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.     

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.     

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

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

Biophysical dynamics of western transition zones: a preliminary synthesis

The nature of the western portions of the biogeographic temperate or transition zones in the North Pacific and North Atlantic is reviewed. The physical transport of nutrients and biomass into them from the Kuroshio and Gulf Stream as well as from the poleward sides are estimated. The conclusion is that the upwelling in the two western boundary currents makes the largest contribution to the nutrient and biomass fluxes into these transition zones. A conservative estimate of the amount of upwelled fluid is derived from absolute velocity sections in the Gulf Stream. The estimate suggests that upwelling into the euphotic zone exceeds 2 × 10⁶ m³ s^–1. This implies that upwelling in these western boundary currents matches or exceeds that in eastern boundary currents such as the California Current. The two western boundary regimes have very different poleward situations. The Oyashio extension flows parallel to the Kuroshio and is a deep current. The North Atlantic Shelf Front flow is to the west where it is ultimately entrained into the edge of the Gulf Stream. There does not seem to be any tendency for this to occur in the Kuroshio. Despite these differences in the northern and western boundaries, the two transition zones are similar with large amplitude meanders, anticyclonic rings and streamers dominating their physical structure. The physical features responsible for the transfer of materials from the boundary current extensions into the transition zones are similar in both systems. Ring formation contributes only ? 10% of the transfer, while ring‐induced streamers contribute 30%. The rest of the transport is contributed by branching of the boundary current front. Both currents have well developed secondary fronts consisting of subtropical surface water pulled into the transition zone. Biologically, the upwelling in both western boundary currents leads to a biomass maximum along the boundary in both secondary producers (copepods) and in small pelagic fish. In the Kuroshio, the latter are the Japanese sardine, Sardinops melanostictus, that spawn in the Kuroshio and then enter the transition zone for the summer and fall months. In the Gulf Stream, the dominate species are menhaden, Brevoortia tyrannus and B. smithi. These species make use of the coastal environments of North America and although the adults spawn in the Gulf Stream, they are not thought to play a major role in the Slope Water, transition zone. The similar differences in the use of the Kuroshio and the Gulf Stream ecosystems occurs in the behaviour of bluefin tuna, squid and other large pelagics. The Gulf Stream system also lacks an equivalent to Pacific saury, Cololabis saira. The biology therefore is at least subtly different, with saury and sardines being replaced by mid‐water fish in the North Atlantic. A fuller comparison of the biology with quantitative methods in both systems should be encouraged.     

Sardine and anchovy regime fluctuations of abundance in four regions of the world oceans: a workshop report

Regimes of high abundance of sardine (Sardinops sagax and Sardina pilchardus) have alternated with regimes of high abundance of anchovy (Engraulis spp.) in each of the five regions of the world where these taxa co-occur and have been extensively fished. When one taxon has been plentiful, the other has usually been at a reduced level of abundance, and vice versa. Changes in the four heavily fished regions that support S. sagax–the Japanese, Californian, Humboldt, and Benguela systems–from a regime dominated by one taxon to a high level of abundance of the other have occurred more or less simultaneously. In the Pacific Ocean, sardines have tended to increase during periods of increasing global air and sea temperatures and anchovies to decrease. The Japanese system is dominated by sardines to a greater extent than the other systems, and sardines off Japan appear to increase as the Kuroshio Current cools. At the eastern edge of the Pacific Ocean, sardines colonize cooler areas during periods of warming. The Benguela system is out of phase with the three Pacific systems. The four systems all appeared to be in a state of flux in the 1980s. Increased abundance of the subdominant taxon is often one of the first signs of change. Sardines are relatively sedentary in refuge areas when scarce but change behavior to become highly migratory and colonize cooler areas when abundant. Anchovies, by contrast, expand around a fixed geographic center.     

Interannual variability of the ecosystem of the Kii Channel, the Inland Sea of Japan, as influenced by bottom intrusion of cold and nutrient-rich water from the Pacific Ocean, and a recent trend of warming and oligotrophication

Interannual variability of the ecosystem of the Kii Channel, productive shelf water on the Pacific side of south‐western Japan, was analysed based on physicochemical environmental variables and abundance of major zooplankton taxa collected monthly for 12 yr from 1987 to 1999. The Kii Channel experienced both short‐term (i.e. 3–4 yr) cyclical changes and a long‐term (i.e. decadal) environmental trend. The short‐term variability was primarily associated with year‐to‐year differences in intrusion of subsurface, cold nutrient‐rich water along the bottom of the Kii Channel from the Pacific Ocean. When this bottom intrusion was intense, the Kii Channel experienced a cold, new production‐dominated ecosystem. The bottom intrusion, however, has become less intense in recent years, because of the closer proximity of the Kuroshio flow axis to the Kii Channel. Hence, there was a consistent trend towards warming and a regenerated production‐dominated ecosystem. In accordance with such environmental changes, the zooplankton community shifted towards more oceanic conditions; major herbivorous calanoids (i.e. Calanus sinicus, Paracalanus parvus (s.l.), Clausocalanus spp. and Acartia omorii) decreased, while carnivores (i.e. hydromedusae and Sagitta spp.) increased. The amount of total fish catch also decreased, while the catch of subtropical species increased. These findings lead us to conclude that the bottom intrusion from the Pacific Ocean plays a key role in determining the biological production in the Kii Channel.     

The abundance of juvenile yellowtail (Seriola quinqueradiata) near the Kuroshio: the roles of drifting seaweed and regional hydrography

We assess the effect of drifting seaweed (Sargassum sp.) biomass, geography and hydrography on juvenile yellowtail (Seriola quinqueradiata) abundance variation off the southeast coast of Japan, near the Kuroshio Current. The amount of drifting seaweed mats progressively increased northeastward into the cooler, coastal waters. Frontal structure indexed using a station‐to‐station ΔSST did not explain spatial variation in the seaweed mat distribution, although the western extent of the Kuroshio Current appeared to act as a boundary. Juvenile yellowtail constituted 51–62% of the fish collected in association with drifting seaweed mats in April 1996 and 1997 and 29% in June 1996. The abundance of juvenile yellowtail was positively correlated with seaweed biomass. The geographic distribution of juvenile yellowtail associated with drifting mats varied among sampling periods, being more southwesterly in April and more northeasterly in June. Simple multiple regression models based on seaweed biomass and geographic distribution (latitude) explained between 35% and 43% of the variation in juvenile yellowtail abundance in spring. Associations with spatial and temporal variations in hydrographic conditions did not contribute to explained variation in a meaningful manner. The results presented here indicate that, off the southeast coast of Japan during April, yellowtail juveniles are likely to be most abundant when seaweed biomass is high, occur offshore, and are bounded by the western extent of the Kuroshio Current near the 19–20°C SST isotherm.     

Effect of horizontal advection induced by intrusion of the Kuroshio water on plankton biomass in the spring fishing period of shirasu on the Pacific coast of Japan

ABSTRACT:   This study explored the fundamental process that controls interannual change in plankton biomass on the Pacific coast of Japan, focusing on the spring period of shirasu fishery. A 1-D model of primary production revealed that the strength of horizontal advection induced by warm water intrusion from the Kuroshio into the Pacific coast is the most critical for plankton biomass compared with other factors such as diffusion and production. This conclusion was also confirmed by a sensitivity analysis of the model.     

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.