The highs and lows of herring: A meta‐analysis of patterns and factors in herring collapse and recovery

Pacific and Atlantic herring populations (genus Clupea) commonly experience episodic collapse and recovery. Recovery time durations are of great importance for the sustainability of fisheries and ecosystems. We collated information from 64 herring populations to characterize herring fluctuations and determine the time scales at low biomass and at high and low recruitment, and use generalized linear models and Random Survival Forests to identify the most important bottom‐up, top‐down and intrinsic factors influencing recovery times. Compared to non‐forage fish taxa, herring decline to lower minima, recover to higher maxima and show larger changes in biomass, implying herring are more prone to booms and busts than non‐forage fish species. Large year classes are more common in herring, but occur infrequently and are uncorrelated among regionally grouped stocks, implying local drivers of high recruitment. Management differs between Pacific and Atlantic herring fisheries, where at similarly low biomass, Pacific fisheries tend to be closed while Atlantic fisheries remain open. This difference had no apparent effect on herring recovery times, which averaged 11 years, although most stocks with longer recovery periods had not yet recovered at the end of the observation period. Biomass recovery is best explained by median recruitment and variability in sea surface height anomalies and sea surface temperatures—higher variability leads to shorter recovery times. In addition, the duration of recruitment failure is closely linked with low biomass. While recovery times rely on the nature of the relationship between spawning biomass and recruitment, they are still largely governed by complex and uncertain processes.     

Sudden change in long‐term ocean climate fluctuations corresponds with ecosystem alterations and reduced recruitment in Norwegian spring‐spawning herring (Clupea harengus,Clupeidae)

Fish stocks vary in abundance. The causes behind the fluctuations may be difficult to determine, especially ones caused by natural fluctuations, but long‐term data series may provide indications of the mechanisms. Assessments show that the recruitment to the Norwegian spring‐spawning herring (Clupea harengus, Clupeidae) has remained low since 2004, a year which produced the last really rich year‐class. Long time‐series of estimated recruitment and mean winter temperature in the ocean showed a significant positive correlation for the period 1921–2004. Here, we show that this positive correlation did not continue from 2005 onwards as the winter temperature increased to high levels while herring recruitment decreased and has remained low. The density of zooplankton in the drift route of the herring larvae dropped significantly after 2004, and their centre of gravity shifted northwards. There may currently be heavy predation on the larvae by Atlanic mackerel (Scomber scombrus, Scombridae), and top‐down regulation is suggested to hamper successful recruitment. Our analysis indicates that the presence of food and overlap with high food concentrations are likely important regulators of survival in herring larvae. The findings may be important for future management and planning of fisheries of this stock because recruitment failure may continue if temperature remains high and food abundance remains low.     

Variation in reproductive potential and influence on Icelandic herring recruitment

Explaining recruitment variation in fish is essential for successful fishery management and is consequently under constant review, with an increasing focus on how maternal factors, relative to environmental influences, operate at the level of individual female spawners and extend from the spawning stock through to recruitment. We estimate total egg production (E) in Icelandic summer‐spawning herring (Clupea harengus) from 1963 through 1999 by using sequential population analyses (SPA) and their estimates of stock biomass and recruitment, various size and maturity metrics, and individual fecundity estimates that rely on total length and the condition of the spawners. Generalized linear models indicate that maternal effects are of significance in explaining SPA‐based recruitment‐at‐age‐3 (R). The best model explained 64% of the variation in R and incorporates E constrained to the repeat spawners (40%), the NAO winter index (18%) and ocean temperature (6%). The latter two represent the winter and spring periods subsequent to year‐class formation. Recruit spawner contributions to E were of no significance in explaining variation in R despite the fact that they could contribute as much as 55% of E when their contribution to E was consistently underestimated by a factor of ～ 2, based only on their contribution to spawning stock biomass. We conclude that the spawning potential of the repeat spawners should replace total spawning stock biomass for determining recruitment potential in stock assessment. In addition to the incorporation of oceanographic factors, this would provided a more cautious and risk‐adverse approach.     

