Temperature-dependent stock-recruitment model for walleye pollock (Theragra chalcogramma) around northern Japan

Changes in fish year‐class strength have been attributed to year‐to‐year variability in environmental conditions and spawning stock biomass (SSB). In particular, sea temperature has been shown to be linked to fish recruitment. In the present study, I examined the relationship between sea surface temperature (SST), SSB and recruitment for two stocks of walleye pollock (Theragra chalcogramma) around northern Japan [Japanese Pacific stock (JPS) and northern Japan Sea stock (JSS)] using a temperature‐dependent stock‐recruitment model (TDSRM). The recruitment fluctuation of JPS was successfully reproduced by the TDSRM with February and April SSTs, and February SST was a better environmental predictor than April SST. In addition, the JPS recruitment was positively related to February SST and negatively to April SST. The JSS recruitment modeled by the TDSRM incorporating February SST was also consistent with the observation, whereas the relationship between recruitment and February SST was negative, that is the opposite trend to JPS. These findings suggest that SST in February is important as a predictor of recruitment for both stocks, and that higher and lower SSTs in February act favorably on the recruitment of JPS and JSS respectively. Furthermore, Ricker‐type TDSRM was not selected for either of the stocks, suggesting that the strong density‐dependent effect as in the Ricker model does not exist for JPS and JSS. I formulate hypotheses to explain the links between SST and recruitment, and note that these relationships should be considered in any future attempts to understand the recruitment dynamics of JPS and JSS.     

Abiotic and biotic factors affecting recruitment variability of walleye pollock (Theragra chalcogramma) off the Pacific coast of Hokkaido,Japan

Abiotic and biotic factors affecting the recruitment variability of the Japanese Pacific stock (JPS) of walleye pollock (Theragra chalcogramma) were examined using a bivariate regression and multivariate combined model. Of the abiotic variables around Funka Bay (spawning ground), February sea surface temperature (SST) and wind direction index showed significant bivariate relationships with recruitment. February SST was positively related to recruitment, suggesting that warmer water temperature in February favors JPS recruitment. On the other hand, the relationship between February wind direction index and recruitment predicts high JPS recruitment under predominant northwest winds in February. For the biotic variables in the Doto area (nursery ground), significant and negative bivariate relationships with recruitment were observed for catch per unit effort of Kamchatka flounder (Atheresthes evermanni), Pacific cod (Gadus macrocephalus), and walleye pollock, implying an important impact of predation by these groundfishes on JPS recruitment. The overall model incorporating these abiotic and biotic factors successfully reproduced the variability in JPS recruitment. Temperature and wind conditions around the spawning ground along with predator condition in the nursery ground appear to play a dominant role in the recruitment dynamics of JPS. Based on these results and prior knowledge, we propose a new hypothesis to explain the processes controlling JPS recruitment.     

Causes of walleye pollock (Theragra chalcogramma) recruitment decline in the northern Sea of Japan: implications for stock management

Recruitment of the northern Japan Sea stock (JSS) of walleye pollock has been decreasing since around 1990. In this study, I analyzed the factors causing this decrease in recruitment by investigating the relationship between recruitment, spawning stock biomass (SSB) and environmental factors using a generalized additive model (GAM). GAM fit to the data showed the importance of SSB, sea surface temperature (SST), ocean current strength (Tsushima Warm Current) and wind intensity (Asian monsoon) in determining the recruitment. Of these, the relationship between SSB and recruitment was positive and not negatively density‐dependent. On the other hand, the recruitment was negatively related to SST and ocean current strength, and a dome‐shaped relationship was observed between wind intensity and recruitment. Since around 1990, the values of SST and ocean current strength have mostly been high and that of wind intensity mostly low. In addition, SSB has been decreasing since the late 1990s. It is likely that the recruitment decline of JSS after approximately 1990 has been caused by warm water temperature, strong Tsushima Warm Current and weak Asian monsoon, and that the recent decrease in SSB has amplified this recruitment decline. According to the model’s estimation, a recruitment recovery due to environmental improvement will be highly restricted as long as SSB remains at its current low level. Significant recovery of SSB is urgently needed for JSS.     

