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.     

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.     

Linking cod (Gadus morhua) and climate: investigating variability in Irish Sea cod recruitment

How climatic variability and anthropogenic pressures interact to influence recruitment is a key factor in achieving sustainable resource management. However, the combined effects of these pressures can make it difficult to detect non‐stationary interactions or shifts in the relationships with recruitment. Here we examine the links between climate and Irish Sea cod recruitment during a period of declining spawning stock biomass (SSB). Specifically, we test for a shift in the relationship between recruitment, SSB and climate by comparing an additive (generalized additive model, GAM) and non‐additive threshold model (TGAM). The relationship between recruitment success, SSB and the climatic driver, sea surface temperature, was best described by the TGAM, with a threshold identified between recruitment and SSB at approximately 7900 t. The analysis suggests a threshold shift in the relationship between recruitment and SSB in Irish Sea cod, with cod recruitment being more sensitive to climatic variability during the recent low SSB regime.     

环境因子对黄海鳀鱼亲体-补充量关系影响的初步研究

鱼类年际资源量的波动可以归因于年间环境条件的变化和该种鱼类亲体数量的变化。文章根据1990～2001年间黄海中南部鳀鱼声学调查评估结果,以及黄海千里岩海区在此年间的表层水温和营养盐统计数据,以Ricker模型（R=αSe^-βS）为基础对黄海鳀鱼（Engraulis japonicas）亲体-补充量关系进行了初步研究。结果表明,黄海千里岩水域表层水温、磷酸盐浓度等环境条件因素对补充量有重要影响。     

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.     

Greenland cod (Gadus morhua): modeling recruitment variation during the second half of the 20th century

Two approaches were used to qualify observed variability in Greenland cod (Gadus morhua) recruitment. In the first analysis, we used the linear trend of the Greenland cod recruitment time series and climatic variables, such as air temperatures from the Denmark Strait and wind conditions off East Greenland and Southwest Greenland, to explain the interannual variation in cod recruitment off Greenland. The model resulting from this ‘trend/environmental approach’, explained 79% of the interannual variation in cod recruitment off Greenland. In the second, analytical approach, the ‘regime approach’, multiple linear regression models were used, with the input data being the time series of cod recruitment and spawning stock biomass (SSB) from Iceland and Greenland, sea surface and air temperatures around Greenland, and zonal wind components between Iceland and Greenland. Model results indicated that, during the decades between 1950 and 1990, there were three different cause–effect regimes which significantly influenced the variability of cod recruitment. The three regimes included: (a) the 1950s and 1960s, a regime with favorable sea surface temperatures and a self‐sustaining cod stock off Greenland with high SSB that produced a series of above‐average, strong year classes; (b) the 1970s and 1980s, a regime of declining SSB and recruitment, with recruitment dependent on advection from Iceland; and (c) the 1990s, when the advective potential for recruitment from the Icelandic cod stock was the only available source for replenishment of the Greenland cod stocks, because cod recruitment in Greenland waters was negligible. The three models explained 76–77% of the observed interannual variation in cod recruitment off Greenland. Both approaches suggested that advective factors were the dominant influences for cod recruitment in the ‘Iceland–Greenland System’.     

Impact of spring upwelling variability off southern‐central Chile on common sardine (Strangomera bentincki) recruitment

Off southern‐central Chile, the impact of spring upwelling variability on common sardine (Strangomera bentincki) recruitment was examined by analyzing satellite and coastal station winds, satellite chlorophyll, and common sardine recruitment from a stock assessment model. In austral spring, the intensity of wind‐driven upwelling is related to sea surface temperature (SST) from the Niño 3.4 region, being weak during warm periods (El Niño) and strong during cold periods (La Niña). Interannual changes in both spring upwelling intensity and SST from the Niño 3.4 region are related to changes in remotely sensed chlorophyll over the continental shelf. In turn, year‐to‐year changes in coastal chlorophyll are tightly coupled to common sardine recruitment. We propose that, in the period 1991–2004, interannual changes in the intensity of spring upwelling affected the abundance and availability of planktonic food for common sardine, and consequently determined pre‐recruit survival and recruitment strength. However, the importance of density‐dependent factors on the reproductive dynamic cannot be neglected, as a negative association exists between spawning biomass and recruitment‐per‐spawning biomass. Coastal chlorophyll, upwelling intensity, and SST anomalies from the Niño 3.4 region could potentially help to predict common sardine recruitment scenarios under strong spring upwelling and El Niño Southern Oscillation (ENSO)‐related anomalies.     