Pacific herring, Clupea pallasi, recruitment in the Bering Sea and north-east Pacific Ocean, II: relationships to environmental variables and implications for forecasting

Previous studies have shown that Pacific herring populations in the Bering Sea and north-east Pacific Ocean can be grouped based on similar recruitment time series. The scale of these groups suggests large-scale influence on recruitment fluctuations from the environment. Recruitment time series from 14 populations were analysed to determine links to various environmental variables and to develop recruitment forecasting models using a Ricker-type environmentally dependent spawner–recruit model. The environmental variables used for this investigation included monthly time series of the following: southern oscillation index, North Pacific pressure index, sea surface temperatures, air temperatures, coastal upwelling indices, Bering Sea wind, Bering Sea ice cover, and Bering Sea bottom temperatures. Exploratory correlation analysis was used for focusing the time period examined for each environmental variable. Candidate models for forecasting herring recruitment were selected by the ordinary and recent cross-validation prediction errors. Results indicated that forecasting models using air and sea surface temperature data lagged to the year of spawning generally produced the best forecasting models. Multiple environmental variables showed marked improvements in prediction over single-environmental-variable models.     

Reconstruction of Atlantic herring (Clupea harengus) recruitment in the North Sea for the past 455 years based on the δ13C from annual shell increments of the ocean quahog (Arctica islandica)

Understanding the recruitment variability of the Atlantic herring North Sea stock remains a key objective of stock assessment and management. Although many efforts have been undertaken linking climatic and stock dynamic factors to herring recruitment, no major attempt has been made to estimate recruitment levels before the 20th century. Here, we present a novel annually resolved, absolutely dated herring recruitment reconstruction, derived from stable carbon isotope geochemistry (δ¹³C), from ocean quahog shells from the Fladen Ground (northern North Sea). Our age model is based on a growth increment chronology obtained from fourteen shells. Ten of these were micromilled at annual resolution for δ¹³C analysis. Our results indicate that the anthropogenically driven relative depletion of ¹³C, the oceanic Suess effect (oSE), became evident in the northern North Sea in the 1850s. We calculated a regression line between the oSE‐detrended δ¹³C results (δ¹³C?) and diatom abundance in the North Sea, the regression being mediated by the effect of phytoplankton on the δ¹³C of the ambient dissolved inorganic carbon. We used this regression to build an equation mediated by a nutritional link to reconstruct herring recruitment using δ¹³C?. The reconstruction suggests that there were five extended episodes of low‐recruitment levels before the 20th century. These results are supported by measured recruitment estimates and historical fish catch and export documentation. This work demonstrates that molluscan sclerochronological records can contribute to the investigation of ecological baselines and ecosystem functioning impacted by anthropogenic activity with implications for conservation and stock management.     

Long-term trends in fish recruitment in the north-east Atlantic related to climate change

This study investigates the temporal correspondence between the main patterns of recruitment variations among north‐east Atlantic exploited fish populations and large‐scale climate and temperature indices. It is of primary importance to know what changes in fish stock productivity can be expected in response to climate change, to design appropriate management strategies. The dominant patterns of recruitment variation were extracted using a standardized principal component analysis (PCA). The first principal component (PC) was a long‐term decline, with a stepwise change occurring in 1987. A majority of Baltic Sea, North Sea, west of Scotland and Irish Sea populations, especially the gadoids, have followed this decreasing trend. On the contrary, some herring populations and the populations of boreal ecosystems have followed an opposite increasing trend. The dominant signal in north‐east Atlantic sea surface temperature, also extracted by a PCA, was highly correlated with the increase in the Northern Hemisphere Temperature anomaly, which is considered to be an index of global warming. The first component of recruitment was inversely correlated with these changes in regional and global temperature. The second PC of recruitment was a decadal scale oscillation, which was not correlated with climate indicators. The analysis of correlations between population recruitment and local temperature also indicated that the dominant pattern of recruitment variation may be related to an effect of global warming. The influence of fishing on recruitment, via its effect on the spawning stock biomass (SSB), was also investigated by the analysis of correlations between fishing mortality, SSB and recruitment. Results indicate that fishing can be another factor explaining recruitment trends, probably acting in combination with the effect of climate, but cannot explain alone the patterns of recruitment variation found here.     

