Mid-latitude wind stress: the energy source for climatic shifts in the North Pacific Ocean

Analyses of atmospheric observations in the North Pacific demonstrate extensive decadal-scale variations in the mid-latitude winter surface wind stress. In the decade after 1976 winter, eastward wind stress doubled over a broad area in the central North Pacific and the winter zero wind stress curl line was displaced about 6° southward. This resulted in increased southward Ekman transport, increased oceanic upwelling, and increased turbulent mixing as well as a southward expansion of the area of surface divergence. All these factors contributed to a decadal winter cold anomaly along the subtropical side of the North Pacific Current. In summer the cold anomaly extended eastward, almost reaching the coast of Oregon. The increased gradient in wind stress curl and southward displacement of the zero curl line also resulted in an increase in total North Pacific Current transport, primarily on the Equator side of this Current. Thus, surface water entering the California Current was of more subtropical origin in the post-1976 decade. Southward (upwelling favourable) wind stress and sea surface temperature (SST) in the area off San Francisco exhibit at least three different types of decadal departures from mean conditions. In association with the 1976 climatic shift, marine fishery production in the Oyashio, California and Alaska Currents altered dramatically, suggesting that these natural environmental variations significantly alter the long-term yields of major North Pacific fisheries.     

The influence of diel vertical migration on zooplankton transport and recruitment in an upwelling region: estimates from a coupled behavioral-physical model

Diel vertical migration (DVM) is a common zooplankton behavior in which organisms reside in surface or near‐surface waters at night and at deeper depths during the day. In many upwelling regions, DVM reduces the transport of organisms away from the region. It is unclear, however, what role DVM plays in recruitment (the arrival of larvae or juveniles to locations where they will become reproducing adults) to upwelling regions. In this study, we estimate the influence of DVM on zooplankton transport, the level of recruitment of locally produced propagules (self‐recruitment), and sources of recruits in the upwelling region near Monterey Bay, California, by simulating the trajectories of fixed‐depth and vertically migrating organisms with a drifter‐tracking algorithm driven by climatological velocity fields from a three‐dimensional hydrodynamic model. Our simulations suggest that DVM into subsurface poleward and onshore currents during the day does not fully compensate for equatorward and offshore transport in the surface Ekman layer at night and does not retain zooplankton in the Monterey Bay region. Our simulations also suggest that DVM decreases the ability of zooplankton to return to the region after being transported away and shifts source regions for recruits closer to the bay. While DVM does not appear to substantially increase the potential for self‐recruitment to the region, this study indicates that other mechanisms, such as transport during non‐upwelling periods, continuous transport below the surface, increases in mean transport depth over time, or seasonal changes in hydrography, may still enable relatively high levels of self‐recruitment to this highly advective region.     

Population dynamics of Calanus chilensis in the Chilean Eastern Boundary Humboldt Current

A study of the population dynamics of Calanus chilensis was carried out in the coastal area of the Mejillones Peninsula (23°S), which is an upwelling site in the Eastern Boundary Humboldt Current. Information was derived from time-series sampling (15 day interval) at three stations, and from two broad-scale cruises: under active upwelling, and under 1997/1998 El Niño conditions. Calanus chilensis showed continuous reproduction throughout the year. In the study area, upwelling takes place intermittently year-round, suggesting that Calanus production is not limited by food. Advection played an important role in structuring the spatial distribution of Calanus , both during upwelling and under El Niño conditions. Population losses from upwelling and offshore transport may be compensated by rapid turnover rates of cohorts, growing at temperature-dependent rates and hence allowing population recovery, even in abnormally warm waters during El Niño conditions.     

Relationships between oceanic conditions and growth of Chinook salmon (Oncorhynchus tshawytscha) from California, Washington, and Alaska, USA

We model age‐specific growth rates of Chinook salmon (Oncorhynchus tshawytscha) with two life‐history behaviors from Alaska (i. Situk and ii.Taku Rivers), Puget Sound, Washington (iii., iv. Skagit River), and California (v. Smith River) relative to oceanic conditions in those regions. By analyzing over 20 yr of biological and physical data from the NE Pacific downwelling, upwelling, and transition zones, we are able to determine the factors affecting growth across much of the species’ range and between life‐history behaviors. With scale increment data from returning fish, we use path analysis and partial least squares regression to quantify the relationships between growth and regional‐ and large‐scale oceanic conditions (e.g., sea level height, sea surface temperature, upwelling). Alaskan fish with both ocean‐ and stream‐type behaviors were fit best by the environmental data from the winter in Alaska waters. Specifically, coastal and gyre factors such as sea surface temperature, river flow, and Ekman pumping positively correlated to growth, indicating a productive and strong Alaska Current promoted growth. Growth of fish from California was fit by local factors such as increased upwelling, lower coastal sea surface temperature, and wind stresses during summer and spring, indicating a productive and strong California Current promoted growth. For Puget Sound, Washington, growth of fish that migrate to sea in their first year was generally negatively correlated to a strong California Current. Puget Sound fish that spend a year in freshwater before migrating to sea were modeled well with environmental data from their source region for the first 2 yr at sea and by data from Alaska waters in their third year at sea. Results suggest that conditions in which the transition zone is dominated by neither the Alaska nor California Currents are best for increased growth of Puget Sound fish.     