Proposal for stock-recruitment relationship for Japanese sardine <Emphasis Type="Italic">Sardinops melanostictus</Emphasis> in North-Western Pacific

ABSTRACT:   This study assessed the stock-recruitment relationship (SRR) for the Japanese sardine Sardinops melanostictus in the North-western Pacific. Of the 20 SRR models investigated, the Akaike information criterion (AIC) was the minimum (AIC = 551.2) when the data were separated into two groups (A and B) and the log-normal distribution was applied as the error term. Group A was constructed with SRR data from 1976–1987 and 1992–2004. Group B consisted of data from 1988–1991. The AIC minimum model was R  = 22.8 S  ×  e ^ε for Group A, where R , S , and ε denote the recruitment of sardine (individual number of 0-year old fish), spawning stock biomass (SSB), and error term, respectively. This model indicated that recruitment was proportional to the SSB and that no density-dependent effect operated over the range of SSB investigated (51 000–11.3 million t). Recruitment was markedly higher (lower) when the sea surface temperature (SST) of the Kuroshio Extension area in February was low (high). The essential SRR can simply be expressed as R  = 22.8 S  ×  e ^ε with the level of recruitment deviating from the model to a greater or lesser degree depending on the environmental conditions.     

Re‐evaluating the effect of wind on recruitment in Gulf of Maine Atlantic Cod (Gadus morhua) using an environmentally‐explicit stock recruitment model

A previous study documented a correlation between Atlantic Cod (Gadus morhua) recruitment in the Gulf of Maine and an annual index of the north component of May winds. This correlation was supported by modeling studies that indicated strong recruitment of Gulf of Maine Atlantic Cod results from high retention of spring‐spawned larvae in years when winds were predominately out of the north, which favor downwelling. We re‐evaluated this relationship using updated recruitment estimates and found that the correlation decreased between recruitment and wind. The original relationship was largely driven by two recruitment estimates, one of which (2005 year‐class) was highly uncertain because it was near the terminal year of the assessment. With additional data, the updated assessment estimated lower recruitment for the 2005 year‐class, which consequently lowered the correlation between recruitment and wind. We then investigated whether an environmentally‐explicit stock recruit function that incorporated an annual wind index was supported by either the original or updated assessment output. Although incorporation of the annual wind index produced a better fitting model, the uncertainty in the estimated parameters and the implied unexploited conditions were not appropriate for providing management advice. These results suggest the need for caution in the development of environmentally‐explicit stock recruitment relationships, in particular when basing relationships and hypotheses on recruitment estimates from the terminal years of stock assessment models. More broadly, this study highlights a number of sources of uncertainty that should be considered when analyzes are performed on the output of stock assessment models.     

Contributions of FOCI research to forecasts of year-class strength of walleye pollock in Shelikof Strait, Alaska

NOAA's Fisheries Oceanography Coordinated Investigations (FOCI) contributes information to help forecast year-class strength of walleye pollock (Theragra chalcogramma ) in the Gulf of Alaska. Quantitative estimates of recruitment are obtained from models of stock assessment and stock projection employing information supplied by FOCI. To generate its information, FOCI convenes specialists in marine biology, physical and fisheries oceanography, meteorology, and statistics to assemble and analyse relevant biological and physical time series with respect to recruitment and processes hypothesized to influence fish survival. Statistical methods encompass linear and nonlinear regression, stochastic simulation modelling, transfer function time series modelling, and tree-modelling regression. The current database consists of 31 years of data, and analyses have identified factors that affect ocean stratification and circulation during spring and summer of the fish's birth year as being important to recruitment. A conceptual model of the recruitment process serves as the framework for a recruitment forecast scheme. A stochastic mathematical simulation model of the conceptual model produces similarities between simulated and observed recruitment time series. FOCI has successfully forecast recruitment observed over the past several years.     

