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.     

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.     

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.     

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.     

Comparison of factors affecting recruitment variability of walleye pollock Theragra chalcogramma in the Pacific Ocean and the Sea of Japan off northern Japan

The Japanese Pacific stock (JPS) and the northern Japan Sea stock (JSS) of walleye pollock Theragra chalcogramma are mainly distributed in the Pacific Ocean and the Sea of Japan off northern Japan, respectively. This paper summarizes and compares the factors affecting the recruitment variability of these two stocks. Spawning season is from December to March for both stocks. JPS recruitment has a positive relationship with the water temperature in January and February, whereas that of JSS has a negative relationship with the water temperature in January, February, and April. One possible reason for this is that pollock larvae have an optimum growth temperature of approximately 5 °C in the field. Drift of early life stages also appears to be an important influence on the recruitment of both stocks. Because the current generated by the northwest wind carries eggs of JPS into the main larval nursery ground, JPS recruitment is enhanced in years when the northwest wind is predominant in February. On the other hand, early life stages of JSS are transported into the nursery ground by the Tsushima Warm Current. However, this current also carries early life stages into the Sea of Okhotsk and offshore, resulting in poor JSS recruitment in years when this current is strong in March. In contrast to JPS, the recruitment of which is significantly impacted by cannibalism, young pollock have not been found in the stomachs of adult JSS. Warm temperatures in the Sea of Japan seem to induce the separation of young and adult pollock, and the shape of the stock–recruitment relationship also suggests that cannibalism is not important for JSS. Based on this knowledge, and on the hatch date distributions of larvae and juveniles, we propose mechanisms that can explain the recruitment fluctuations for JPS and JSS pollock.     

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.     

Interannual variability in growth of walleye pollock, Theragra chalcogramma, in the central Bering Sea

Interannual variability in growth of walleye pollock, Theragra chalcogramma, was examined. Adult walleye pollock were collected from the central Bering Sea (Aleutian Basin) from 1978 to 1999. Average fork lengths were found to be approximately 47 cm during the 1970–80s, this increased to 56 cm in the late 1990s. Age was determined for 4805 individuals using the otolith break and burn method. Ages ranged from 5–23 years and the year classes of 1978 and 1989 were dominant in the 1980s and the 1990s, respectively. Fish had significantly larger length-at-age in the 1990s compared to the 1970–80s, and interannual variability in age–length relationship was clearly observed. Taking into consideration a recent decrease of the walleye pollock biomass in the central Bering Sea, density-dependent growth was supported as one possibility of the growth variability. At the same time, we could not rule out the possibility that oceanographic variability affected the growth of walleye pollock in the area.     

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.     

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.     

Acoustic identification of isada krill, Euphausia pacifica Hansen, off the Sanriku coast, north-eastern Japan

In this study, the feasibility of identification and abundance estimation of isada krill, Euphausia pacifica , was examined using a two-frequency (38 and 120 kHz) method. The acoustic survey and midwater trawling were conducted off the Sanriku coast, northern part of Honshu, Japan. In one pattern, the SV difference between the two frequencies was large, and in the other, the SV difference was small. These results were interpreted by scattering models, and showed that discrimination between isada krill and walleye pollock was possible.     

Effect of environmental factors and prey species on daytime and nighttime vertical distribution of juvenile walleye pollock in and around Funka Bay

We examined factors affecting the vertical distribution of juvenile walleye pollock during “the second critical period” (i.e. the transition period of prey size) in and around Funka Bay, Hokkaido, Japan. During the day, small juveniles (<30 mm) were distributed mainly in shallow, warm, and low-salinity water within the bay, whereas larger ones (≥30 mm) occurred in deeper, cold and saline water. During the night, the large juveniles migrated to shallower depths. Outside the bay, small- and large-sized juveniles were distributed in water masses of similar physical properties during the daytime, whereas during the nighttime, most of the juveniles moved to shallower depth, and some juveniles dived to deeper water mass. Dietary analysis revealed that large juveniles fed on large copepods (≥0.5 mm in body width) more than small juveniles did. The composition of zooplankton in our survey area showed that large-sized copepods often occurred in deeper saline water. From these results, there is a possibility that juvenile pollock inside and outside Funka Bay occupied different water masses preferable for each feeding advantage.     

