Combined effects of temperature and fishing mortality on the Barents Sea ecosystem stability

Temporal variability in abundance and composition of species in marine ecosystems results from a combination of internal processes, external drivers, and stochasticity. One way to explore the temporal variability in an ecosystem is through temporal stability, measured using the inverse of the coefficient of variation for biomass of single species. The effect of temperature and fisheries on the variability of the Barents Sea food web is still poorly understood. To address this question, we simulate the possible dynamics of Barents Sea food web under different temperature and fishery scenarios using a simple food-web model (Non-Deterministic Network Dynamic [NDND]). The NDND model, which is based on chance and necessity (CaN), defines the state space of the ecosystem using its structural constraints (necessity) and explores it stochastically (chance). The effects of temperature and fisheries on stability are explored both separately and combined. The simulation results suggest that increasing temperature has a negative effect on species biomass and increasing fisheries triggers compensatory dynamics of fish species. There is a major intra-scenario variability in temporal stability, while individual scenarios of temperature and fisheries display a weak negative impact and no effect on stability, respectively. However, combined scenarios indicate that fisheries amplify the effects of temperature on stability, while increasing temperature leads to a shift from synergistic to antagonistic effects between these two drivers.     

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.     

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.     

Effects of temperature, wind and spawning stock biomass on recruitment of Arcto-Norwegian cod

Sea temperature has earlier been shown to have a large influence on the recruitment of Arcto-Norwegian cod, Gadus morhua. We here hypothesize that this linkage is partly due to the direct effect of temperature on larval and juvenile growth. Secondly, temperature acts as a proxy for both biotic and abiotic factors influencing recruitment. Indices of abundance of early juvenile cod (2–3 months old), 0-group cod (4–5 months old) and 3-year-old cod are analysed in more detail against the environmental temperature, wind stress components, wind-induced turbulent energy and the spawning stock biomass. To deal with autocorrelation, non-stationar-time and nun-normality, which complicate a statistical time series analysis, randomization and Box-Jenkins methods are applied. In addition to the important effect of high sea temperature during the early life stage in forming strong year classes, the results show that the spawning stock biomass is nearly as important. Also, alongshore southerly wind stress anomalies during the period of pelagic drift (from April through summer) and offshore wind stress anomalies during egg and early larval stages (in April) act favourably on recruitment. The beneficial effect of southerly wind anomalies could he linked partly to high temperature, but the flux of zooplankton-rich water from the Norwegian Sea into the feeding areas of the Barents Sea may also be increased. The favourable influence of offshore winds in April is less predominant and causal links are also less clear; possible explanations for this might be increased offshore spreading of eggs and early larvae, resulting in reduced risk of predation, and increased compensation inflow of intermediate Norwegian Sea water which, in this restricted period of time, has a high concentration of spawning copepods suitable as prey for the developing cod larvae.     

Trophic role of Atlantic cod in the ecosystem

As the world's oceans continue to undergo drastic changes, understanding the role of key species therein will become increasingly important. To explore the role of Atlantic cod ( Gadus morhua Gadidae) in the ecosystem, we reviewed biological interactions between cod and its prey, predators and competitors within six ecosystems taken from a broad geographic range: three are cod-capelin ( Mallotus villosus Osmeridae) systems towards cod's northern Atlantic limit (Barents Sea, Iceland and Newfoundland–Labrador), two are more diverse systems towards the southern end of the range (North Sea and Georges Bank–Gulf of Maine), and one is a species-poor system with an unusual physical and biotic environment (Baltic Sea). We attempt a synthesis of the role of cod in these six ecosystems and speculate on how it might change in response to a variety of influences, particularly climate change, in a fashion that may apply to a wide range of species. We find cod prey, predators and competitors functionally similar in all six ecosystems. Conversely, we estimate different magnitudes for the role of cod in an ecosystem, with consequently different effects on cod, their prey and predator populations. Fishing has generally diminished the ecological role of cod. What remains unclear is how additional climate variability will alter cod stocks, and thus its role in the ecosystem.     

