Variations in spawning ground area and egg density of the Japanese sardine in Pacific coastal and oceanic waters

The spawning ground of the Japanese sardine, Sardinops melanostictus (Schlegel), was distributed over the oceanic waters as well as the coastal waters along the Pacific coasts of western and eastern Japan during 1978–1992. The area of the spawning ground in the coastal waters on the continental shelf has ranged from 95 000 km² in 1992 to 143 000 km² in 1988, constituting 44–77% of the total area of the spawning ground. The area of the coastal spawning ground was relatively constant in spite of the large fluctuations in egg abundance, i.e. size of the spawning population, from 88 trillion (1987) to 668 trillion (1989) in the waters. Spawning adults seemed to extend over the coastal waters irrespective of the size of the spawning population. In contrast to the coastal waters, the spawning area in the oceanic waters offshore of the continental shelf increased from 31 000 km² in 1978 to 183 000 km² in 1988 and then shrank to 83 000 km² in 1992, as a function of the spawning population size. The egg distribution density in the coastal waters stayed less than 6000 m⁻² mo⁻¹, but it reached as high as 27 400 m⁻² mo⁻¹ in the expanded spawning ground in the oceanic waters. The oceanic waters seemed to function as a reserve spawning ground for the sardine in years of extremely high spawning population.     

Importance of the Shatsky Rise Area in the Kuroshio Extension as an offshore nursery ground for Japanese anchovy (Engraulis japonicus) and sardine (Sardinops melanostictus)

Recent findings suggest that recruitment of Japanese anchovy (Engraulis japonicus) and sardine (Sardinops melanostictus) depends on survival during not only the first feeding larval stage in the Japanese coastal waters and the Kuroshio front but also during the post‐larval and juvenile stages in the Kuroshio Extension. Spatial distributions of juvenile anchovy and sardine around the Shatsky Rise area in the Kuroshio Extension region and the Kuroshio–Oyashio transition region are described, based on a field survey in the late spring using a newly developed mid‐water trawl for sampling juveniles. All stages of anchovy from post‐larvae to juveniles were obtained in the northern Shatsky Rise area. The Kuroshio Extension bifurcates west of the Shatsky Rise area and eddies are generated, leading to higher chlorophyll concentrations than in the surrounding regions in April and May. When Japanese anchovy and sardine spawn near the Kuroshio front or the coastal waters south‐east of Japan, their larvae are transported by the Kuroshio Extension and are retained in the Shatsky Rise area, which forms an important offshore nursery ground, especially during periods of high stock abundance.     

Transport and survival of larvae of pelagic fishes in Kuroshio system region estimated with Lagrangian drifters

ABSTRACT:   Transport and survival of larvae of pelagic fishes in the Kuroshio system region were studied using Lagrangian drifter data recorded from 1990 to 2003. A large portion of the drifters from the Kuroshio area south of Japan spread around the Kuroshio Extension up to 170°E, while some moved south to the offshore area of the Kuroshio because of a recirculation gyre. The monthly mean eastward movement from areas south of Japan was approximately 800–900 km, which was smaller than previous numerical estimates. The results of a survival model assuming optimal temperatures for larvae suggest that surface waters during the observation period were too warm for larval Japanese sardine, which has an optimal temperature of 16°C, and the adult abundances did not increase during the observation period. In contrast, the spawning ground temperatures and transport conditions from an area south-west of Japan in April–June matched the requirement for the larval Japanese anchovy, which has an optimal temperature of 22°C. The combined effects of temperature variations due to seasonality and water mass mixing are suggested to play an important role in determining the environmental temperatures that occur in an area.     

Environmental variability and growth histories of larval Japanese sardine (Sardinops melanostictus) and Japanese anchovy (Engraulis japonicus) near the frontal area of the Kuroshio