Migrations and hydrography determine the abundance fluctuations of blue whiting (Micromesistius poutassou) in the Barents Sea

The Barents Sea is the north‐eastern fringe of the distribution of blue whiting (Micromesistius poutassou). Fluctuations in distribution and abundance of blue whiting in the area have been marked. Two hypotheses are put forward to explain these fluctuations. First, rich year classes in the main Atlantic stock of blue whiting may contribute to increased abundance in the Barents Sea. Second, variations in hydrography, such as influx of warm Atlantic water, may be particularly important in this fringe area. We investigated these hypotheses using data from bottom trawl surveys conducted during the period 1981–2006. Variations in abundance (measured either as incidence or density) and distribution were correlated with recruitment in the Atlantic stock of blue whiting as well as hydrographic conditions. Regression analyses indicated that the abundance fluctuations are primarily determined by variations in recruitment of Atlantic blue whiting, a strong year class leading to high abundance in the Barents Sea the year after spawning. However, salinity anomaly in the Fugløya–Bear Island transect during the previous year, an indicator of high inflow of Atlantic water, had also a significant, positive effect. Thus, the data suggested a climatic modulation of dynamics that were primarily determined by recruitment of blue whiting in the main Atlantic stock. Analyses of size structure as well as earlier studies on population genetics supported this conclusion.     

Long-term variability in spawning stock age structure influences climate–recruitment link for Barents Sea cod

Fish populations may spawn a vast number of offspring, while only a small and highly variable fraction of a new cohort survives long enough to enter into the fisheries as recruits. It is intuitive that the size and state of the spawning stock, the adult part of the fish population, is important for recruitment. Additionally, environmental conditions can greatly influence survival through vulnerable early life stages until recruitment. To understand what regulates recruitment, an essential part of fish population dynamics, it is thus necessary to explain the impact of fluctuations in both spawning stock and environment, including interactions. Here, we examine if the connection between the environment and recruitment is affected by the state of the spawning stock, including biomass, mean age and age diversity. Specifically, we re-evaluate the hypothesis stating that recruitment from a spawning stock dominated by young fish and few age classes is more vulnerable to environmental fluctuations. We expand upon earlier work on the Barents Sea stock of Atlantic cod, now with data series extended in time both backwards and forwards to cover the period 1922–2019. While our findings are correlative and cannot prove a specific cause and effect mechanism, they support earlier work and strengthen the evidence for the hypothesis above. Furthermore, this study supports that advice to fisheries management should include considerations of environmental status.     

Observation and quantification of two incidents of mass fish kill of Icelandic summer spawning herring (Clupea harengus) in the winter 2012/2013

The Icelandic summer‐spawning herring (Clupea harengus) stock overwinters in large, dense schools like other herring stocks. In the winter of 2012/2013 around 300 thousand tonnes, or ~70% of the spawning stock, overwintered in a fjord west of Iceland. The inner part of the fjord, where the herring was located, is separated from the outer part with a natural barrier and a built‐up road with a 210 m long bridge. This creates strong tidal currents under the bridge. On 14 December and again on 1 February mass mortalities of herring took place in this location, and the sea floor and the shores were covered with dead herring. Fieldwork, including camera and video recordings on the shore and on a small boat, was conducted 3 and 4 days after the incidents. Results from this indicated that a total of 55 thousand tonnes had died during these two incidents, an amount nearly equal to the total catch from the stock that year. Measurements of environmental conditions in the days following the incidents showed that the oxygen saturation was generally 20%–40% but was as low as 15% (1.1 ml/L). The most likely explanation for the mortalities was oxygen depletion resulting from respiration of the large herring biomass, limited atmospheric‐water gas exchange due to calm and cold weather prior to both incidents and sea ice on part of the fjord, and limited renewal of water coming in and out via tidal currents. Aerobic decomposition of dead herring came additionally in the latter incident.     

Does larval mortality influence population dynamics? An analysis of North Sea herring (Clupea harengus) time series

The daily mortality rates of North Sea herring early‐stage larvae are found to vary over decades. Larval abundance data were used with a spatio‐temporal oceanographic model to reconstruct temperature histories of the observed larvae. The histories were used in conjunction with a temperature‐based growth model to estimate larval age. Mean daily mortality rates were then estimated for the four spawning components (Downs, Banks, Buchan and Orkney/Shetland) using the vertical life table approach, which considers instantaneous abundances across all ages rather than following distinct cohorts. All spawning components, but especially Downs (in the south), exhibited a steady rise in mortality associated with increasing population size. In addition, the three northern components shared a distinct trend in mortality that was significantly correlated with ambient water temperatures experienced by the larvae during the respective time periods after hatching. This trend was also significantly negatively correlated with the residuals of the whole stock‐recruitment relationship. These findings were generally robust to assumptions about growth and hatch length of larvae. The compensatory increase in productivity in the late 1980s and poor recruitment since 2000 coincide with changes in the mortality of larvae younger than 30 days post hatch and covary with larval density and temperature. Thus we suggest that the mortality of early‐stage larvae does impact on the population dynamics in North Sea herring in its current productivity regime, implying a critical period in the determination of year class strength.     

