Density dependence in total egg production per spawner for marine fish

A paradigm of fisheries science holds that spawning stock biomass (SSB) is directly proportional to total egg production (TEP) of fish stocks. This “SSB–TEP proportionality” paradigm has been a basic premise underlying the spawner–recruitment models for fisheries management and numerous studies on recruitment mechanisms of fish. Studies on maternal effects on reproductive potential of a stock have progressed during the last few decades, leading to doubt concerning the paradigm. Nonetheless, a direct test of the paradigm at multidecadal scales has been difficult because of data limitations in the stock assessment systems worldwide. Here, we tested the paradigm for marine fish based on a novel combination of two independent 38‐year time series: fishery‐dependent stock assessment data and fishery‐independent egg survey data. Through this approach, we show that the SSB–TEP proportionality is distorted by density dependence in total egg production per spawner individual (TEPPS) or spawner unit weight (TEPPSW) at a multidecadal scale. The TEPPS/TEPPSW exponentially declined with biomass and thus was density‐dependent for Japanese sardine, a small pelagic species exhibiting a high level of population fluctuation, in the western North Pacific. By contrast, the TEPPS/TEPPSW was sardine‐density‐dependent for Japanese anchovy, another small pelagic species exhibiting a moderate level of population fluctuation well‐known for being out of phase with sardine. Our analysis revealed intraspecific (sardine) and interspecific (anchovy) density dependence in TEPPS/TEPPSW, which was previously unaccounted for in spawner–recruitment relationships. Such density‐dependent effects at the time of spawning should be considered in fisheries management and studies on recruitment mechanisms.     

Disentangling density‐dependent effects on egg production and survival from egg to recruitment in fish

Understanding of density‐dependent effects is key to achieving sustainable management of self‐regulating biological resources such as fish stocks. Traditionally, density‐dependent effects on population abundance in fish have been considered to occur from hatching to recruitment, based on the paradigm of proportionality between spawning stock biomass and total egg production. Here, we demonstrate how the existence of intraspecific and interspecific density dependence in egg production changes the current understanding of density‐dependent processes in the life history of fish, by disentangling density‐dependent effects on egg production and survival from egg to recruitment, using sardine (Sardinops melanostictus, Clupeidae) and anchovy (Engraulis japonicus, Engraulidae) as model species. For sardine, strong intraspecific density‐dependent effects occurred in egg production, but no density‐dependent effects occurred or if any they were weak enough to be masked by environmental factors from hatching to recruitment. In contrast, for anchovy, interspecific density‐dependent effects occurred in egg production. In the survival after hatching, anchovy experienced stronger intraspecific density‐dependent effects than currently recognized. This analysis could overturn the current understanding of density‐dependent effects in the life history, highlighting contrasts between the effects on individual quality and population abundance and between the model species. We propose to reconsider the basis of fisheries management and recruitment studies based on the revised understanding of density‐dependent effects in the life history of the respective species.     

When in life does density dependence occur in fish populations?

Fisheries advice is based on demographic calculations, which assume that density‐dependent processes regulating recruitment occur only in early life. This assumption is challenged by laboratory and lake studies and some recent indications from marine systems that demonstrate density‐dependent regulation late in life. By accounting for spatial dynamics of a population, something that has previously been ignored in models of fish, we show that density‐dependent regulation is determined by the size of the habitat: in small habitats, for example small lakes, regulation occurs late in life, while it can occur early in large habitats. When regulation happens late in life, fisheries yield is maximized by exploitation of mainly juvenile fish, while exploiting mature fish maximizes yield if regulation happens early. We review and interpret observations of density dependence in the light of the theory. Our results challenge the current assumption that density dependence always occurs early in life and highlights the need for an increased understanding of density‐dependent processes. This can only come about by a change of focus from determining stock‐recruitment relationships towards understanding when and how density‐dependent regulation occurs in nature.     