Simulation of the impact of climate change on the egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island,the western coast of Japan

In order to investigate the impact of climate change on egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island western Japan, we conducted particle‐tracking simulations on transport success/failure to fishing grounds from 1960 to 2007. The modeled transport success since the mid‐1990s increased and decreased in the offshore and coastal zones, respectively, compared with the 1960s and 1970s. The estimated northward shift of the spawning ground and weakened Tsushima Warm Current contributed to increase in modeled transport success to the offshore zone. Conversely, the weakening trend of the modeled onshore current in the Goto‐Nada Sea combined with the northward shift of the spawning ground resulted in unsuccessful larval transport. These results suggest that fluctuations in juvenile and subadult anchovy catches in this area may be attributable to changes in the physical environment. The present study showed that changes in transport success induced by oceanographic fluctuations related to climate change, have the potential to affect anchovy recruitment off the western coast of Japan.     

Spatiotemporal catch distribution of age-0 Pacific bluefin tuna Thunnus orientalis caught by the Japanese troll fishery in relation to surface sea temperature and seasonal migration

This study used a delta-lognormal model to analyze monthly catches of age-0 Pacific bluefin tuna by the troll fishery. The model included fixed effects of month, area, and month–area interaction, and random effects of port, year and port–year interaction. The catch patterns by month and area predicted by the statistical model (standardized catch) revealed that main fishing grounds along the Tsushima Warm Current generally shifted from north to south as the season turned from autumn to winter. In contrast, the standardized catch along the Kuroshio Current did not show such clear spatiotemporal patterns. The standardized catch along the Tsushima Warm Current is significantly associated with average monthly sea surface temperatures in the fishing grounds and consistent with migration routes revealed by tagging experiments in previous studies. These associations indicate the spatiotemporal catch pattern in the Tsushima Warm Current region partly reflects seasonal migration. Knowledge of the possible associations among fish migration, environmental factors and spatiotemporal distribution of the catch will contribute to future management of this species.     

Length-based Beverton and Holt spawning stock biomass per-recruit-models, with application to the Lates stappersii (Boulenger) stock in Lake Tanganyika

Two kinds of length-based Beverton and Holt spawning stock biomass per-recruit ( SSB/R )-models are presented. The first allows the computation of the absolute value of SSB/R . The second estimates the index of SSB/R , also called 'relative SSB/R '. Both models are extensions of the original age-based Beverton and Holt SSB/R -models. The SSB/R and its index can be assessed with reference to variations in the exploitation characteristics. These models were applied to the Lates stappersii (Boulenger) stock in Lake Tanganyika using fishing mortality as the independent variable. They produce an absolute value of SSB/R of 12.95 g, representing 22.4% of that of the 'virgin' stock for the 1993–1994 equilibrium-fishing regime. This level is indicative of an unsafe state of the stock in question. Moreover, the SSB for L. stappersii seems to be fragile under increased fishing pressure as, for example, doubling or tripling the current fishing regime would reduce the current SSB/R by 57.3 and 77.4%, respectively. In general, the use of length-based Beverton and Holt SSB/R -models is handicapped by underlying hypotheses and sub-models which are restrictive.     

Is spawning stock biomass a robust proxy for reproductive potential?

Reference points based on fishing mortality (F) and spawning stock biomass (SSB) are a requirement of many fisheries management frameworks. SSB is assumed to be a proxy for stock reproductive potential (SRP). Limit reference points based on SSB are used to indicate the level of biomass below which productivity is affected. SSB fails to account for changes in fecundity, egg viability and sex ratio, and it has been argued that total egg production (TEP) provides a better reflection of SRP. We explore how accounting for TEP impacts limit reference points and evidence for a relationship between stock and recruit. Time series of SSB and TEP are compared for three North Sea stocks: cod (Gadus morhua), herring (Clupea harengus) and plaice (Pleuronectes platessa). Dynamics based on TEP are different from those based on SSB for cod and plaice, but the stock–recruit relationships were not ‘improved’ using TEP. Shifts in productivity (spawner per recruit) occur in all three time series and SSB underestimated uncertainty. Yet again, it was shown that assumptions of stationarity about fish population productivity are incorrect. We argue that the use of TEP does improve the realism in our understanding of stock dynamics, and demographically, more complex management strategy evaluation is required to develop management procedures that are robust to uncertainty and integrate F and the demographic health of a stock. Empirical feedback control systems based on fisheries independent indices including surveys of eggs, larvae, recruits, juveniles or spawning adults should be evaluated and compared to traditional approaches.     

