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.     

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.     

Empirical biomass model for the Japanese sardine, Sardinops melanostictus, with sea surface temperature in the Kuroshio Extension

An Empirical Biomass Model for the Japanese sardine, Sardinops melanostictus, was developed on the basis of the relationship between February sea surface temperature (SST) in the Kuroshio Extension (30–35°N, 145–180°E) and the mortality coefficient during the period from egg to age 1, observed in 1979–94, to examine the long‐term variation of biomass. The periods of the good and bad catch, the year of the biomass peak, and the speed of the biomass decline in the period from 1957 to 1994 were successfully reproduced, except for the biomass increase in the early 1970s. When the model also included with a density‐dependent effect, the whole history of the observed catch during 1957–94 was almost perfectly reproduced. These results suggest that the environment in the Kuroshio Extension region, represented by winter–spring SST, is regarded as a leading factor for determining fluctuations of the sardine biomass in the long term, and that the density effect has a secondary contribution.     

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.     

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.     

The effect of vertical and horizontal distribution on retention of sardine (Sardinops sagax) larvae in the Northern Benguela – observations and modelling

In the present study, a modelling experiment is conducted to simulate the transport of sardine (Sardinops sagax) eggs and larvae in the Northern Benguela. Based on historical and newly obtained data, different scenarios of vertical and horizontal distribution are applied and the effects on retention are discussed. The simulations showed that vertical and horizontal distribution were important for retention of sardine larvae in the Northern Benguela. By using age‐dependent data on vertical distribution, it was shown that retention of particles in the simulation was substantially enhanced compared with a scenario where particles were distributed in the offshore moving Ekman layer. Retention was lowest during October–December (when upwelling intensity is high) and highest during February–April (when upwelling intensity is somewhat lower). When different spawning areas were considered, highest retention was observed in an area near Walvis Bay. It is concluded that the behaviour of sardine larvae is adapted to the circulation system in the Northern Benguela in a way that promotes retention of the larvae in inshore nursery areas.     

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.     

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.     

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

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

Oil sardine (Sardinella longiceps) off the Malabar Coast: density dependence and environmental effects

Marine fish stocks are known for extensive variation in landings, with temporal fluctuations attributable to density-dependent as well as environmental effects. In this paper we analysed a 44-yr time-series of oil sardine Sardinella longiceps landings from the Arabian Sea off the Malabar Coast of India. Density dependence was detected in the landings of oil sardine, reinforcing the potential for sustainable yields. Significant environmental factors (precipitation and sea level reflecting the strength of upwelling) during the monsoon period (June–August) are hypothesized to affect the dynamics of landings by influencing spawning and recruitment success. Together, density dependence and environmental variability during the monsoon explained 80% of the among-years variance in landings. Our results have important consequences for understanding catch variability and are potentially useful for facilitating management of this commercially important fishery.     

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).     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Assessing the abundance and distribution of eggs of sardine, Sardinops sagax, and round herring, Etrumeus whiteheadi, on the western Agulhas Bank, South Africa, using a continuous, underway fish egg sampler

A continuous, underway fish egg sampler (CUFES) was employed to assess the abundance and distribution of eggs of both sardine, Sardinops sagax , and round herring, Etrumeus whiteheadi , on the Western Agulhas Bank, South Africa, during September 1996. Samples were collected while underway along six inshore/offshore transects, and at stations along the transects. Volumetric estimates of egg density (eggs m⁻³) from on-station CUFES samples were highly correlated with both volumetric and areal (eggs m⁻²) estimates of egg density from samples collected from CalVET net hauls at these stations, demonstrating the validity of this novel sampling technique. Sardine and round herring eggs were encountered in a band running parallel to the coast and extending from 10 to 30 nautical miles offshore to the shelf edge, and highest egg densities were associated with strong north-west-flowing currents in the region of the shelf edge. Collecting samples while underway increased the precision of the estimate of mean egg density for sardine eggs but not for round herring eggs. The use of CUFES in obtaining a fine-scale resolution of sardine egg distribution, and as a tool for stock assessment, are discussed.