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1.
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.  相似文献   

2.
Important environmental processes for the survival and recruitment of early life stages of pelagic fishes have been synthesized through Bakun's fundamental triad as enrichment, concentration and retention processes (A. Bakun, 1996, Patterns in the Ocean. Ocean Processes and Marine Population Dynamics. San Diego, CA, USA: University of California Sea Grant). This conceptual framework states that from favourable spawning habitats, eggs and larvae would be transported to and/or retained in places where food originating from enrichment areas would be concentrated. We propose a method for quantifying two of the triad processes, enrichment and retention, based on the Lagrangian tracking of particles transported within water velocity fields generated by a three‐dimensional hydrodynamic model. We apply this method to the southern Benguela upwelling ecosystem, constructing putative maps of enrichment and retention. We comment on these maps regarding main features of the circulation in the region, and investigate seasonal variability of the processes. We finally discuss the results in relation to available knowledge on the reproductive strategies of two pelagic clupeoid species abundant in the southern Benguela, anchovy (Engraulis encrasicolus) and sardine (Sardinops sagax). Our approach is intended to be sufficiently generic so as to allow its application to other upwelling systems.  相似文献   

3.
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.  相似文献   

4.
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.  相似文献   

5.
Generalized additive models (GAMs) were fitted to sardine (Sardina pilchardus) egg distribution data from three daily egg production method surveys. The results showed that the area of egg cover off Portugal decreased significantly from 11 800 km2 in 1988 to 7000 km2 in 1997 and 7400 km2 in 1999. This is because of a significant reduction in sardine egg presence off northern Portugal, GAM estimated areas being similar or higher in the late 1990s for southwestern and southern Portugal. The distributional area covered by larvae was not estimated for 1988 (larval distribution extended beyond the survey area), although it was probably higher than the 9600 km2 for 1997 and 5500 km2 for 1999. In 1997 and 1999, the Gulf of Cadiz was also sampled, indicating extensive areas with sardine eggs and larvae (more than 50% of the total area of distribution off Portugal). Standardized data from 15 ichthyoplankton surveys between 1985 and 2000 show a decline in the mean probability of egg presence within the Portuguese continental shelf from the mid‐1980s to the late‐1990s, because of a marked reduction in egg presence off northern Portugal. Sardine larval data from the same surveys suggest that the reduction in mean probability of presence in the north is less marked than for eggs (although this comparison ignores the presence of sardine larvae beyond the continental shelf in the 1980s). Similar changes off northern Portugal and western Galicia are observed in commercial sardine catches and the acoustically estimated area of fish distribution. It is possible that the observed decline in spawning area off northwestern Iberia during the 1990s is indirectly reflecting the prevalence of environmental conditions detrimental to sardine recruitment (northerly winds during winter that favour coastal upwelling and offshore transport), which have reduced the spawning contribution of young fish in that area.  相似文献   

6.
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).  相似文献   

7.
Sardine fisheries in the Iberian Atlantic shelf (36°N–44.5°N) show decadal‐scale cycles. In the late 1990s, a positive phase in sardine stock was expected; on the contrary, catches have declined until now. Regime shifts in climatic and oceanographic variables on different scales (as forcing factor) and shifts in sardine stock (as result) have been used with the aim of identifying the physical variables that explain most of the sardine population variance in the region. Circa 1998, when last sardine regime shift was detected, the main patterns of large‐scale atmospheric circulation in the Northern Hemisphere with influence in the study area namely Northern Atlantic Oscillation (NAO) and East Atlantic (EA) pattern changed and coupled in a combination that led to a rise in sea surface temperature and a decline in the coastal upwelling intensity. Several years with a downwelling situation in average in the main spawning and feeding Iberian sardine areas would have affected the stock abundance, averting the return to the projected positive regime. The sardine negative regime shift was detected first in the regions of the study located further north. The regional variable latent heat flux that groups a set of environmental processes related to the ocean–atmosphere heat exchanges and so with the turbulence manages to explain the 72% of sardine recruitment.  相似文献   

