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.     

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.     

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.     

Growth variability of Pacific saury Cololabis saira larvae under contrasting environments across the Kuroshio axis: survival potential of minority versus majority

Growth variability was examined for Pacific saury Cololabis saira larvae under contrasting environments across the Kuroshio axis, based on samples collected during the winter spawning season in 2013 and 2014. The growth rate index (residual of the otolith marginal 3‐day mean increment width from the linear regression on knob length) of larvae was compared among three areas: the inshore side of the Kuroshio axis, the Kuroshio axis, and the offshore side of the Kuroshio axis in relation to sea surface temperature (SST), salinity (SSS) and chlorophyll‐a (CHL) concentration. The larvae were more densely distributed in the Kuroshio axis and offshore areas of higher temperature and salinity and lower chlorophyll‐a concentration than in the inshore areas of lower temperature and salinity and higher chlorophyll‐a concentration. No marked differences in the growth rate index were found among the three areas, even though the larvae in the inshore areas showed slightly higher growth rates in 2013. Despite the broad ranges of environmental factors, no clear relationship between the growth rate index and any environmental factor was detected. The survival potential of Pacific saury larvae was considered to be at least comparable under contrasting environments across the Kuroshio axis. Such a geographical homogeneity is concluded to be attributable to compensable effects of physical and biological factors. We hypothesize that the minority under physically‐unfavorable but biologically‐favorable conditions on the inshore side of the Kuroshio axis could survive equally well as the majority under physically‐favorable but biologically‐unfavorable conditions around the Kuroshio axis and on the offshore side of the Kuroshio axis.     

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.     

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.     

Variability in growth rates of larval haddock in the northern North Sea

The large-scale distribution of haddock ( Melanogrammus aeglefinus ) larvae in the northern North Sea was mapped in a grid survey carried out in late April 1996. A drifting buoy was deployed in the centre of one of the areas of concentration of larvae located off the east coast of the Shetland Isles, where intensive sampling was carried out for ≈ 10 days. Daily larval haddock growth variability, estimated from otolith microstructure analysis, was independent of the measured variability of the physical and biological environment of the larvae. The survey coincided with the onset of the spring plankton production bloom, and a likely explanation for the absence of environmental effects on larval growth was high food availability and larval feeding rates. Nevertheless, differences in growth were observed between cohorts, with larvae hatched later in the spring displaying higher growth at age than those hatched earlier. Particle-tracking modelling suggested that differences in temperature history between cohorts, on their own or compounded by a potential interaction between temperature and the development of plankton production, may explain the higher growth rate of the larvae hatched later in the season.     

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.     

Growth and mortality of larval anchoveta Engraulis ringens,in northern Chile during winter and their relationship with coastal hydrographic conditions

Larval growth, age, growth effect and instantaneous mortality were estimated in anchoveta, Engraulis ringens, collected biweekly during the austral winter of 2014 in nearshore waters off Bay of Antofagasta (23°41′W–70°30′S), northern Chile. Through measuring standard length (SL) and sagitta microstructure analysis, it was estimated that the growth rate of E. ringens larvae decreased from June (0.85 mm day^?1) to August (0.50 mm day^?1). However, the water temperature was homogeneous during the sampling dates (14.6, 15.2, 14.4, and 14.6°C), suggesting that the decelerating larval growth was not linked to changes in sea temperature. Additionally, larvae with slow growth have larger otoliths compared with conspecifics with fast growth (growth effect). Larval mortality rates tended to decrease until the middle of July (0.18 per day) but increased to 0.25 per day in early August, which coincided with lower food availability (i.e., chlorophyll‐a, 2.7–5.6 mg m^?3) and a high occurrence of smaller larvae (1.58–11.5 mm). Partial least squares analysis indicates low covariance between the biological and oceanographic variables (PLS: 11.71%), suggesting that other factors, such as parental effects, may explain the abrupt decrease in the larval growth rates.     

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.     

