Occurrence and density of Pacific saury Cololabis saira larvae and juveniles in relation to environmental factors during the winter spawning season in the Kuroshio Current system

The occurrence and density of Pacific saury Cololabis saira larvae and juveniles were examined in relation to environmental factors during the winter spawning season in the Kuroshio Current system, based on samples from extensive surveys off the Pacific coast of Japan in 2003–2012. Dense distributions of larvae and juveniles were observed in areas around and on the offshore side of the Kuroshio axis except during a large Kuroshio meander year (2005). The relationships of larval and juvenile occurrence and density given the occurrence to sea surface temperature (SST), salinity (SSS), and chlorophyll‐a concentration (CHL) were examined by generalized additive models for 10‐mm size classes up to 40 mm. In general, the optimal SST for larval and juvenile occurrence and density given the occurrence was consistently observed at 19–20°C. The patterns were more complex for SSS, but a peak in occurrence was observed at 34.75–34.80. In contrast, there were negative relationships of occurrence and density given the occurrence to CHL. These patterns tended to be consistent among different size classes, although the patterns differed for the smallest size class depending on environmental factors. Synthetically, the window for spawning and larval and juvenile occurrence and density seems to be largely determined by physical factors, in particular temperature. The environmental conditions which larvae and juveniles encounter would be maintained while they are transported. The survival success under the physically favorable but food‐poor conditions of the Kuroshio Current system could be key to their recruitment success.     

Chlorophyll a variation in the Kuroshio Extension revealed with a mixed-layer tracking float: implication on the long-term change of Pacific saury (Cololabis saira)

Variation of chlorophyll a from March 2004 to July 2005 in the formation region of Subtropical Mode Water in the Kuroshio Extension was observed with a mixed‐layer tracking profiling float parking at around 40 m depth. Chlorophyll a concentration in the mixed‐layer is seasonally high from winter to early spring (January–April) even in the deep mixed layer; whereas during winter it tends to be lower for the corresponding deeper mixed layer. The chlorophyll a integrated over the winter mixed‐layer depth (MLD) increases with MLD for MLD <200 m and for MLD > 250 m the integrated chlorophyll a almost disappears, probably because of the large MLD exceeds in the critical depth with light limitation. These results suggest that the wintertime MLD influences the recruitment of Pacific saury (Cololabis saira) in the Kuroshio Extension because the wintertime food environment for larvae and juvenile of the Pacific saury could be better in the shallow wintertime MLD period of the 1950s and 1990s and worse in the deep MLD period of the 1970s and early 1980s.     

Oceanographic factors affecting interannual recruitment variability of Pacific saury (Cololabis saira) in the central and western North Pacific

Pacific saury (Cololabis saira) has a short life span of 2 years and tends to exhibit marked population fluctuations. To examine the importance of sea surface temperature (SST) and mixed layer depth (MLD) as oceanographic factors for interannual variability of saury recruitment in early life history, we analyzed the relationship between abundance index (survey CPUE (catch per unit of effort)) of age‐1 fish and the oceanographic factors in the spawning and nursery grounds of the previous year when they were born, for the period of 1979–2006, in the central and western North Pacific. Applying the mixture of two linear regression models, the variability in the survey CPUE was positively correlated with previous year's winter SST in the Kuroshio Recirculation region (KR) throughout the survey period except 1994–2002. In contrast, the survey CPUE was positively correlated with the previous year's spring MLD (a proxy of spring chlorophyll a (Chl‐a) concentration) in the Kuroshio‐Oyashio Transition and Kuroshio Extension (TKE) during 1994–2002. This period is characterized by unusually deep spring MLD during 1994–1997 and anomalous climate conditions during 1998–2002. We suggest that saury recruitment variability was generally driven by the winter SST in the KR (winter spawning/nursery ground), or by the spring Chl‐a concentration (a proxy of prey for saury larvae) in the TKE (spring spawning/nursery ground). These oceanographic factors could be potentially useful to predict abundance trends of age‐1 saury in the future if the conditions leading to the switch between SST and MLD as the key input variable are elucidated further.     