Comparison of factors affecting recruitment variability of walleye pollock Theragra chalcogramma in the Pacific Ocean and the Sea of Japan off northern Japan

The Japanese Pacific stock (JPS) and the northern Japan Sea stock (JSS) of walleye pollock Theragra chalcogramma are mainly distributed in the Pacific Ocean and the Sea of Japan off northern Japan, respectively. This paper summarizes and compares the factors affecting the recruitment variability of these two stocks. Spawning season is from December to March for both stocks. JPS recruitment has a positive relationship with the water temperature in January and February, whereas that of JSS has a negative relationship with the water temperature in January, February, and April. One possible reason for this is that pollock larvae have an optimum growth temperature of approximately 5 °C in the field. Drift of early life stages also appears to be an important influence on the recruitment of both stocks. Because the current generated by the northwest wind carries eggs of JPS into the main larval nursery ground, JPS recruitment is enhanced in years when the northwest wind is predominant in February. On the other hand, early life stages of JSS are transported into the nursery ground by the Tsushima Warm Current. However, this current also carries early life stages into the Sea of Okhotsk and offshore, resulting in poor JSS recruitment in years when this current is strong in March. In contrast to JPS, the recruitment of which is significantly impacted by cannibalism, young pollock have not been found in the stomachs of adult JSS. Warm temperatures in the Sea of Japan seem to induce the separation of young and adult pollock, and the shape of the stock–recruitment relationship also suggests that cannibalism is not important for JSS. Based on this knowledge, and on the hatch date distributions of larvae and juveniles, we propose mechanisms that can explain the recruitment fluctuations for JPS and JSS pollock.     

Climate impacts on the landings of Indian oil sardine over the south‐eastern Arabian Sea

The landings of Indian oil sardine (Sardinella longiceps, Clupeidae) along the south‐eastern Arabian Sea are about 43.8% of total Indian oil sardine production. The annual landings of this species exhibit large‐scale variability with prolonged years of surplus or deficit landings without identified reason. Evaluating Indian oil sardine landings along the Kerala coast during 1961–2017 in relation to environmental variations, we have elucidated a putative link between variability in landings versus environmental parameters and climate indices. The variables examined in this study, such as salinity and temperature along with physical indices such as upwelling and mixed layer depth (MLD) of the ocean help to propose a mechanism to temporal variability in the landings of Indian oil sardine. Colder temperature and timely intense upwelling lead to nutrient enrichment in the surface water, which promotes the growth of phytoplankton (chl‐a) and thereby food availability to Indian oil sardine are found during years with surplus catch. Less saline surface waters and shoaling of MLD at these times could lead to the aggregation of fish at particular depths and thereby a good catches. The reverse mechanism, such as more surface saline water, warm temperature, downwelling or weak upwelling, and less nutrient enrichment, leads to deficit landings. Further, it was noticed that the Pacific decadal oscillation and Atlantic multidecadal oscillation have a more pronounced impact on Indian oil sardine landings over the coast of south‐eastern Arabian Sea than previously reported ENSO associated impacts. All these point towards climate change implications for the Indian oil sardine fishery.     

Population structure of the deep-sea shrimp (Pandalus borealis) in the north-east Atlantic based on allozyme variation

In order to elucidate the population structure of the deep-sea shrimp (Pandalus borealis) in the NE Atlantic, 32 subsamples and 3 865 individuals were analysed for allozymic variation. They were caught at various locations in the Barents Sea, in waters off Svalbard, Jan Mayen and Iceland, and in fjords along the Norwegian coast. Only three enzymes (malate dehydrogenase, phosphoglucomutase and glucosephosphate isomerase) of the 22 initially tested showed a combination of gel images that could be interpreted with confidence and allozymic variation. The locus coding for malate dehydrogenase was by far the most polymorphic. Samples caught within the Barents Sea and in the Svalbard area showed no significant heterogeneity in allele frequencies, supporting earlier suggestions of only one population of P. borealis in the Barents Sea. Genetic differentiation was found, however, between Norwegian fjords and the Barents Sea, and among fjords.     

