Long-term variation in the abundance of the brown shrimp Crangon crangon (L.) population of the German Bight and possible causes for its interannual variability

Time series on Crangon crangon densities in the German Wadden Sea show a considerable degree of interannual variability, for the entire region in spring and in autumn as well as for three subareas, North Frisia, East Frisia and Elbe estuary. Across the entire survey area C. crangon density was inversely related to water depth. In autumn after the recruitment, settlement shrimp densities are correlated across a larger spatial scale (>100 km longshore), whereas in spring the overwintering adult spawning stock is uncorrelated across subareas. Interannually, extreme density variations can develop even over a short time span of 1 yr. On a large spatial scale shrimp abundance in autumn was correlated with year‐to‐year changes in physical environmental and biological parameters, winter water temperature, autumn river runoff and the winter NAO index. On a regional scale (North Frisia), density of gadoid predators was an additional component affecting shrimp stock abundance. No correlations to such parameters could be detected for the C. crangon stock in spring. Possible causes for the spring situation and additional sources for variability are discussed.     

Population dynamics of the brown shrimp Crangon crangon L., in shallow areas of the German Wadden Sea

Settlement, growth and mortality of the brown shrimp, Crangon Crangon L., were studied in the north Frisian Wadden Sea over a period of 19 months to fill existing gaps regarding the species' life cycle, and thereby to clarify points related to the fishery. Settlement of postlarval shrimp in the shallow areas occurred in three batches: in May; during summer; and an overwintering batch. Compared with an earlier model based on laboratory-derived data, a new growth model based on field data predicts similar growth rates for the earlier phase of the shrimp life cycle, but higher rates for the later stages of the cohorts. Mean growth rate of the different cohorts over the size range sampled was 0.23 ± 0.08 mm day^?1. Of the factors tested, only temperature significantly influenced growth rates. Preliminary estimates of instantaneous total mortality values for shrimp cohorts from these shallow areas ranged from 0.03 to 0.12 day^?1. Salinity explained most of the variation in mortality rates, but this most likely indicates an indirect effect of depth on emigration. The cumulative effects of growth rates estimated from the new model and lower losses through predation could imply a higher contribution of winter eggs to the autumn fishery. This, however, needs further evaluation to properly quantify the trade-offs between settlement and recruitment to the fishery, as well as the growth and mortality rates.     

Life cycle of Crangon crangon in the North Sea: a simulation of the timing of recruitment as a function of the seasonal temperature signal

The brown shrimp C. crangon supports a large commercial fishery in British, Belgian, Dutch, German and Danish waters. It produces eggs throughout the year with two seasonal peaks in summer and winter, respectively. Uncertainty exists with regard to the relative importance of the two egg production seasons for the mass invasion of juvenile recruits of 10–20 mm length in May/June, which is the dominant seasonal signal in the German and Dutch Wadden Sea, and which is presumed to grow into the exploitable stock by the autumn, causing the typical rise of commercial catches at that time of the year. A simulation model was developed that predicts the daily abundance of juvenile recruits attaining a given length, typically in the range of 10–20 mm. The model uses: (i) experimental data on the development times of eggs and larvae and the growth rates of juveniles; (ii) field data on the seasonal temperature cycle in different years (1986, 1992 and 1993 or mean conditions) and areas (German and Dutch Wadden Sea) of the North Sea; and (iii) a calculated index of the relative seasonal egg production intensity of adult C. crangon . Predictions of the simulations are compared with field observations on the seasonal occurrence of juvenile recruits in the German and Dutch Wadden Sea. Using temperature data from German waters, in the simulations peak recruitment was predicted to occur 1–2 months later than that observed in the field. However, if seasonal temperature data from Dutch waters were used, the predicted time of the first mass occurrence of recruits matched the field observations more closely. Simulations revealed that the first mass invasion of juveniles originates almost entirely from the winter egg production. It was also found that the simulated recruitment is condensed into a peak, which is narrower after a cold winter.     

