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.     

Response of Scotian Shelf silver hake (Merluccius bilinearis) to environmental variability

Commercially‐exploited fish populations are not only shaped by fishing pressure, but also by the dynamics of their environment. By quantifying the influence of environmental variability, fisheries management advice can be improved and uncertainties reduced. To this end, we developed statistical models of the response of Scotian Shelf silver hake stock metrics to variability in zooplankton community composition and phenology over the past 18 years and in the physical environment since 1985. Dominant modes of variability in these pelagic habitat indicators were characterized using principal component analysis, and the relationships of silver hake condition, abundance, and recruitment to pelagic habitat variability were assessed using generalized additive models. Condition was largely modulated by the onset and duration of the spring bloom, which controls food availability. In contrast, adult abundance was governed by composition of the zooplankton community and bottom‐water temperature, which dictates the distribution of silver hake. Finally, recruitment was affected by both thermal conditions and food availability. Our results presented here form the basis for qualitative assessment of ecosystem attributes and the influence on silver hake stock productivity.     

The recruitment process of the Barents Sea capelin (Mallotus villosus) stock, 2001–2003

Oceanographic and predation processes are important modulators of fish larvae survival and mortality. This study addresses the hypothesis that immature Norwegian spring‐spawning herring (Clupea harengus), when abundant in the Barents Sea, determine the capelin reproduction success through consumption of Barents Sea capelin (Mallotus villosus) larvae. Combining a hydrodynamic model and particle‐tracking individual‐based model, a realistic spatio‐temporal overlap between capelin larvae and predatory immature herring was modelled for the summer seasons of 2001–2003. Capelin larvae originating from western spawning grounds became widely dispersed during the summer season, whereas those originating from eastern spawning grounds experienced a rapid drift into the southeastern Barents Sea. Herring caused a 3% mortality of the capelin larvae population in 2001 and a 16% mortality in 2003, but the effect of predation from herring on capelin larvae was negligible in 2002. Despite a strong capelin larvae cohort and a virtual absence of predatory herring, the recruitment from the capelin 2002 year class was relatively poor from a long‐term perspective. We show that the choice of capelin spawning grounds has a major impact on the subsequent capelin larvae drift patterns, constituting an important modulator of the capelin larvae survival. Variation in drift patterns during the summer season is likely to expose the capelin larvae to a wide range of hazards, including predation from young cod, sandeel and other predators. Such alternative predators might thus have contributed to the poor capelin recruitment during 2001–2003, leading to the collapse of the capelin stock in the subsequent years.     

Effect of environmental variability on body condition and recruitment success of Atlantic Mackerel (Scomber scombrus L.) in the Gulf of St. Lawrence

The objective of this study was to assess the effect of environmental variability on the dynamics of the Atlantic mackerel (Scomber scombrus L.) stock in the Gulf of St. Lawrence (GSL). We first described the dominant modes of physical and biological (zooplankton) variability using Principal Components Analyses of 40 variables. Two principal modes of variability were identified, a long‐term mode (15–20 yr) associated with a warming of the GSL and a second mode describing alternating cold and warm periods at a higher frequency (5–10 yr). A strong link between physical forcing and the dynamics of zooplankton species known to be important for mackerel was shown. Second, a set of Generalized Additive Models (GAM) was developed to explore how these environmental variations could influence mackerel condition (Fulton's K) and recruitment success (R_s). Optimal GAMs including variations in abundance, species composition and phenology of key copepods improved model performance by 40–50% relative to those considering only physical environmental conditions. The results are consistent with the match–mismatch hypothesis and illustrate the key role of zooplankton dynamics in modulating variations in mackerel K and R_s. Finally, this study showed that large variations in R_s could be caused by varying environmental conditions independently of the influence of stock biomass. Our results strongly indicate that the effect of environmental variability should be considered in the implementation of an ecosystem‐based approach to Atlantic mackerel stock management.     