Stock–environment–recruitment models for North Atlantic albacore (Thunnus alalunga)

Different stock–recruitment models were fitted to North Atlantic albacore (Thunnus alalunga) recruitment and spawning stock biomass data. A classical density dependence hypothesis, a recent environmental‐dependence hypothesis and a combination of both were considered. For the latter case, four stock–environment–recruitment models were used: Ricker, Beverton‐Holt, Deriso's General Model (modified to take into account environmental effects) and conditioned Neural Networks. Cross‐validation analysis showed that the modified Deriso model had the best predictive capability. It detected an inverse effect of the North Atlantic Oscillation (NAO) on recruitment, a Ricker‐type behaviour with density dependent overcompensation when environmental conditions are unfavourable and a Beverton–Holt‐type behaviour towards an asymptotic recruitment carrying capacity with favourable environmental conditions. The Neural Network model also detected that under favourable environmental conditions high spawning stock biomass does not necessarily have a depensatory effect on recruitment. Moreover, they suggest that under extremely favourable environmental conditions, albacore recruitment could increase well above the asymptotic carrying capacity predicted by Beverton–Holt‐type models. However, the general decrease in spawning stock biomass in recent years and increasing NAO trends suggest that there is low probability of exceptionally large recruitment in the future and instead there is a danger of recruitment overfishing.     

Relationship between recruitment of Japanese sardine (Sardinops melanostictus) and environment of larval habitat in the low‐stock period (1995–2010)

Stock level of Japanese sardine (Sardinops melanostictus) was high from 1980s to early 1990s and low from late 1990s to 2000s. The warm and cold water masses in the vicinity of the Kuroshio axis from winter to early spring used to be critical for the recruitment in the high‐stock period, because most of the larvae were distributed there. However, the environmental fluctuation might not affect the recruitment in the low‐stock period. Some studies reported that spawning location and spawning season, and hence the larval habitat, differ depending on the stock level. Three points were investigated in this study: (a) how spawning location and spawning season shifted from the late 1990s, (b) confirmation of the distribution area of larvae in the recent low‐stock period and (c) whether the water temperature in the vicinity of the Kuroshio axis was still related to the recruitment in the low‐stock period. The spawning location and spawning season clearly changed after 1995. Consequently, particle tracking experiments suggested that the larvae appeared in the vicinity of the Kuroshio axis from winter to early spring decreased. Nevertheless, only the ambient temperature of larvae that appeared in the vicinity of the Kuroshio axis from winter had a significant negative correlation with an index of the recruitment in the low‐stock period. It is suggested that the warm and cold masses in the vicinity of the Kuroshio axis are critical for the recruitment regardless of the stock level.     

Forecasting fish stock dynamics under climate change: Baltic herring (Clupea harengus) as a case study

Climate change and anthropogenic disturbances may affect marine populations and ecosystems through multiple pathways. In this study we present a framework in which we integrate existing models and knowledge on basic regulatory processes to investigate the potential impact of future scenarios of fisheries exploitation and climate change on the temporal dynamics of the central Baltic herring stock. Alternative scenarios of increasing sea surface temperature and decreasing salinity of the Baltic Sea from a global climate model were combined with two alternative fishing scenarios, and their direct and ecosystem‐mediated effects (i.e., through predation by cod and competition with sprat) on the herring population were evaluated for the period 2010–2050. Gradual increase in temperature has a positive impact on the long‐term productivity of the herring stock, but it has the potential to enhance the recovery of the herring stock only in combination with sustainable fisheries management (i.e., F_msy). Conversely, projections of herring spawning stock biomass (SSB) were generally low under elevated fishing mortality levels (F_high), comparable with those experienced by the stock during the 1990s. Under the combined effects of long‐term warming and high fishing mortality uncertainty in herring SSB projections was higher and increasing for the duration of the forecasts, suggesting a synergistic effect of fishery exploitation and climate forcing on fish populations dynamics. Our study shows that simulations of long‐term fish dynamics can be an informative tool to derive expectations of the potential long‐term impact of alternative future scenarios of exploitation and climate change.     