Early life history and recruitment of black crappie (Pomoxis nigromaculatus) in two South Dakota waters

Abstract— We compared the early life history of black crappie ( Pomoxis nigromaculatus ) in Richmond (an impoundment) and Brant (a natural lake) lakes during 1994-1996. We expected variable recruitment (i. e., missing year classes) in the natural lake and more consistent recruitment in the impoundment. Larval black crappie abundance was always higher in Richmond Lake than Brant Lake. Peak abundance of larval black crappie was highest during 1994 in both waters. However, peak larval abundance did not correspond with fall trap-net catch per unit effort (CPUE) of age-0 black crappie, which was highest during 1995. Thus, recruitment of black crappie was not simply a function of the number of larvae hatched. Similar trends in catches of age-0 black crappie with the ichthyoplankton trawl and trap nets for both water bodies during 1994-1996 suggest that broad environmental factors similarly affected reproduction and recruitment in Richmond and Brant lakes.     

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.     

Linking cod (Gadus morhua) and climate: investigating variability in Irish Sea cod recruitment

How climatic variability and anthropogenic pressures interact to influence recruitment is a key factor in achieving sustainable resource management. However, the combined effects of these pressures can make it difficult to detect non‐stationary interactions or shifts in the relationships with recruitment. Here we examine the links between climate and Irish Sea cod recruitment during a period of declining spawning stock biomass (SSB). Specifically, we test for a shift in the relationship between recruitment, SSB and climate by comparing an additive (generalized additive model, GAM) and non‐additive threshold model (TGAM). The relationship between recruitment success, SSB and the climatic driver, sea surface temperature, was best described by the TGAM, with a threshold identified between recruitment and SSB at approximately 7900 t. The analysis suggests a threshold shift in the relationship between recruitment and SSB in Irish Sea cod, with cod recruitment being more sensitive to climatic variability during the recent low SSB regime.     

The missing link: tidal‐influenced activity a likely candidate to close the migration triangle in brown shrimp Crangon crangon (Crustacea,Decapoda)

Life cycle closure for species inhabiting areas with daily varying currents but directed net water transport requires specific behavior to minimize losses due to advection of passive drifting life stages. Variations in swimming activity of different‐sized Crangon crangon (15–65 mm total length) were therefore monitored under constant laboratory conditions immediately after being caught in the German Wadden Sea. Activity of shrimps of different sizes, caught at different seasons, always peaked at times corresponding with ebb tide in the habitat from where they were taken. This behavior was maintained for several days if no external stimuli were present but shifted to night activity if a light–dark cycle was provided. The observed behavior/activity pattern was included in a coupled hydrodynamic and individual‐based model (IBM) and the shift in the location of a shrimp cohort was monitored over time. Performance of ebb tide activity not only allowed the shrimps to reach the preferred deeper winter and spawning areas but also allowed them to migrate against the dominating current from eastern nurseries to more western located spawning areas. Passively drifting larvae released at these locations and later larval and juvenile stages that perform flood tide transport can reach the nurseries again. This links the nurseries and adult spawning grounds and closes the migration triangle.     

Causes of walleye pollock (Theragra chalcogramma) recruitment decline in the northern Sea of Japan: implications for stock management

Recruitment of the northern Japan Sea stock (JSS) of walleye pollock has been decreasing since around 1990. In this study, I analyzed the factors causing this decrease in recruitment by investigating the relationship between recruitment, spawning stock biomass (SSB) and environmental factors using a generalized additive model (GAM). GAM fit to the data showed the importance of SSB, sea surface temperature (SST), ocean current strength (Tsushima Warm Current) and wind intensity (Asian monsoon) in determining the recruitment. Of these, the relationship between SSB and recruitment was positive and not negatively density‐dependent. On the other hand, the recruitment was negatively related to SST and ocean current strength, and a dome‐shaped relationship was observed between wind intensity and recruitment. Since around 1990, the values of SST and ocean current strength have mostly been high and that of wind intensity mostly low. In addition, SSB has been decreasing since the late 1990s. It is likely that the recruitment decline of JSS after approximately 1990 has been caused by warm water temperature, strong Tsushima Warm Current and weak Asian monsoon, and that the recent decrease in SSB has amplified this recruitment decline. According to the model’s estimation, a recruitment recovery due to environmental improvement will be highly restricted as long as SSB remains at its current low level. Significant recovery of SSB is urgently needed for JSS.     