Biophysical dynamics of western transition zones: a preliminary synthesis

The nature of the western portions of the biogeographic temperate or transition zones in the North Pacific and North Atlantic is reviewed. The physical transport of nutrients and biomass into them from the Kuroshio and Gulf Stream as well as from the poleward sides are estimated. The conclusion is that the upwelling in the two western boundary currents makes the largest contribution to the nutrient and biomass fluxes into these transition zones. A conservative estimate of the amount of upwelled fluid is derived from absolute velocity sections in the Gulf Stream. The estimate suggests that upwelling into the euphotic zone exceeds 2 × 10⁶ m³ s^–1. This implies that upwelling in these western boundary currents matches or exceeds that in eastern boundary currents such as the California Current. The two western boundary regimes have very different poleward situations. The Oyashio extension flows parallel to the Kuroshio and is a deep current. The North Atlantic Shelf Front flow is to the west where it is ultimately entrained into the edge of the Gulf Stream. There does not seem to be any tendency for this to occur in the Kuroshio. Despite these differences in the northern and western boundaries, the two transition zones are similar with large amplitude meanders, anticyclonic rings and streamers dominating their physical structure. The physical features responsible for the transfer of materials from the boundary current extensions into the transition zones are similar in both systems. Ring formation contributes only ? 10% of the transfer, while ring‐induced streamers contribute 30%. The rest of the transport is contributed by branching of the boundary current front. Both currents have well developed secondary fronts consisting of subtropical surface water pulled into the transition zone. Biologically, the upwelling in both western boundary currents leads to a biomass maximum along the boundary in both secondary producers (copepods) and in small pelagic fish. In the Kuroshio, the latter are the Japanese sardine, Sardinops melanostictus, that spawn in the Kuroshio and then enter the transition zone for the summer and fall months. In the Gulf Stream, the dominate species are menhaden, Brevoortia tyrannus and B. smithi. These species make use of the coastal environments of North America and although the adults spawn in the Gulf Stream, they are not thought to play a major role in the Slope Water, transition zone. The similar differences in the use of the Kuroshio and the Gulf Stream ecosystems occurs in the behaviour of bluefin tuna, squid and other large pelagics. The Gulf Stream system also lacks an equivalent to Pacific saury, Cololabis saira. The biology therefore is at least subtly different, with saury and sardines being replaced by mid‐water fish in the North Atlantic. A fuller comparison of the biology with quantitative methods in both systems should be encouraged.     

2015年超强厄尔尼诺事件对西北太平洋柔鱼渔场变动的影响

为研究超强厄尔尼诺事件对西北太平洋海域柔鱼（Ommastrephes bartarmii）资源量变动的影响,并分析柔鱼栖息地在极端气候条件下的变化规律,根据上海海洋大学鱿钓科学技术组提供的中国柔鱼生产捕捞数据,比较2008年正常气候年份与2015年超强厄尔尼诺年份的单位捕捞努力量渔获量（CPUE）、产量、捕捞努力量以及渔场纬度重心（LATG）的变化;利用栖息地适宜性指数模型对西北太平洋柔鱼栖息地的海表温度（SST）、光合有效辐射范围（PAR）和海表面高度距平（SSHA）3个关键环境因子进行分析。渔业数据时间为2008年和2015年9—11月,数据覆盖范围为36°N~48°N、150°E~170°E。结果发现,相对于2008年正常年份,2015年超强厄尔尼诺事件下的CPUE明显降低,且LATG向南偏移;此外,2015年适宜的SST和PAR范围均显著降低,导致适宜的栖息地面积与正常年份相比大幅减少;最适宜的SST和PAR等值线向南偏移,导致有利的栖息地纬度位置向南移动。研究认为,2015年超强厄尔尼诺事件发生时,柔鱼渔场环境不适于柔鱼生长,适宜栖息地面积减少且向南移动,导致该年份柔鱼资源丰度骤减,渔场向南偏移。     

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.     

