Effects of temperature and food availability on growth rate during late larval stage of Japanese anchovy (Engraulis japonicus) in the Kuroshio–Oyashio transition region

During periods when the population size of Japanese anchovy Engraulis japonicus is large, the abundance of 1‐yr olds has been considered to be dependent on the growth and survival processes in the late larval and early juvenile stages in the Kuroshio–Oyashio transition region off northern Japan. Recent growth rates for 10 days before capture of larval and early juvenile E. japonicus were estimated and examined in relation to the surface water temperature and the available copepod density in 1997, 1998 and 1999. Late larval and early juvenile E. japonicus were distributed in the waters with temperature from 15 to 19°C and available prey density from 10 to 1000 mg dry weight (DW) m^?2 in the transition region. The late larval growth rates were found to be regulated more strongly by water temperature than by copepod density in the waters <16°C, and more strongly by copepod density than water temperature in the waters <100 mg DW m^?2 in the Kuroshio–Oyashio transition region. The recent growth rates decreased from the western waters to the eastern waters in the survey area 140–170°E in 1998, correlating with decreases of food availability to 50–100 mg DW m^?2. While in 1999, the recent growth rates were faster in the waters east of 150°E, resulting from eastward expansion of warm water ranges and high available prey density 100–400 mg DW m^?2. The key environmental factors regulating late larval growth rate of E. japonicus in the transition region seem to be spatially different between years.     

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.     

Effects of juvenile size at release and early marine growth on adult return rates for Hokkaido chum salmon (Oncorhynchus keta) in relation to sea surface temperature

Using path analyses, we investigated relationships between size at release from hatcheries, the early marine growth of juveniles, and adult return rates for chum salmon from five river stocks of Hokkaido, Japan, in relation to sea surface temperature during ocean residence. Marine growth was estimated using scales collected from 11 760 adults of age 0.3 (1980–2004). The growth and survival of each stock appeared to have a different suite of regulatory processes. Interannual variability in return rates was mainly regulated by size at release in two stocks from the Sea of Okhotsk. A similar relationship was found in one stock from the Sea of Japan, but growth during coastal residency also affected their return rates. In two stocks from the Pacific coast of Hokkaido, variability in return rates was not related to size at release or to the coastal growth of juveniles, but with offshore growth in the Sea of Okhotsk, the nursery area for juveniles after leaving Japanese coastal waters. Whereas coastal growth tended to be negatively correlated with size at release in some stocks, offshore growth was positively associated with the August–November sea surface temperature in all stocks. This study confirmed that mortality of juvenile salmon occurred in two phases, during the coastal residency and the late period of the growing season, but the relative importance of both phases varied by stock and region, which probably regulated year‐class strength of Hokkaido chum salmon.     

Transport and survival of larvae of pelagic fishes in Kuroshio system region estimated with Lagrangian drifters

ABSTRACT:   Transport and survival of larvae of pelagic fishes in the Kuroshio system region were studied using Lagrangian drifter data recorded from 1990 to 2003. A large portion of the drifters from the Kuroshio area south of Japan spread around the Kuroshio Extension up to 170°E, while some moved south to the offshore area of the Kuroshio because of a recirculation gyre. The monthly mean eastward movement from areas south of Japan was approximately 800–900 km, which was smaller than previous numerical estimates. The results of a survival model assuming optimal temperatures for larvae suggest that surface waters during the observation period were too warm for larval Japanese sardine, which has an optimal temperature of 16°C, and the adult abundances did not increase during the observation period. In contrast, the spawning ground temperatures and transport conditions from an area south-west of Japan in April–June matched the requirement for the larval Japanese anchovy, which has an optimal temperature of 22°C. The combined effects of temperature variations due to seasonality and water mass mixing are suggested to play an important role in determining the environmental temperatures that occur in an area.     

Habitat shift of ayu Plecoglossus altivelis altivelis in early stages from waters adjacent to the bank to the center of flow in the Shimanto Estuary

ABSTRACT: To clarify the habitats of the ayu in the early stages of development in the Shimanto Estuary, the size and hatching date of the larval and juvenile ayu were examined. Sizes of larvae and juveniles were larger in the center of the flow area than in the waters adjacent to the estuary's banks, which suggests that larvae shift habitat to the center of flow from the waters along the estuary's banks. This shift began at approximately 20 mm body length. However, the resident term in the waters adjacent to the estuary's banks changed with the hatching dates; that is, the early and late-hatched larvae became short and long-term residents, respectively. A similar phenomenon was found also in the juveniles collected in the freshwater section that were migrating upstream. The growth rates of larval ayu in the waters adjacent to the banks tended to decrease with hatching dates. The fluctuations in growth rate with hatching date appear to be a factor leading to the variation of resident term.     

