Recruitment,distribution boundary and habitat temperature of an arcto‐boreal gadoid in a climatically changing environment: a case study on Northeast Arctic haddock (Melanogrammus aeglefinus)

Climate change is expected to have major effects on the distribution and abundance of fish. In spite of extensive research on the topic in high‐latitude marine ecosystems, the mechanistic understanding of how temperature impacts recruitment and distribution of arcto‐boreal fish stocks remains elusive. Exemplified by an arcto‐boreal gadoid in the Barents Sea, the Northeast Arctic (NEA) haddock (Melanogrammus aeglefinus), we investigate the effect of ecosystem temperature (here temperature from a fixed reference section) on abundance and distribution boundaries between 1981 and 2008. During this time interval there has been a trend of increasing temperature in the ecosystem. We compare the ecosystem temperature with the species habitat temperature of NEA haddock (i.e., ambient temperature of the population) – two temperature approaches representing the indirect and direct environmental impacts on fish, respectively. In addition to the temperature effects, density‐dependent effects on distribution boundaries are considered. The study is based on swept area density estimates and spatial temperature data collected annually in winter surveys. We found a positive relationship between ecosystem temperature and abundance, a connection related to both direct and indirect mechanisms with short‐term and long‐term pathways. Distribution boundaries are, on a year‐to‐year basis, more related to abundance than ecosystem temperature. The long‐term trends, however, indicate a north‐eastward shift in distribution boundaries, probably indirectly related to the coinciding ecosystem temperature increase. In spite of the gradual increase in ecosystem temperature, the abundance of 4‐ to 7‐ year old NEA haddock expanded into colder waters. Thus, our results show how different the two temperature approaches may be.     

The growth of larval cod and haddock in the Irish Sea: a model with temperature,prey size and turbulence forcing

We applied a physiological individual‐based model for the foraging and growth of cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae, using observed temperature and prey fields data from the Irish Sea, collected during the 2006 spawning season. We used the model to estimate larval growth and survival and explore the different productivities of the cod and haddock stocks encountered in the Irish Sea. The larvae of both species showed similar responses to changes in environmental conditions (temperature, wind, prey availability, daylight hours) and better survival was predicted in the western Irish Sea, covering the spawning ground for haddock and about half of that for cod. Larval growth was predicted to be mostly prey‐limited, but exploration of stock recruitment data suggests that other factors are important to ensure successful recruitment. We suggest that the presence of a cyclonic gyre in the western Irish Sea, influencing the retention and/or dispersal of larvae from their spawning grounds, and the increasing abundance of clupeids adding predatory pressure on the eggs and larvae; both may play a key role. These two processes deserve more attention if we want to understand the mechanisms behind the recruitment of cod and haddock in the Irish Sea. For the ecosystem‐based management approach, there is a need to achieve a greater understanding of the interactions between species on the scale a fish stock is managed, and to work toward integrated fisheries management in particular when considering the effects of advection from spawning grounds and prey–predator reversal on the recovery of depleted stocks.     

Variability in the spawning habitat of Pacific sardine (Sardinops sagax) off southern and central California

The spatial pattern of sardine spawning as revealed by the presence of sardine eggs is examined in relation to sea surface temperature (SST) and mean volume backscatter strength (MVBS) measured by a 150 kHz acoustic Doppler current profiler (ADCP) during four spring surveys off central and southern California in 1996–99. Studies in other regions have shown that MVBS provides an excellent measure of zooplankton distribution and density. Zooplankton biomass as measured by survey net tows correlates well with concurrently measured MVBS. The high along‐track resolution of egg counts provided by the Continuous Underway Fish Egg Sampler (CUFES) is a good match to the ADCP‐based data. Large interannual differences in the pattern and density of sardine eggs are clearly related to the concurrently observed patterns of surface temperature and MVBS. The strong spatial relationship between sardine eggs and MVBS is particularly evident because of the large contrast in zooplankton biomass between the 1998 El Niño and 1999 La Niña. The inshore distribution of sardine spawning appears to be limited by the low temperatures of freshly upwelled waters, although the value of the limiting temperature varies between years. Often there is an abrupt offshore decrease in MVBS that is coincident with the offshore boundary of sardine eggs. Possible reasons for this association of sardine eggs and high zooplankton biomass include an evolved strategy that promotes improved opportunity of an adequate food supply for subsequent larval development, and/or adult nutrient requirements for serial spawning. Hence, the distribution of these parameters can be used as an aid for delineating the boundaries of sardine spawning habitat.     

