Investigating the ‘northern Humboldt paradox’ from model comparisons of small pelagic fish reproductive strategies in eastern boundary upwelling ecosystems

Eastern boundary upwelling ecosystems are highly productive and sustain the world’s largest fisheries, usually dominated by sardine and anchovy species. Stock size is highly variable from year to year due to the impact of the unstable physical environment on fish early stages. Biophysical models of early life‐stage dispersal of marine organisms have been built by coupling (i) hydrodynamic models and (ii) life history models (i.e. egg and larva stages), and are therefore useful tools to investigate physical–biological interactions. Here, we review biophysical models of anchovy and sardine ichthyoplankton dispersals developed in the Benguela, Humboldt and Canary Current upwelling ecosystems. We also include a similar study conducted in the California Current upwelling on zooplankton. We then integrate this information into a comparative analysis of sardine and anchovy reproductive strategies in the different systems. We found that the main spawning periods match the season of (i) maximal simulated ichthyoplankton retention over the continental shelf in the northern Benguela, southern Humboldt and Canary (for sardine); (ii) maximal food concentration in the southern Benguela, California and Canary (for anchovy); and (iii) maximal shelf retention of ichthyoplankton and food concentration in the northern Humboldt (for both anchovy and sardine). This specificity of the northern Humboldt ecosystem could explain why it sustains the largest small pelagic fish stock. Finally, the possible effects of climate change on these patterns are discussed.     

Investigating connectivity between two sardine stocks off South Africa using a high‐resolution IBM: Retention and transport success of sardine eggs

This study applied a previously used Lagrangian individual‐based model (IBM) for sardine in the Southern Benguela to an improved and more robust hydrodynamic model to investigate whether a more representative spatial coverage, greater horizontal and vertical resolution, more realistic winds and improved representation of mesoscale features such as eddies and filaments would give different results for transport and retention of early life stages. Despite major differences between the old and new hydrodynamic models, overall the IBM results were quite similar to the previous southern Benguela sardine IBM study. This surprising result indicates that it is the macroscale circulation features resolved by the two hydrodynamic models that are controlling transport and retention of sardine early life stages. The contribution of transient mesoscale features such as eddies and filaments appears to be less important when transport patterns are averaged over the 21‐year‐long experiment. Another aim of this study was to better estimate the contribution of south coast spawning to west coast sardine recruitment. This was possible because of an eastward extension of the geographical domain of the new hydrodynamic model which provided a more realistic representation of the south coast spawning ground. Three main spawning and nursery area systems, similar to those identified in the previous sardine IBM, were identified: west coast and west coast (WC‐WC), south coast and west coast (SC‐WC), and south coast and south coast (SC‐SC). Spawning area proved to be an important determinant of modelled retention and transport success, with spawning depth also playing an important role on the west coast. The main difference observed from the previous study was an increase in the average percentage of particles released on the south coast and transported to the west coast (P₀, 17.4%). This indicates more connectivity between the southern and western sardine stocks than previously thought and is therefore important for fishery management. Standardized anomalies from the modelled retention/transport were compared with recruitment estimates from stock assessment models but there was no correlation between the two sets of anomalies. However, a significant correlation was observed between the modelled retention/transport anomalies for the west coast and total cumulative upwelling anomalies for the Southern Benguela (r = ?0.67, p < .001).     

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.     

The selection of spawning location of sardine (Sardinops sagax) in the northern Benguela after changes in stock structure and environmental conditions

Most reports on the distribution of spawning areas of sardine (Sardinops sagax) in the northern Benguela originate from the 1970s and 1980s. The northern Benguela system was in a high upwelling regime during those decades. Since the early 1990s upwelling favourable winds have decreased and a trend of increasing sea surface temperature (SST) has been observed. Changes in the structure of sardine stock in the northern Benguela have been observed and it has been suggested that a reduced biomass and changes in stock structure has led to decreased spawning in the favourable southern locations, thus preventing a recovery of the sardine stock. The present paper on the contrary shows that there has been a shift in spawning location from the less favourable northern areas in the early 1980s to spawning areas further south in the 2000s. Thus, the failure of the northern Benguela sardine stock to recover since its collapse in the late 1960s cannot be explained by spawning in less favourable areas. The shift in preferred spawning location to more southern areas since the 1980s was to be expected with a general warming of the northern Benguela system. Alternative explanations for the failure of the sardine stock to recover such as a reduction in average length as well as length at 50% maturity, leading to a reduction in reproductive output, increased predation pressure, and increased low oxygen waters are proposed.     

