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.     

Complete nucleotide sequence and variation of mitochondrial DNA from 10 individuals of walleye pollock, Theragra chalcogramma

ABSTRACT:   The complete mitochondrial genome sequence of 10 walleye pollocks, Theragra chalcogramma , from the Japan Sea and Bering Sea was determined. The 16 568–16 571 bp genome contains the same 37 mitochondrial structural genes (two ribosomal RNA, 22 transfer RNA, and 13 protein-coding genes) as found in all other vertebrates analyzed, in an organization identical to that of other bony fish. The major non-coding region had several conserved sequence features. Nucleotide variations of ND1, ND5, and control region were high, and these regions appear to be good candidates for high-resolution markers in population studies.     

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.     

Mortality of larval walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1988–91

Mortality rates of larval walleye pollock Theragra chakogramma were estimated from larval survey data from 1988 to 1991. Mortality estimates were based on cohort-specific losses between occupations of survey grids. Interannually, estimates of early feeding stage larval mortality rates ranged over an order of magnitude, from 0.045–0.43 day^-1, and declined sharply with age. There is some evidence that mortality rates of early feeding larvae tend to be negatively correlated with temperature and postively correlated with wind mixing.     

Comparative analysis of ichthyoplankton sampling gear for early life stages of walleye pollock (Theragra chalcogramma)

Comparative gear tows were conducted during the summers of 1987–1990 in the Gulf of Alaska. Efficiencies of four types of ichthyoplankton sampling gear were compared with respect to total catch and size distribution of larval fish. Fish were sampled using the bongo net (333 and 505 μm mesh), 1 m Tucker net, 3 m Tucker net and the Methot frame net. Results indicate that the bongo and the 1 m Tucker nets caught similar numbers and size distribution of fish. The larger 3 m Tucker and Method frame nets caught a larger size class of fish but a smaller total catch. Although there was a slightly significant diel difference in total catch detected in the smaller nets, no diel differences in size distribution of fish caught were detected for any of the nets.     

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.     

Cohort survival patterns of walleye pollock, Theragra chalcogramma, in Shelikof Strait, Alaska: a critical factor analysis

A series of age-specific life tables for walleye pollock ( Theragra chalcogramma ) in the western Gulf of Alaska was compiled for the 1980-91 year classes. The life tables were utilized to perform an exploratory key factor analysis to examine the timing of critical periods in the recruitment process, evidence of density-dependence at different stages and trends in mortality rates. Early larval mortality was significantly correlated with generational mortality (In recruits/spawning bio-mass), but patterns in juvenile mortality also were similar to generational mortality and in some years were clearly dominant in determining the fate of a cohort. Density-dependent mortality, based on the correlation between mortality and initial abundance, was indicated only for the late larval to early juvenile stage. Time trends were marginally significant for juvenile mortality. It is speculated that the observed increase in juvenile mortality is associated with increasing abundance of arrowtooth flounder. Weaknesses in the data base are discussed; these along with the short time series involved make our conclusions tentative and subject to further study. We hypothesize that pollock recruitment levels can be established at any life stage depending on sufficient supply from prior stages, a type of dynamics which can be termed supply dependent multiple life stage control.     

Effect of ocean conditions on the cross-shelf distribution of walleye pollock (Theragra chalcogramma) and capelin (Mallotus villosus)