Seasonal potential fishing ground prediction of neon flying squid (Ommastrephes bartramii) in the western and central North Pacific

We explored the seasonal potential fishing grounds of neon flying squid (Ommastrephes bartramii) in the western and central North Pacific using maximum entropy (MaxEnt) models fitted with squid fishery data as response and environmental factors from remotely sensed [sea surface temperature (SST), sea surface height (SSH), eddy kinetic energy (EKE), wind stress curl (WSC) and numerical model‐derived sea surface salinity (SSS)] covariates. The potential squid fishing grounds from January–February (winter) and June–July (summer) 2001–2004 were simulated separately and covered the near‐coast (winter) and offshore (summer) forage areas off the Kuroshio–Oyashio transition and subarctic frontal zones. The oceanographic conditions differed between regions and were regulated by the inherent seasonal variability and prevailing basin dynamics. The seasonal and spatial extents of potential squid fishing grounds were largely explained by SST (7–17°C in the winter and 11–18°C in the summer) and SSS (33.8–34.8 in the winter and 33.7–34.3 in the summer). These ocean properties are water mass tracers and define the boundaries of the North Pacific hydrographic provinces. Mesoscale variability in the upper ocean inferred from SSH and EKE were also influential to squid potential fishing grounds and are presumably linked to the augmented primary productivity from nutrient enhancement and entrainment of passive plankton. WSC, however, has the least model contribution to squid potential fishing habitat relative to the other environmental factors examined. Findings of this work underpin the importance of SST and SSS as robust predictors of the seasonal squid potential fishing grounds in the western and central North Pacific and highlight MaxEnt's potential for operational fishery application.     

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.     

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.     

Daily growth rate model of Japanese anchovy larvae Engraulis japonicus in Hiuchi-nada Sea, central Seto Inland Sea

In the present study, we developed a larval anchovy growth model in relation to sea temperature and food availability via food consumption and metabolic process terms, based on biological data from previous laboratory experiments and field surveys from 2003 to 2006 in Hiuchi-nada Sea, central part of Seto Inland Sea, Japan. To investigate when food shortage for larval anchovy and then recruitment failure occur in Hiuchi-nada Sea, anchovy food requirements were estimated by using the growth model, and we compared the food requirement with anchovy food availability. We applied an estimation method for growth model parameters, Hewett?CJohnson p and Q ₁₀, by minimizing the sum of squares of difference between mass-specific growth rates estimated by the models and those by otolith growth analysis. Parameter p was 0.86, slightly higher than typical values, and Q ₁₀ was 2.11, close to the value used for the biological model of larval northern anchovy. Food shortage for anchovy larvae did not occur in Hiuchi-nada Sea, although it was indicated that low food availability led to a low reproductive success rate. The newly developed growth model is considered optimal at present and useful to link environmental conditions and larval growth.     

Increasing the stocking density in Paracentrotus lividus larviculture: Effects on survival and metamorphosis rates

The development of sustainable methods for sea urchin juvenile production is currently constrained by high mortality rates during larval growth and the high costs of larval rearing systems management. With the aim of developing a method for the production of juveniles of the purple sea urchin Paracentrotus lividus in a medium‐scale recirculating system, the present study focused on the effects of high stocking densities on larval growth. Plutei larvae were reared at three different densities (up to 7 ind/ml) in a semi‐static culture system. Larval survival and metamorphosis success were evaluated in order to identify the most effective density range. The highest metamorphosis rates (80%–95%) were obtained at 4 and 7 larvae/ml. These results are comparable with (and in some cases higher than) those reported for the same species at much lower larval densities. In conclusion, the rearing conditions tested here show for the first time that a significantly higher (4 ind/ml) stocking density than those of traditional P. lividus rearing methods (based on large volumes and low densities) can be adopted, thus supporting the feasibility of an increase in the final output of competent larvae with no increase in rearing volumes.     