Pelagic ecology of a northern boundary current system: effects of upwelling on the production and distribution of sardine (Sardinops sagax), anchovy (Engraulis australis) and southern bluefin tuna (Thunnus maccoyii) in the Great Australian Bight

Shelf waters of southern Australia support the world's only northern boundary current ecosystem. Although there are some indications of intense nitrate enrichment in the eastern Great Australian Bight (GAB) arising from upwelling of the Flinders Current, the biological consequences of these processes are poorly understood. We show that productivity in the eastern GAB is low during winter, but that coastal upwelling at several locations during the austral summer–autumn results in localized increases in surface chlorophyll a concentrations and downstream enhancement of zooplankton biomass. Sardine (Sardinops sagax) and anchovy (Engraulis australis) eggs and larvae are abundant and widely distributed in shelf waters of the eastern and central GAB during summer–autumn, with high densities of sardine eggs and larvae occurring in areas with high zooplankton biomass. Egg densities and distributions support previous evidence suggesting that the spawning biomass of sardine in the waters off South Australia is an order of magnitude higher than elsewhere in southern Australia. Sardine comprised >50% of the identified prey species of juvenile southern bluefin tuna (SBT, Thunnus maccoyii) collected during this study. Other studies have shown that the lipid content of sardine from the GAB is relatively high during summer and autumn. We suggest that juvenile SBT migrate into the eastern and central GAB during each summer–autumn to access the high densities of lipid‐rich sardines that are available in the region during the upwelling period. Levels of primary, secondary and fish production in the eastern GAB during summer–autumn are higher than those recorded in other parts of Australia, and within the lower portion of ranges observed during upwelling events in the productive eastern boundary current systems off California, Peru and southern Africa.     

Does upwelling intensity determine larval fish habitats in upwelling ecosystems? The case of Senegal and Mauritania

European sardine (Sardina pilchardus) and round sardinella (Sardinella aurita) comprise two‐thirds of total landings of small pelagic fishes in the Canary Current Eastern Boundary Ecosystem (CCEBE). Their spawning habitat is the continental shelf where upwelling is responsible for high productivity. While upwelling intensity is predicted to change through ocean warming, the effects of upwelling intensity on larval fish habitat expansion is not well understood. Larval habitat characteristics of both species were investigated during different upwelling intensity regimes. Three surveys were carried out to sample fish larvae during cold (permanent upwelling) and warm (low upwelling) seasons along the southern coastal upwelling area of the CCEBE (13°–22.5°N). Sardina pilchardus larvae were observed in areas of strong upwelling during both seasons. Larval habitat expansion was restricted from 22.5°N to 17.5°N during cold seasons and to 22.5°N during the warm season. Sardinella aurita larvae were observed from 13°N to 15°N during cold seasons and 16–21°N in the warm season under low upwelling conditions. Generalized additive models predicted upwelling intensity driven larval fish abundance patterns. Observations and modeling revealed species‐specific spawning times and locations, that resulted in a niche partitioning allowing species' co‐existence. Alterations in upwelling intensity may have drastic effects on the spawning behavior, larval survival, and probably recruitment success of a species. The results enable insights into the spawning behavior of major small pelagic fish species in the CCEBE. Understanding biological responses to physical variability are essential in managing marine resources under changing climate conditions.     

Do Sardinella aurita spawning seasons match local retention patterns in the Senegalese–Mauritanian upwelling region?