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.     

Marine biogeochemical response to a rapid warming in the main stream of the Kuroshio in the western North Pacific

Impact of climate change on marine biogeochemical parameters and ecosystem is one of the important issues of our environment. Direct evidence of marine pelagic ecosystem changes is found with warming of sea water and sea‐level rise in the main stream of the Kuroshio in the East China Sea and the western North Pacific during these three decades based on the analysis of long‐term comprehensive hydrographic observations. In terms of annual mean, the warming rate of surface air temperature and sea surface temperature ranged from 0.15 to 0.21°C per decade in and around the main stream of the Kuroshio in the East China Sea, which exceed the global mean warming rate of 0.128 ± 0.026°C per decade during the period from 1956 to 2005 reported in IPCC 2007. One of the features in this rapid warming region is an increase of number of Pterosagitta draco, a cosmopolitan warm‐water zooplankton. Biogeochemical parameters, such as wet weight of zooplankton, plant pigment and nutrients concentration in the upper 200 m have been decreasing while dissolved oxygen content and seawater temperature have been increasing in the upper 200 m in the main stream of the Kuroshio in the East China Sea. These observed linear trends of the biogeochemical parameters would be foresights for temperate oceans in the future.     

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。     

Modelling the advection and diffusion of eggs and larvae of Greenland halibut (Reinhardtius hippoglossoides) in the north-east Arctic

Since the late 1980s there has been considerable uncertainty in recruitment levels of the north‐east Arctic stock of Greenland halibut (Reinhardtius hippoglossoides). The abundance of several year classes, originally considered very low at 0–3 yr age, appeared higher than expected at the age of 6+. This may be due to poor targeting of recruitment surveys of the younger year classes. The present work considers the transport and dispersion of eggs and larvae of Greenland halibut by numerical modelling in order to predict the locations of the initial recruitment grounds. Current fields from a 3D baroclinic hydrodynamic model are fed into a Lagrangian particle‐tracking model developed for the Barents Sea area. The particles are released into the current at the spawning field along the shelf slope from Lofoten to Bear Island (69–75°N). Vertically, the particles can follow a predefined depth‐by‐age curve or be kept at a fixed depth. This model system is used for different years to examine changes in the drift pattern. The results indicate that spawning location, transport depth and inflowing activity to the Barents Sea are important factors influencing the distribution of juveniles.     

Salish Sea Chinook salmon exhibit weaker coherence in early marine survival trends than coastal populations

Identifying factors that influence anadromous Pacific salmon (Oncorhynchus spp.) population dynamics is complicated by their diverse life histories and large geographic range. Over the last several decades, Chinook salmon (O. tshawytscha) populations from coastal areas and the Salish Sea have exhibited substantial variability in abundance. In some cases, populations within the Salish Sea have experienced persistent declines that have not rebounded. We analyzed a time series of early marine survival from 36 hatchery Chinook salmon populations spanning ocean entry years 1980–2008 to quantify spatial and temporal coherence in survival. Overall, we observed higher inter‐population variability in survival for Salish Sea populations than non‐Salish Sea populations. Annual survival patterns of Salish Sea populations covaried over smaller spatial scales and exhibited less synchrony among proximate populations relative to non‐Salish Sea populations. These results were supported by multivariate autoregressive state space (MARSS) models which predominantly identified region‐scale differences in survival trends between northern coastal, southern coastal, Strait of Georgia, and Puget Sound population groupings. Furthermore, Dynamic Factor Analysis (DFA) of regional survival trends showed that survival of southern coastal populations was associated with the North Pacific Gyre Oscillation, a large‐scale ocean circulation pattern, whereas survival of Salish Sea populations was not. In summary, this study demonstrates that survival patterns in Chinook salmon are likely determined by a complex hierarchy of processes operating across a broad range in spatial and temporal scales, presenting challenges to the management of mixed‐stock fisheries.     