Survival and growth of sardine larvae in the offshore side of the Kuroshio

After 1984 the major spawning of the Japanese sardine, Sardinops melanostictus, has been observed to occur in the offshore waters, where the survival of early-stage larvae (4–10 mm in length) is questionable. The main objective of this study is to estimate the growth and survival rates of the early-stage larval cohort in the offshore side of the Kuroshio Current. A radar-reflecting buoy with a surface drogue was launched to tag a patch of larvae, and the patch was traced for 3 days in March 1991. The survival rate of the early-stage larval cohort was calculated from the change in density during the survey. The range of the instantaneous mortality rate was from 0.83 to 1.11 day, a survival rate of32-44%day-¹.The mean density of the smaller larvae (4–6 mm in length) decreased more rapidly than that of the larger larvae (6–10 mm in length) during the sampling period. The survival rate of the first-feeding larvae in the offshore region seemed to be lower than that of the post-flrst-feeding larvae. However, the growth rates of the first-feeding larvae's survivors in the region were higher than those of the post-first-feeding larvae's survivors. The first-feeding larvae in the offshore region seemed to survive when the growth rate of the larvae was high.     

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.     

Distribution of jack mackerel (Trachurus japonicus) larvae and juveniles in the East China Sea, with special reference to the larval transport by the Kuroshio Current

Horizontal distribution patterns of jack mackerel (Trachurus japonicus) larvae and juveniles were investigated in the East China Sea between 4 February and 30 April 2001. A total of 1549 larvae and juveniles were collected by bongo and neuston nets at 357 stations. The larvae were concentrated in the frontal area between the Kuroshio Current and shelf waters in the upstream region of the Kuroshio. The abundance of small larvae (<3 mm notochord length) was highest in the southern East China Sea (SECS) south of 28°N, suggesting that the principal spawning ground is formed in the SECS from late winter to spring. Jack mackerel also spawned in the northern and central East China Sea (NECS and CECS, respectively), as some small larvae were also collected in these areas. In the SECS, the abundance of small larvae was highest in February and gradually decreased from March to April. The habitat temperature of small larvae in the SECS and CECS (20–26°C) was higher than that in the NECS (15–21°C), suggesting higher growth rates in the SECS and CECS than in the NECS. The juveniles (10‐ to 30‐mm standard length) became abundant in the NECS off the west coast of Kyushu Island and CECS in April and were collected in association with scyphozoans typical of the Kuroshio waters. However, juveniles were rarely collected in the SECS, where the small larvae were concentrated. Considering the current systems in the study area, a large number of the eggs and larvae spawned and hatched in the SECS would be transported northeastward by the Kuroshio and its branches into the jack mackerels’ nursery grounds, such as the shallow waters off the west coast of Kyushu and the Pacific coast of southern Japan.     

Temperature determines growth rates of larval round herring Etrumeus teres in the Pacific coastal waters off southern Japan

Seasonal variation in daily growth rates in the early and middle larval stages of round herring Etrumeus teres were largely determined by the sea temperatures experienced by hatch-date cohorts in the Pacific coastal waters off southern Japan. Round herring larvae were collected by purse seining in the coastal waters of central Tosa Bay. A total of 451 larvae were aged by reading daily rings in otoliths. Individuals within a range of 2–5 hatch dates were grouped as hatch-date cohorts. We selected 16 cohorts that hatched during September 2000 and March 2002 and calculated mean widths of otolith growth increments for each cohort during the first feeding stage (W _FF, increments 1–5) and the maximum increment width in the middle larval stage (W _MAX). Seasonal variation in mean W _FF and W _MAX among the 16 cohorts was largely (80–90 %) explained by the sea temperature in the bay. These results indicate that temperature was a predominant determinant of larval growth rates; other environmental factors, such as food availability, did not substantially affect growth rates of round herring larvae in coastal waters along the subtropical Kuroshio Current off southern Japan.     

Carbon source and trophic position of pelagic fish in coastal waters of south-eastern Izu Peninsula,Japan, identified by stable isotope analysis

ABSTRACT:   It is important to clarify trophic dynamics in marine ecosystems for management of the fishing ground. Organic carbon sources and trophic position of pelagic fishes in the coastal waters of the south-eastern Izu Peninsula, Japan, were examined on the basis of carbon and nitrogen stable isotope distributions. The δ¹³C of the fishes was mostly distributed from −19 to −16‰ for nektonic fishes (13 species of adults and immatures) and planktonic fishes (10 species of larvae and juveniles), close to the δ¹³C values of particulate organic matter and planktonic decapods. These δ¹³C signatures for the inhabitants of the water column were in contrast with the high δ¹³C values (mainly −16 to −13‰) for demersal fishes of Scorpaeniformes and benthic polychaetes collected in the surf zone. These results indicate that nektonic and planktonic fishes depend on phytoplankton for carbon supply. The δ¹⁵N signatures suggest that the trophic position ranged 3.1–4.5 for the nektonic fishes and 2.9–3.7 for the planktonic fishes, premised on trophic level 3 for larval Japanese anchovy Engraulis japonicus . Thus, planktivorous fishes should be mainly assigned to trophic levels 3 and 4 in this area.     