Environmental variability and growth‐rate histories from hatching to capture were investigated for larval Japanese sardine (Sardinops melanostictus) and Japanese anchovy (Engraulis japonicus). Larvae collected around the front of the Kuroshio Current were examined using otolith microstructure analysis, and their movement was estimated from numerical particle‐tracking experiments. Sardine larvae collected inshore of the Kuroshio front originated from a coastal area near the sampling site, while those collected in the offshore area originated from an area 500–800 km west‐southwest of the sampling site. Anchovy larvae collected both inshore and offshore had been transported from widely distributed spawning areas located west of the sampling area. At the age of 13–14 days for sardine and 19–20 days for anchovy, the offshore group exhibited significantly higher mean growth rates than did the inshore group. Although the offshore area was generally warmer than the inshore area, temporal variations in growth rate are not attributable solely to fluctuations in environmental temperature. While previous studies have examined the relationship between larval growth rates and environment based solely on data at capture, the methods used in the present study, combining otolith analysis and numerical particle‐tracking experiments, utilize data up until hatching. Although the relationship between growth rate and environment was not fully confirmed, this approach will greatly advance our understanding of fish population dynamics.     

Transport and environmental temperature variability of eggs and larvae of the Japanese anchovy (Engraulis japonicus) and Japanese sardine (Sardinops melanostictus) in the western North Pacific estimated via numerical particle-tracking experiments

Numerical particle-tracking experiments were performed to investigate the transport and variability in environmental temperature experienced by eggs and larvae of Pacific stocks of the Japanese anchovy ( Engraulis japonicus ) and Japanese sardine ( Sardinops melanostictus ) using high-resolution outputs of the Ocean General Circulation Model for the Earth Simulator (OFES) and the observed distributions of eggs collected from 1978 to 2004. The modeled anchovy individuals tend to be trapped in coastal waters or transported to the Kuroshio–Oyashio transition region. In contrast, a large proportion of the sardines are transported to the Kuroshio Extension. The egg density-weighted mean environmental temperature until day 30 of the experiment was 20–24°C for the anchovy and 17–20°C for the sardine, which can be explained by spawning areas and seasons, and interannual oceanic variability. Regression analyses revealed that the contribution of environmental temperature to the logarithm of recruitment per spawning (expected to have a negative relationship with the mean mortality coefficient) was significant for both the anchovy and sardine, especially until day 30, which can be regarded as the initial stages of their life cycles. The relationship was quadratic for the anchovy, with an optimal temperature of 21–22°C, and linear for the sardine, with a negative coefficient. Differences in habitat areas and temperature responses between the sardine and anchovy are suggested to be important factors in controlling the dramatic out-of-phase fluctuations of these species.     

Differences in food availability for Japanese sardine larvae between the frontal region and the waters on the offshore side of Kuroshio

The availability of food for larvae of the Japanese sardine, Sardinops melanostictus , was investigated in the Kuroshio frontal region and the waters on the offshore side of the Kuroshio, the Pacific coast of central Japan, in March 1990 and 1991, respectively. Food availability was assessed by changes in biomass and production of nauplii and small copepods, and RNA/DNA ratios of the larvae during about 2.5 days (the frontal region) or 3 days (the offshore waters) of tracking a drifter released in a pitch of the larvae. The biomas of the nauplii tended to increase with time in the frontal region and to decrease in the waters on the offshore side of the Kuroshio during the drifter tracking periods. The production of small copepods including nauplii in the waters on the offshore de of the kuroshio was 14% of that in the frontal region. The sum of the mean food requirements of the carnivorous macrozooplankters and sardine larvae was 11% of the production of small copepods including nauplii in the frontal region, compared with 136% in the waters offshore of the Kuroshio. The RNA/DNA ratios of postlarvae smaller than 8 mm in the frontal reqion were significantly higher than those in the waters on the offshore side of the Kuroshio ( P < 0.001) It is considered that the food availability for sardine larvae was relatively high in the frontal region and low in the waters on the offshore side of the Kuroshio. The food availability for the larvae probably deteriorated with the offshore shift of the main spawning ground from the frontal region to the waters on the offshore side of the Kuroshio in the latter half of the 1980s.     

