration occurs because life history stages have different environmental requirements. This study illustrates the need to consider species responses to climate change over different life history stages, and that studies on ecosystem responses should take ontogenetic differences into consideration when assessing impacts.     

Marine biogeochemical response to a rapid warming in the main stream of the Kuroshio in the western North Pacific

Impact of climate change on marine biogeochemical parameters and ecosystem is one of the important issues of our environment. Direct evidence of marine pelagic ecosystem changes is found with warming of sea water and sea‐level rise in the main stream of the Kuroshio in the East China Sea and the western North Pacific during these three decades based on the analysis of long‐term comprehensive hydrographic observations. In terms of annual mean, the warming rate of surface air temperature and sea surface temperature ranged from 0.15 to 0.21°C per decade in and around the main stream of the Kuroshio in the East China Sea, which exceed the global mean warming rate of 0.128 ± 0.026°C per decade during the period from 1956 to 2005 reported in IPCC 2007. One of the features in this rapid warming region is an increase of number of Pterosagitta draco, a cosmopolitan warm‐water zooplankton. Biogeochemical parameters, such as wet weight of zooplankton, plant pigment and nutrients concentration in the upper 200 m have been decreasing while dissolved oxygen content and seawater temperature have been increasing in the upper 200 m in the main stream of the Kuroshio in the East China Sea. These observed linear trends of the biogeochemical parameters would be foresights for temperate oceans in the future.     

Ecological processes influencing mortality of juvenile pink salmon (Oncorhynchus gorbuscha) in Prince William Sound,Alaska

Our collaborative work focused on understanding the system of mechanisms influencing the mortality of juvenile pink salmon (Oncorhynchus gorbuscha) in Prince William Sound, Alaska. Coordinated field studies, data analysis and numerical modelling projects were used to identify and explain the mechanisms and their roles in juvenile mortality. In particular, project studies addressed the identification of major fish and bird predators consuming juvenile salmon and the evaluation of three hypotheses linking these losses to (i) alternative prey for predators (prey‐switching hypothesis); (ii) salmon foraging behaviour (refuge‐dispersion hypothesis); and (iii) salmon size and growth (size‐refuge hypothesis). Two facultative planktivorous fishes, Pacific herring (Clupea pallasi) and walleye pollock (Theragra chalcogramma), probably consumed the most juvenile pink salmon each year, although other gadids were also important. Our prey‐switching hypothesis was supported by data indicating that herring and pollock switched to alternative nekton prey, including juvenile salmon, when the biomass of large copepods declined below about 0.2 g m^?3. Model simulations were consistent with these findings, but simulations suggested that a June pteropod bloom also sheltered juvenile salmon from predation. Our refuge‐dispersion hypothesis was supported by data indicating a five‐fold increase in predation losses of juvenile salmon when salmon dispersed from nearshore habitats as the biomass of large copepods declined. Our size‐refuge hypothesis was supported by data indicating that size‐ and growth‐dependent vulnerabilities of salmon to predators were a function of predator and prey sizes and the timing of predation events. Our model simulations offered support for the efficacy of representing ecological processes affecting juvenile fishes as systems of coupled evolution equations representing both spatial distribution and physiological status. Simulations wherein model dimensionality was limited through construction of composite trophic groups reproduced the dominant patterns in salmon survival data. In our study, these composite trophic groups were six key zooplankton taxonomic groups, two categories of adult pelagic fishes, and from six to 12 groups for tagged hatchery‐reared juvenile salmon. Model simulations also suggested the importance of salmon density and predator size as important factors modifying the predation process.     

Springtime ichthyoplankton of the slope region off the north-eastern United States of America: larval assemblages, relation to hydrography and implications for larval transport

Larval transport in the slope region off north‐eastern North America influences recruitment to juvenile habitats for a variety of fishes that inhabit the continental shelf. In this study, collections of larval fishes were made during springtime over the continental slope to provide insights into larval distributions and transport. Ichthyoplankton composition and distribution mirrored the physical complexity of the region. Three larval fish assemblages were defined, each with different water mass distributions. A Gulf Stream assemblage was found predominantly in the Gulf Stream and associated with filaments of discharged Gulf Stream water in the Slope Sea. Larvae of this assemblage originated from oceanic and shelf regions south of Cape Hatteras. Several members of this assemblage utilize habitats in the Middle Atlantic Bight (MAB) as juveniles (Pomatomus saltatrix, Peprilus triacanthus) and other members of the assemblage may share this life cycle (Mugil curema, Sphyraena borealis, Urophycis regia). A Slope Sea assemblage was found in all water masses, and was composed of epi‐ and mesopelagic fish larvae, as well as larvae of benthic shelf/slope residents. Larvae of one member of this assemblage (U. tenuis) are spawned in the Slope Sea but cross the shelf‐slope front and use nearshore habitats for juvenile nurseries. A MAB shelf assemblage was found in MAB shelf water and was composed of larvae that were spawned on the shelf. Some of these species may cross into the Slope Sea before returning to MAB shelf habitats (e.g. Enchelyopus cimbrius, Glyptocephalus cynoglossus). Previous studies have examined the effect of warm‐core rings on larval distributions, but this study identifies the importance of smaller‐scale features of the MAB shelf/slope front and of filaments associated with Gulf Stream meanders. In combination with these advective processes, the dynamic nature of larval distributions in the Slope Sea appears to be influenced, to varying degrees, by both vertical and horizontal behaviour of larvae and pelagic juveniles themselves.     