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.     

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.     

Study on the potential of suspended longline mariculture of the scallop Chlamys farreri in offshore areas

The growth and survival of the scallop Chlamys farreri exposed to different initial densities were evaluated in culture systems from inshore and offshore areas of Sungo Bay, China, from May 2007 to March 2008. Water quality data (e.g. temperature, salinity, current speed, chlorophyll a and total particulate material) were measured concurrently. The C. farreri reached the market size after 10 months of cultivation. Average survival rates for the same initial densities were lower in the inshore area than those in the offshore area. Survival rates also were inversely proportional to the initial densities. Scallops in offshore area with initial densities of 20 and 30 individuals per disc resulted in significantly greater shell heights, soft tissue weights and muscle weight compared with the other experimental groups. Tissue weight increased rapidly from May to September. However, scallops in offshore area had a second growth peak during October to November. Scallops in the offshore area had higher growth and survival rates, which may be attributed to high water current speed resulting in increased food supplies. Results indicate that the scallop C. farreri has an aquaculture potential in offshore cultivation systems.     

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

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

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.     

Habitat use and movement patterns of small (age‐0) juvenile Pacific bluefin tuna (Thunnus orientalis) relative to the Kuroshio

The habitat use of Pacific bluefin tuna (Thunnus orientalis; PBF) in nursery waters off the southern coast of Japan was investigated using archival tags over a 3 year study period (2012–2015), and the data were used to examine the free‐ranging habitat preferences of PBF and the relationship between their horizontal movements and the path of the Kuroshio off the Pacific coast of Japan. The path of the Kuroshio fluctuated seasonally, leading to changes in water temperature that strongly influenced the habitat use of small PBF (2–3 months after hatching). Most PBF were present in coastal waters inshore of the path of the current, and their habitat use changed in response to the distance of the current from the coast. The Kuroshio typically flowed along the coast from summer to autumn, and PBF remained in the coastal waters off Kochi Prefecture during this period. In contrast, PBF quickly moved eastward in winter when the current moved away from the coast. Throughout the winter and spring, the area of habitat use extended widely from the eastern end of the southern coast of Japan (the Boso Peninsula) to the offshore Kuroshio‐Oyashio transition region. These findings suggest that the seasonal habitat use and movement behavior of juvenile PBF are influenced by the distance of the Kuroshio axis from the coast, and the ultimate drivers are likely variations in oceanographic conditions and prey availability along the southern coast of Japan.     

Variations in the abundance of Pacific saury (Cololabis saira) from the northwestern Pacific in relation to oceanic-climate changes

Effects of oceanic-climate changes on the abundance of Pacific saury (Cololabis saira) in the northwestern Pacific during the last half century were investigated. Abundance indices of both large and medium size groups exhibit interannual–decadal variations, but their patterns were different. The large and medium size groups of saury are corresponding to the recruitments of winter- and spring-cohort, respectively. The abundance of large size group saury was significantly correlated with the winter sea surface temperature (SST) in the Kuroshio region, whereas the medium size group saury showed high correlations with SST in the Kuroshio–Oyashio transition zone and the Oyashio region, indicating that the two size groups are affected by subtropical and subarctic environment, respectively. Significant negative correlation between the abundance index and the southern oscillation index (SOI) suggested that El Niño-southern oscillation (ENSO) events have marked impacts on the large size group saury. Subtropical high pressure index and far east zonal index also show high correlations with the abundance of both large and medium size group saury, indicating a linkage between large-scale atmospheric circulation and the abundance of saury. These correlations demonstrate that the abundance of Pacific saury is directly affected by the SST fields through large-scale atmosphere–ocean interactions from the equatorial Pacific to mid- and high-latitude areas such as El Niño events.     