Alternating dominance of postlarval sardine and anchovy caught by coastal fishery in relation to the Kuroshio meander in the Enshu-nada Sea

In the mid 1970s, the fishery catch of postlarval Japanese anchovy (Engraulis japonica) in a shelf region of the Enshu‐nada Sea, off the central Pacific coast of Japan, started to decline corresponding to a rapid increase of postlarval sardine (Sardinops melanostictus). In late 1980s, sardine started to decline, and it was replaced by anchovy in the 1990s. This alternating dominance of postlarval sardine and anchovy corresponded to the alternation in egg abundance of these two species in the spawning habitat of this sea. It was also noteworthy that during the period of sardine decline, sardine spawning occurred in April–May, a delay of two months compared with spawning in the late 1970s. The implication of oceanographic changes in the spawning habitat for the alternating dominance of sardine and anchovy eggs was explored using time‐series data obtained in 1975–1998, focusing on the effect of the Kuroshio meander. Large meanders of the Kuroshio may have enhanced the onshore intrusion of the warm water into the shelf region and contributed to an increase in temperature in the spawning habitat. This might favour sardine, because its egg abundance in the shelf region was more dependent on the temperature in early spring than was that of anchovy. In addition, enhanced onshore intrusion could contribute to transport of sardine larvae from upstream spawning grounds of the Kuroshio region. On the other hand, anchovy egg abundance was more closely related to lower transparency at the shelf edge, which may indicate the prevalence and prolonged residence of the coastal water, and therefore higher food availability, frequently accompanying non‐meandering Kuroshio. The expansion/shrinkage of the spawning habitat of sardine and anchovy in the shelf region, apparently responding to the change in the Kuroshio, possibly makes the alternation in dominance of postlarval sardine and anchovy most prominent in the Enshu‐nada Sea, in combination with changes in the abundance of spawning adults, which occurred almost simultaneously in the overall Kuroshio region. The implication of this rather regional feature for the alternating dominance of sardine and anchovy populations on a larger spatial scale is also discussed.     

Modelling the dispersal of larval Japanese sardine, Sardinops melanostictus, by the Kuroshio Current in 1993 and 1994

Acoustic Doppler current profiler (ADCP) data collected during routine monitoring surveys of the distribution and abundance of Japanese sardine larvae ( Sardinops melanostictus ) off the Pacific coast of Japan in February 1993 and 1994 were used to construct stationary average flowfields for three levels in the upper 100 m in each year. No large-scale meanders in the path of the Kuroshio Current were present in either year, but the axis of the current was closer to the coast in 1993 than in 1994. The flowfields were used to drive a particle-tracking model representing the dispersal of sardine eggs and larvae. Particles were released in accordance with the observed distribution of eggs, and their positions tracked for up to 40 days. In 1993, the model indicated that ≈ 50% of the egg production was carried north-eastwards out of the survey area into the area of the NW Pacific referred to as the Kuroshio Extension Zone. In contrast, only 5% of the egg production was exported to the Extension Zone in 1994, the remainder being retained in Japanese coastal waters. The consequences of the different dispersal patterns are discussed in relation to subsequent recruitment to the sardine stock. Based on commercial catch data, survival of the 1993 year class was 15% of that for the 1994 class. Hence, the results indicate that export of larvae to the Kuroshio Extension cannot in itself lead to successful recruitment.     

Environmental characteristics of spawning and nursery grounds of Japanese sardine and mackerels in the Kuroshio and Kuroshio Extension area

The sustainable use of marine resources requires understanding the surrounding ecosystem and elucidating mechanisms of variation. However, we still lack a comprehensive understanding of environmental variation in the spawning and nursery grounds of important fisheries species Japanese sardine (Sardinops melanostictus) and mackerels (Scomber japonicus and Scomber australasicus) in the northwest Pacific. Here, we investigate detailed physical, chemical, and biological environment variations in the spawning and nursery grounds along the Kuroshio and Kuroshio Extension area from intensive investigation in spawning season (April) of 2013. We found similar water mass property and copepod community in the egg‐rich Kuroshio area and the larvae‐rich downstream Kuroshio Extension area, indicating environmental variability is small during transportation and development processes. The egg‐rich northern Izu Islands region showed high copepod abundance, although low nutrient and chlorophyll concentrations were observed. Eggs were scarce or absent in the second survey 10 days after abundant eggs were observed in the region, along with differences in water property and copepod community. This indicates that not only the location but also the specific water characteristic and copepod community are a determining factor for spawning. Indicator communities of copepod found in our study (indicator community of transportation process from spawning ground, of non‐spawning ground, and of reproductive area in the Kuroshio Extension area) would be a key factor for recruitment prediction.     