Multiple spawning events promote increased larval dispersal of a predatory fish in a western boundary current

Transport of larvae by ocean currents is an important dispersal mechanism for many species. The timing and location of spawning can have a large influence on settlement location. Shifts in the known spawning habitat of fish, whether due to climate or the discovery of new spawning stock, can influence the distribution of juveniles and our understanding of connectivity. The globally distributed species; Pomatomus saltatrix, is one such example where a previously unrecognised summer spawning event and a more southern latitudinal extent was recently reported for the southwest Pacific population. Although restrictions are in place to protect the traditional spawning event, the importance of the newly recognised summer spawning event is uncertain. Here, we investigate larval dispersal of P. saltatrix using particle tracking simulations to identify the contributions of the different spawning events to settlement. By modelling dispersal of larvae released in northern and mid‐latitude regions over the Austral spring and summer, we show that the newly recognised mid‐latitude summer spawning event contributes over 50% of the larvae reaching southern latitudes. This is due to a reduced (1–2 days) pelagic larval duration (associated with temperature), resulting in reduced larval mortality, and the seasonal (summer) strengthening of the East Australian Current (EAC) transporting particles ~50 km further south. These findings demonstrate that in dynamic boundary current systems such as the EAC, the final settlement location of larvae that are transported by ocean currents can vary considerably depending on the timing and location of spawning and that multiple spawning events are important for maximum dispersal.     

Evaluation of a spatially sex‐specific assessment method incorporating a habitat preference model for blue marlin (Makaira nigricans) in the Pacific Ocean

Blue marlin, widely distributed throughout the Pacific Ocean, are sexually dimorphic, have certain preferred habitats, and migrate seasonally. These characteristics have been ignored in previous stock assessment models. A population dynamics model that includes spatial structure, and sex and age structure was therefore constructed and fitted to fisheries data for blue marlin, along with information on the relative density of the population over space derived from a habitat preference model that uses the oceanographic and biological variables sea‐surface temperature, mixed layer depth, sea‐surface height anomaly, and chlorophyll‐a concentration. Monte Carlo simulation was then used to examine the estimation performance of the stock assessment method. Estimates of management‐related quantities including current spawning stock biomass are substantially biased when the assessment method ignores seasonal movement and sexual dimorphism. We also found that (i) uncertainty about the relationship between catch rate and abundance influences estimation performance to a larger extent than uncertainty in catches, (ii) the outcomes of the assessment are sensitive to the values assumed for natural mortality and stock‐recruitment steepness, and (iii) the ratio of current spawning stock biomass to that at pre‐exploitation equilibrium appears to be the most robust among the quantities considered. We conclude that assessment methods for blue marlin in the Pacific Ocean need to take account of seasonal migration and sex structure to improve stock assessments.     

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.     

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.     

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.     

Common trends in recruitment dynamics of north‐east Atlantic fish stocks and their links to environment,ecology and management

Recruitment dynamics are challenging to assess or predict because of the many underlying drivers that vary in their relevance over time and space. Stock size, demographic and trait composition, condition and distribution of spawning fish and the spatio‐temporal dynamics of trophic and environmental interactions all influence recruitment processes. Exploring common patterns among stocks and linking them to potential drivers may therefore provide insights into key mechanisms of recruitment dynamics. Here, we analysed stock‐recruitment data of 64 stocks from the north‐east Atlantic Ocean for common trends in variation and synchrony among stocks using correlation, cluster and dynamic factor analyses. We tested common trends in recruitment success for relationships with large‐scale environmental processes as well as stock state indicators, and we explored links between recruitment success and demographic, environmental and ecological variables for a subset of individual stocks. The results revealed few statistically significant correlations between stocks but showed that underlying common trends in recruitment success are linked to environmental indices and management indicators. Statistical analyses confirmed previously suggested relationships of environmental–ecological factors such as the subpolar gyre and Norwegian coastal current with specific stocks, and indicated a large relevance of spawning stock biomass and demographics, as well as predation, whereas other suggested relationships were not supported by the data. Our study shows that despite persistent challenges in determining drivers of recruitment due to poor data quality and unclear mechanisms, combining different data analysis techniques can improve our understanding of recruitment dynamics in fish stocks.     