Stock assessment and management for walleye pollock in Japan

We review the stock assessment strategies and management procedures for walleye pollock Theragra chalcogramma in Japan. In Japan, walleye pollock is classified into 4 stocks. Because biological data, fishing conditions, etc. are different for each stock, the stocks are assessed by different methods. Harvest strategies aiming at stock recovery are proposed for the Northern Japan Sea stock and the Nemuro Strait stock, which are currently in poor condition. For the Japanese Pacific stock and the Southern Okhotsk Sea stock, which are in good condition, harvest strategies for current fishery operations are proposed. In Japan, fisheries co-management has traditionally been carried out, and in recent years a total catch limitation system called the total allowable catch, a resource recovery plan, and a resource management plan have also been implemented. Although a plan is devised that accounts for the stock conditions of walleye pollock, it is also necessary to consider socioeconomic factors, ecosystem factors, and so on. However, we consider that the main focus of stock management for walleye pollock will still be maintaining fishing pressure at an appropriate level, which includes regulating fish size and price during the fishing season.     

Geographical variations in infection by larval Anisakis simplex and Contracaecum osculatum (Nematoda, Anisakidae) in walleye pollock Theragra chalcogramma stocks off Hokkaido, Japan

ABSTRACT: Stocks of walleye pollock Theragra chalcogramma collected from: (i) the Sea of Japan (off Rebun Island and Kumaishi); (ii) the Pacific coast (off Shikabe and eastern Hokkaido); and (iii) Nemuro Strait off Hokkaido, northern Japan, were examined for anisakid nematodes during December 1999 to February 2000, and the prevalence and abundance of Anisakis simplex and Contracaecum osculatum larvae were compared among the various sampling sites for fish of the same size and age. Anisakis simplex was generally more abundant than C. osculatum . Infection by A. simplex varied between the aforementioned stocks of walleye pollock as well as within stocks, whereby fish from off Rebun Island and Nemuro Strait were infected the most, followed by those from off the Pacific coast and Kumaishi. Infection by C. osculatum differed between the host stocks, and C. osculatum was the most abundant among the fish from Nemuro Strait. The infection variations seemed to be due to differences in host growth rate, host feeding habit, and the distribution of marine mammal final hosts. The results indicate that these two larval nematodes are useful biological indicators for the population study of walleye pollock in Japanese waters.     

Linking Northeast Pacific recruitment synchrony to environmental variability

We investigated the hypothesis that synchronous recruitment is due to a shared susceptibility to environmental processes using stock–recruitment residuals for 52 marine fish stocks within three Northeast Pacific large marine ecosystems: the Eastern Bering Sea and Aleutian Islands, Gulf of Alaska, and California Current. There was moderate coherence in exceptionally strong and weak year‐classes and correlations across stocks. Based on evidence of synchrony from these analyses, we used Bayesian hierarchical models to relate recruitment to environmental covariates for groups of stocks that may be similarly influenced by environmental processes based on their life histories. There were consistent relationships among stocks to the covariates, especially within the Gulf of Alaska and California Current. The best Gulf of Alaska model included Northeast Pacific sea surface height as a predictor of recruitment, and was particularly strong for stocks dependent on cross‐shelf transport during the larval phase for recruitment. In the California Current the best‐fit model included San Francisco coastal sea level height as a predictor, with higher recruitment for many stocks corresponding to anomalously high sea level the year before spawning and low sea level the year of spawning. The best Eastern Bering Sea and Aleutian Islands model included several environmental variables as covariates and there was some consistent response across stocks to these variables. Future research may be able to utilize these across‐stock environmental influences, in conjunction with an understanding of ecological processes important across early life history stages, to improve identification of environmental drivers of recruitment.     

Integrating seabird dietary and groundfish stock assessment data: Can puffins predict pollock spawning stock biomass in the North Pacific?