Sardinella aurita is the most abundant small pelagic fish in the Senegalese–Mauritanian region. The success of its reproduction crucially depends on the local circulation as this determines whether larvae reach coastal nursery areas favorable to their survival or are dispersed into the open ocean. As a first step towards evaluating sardinella vulnerability to climate‐driven hydrodynamical changes, this study aims at underpinning how transport pathways drive optimal spatial and seasonal patterns for sardinella reproduction. We have used two estimates of the Senegalese–Mauritanian coastal seasonal circulation simulated by two hydrodynamical model configurations that differ in their forcing and topography. Nursery areas are determined by evaluating coastal retention with a Lagrangian individual‐based model that accounts for processes such as diel vertical migration and mortality as a result of lethal temperature exposure. Our results suggest that the shelf zones located at the Arguin Bank (19.5°N–21°N) and south of Senegal (12°N–14.75°N) are highly retentive. We find maximum retention rates in July–August and November–December over the Arguin Bank; from February–July and November–December over the southern Senegalese shelf; and lower retention rates over the central region (14.75°N–19.5°N) that are locally maximum in June–July when the upwelling weakens. These retention areas and their seasonality are in agreement with previously reported spawning patterns, suggesting that the Sardinella aurita spawning strategy may result from a trade‐off between retention patterns associated with the seasonal circulation and food availability. Exposure to lethal temperatures, although not well studied, could be a further limiting factor for spawning. The Lagrangian analysis reveals important connectivity between sub‐regions within and south of the system and hence underlines the importance for joint management of the Sardinella aurita stock.     

Clupeoid larval growth and plankton production in the Benguela upwelling system

Upwelling is a characteristic feature of the Benguela system. Strong upwelling leads to high primary production, which, in turn, supports large biomass of higher trophic levels. To analyse intra‐, interannual and decadal variability of fish recruitment, growth, and distribution, different methods to quantify this upwelling have been used. In the present work, an upwelling index, modelled diatom primary production, based on outputs from a numerical model for the Benguela upwelling system, is proposed. Upwelled water brings large amounts of nutrients into the euphotic zone and this supports primary productivity, which can be considered a preconditioner for the zooplankton that clupeoid larvae feed on. Therefore, it is assumed that high upwelling gives the potential for good feeding conditions for these larvae. This fact has motivated the use of the modelled index for upwelling intensity as a proxy for food availability and an investigation of the interannual variability of observed mean individual weight of Cape anchovy (Engraulis capensis) and sardine (Sardinops sagax) recruits. The study shows that the mean Cape anchovy recruit weight has a linear relationship with upwelling activity along the transport route (from the spawning to the nursery areas) and inside the nursery areas, while sardine have an Optimal Environmental Window (OEW) relationship to the upwelling on the transport route. A possible explanation for these observed differences is the prey the larvae feed on and the change in the plankton community as a function of upwelling activity.     

Evaluating the effects of climate change in the southern Benguela upwelling system using the Atlantis modelling framework

The ocean is affected by multiple anthropogenic stressors including climate change, the effects of which are already evident in many ocean ecosystems. The ABACuS v2 end‐to‐end model together with climate projections from the NEMO‐MEDUSA 2.0 model were used to evaluate the effects of fishing, warming and horizontal and vertical mixing on the southern Benguela upwelling system. Of the drivers examined in this study, warming had the greatest effect on species biomass, with mainly negative effects. The magnitude of the impacts of warming intensified from the RCP 2.6 to the 8.5 emission scenario. Fishing negatively affected demersal and large pelagic fish, which in turn resulted in a biomass increase of forage fish due to a decrease in predation pressure. Water mixing was found to have minor indirect effects on zooplankton biomass and fish. The responses of species and species groups to the combined effects of fishing and warming were approximately equally divided between additive, synergistic and antagonistic. Interpretation of our model results suggests that the southern Benguela system is likely to be affected by climate change, including substantial changes in the abundance of some species important to the region's fisheries. Future planning for fisheries needs to take this into account, including through management that strives to maintain the resilience of key species and the system as a whole. In line with previous studies on the southern Benguela, the results reinforce the importance of including consideration of the indirect and combined impacts of climate change and fishing in management and planning.     