8.
Gonosomatic indices and egg and larval densities observed from 1986 to 2001 suggest that the peak spawning season of the Australian anchovy (Engraulis australis) in South Australia occurs during January to March (summer and autumn). This coincides with the spawning season of sardine (Sardinops sagax) and the period when productivity in shelf waters is enhanced by upwelling. Anchovy eggs were abundant throughout gulf and shelf waters, but the highest densities occurred in the northern parts of Spencer Gulf and Gulf St Vincent where sea surface temperatures (SST) were 24–26°C. In contrast, larvae >10 mm total length (TL) were found mainly in shelf waters near upwelling zones where SSTs were relatively low (<20°C) and levels of chlorophyll a (chl a) relatively high. Larvae >15 mm TL were collected only from shelf waters near upwelling zones. The high levels of larval abundance in the upwelling zones may reflect higher levels of recruitment to later stages in these areas compared with the gulfs. The sardine spawns mainly in shelf waters; few eggs and no larvae were collected from the northern gulfs. The abundance of anchovy eggs and larvae in shelf waters increased when sardine abundance was reduced by large‐scale mortality events, and decreased as the sardine numbers subsequently recovered. We hypothesize that the upwelling zones provide optimal conditions for the survival of larval anchovy in South Australia, but that anchovy can only utilize these zones effectively when the sardine population is low. At other times, northern gulf waters of South Australia may provide a refuge for the anchovy that the sardine cannot utilize.  相似文献   

9.
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.  相似文献   

10.
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.  相似文献   

11.
Stock level of Japanese sardine (Sardinops melanostictus) was high from 1980s to early 1990s and low from late 1990s to 2000s. The warm and cold water masses in the vicinity of the Kuroshio axis from winter to early spring used to be critical for the recruitment in the high‐stock period, because most of the larvae were distributed there. However, the environmental fluctuation might not affect the recruitment in the low‐stock period. Some studies reported that spawning location and spawning season, and hence the larval habitat, differ depending on the stock level. Three points were investigated in this study: (a) how spawning location and spawning season shifted from the late 1990s, (b) confirmation of the distribution area of larvae in the recent low‐stock period and (c) whether the water temperature in the vicinity of the Kuroshio axis was still related to the recruitment in the low‐stock period. The spawning location and spawning season clearly changed after 1995. Consequently, particle tracking experiments suggested that the larvae appeared in the vicinity of the Kuroshio axis from winter to early spring decreased. Nevertheless, only the ambient temperature of larvae that appeared in the vicinity of the Kuroshio axis from winter had a significant negative correlation with an index of the recruitment in the low‐stock period. It is suggested that the warm and cold masses in the vicinity of the Kuroshio axis are critical for the recruitment regardless of the stock level.  相似文献   

12.
Interannual variation of some biological parameters for the Japanese sardine, Sardinops melanosticta, population in Korean waters was investigated using scientific surveys and fisheries information since the late 1970s. The abundance and geographical coverage of sardine eggs were high (peak in 1986) and broad when spawning biomasses were high in the mid 1980s, and vice versa in the late 1970s and the early 1990s. Also, feeding and spawning areas based on fisheries information exhibited the same pattern of expansion/contraction as seen in ichthyoplankton surveys. Annual Gonadal Somatic Index (GSI) in spawning season (February to April) and the size at age 1 of sardine were reduced during the high abundance period. It is suggested that density‐dependent effects on the reproduction and growth of the sardine population in Korean waters existed.  相似文献   