Information on the annual variability in abundance and growth of juvenile groundfish can be useful for predicting fisheries stocks, but is often poorly known owing to difficulties in sampling fish in their first year of life. In the Western Gulf of Alaska (WGoA) and Eastern Bering Sea (EBS) ecosystems, three species of puffin (tufted and horned puffin, Fratercula cirrhata, Fratercula corniculata, and rhinoceros auklet, Cerorhinca monocerata, Alcidae), regularly prey upon (i.e., “sample”) age-0 groundfish, including walleye pollock (Gadus chalcogramma, Gadidae) and Pacific cod (Gadus microcephalus, Gadidae). Here, we test the hypothesis that integrating puffin dietary data with walleye pollock stock assessment data provides information useful for fisheries management, including indices of interannual variation in age-0 abundance and growth. To test this hypothesis, we conducted cross-correlation and regression analyses of puffin-based indices and spawning stock biomass (SSB) for the WGoA and EBS walleye pollock stocks. For the WGoA, SSB leads the abundance of age-0 fish in the puffin diet, indicating that puffins sample the downstream production of the WGoA spawning stock. By contrast, the abundance and growth of age-0 fish sampled by puffins lead SSB for the EBS stock by 1–3 years, indicating that the puffin diet proxies incoming year class strength for this stock. Our study indicates connectivity between the WGoA and EBS walleye pollock stocks. Integration of non-traditional data sources, such as seabird diet data, with stock assessment data appears useful to inform information gaps important for managing US fisheries in the North Pacific.     

Effects of juvenile size at release and early marine growth on adult return rates for Hokkaido chum salmon (Oncorhynchus keta) in relation to sea surface temperature

Using path analyses, we investigated relationships between size at release from hatcheries, the early marine growth of juveniles, and adult return rates for chum salmon from five river stocks of Hokkaido, Japan, in relation to sea surface temperature during ocean residence. Marine growth was estimated using scales collected from 11 760 adults of age 0.3 (1980–2004). The growth and survival of each stock appeared to have a different suite of regulatory processes. Interannual variability in return rates was mainly regulated by size at release in two stocks from the Sea of Okhotsk. A similar relationship was found in one stock from the Sea of Japan, but growth during coastal residency also affected their return rates. In two stocks from the Pacific coast of Hokkaido, variability in return rates was not related to size at release or to the coastal growth of juveniles, but with offshore growth in the Sea of Okhotsk, the nursery area for juveniles after leaving Japanese coastal waters. Whereas coastal growth tended to be negatively correlated with size at release in some stocks, offshore growth was positively associated with the August–November sea surface temperature in all stocks. This study confirmed that mortality of juvenile salmon occurred in two phases, during the coastal residency and the late period of the growing season, but the relative importance of both phases varied by stock and region, which probably regulated year‐class strength of Hokkaido chum salmon.     

Straddling the line: cooperative and non-cooperative strategies for management of Bering Sea pollock

The eastern Bering Sea fishery for pollock, Theragra chalcogramma, yields a first wholesale value over $1 billion; it is the premier US fishery. While there is general agreement that this fishery is managed under principles that foster sustainability, the stock is not wholly contained within the US Exclusive Economic Zone. Management of straddling stocks can be highly contentious, particularly when, as is the case for pollock, the spatial distribution varies considerably. When the center of pollock abundance shifts to the northwest, an increased portion of the stock is exposed to harvest by vessels operating in the Russian Federation Exclusive Economic Zone. The lack of coordination in the management of this transboundary stock presents a risk that is not reflected in current management strategies. We use a multiple product/multiple market bioeconomic model to characterize optimal cooperative and non-cooperative harvest management strategies from the perspective of US and Russian pollock fisheries under environmentally induced changes in pollock abundance and the distribution of that abundance.     

Relationship between sea-surface temperature and catch fluctuations in the Pacific stock of walleye pollock in Japan

ABSTRACT:   This paper investigates the relationship between sea-surface temperature (SST) and catch fluctuations in the Pacific stock of walleye pollock Theragra chalcogramma in Japan. Incorporating time lags between years of birth and harvest, the correlation coefficients between the catch and SST in two regions off the east coast of Hokkaido were calculated. The catch in year t had a high negative correlation with the SST during January–April and November–December of the years t- 2 and t- 3 in the spawning area. These results coincided well with the correlation observed in the northern 'Sea of Japan' stock. Both analyses suggested that the long-term catch fluctuations of the two stocks could be explained by the same mechanism, that is, the fluctuations would be explained by the SST in their spawning area during the spawning season using 2–3 or 3–5 years time lags, which corresponded to the dominant age of the catch within these two stocks.     

