Spawning patterns provide further evidence for multiple stocks of sardine (Sardinops sagax) off eastern Australia

Previous studies have suggested that sardine, Sardinops sagax, off eastern Australia spawns across its entire range when habitat conditions are suitable. However, recent studies have suggested that separate sub‐populations and spawning groups may occur in the region. Spawning patterns off eastern Australia were investigated using data collected during nine ichthyoplankton surveys conducted between 1997–2015, and adult reproductive data obtained from ad hoc commercial sampling off New South Wales (NSW). The egg surveys covered the known distribution of sardine off eastern Australia and included year‐round sampling in the northern and southern parts of this range. Egg distributions and analysis of gonadosomatic indices identified two spatio‐temporally separate spawning groups; one occurring off southern Queensland to northern NSW during late winter and early spring, and a smaller group off eastern Tasmania to southern NSW during summer. Most eggs were collected from waters 50–90 m deep, with sea surface temperatures of 18–23°C. Additive modelling indicated depth was the most significant factor driving selection of spawning habitat, followed by the interaction of month and latitude. Low egg densities were recorded in waters between 34–37°S, despite conditions within the ranges suitable for spawning. The presence of two spawning groups of sardine off eastern Australia supports recent findings that two sub‐populations occur in the region. Findings of this study will help to optimise the spatio‐temporal extent of future egg surveys and further confirm the need to coordinate future management of each sub‐population among relevant jurisdictions.     

Remotely sensed spawning habitat of Pacific sardine (Sardinops sagax) and Northern anchovy (Engraulis mordax) within the California Current

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.     

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.     

Spatial and temporal distribution and abundance of European hake, Merluccius merluccius, eggs and larvae in the Catalan coast (NW Mediterranean)

Plankton hauls were conducted on five surveys from November 1998 to November 1999 to study the spatial distribution of eggs and larvae of European hake off Catalonia (NW Mediterranean Sea). Also CTD casts to record hydrographic parameters were carried out on a closely spaced station grid. Merluccius merluccius eggs and larvae appeared mainly in late spring, summer and autumn surveys and were very scarce in winter. Strong differences in terms of egg and larval densities were observed between the two November surveys, which could be attributed to the anomalous hydrographic situation during November 1998. M. merluccius egg and larvae were mainly distributed over the continental shelf, with peak abundances between the 100 m isobath and the edge of the shelf. On the evidence of larval size frequency distributions in the different sampling sectors and the closely overlapping distribution patterns for the eggs and the adult spawning stock, drifting of hake eggs and larvae was not a major factor. The larval distribution extended only slightly further offshore than the egg distribution. Using the hydrographic information and the larval distribution data, an attempt was made to relate the different seasonal productivity levels over the spawning period and the distribution of the larvae.     

Habitat associations of dolphinfish larvae in the Gulf of Mexico

The aim of this study was to investigate the role of the northern Gulf of Mexico (Gulf) as essential habitat of common dolphinfish (Coryphaena hippurus) and pompano dolphinfish (C. equiselis) during early life and to characterize conditions in which they were found. Eight ichthyoplankton surveys were conducted during the summer months from 2007 to 2010 in northern Gulf waters off Texas and Louisiana (26–28°N, 87–93°W), during which 1145 dolphinfish larvae were collected. C. hippurus, the dominant dolphinfish species, were collected in 57% of sampling sites with an overall mean density of 0.73 larvae 1000 m^?3, whereas C. equiselis were only collected at 18% of sites with a mean density of 0.12 larvae 1000 m^?3. Generalized additive models (GAMs) were used to examine the relationship between several environmental parameters and C. hippurus presence/absence and density. Both presence/absence and density GAMs indicated that catches of C. hippurus larvae increased near fronts and eddies and that increased abundances were most strongly associated with higher salinities and cooler temperatures. In addition, our models showed that C. hippurus larvae were positively associated with C. equiselis larvae, suggesting that C. hippurus and C. equiselis use similar habitats. Results of this study indicate that the Gulf may represent important spawning and/or nursery habitat of dolphinfishes and that mesoscale features and physicochemical conditions influence the distribution and abundance of C. hippurus larvae in this region.     

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.     

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.     

Egg and larval distributions of seven fish species in north-east Atlantic waters

The distribution of egg and larvae of mackerel, horse mackerel, sardine, hake, megrim, blue whiting and anchovy along the European Atlantic waters (south Portugal to Scotland) during 1998 is described. Time of the year, sea surface temperature and bottom depth are used to define the spawning habitat of the different species. Mackerel, horse mackerel, and sardine eggs and larvae presented the widest distribution, whereas megrim and anchovy showed a limited distribution, restricted to the Celtic Sea and the Bay of Biscay respectively. Correspondingly mackerel, horse mackerel and sardine showed the highest aggregation indices. Blue whiting larvae were found at the lowest temperatures, whereas anchovy eggs and larvae were found in the warmest waters. The analysis is a basis for evaluation of ongoing changes in the pelagic ecosystem of the north‐east Atlantic.     

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.     

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

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

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.     