Density‐dependent egg production in chub mackerel in the Kuroshio Current system

A paradigm of proportionality between spawning stock biomass (SSB) and total egg production (TEP) has been largely untested at multidecadal scales mainly because of difficulty in estimating annual TEP. Recently, this paradigm was directly tested for sardine (Sardinops melanostictus) and anchovy (Engraulis japonicus) at a multidecadal scale to reveal that SSB–TEP proportionality was partially distorted by intraspecific (sardine) and interspecific (anchovy) density dependence in total egg production per spawner individual (TEPPS) or unit weight (TEPPSW). In the present study, we demonstrate intraspecific density dependence in TEPPS/TEPPSW for chub mackerel (Scomber japonicus) in the Kuroshio Current system, using a proxy for TEPPS/TEPPSW, calculated from snapshot abundance data based on fishery‐independent egg surveys in combination with fishery‐dependent stock assessment data, at a multidecadal scale (38 years). TEPPS/TEPPSW exponentially declined with SSB, indicating a strong intraspecific density dependence in TEPPS/TEPPSW in chub mackerel. The observed phenomenon for chub mackerel was similar to that for sardine. Hence, intraspecific density dependence in TEPPS/TEPPSW may be a phenomenon that is generally applicable for species with a high maximum biomass and large population fluctuations. Lastly, we recommend the application of a TEP‐based framework to studies on recruitment mechanisms of fish.     

Density‐ and size‐dependent mortality in fish early life stages

The importance of survival and growth variations early in life for population dynamics depends on the degrees of compensatory density dependence and size dependence in survival at later life stages. Quantifying density‐ and size‐dependent mortality at different juvenile stages is therefore important to understand and potentially predict the recruitment to the population. We applied a statistical state‐space modelling approach to analyse time series of abundance and mean body size of larval and juvenile fish. The focus was to identify the importance of abundance and body size for growth and survival through successive larval and juvenile age intervals, and to quantify how the dynamics propagate through the early life to influence recruitment. We thus identified both relevant ages and mechanisms (i.e. density dependence and size dependence in survival and growth) linking recruitment variability to early life dynamics. The analysis was conducted on six economically and ecologically important fish populations from cold temperate and sub‐arctic marine ecosystems. Our results underscore the importance of size for survival early in life. The comparative analysis suggests that size‐dependent mortality and density‐dependent growth frequently occur at a transition from pelagic to demersal habitats, which may be linked to competition for suitable habitat. The generality of this hypothesis warrants testing in future research.     

Environmental determinants of growth rates for larval Japanese anchovy Engraulis japonicus in different waters

Do disparate mechanisms determine growth rates of fish larvae in the different regions? The relationship between growth rates and environmental factors (sea temperature and food availability) was examined for larval Japanese anchovy Engraulis japonicus in geographically and environmentally different waters, through sagittal otolith microstructure analysis. Recent 3‐day mean growth rates directly before capture were positively related with sea‐surface temperature (SST) but not with food availability (plankton density) for the larvae in the Kuroshio Extension and Kuroshio–Oyashio transition regions of the western North Pacific. On the contrary, variations in recent growth rates were attributed to food availability (plankton density) as well as SST for the larvae in the East China Sea. In the shirasu fishing ground in Sagami Bay, larval growth rates were variable under the influences of both SST and food availability (feeding incidence). On the surface, the growth–environment relationships seemed to differ among regions. However, a definite general pattern of the dome‐shaped relationship between recent growth rates and SST was observed when all the regions were combined. Growth rates were similar even among clearly different regions if at the same SST. Overall, growth rates roughly increased with SST until they reached the maximum at SST of 21–22°C (i.e. optimal growth temperature), and declined when SST went over 21–22°C. On the contrary, no clear relationship was observed between growth rate and plankton density or between SST and plankton density. Therefore, the apparent among‐region differences would be firstly caused by the differences in regional SST range. The systematic mechanism of growth determination for widespread pelagic fish species larvae would be run by primarily sea temperature and secondarily food availability, at the species level.     

Saving large fish through harvest slots outperforms the classical minimum‐length limit when the aim is to achieve multiple harvest and catch‐related fisheries objectives

We address the problem of optimal size‐selective exploitation in an age‐structured fish population model by systematically examining how density and size dependency in growth, mortality and fecundity affect optimal harvesting patterns when judged against a set of fisheries objectives. The study offers five key insights. First, while minimum‐length limits often maximize the biomass yield, exploitation using harvest slots (i.e. regulations that protect both immature and very large individuals) can generate within 95% of maximum yield; harvest slots also generally maximize the number of fish that are harvested. Second, density dependence in growth and size‐dependent mortality predict more liberal optimal size limits than those derived under assumptions of no density and size dependence. Third, strong density dependence in growth maximizes the catch of trophy fish only when modest harvest is introduced; the same holds for numbers harvested, when the stock–recruitment function follows the Ricker type. Fourth, the inclusion of size‐dependent maternal effects on fecundity or egg viability has only limited effects on optimal size limits, unless the increase in fecundity with mass (“hyperallometry”) is very large. However, large hyperallometry in fecundity shifts the optimal size limit for biomass yield from the traditional minimum‐length limit to a harvest slot. Fifth, harvest slots generally provide the best compromises among multiple objectives. We conclude that harvest slots, or more generally dome‐shaped selectivity to harvest, can outperform the standard minimum‐length selectivity. The exact configuration of optimal size limits crucially depends on objectives, local fishing pressure, the stock–recruitment function, and the density and size dependency of growth, mortality and fecundity.     