Long-term stock assessment and growth changes of the Japanese sardine (Sardinops melanostictus) in the Sea of Japan and East China Sea from 1953 to 2006

We estimated the stock size of Japanese sardine ( Sardinops melanostictus ) in the Sea of Japan and East China Sea since 1953 using cohort analysis based on the changes of growth patterns. Growth of Japanese sardine, estimated by using annual rings on archived scales since 1961, showed that body lengths were extremely stunted in the 1980–1987 year-classes. The body length at age 3 from February to April in the 1980–1987 year-classes, a period when the stock size exceeded 4 million tons, was 180.0 ± 2.6 mm (mean ± SD), and in the other year-classes was 195.1 ± 7.6 mm. The body length at age 3 and wet weight of zooplankton in August in the offshore area of the Sea of Japan had a significantly positive correlation. We assumed three scenarios for maturation ratios, and estimated Ricker's spawner–recruitment relationships. We analyzed the correlations between logarithmic recruitment residuals (LNRR) and environmental factors in winter, represented by the North Pacific index (NPI), Aleutian low pressure index (ALPI), Pacific decadal oscillation (PDO), monsoon index (MOI), Arctic oscillation (AO) and Southern oscillation index (SOI). Significant correlations were observed between MOI and LNRR and between AO and LNRR. A combination of strong MOI and weak AO would increase the biomass of phytoplankton and zooplankton and subsequently increase the recruitment of Japanese sardine.     

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.     

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.     

Life-history traits of walleye pollock, Theragra chalcogramma, in the northeastern Japan Sea during early to mid 1990s

I examined the age, growth, maturity, mortality, and body condition of walleye pollock, Theragra chalcogramma, in the northeastern Japan Sea (northern Japan Sea population) and evaluated their resilience to exploitation. Walleye pollock were collected in pre-spawning (October 1991-1995) and post-spawning (April 1990-1996) seasons. Estimated ages ranged from 3 to 18 years for both sexes. A von Bertalanffy growth model showed that females had longer asymptotic fork length (460 mm) than males (425 mm). Fifty percent of females and males were mature at 348 mm (4.6 years) and 322 mm (3.9 years), respectively. The instantaneous natural mortality rate was estimated to be 0.22. These life-history traits in the northern Japan Sea population were compared to those in the Bering Sea, the Gulf of Alaska, and the Japan Pacific populations. As a result, female walleye pollock in this population matured at small body sizes, grew rapidly toward small maximum sizes, and had short reproductive lifespans with low size-specific fecundity and poor body condition. Low prey availability and habitat temperatures are considered as a possible mechanism for the small maximum sizes in this population. The potential rate of population increase of both the northern Japan Sea population and other pollock populations tended to be lower than other exploited populations of non-viviparous marine fishes, suggesting potentially lower resilience to exploitation in this population and walleye pollock populations in general.     

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.     

Evaluation of stock management procedures for walleye pollock in northern waters of the Sea of Japan using a simulation

To evaluate walleye pollock stock management procedures in the northern waters of the Sea of Japan, 30-year population dynamics, including uncertainties, were forecast. Errors in current stock size estimation, variability in future recruitment and changes in future fishing mortalities were incorporated. Results of virtual population analysis (VPA) from resampled catch-at-age data with bootstrap methods was used as the current stock size estimation with uncertainty. Performances of each scenario were evaluated using conservation, utilization, stability and reliability factors. Twenty-two management scenarios and continuing the current fishing mortality were evaluated. Scenarios with minor regulation changes and continuing the current fishing mortality showed poor stock conservation performances. Scenarios with minor regulation changes produced good short-term but poor long-term utilization. Stabilities were poor in continuing the current fishing mortality and fishing ban scenarios. Reliability in all scenarios after 30 years was smaller than in continuing the current fishing mortality; however, small differences among scenarios were observed. The simulation results indicated that multilateral assessment is needed to evaluate the management candidates. Uncertainty caused by recruitment variability mostly affected future population dynamics. The role of simulations in the production of effective scientific advice is discussed.     