The importance of episodic weather events to the ecosystem of the Bering Sea shelf

Climate variability on decadal time scales is generally recognized to influence high‐latitude marine populations. Our recent work in studying air–sea interactions in the Bering Sea suggests that interannual to decadal climate variability is important through its modulation of the frequencies and magnitudes of weather events on intraseasonal time scales. We hypothesize that it is these weather events that directly impact the marine ecosystem of the Bering Sea shelf. The linkages between the event‐scale weather and the ecosystem are illustrated with three examples: walleye pollock (Theragra chalcogramma), Tanner crabs (Chionoecetes bairdi), and coccolithophorid phytoplankton (Emiliania huxleyi). We hypothesize that the strong recruitment of walleye pollock that occurred in 1978, 1982, and 1996 can be attributed in part due to the seasonably strong storms that occurred in the early summer of those years. These storms caused greater than normal mixing of nutrients into the euphotic zone which presumably led to sustained primary productivity after the spring bloom and, possibly, enhanced prey concentrations for pollock larvae and their competitors. Recruitment of Tanner crab was particularly strong for the 1981 and 1984 year‐classes. These years had periods of prominent east wind anomalies along the Alaska Peninsula during the previous winter. Such winds promote flow through Unimak Pass, and hence an enhanced flux of nutrient‐rich water onto the shelf. This mechanism may have ultimately resulted in favorable feeding conditions for Tanner crab larvae. Finally, an unprecedented coccolithophorid bloom occurred over the Bering Sea shelf in the summer of 1997. This summer featured lighter winds and greater insolation than usual after a spring that included a very strong May storm. This combination brought about a warm, nutrient‐poor upper mixed layer by mid‐summer. This provided a competitive advantage for coccolithophorid phytoplankton in 1997 and to a lesser extent in 1998. Unusually high concentrations of coccolithophores persisted for the following two years although physical environmental conditions did not remain favorable. While slow variations in the overall aspects of the physical environment may be important for setting the stage, we propose that the significant multi‐year adjustments in the marine ecosystem of the Bering Sea shelf are more directly caused by major air–sea interaction events on intraseasonal time scales.     

Modeling the impact of climate variability on Black Sea anchovy recruitment and production

The connection of climate variability with anchovy spawning and recruitment in the Black Sea in particular, and other ecosystems in general, was studied using a two‐way coupled lower trophic level and anchovy bioenergetics model. Climate variability was represented by a 50‐yr time series of daily temperature and vertical mixing rates with stochastic variations. Temperature was found to be the dominant factor influencing early life stages and hence population dynamics of Black Sea anchovy as marked by a high correlation of anchovy egg production and recruitment success in response to changes in temperature. Each decrease of 2°C in summer mean temperatures resulted in a delay in the timing of egg production of between 12 and 19 days. Water temperatures in the spawning season had a greater influence than the number of available spawning females on the intensity of egg production. Anchovy recruitment was similarly influenced by temperature, with decreased temperatures resulting in a significant delay in the onset of peak recruitment during the fall by 21–38 days. Also, recruitment numbers in December decreased by about 20% with decreasing temperatures. The impact of temperature on production was slightly diminished by the impact of vertical mixing. The strong linkage of climate variability with anchovy spawning and recruitment has an important prediction potential for short‐term anchovy stock estimations, which may serve fisheries management purposes.     

Use of reflex indicators for measuring vitality and mortality of the snow crab (Chionoecetes opilio) in captivity

The snow crab (Chionoecetes opilio) is one of the most important commercial crabs in the world; it is heavily exploited in Atlantic Canada, Alaska, the Sea of Japan and the Barents Sea. Catches in the Barents Sea north of Norway have increased dramatically in the last decade. Most of the world's catch is processed, frozen and exported overseas. However, recently there has been considerable interest in exporting live snow crab, particularly in Norway. The stress of capture and live transport can result in significant mortalities. In order to establish a live export industry for snow crab, the welfare of the animal must be monitored throughout all the steps of the live transport process. In this study the reactions of snow crabs exposed to increasing periods of air exposure were measured in terms of reflex indicators, incidence of mortality, blood lactate levels and blood protein and haemocyanin. Although this is not a specific live holding or live transport process the aim was to test the suitability of reflex indicators to reflect vitality (stress) and not just to predict mortality. This would be compared with traditional blood biochemistry techniques for measuring crustacean stress. The study demonstrated that the reflex index score (RIS) is suitable to assess the vitality of snow crab. Longer air exposure periods render higher mortality rates and less vital individuals. The authors believe using vitality reflex indicators would be a suitable way of measuring crab welfare during the live holding and transport process.     