Recruitment of common sardine (Strangomera bentincki) and anchovy (Engraulis ringens) off central-south Chile in the 1990s and the impact of the 1997–1998 El Niño

The recruitment of Strangomera bentincki (common sardine) and Engraulis ringens (anchovy) and the relationships with oceanographic conditions in the upwelling ecosystem of central-south Chile were investigated from 1990 to 1998, with emphasis on the 1997–1998 El Niño. Time series of recruitment, biomass, local sea surface temperature, and a coastal upwelling index were used to explore relationships during the spawning (July–August) and pre-recruitment (August–December) periods. The 1997–1998 El Niño caused physical changes in the small pelagic fish habitat off central-south Chile. Anomalous sea surface temperatures (SST) and upwelling indexes began to be detected from May 1997 and persisted into 1998. Recruitment of common sardine showed significantly negative relationship with SST anomalies during the pre-recruitment period, as well as with the upwelling index during the peak of spawning. However, the recruitment of anchovy did not seem to be affected by the environmental changes observed in the 1990s. Instead, the recruitment rate of anchovy showed negative relationship with the recruitment rate of common sardine. We conclude that the conditions of the 1997–1998 El Niño off central-south Chile affected the survival of common sardine offspring, and that the recruitment success of anchovy could be determined by less-abundant cohorts of common sardine through a biological mechanism of interaction.     

Impact of freshwater input and wind on landings of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in shelf waters surrounding the Ebre (Ebro) River delta (north-western Mediterranean)

Time series analyses (Box–Jenkins models) were used to study the influence of river runoff and wind mixing index on the productivity of the two most abundant species of small pelagic fish exploited in waters surrounding the Ebre (Ebro) River continental shelf (north‐western Mediterranean): anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus). River flow and wind were selected because they are known to enhance fertilization and local planktonic production, thus being crucial for the survival of fish larvae. Time series of the two environmental variables and landings of the two species were analysed to extract the trend and seasonality. All series displayed important seasonal and interannual fluctuations. In the long term, landings of anchovy declined while those of sardine increased. At the seasonal scale, landings of anchovy peaked during spring/summer while those of sardine peaked during spring and autumn. Seasonality in landings of anchovy was stronger than in sardine. Concerning the environmental series, monthly average Ebre runoff showed a progressive decline from 1960 until the late 1980s, and the wind mixing index was highest during 1994–96. Within the annual cycle, the minimum river flow occurs from July to October and the wind mixing peaks in winter (December–April, excluding January). The results of the analyses showed a significant correlation between monthly landings of anchovy and freshwater input of the Ebre River during the spawning season of this species (April–August), with a time lag of 12 months. In contrast, monthly landings of sardine were significantly positively correlated with the wind mixing index during the spawning season of this species (November–March), with a lag of 18 months. The results provide evidence of the influence of riverine inputs and wind mixing on the productivity of small pelagic fish in the north‐western Mediterranean. The time lags obtained in the relationships stress the importance of river runoff and wind mixing for the early stages of anchovy and sardine, respectively, and their impact on recruitment.     

Do albacore exert top‐down pressure on northern anchovy? Estimating anchovy mortality as a result of predation by juvenile north pacific albacore in the California current system

Quantifying the mortality of marine fishes is important for understanding spawner–recruit relationships, predicting year‐class strength, and improving fishery stock assessment models. There is increasing evidence that pelagic predators can exert a top‐down influence on prey, especially during critical early life‐history stages. The objective of this study was to quantify predation by North Pacific albacore on Northern anchovy in the California current system (CCS). I estimated the abundance of juvenile albacore in the CCS from 1966–2005 using stock assessment models and spatially explicit catch‐per‐unit‐effort time series. Anchovy abundance (1966–93), both recruitment and total biomass, was obtained from a stock assessment model. Annual rates of anchovy consumption by albacore were calculated using diet studies of albacore in the CCS, an age‐structured bioenergetics model, and regional estimates of albacore abundance. The range of estimates was large: albacore may remove from less than 1% to over 17% of anchovy pre‐recruitment biomass annually. Relationships between predation and recruitment biomass were consistent with expectations from top‐down effects, but further study is required. This is the first attempt to quantify a specific source of mortality on anchovy recruits and to demonstrate potential top‐down effects of predation on anchovy.     