Dual effects of reversed winter–spring temperatures on year‐to‐year variation in the recruitment of chub mackerel (Scomber japonicus)

To clarify the effects of temperature on the recruitment of chub mackerel (Scomber japonicus) in the North Pacific, we investigated the influence of winter surface temperature (WST) on spawners at the time of maturity around the spawning grounds and the influence of ambient spring temperature on larvae using estimated temperature (ET) obtained from particle tracking experiments. We found a significant positive correlation between ET approximately 10 days following hatching and the recruitment per spawning stock biomass (RPS) after 2000. The closer (more meandering) the Kuroshio Current (KC) axis was in relation to the spawning ground, the higher (lower) the spring surface temperature and the higher (lower) RPS was in the spawning ground. In contrast, WST inside KC near the maturity/spawning ground was significantly negatively correlated with RPS. A significant negative correlation between the temperatures in winter and spring was detected in the area after 2000, when the conditions of the Pacific decadal oscillation index and the stability of the Kuroshio Extension were synchronous, indicating that KC shifted northward during this time. The reversed temperature pattern was consistent with the winter–spring movement of KC axis in the offshore direction and was correlated with the winter–spring difference in the intensity of the Aleutian low. These results suggest that the annual variation in chub mackerel recruitment after 2000 was strongly affected by the combined effects of ambient temperature because of the reversal of conditions that occurred between winter and spring around the maturity/spawning ground, which was related to the KC path.     

Environmentally driven forecasts of northern rock sole (Lepidopsetta polyxystra) recruitment in the eastern Bering Sea

Northern rock sole recruitment in the eastern Bering Sea has been hypothesized to (a) depend on wind‐driven surface currents linking spawning and nursery areas, (b) be density‐dependent, and (c) be negatively impacted by cold bottom temperatures over a large nursery area during the first summer of life. A suite of models was developed to test these hypotheses. Data included 32 years of recruitment and spawning biomass estimates derived from a stock assessment model and wind and temperature indices customized to the environmental exposure of age‐0 northern rock sole in the eastern Bering Sea. The predictive ability of the models was evaluated, and the models were used to forecast recruitment to age‐4 for recent year classes which are poorly retained by the standard multi‐species bottom trawl survey gear. Models which included wind and temperature indices performed better than a naïve forecast based on the running mean. The best‐performing model was a categorical model with wind and temperature thresholds, which explained 49% of the variation in recruitment. Ricker models performed more poorly than models without a spawning biomass term, providing no evidence that recruitment is related to stock size. The models forecast higher recruitment for the most recent year classes (2015–2018) than for prior year classes with observed poor recruitment (2006–2013). These environment‐based recruitment forecasts may improve recruitment estimates for the most recent year classes and facilitate study of the effects of future climate change on northern rock sole population dynamics.     

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.     

Using a biophysical model to investigate connectivity between spawning grounds and nursery areas of King George whiting (Sillaginodes punctatus: Perciformes) in South Australia’s gulfs

Many demersal marine fish species depend on a dispersive larval stage that connects geographically discrete sub‐populations. Understanding connectivity between these sub‐populations is necessary to determine stock structure, which identifies the appropriate spatial scale for fishery management. Such connectivity is poorly understood for King George whiting (Sillaginodes punctatus; Perciformes) in South Australia's gulf system, even though spawning grounds and nursery areas are adequately defined. In response to declines in commercial catches and estimated biomass, this study aimed to determine the most important spawning grounds and nursery areas to recruitment, and the connectivity between them. A biophysical model was seeded with particles according to the distribution and density of eggs throughout the spawning area in 2017 and 2018. Despite inter‐annual differences in the origins of particles, dispersal pathways and predicted settlement areas remained consistent between years. Predicted settlement was generally highest to nursery areas only short distances from regional spawning grounds, consistent with previous hydrodynamic models. However, the model also predicted that spawning in one region could contribute to recruitment in an adjacent region later in the spawning season, which aligned with the breakdown of thermohaline fronts at the entrance of each gulf. The connectivity between spawning grounds and nursery areas predicted by the model is supported by spatio‐temporal patterns in the otolith chemistry of pre‐flexion larvae and settled juveniles. Consequently, the most parsimonious explanation is that the populations of King George whiting in South Australia's gulf system constitute a single, panmictic stock, which has implications for fishery management.     