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

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

Impact of winter‐to‐spring environmental variability along the Kuroshio jet on the recruitment of Japanese sardine (Sardinops melanostictus)

Winter‐to‐spring variability in sea surface temperature (SST) and mixed layer depth (MLD) around the Kuroshio current system and its relationship to the survival rate (ln [recruit per spawning stock biomass], LNRPS) of Japanese sardine (Sardinops melanostictus) were investigated based on a correlation analysis of data from 1980 to 1995. The data were from a high‐resolution ocean general circulation model using the ‘Kuroshio axis coordinates’, in which the meridional positions are relocated to a latitude relative to the Kuroshio axis at each longitude, rather than the geographically fixed coordinates. A significant positive (negative) correlation between LNRPS and winter MLD (winter–spring SST) was detected near the Kuroshio axis from areas south of Japan (where eggs are spawned) to the Kuroshio Extension (where larvae are transported). This result is in contrast to previous studies using geographically fixed coordinates, which showed a significant correlation predominantly in the area south of the Kuroshio Extension in winter, where at this time few larvae have been found. From the late 1980s to early 1990s, when the survival rate was remarkably low, MLD around the axis was shallow and SST was high. Although MLD and SST show a significant correlation, significant partial correlations were also observed between February MLD and LNRPS when the contribution of SST was excluded, and between March SST and LNRPS when the contribution of MLD was excluded. We presume that MLD shoaling reduced the nutrient supply from deep layers, resulting in less productivity in the spring, and SST warming could have a negative influence on larval growth.     

Variations in food abundance for Japanese sardine larvae related to the Kuroshio meander

Long-term fluctuations of chlorophyll a concentration, and abundance of herbivorous or omnivorous small copepods during the spawning season of the Japanese sardine, Sardinops melanosticta , were examined in relation to the types of Kuroshio meander. The purpose was to clarify the influence of the meander on the production of food organisms for the sardine larvae and their survival in the Pacific coastal region of central Japan. During an A-type meander of the Kuroshio, when the offshore and inshore movements of the Kuroshio path were small except at the beginning of the meander, the surface chlorophyll a concentration in the coastal region was lower than that during a non-A-type meander with frequent and prompt fluctuations of the Kuroshio path. The abundance of small copepods was also low during the A-type meander except in the subsequent spring just after the beginning of an A-type meander, but often high during the non-A-type meander. Prompt fluctuations of the Kuroshio path during the non-A-type meander probably cause local upwellings which stimulate primary and secondary production near the Kuroshio, and may enhance the survival of the sardine larvae. On the contrary, the quasi-stationary state of an A-type meander may be unfavourable for the production of phytoplankton and nauplii of herbivorous or omnivorous small copepods, and therefore the survival of the sardine larvae becomes poor.     

Effect of wind stress on the catch of Japanese anchovy Engraulis japonicus off northwestern Kyushu, Japan

Effect of wind stress on the annual catch of Japanese anchovy Engraulis japonicus off northwestern Kyushu for the period between 1963 and 2009 was investigated. Regime shift analysis detected several step changes in catch and environmental variables. Since the mid-1980s, the anchovy catch in the coastal fishery zones has declined, while the catch in the offshore zone has increased. The decline of catch in the coastal zones showed a significant correlation with the long-term variations in prevailing north-northeastward wind stress over the Goto-Nada Sea during spring spawning season. The results indicated that weakened north-northeastward winds caused the recent low recruitment of anchovy through low levels of wind-induced eggs and larval transport from the offshore spawning ground to the coastal nursery areas, resulting in the potential shift of nursery area to the northwestern offshore region. Thus, as well as the growth-favorable ambient temperature, transport process would play a key role on long-term fluctuations in anchovy abundance in these coastal seas.     

Relationship between recruitment of Japanese sardine (Sardinops melanostictus) and environment of larval habitat in the low‐stock period (1995–2010)

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

Larval fish distribution and retention in the Canary Current system during the weak upwelling season

