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.     

Transport and survival of Japanese sardine (Sardinops melanostictus) eggs and larvae via particle‐tracking experiments

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).     

A modeling approach to evaluate growth and movement for recruitment success of Japanese sardine (Sardinops melanostictus) in the western Pacific

A two‐dimensional individual‐based fish movement model coupled with fish bioenergetics was developed to simulate the observed migration and growth of Japanese sardine (Sardinops melanostictus) in the western North Pacific. In the model, derived from the observed ocean–environmental data as the driving force, fish movement was adapted as a kinesis behavior. The model successfully simulated the observed transport patterns during the egg and larval stages and the northward migrations during the juvenile stage in 2005, 2006 and 2007. The model results showed that both temperature during the larval stage in the Kuroshio Extension and the prey availability during the early juvenile stage in the Kuroshio–Oyashio transitional area are important factors for growth of Japanese sardine. In autumn, the observed juvenile sardine were mainly distributed in the subarctic water region off the Kuril Islands, which is an area (158–165°E, 43–47°N) with a high chlorophyll‐a (Chl‐a) concentration. The model reproduced the fish distribution, which has a high density in this region. The high Chl‐a concentration area in autumn may contribute to increasing the survival rate of Japanese sardine by cascading up the food chain, from the high primary production, and is an important habitat for recruitment success of Japanese sardine.     

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.     

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.     

Non-stationary effects of multiple drivers on the dynamics of Japanese sardine (Sardinops melanostictus,Clupeidae)

Non-stationary driver-response relationships are increasingly being recognized by scientists, underlining that a paradigm shift out of conventional stationary relationships is crucial. Japanese sardine (Sardinops melanostictus, Clupeidae) is a typical small pelagic fish in the northwestern Pacific with considerable fluctuations in productivity, bringing about great economic and ecological concerns. Numerous studies suggest that the population dynamics of Japanese sardine is an integrated process affected by multiple density-dependent, fishing and climatic drivers. However, little has hitherto been done to incorporate the non-stationary effects of multiple drivers, impeding progresses in understanding the population dynamics and in developing management strategies. In this study, we adopted variable coefficients generalized additive models to reveal the non-stationary effects of density dependence, fishing pressure and climatic conditions on the population dynamics of Japanese sardine. Results suggest that the dynamics of Japanese sardine from 1976 to 2018 could be divided into four periods: the 1980s when suitable climatic conditions from strong Siberian High pressure system sustained high abundance; the 1990s when negative density-dependent effects and degrading climatic conditions due to temperature increase led to population collapse; the 2000s when negative triple effects, particularly high fishing pressure, restricted the population increase; and the 2010s when favourable climatic conditions with re-strengthening Siberian High pressure system accompanied by low fishing pressure contributed to the population recovery. The study highlights that precise identifications of population status and climatic conditions are helpful to achieve good trade-offs between resource exploitation and protection and to facilitate ecosystem-based management for Japanese sardine fisheries.     

Environmental effects on recruitment and productivity of Japanese sardine Sardinops melanostictus and chub mackerel Scomber japonicus with recommendations for management

We compared a wide range of environmental data with measures of recruitment and stock production for Japanese sardine Sardinops melanostictus and chub mackerel Scomber japonicus to examine factors potentially responsible for fishery regimes (periods of high or low recruitment and productivity). Environmental factors fall into two groups based on principal component analyses. The first principal component group was determined by the Pacific Decadal Oscillation Index and was dominated by variables associated with the Southern Oscillation Index and Kuroshio Sverdrup transport. The second was led by the Arctic Oscillation and dominated by variables associated with Kuroshio geostrophic transport. Instantaneous surplus production rates (ISPR) and log recruitment residuals (LNRR) changed within several years of environmental regime shifts and then stabilized due, we hypothesize, to rapid changes in carrying capacity and relaxation of density dependent effects. Like ISPR, LNRR appears more useful than fluctuation in commercial catch data for identifying the onset of fishery regime shifts. The extended Ricker models indicate spawning stock biomass and sea surface temperatures (SST) affect recruitment of sardine while spawning stock biomass, SST and sardine biomass affect recruitment of chub mackerel. Environmental conditions were favorable for sardine during 1969–87 and unfavorable during 1951–67 and after 1988. There were apparent shifts from favorable to unfavorable conditions for chub mackerel during 1976–77 and 1985–88, and from unfavorable to favorable during 1969–70 and 1988–92. Environmental effects on recruitment and surplus production are important but fishing effects are also influential. For example, chub mackerel may have shifted into a new favorable fishery regime in 1992 if fishing mortality had been lower. We suggest that managers consider to shift fishing effort in response to the changing stock productivity, and protect strong year classes by which we may detect new favorable regimes.     