北部湾带鱼空间聚散变化特征

本研究利用 2006―2018 年北部湾底拖网渔业资源调查的带鱼(Trichiurus haumela)资源密度数据, 通过标准差椭圆(standard deviational ellipse, SDE)分析带鱼空间分布格局, 利用广义可加模型(generalized additive model, GAM)研究带鱼资源密度与环境因子叶绿素 a (Chl-a)、海表水温异常值、水深、离岸距离、经度和纬度的关系, 为北部湾带鱼资源的可持续利用和保护提供科技支撑。研究结果显示, 北部湾带鱼资源密度分布呈逐年集聚现象, 渔场分布方向受海底地貌影响呈现西南–东北向, 夏季资源密度重心呈西南–东北向移动, 冬季资源密度重心呈南–北向移动, 且冬季带鱼资源密度重心较夏季更偏东北向; 春、秋季为带鱼主要的产卵季节, 其资源密度重心位于夏、冬季带鱼渔场中部。研究表明北部湾带鱼资源密度的逐年下降是导致其空间分布逐渐集聚的主要原因。Chl-a 对带鱼资源密度和空间分布均有影响, 水深、经度和海表水温异常值对带鱼资源密度有影响, 但对其空间分布无直接影响。     

福建平潭岛海域游泳生物生态和资源保护

1990年4个季节月调查，平潭岛海域有游泳生物88种，其中鱼类62种，甲壳类22种，头足类4种。鱼类区系属于印度－西太平洋的中国－日本亚区。种类数的季节分布呈秋季＞夏季＝冬季＞春季，生物量匠优势种均为龙头鱼和带鱼，生物量的季节分布呈冬季＞秋季＞夏季＞春季，密度呈夏季＞春季＞冬季＞秋季。平均月生物量和密度岛北部海域分别是岛南部海域的2.55倍和1.32倍。本文分析了种类组成与环境的关系，并提出了渔业资源保护的建议：（1）保护幼鱼的索饵场和生殖群体的产卵场；（2）适当适长伏季休渔期；（3）定置网的最适捕捞力量估算为12912张。     

Quantifying the influence of geography and environment on the northeast Atlantic mackerel spawning distribution

Mackerel (Scomber scombrus) in the northeast Atlantic have shown changes in distribution at certain times of the year, which have impacted their exploitation and management. In this study, mackerel spawning habitat over 21 recent years was characterised using generalised additive modelling, based on spatial egg density data collected every third year during targeted ichthyoplankton surveys. Mackerel spawning distribution was found to depend primarily on geographical variables (coordinates and bottom depth), with preferred spawning locations on the shelf‐edge from the north of the Iberian peninsula to the west of Scotland, with a maximum west of Ireland. Environmental drivers had a lesser influence on egg distribution. Dome shaped relationships were found with temperature and mixed layer depth, with respective optimum at 13°C and around –300 m. The model was used to reconstruct maps of the potential habitat (areas where conditions were suitable, but not necessarily used, for spawning). Little changes were observed over the years in the potential habitat, suggesting that the expansion of the egg distribution (realised habitat) was not triggered by changes in the environmental variables investigated. Little evidence was found for density‐dependent habitat selection. There was a tendency for mackerel to make more use of areas of lesser suitability in years with large stock size (1992 and 2010). This pattern, however, broke down in 2013, when stock size was the highest, as spawning occurred very south and concentrated in the most suitable habitat.     

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.     

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.     

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.     

Transport of larvae and juvenile fish into central and northern Norwegian waters

The distribution of juvenile fish in Norwegian waters in spring and summer provides evidence that some of these are transported into Norwegian coastal waters from spawning grounds outside the ecosystems which comprise the central and northern Norwegian coast, the Barents Sea and the Spitsbergen area. The relevant species are capelin (Mallotus villosus Müller), haddock (Melanogrammus aeglefinus L.), herring (Clupea hareri' gus L.) and saithe (Pollachius virens L.). A substantial part of the saithe and haddock stock along the central and northern Norwegian coast is considered to be recruited from spawning grounds in the northern North Sea. A contribution of these species from the spawning grounds around the Faeroe Islands and west of Scotland may also occur in some years. Herring from the northern North Sea are found scattered off the Norwegian coast north to 70^oN. It seems likely that pelagic species such as capelin and herring are transported by the prevailing currents from Iceland.