Distribution and abundance of South-west Atlantic anchovy spawners (Engraulis anchoita) in relation to oceanographic processes in the southern Brazilian shelf

The reproductive environment of the South-west Atlantic anchovy, Engraulis anchoita , in the southern Brazilian coast was investigated using maritime weather reports from the US National Climatic Center. These reports were summarized to yield seasonal distributions of sea surface temperature, wind stress and Ekman transport for 2-month segments of the seasonal cycle. The vertical oceanographic structure and dynamics were studied using temperature and salinity data collected from oceanographic cruises. The seasonal distribution and biomass of South-west Atlantic anchovy spawners were estimated by acoustic surveys. Anchoita spawns intensively off southern Brazil during winter and early spring when the Ekman transport is directed onshore, and the combined effects of freshwater run-off and the flow of cold water near the bottom result in a strong vertical stability over the continental shelf. During this season, primary production peaks due to nutrient input from Sub-Antarctic waters and freshwater run-off. These conditions would avoid dispersal of eggs and larvae offshore and favour the production and maintenance of fine-scale food particle aggregations required for successful first feeding of newly hatched larvae. In summer, the conditions are almost reversed and anchoita spawners are virtually absent from the area. Thus, the spawning strategy of anchoita in southern Brazil seems to be tuned in a way to optimize larval retention, minimize exposure to turbulent mixing and take advantage of enhanced plankton productivity.     

Influence of episodic upwelling on capelin, Mallotusvillosus, and Atlantic cod, Gadusmorhua, catches in Newfoundland coastal waters

Wind-induced upwelling in the north-west Atlantic has been hypothesized to influence catch rates of fish by gear fishing at fixed locations. Passive movement to shallow depths during upwelling has been proposed to increase encounter rates of fish with net leaders set at the coast. However, supporting evidence is not conclusive, possibly because fish respond to strong events rather than all events. We investigated whether catch rates of capelin, Mallotusvillosus, and Atlantic cod, Gadusmorhua, were related to the strength of upwelling, as measured by the rates of water temperature change, or to stage of upwelling. Capelin trap catches were positively related to increases in water temperature, representing the relaxation phase. This result was due primarily to increases in catch after strong (> 4°C change) rather than after typical (< 4°C change) upwelling events. Cod trap catches were not related to upwelling strength but did increase 1–2 days after typical events. The data suggest that upwelling increases capelin movement, while the return of warmer surface water after an upwelling event increases cod movement.     

Offshore currents explain the discontinuity of a fish community in the seagrass bed along the Japanese archipelago

Oceanographic conditions can affect spatial variability in fish community structures by influencing the temperature‐dependent latitudinal distribution of adult fishes and transport during their young stages. In order to examine latitudinal variability in the fish community structure within a single coastal ecosystem, quantitative sampling was conducted in the sub‐tidal zone of seagrass Zostera marina beds over a broad latitudinal scale (31.31–43.0°N: from subtropical to sub‐boreal zones, covering 80% of the latitudinal range of seagrass distribution in Japan) in the western North Pacific based on a uniform methodology. Cluster analysis with the similarity of fish communities showed that 13 sampling sites were divided into two clusters. The border between the two clusters corresponded with the area of mixing of two dominant currents, Oyashio and Kuroshio, which form a border between the warm temperate zone and the cool temperate zone off the Pacific coast of Japan. Oceanographic properties, such as major currents off the coast, are suggested to affect the latitudinal variability in the fish communities in the coastal ecosystem in the western North Pacific.     

Hypoxia-based habitat compression of tropical pelagic fishes

Large areas of cold hypoxic water occur as distinct strata in the eastern tropical Pacific (ETP) and Atlantic oceans as a result of high productivity initiated by intense nutrient upwelling. We show that this stratum restricts the depth distribution of tropical pelagic marlins, sailfish, and tunas by compressing the acceptable physical habitat into a narrow surface layer. This layer extends downward to a variable boundary defined by a shallow thermocline, often at 25 m, above a barrier of cold hypoxic water. The depth distributions of marlin and sailfish monitored with electronic tags and average dissolved oxygen (DO) and temperature profiles show that this cold hypoxic environment constitutes a lower habitat boundary in the ETP, but not in the western North Atlantic (WNA), where DO is not limiting. Eastern Pacific and eastern Atlantic sailfish are larger than those in WNA, where the hypoxic zone is much deeper or absent. Larger sizes may reflect enhanced foraging opportunities afforded by the closer proximity of predator and prey in compressed habitat, as well as by the higher productivity. The shallow band of acceptable habitat restricts these fishes to a very narrow surface layer and makes them more vulnerable to over‐exploitation by surface gears. Predictably, the long‐term landings of tropical pelagic tunas from areas of habitat compression have been far greater than in surrounding areas. Many tropical pelagic species in the Atlantic Ocean are currently either fully exploited or overfished and their future status could be quite sensitive to increased fishing pressures, particularly in areas of habitat compression.     