Bonga shad (Ethmalosa fimbriata) spawning tactics in an upwelling environment

Successful recruitment in small pelagic fish populations inhabiting upwelling zones is subject to variation in fecundity and is driven by spatial and temporal fluctuations in environmental conditions, that is, mainly sea surface temperature, salinity and food availability. These fluctuations in abiotic factors have stimulated small pelagic fish populations to exhibit specifically adapted spawning tactics. To better understand to what extent a short‐lived exploited fish species such as bonga shad Ethmalosa fimbriata has adapted to an upwelling environment, we have investigated the interrelationship between upwelling intensity as a proxy for productivity and population fecundity by means of a virtual population analysis. We found that females of intermediate size contributed significantly more eggs to the population's fecundity than smaller or larger ones. Our model results further indicate that E. fimbriata exhibits a spawning preference at water temperatures of around 25°C and upwelling intensities of around 2.5 m³ s^?1 m^?1. Hence, we hypothesize that climate change‐driven increases in sea temperatures and modifications of upwelling‐favourable winds could significantly impact the species’ reproductive biology. To understand how climate change might impact fisheries, spawning tactics of small pelagic fishes are important to assess as well as their recruitment success. Such information is particularly relevant in countries where the fishery is critical at socio‐economic level, to better implement fisheries management addressing multiple stressors.     

Turbulent mixing and mesoscale distributions of late-stage fish larvae on the NW Shelf of Western Australia

Light traps were deployed to describe vertical and cross‐shelf distributions of late‐stage larval fishes during five cruises in each of the 1997/98 and 1998/99 summers in the region of the Gulf of Exmouth on the southern North West Shelf of Western Australia. At each light trap station on a cross‐shelf transect we measured water temperature, salinity and chlorophyll a and used vertical plankton tows to estimate zooplankton biomass and copepod abundance. Current meters were deployed on moorings near the transect and the data used to model flows and mixing on the NW Shelf and in the Gulf. The majority of reef, pelagic and baitfish larvae (81, 83 and 66% respectively) were collected at only two stations that marked the boundary between stratified waters offshore and well‐mixed water within the Gulf. Most baitfishes (primarily Spratelloides spp.) were captured by traps deployed near the seabed, while reef fishes (mostly pomacentrids, lethrinids and siganids) and pelagic species (mostly scombrids and carangids) were captured in traps deployed near surface. Catch composition varied between summers with 64% of baitfishes collected in the first summer, while the majority of reef and pelagic fishes (81 and 80% respectively) were captured in the second summer. Modelling of circulation showed that the velocity of tidal currents was enhanced by constriction of flow between NW Cape and South Muiron Island and by shallowing of the shelf. Flood‐tide intrusions of water allowed the thermocline to move up the continental shelf, upwelling cool nutrient‐rich water that was then mixed throughout the entire water column at stations in the mouth of the Gulf. This upwelling and mixing resulted in higher chlorophyll a concentrations and copepod abundances either as a result of local in situ growth or advection/aggregation processes, and may account for the great abundances of late‐stage fish larvae in the mouth of the Gulf.     

A numerical study of inferred rockfish (Sebastes spp.) larval dispersal along the central California coast

Successful recruitment of marine fishes depends on survival during early life-history stages, which is influenced by oceanic advection due to its impact on coastal trophodynamics and transport processes. Here we evaluate the influence of ocean circulation on the dispersal of rockfish ( Sebastes spp.) larvae along the central California coast using an implementation of the Regional Ocean Modeling System, driven at the surface by output from the Coupled Ocean Atmosphere Mesoscale Prediction System. Thousands of floats simulating rockfish larval propagules, constrained to follow fixed depths, were released over a broad coastal area at 2-day intervals, and transported by simulated ocean currents at depths of 1, 7, 20, 40, and 70 m. Trajectory statistics are averaged across the 4-yr period from January 2000 through December 2003 to reveal mean trajectory direction, net displacement, fractional cross-shore loss, and duration of retention for different seasons. On average, near-surface propagules originating nearshore are transported offshore during the upwelling season, whereas deeper propagules move alongshore to the north. This vertical shear vanishes during winter, with most floats moving alongshore to the north, regardless of depth. After 35 days in the water column, typical transport distances were ∼50 km for floats remaining nearshore and ∼150 km for floats over the midshelf and slope. Implications for performance of marine reserves for rockfish conservation are discussed. Our results also provide evidence for a strong semiannual pattern of coastal retention rates, with high export of near-surface drifters during the upwelling season. In contrast, high rates of shelf retention occurred for releases at 20 m and deeper during summer, and at all depths during winter.     