Temperature-dependent stock-recruitment model for walleye pollock (Theragra chalcogramma) around northern Japan

Changes in fish year‐class strength have been attributed to year‐to‐year variability in environmental conditions and spawning stock biomass (SSB). In particular, sea temperature has been shown to be linked to fish recruitment. In the present study, I examined the relationship between sea surface temperature (SST), SSB and recruitment for two stocks of walleye pollock (Theragra chalcogramma) around northern Japan [Japanese Pacific stock (JPS) and northern Japan Sea stock (JSS)] using a temperature‐dependent stock‐recruitment model (TDSRM). The recruitment fluctuation of JPS was successfully reproduced by the TDSRM with February and April SSTs, and February SST was a better environmental predictor than April SST. In addition, the JPS recruitment was positively related to February SST and negatively to April SST. The JSS recruitment modeled by the TDSRM incorporating February SST was also consistent with the observation, whereas the relationship between recruitment and February SST was negative, that is the opposite trend to JPS. These findings suggest that SST in February is important as a predictor of recruitment for both stocks, and that higher and lower SSTs in February act favorably on the recruitment of JPS and JSS respectively. Furthermore, Ricker‐type TDSRM was not selected for either of the stocks, suggesting that the strong density‐dependent effect as in the Ricker model does not exist for JPS and JSS. I formulate hypotheses to explain the links between SST and recruitment, and note that these relationships should be considered in any future attempts to understand the recruitment dynamics of JPS and JSS.     

Modelling potential spawning habitat of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus) in the Bay of Biscay

Large amplitude variations in recruitment of small pelagic fish result from interactions between a fluctuating environment and population dynamics processes such as spawning. The spatial extent and location of spawning, which is critical to the fate of eggs and larvae, can vary strongly from year to year, as a result of changing population structure and environmental conditions. Spawning habitat can be divided into ‘potential spawning habitat’, defined as habitat where the hydrographic conditions are suitable for spawning, ‘realized spawning habitat’, defined as habitat where spawning actually occurs, and ‘successful spawning habitat’, defined as habitat from where successful recruitment has resulted. Using biological data collected during the period 2000–2004, as well as hydrographic data, we investigate the role of environmental parameters in controlling the potential spawning habitat of anchovy and sardine in the Bay of Biscay. Anchovy potential spawning habitat appears to be primarily related to bottom temperature followed by surface temperature and mixed‐layer depth, whilst surface and bottom salinity appear to play a lesser role. The possible influence of hydrographic factors on the spawning habitat of sardine seems less clear than for anchovy. Modelled relationships between anchovy and sardine spawning are used to predict potential spawning habitat from hydrodynamical simulations. The results show that the seasonal patterns in spawning are well reproduced by the model, indicating that hydrographic changes may explain a large fraction of spawning spatial dynamics. Such models may prove useful in the context of forecasting potential impacts of future environmental changes on sardine and anchovy reproductive strategy in the north‐east Atlantic.     

Predation dynamics of mackerel on larval and juvenile anchovy: is capture success linked to prey condition?

We tested whether the predation dynamics of chub mackerel Scomber japonicus and spotted mackerel S. australasicus on young anchovy Engraulis japonicus relates to individual growth characteristics of the prey and could account for the growth-selective survival predicted by recruitment hypotheses. Juvenile and adult mackerel were sampled along with their young anchovy prey field in 2004 (juvenile mackerel and larval anchovy) and 2005 (adult mackerel and juvenile anchovy) off the Pacific coast of Honshu, Japan. The recent 5-day mean growth rate of larval and juvenile survivors and prey found in the stomach of mackerel was estimated from the otolith microstructure. No significant difference was found between the recent growth of larval or juvenile survivors and that of preyed individuals. We conclude that despite a relatively small body size, the high activity level and predation skills displayed by mackerel prevent fast-growing larvae and early juveniles from benefitting in terms of the expected survival advantage over slow-growers. Hence, growth-selective predation mortality of larval fish would depend on the feeding ecology of the predator rather than predator size. Selection for fast growth is more likely to occur under predation pressure from invertebrate organisms and small pelagic fish specialized on zooplankton, such as herring and anchovy.     