Towards a more complete understanding of the life cycle of brown shrimp (Crangon crangon): modelling passive larvae and juvenile transport in combination with physically forced vertical juvenile migration

In this study we developed and utilized a complex model approach to investigate the impact of stage‐specific transport processes on the development and spatial distribution of brown shrimp (Crangon crangon) post‐larvae and juveniles in the German Bight. First, we focused on drift processes during the pelagic larval stage by coupling an individual‐based model for egg and larval development ‘off‐line’ to a 3D hydrodynamic model utilizing the Lagrangian method. Secondly, we investigated tidal‐induced transport processes after juvenile settlement. To determine the tidal cycle, the model coupling was accomplished ‘on‐line’ by resolving the individual‐based model and hydrodynamic model with the same time step. The vertical migration of juveniles, a prerequisite for the selective tidal stream transport (STST), was modelled as a sub‐grid scale physical process (balance of forces: gravitation, buoyancy, Stoke’s friction and dynamic uplift) and considered complex particle dynamics. We applied the model to test temperature and salinity cues as possible tidal indicators utilized by juvenile brown shrimp. Our results indicated that transport processes could significantly change the timing and spatial distribution of post‐larval abundance. We also showed that the small‐scale hydrodynamic forcing acting on the bodies of juvenile brown shrimps was sufficient to account for the vertical migration required to use STST. For both investigated tidal cues STST performing juvenile brown shrimp were transported on‐shore. A faster and more continuous STST was calculated for the salinity cue, resulting in larger abundances of brown shrimp in estuarine areas.     

Influence of larval transport and temperature on recruitment dynamics of North Sea cod (Gadus morhua) across spatial scales of observation

The survival of fish eggs and larvae, and therefore recruitment success, can be critically affected by transport in ocean currents. Combining a model of early‐life stage dispersal with statistical stock–recruitment models, we investigated the role of larval transport for recruitment variability across spatial scales for the population complex of North Sea cod (Gadus morhua). By using a coupled physical–biological model, we estimated the egg and larval transport over a 44‐year period. The oceanographic component of the model, capable of capturing the interannual variability of temperature and ocean current patterns, was coupled to the biological component, an individual‐based model (IBM) that simulated the cod eggs and larvae development and mortality. This study proposes a novel method to account for larval transport and success in stock–recruitment models: weighting the spawning stock biomass by retention rate and, in the case of multiple populations, their connectivity. Our method provides an estimate of the stock biomass contributing to recruitment and the effect of larval transport on recruitment variability. Our results indicate an effect, albeit small, in some populations at the local level. Including transport anomaly as an environmental covariate in traditional stock–recruitment models in turn captures recruitment variability at larger scales. Our study aims to quantify the role of larval transport for recruitment across spatial scales, and disentangle the roles of temperature and larval transport on effective connectivity between populations, thus informing about the potential impacts of climate change on the cod population structure in the North Sea.     

The influence of life history dynamics and environment on the determination of year class strength in North Sea herring (Clupea harengus L.)

The inter-annual variability in year class strength (1976–2000) of North Sea herring (Clupea harengus) was investigated using Paulik diagrams based on survey data and Virtual Population Analysis. The herring life cycle was split into five stages: spawning stock biomass (SSB), egg production, larvae, fish with 0 winter rings on the otolith (0-wr), 1-wr and 2-wr. Surveys were used as indices and Paulik analysis revealed relationships between stages. In 80% of the years, year class strength reflected SSB. Poorer than expected year classes were determined during the larva to 0-wr phase, whilst stronger than expected year classes were apparently determined during the 0-wr to 1-wr stage. There was no clear relationship between survival of young stages of herring and the abundance of Calanus finmarchicus but the year class strength of 0-wr and 1-wr had a negative relationship to bottom water temperature. Lower sea water temperatures in the North Sea are associated with higher Calanus abundance. The analysis shows that the strength of aberrant year classes of North Sea herring is determined between the pelagic larval and the juvenile stages.     