Historical DNA as a tool to genetically characterize the Mediterranean sand tiger shark (Carcharias taurus,Lamniformes: Odontaspididae): A species probably disappeared from this basin

1. The sand tiger shark (Carcharias taurus) is a coastal species distributed in temperate and sub‐tropical waters, classified as ‘Vulnerable’ at global level and ‘Critically endangered’ in eastern Australia, south‐western Atlantic Ocean, and Mediterranean Sea. Six populations (north‐western Atlantic, Brazil, South Africa, Japan, eastern Australia, and western Australia) with low genetic diversity and limited gene flow were identified worldwide, but genetic information for many other geographic areas are still missing. Specifically, this species is listed in several reports as part of the Mediterranean fauna, even if there has been a lack of catches and sightings in recent years in this basin. To clarify the origin of C. taurus individuals caught in the past in the Mediterranean Sea, historical samples were genetically analysed.
2. Nine samples with certain Mediterranean origin were collected from different European museums. DNA was extracted and ~600 bp of the mitochondrial DNA control region were amplified using eight overlapping species‐specific primer pairs. Sequences obtained were aligned with all the haplotypes globally known so far.
3. Genetic analysis revealed the misidentification of one museum specimen. Among the remaining Mediterranean historical samples, three different haplotypes were recovered. Two of them previously observed only in South Africa and one described in both South African and Brazilian populations.
4. Results suggest a genetic relationship between Mediterranean sand tiger sharks and those from the western Indian Ocean. According to previous studies, we hypothesized that, during the Pleistocene, the cold Benguela upwelling barrier was temporarily reduced allowing the passage of C. taurus individuals from the Indian to Atlantic Ocean. After the restoration of this phylogeographic barrier some individuals were trapped in the Atlantic Ocean and probably migrated northward colonizing the western African coasts and the Mediterranean Sea.

     

Recruitment ecology of pelagic‐broadcast spawning minnows: paradigms from the ocean advance science and conservation of an imperilled freshwater fauna

Pelagic‐broadcast spawning is a dominant mode of fish reproduction in the oceans and is associated with Fundamental Triad (i.e. nutrient enrichment, nutrient concentration, propagule retention) and Loophole (i.e. disruption of larval competition and predation) processes that are mediated by water currents. Pelagic‐broadcast spawning is uncommon in freshwater, but is employed by an evolutionarily convergent spawning guild in rivers on the Great Plains, North America. We reviewed ecological studies of pelagic‐broadcast spawning minnows to evaluate whether Fundamental Triad and Loophole concepts explain the adaptive significance of this behaviour. Pelagic‐broadcast spawning minnows exhibit spawning periodicity dependent on floods. Nutrient enrichment, nutrient concentration and propagule retention are enhanced during flood recession. Retention is also enhanced by high fecundity and rapid development of pelagic‐broadcast propagules. Predation and competition threats are relatively low in temporary, patchily distributed nurseries present during flood recession. Extended spawning periods and diffusive spread of juveniles and adults ensure annual recruitment despite the unpredictable and often harsh conditions in rivers on the plains. Over the last century, pelagic‐broadcast spawners have suffered declines due to dewatering, flood control and sediment deprivation that disrupt natural Fundamental Triad and Loophole processes. We propose that restoration of conditions necessary for Fundamental Triad and Loophole processes to operate will be most successful at stopping decline and extinction of pelagic‐broadcast spawning minnows of the Great Plains.     