Recruitment of common sardine (Strangomera bentincki) and anchovy (Engraulis ringens) off central-south Chile in the 1990s and the impact of the 1997–1998 El Niño

The recruitment of Strangomera bentincki (common sardine) and Engraulis ringens (anchovy) and the relationships with oceanographic conditions in the upwelling ecosystem of central-south Chile were investigated from 1990 to 1998, with emphasis on the 1997–1998 El Niño. Time series of recruitment, biomass, local sea surface temperature, and a coastal upwelling index were used to explore relationships during the spawning (July–August) and pre-recruitment (August–December) periods. The 1997–1998 El Niño caused physical changes in the small pelagic fish habitat off central-south Chile. Anomalous sea surface temperatures (SST) and upwelling indexes began to be detected from May 1997 and persisted into 1998. Recruitment of common sardine showed significantly negative relationship with SST anomalies during the pre-recruitment period, as well as with the upwelling index during the peak of spawning. However, the recruitment of anchovy did not seem to be affected by the environmental changes observed in the 1990s. Instead, the recruitment rate of anchovy showed negative relationship with the recruitment rate of common sardine. We conclude that the conditions of the 1997–1998 El Niño off central-south Chile affected the survival of common sardine offspring, and that the recruitment success of anchovy could be determined by less-abundant cohorts of common sardine through a biological mechanism of interaction.     

Interannual changes in sablefish (Anoplopoma fimbria) recruitment in relation to oceanographic conditions within the California Current System

Sablefish (Anoplopoma fimbria) supports substantial fisheries in both the eastern and western Pacific Oceans. Juvenile recruitment along the west coast of the continental United States has been highly variable over the past three decades. Using a generalized additive model, we demonstrate that physical oceanographic variables within the California Current System have significant effects on sablefish recruitment. Significant relationships were found between juvenile recruitment and northward Ekman transport, eastward Ekman transport, and sea level during key times and at key locations within the habitat of this species. The model explains nearly 70% of the variability in sablefish recruitment between the years 1974 and 2000. The predictive power of the model was demonstrated by refitting without the last 5 yr of data and subsequent prediction of those years. Bootstrap assessments of bias associated with parameter estimates and jackknife‐after‐bootstrap assessments of the influence of individual data on parameter estimates are presented and discussed. Using this model, it is possible to draw preliminary conclusions concerning year‐class strength of cohorts not yet available to the survey gear as well as historic year‐class strengths. We discuss changes in zooplankton abundance and shifts in species of copepods associated with fluctuations in the physical variables that appear to have a major influence on sablefish recruitment.     

Synchronicity in southern hemisphere squid stocks and the influence of the Southern Oscillation and Trans Polar Index

Squid are short lived, with highly labile populations that respond rapidly to changes in environmental conditions. This makes them a good model for studying the response of recruitment processes to environmental signals. This study examines the influence of the Southern Oscillation Index (SOI) and Trans Polar Index (TPI) on the environment and abundance of six species of commercially important squid from the southern hemisphere, all linked to major current systems connected by the Antarctic Circumpolar Current: Dosidicus gigas (Southeast Pacific), Loligo vulgaris reynaudii (Southeast Atlantic), Nototodarus sloanii, N. gouldi (Southwest Pacific), Illex argentinus and L. gahi (Southwest Atlantic). All fisheries displayed a high level of inter‐annual variability and a degree of synchronicity was seen to occur in the abundance of the three Pacific species. The SOI signal was reflected in the environment of each fishery, particularly in Pacific regions. Both indices are correlated with squid abundance, particularly during the early life history stages (SOI) and adult stages (TPI), suggesting some degree of latitudinal separation, with juveniles potentially influenced by environmental variability at lower latitudes and adults at higher latitudes.     