Abiotic and biotic factors affecting recruitment variability of walleye pollock (Theragra chalcogramma) off the Pacific coast of Hokkaido,Japan

Abiotic and biotic factors affecting the recruitment variability of the Japanese Pacific stock (JPS) of walleye pollock (Theragra chalcogramma) were examined using a bivariate regression and multivariate combined model. Of the abiotic variables around Funka Bay (spawning ground), February sea surface temperature (SST) and wind direction index showed significant bivariate relationships with recruitment. February SST was positively related to recruitment, suggesting that warmer water temperature in February favors JPS recruitment. On the other hand, the relationship between February wind direction index and recruitment predicts high JPS recruitment under predominant northwest winds in February. For the biotic variables in the Doto area (nursery ground), significant and negative bivariate relationships with recruitment were observed for catch per unit effort of Kamchatka flounder (Atheresthes evermanni), Pacific cod (Gadus macrocephalus), and walleye pollock, implying an important impact of predation by these groundfishes on JPS recruitment. The overall model incorporating these abiotic and biotic factors successfully reproduced the variability in JPS recruitment. Temperature and wind conditions around the spawning ground along with predator condition in the nursery ground appear to play a dominant role in the recruitment dynamics of JPS. Based on these results and prior knowledge, we propose a new hypothesis to explain the processes controlling JPS recruitment.     

Environmentally driven forecasts of northern rock sole (Lepidopsetta polyxystra) recruitment in the eastern Bering Sea

Northern rock sole recruitment in the eastern Bering Sea has been hypothesized to (a) depend on wind‐driven surface currents linking spawning and nursery areas, (b) be density‐dependent, and (c) be negatively impacted by cold bottom temperatures over a large nursery area during the first summer of life. A suite of models was developed to test these hypotheses. Data included 32 years of recruitment and spawning biomass estimates derived from a stock assessment model and wind and temperature indices customized to the environmental exposure of age‐0 northern rock sole in the eastern Bering Sea. The predictive ability of the models was evaluated, and the models were used to forecast recruitment to age‐4 for recent year classes which are poorly retained by the standard multi‐species bottom trawl survey gear. Models which included wind and temperature indices performed better than a naïve forecast based on the running mean. The best‐performing model was a categorical model with wind and temperature thresholds, which explained 49% of the variation in recruitment. Ricker models performed more poorly than models without a spawning biomass term, providing no evidence that recruitment is related to stock size. The models forecast higher recruitment for the most recent year classes (2015–2018) than for prior year classes with observed poor recruitment (2006–2013). These environment‐based recruitment forecasts may improve recruitment estimates for the most recent year classes and facilitate study of the effects of future climate change on northern rock sole population dynamics.     

Long-term stock assessment and growth changes of the Japanese sardine (Sardinops melanostictus) in the Sea of Japan and East China Sea from 1953 to 2006

We estimated the stock size of Japanese sardine ( Sardinops melanostictus ) in the Sea of Japan and East China Sea since 1953 using cohort analysis based on the changes of growth patterns. Growth of Japanese sardine, estimated by using annual rings on archived scales since 1961, showed that body lengths were extremely stunted in the 1980–1987 year-classes. The body length at age 3 from February to April in the 1980–1987 year-classes, a period when the stock size exceeded 4 million tons, was 180.0 ± 2.6 mm (mean ± SD), and in the other year-classes was 195.1 ± 7.6 mm. The body length at age 3 and wet weight of zooplankton in August in the offshore area of the Sea of Japan had a significantly positive correlation. We assumed three scenarios for maturation ratios, and estimated Ricker's spawner–recruitment relationships. We analyzed the correlations between logarithmic recruitment residuals (LNRR) and environmental factors in winter, represented by the North Pacific index (NPI), Aleutian low pressure index (ALPI), Pacific decadal oscillation (PDO), monsoon index (MOI), Arctic oscillation (AO) and Southern oscillation index (SOI). Significant correlations were observed between MOI and LNRR and between AO and LNRR. A combination of strong MOI and weak AO would increase the biomass of phytoplankton and zooplankton and subsequently increase the recruitment of Japanese sardine.     