13.
We use trivariate kernel density estimation to define spawning habitat of northern anchovy ( Engraulis mordax ) and Pacific sardine ( Sardinops sagax ) in the California Current using satellite data and in situ egg samples from the Continuous Underway Fish Egg Sampler (CUFES) deployed during surveys in April by the California Cooperative Oceanic Fisheries Investigations (CalCOFI). Observed egg distributions were compared with monthly composite satellite sea surface temperature (SST) and surface chlorophyll a (chl a ) data. Based on the preferred spawning habitat, as defined in SST and chl a space, the satellite data were used to predict potential spawning habitat along two areas of the west coast of North America. Data from the southern area (21.5 to 39°N) were compared to observations from the CUFES data for the period 1998–2005. Northern anchovy and Pacific sardine exhibited distinctly different spawning habitat distributions. A significant relationship was found between satellite-based spawning area and that measured during surveys for sardine. CUFES area estimated for sardine was similar in magnitude to that estimated from satellite data (∼60 000 km2). In contrast, spawning habitat of anchovy averaged between 1000 and 200 000 km2 for the period 1998–2005, for CUFES and satellite estimates, respectively. Interannual variability in the area (km2) and duration (months) of estimates of suitable habitat varied between species and between the northern (39 to 50.5°N) and southern portions of the California Current. Long-term monitoring of habitat variability using remote sensing data is possible in the southern portion of the California Current, and could be improved upon in the northern area with the addition of surveys better timed to describe relationships between observed and estimated spawning habitats.  相似文献   

14.
A geostatistical analysis has been undertaken on the spatial structure of co-occurring adult and recruit populations of anchovy, Engraulis capensis , and sardine, Sardinops sagax , in the southern Benguela upwelling region, using information from two acoustic surveys. The study was prompted by the need for a more efficient design for surveying sardine abundance, which is increasing in relation to that of anchovy; the current acoustic survey design is based on the distribution of anchovy. Variograms of fish density and density indicator variables were computed, as well as cross-variograms between the indicator variables. The sardine variograms were less structured than the anchovy variograms, with slightly greater nugget effects, indicating greater randomness in space at small scales. The indicator variograms showed progressive loss of structure with increasing density. Anchovy formed high-density schools during the day, breaking into larger, low-density aggregations at night. Sardine, on the other hand, remained in relatively high-density schools throughout the day. The cross-variograms revealed some spatial continuity between low- and high-density areas for anchovy, but no such transitional structures were evident for sardine. It was concluded that sardine are more patchily distributed than anchovy and, unlike anchovy, they may not have a single common way of occupying space at the population level, perhaps partly because of the broader age structure of the population. It is concluded that, while the current acoustic survey design is well suited to the spatial distribution of anchovy, it is not as well suited to that of sardine. Future survey designs should be more tailored to the spatial distribution of sardine, possibly by using sampling methods to cope with the highly patchy distributions expected.  相似文献   

15.
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16.
Particle‐tracking experiments were performed to infer the distribution of larvae of the Japanese sardine (Sardinops melanostictus) and to detect effects of transport environment on sardine recruitment, using the output of a high‐resolution ocean general circulation model and observed data of sardine spawning grounds during 1978–2004. By the 60th day following spawning, approximately 50% of the larvae had been transported to the Kuroshio Extension (KE). Whereas the spawning period and grounds changed markedly in relation to the stock level, the proportion of larvae transported to the KE remained relatively constant and no significant correlations were found between sardine recruitment and the transport proportion. Instead, the recruitment was found to be correlated with physical parameters including the mixed layer depth and the sea surface temperature along several major transport trajectories of sardine larvae. The correlations were most significant for the trajectories in the region 0.5° south to 1° north of the Kuroshio axis (defined as the location of velocity maxima at each longitude) and for larvae spawned in February and March during the high stock period (1978–94), and for larvae spawned in March and April during the low stock period (1995–2004).  相似文献   