Stock–environment–recruitment models for North Atlantic albacore (Thunnus alalunga)

Different stock–recruitment models were fitted to North Atlantic albacore (Thunnus alalunga) recruitment and spawning stock biomass data. A classical density dependence hypothesis, a recent environmental‐dependence hypothesis and a combination of both were considered. For the latter case, four stock–environment–recruitment models were used: Ricker, Beverton‐Holt, Deriso's General Model (modified to take into account environmental effects) and conditioned Neural Networks. Cross‐validation analysis showed that the modified Deriso model had the best predictive capability. It detected an inverse effect of the North Atlantic Oscillation (NAO) on recruitment, a Ricker‐type behaviour with density dependent overcompensation when environmental conditions are unfavourable and a Beverton–Holt‐type behaviour towards an asymptotic recruitment carrying capacity with favourable environmental conditions. The Neural Network model also detected that under favourable environmental conditions high spawning stock biomass does not necessarily have a depensatory effect on recruitment. Moreover, they suggest that under extremely favourable environmental conditions, albacore recruitment could increase well above the asymptotic carrying capacity predicted by Beverton–Holt‐type models. However, the general decrease in spawning stock biomass in recent years and increasing NAO trends suggest that there is low probability of exceptionally large recruitment in the future and instead there is a danger of recruitment overfishing.     

The diet of Chesapeake Bay striped bass in the late 1950s

The diet of Chesapeake Bay striped bass, (Morone saxatilis) Walbaum, based on unpublished stomach content data from 916 fish collected between 1955 and 1959 was described. The diet in the late 1950s, quantified using an index of relative importance (IRI), was dominated by Atlantic menhaden, Brevoortia tyrannus Latrobe. Atlantic menhaden (66%) and bay anchovy, Anchoa mitchilli Valenciennes, (19%) had the highest IRI value overall. Small striped bass (<600 mm total length) ate predominantly bay anchovy (IRI = 67%). Large striped bass (≥ 600 mm total length) ate predominantly Atlantic menhaden (IRI = 93%). Since 1990 small striped bass rely more on invertebrate prey and larger fish now rely more on small pelagic prey, such as bay anchovy and 0‐age clupeids. Analysis of historical data using current techniques provided a valuable tool for comparison to help in understanding the current striped bass predator–prey relationship in Chesapeake Bay.     

Oceanographic drivers of sablefish recruitment in the California Current

Oceanographic processes and ecological interactions can strongly influence recruitment success in marine fishes. Here, we develop an environmental index of sablefish recruitment with the goal of elucidating recruitment‐environment relationships and informing stock assessment. We start with a conceptual life‐history model for sablefish Anoplopoma fimbria on the US west coast to generate stage‐ and spatio‐temporally‐specific hypotheses regarding the oceanographic and biological variables likely influencing sablefish recruitment. Our model includes seven stages from pre‐spawn female condition through benthic recruitment (age‐0 fish) for the northern portion of the west coast U.S. sablefish stock (40°N–50°N). We then fit linear models and use model comparison to select predictors. We use residuals from the stock‐recruitment relationship in the 2015 sablefish assessment as the dependent variable (thus removing the effect of spawning stock biomass). Predictor variables were drawn primarily from ROMS model outputs for the California Current Ecosystem. We also include indices of prey and predator abundance and freshwater input. Five variables explained 57% of the variation in recruitment not accounted for by the stock‐recruitment relationship in the sablefish assessment. Recruitment deviations were positively correlated with (i) colder conditions during the spawner preconditioning period, (ii) warmer water temperatures during the egg stage, (iii) stronger cross‐shelf transport to near‐shore nursery habitats during the egg stage, (iv) stronger long‐shore transport to the north during the yolk‐sac stage, and (v) cold surface water temperatures during the larval stage. This result suggests that multiple mechanisms likely affect sablefish recruitment at different points in their life history.     