The spatial distribution of fish larvae was studied in the Canaries‐African Coastal Transition Zone, outside the strong upwelling season. An onshore–offshore transition in the larval fish community structure was observed, from a coastal assemblage dominated by small pelagics (sardine, anchovy, mackerel), bounded by the upwelling front, to an offshore assemblage dominated by mesopelagic species (mainly Myctophidae, Phosichthydae, Gonostomatidae). Distribution of the neritic larvae was deeply influenced by the intense mesoscale activity found in the area, both horizontally (larvae were advected offshore but were always retained within the upwelling area) and vertically (larvae were deepened in the vicinity of two anticyclonic eddies). A combined effect of the upwelling front and a cyclonic–anticyclonic eddy dipole is likely the successful retention mechanism for these larvae. These results support the current belief that retention may be higher than previously thought in upwelling areas. Oceanic larvae were also collected in higher abundances near the front and an anticyclonic eddy. Neritic and oceanic larvae frequently showed a differentiated position in the water column, although they sometimes coexisted. Finally, larval connectivity between Islands within the Canary archipelago is suggested. The present study thus contributes to the understanding of the complex dispersal and retention processes in the Canaries‐African Coastal Transition Zone. However, results also highlight the poor knowledge of this region compared with the other three main Eastern Boundary Upwelling Systems in terms of ichthyoplankton dynamics. The importance of routine monitoring programs of commercial and non‐commercial species in the area is emphasized.     

Simulation of the impact of climate change on the egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island,the western coast of Japan

In order to investigate the impact of climate change on egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island western Japan, we conducted particle‐tracking simulations on transport success/failure to fishing grounds from 1960 to 2007. The modeled transport success since the mid‐1990s increased and decreased in the offshore and coastal zones, respectively, compared with the 1960s and 1970s. The estimated northward shift of the spawning ground and weakened Tsushima Warm Current contributed to increase in modeled transport success to the offshore zone. Conversely, the weakening trend of the modeled onshore current in the Goto‐Nada Sea combined with the northward shift of the spawning ground resulted in unsuccessful larval transport. These results suggest that fluctuations in juvenile and subadult anchovy catches in this area may be attributable to changes in the physical environment. The present study showed that changes in transport success induced by oceanographic fluctuations related to climate change, have the potential to affect anchovy recruitment off the western coast of Japan.     

Entrainment and advection of larval sardine,Sardinops sagax,by the East Australian Current and retention in the western Tasman Front

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.     

A spatial model of population dynamics of the early life stages of Japanese sardine, Sardinops melanostictus, off the Pacific coast of Japan

We constructed a numerical model reproducing the transport, survival and individual growth of the early life stages of Japanese sardine, Sardinops melanostictus, off the Pacific coast of Japan during 1978–93. The causes of early life stage mortality, including the influence of the effects of the spatial relationship between the spawning grounds and the Kuroshio on the mortality rate, were investigated. Survival and transport from egg stage to 60 days after spawning were modelled daily in a 1 × 1 degree mesh cell and individual growth in the period was modelled in each region (Kuroshio, Inshore, Offshore and Transition regions). Individual growth and survival from 60 to 180 days after spawning were modelled daily in the Transition region. Environmental data were taken from outside the model system. Our simulation indicates that survival variability in the larval stage (5–25 mm in standard length) is the key factor in determining the year‐class strength. The simulation revealed that strong year classes occurred with good survival in the spawning ground and whilst entrained in the Kuroshio current being transported to the main feeding grounds in the Transition region. The simulation also indicated that survival rates in 1988–93 were low in the Inshore, Kuroshio and Offshore regions, which depressed the year‐class strength during that period.     

Changes in the distribution of sardine eggs and larvae off Portugal, 1985–2000

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.     

Mesoscale eddies and survival of late stage Pacific sardine (Sardinops sagax) larvae

We examined the distribution of sardine larvae relative to environmental conditions with the purpose of identifying and characterizing habitat that encourages high larval growth and survival, based on the 1983–1998 surveys of the California Cooperative Oceanic Fisheries Investigations (CalCOFI). Long-term averages show that sardine 'survivors' (spatially aggregated larvae ≥ 18 days old) were most abundant offshore, whereas sardine egg density, chlorophyll biomass and zooplankton volume were greatest inshore. In contrast, mesoscale eddies, observed in remotely sensed sea surface temperature imagery, were found only in offshore regions. To further examine the link between eddies – which often result in locally elevated chlorophyll and zooplankton – and sardine survival, we compared the distribution of larvae and eddies survey by survey. Sardine survivors were most abundant offshore in only one-quarter of the research surveys, and when they were most abundant offshore they were associated with eddies. This indicates that the offshore eddy habitat produced exceptionally large numbers of survivors, as evidenced by the disproportionate effect on the long-term average.