Sardine and anchovy regime fluctuations of abundance in four regions of the world oceans: a workshop report

Regimes of high abundance of sardine (Sardinops sagax and Sardina pilchardus) have alternated with regimes of high abundance of anchovy (Engraulis spp.) in each of the five regions of the world where these taxa co-occur and have been extensively fished. When one taxon has been plentiful, the other has usually been at a reduced level of abundance, and vice versa. Changes in the four heavily fished regions that support S. sagax–the Japanese, Californian, Humboldt, and Benguela systems–from a regime dominated by one taxon to a high level of abundance of the other have occurred more or less simultaneously. In the Pacific Ocean, sardines have tended to increase during periods of increasing global air and sea temperatures and anchovies to decrease. The Japanese system is dominated by sardines to a greater extent than the other systems, and sardines off Japan appear to increase as the Kuroshio Current cools. At the eastern edge of the Pacific Ocean, sardines colonize cooler areas during periods of warming. The Benguela system is out of phase with the three Pacific systems. The four systems all appeared to be in a state of flux in the 1980s. Increased abundance of the subdominant taxon is often one of the first signs of change. Sardines are relatively sedentary in refuge areas when scarce but change behavior to become highly migratory and colonize cooler areas when abundant. Anchovies, by contrast, expand around a fixed geographic center.     

Climate‐driven winter variations of Calanus sinicus abundance in the East China Sea

Global environmental changes threaten the sustainable use of resources and raise uncertainties regarding marine populations' responses in a changing Ocean. The pelagic copepods of the genus Calanus play a central role in shelf ecosystems transferring phytoplankton carbon to harvested populations, from boreal to temperate regions. Here we examined a 15‐yr time series of Calanus sinicus abundance in regards to climate forcing in the East China Sea. We identified a compound effect of the Pacific Decadal Oscillation (PDO) and the East Asian Winter Monsoon (EAWM) on environmental conditions in the East China Sea. Such climate influences not only a southward transport of C. sinicus from its population centres into the Taiwan area, but favours advantageous thermal conditions for the species as well. On the interannual scale, our results show that the population size of C. sinicus echoes climate‐driven temperature changes. Hence, the possibility of using the PDO and EAWM variability for assessing and predicting interannual abundance changes of C. sinicus in the East China Sea is considered. The observed close relationship between climate and C. sinicus may promote bottom‐up controls in the pelagic food web, further influencing the southern edge of the species' geographic distribution. Owing to the prominent role this species plays in food web dynamics these results might help integrative fisheries management policies in the heavily exploited East China Sea.     

The importance of retention processes in upwelling areas for recruitment of Octopus vulgaris: the example of the Arguin Bank (Mauritania)

The relationship between recruitment of octopus ( Octopus vulgaris ) populations and environmental conditions off the Arguin Bank (Mauritania) in the main nursery ground was investigated between 1990 and 1996. Three environmental indices (coastal upwelling intensity, coastal retention and wind-induced turbulence indices) are derived from satellite infrared imagery (METEOSAT) and from the COADS (Comprehensive Ocean–Atmosphere Data Set) data to quantify the coastal retention process and productivity, both believed to be important factors for recruitment success. The octopus recruitment index as well as environmental indices fluctuate annually and seasonally. Linear regression and generalized additive models (GAM) are used to relate recruitment indices and the environmental conditions prevailing during the early-life planktonic stage, known to be the main critical period of the octopus life cycle. Coastal retention especially appears to be a key factor for recruitment success. Owing to seasonal variations in enrichment and mixing processes, larvae benefit from retention in the spring but are negatively affected by a breakdown in retention in the autumn. Increasing upwelling intensity is beneficial for octopus recruitment, as the Arguin Bank limits the detrimental dispersive effects linked to upwelling.     