Chum salmon (Oncorhynchus keta) growth and temperature indices as indicators of the year–class strength of age‐1 walleye pollock (Gadus chalcogrammus) in the eastern Bering Sea

Ecosystem‐based fisheries management requires the development of physical and biological time series that index ocean productivity for stock assessment and recruitment forecasts for commercially important species. As recruitment in marine fish is related to ocean condition, we developed proxies for ocean conditions based on sea surface temperature (SST) and biometric measurements of chum salmon (Oncorhynchus keta) captured in the walleye pollock (Gadus chalcogrammus) fishery in the eastern Bering Sea in three periods (July 16–30, September 1–15 and September 16–30). The main purpose of this paper was to evaluate Pacific salmon (Oncorhynchus spp.) growth as a possible indicator of ocean conditions that, in turn, may affect age‐1 walleye pollock recruitment. Marine growth rates of Pacific salmon are the result of a complex interplay of physical, biological and population‐based factors that fish experience as they range through oceanic habitats. These growth rates can, therefore, be viewed as indicators of recent ocean productivity. Thus, our hypothesis was that estimated intra‐annual growth in body weight of immature and maturing age‐4 male and female chum salmon may be used as a biological indicator of variations in rearing conditions also experienced by age‐0 walleye pollock; consequently, they may be used to predict the recruitment to age‐1 in walleye pollock. Summer SSTs and chum salmon growth at the end of July and September explained the largest amount of variability in walleye pollock recruitment indicating that physical and biological indices of ocean productivity can index fish recruitment.     

Recent decline in cod stocks in the North Sea–Skagerrak–Kattegat shifts the sources of larval supply

Cod stocks in the North Sea, including the Kattegat and the Skagerrak, have declined dramatically since the 1970s. Occasionally there is a high recruitment of juveniles in Kattegat/Skagerrak, without leading to the rebuilding of adult cod stocks despite reduced fishing mortality. In a biophysical model of egg and larval drift, we examined the potential importance of extant and historical spawning grounds for recruitment of cod in the Kattegat/Skagerrak seas using data of spawning stock biomass from the 1970s and from today's reduced stocks. The results suggest that Kattegat in the 1970s relied on largely locally retained (83%) larvae with little annual variation in recruitment. Kattegat also provided a substantial proportion of larvae recruiting in Swedish Skagerrak (72%). This is in contrast to present conditions where the Kattegat spawning stock has been reduced by 94%, and Kattegat only provides 34% of locally retained larvae and 30% to Swedish Skagerrak. Instead, the protected area in the Öresund and the Belt Sea are expected today to provide most larvae recruiting in Kattegat. Also, the inflow of larvae from the North Sea to Skagerrak and Kattegat can be significant although highly variable between years, with a positive correlation to the North‐Atlantic Oscillation index (NAO). The rebuilding of healthy spawning areas in the Kattegat may be key for restoring local cod stocks in both Kattegat and along the Skagerrak coast. This poses a management challenge if cod with local ‘Kattegat’ adaptations, e.g., in terms of egg density and migration patterns, are lost or reduced to non‐resilient densities.     

Climate change in the southeastern Bering Sea: impacts on pollock stocks and implications for the oscillating control hypothesis

Concern about impacts of climate change in the Bering Sea prompted several research programs to elucidate mechanistic links between climate and ecosystem responses. Following a detailed literature review, Hunt et al. (2011) (Deep‐Sea Res. II, 49, 2002, 5821) developed a conceptual framework, the Oscillating Control Hypothesis (OCH), linking climate‐related changes in physical oceanographic conditions to stock recruitment using walleye pollock (Theragra chalcogramma) as a model. The OCH conceptual model treats zooplankton as a single box, with reduced zooplankton production during cold conditions, producing bottom‐up control of apex predators and elevated zooplankton production during warm periods leading to top‐down control by apex predators. A recent warming trend followed by rapid cooling on the Bering Sea shelf permitted testing of the OCH. During warm years (2003–06), euphausiid and Calanus marshallae populations declined, post‐larval pollock diets shifted from a mixture of large zooplankton and small copepods to almost exclusively small copepods, and juvenile pollock dominated the diets of large predators. With cooling from 2006–09, populations of large zooplankton increased, post‐larval pollock consumed greater proportions of C. marshallae and other large zooplankton, and juvenile pollock virtually disappeared from the diets of large pollock and salmon. These shifts in energy flow were accompanied by large declines in pollock stocks attributed to poor recruitment between 2001 and 2005. Observations presented here indicate the need for revision of the OCH to account for shifts in energy flow through differing food‐web pathways due to warming and cooling on the southeastern Bering Sea shelf.