Variation in abundance of Norwegian spring-spawning herring (Clupea harengus, Clupeidae) throughout the 20th century and the influence of climatic fluctuations

A long-term (1907–98) virtual population analysis (VPA) was made for Norwegian spring-spawning herring (NSSH), which is a huge pelagic fish stock in the north-east Atlantic. It shows that this herring stock has had large fluctuations during the last century; these fluctuations have mainly been determined by variations in the temperature of the inflowing water masses to the region. The spawning stock biomass (SSB) increased from a rather low level in the early years of this century and reached a high level of around 14 million tons by 1930. The spawning stock biomass then decreased to a level of around 10 million tons by 1940, but increased again to a record high level of 16 million tons by 1945. The stock then started to decrease and during the next 20-year period fell to a level of less than 50 000 tons by the late 1960s. Through the 1970s and 1980s, the stock slowly recovered and after the recruitment of strong year classes in 1983 and 1990–1992 the stock recovered to a spawning stock biomass of about 10 million tons. The long-term fluctuation in spawning stock biomass is caused by variations in the survival of recruits. It is found that the long-term changes in spawning stock abundance are highly correlated with the long-term variations in the mean annual temperature of the inflowing Atlantic water masses (through the Kola section) into the north-east Atlantic region. The recruitment is positively correlated with the average temperature in the Kola section in the winter months, January–April, which indicates that environmental factors govern the large-scale fluctuations in production for this herring stock.     

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.     

It’s about time: A synthesis of changing phenology in the Gulf of Maine ecosystem

The timing of recurring biological and seasonal environmental events is changing on a global scale relative to temperature and other climate drivers. This study considers the Gulf of Maine ecosystem, a region of high social and ecological importance in the Northwest Atlantic Ocean and synthesizes current knowledge of (a) key seasonal processes, patterns, and events; (b) direct evidence for shifts in timing; (c) implications of phenological responses for linked ecological‐human systems; and (d) potential phenology‐focused adaptation strategies and actions. Twenty studies demonstrated shifts in timing of regional marine organisms and seasonal environmental events. The most common response was earlier timing, observed in spring onset, spring and winter hydrology, zooplankton abundance, occurrence of several larval fishes, and diadromous fish migrations. Later timing was documented for fall onset, reproduction and fledging in Atlantic puffins, spring and fall phytoplankton blooms, and occurrence of additional larval fishes. Changes in event duration generally increased and were detected in zooplankton peak abundance, early life history periods of macro‐invertebrates, and lobster fishery landings. Reduced duration was observed in winter–spring ice‐affected stream flows. Two studies projected phenological changes, both finding diapause duration would decrease in zooplankton under future climate scenarios. Phenological responses were species‐specific and varied depending on the environmental driver, spatial, and temporal scales evaluated. Overall, a wide range of baseline phenology and relevant modeling studies exist, yet surprisingly few document long‐term shifts. Results reveal a need for increased emphasis on phenological shifts in the Gulf of Maine and identify opportunities for future research and consideration of phenological changes in adaptation efforts.     

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。     

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.     

Long-term variation in the abundance of the brown shrimp Crangon crangon (L.) population of the German Bight and possible causes for its interannual variability

Time series on Crangon crangon densities in the German Wadden Sea show a considerable degree of interannual variability, for the entire region in spring and in autumn as well as for three subareas, North Frisia, East Frisia and Elbe estuary. Across the entire survey area C. crangon density was inversely related to water depth. In autumn after the recruitment, settlement shrimp densities are correlated across a larger spatial scale (>100 km longshore), whereas in spring the overwintering adult spawning stock is uncorrelated across subareas. Interannually, extreme density variations can develop even over a short time span of 1 yr. On a large spatial scale shrimp abundance in autumn was correlated with year‐to‐year changes in physical environmental and biological parameters, winter water temperature, autumn river runoff and the winter NAO index. On a regional scale (North Frisia), density of gadoid predators was an additional component affecting shrimp stock abundance. No correlations to such parameters could be detected for the C. crangon stock in spring. Possible causes for the spring situation and additional sources for variability are discussed.     