Modelling the effect of buoyancy on the transport of anchovy (Engraulis capensis) eggs from spawning to nursery grounds in the southern Benguela: an IBM approach

An individual‐based model (IBM) was used to investigate the effects of physical and biological variables on the transport via a jet current of anchovy (Engraulis capensis) eggs from spawning to the nursery grounds in the southern Benguela ecosystem. As transport of eggs and early larvae is considered to be one of the major factors impacting on anchovy recruitment success, this approach may be useful to understand further the recruitment variability in this economically and ecologically important species. By coupling the IBM to a 3D hydrodynamic model of the region called Plume, and by varying parameters such as the spatial and temporal location of spawning, particle buoyancy, and the depth range over which particles were released, we could assess the influences of these parameters on transport success. A sensitivity analysis using a General Linear Model identified the primary determinants of transport success in the various experimental simulations, and model outputs were examined and compared with patterns observed in field studies. Model outputs compared well with observed patterns of vertical and horizontal egg distribution. Particle buoyancy and area of particle release were the major single determinants of transport success, with an egg density of 1.025 g cm^?3 maximizing average particle transport success and the western Agulhas Bank being the most successful spawning area. This IBM may be useful as a generic prototype for other upwelling ecosystems.     

Variation in the distribution of ocean shrimp (Pandalus jordani) recruits: links with coastal upwelling and climate change

In this analysis, an atypical northward shift in the distribution of age‐1 ocean shrimp (Pandalus jordani) recruits off Oregon in 2000 and 2002–2004 was linked to anomolously strong coastal upwelling winds off southern Oregon (42°N latitude) in April–July of the year of larval release (t?1). This is the first clear evidence that strong upwelling winds can depress local recruitment of ocean shrimp. Regression analysis confirmed a long‐term negative correlation between log_e of ocean shrimp recruitment and April sea level height (SLH) at Crescent City, California, in the year of larval release, for both northern and southern Oregon waters. The regional pattern of ocean shrimp catches and seasonal upwelling winds showed that, although the timing of the spring transition as reflected in April SLH drives ocean shrimp recruitment success off Oregon generally, the strength and consistency of spring upwelling limits the distribution of large concentrations of ocean shrimp at the southern end of the northern California/Oregon/Washington area. A northward shift in 1999 and 2001–03 in the northern edge of this ‘zone of maximum upwelling’ is the likely cause of the weak southern Oregon recruitment and resulting atypical distribution of ocean shrimp observed off Oregon in 2000 and 2002–04, with a return to a more typical catch distribution as spring upwelling moderated in subsequent years. It is noted that a northward shift in the conditions that produce strong and steady spring upwelling winds is consistent with many predictions of global climate models under conditions of global warming.     

北太平洋小型中上层鱼类资源对气候–海洋变化的响应研究进展

小型中上层鱼类是北太平洋海域重要的渔业资源,具有生命周期短、生长速度快、高集群性等特点,其资源年间波动显著,且受气候-海洋变化的影响。本文围绕秋刀鱼（Coloabis saira）、鲣（Katsuwonus pelamis）、鲐（Scomber japonicus）、鳀（Engraulis japonious）、竹䇲鱼（Trachurus japonicus）、沙丁鱼（Sardinops sagax）6种主要的小型中上层鱼类,回顾了厄尔尼诺-南方涛动（El Niño/La Niña-southern oscillation,ENSO）、太平洋年代际振荡（the Pacific decadal oscillation,PDO）、黑潮-亲潮（Kuroshio-Oyashio,KR-OY）等关键气候-海洋指数的特点及对鱼类栖息地环境和资源变动的影响。概括了气候-海洋变化对小型中上层鱼类的洄游分布和资源丰度的直接影响过程,以及对亲体繁殖产卵、仔稚体成活率和资源量波动间接的滞后影响过程。建议：（1）在多种气候-海洋指数基础上添加种群动态过程、捕捞方式系数、自然死亡率等参数构建生物量动态模型,揭示气候-海洋变化对渔业资源量的影响过程;（2）结合北太平洋涛动（North Pacific oscillation,NPO）、北极涛动（Arctic oscillation,AO）、北太平洋环流振荡（North Pacific gyre oscillation,NPGO）等其他北太平洋主要气候,基于物理海洋模型及空间耦合水动力学模型研究大尺度海流、中尺度涡旋对小型中上层鱼类影响。