Marketing and distribution of walleye pollock: past,current and future developments

In this paper, we examine the structure and change of the walleye pollock industry from a marketing and distribution perspective. Our focus is not only on industrial internationalisation, but also on local industry and community utilisation of walleye pollock. This is a subject of interest because its diverse utilisation has the potential to promote industrial management as well as the livelihood of local communities. We reason that Japan’s weakness in marketing, distribution and price setting of walleye pollock is largely due to its lack of competitiveness in the world market. Given the rapid decline of walleye pollock utilisation by local industry and society, it is of critical importance to take steps towards a long-term solution considering the rapid decline of walleye pollock use by our local industry and society. Our analysis indicates that Japan’s walleye pollock industry is inherently resilient, with diverse commodity chains, business flexibility and traditional food culture. However, as local industry and community have been reducing use of walleye pollock over time, there is a need to restore and reactivate the local society.     

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.     

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.     

Interannual variation in the coastal distribution of a juvenile gadid in the northeast Pacific Ocean: The relevance of wind and effect on recruitment

Drift of propagules occurs within many populations inhabiting flow fields. This affects the number of propagules that rejoin their source population (recruitment) and plays a role in adaptive spatial redistribution. We focus on the cause and consequence of interannual variation in geographic distribution of population density among five cohorts of young‐of‐the‐year (age‐0) juvenile walleye pollock Gadus chalcogrammus in the western Gulf of Alaska (GOA). The coastal GOA is a wind‐driven advective system. Walleye pollock spawn during spring and their eggs and larvae drift southwestward; by late summer, age‐0 juveniles are variously distributed over the shelf. We found that high population densities of age‐0 juveniles (ca. 6 months old) near the southwestward exit of the Alaska Coastal Current from the GOA corresponded with high abundance of larvae from the major spawning area upstream, but did not translate into high abundance at older ages. Further, offshore and upwelling‐favorable winds were associated with the high downstream abundance and presumed export. In contrast, downwelling‐favorable (northeasterly) wind during and shortly after spawning (April–May) was associated with high recruitment at age 1. Finally, we found that recruitment also increased with apparent retention of age‐0 juveniles in favorable habitat upstream near the main spawning area. We hypothesize that wind‐related retention in superior upstream habitat favors recruitment. Our results argue for including wind‐driven transport in future walleye pollock recruitment models. We encourage more work on the juvenile stage of marine fishes aimed at understanding how transport and species‐specific habitat suitability interact to affect population response to large‐scale forcing.     

Interannual changes in the timing of walleye pollock spawning migration and their impacts on the gillnet fishery in the southwestern Pacific coast of Hokkaido,Donan area,Japan

The southwestern Pacific coast of Hokkaido is the main spawning ground for the Japanese Pacific stock of walleye pollock Theragra chalcogramma. A commercial gillnet fishery targeting spawning adult pollock in this area mainly operates from October to January to coincide with the migration of adult pollock from the feeding ground. Given the results of acoustic surveys, and changes in the proportion of the monthly total catch that was monthly walleye pollock caught by the commercial gillnet fishery, it is thought that the timing of walleye pollock spawning migration to the Donan area varies among years and that the pollock catch of the gillnet fishery clearly reflects changes in pollock abundance in this area. A time series of interannual variability in catch data from 1980 to 2005 suggested that adult pollock migrated and concentrated on their spawning ground later in the 1980s and after 2000 than in the 1990s. Such decadal-scale shifts are presumably caused by climatic changes (e.g., in water temperature) in the Oyashio region. These shifts affect the gillnet fishery through differences in monthly unit prices of pollock and changes in the formation of fishing grounds. These scientific findings can aid the establishment of rules for more efficient walleye pollock resource management under the total allowable catch system.     