Density dependence in total egg production per spawner for marine fish

A paradigm of fisheries science holds that spawning stock biomass (SSB) is directly proportional to total egg production (TEP) of fish stocks. This “SSB–TEP proportionality” paradigm has been a basic premise underlying the spawner–recruitment models for fisheries management and numerous studies on recruitment mechanisms of fish. Studies on maternal effects on reproductive potential of a stock have progressed during the last few decades, leading to doubt concerning the paradigm. Nonetheless, a direct test of the paradigm at multidecadal scales has been difficult because of data limitations in the stock assessment systems worldwide. Here, we tested the paradigm for marine fish based on a novel combination of two independent 38‐year time series: fishery‐dependent stock assessment data and fishery‐independent egg survey data. Through this approach, we show that the SSB–TEP proportionality is distorted by density dependence in total egg production per spawner individual (TEPPS) or spawner unit weight (TEPPSW) at a multidecadal scale. The TEPPS/TEPPSW exponentially declined with biomass and thus was density‐dependent for Japanese sardine, a small pelagic species exhibiting a high level of population fluctuation, in the western North Pacific. By contrast, the TEPPS/TEPPSW was sardine‐density‐dependent for Japanese anchovy, another small pelagic species exhibiting a moderate level of population fluctuation well‐known for being out of phase with sardine. Our analysis revealed intraspecific (sardine) and interspecific (anchovy) density dependence in TEPPS/TEPPSW, which was previously unaccounted for in spawner–recruitment relationships. Such density‐dependent effects at the time of spawning should be considered in fisheries management and studies on recruitment mechanisms.     

Population reproductive potential: Evaluation of long-term stock productivity

In order to avoid recruitment overfishing, fish stocks must have sufficient reproductive ability. The spawning stock biomass (SSB), which ignores the value of immature fish, is widely used as an index of stock sustainability. From the perspective of sustainability, immediate reproduction, as well as future spawning, must be considered. We developed an index of long-term stock productivity, called the population reproductive potential (PRP). PRP is defined as the expected total reproductive value of the standing stock. We used PRP to assess the western Atlantic bluefin tuna (WBT) stock. The trends in SSB, numbers (N), biomass and PRP of WBT are inconsistent when compared to each other, due to fluctuation in age composition. We evaluated the long-term productivity of WBT by computer simulation and compared the result with trends in the abundance indices. The result of the computer simulation was highly consistent with the trend in the PRP. Short-term trends in SSB and N often do not reflect long-term stock trends, because they are highly sensitive to age-composition dynamics. The PRP is useful for evaluating stock trends, especially when the age composition is unstable.     

Density‐dependent egg production in chub mackerel in the Kuroshio Current system

A paradigm of proportionality between spawning stock biomass (SSB) and total egg production (TEP) has been largely untested at multidecadal scales mainly because of difficulty in estimating annual TEP. Recently, this paradigm was directly tested for sardine (Sardinops melanostictus) and anchovy (Engraulis japonicus) at a multidecadal scale to reveal that SSB–TEP proportionality was partially distorted by intraspecific (sardine) and interspecific (anchovy) density dependence in total egg production per spawner individual (TEPPS) or unit weight (TEPPSW). In the present study, we demonstrate intraspecific density dependence in TEPPS/TEPPSW for chub mackerel (Scomber japonicus) in the Kuroshio Current system, using a proxy for TEPPS/TEPPSW, calculated from snapshot abundance data based on fishery‐independent egg surveys in combination with fishery‐dependent stock assessment data, at a multidecadal scale (38 years). TEPPS/TEPPSW exponentially declined with SSB, indicating a strong intraspecific density dependence in TEPPS/TEPPSW in chub mackerel. The observed phenomenon for chub mackerel was similar to that for sardine. Hence, intraspecific density dependence in TEPPS/TEPPSW may be a phenomenon that is generally applicable for species with a high maximum biomass and large population fluctuations. Lastly, we recommend the application of a TEP‐based framework to studies on recruitment mechanisms of fish.     