Long‐term trends and drivers of larval phenology and abundance of dominant brachyuran crabs in the Gulf of St. Lawrence (Canada)

Climate change has led to major shifts in the timing of biological events, with many studies demonstrating earlier phenology in response to warming. However, few of these studies have investigated the effects of climate change on the phenology of larvae in marine species. Phenological shifts can result in mismatches between consumers and prey and hence affect growth and survival of individuals, and ultimately population demography. We investigated the temporal changes in phenology and abundance of the larvae of dominant brachyuran crabs in the southern Gulf of St. Lawrence (eastern Canada) based on plankton collections spanning 1982–2012. The Gulf of St. Lawrence has warmed since the early 1990s, and our analyses revealed that larvae of snow crab (Chionoecetes opilio) and toad crabs (Hyas spp.) exhibited a significant trend towards earlier phenology over the 30‐year study period. This shift in phenology appeared to be a consequence of the effect of climate warming on both the timing of hatching and larval development rate. Larval abundance responded differently by crab taxon to climate warming, likely due to differences in thermal tolerance. The warming trend was unfavourable to snow crab, which is the most cold‐adapted and stenothermic of the taxa examined in this study. The abundance of snow crab larvae was lower when sea ice retreat occurred earlier than day 110 of the year and sea surface temperature was higher than 8.5°C. On the other hand, larval abundance of rock crab (Cancer irroratus), which prefers higher temperatures, was positively related to surface temperature.     

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.     

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.     

Pelagic larval traits of the amphidromous goby Sicyopterus lagocephalus display seasonal variations related to temperature in La Réunion Island

Amphidromous gobies represent a substantial part of freshwater fish diversity throughout islands of the Indo‐Pacific region. They display a marine pelagic phase during several months before recruiting in rivers. Understanding the relationship between larval traits and environmental conditions is a major challenge for the evaluation of a spatial scale of connectivity and populations’ dynamics, especially in a climate change context. In this study, the larval traits of Sicyopterus lagocephalus were examined and related to the sea surface temperature (SST), over three consecutive years in La Réunion Island (Mascarene archipelago). The pelagic larval duration (PLD, range from 96 to 293 days), the size‐at‐recruitment (range from 26.5 to 37 mm TL) and the larval growth rate (range from 0.112 to 0.293 mm·day^?1 TL) varied seasonally depending on hatching date. The larval growth rate was inversely correlated to the fluctuations of PLD and size. Larvae living in high sea water temperatures exhibited a faster growth, shorter PLDs and smaller sizes‐at‐recruitment than those living in cool water temperatures. The instantaneous daily growth was assessed by the study of otolith increment widths. The daily growth was not linear throughout larval life and was positively correlated to SST. We showed high amplitude of PLD periodic fluctuations (170.39 ± 43.75 days) related to temperature (12.8% PLD drop per degree), which could affect dispersal and induce intermittent connectivity between distant populations. This high plasticity of larval traits is likely to be advantageous to respond to the wide range of environmental conditions encountered throughout the species distribution range.     

Relationship between daily survival rates of larval Japanese anchovy (Engraulis japonicus) and concentrations of copepod nauplii in the Seto Inland Sea, Japan

We attempted to estimate the survival rates of larval cohorts (3–15 mm in body length) of the Japanese anchovy Engraulis japonicus population in Hiuchi‐nada, Seto Inland Sea, Japan, and examined the relationship to their food availability. The survival rates were directly calculated from the change in larval density during the survey. The estimated daily survival rate ranged between 0 and 89% d⁻¹, but increased with the increase in the mean concentration of small‐sized copepod nauplii (<100 μm in body length) sampled at 10 m depth. When the food concentration was higher than about 5 nauplii L⁻¹, the daily survival rate of larval anchovy reached an asymptote of approximately 89% day⁻¹. It might be possible to provide a framework for the forecast for larval abundance immediately prior to recruitment size (10–15 mm SL), based on larval abundance of the monitored size (3–8 mm SL) and the survival rate which is estimated from the concentration of small‐sized nauplii.