Predicting skipjack tuna forage distributions in the equatorial Pacific using a coupled dynamical bio-geochemical model

Skipjack tuna ( Katsuwonus pelamis ) contributes ≈70% of the total tuna catch in the Pacific Ocean. This species occurs in the upper mixed-layer throughout the equatorial region, but the largest catches are taken from the warmpool in the western equatorial Pacific. Analysis of catch and effort data for US purse seine fisheries in the western Pacific has demonstrated that one of the most successful fishing grounds is located in the vicinity of a convergence zone between the warm (>28–29°C) low-salinity water of the warmpool and the cold saline water of equatorial upwelling in the central Pacific (Lehodey et al ., 1997). This zone of convergence, identified by a well-marked salinity front and approximated by the 28.5°C isotherm, oscillates zonally over several thousands of km in correlation with the El Niño–Southern Oscillation. The present study focuses on the prediction of skipjack tuna forage that is expected to be a major factor in explaining the basin-scale distribution of the stock. It could also explain the close relation between displacements of skipjack tuna and the convergence zone on the eastern edge of the warmpool. A simple bio-geochemical model was coupled with a general circulation model, allowing reasonable predictions of new primary production in the equatorial Pacific from mid-1992 to mid-1995. The biological transfer of this production toward tuna forage was simply parameterized according to the food chain length and redistributed by the currents using the circulation model. Tuna forage accumulated in the convergence zone of the horizontal currents, which corresponds to the warmpool/equatorial upwelling boundary. Predicted forage maxima corresponded well with high catch rates.     

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

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

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.     

Distribution and community structure of ichthyoplankton from the northern and central California Current in May 2004–06

The distributions, concentrations, and community structure of pelagic larval fishes collected from the central and northern California Current in the northeast Pacific Ocean during May 2004, 2005, and 2006 were analyzed to investigate inter-annual, latitudinal, cross-shelf, and depth-stratified variability. The inter-annual climate-induced variability during the sampling period provided a unique opportunity to observe how larval fish communities adjust to rapidly changing environmental conditions. The 170 depth-stratified samples collected from three cruises yielded 14 819 fish larvae from 56 taxa representing 23 families. Dominant larval taxa were Engraulis mordax , Citharichthys spp., Sebastes spp., and Stenobrachius leucopsarus . Larval concentrations decreased significantly in 2006 from 2004 and 2005 levels following the anomalous oceanic conditions observed in 2005 and decreased water temperature in 2006. Larvae were generally found in higher concentrations at northern (>43°N) versus southern (<43°N) stations, with larval E. mordax and Citharichthys spp. found almost exclusively in the north during all sampled years. Inter-annual variability related to dynamic upwelling intensity was observed in cross-shelf larval distributions, although concentrations of S. leucopsarus larvae consistently increased in the offshore direction, while larval Sebastes spp. were generally found in highest concentrations at intermediate stations along the shelf. Multivariate analyses revealed that latitude, station depth, and sea-surface temperature were the most important factors explaining variability in larval concentrations. The present study shows that the ichthyoplankton community of the central and northern California Current changed dramatically in response to the variable environmental conditions of 2004–06.     

The influence of ocean climate on coastal plankton and fish production

This study evaluates the influence of oceanic variability on plankton and fish production in a coastal upwelling system off southern Vancouver Island, British Columbia, Canada. A trophodynamics model is used to hind-cast plankton and fish production as estimated by empirical seasonal patterns in upwelling, sea surface temperature, solar radiation, and by biomasses of Pacific hake (Merluccius productus) and Pacific herring (Clupea pallasi). Simulation results indicate that interannual and longer-term variability in coastal plankton production occurs, and that the patterns are primarily determined by the dynamics in the initiation and intensity of coastal upwelling. Fish properties are greatly influenced by zooplankton, with simulated hake production being correlated with summer euphausiid production, and measured herring condition being correlated with autumn zooplankton production. An evaluation of transfer efficiencies indicates that interannual variability in the diatom-to-fish transfer is high (CV = 78%), with the majority of this variability occurring in the diatom-to-zooplankton component of the transfer. Seasonal dynamics of the diatom-to-zooplankton transfer result from the effects of upwelling on diatom production, and subsequent trophodynamic phasing with copepods and euphausiids.     

The effect of vertical and horizontal distribution on retention of sardine (Sardinops sagax) larvae in the Northern Benguela – observations and modelling

In the present study, a modelling experiment is conducted to simulate the transport of sardine (Sardinops sagax) eggs and larvae in the Northern Benguela. Based on historical and newly obtained data, different scenarios of vertical and horizontal distribution are applied and the effects on retention are discussed. The simulations showed that vertical and horizontal distribution were important for retention of sardine larvae in the Northern Benguela. By using age‐dependent data on vertical distribution, it was shown that retention of particles in the simulation was substantially enhanced compared with a scenario where particles were distributed in the offshore moving Ekman layer. Retention was lowest during October–December (when upwelling intensity is high) and highest during February–April (when upwelling intensity is somewhat lower). When different spawning areas were considered, highest retention was observed in an area near Walvis Bay. It is concluded that the behaviour of sardine larvae is adapted to the circulation system in the Northern Benguela in a way that promotes retention of the larvae in inshore nursery areas.