东海区小黄鱼繁殖模型优化选择及其管理应用研究

根据 1999-2008 年东海区渔业资源底拖网大面定点调查获取的小黄鱼(Larimichthys polyactis)渔业生物学资料,利用AIC(Akaike Information Criterion)与BIC(Bayesian Information Criterion),对小黄鱼的Ricker、Beverton-Holt和Cushing繁殖模型进行了拟合优度检验.针对选择的繁殖模型,经单因子相关分析和逐步回归分析,筛选对繁殖模型有重要影响的环境因子,经模型的拟合和检验,确定东海区小黄鱼的适用繁殖模型.结果表明:3种繁殖模型中,Ricker繁殖模型更适合模拟小黄鱼亲体与补充量关系,但吕泗渔场海域3-4月平均海水表温、7月海水表温和5月长江径流量、7-8月长江平均径流量以及当年夏季风速对模型中的补充量有着重要的影响;优化后的Ricker繁殖模型不仅可以提高东海区小黄鱼亲鱼量与补充量的拟合精度,而且可参考该模型修正当年度小黄鱼的资源管理目标,提高资源管理的科学性.     

Impact of winter‐to‐spring environmental variability along the Kuroshio jet on the recruitment of Japanese sardine (Sardinops melanostictus)

Winter‐to‐spring variability in sea surface temperature (SST) and mixed layer depth (MLD) around the Kuroshio current system and its relationship to the survival rate (ln [recruit per spawning stock biomass], LNRPS) of Japanese sardine (Sardinops melanostictus) were investigated based on a correlation analysis of data from 1980 to 1995. The data were from a high‐resolution ocean general circulation model using the ‘Kuroshio axis coordinates’, in which the meridional positions are relocated to a latitude relative to the Kuroshio axis at each longitude, rather than the geographically fixed coordinates. A significant positive (negative) correlation between LNRPS and winter MLD (winter–spring SST) was detected near the Kuroshio axis from areas south of Japan (where eggs are spawned) to the Kuroshio Extension (where larvae are transported). This result is in contrast to previous studies using geographically fixed coordinates, which showed a significant correlation predominantly in the area south of the Kuroshio Extension in winter, where at this time few larvae have been found. From the late 1980s to early 1990s, when the survival rate was remarkably low, MLD around the axis was shallow and SST was high. Although MLD and SST show a significant correlation, significant partial correlations were also observed between February MLD and LNRPS when the contribution of SST was excluded, and between March SST and LNRPS when the contribution of MLD was excluded. We presume that MLD shoaling reduced the nutrient supply from deep layers, resulting in less productivity in the spring, and SST warming could have a negative influence on larval growth.     

Population trends of bycatch species reflect improving status of target species

Synthesis studies of fish stocks worldwide suggest improving status of mainly target species that are fully assessed. Other analyses, primarily based on catch data alone, but which include a wider range of species as well as bycatch, present a different view. Catch‐only analyses could be more robust if fishery‐independent data were used and discards accounted for. We develop a model that uses only survey biomass at length and landings data to estimate fishing mortality, spawning stock biomass (SSB) and discards. An analysis of species from the North Sea shows the model results compare well with most fully assessed stocks. When applied to bycatch species with limited data, trends in fishing mortality and SSB typically reflect those of the target species. In the last decade, mean fishing mortality rates have tended to decline, while mean SSB has increased. Despite increasing SSB, recent mean recruitment appears to have been lower than previously which may limit future biomass recovery. Species usually associated with more northerly distributions appear to show the greatest effect of weaker recruitment, which may be linked to climate. Estimated discards have tended to decline in magnitude as a result of reduced fishing mortality and associated lower total catches. The model offers a simple way to use both landings and survey data to obtain more detailed population trends for data limited species.