17.
This study describes broad-scale spatial variations in sardine growth across the northeastern Atlantic and Mediterranean waters using opportunistic samples collected in recent years. More detailed information on spatial, decadal and seasonal growth variations is provided for the Iberian-Biscay region using data collected in acoustic surveys since the mid-1980s. Growth curves are fitted to annual or monthly length-at-age data using a robust Von Bertalanffy model; parameters for recent samples are compared with literature information using an auximetric plot while differences between areas within the Iberian-Biscay region are tested by log-likelihood ratio tests. Sardine growth performance is generally lower in the Mediterranean and declines across the northeastern Atlantic from the English Channel to north Morocco but increases sharply off Mauritania. Lower growth of Mediterranean sardines is possibly associated to the overall oligotrophy of this Sea while differentiation from the Atlantic is likely sustained by reproductive isolation between populations from the two areas. Within the northeastern Atlantic, size- and age related migrations may partly explain differences in maximum length/age and mean length-at-age between neighbouring areas but the broad-scale latitudinal decline in growth is consistent with adaptation to the north–south decline in seasonal temperature gradients and to the annual cycles of plankton production. Within the Atlantic Iberian waters, sardine grows and improves in condition during spring and summer when the allocation of energetic resources for gonad development cease, temperature is close to the annual maxima and plankton production is high. Variation in sardine length-at-age and growth within the Atlanto-Iberian stock area has implications for stock structure and needs to be taken into account in the calculation of weight and maturity-at-age for assessment purposes. No evidence of broad temporal changes in sardine growth within the Iberian-Biscay region is obtained.  相似文献   

18.
The poleward flowing East Australian Current (EAC) drives sporadic upwelling, entrains coastal water and forms the western Tasman Front (wTF), creating a mosaic of water types and larval transport routes along south eastern Australia. The spatial distribution, otolith chemistry and growth rates of larval sardine (Sardinops sagax) were examined to infer spawning location and larval transport. A gradient of increasing larval size from north to south along the shelf was not detected but was evident between the shelf and offshore in the wTF. Here larvae were larger and older. Based on the occurrence of newly hatched larvae, spawning by S. sagax between southern Queensland and mid New South Wales (NSW) was more extensive than previously reported. The otolith chemistry from two wTF larval size classes differed, implying different origins. The otolith chemistry of wTF post‐flexion larvae was similar to larvae from northern NSW, whereas wTF flexion larvae were similar to larvae observed nearby from mid‐NSW. Two possible larval transport routes, direct and indirect, are inferred from otolith chemistry, current velocities and a previously published particle tracking study. Either larvae from northern NSW were advected south and entrained with younger larvae directly into the wTF, or larvae from a range of shelf regions were advected around the southern edge of an anticyclonic eddy, to join younger larvae directly entrained into the wTF. Based on the co‐occurrence of larval ages and sizes in the wTF and their advection routes, the wTF appears to be an important larval retention zone.  相似文献   

19.
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.  相似文献   

20.
From 1998 to 2011, the effects of environmental conditions on the spatial and temporal trends of sardine and sardinella catch rates in the Mauritanian waters were investigated using generalized additive models. Two models were used: a global model and an oceanographic model. The global models explained more of the variability in catch rates (60.4% for sardine and 40% for sardinella) than the oceanographic models (42% for sardine and 32.4% for sardinella). Both species showed clear and inverse seasonal variations in abundances corresponding to their main spawning activities and the hydrologic seasons off the Mauritanian waters. Sardine prefer colder waters and seem to occupy the ‘gap’ in the northern part of the Mauritanian waters as soon as sardinella has left the area because of to lower water temperatures. Unlike sardinella, sardine showed a gradual southward extension between 2002 and 2009. The oceanographic model revealed that a high proportion of catch variability for the two species could be explained by environmental variables. The main environmental parameters explaining the variability are sea surface temperature (SST) and the upwelling index for sardinella, and the chlorophyll‐a (Chl‐a) concentration, the upwelling index and SST for sardine. The sardinella spatio‐temporal variations off Mauritania seem to be more controlled by thermal than productivity gradients, probably linked to the species physiological constraints (thermal tolerance) whereas sardine seems to be more controlled by an ‘optimal upwelling and temperature’ windows. The results presented herein may be useful for understanding the movement of these species along the Mauritanian coast and hence their management under a changing climate.  相似文献   

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