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.     

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.     

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.     

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.     

Fluctuation in stock properties of north-east Arctic cod related to long-term environmental changes

From a historic perspective, the north‐east Arctic cod stock, which is found in the Barents Sea–Svalbard region, has been the most productive gadoid stock in the Atlantic. Variation in catch has always been large, but during the last 10–15 years catch and stock abundance have reached the lowest level on record. Three major causes of variation have been discussed: (i) stock reduction through exploitation; (ii) environmental influences on recruitment; and (iii) species interaction effects on maturation, growth and mortality. In addition, interactions among these three sources might be important. The influence of each specific factor is difficult to evaluate from incidental observations and short‐term time series. In this respect, the time series on catches and on biological and environmental information of this stock, which partly extend back to the 19th century, occupy a unique position in comparison to data on most other stocks. In this paper, fluctuations in catch and stock abundance are compared with changes in recruitment, size/age and growth. This information is discussed in view of historic variation in ecology and environment. The stock has been under particularly high exploitation pressure since the mid‐1970s. Further, large changes in growth rates and poor recruitment to the commercially exploited stock are characteristic of late 1980s and throughout the 1990s. The analysis shows that substantial long‐term variation might underlie short‐term variability, and more importantly, that long‐term changes roughly coincide with similar fluctuations in the environment. Such factors might substantially affect the relationship between spawning stock and recruitment, which is also apparent from the difference in conclusions reached by various published studies. Consequently, it is suggested that using a steady‐state perspective for the population dynamics may lead to mismanagement and to a reduction of the long‐term yield from this stock.     

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’.     

The influence of life history dynamics and environment on the determination of year class strength in North Sea herring (Clupea harengus L.)

The inter-annual variability in year class strength (1976–2000) of North Sea herring (Clupea harengus) was investigated using Paulik diagrams based on survey data and Virtual Population Analysis. The herring life cycle was split into five stages: spawning stock biomass (SSB), egg production, larvae, fish with 0 winter rings on the otolith (0-wr), 1-wr and 2-wr. Surveys were used as indices and Paulik analysis revealed relationships between stages. In 80% of the years, year class strength reflected SSB. Poorer than expected year classes were determined during the larva to 0-wr phase, whilst stronger than expected year classes were apparently determined during the 0-wr to 1-wr stage. There was no clear relationship between survival of young stages of herring and the abundance of Calanus finmarchicus but the year class strength of 0-wr and 1-wr had a negative relationship to bottom water temperature. Lower sea water temperatures in the North Sea are associated with higher Calanus abundance. The analysis shows that the strength of aberrant year classes of North Sea herring is determined between the pelagic larval and the juvenile stages.     

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.     

Impact of environmental deviations on the larval and year-class abundances in the spring spawning herring (Clupea harengus membras L.) of the Gulf of Riga (Baltic Sea) in 1947–2004

Regular weekly sampling of larvae with a Hensen net in the stationary stations on the main spawning grounds of the brackishwater herring population in the Gulf of Riga was carried out in 1947–2004. Probably due to the rapidly increasing anthropogenic eutrophication and the rising winter temperature the yearly abundance of herring larvae has significantly increased with a shift in the average abundance in 1972/1973. Correlation of the average number of herring larvae with the year-class strength was significant for the intermediate (10–16 mm, P < 0.05) and large (≥17 mm, P < 0.001) larval size groups. The monthly average air temperature in the period from January to March was the main environmental variable regulating the abundance of the year classes (P < 0.001). The dependence between the spawning stock biomass (SSB) and year-class abundance differed by climate periods. In the period of cold winters environmental conditions dominated in determination of the year-class abundance whereas the role of the SSB was nonsignificant (P > 0.05). In the period of milder winters the importance of the SSB in determination of the abundance of year classes considerably increased and it significantly correlated with the abundance of year classes (P < 0.05). The winter–spring temperatures and other climatic variables presumably control the abundance of the forming year-class during the period of the highest larval mortality mainly via the production of their food.