Variability of oceanographic and meteorological conditions in the northern Alboran Sea at seasonal,inter‐annual and long‐term time scales and their influence on sardine (Sardina pilchardus Walbaum 1792) landings

Time series of European sardine (Sardina pilchardus) landings from 1962 and environmental variables from 1978 in the northern Alboran Sea are analysed. European sardine spawns in the northern Alboran Sea from mid‐autumn to late winter at a temperature range slightly higher than the one observed in the nearby Eastern North Atlantic and the North Western Mediterranean. Individuals hatched during autumn and winter are incorporated to the fishery during the following summer and autumn producing the maximum annual landings. These landings show both a decreasing long‐term trend and a strong inter‐annual variability. Although further research is needed, the warming trend of sea surface temperature and the decrease in upwelling intensity inferred from empirical orthogonal function (EOF) analyses could have some influence on the negative trends of sardine landings. The inter‐annual variability of sardine abundance seems to be related to the wind intensity at a local scale, the second principal component of the chlorophyll concentration and the sardine abundance during the preceding year. If the inter‐annual variability is considered, a linear model including these three variables with a one‐year time lag allows to explain 79% of the sardine landings variance. If the negative linear trend is also considered, the model explains 86% of the variance. These results indicate that the body condition of spawners, linked to the food availability during the preceding year, is the main factor controlling the recruitment success. The possibility of predicting sardine landings 1 year in advance could have important implications for fishery management.     

冬季黄、东海水文状况及其变化

IntroductionThe China Sea is divided into four sea regions(Fig.1) :the Bohai Sea(BS) ,the Yellow Sea(YS) ,theEast China Sea(ECS) and South China Sea(SCS) .The Bohai Sea,the Yellow Sea and the shelf of the EastChina Sea together form a broad epicontinental Sea.The average depth of Bohai Sea is18m and that of Yellow Sea is4 4m.Together,the Bohai Sea and Yel-low Sea form a gulf opening to the ECS.A shallow trough runs through the gulf and can be traced south tothe northern end o…     

不同强度拉尼娜事件下东海鲐栖息地的时空变动

鲐(Scomber japonicus)是一种重要的经济鱼种, 其栖息地环境易受到气候变化的影响。本研究依据 1950— 2015 年 2.5 m、25 m、50 m 水层温度数据以及 Ni?o3.4 指数, 计算东海鲐栖息地适宜性指数(HSI), 对比分析不同强度拉尼娜事件期间鲐栖息地的时空变动。结果表明, 拉尼娜事件期间鲐渔场适宜栖息地面积高于正常气候条件。 弱拉尼娜事件和中强度拉尼娜事件期间, 鲐渔场各水层温度偏低, 适宜的温度面积比例较高, 栖息地适宜性较高; 而强拉尼娜事件期间, 鲐渔场各水层温度偏高, 适宜的温度面积比例较低, 栖息地适宜性较低。相较于强拉尼娜事件, 弱拉尼娜事件和中强度拉尼娜事件期间, 适宜栖息地的经度重心偏东, 纬度重心偏南, 不适宜栖息地的经度重心偏西, 纬度重心偏北。此外, 鲐渔场各水层最适宜温度等值线在强拉尼娜事件下偏西北方向, 推测不同强度拉尼娜下鲐栖息地空间分布差异, 可能是由于各水层最适宜温度显著的月间分布差异所致。研究表明, 不同强度拉尼娜事件对东海鲐栖息地的影响具有显著差异。     

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.