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.     

Simulation of the impact of climate change on the egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island,the western coast of Japan

In order to investigate the impact of climate change on egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island western Japan, we conducted particle‐tracking simulations on transport success/failure to fishing grounds from 1960 to 2007. The modeled transport success since the mid‐1990s increased and decreased in the offshore and coastal zones, respectively, compared with the 1960s and 1970s. The estimated northward shift of the spawning ground and weakened Tsushima Warm Current contributed to increase in modeled transport success to the offshore zone. Conversely, the weakening trend of the modeled onshore current in the Goto‐Nada Sea combined with the northward shift of the spawning ground resulted in unsuccessful larval transport. These results suggest that fluctuations in juvenile and subadult anchovy catches in this area may be attributable to changes in the physical environment. The present study showed that changes in transport success induced by oceanographic fluctuations related to climate change, have the potential to affect anchovy recruitment off the western coast of Japan.     

Late winter larval mesopelagic fish assemblage in the Kuroshio waters of the western North Pacific

The larval mesopelagic fish assemblage and its distribution patterns were investigated in the Kuroshio region off southern Japan in late winter. A total of 8690 fish larvae was collected, 85.8% of which were mesopelagic fish larvae. Mesopelagic fish larvae were significantly more abundant in the area east of the Kuroshio axis than west of the Kuroshio axis (660.6 versus 194.5 ind 10 m^?2). Sigmops gracile, Bathylagus ochotensis, Notoscopelus japonicus, Diaphus slender type and Myctophum asperum were the five most abundant larvae and accounted for 16.9, 16.4, 15.2, 13.9 and 9.3% of the total catch in numbers, respectively. We conclude that these larvae were transported by the Kuroshio Current to the more productive transition waters, where they spend their juvenile stage from spring to early summer. The possibility of expatriation and southward long‐distance spawning migrations of N. japonicus and B. ochotensis are discussed, based on the geographic distribution patterns of their larvae, juveniles and adults.     

基于鱼卵仔鱼数据的东海中南部日本鲭产卵场分析

利用2016年4—5月在东海中南部海域的调查数据,对该海域日本鲭(Scomber japonicus)鱼卵仔鱼的分布特征进行了研究,并对产卵场的划分展开讨论及提出保护措施和管理建议。结果显示:4月,日本鲭鱼卵仔鱼主要出现在台湾海峡和东海南部海域,出现率均大于25%,但仔鱼在东海南部外海的出现率和丰度均较低,分别为5.00%和0.01个·100m-3。5月,日本鲭鱼卵仔鱼在台湾海峡的出现率和丰度均显著降低,分别下降至5.88%和0.22个·100m^-3;东海南部外海日本鲭鱼卵仔鱼的出现率和丰度则均显著升高,分别升至50%和36.48个·100m^-3,其中鱼卵集中出现在南部外海,出现率和丰度分别达到了40%和36.46个·100m^-3;东海中部外海从5月开始出现日本鲭鱼卵和仔鱼。结合历史资料,可以判断东海中南部日本鲭产卵场主要有台湾海峡、东海中南部近海、东海南部外海、东海中部外海等4个。日本鲭鱼卵丰度与表层水温呈显著相关关系(R=0.37,P<0.05),与表层盐度无显著相关关系(P>0.05);仔鱼丰度与表层水温及表层盐度均无显著相关关系(P>0.05)。结果表明,水温对日本鲭的产卵活动影响较大,随着水温的逐步升高,性腺发育速度加快,开始产卵的亲体数量也大量增加,这从鱼卵仔鱼的数量大量增加得以体现;而盐度变化对日本鲭的产卵活动影响相对较小。针对日本鲭不同的产卵场,建议制定差异化的繁殖亲体和幼鱼保护措施。     

Morphological and habitat characteristics of settling and newly settled Roughscale Sole Clidoderma asperrimum collected in the coastal waters of northeastern Japan