Japanese sardine (Sardinops melanostictus) mortality in relation to the winter mixed layer depth in the Kuroshio Extension region

A drastic population change in Japanese sardine (Sardinops melanostictus) has been noted as being related to winter sea surface temperature (SST) in the Kuroshio Extension region. The former studies suggest two possible explanations. One is that temperature itself affects sardine. The other is that SST represents the environmental change of the Kuroshio Extension region and other causes directly affecting sardine. In this study, we found that sardine mortality from post‐larva to age 1 negatively correlated with the winter mixed layer depth (MLD) in the Kuroshio Extension region from 1979 to 1993. During the period of a deep winter mixed layer (during the early 1980s), sardine mortality was low, whereas mortality was high when the winter mixed layer was shallow (during the late 1980s to early 1990s). By using a lower trophic‐level ecosystem model forced by the observed time series of MLD, SST, light intensity and nutrient data, we found that the estimated spring zooplankton density drastically varies from year to year and has a significant negative correlation with sardine mortality. The inter‐annual variation of spring zooplankton density is caused by the winter MLD variation. During the deep winter mixed layer years, a phytoplankton bloom occurs in spring, whereas during the shallow winter mixed layer years, the bloom occurs in winter. The results of our study suggest that the decline in the Japanese sardine population during the late 1980s to early 1990s was due to an insufficient spring food supply in the Kuroshio Extension region where sardine larvae and juvenile are transported.     

Long-term stock assessment and growth changes of the Japanese sardine (Sardinops melanostictus) in the Sea of Japan and East China Sea from 1953 to 2006

We estimated the stock size of Japanese sardine ( Sardinops melanostictus ) in the Sea of Japan and East China Sea since 1953 using cohort analysis based on the changes of growth patterns. Growth of Japanese sardine, estimated by using annual rings on archived scales since 1961, showed that body lengths were extremely stunted in the 1980–1987 year-classes. The body length at age 3 from February to April in the 1980–1987 year-classes, a period when the stock size exceeded 4 million tons, was 180.0 ± 2.6 mm (mean ± SD), and in the other year-classes was 195.1 ± 7.6 mm. The body length at age 3 and wet weight of zooplankton in August in the offshore area of the Sea of Japan had a significantly positive correlation. We assumed three scenarios for maturation ratios, and estimated Ricker's spawner–recruitment relationships. We analyzed the correlations between logarithmic recruitment residuals (LNRR) and environmental factors in winter, represented by the North Pacific index (NPI), Aleutian low pressure index (ALPI), Pacific decadal oscillation (PDO), monsoon index (MOI), Arctic oscillation (AO) and Southern oscillation index (SOI). Significant correlations were observed between MOI and LNRR and between AO and LNRR. A combination of strong MOI and weak AO would increase the biomass of phytoplankton and zooplankton and subsequently increase the recruitment of Japanese sardine.     