Acoustic trawl surveys were conducted in 2000 and 2001 in two troughs located off the eastern coast of Kodiak Island in the Gulf of Alaska as part of a multiyear, multidisciplinary experiment to examine the influence of environmental conditions on the spatial distribution of adult and juvenile walleye pollock (Theragra chalcogramma) and capelin (Mallotus villosus). Continuous underway sea surface temperature samples and water column profiles collected in 2000 and 2001 showed the presence of a sharp shelf‐break front in Chiniak Trough and a mid‐trough front in Barnabas Trough. At distances <22 km from shore, the water column was well mixed, whereas a well‐defined mixed layer was present beyond approximately 22 km from shore. Satellite drifter tracks in Barnabas Trough entered along the upstream edge of the trough and appeared to follow the frontal boundary across the middle portion of the trough. A storm in 2001 weakened stratification and cooled surface water temperature by 1.6–2.1°C. Wind mixing associated with the storm event mixed subsurface chlorophyll a to the surface and enhanced nutrients in the surface waters. The storm event revealed spatial partitioning of summer production in Barnabas Trough, with production concentrated in regions inside the mid‐trough front. In contrast, post‐storm summer production was distributed throughout Chiniak Trough. The spatial distribution of walleye pollock and capelin differed and appeared to be related to differences in habitat characteristics. Acoustic survey data identified four acoustic sign types: age‐1 pollock, adult pollock, capelin, capelin–age‐0 pollock mix. The spatial distribution of these four sign types appears to be influenced by the oceanographic and topographic features of the two troughs. Adult pollock were broadly distributed throughout Chiniak Trough, whereas adult pollock were aggregated on the coastal side of the frontal system in Barnabas Trough. In 2000, capelin occurred with age‐0 pollock. In Chiniak Trough, capelin were most abundant along steep topographic gradients at the edges of the trough and in a deep region near Cape Chiniak, whereas the capelin–age‐0 mix (2000) or capelin (2001) concentrations were observed in slope water intrusions over the outer shelf in Barnabas Trough. Results suggest that habitat selection of walleye pollock and capelin are controlled by different processes. Capelin distributions appear to be limited by oceanographic conditions while other factors appear to be more important for pollock.     

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.     

Utilization of free amino acids, yolk proteins and lipids in developing eggs and yolk-sac larvae of walleye pollock Theragra chalcogramma

ABSTRACT:   To elucidate the utilization of the major yolk nutrient stocks in eggs and larvae of walleye pollock Theragra chalcogramma , the contents of free amino acids (FAA), the major yolk protein (180 kDa lipovitellin originated from vitellogenin B in ovulated eggs: oLv B), and lipids were measured. Most eggs hatched 18 days after fertilization at 5°C, and all larvae absorbed almost all their yolk mass by 28 days. The total FAA content showed no change during the first 6 days, and then decreased to 28% of the initial level by 18 days. The oLv B contents, measured by an enzyme-linked immunosorbent assay using a specific antiserum against oLv B, gradually decreased from 6 to 18 days, followed by a rapid decline. The content of phospholipids (PL) and triacylglycerols (TG) showed no marked change until hatching, and then decreased until disappearance of yolk sac. From these results, it is proposed that there are two main periods for nutrient utilization in embryos and larvae of walleye pollock. In the first period, FAA was mainly utilized until 18 days after fertilization. Active utilization of oLv B and lipids (PL and TG) instead of FAA occurred during the second period from 18 to 28 days.     

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.     

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.     

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.     

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.     

Interannual variability in growth of larval and juvenile walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1983–91

Interannual variability in growth of larval walleye pollock Theragra chalcogramma was examined from 1983 to 1991 and of juveniles from 1985 to 1990. ANCOVA was used to assess differences in population growth rates, and an alternate method was developed to examine variations between annual length-at-age data and average 'expected' values over different age groupings. For larvae, the years 1986, 1987, 1989 and 1990 had higher than average length-at-age, and 1988 and 1991 had lower than average values. Relationships between growth and SST and larval density were not clear. A tentative relationship between copepod nauplii abundance and larval length-at-age was noted. The consequence of larval growth to larval mortality, late larval abundance or recruitment was not clear. We conclude that larval mortality rates are highly variable and tend to mask effects of moderate variability in growth on later abundance. For juveniles, 1987 had significantly lower than average length-at-age and 1988 had higher than average values. Although there are few years of data, they tend to support the importance of juvenile growth in the recruitment process. Conditions for the large 1988 year class are documented and discussed, including warm SST, calm winds, relatively low larval growth rates, low abundances of potential predators on larvae, low larval mortality rates, and high juvenile growth rates.