Relative importance of gulf and shelf waters for spawning and recruitment of Australian anchovy, Engraulis australis, in South Australia

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.     

Fine scale spatial pattern of Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs

Pacific sardine (Sardinops sagax) and northern anchovy (Engraulis mordax) eggs exhibited different spatial structure on the scale of 0.75–2.5 km in two egg patches sampled in the Southern California Bight in April 2000. Plankton samples were collected at 4‐min intervals with a Continuous Underway Fish Egg Sampler (CUFES) on 5 × 5 km grids centered on surface drifters. Variograms were calculated for sardine and anchovy eggs in Lagrangian coordinates, using abundances of individual developmental stages grouped into daily cohorts. Model variograms for sardine eggs have a low nugget effect, about 10% of the total variance, indicating high autocorrelation between adjacent samples. In contrast, model variograms for anchovy eggs have a high nugget effect of 50–100%, indicating that most of the variance at the scales sampled is spatially unstructured. The difference between observed spatial patterns of sardine and anchovy eggs on this scale may reflect the behavior of the spawning adults: larger, faster, more abundant fish may organize into larger schools with greater structure and mobility that create smoother egg distributions. Size and mobility vary with population size in clupeoids. The current high abundance of sardines and low abundance of anchovy off California agree with the greater autocorrelation of sardine egg samples and the observed tendency for locations of anchovy spawning to be more persistent on the temporal scale of days to weeks. Thus the spatial pattern of eggs and the persistence of spawning areas are suggested to depend on species, population size and age structure, spawning intensity and characteristic physical scales of the spawning habitat.     

Historical fluctuations in spawning location of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during 1967–73 and 2000–2004

The spatial extent of small pelagic fish spawning habitat is influenced by environmental factors and by the state of the adult population. In return, the configuration of spawning habitat affects recruitment and therefore the future structure of the adult population. Interannual changes in spatial patterns of spawning reflect variations in adult population structures and their environment. The present study describes the historical changes in the spatial distribution of spawning of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during two periods: 1967–72 and 2000–2004. Using data from egg surveys conducted in spring, the spatial distributions of anchovy and sardine eggs are characterized by means of geostatistics. For each survey, a map of probability of egg presence is constructed. The maps are then compared to define (1) recurrent spawning areas, (2) occasional spawning areas and (3) unfavourable spawning areas during each period. Sardine spawning habitat is generally fragmented and appears spatially limited by the presence of cold bottom water. It is confined to coastal or shelf break refuge areas in years of restricted spawning extent. For anchovy, recurrent spawning sites are found in Gironde and Adour estuaries whilst spawning can extend further offshore in years of more intense spawning. For both species, the mean pattern of spawning has changed between 1967–72 and 2000–2004. Noticeably, the spatial distribution of anchovy eggs in spring has expanded northward. This trend possibly results from changes in environmental conditions during the last four decades.     

Evidence of density-dependent effects on population variation of Japanese sardine (Sardinops melanosticta) off Korea

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.     

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.     

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.     

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.     

Transport and survival processes of eggs and larvae of jack mackerel <Emphasis Type="Italic">Trachurus japonicus</Emphasis> in the East China Sea

ABSTRACT:   Recent surveys showed substantial aggregation of larvae of jack mackerel in the southern East China Sea, indicating intensive spawning grounds near Taiwan. A numerical model was applied to investigate transport and survival processes of eggs and larvae of jack mackerel from the spawning area to the nurseries. The results show that: (i) the distributions of larvae simulated by the model agreed well with those obtained by field survey; (ii) the stock of jack mackerel in the Sea of Japan is composed of both groups from north of Taiwan and from the western coast of Kyushu. It takes more than two months for the former to reach the Sea of Japan, while it is within 40 days for the latter; and (iii) large proportions of the eggs and larvae spawned off the north of Taiwan are transported rapidly to the Pacific side of Kyushu by the Kuroshio Current, and the rest slowly to the east or north-east along the continental slope in the East China Sea. In contrast to the larval flux, survivors are more abundant in the northern East China Sea than in the Pacific Ocean, indicating that survival in the northern East China Sea would determine the jack mackerel stock in Japan.     

Modeling the transport and survival of Japanese sardine larvae in and around the Kuroshio Current

We have numerically modeled the advection and diffusion of sardine eggs and larvae to investigate the larval transport processes of Japanese sardine from the spawning grounds by the Kuroshio.
The results indicated that the offshore drift current induced by the winter monsoon and the location of the spawning ground have significant effects on the survival of the Japanese sardine. The contribution of the drift current, the distance of the spawning ground from the Kuroshio axis, and the eddy diffusivity to the larval retention in the coastal area is approximately expressed by the following equation: where R is the retention rate in the coastal area, a the variance of initial distribution of eggs, T the time after the eggs were spawned, – V₀ the velocity of the wind-induced offshore current, y₀ the distance of the center of the spawning area from the Kuroshio axis, and K the coefficient of horizontal eddy diffusivity.
The year-to-year variation in larval survival rates stimulated by the two-dimensional model are consistent with those estimated previously by using field data of egg and larval abundance during 1978–1988.