Vertical distributions of Japanese sardine (Sardinops melanostictus) eggs: comparison of observations and a wind-forced Lagrangian mixing model

To understand how Japanese sardine eggs vertically disperse in water, we acquired a series of depth‐resolved samples of eggs while tracking a drogued buoy to follow the water mass in which the eggs were spawned. This survey was conducted in early March of 1991, when the sardine population was abundant and actively spawning in the Pacific off Kyushu. Buoyancy of the eggs was estimated by measuring the specific gravity, and was 1–4 × 10^?3 less than that of the water in which they were spawned, giving a rising speed of 1–2 mm s^?1. In an attempt to diagnose the dynamics underlying the observed vertical profiles of the eggs, a one‐dimensional egg distribution model incorporating the egg rising speed and wind‐ and depth‐dependent vertical diffusivity was formulated. Observed eggs were mainly in the surface mixed layer, and their vertical distribution changed with wind‐induced surface mixing: during strong winds, the eggs were distributed relatively evenly over the upper 60–75 m, whereas during weak winds and shallow pycnocline, the eggs formed a subsurface peak around 25 m depth. The model accurately reproduced these variations in the vertical distributions of eggs, suggesting that the gradients of egg concentration are sensitive to the strength of wind and the structure of underlying hydrography. Further tests with the model showed the importance of positive buoyancy: neutrally buoyant eggs formed subsurface maxima below the food‐rich euphotic zone. The balance of vertical mixing and floating, which allows the eggs to remain in the euphotic zone but away from the near‐surface waters, may be an adaptation to minimize surface‐enhanced predation, while allowing the eggs to hatch in the food‐rich euphotic zone.     

Naupliar copepod concentrations in the spawning grounds of Japanese sardine, Sardinops melanostictus, along the Kuroshio Current

Naupliar copepods were distributed at similar concentrations over the waters inshore and offshore of the Kuroshio Current off central Japan in early spring 1993 and 1994, overlapping with the distribution of early feeding larvae of Sardinops melanostictus . Although N, P, Si and chlorophyll a concentrations were higher in the waters inshore of the Kuroshio axis than in the offshore waters, the mean concentrations of nauplii were not statistically different between the two waters. Food availability for larval S. melanostictus did not seem to be different between the two waters in terms of the mean food concentrations. Using the critical food concentration (>9 nauplii L^–1) for 25% survival during 3 days after first feeding derived from a rearing experiment, percentages in number of stations or water samples with nauplii concentrations >9 nauplii L^–1 were higher in the inshore waters than in the offshore waters in both years examined. Considering that fish larvae may depend on small-scale patchiness of food for their survival, the inshore waters seemed to be more favourable for first-feeding larvae than the offshore waters.     

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.     

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.     

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.     

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.     

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.     

The recruitment process of the Barents Sea capelin (Mallotus villosus) stock, 2001–2003

Oceanographic and predation processes are important modulators of fish larvae survival and mortality. This study addresses the hypothesis that immature Norwegian spring‐spawning herring (Clupea harengus), when abundant in the Barents Sea, determine the capelin reproduction success through consumption of Barents Sea capelin (Mallotus villosus) larvae. Combining a hydrodynamic model and particle‐tracking individual‐based model, a realistic spatio‐temporal overlap between capelin larvae and predatory immature herring was modelled for the summer seasons of 2001–2003. Capelin larvae originating from western spawning grounds became widely dispersed during the summer season, whereas those originating from eastern spawning grounds experienced a rapid drift into the southeastern Barents Sea. Herring caused a 3% mortality of the capelin larvae population in 2001 and a 16% mortality in 2003, but the effect of predation from herring on capelin larvae was negligible in 2002. Despite a strong capelin larvae cohort and a virtual absence of predatory herring, the recruitment from the capelin 2002 year class was relatively poor from a long‐term perspective. We show that the choice of capelin spawning grounds has a major impact on the subsequent capelin larvae drift patterns, constituting an important modulator of the capelin larvae survival. Variation in drift patterns during the summer season is likely to expose the capelin larvae to a wide range of hazards, including predation from young cod, sandeel and other predators. Such alternative predators might thus have contributed to the poor capelin recruitment during 2001–2003, leading to the collapse of the capelin stock in the subsequent years.     

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.