This report first describes the morphological and habitat characteristics of larval and juvenile Roughscale Sole Clidoderma asperrimum. Five metamorphosing larvae (22.1–31.7 mm standard length, SL) and 12 juveniles (23.7–46.4 mm SL) were collected from two locations in coastal waters of northeastern Japan, Shizugawa Bay (rocky habitats, 6–12 m depth) and Iwaki (sandy habitats, 10–50 m depth), respectively, using a hand net while SCUBA diving and a small trawl net during May–August in 2008 and 2010. Underwater observation in Shizugawa Bay revealed that metamorphosing larvae and juveniles that had settled on rocks or stones showed transparent and variegated body coloration, which is cryptic to a rocky habitat. Juveniles in Iwaki showed monotonic coloration. The settlement would occur during metamorphosis because the same larva observed on a stone subsequently swam with a left-lateralized body posture. The larval morphology was characterized by a dense covering of fine spines on the whole body surface. Spines on the ocular side developed steadily, gradually forming bony tubercles with growth, whereas those on the blind side regressed with development and finally disappeared in metamorphosed juveniles. Our results indicate that Roughscale Sole can settle and adapt to rocky and sandy habitats in coastal waters with wide depth range.     

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.     

Growth and movement patterns of early juvenile European anchovy (Engraulis encrasicolus L.) in the Bay of Biscay based on otolith microstructure and chemistry

Various hypotheses have been put forward to explain the mechanisms in the Bay of Biscay that result in a good recruitment of European anchovy. Anchovy larvae from the spawning area in the Gironde River plume are advected towards off-shelf waters, where juveniles are commonly observed. Otolith microstructural and chemical analysis were combined to assess the importance of this off-shelf transport and to determine the relative contribution of these areas for anchovy survival. Chemical analysis of otoliths showed that anchovy juveniles in the Bay of Biscay can be divided into two groups: a group that drifts towards off-shelf waters early in their life and returns later, and a group that remains in the low salinity waters of the coastal area. The first group presents significantly faster growth rates (0.88 mm day⁻¹) than those remaining in the coastal waters (0.32 mm day⁻¹). This may be due to off-shelf waters being warmer in spring/summer, and to the fact that the lower food concentration is compensated for by higher prey visibility. Furthermore, the group of juveniles that drifted off the spawning area and had faster growth rates represents 99% of the juvenile population. These findings support the hypothesis that anchovy in the Bay of Biscay may use off-shelf waters as a spatio-temporal loophole, suggesting that transport off the shelf may be favourable for recruitment.     

Interannual variability in growth of larval and juvenile walleye pollock Theragra chalcogramma in the western Gulf of Alaska, 1983–91

Interannual variability in growth of larval walleye pollock Theragra chalcogramma was examined from 1983 to 1991 and of juveniles from 1985 to 1990. ANCOVA was used to assess differences in population growth rates, and an alternate method was developed to examine variations between annual length-at-age data and average 'expected' values over different age groupings. For larvae, the years 1986, 1987, 1989 and 1990 had higher than average length-at-age, and 1988 and 1991 had lower than average values. Relationships between growth and SST and larval density were not clear. A tentative relationship between copepod nauplii abundance and larval length-at-age was noted. The consequence of larval growth to larval mortality, late larval abundance or recruitment was not clear. We conclude that larval mortality rates are highly variable and tend to mask effects of moderate variability in growth on later abundance. For juveniles, 1987 had significantly lower than average length-at-age and 1988 had higher than average values. Although there are few years of data, they tend to support the importance of juvenile growth in the recruitment process. Conditions for the large 1988 year class are documented and discussed, including warm SST, calm winds, relatively low larval growth rates, low abundances of potential predators on larvae, low larval mortality rates, and high juvenile growth rates.     

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.     

Skeletal development and deformities in cultured larval and juvenile seven-band grouper, Epinephelus septemfasciatus (Thunberg)

The seven‐band grouper, Epinephelus septemfasciatus (Thunberg), is currently recognized as a potential new species for aquaculture in Japan. This study describes normal and abnormal skeletal development of the jaw and vertebrae in cultured larvae and early juveniles of E. septemfasciatus. The ontogenetic stages at which skeletal deformities of jaw and vertebra developed were also described for this species. Osteological observations were made using a clearing and staining method for larvae and soft X‐ray photographs for juveniles. A high incidence of skeletal deformities was observed in the jaws and vertebral column during the larval and juvenile stages. Most of the jaw deformities were explained by an abnormal maxilla curvature. Jaw deformities were visually evident from flexion stage after ossification of the deformed elements. Deformities in the vertebral column (mostly lordosis) were observed from the post‐larval stage and became more evident as growth proceeded. The lordosis generally occurred on the positions of the 8–11th vertebra. These types of deformities are compared with those of other species, and possible causative factors of the skeletal deformities are discussed.