Modelling the impacts of environmental variation on habitat suitability for Pacific saury in the Northwestern Pacific Ocean

We developed habitat suitability index (HSI) models for two size classes of Pacific saury Cololabis saira in the Northwestern Pacific Ocean. Environmental data, including sea surface temperature, sea surface height, salinity, and net primary production, and catch and effort data from Taiwanese distant‐water stick‐held dip net fisheries during the main fishing season (August–October) during 2002–2015 were used. Habitat preferences and suitable habitat area differed between size classes. The suitable habitat was located between 40–47.5°N and 145–165°E for large‐sized Pacific saury but encompassed a greater area (35–47°N and 140–165°E) for medium‐sized Pacific saury. Both size classes were affected by substantial interannual variation in the environmental variables, which in turn can be important in determining the potential fishing grounds. We found a significant negative relationship between the suitable habitat area and the Niño3.4 indices with a time‐lag of 6 months for the large‐sized (r = ?0.68) and medium‐sized (r = ?0.42) Pacific saury, respectively, as well as the total landings of Pacific saury by all fishing fleets (r = ?0.46). As remotely‐sensed environmental data become increasingly available, HSI models may prove useful for evaluation of possible changes in habitat suitability resulting from climate change or other environmental phenomena and in formulating scientific advice for management.     

On the relationship between the Oyashio front and saury fishing grounds in the north-western Pacific: a forecasting method for fishing ground locations

The relationships between fishing ground locations of Pacific saury (Cololabis saira) and the two Oyashio fronts, the offshore front between 146^oE and 155^oE and the coastal Oyashio intrusion, before the saury fishing season, were examined using data from 1971 to 1991. Interannual geographical shifts of both the offshore Oyashio front and saury fishing grounds had a dominant interdecadal fluctuation. In years when the offshore front shifted north (south), the fishing grounds were formed relatively nearshore (offshore). When the offshore front shifted north, the fishing grounds were formed further nearshore in years the coastal Oyashio intrusion extended south. When the offshore front shifted south, on the other hand, the southward extension of the coastal intrusion did not necessarily cause formation of coastal fishing grounds. These results showed that locations of the fishing grounds depend not only on local and instantaneous oceanographic environments around the fishing grounds, but also on oceanographic conditions over an extensive range of the Oyashio area. This might indicate that the ecology of the saury's northward migration, through mixed water regions between the Kuroshio and Oyashio fronts in spring and summer, has a close relation to the shift of the offshore Oyashio front over a time range of months. A practical forecasting method for locations of saury fishing grounds is proposed based on the oceanography before the fishing season.     

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.     

Amazon River water in the northeastern Caribbean Sea and its effect on larval reef fish assemblages during April 2009

During April to June 2009, a large bolus of Amazon River water impacted the northeastern Caribbean Sea. Shipboard observations collected near Saba Bank, the U.S. and British Virgin Islands, and the Anegada Passage showed low surface salinity (35.76 ± 0.05 Practical Salinity Unit (PSU)), elevated surface temperature (26.77 ± 0.14°C), high chlorophyll‐a (1.26 ± 0.21 mg m^?3) and high dissolved oxygen (4.90 ± 0.06 mL L^?1) in a 20‐ to 30‐m thick surface layer in the riverine plume. The water was ~1°C warmer, 1 PSU fresher, 0.3 mL L^?1 higher in oxygen and 1.2 mg m^?3 higher in chlorophyll‐a than Atlantic Ocean waters to the north, with Caribbean surface waters showing intermediate values. Plankton net tows obtained in the upper 100 m of the water column revealed larval fish assemblages within the plume that were significantly different from those of the surrounding waters and from those encountered in the area in previous years. The plume waters contained higher concentrations of mesopelagic fish larvae from the families Myctophidae and Nomeidae, which as adults typically inhabit offshore, deep water habitats. Concentrations of larvae from inshore and reef‐associated families such as Scaridae, Serranidae, Labridae and Clupeidae were lower than those found outside the plume in similar shallow areas, particularly in near‐surface waters. An event like the one observed in 2009 had not been documented in at least the past 30 yr, and yet it was followed by another similarly extreme event in 2010. The ecological implications, including any long‐term consequences of such recent extreme events, are important and merit further study.     