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.     

First description and collection records of juvenile slime flounder <Emphasis Type="Italic">Microstomus achne</Emphasis> from the eastern and western coastal waters off Honshu Island,Japan

Morphological characteristics and body pigmentations of juvenile slime flounder Microstomus achne were described from one specimen [27.8 mm total length (TL)] collected off Niigata Prefecture from the Sea of Japan (35 m depth) in May 1993 and from three specimens (30.1, 36.2, and 40.9 mm TL) collected off Fukushima Prefecture on the Pacific coast of northern Japan (15 and 50 m depths) during April–May 2008 and 2009. These age-0 juveniles had a characteristically slenderly oval-shaped body and slightly protruded front nostrils. Sparsely distributed melanophores and xanthophores were clearly visible throughout the ocular side of one specimen, which was judged as a newly settled juvenile, whereas several spots aggregated by small melanophores were found in the others with elongated pectoral fins and small cycloid scales. Melanophores were only slightly detectable on the blind side. Comparisons of the depth distributions of slime flounder juveniles and sympatrically collected age-0 pleuronectids (Pseudopleuronectes herzensteini, Pseudopleuronectes yokohamae, Platichthys bicoloratus, and Dexistes rikuzenius) suggested that slime flounder can settle on broad nursery areas of coastal and offshore pleuronectid flatfishes, probably depending on local oceanic conditions during the protracted pelagic phase.     

Effects of sampling errors on abundance estimates from virtual population analysis for walleye pollock in northern waters of Sea of Japan

ABSTRACT:   Effects of sampling errors on abundance estimates from virtual population analysis (VPA) were quantified with the bootstrap method for stock of walleye pollock in the the Sea of Japan. In the bootstrap method, individual fish measurements were resampled. A total of 1000 bootstrap samples were produced for each year from 1991 to 2001. The coefficients of variation (CV) of catch at age in 2001 ranged 6.1–33.1%. The CV of an abundance estimate in 2001 ranged 9.0–35.7%. Abundance estimates of the oldest age and the latest year, which had larger CVs than the other estimates, were sensitive to sampling errors. Effects of sampling errors became smaller when the catch at age had been accumulated over a few years. Although VPA includes various types of errors, only the sampling errors have room for improvement in reality. Quantifying the effect of sampling error on VPA estimates is essential for sound and efficient stock management, and is emphasized in this study.     

A review of the distribution, ecology and population dynamics of age-0 walleye pollock in the Gulf of Alaska

This review paper synthesizes published research and unpublished data on the abundance and distribution patterns, ecology and population dynamics of walleye pollock ( Theragra chalcogramma ) during their first year of life (age-0) in the Gulf of Alaska. Distribution patterns have been described using mainly trawl catches, but recently, acoustic methodology has been employed, especially in examining vertical distributions. Although age-0 pollock are found throughout the Gulf, the highest catches occurred west of Kodiak Island. Pollock are pelagic for at least their first 6 months of life but show an ontogenetic increase in depth distribution superimposed on a pronounced diel vertical migration at a larger size. Daily growth rates are variable depending on year, season and area, and growth generally ceases during the winter. The diet of age-0 pollock shifts from mainly copepods in early juveniles to euphausiids by fall, with epibenthic organisms becoming important during the winter months. Feeding occurs mainly at night in surface waters. Age-0 pollock are most frequently associated with gelatinous zooplankton (medusae) and older pollock. Many predators on age-0 pollock have been identified; the most important are arrowtooth flounder ( Atheresthes stomias ), adult pollock, puffins ( Fratercula spp.), murres ( Una spp.), harbor seals ( Phoca vitulina richardsi ) and Steller sea lions ( Eumetopiasjubatus ). Modelling provides some insight into the population dynamics of these juveniles and environmental conditions which interannually affect their survival. These results are discussed relative to the importance of age-0 pollock in the recruitment of this species and to their role in the pelagic ecosystem.