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.     

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.     

A synthesis of biological and physical processes affecting the feeding environment of larval walleye pollock (Theragra chalcogramma) in the eastern Bering Sea

Biological and physical phenomena that affect conditions for larval survival and eventual recruitment differ in the oceanic and shelf regions. In the oceanic region, eddies are a common feature. While their genesis is not well known, eddies have unique biophysical characteristics and occur with such regularity that they likely affect larval survival. High concentrations of larval pollock often are associated with eddies. Some eddies are transported onto the shelf, thereby providing larvae to the Outer Shelf Domain. Advection, rather than local production, dominated the observed springtime increase in chlorophyll (often a correlate of larval food) in the oceanic region. Over two-thirds of the south-eastern shelf, eddies are absent and other phenomena are important. Sea ice is a feature of the shelf region: its interannual variability (time of arrival, persistence, and areal extent) affects developmental rate of larvae, timing of the phytoplankton bloom (and potentially the match/mismatch of larvae and prey), and abundance and distribution of juvenile pollock. In the oceanic region, interannual variation in food for first-feeding pollock larvae is determined by advection; in the shelf region, it is the coupled dynamics of the atmosphere–ice–ocean system.     

Copepods and salmon: characterizing the spatial distribution of juvenile salmon along the Washington and Oregon coast,USA

Yearling Chinook (Oncorhynchus tshawytscha) and coho salmon (Oncorhynchus kisutch) were sampled concurrently with physical variables (temperature, salinity, depth) and biological variables (chlorophyll a concentration and copepod abundance) along the Washington and Oregon coast in June 1998–2008. Copepod species were divided into four different groups based on their water‐type affinities: cold neritic, subarctic oceanic, warm neritic, and warm oceanic. Generalized linear mixed models were used to quantify the relationship between the abundance of these four different copepod groups and the abundance of juvenile salmon. The relationships between juvenile salmon and different copepod groups were further validated using regression analysis of annual mean juvenile salmon abundance versus the mean abundance of the copepod groups. Yearling Chinook salmon abundance was negatively correlated with warm oceanic copepods, warm neritic copepods, and bottom depth, and positively correlated with cold neritic copepods, subarctic copepods, and chlorophyll a concentration. The selected habitat variables explained 67% of the variation in yearling Chinook abundance. Yearling coho salmon abundance was negatively correlated with warm oceanic copepods, warm neritic copepods, and bottom depth, and positively correlated with temperature. The selected habitat variables explained 40% of the variation in yearling coho abundance. Results suggest that copepod communities can be used to characterize spatio‐temporal patterns of abundance of juvenile salmon, i.e., large‐scale interannual variations in ocean conditions (warm versus cold years) and inshore‐offshore (cross‐shelf) gradients in the abundance of juvenile salmon can be characterized by differences in the abundance of copepod species with various water mass affinities.     

A review: Walleye pollock in the North Pacific–how much difference do they really make?

Populations of several species of marine birds and mammals in the Bering Sea and Gulf of Alaska have been declining since the mid-1970s, with numbers of one, the Steller sea lion (Eumetopias jubatus) , so depressed it was listed as threatened under the Endangered Species Act in spring 1990. All of the declining populations depend to an important extent on walleye pollock (Theragra chakogramma) for food, although they eat numerous other species as well. In contrast, certain animals that compete with pollock for common prey have been increasing in abundance. All of these changes could be related through food web connections mediated by pollock. Pollock is also important to people–it presently supports the largest single-species commercial fishery in the world, in large part because of its great biomass, which has averaged about 15 × 10⁶t in the Bering Sea over the past 15 years. Pollock consume an inordinate proportion of the pelagic production in the Bering Sea, which further supports the conclusion that it is a key species in the ecosystem. However, there are conflicting hypotheses about the importance of the roles played by pollock as predator and prey, and about the effect that changes in pollock abundance might have on biomass yield at higher trophic levels.     

Spatio‐temporal analyses of marine predator diets from data‐rich and data‐limited systems

Accounting for variation in prey mortality and predator metabolic potential arising from spatial variation in consumption is an important task in ecology and resource management. However, there is no statistical method for processing stomach content data that accounts for fine‐scale spatio‐temporal structure while expanding individual stomach samples to population‐level estimates of predation. Therefore, we developed an approach that fits a spatio‐temporal model to both prey‐biomass‐per‐predator‐biomass data (i.e. the ratio of prey biomass in stomachs to predator weight) and predator biomass survey data, to predict “predator‐expanded‐stomach‐contents” (PESCs). PESC estimates can be used to visualize either the annual landscape of PESCs (spatio‐temporal variation), or can be aggregated across space to calculate annual variation in diet proportions (variation among prey items and among years). We demonstrated our approach in two contrasting scenarios: a data‐rich situation involving eastern Bering Sea (EBS) large‐size walleye pollock (Gadus chalcogrammus, Gadidae) for 1992–2015; and a data‐limited situation involving West Florida Shelf red grouper (Epinephelus morio, Epinephelidae) for 2011–2015. Large walleye pollock PESC was predicted to be higher in very warm years on the Middle Shelf of the EBS, where food is abundant. Red grouper PESC was variable in north‐western Florida waters, presumably due to spatio‐temporal variation in harmful algal bloom severity. Our approach can be employed to parameterize or validate diverse ecosystem models, and can serve to address many fundamental ecological questions, such as providing an improved understanding of how climate‐driven changes in spatial overlap between predator and prey distributions might influence predation pressure.