Biological investigations of anchovy and some selected fish species caught during the R／V “Bei Dou”surveys 1984—1999

IntroductionProductivity of an ecosystem may be measured in terms of structure,abundance and growth of keyspecies.Stock structure,measured as age or size composition in a given stock may indicate whether the stockis in a state of growing,stable or declining.Individual growth may be measured in terms of annual growth ofindividual age groups and maturity of key species will similarly reflect the potential of the species to copewith the influence of a changing environment of predators and human i…     

Investigating the ‘northern Humboldt paradox’ from model comparisons of small pelagic fish reproductive strategies in eastern boundary upwelling ecosystems

Eastern boundary upwelling ecosystems are highly productive and sustain the world’s largest fisheries, usually dominated by sardine and anchovy species. Stock size is highly variable from year to year due to the impact of the unstable physical environment on fish early stages. Biophysical models of early life‐stage dispersal of marine organisms have been built by coupling (i) hydrodynamic models and (ii) life history models (i.e. egg and larva stages), and are therefore useful tools to investigate physical–biological interactions. Here, we review biophysical models of anchovy and sardine ichthyoplankton dispersals developed in the Benguela, Humboldt and Canary Current upwelling ecosystems. We also include a similar study conducted in the California Current upwelling on zooplankton. We then integrate this information into a comparative analysis of sardine and anchovy reproductive strategies in the different systems. We found that the main spawning periods match the season of (i) maximal simulated ichthyoplankton retention over the continental shelf in the northern Benguela, southern Humboldt and Canary (for sardine); (ii) maximal food concentration in the southern Benguela, California and Canary (for anchovy); and (iii) maximal shelf retention of ichthyoplankton and food concentration in the northern Humboldt (for both anchovy and sardine). This specificity of the northern Humboldt ecosystem could explain why it sustains the largest small pelagic fish stock. Finally, the possible effects of climate change on these patterns are discussed.     

Reproductive success of the Argentine anchovy,Engraulis anchoita,in relation to environmental variability at a mid‐shelf front (Southwestern Atlantic Ocean)

The mid‐shelf front (MSF) of the Buenos Aires province continental shelf in the Southwestern Atlantic Ocean plays a central role in the pelagic ecosystem of the region acting as the main spring reproductive area for the northern population of the Argentine anchovy Engraulis anchoita and supporting high concentrations of chlorophyll as well as zooplankton, the main food of anchovy. To investigate the influence of environmental variability on the reproductive success of E. anchoita, we analyzed a 13‐yr time series (1997–2009) of environmental data at MSF including chlorophyll dynamics, as well as zooplankton composition and abundance, ichthyoplankton distributions, and recruitment of E. anchoita. Spring chlorophyll concentrations showed high interannual variability and were mainly influenced by changes in water temperature and vertical stratification, which in turn control nutrient supply to the surface. Chlorophyll dynamics (magnitude, timing, and duration of the spring bloom) explained most of the variability observed in E. anchoita recruitment, most likely via fluctuations in the availability of adequate food for the larvae. Our results suggest that satellite ocean color products can be valuable tools for understanding variability in ecosystem dynamics and its effects on the recruitment of fish.     

Effects of temperature,food availability,and body size on daily growth rate of Japanese anchovy <Emphasis Type="Italic">Engraulis japonicus</Emphasis> larvae in Hiuchi-nada

With data sampled from 2003 to 2006, this study presents the effects of temperature, food availability (C), and body size on the somatic growth rate [^(g)] \hat{g} (mm day⁻¹) and on the size-specific growth rates [^(G)] \hat{G} (day⁻¹) of larval Japanese anchovy Engraulis japonicus caught in Hiuchi-nada, Seto Inland Sea, Japan, a semi-enclosed narrow sea. C was calculated using an individual-based model, which is an encounter model between copepods and anchovy where the anchovy resulted in a successful capture of the copepod. [^(g)] \hat{g} decreased with increasing anchovy length, but increased with increasing C. No significant relationship was found between [^(g)] \hat{g} and temperature. [^(G)] \hat{G} decreased with increasing temperature, but anchovy weight, on the other hand, increased with increasing C. In the literature, the mechanism of growth determination for larvae of pelagic fish species is considered to be primarily determined by sea temperature and secondarily by food availability. However, [^(g)] \hat{g} and [^(G)] \hat{G} were found to be dependent on C rather than temperature. [^(G)] \hat{G} was dependent on anchovy weight rather than C and temperature. For forecasting the number of days required for development from an arbitrary pre-recruitment size to the recruitment size, we address not only temperature but also food availability and the size effect when considering the mechanism of growth determination for larvae.