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.     

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.     

Initial design for a fish bioenergetics model of Pacific saury coupled to a lower trophic ecosystem model

A fish bioenergetics model coupled with an ecosystem model was developed to reproduce the growth of Pacific saury. The model spatially covers three different oceanographic spatial domains corresponding to the Kuroshio, Oyashio, and interfrontal (mixed water) regions. In this coupled model, three (small, large, and predatory) zooplankton densities which were derived from the lower trophic level ecosystem model were input to the bioenergetics model of saury as the prey densities. Although certain model parameters were imposed from other species’ bioenergetics, several model parameters were estimated from observational data specific to Pacific saury. The integrated model results reproduced appropriate growth rates of Pacific saury. Model sensitivities to water temperature and prey density are examined and observational methods to evaluate the model parameters are discussed.     

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.     

Growth and survival of the winged pearl oyster Pteria sterna (Gould, 1851) in suspended culture in the tropical Eastern Pacific: Influence of environmental factors

The growth, survival and influence of environmental factors were analysed in two cohorts of cultured Pteria sterna in Ayangue Bay, Province of Santa Elena, Ecuador (tropical Eastern Pacific). Juveniles representing cohorts I and II (8.4 ± 0.54 and 5.0 ± 0.17 mm in dorso‐ventral axis) were deployed in November 2015 and February 2016, and grown in pearl nets suspended in a long line for 12 and 10 months respectively. The stocking density was monthly and bi‐monthly reduced during sampling of individuals to determine growth in dorso‐ventral shell axis, dry mass of shell, soft tissues and dry mass of fouling on shell. Water temperature, salinity, total seston and phytoplankton biomass (chlorophyll a) were determined at the culture site. Results showed that P. sterna reached ~100 mm in length during the first year of culture. Although little negative influence of environmental factors was detected, high temperatures during the reproduction period can be the most negative influential trait. The highest tissue mass (6 g), which occurred at the 10th month of cultivation, as well as a high availability of spat by artificial collectors in the coastal waters, showed that the species can be considered a good candidate for aquaculture in the tropical eastern Pacific.     

Dual effects of reversed winter–spring temperatures on year‐to‐year variation in the recruitment of chub mackerel (Scomber japonicus)

To clarify the effects of temperature on the recruitment of chub mackerel (Scomber japonicus) in the North Pacific, we investigated the influence of winter surface temperature (WST) on spawners at the time of maturity around the spawning grounds and the influence of ambient spring temperature on larvae using estimated temperature (ET) obtained from particle tracking experiments. We found a significant positive correlation between ET approximately 10 days following hatching and the recruitment per spawning stock biomass (RPS) after 2000. The closer (more meandering) the Kuroshio Current (KC) axis was in relation to the spawning ground, the higher (lower) the spring surface temperature and the higher (lower) RPS was in the spawning ground. In contrast, WST inside KC near the maturity/spawning ground was significantly negatively correlated with RPS. A significant negative correlation between the temperatures in winter and spring was detected in the area after 2000, when the conditions of the Pacific decadal oscillation index and the stability of the Kuroshio Extension were synchronous, indicating that KC shifted northward during this time. The reversed temperature pattern was consistent with the winter–spring movement of KC axis in the offshore direction and was correlated with the winter–spring difference in the intensity of the Aleutian low. These results suggest that the annual variation in chub mackerel recruitment after 2000 was strongly affected by the combined effects of ambient temperature because of the reversal of conditions that occurred between winter and spring around the maturity/spawning ground, which was related to the KC path.