The effects of survey design and circulation pattern on the perceived abundance of herring larvae: a case study for Norwegian spring spawning herring (Clupea harengus)

A larval survey is used in the annual assessment as an index of the spawning stock size of Norwegian spring spawning herring (Clupea harengus). To test how inter‐annual fluctuations in circulation pattern, survey design and execution of the survey affected the larval abundance estimate we conducted simulated surveys using a model framework with idealized assumptions to model larval drift and sampled larvae using several realistic survey scenarios. The results suggest that inter‐annual variations in circulation pattern alone can have a profound effect on the perception of larvae abundance and that the direction of the survey (north to south versus south to north) can have a significant effect on the estimated abundance, particularly if hatching occurs over a short period of time. Additionally, disruptions to a continuous survey schedule also have an effect and, as such, sampling strategies in case of disruption to the survey are proposed.     

澳洲鲭太平洋群系的资源评估与管理策略

澳洲鲭是西北太平洋重要的经济种类,了解和掌握澳洲鲭太平洋群系资源开发状况对确保其可持续利用具有重要的意义。根据日本中央水产研究所提供的1995—2015年澳洲鲭太平洋群系的生产统计和资源调查资料,利用基于年龄结构的实际种群模型和单位补充量产量模型等进行资源量评估,分析澳洲鲭太平洋群系资源利用情况及其管理策略。结果显示,历年澳洲鲭太平洋群系资源量虽有波动但仍保持在较高水平,2015年资源量最高约为65万t;年平均捕捞死亡系数呈波动下降趋势,2015年捕捞死亡系数只有0.15,近五年平均捕捞死亡系数Fcur=0.33,单位补充量亲体量是未开发时的32.7%,不存在生长型捕捞过度,也不存在补充型捕捞过度,处于可持续开发状态。研究还探讨了水温变化引起自然死亡波动以及不同开捕年龄对澳洲鲭太平洋群系资源状况的影响。研究表明,该渔业目前开发和利用程度合理,建议使用F0.1做为管理参考点进行渔业资源的管理。     

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.     

Pacific herring, Clupea pallasi, recruitment in the Bering Sea and north-east Pacific Ocean, I: relationships among different populations

We examined recruitment and average weight-at-age time series for Pacific herring ( Clupea pallasi ) populations from the Bering Sea and north-east Pacific Ocean to determine similarities. Statistical correlation and multivariate clustering methods indicated Pacific herring populations form large-scale groups. Large year classes occur synchronously among several Pacific herring populations. Multivariate cluster analyses of recruitment and weight-at-age data indicated that Bering Sea herring populations are distinct from north-east Pacific Ocean populations. Within the NE Pacific Ocean, there appear to be three groups of herring populations: a British Columbia group, a south-east Alaska coastal group, and an outer Gulf of Alaska group. Jackknife and randomization tests indicate these groups are robust and not the result of random chance. Deviations from observed herring population groups were examined for indications of anthropogenic perturbations. The Prince William Sound herring populations did not show any strong deviations corresponding to the oil spill of 1989. There might not yet be enough data since the spill to detect changes in the recruitment or weight-at-age data since that time, particularly if oil spill effects were concentrated on the early life history stages.     

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.     

Changes in spawning stock structure strengthen the link between climate and recruitment in a heavily fished cod (Gadus morhua) stock

Atlantic cod (Gadus morhua) is one of the commercially most important fish species in the North Atlantic and plays a central role in several ecosystems. Fishing pressure has been heavy over a prolonged period and the recent decades have shown dramatic decline in abundance of many stocks. The Arcto‐Norwegian (or North‐east Arctic) cod stock in the Barents Sea is now the largest stock of Atlantic cod. Recruitment to this stock has varied extensively during the last 60 yr. There is evidence for fluctuations in climate, particularly sea temperature, being a main cause for this variability, higher temperatures being favourable for survival throughout the critical early life stages. Our studies of time series present compelling evidence for a strengthening of the climate–cod recruitment link during the last decades. We suggest this is an effect of the age and length composition of the spawning stock having changed distinctly. The age of the average spawner has decreased by more than 3 yr from between 10 and 11 in the late 1940s to 7–8 in the 1990s, average length from just above 90 cm to around 80 cm. The number of age classes contributing to the spawning stock has also decreased, while the number of length groups present increased slightly. Significant decrease in age of spawners has frequently been described for other heavily fished stocks worldwide. We therefore find it likely that the proposed mechanism of increased influence of climate on recruitment through changes in the spawning stock age and size composition is of a general nature and might be found in other systems.