东、黄海星康吉鳗生长、死亡和单位补充量渔获量

依据2011年、2016年和2017年在东、黄海进行的底拖网调查数据,研究了星康吉鳗的生长参数、死亡系数和单位补充量渔获量(YPR)。星康吉鳗的体长体质量关系和生长方程的拟合结果表明,其条件因子a估计值为4.5×10–4,其异速生长系数b的估计值为3.3。其渐近体长L∞的估计均值为102 cm,生长速率K的均值为0.21/年,理论上体长为零时的年龄t0估计均值为–0.19。通过Pauly经验公式求得星康吉鳗的自然死亡系数(M)为0.33。体长转换的渔获曲线求得总死亡系数为3.36,进而求得现阶段的捕捞死亡系数(F)为3.03,开发率高达90%。此外求得现阶段星康吉鳗的开捕体长为30 cm,对应的开捕年龄为1.47龄,远小于其体质量生长的拐点年龄(3.70龄)和临界年龄(3.39龄)。本研究根据单位补充量渔获量模型,估计其在不同自然死亡系数和开捕体长的情况下随捕捞死亡系数的变化曲线,并估算其生物学参考点。随F增加,YPR先增加到最大值,再逐渐减小。现阶段星康吉鳗的YPR为27.14,而当M为0.33时Fmax估计值为0.38,YPRmax为52.89,即将F降低为1/8,可获得将近2倍的YPR;对应的F0.1为0.255,比Fmax降低了三分之一,YPR0.1(50.38)只比最大值降低了不足5%。M越大,求得的生物学参考点越大,可获得的YPR越小;当开捕体长增加时,F和YPR都会有不同程度的增加。因此,现阶段中国东黄海星康吉鳗的开发率过高,开捕体长过小,处于过度捕捞的状态。为了维持星康吉鳗种群的生态健康、实现该渔业的可持续发展,建议大幅削减其捕捞强度,降低捕捞死亡系数,同时增加开捕体长。     

Reconstruction of Atlantic herring (Clupea harengus) recruitment in the North Sea for the past 455 years based on the δ13C from annual shell increments of the ocean quahog (Arctica islandica)

Understanding the recruitment variability of the Atlantic herring North Sea stock remains a key objective of stock assessment and management. Although many efforts have been undertaken linking climatic and stock dynamic factors to herring recruitment, no major attempt has been made to estimate recruitment levels before the 20th century. Here, we present a novel annually resolved, absolutely dated herring recruitment reconstruction, derived from stable carbon isotope geochemistry (δ¹³C), from ocean quahog shells from the Fladen Ground (northern North Sea). Our age model is based on a growth increment chronology obtained from fourteen shells. Ten of these were micromilled at annual resolution for δ¹³C analysis. Our results indicate that the anthropogenically driven relative depletion of ¹³C, the oceanic Suess effect (oSE), became evident in the northern North Sea in the 1850s. We calculated a regression line between the oSE‐detrended δ¹³C results (δ¹³C?) and diatom abundance in the North Sea, the regression being mediated by the effect of phytoplankton on the δ¹³C of the ambient dissolved inorganic carbon. We used this regression to build an equation mediated by a nutritional link to reconstruct herring recruitment using δ¹³C?. The reconstruction suggests that there were five extended episodes of low‐recruitment levels before the 20th century. These results are supported by measured recruitment estimates and historical fish catch and export documentation. This work demonstrates that molluscan sclerochronological records can contribute to the investigation of ecological baselines and ecosystem functioning impacted by anthropogenic activity with implications for conservation and stock management.     

Environmental control of Northeast Atlantic mackerel (Scomber scombrus) recruitment in the southern Bay of Biscay: case study of failure in the year 2000

We investigate the effect of strong meteorological perturbations in early spring on the success of mackerel (Scomber scombrus) recruitment in the N/NW Iberian area (southern Bay of Biscay) for the period 1999–2008. In 2000, the year of the most pronounced recruitment failure on record, two consecutive multidisciplinary surveys sampled hydrographic conditions and mackerel eggs, larvae and post‐larvae over the main mackerel spawning grounds of the north and northwest coast of the Iberian Peninsula. Analysis of egg and larval abundance and birthdates based on the otoliths of mackerel juveniles caught between July and October 2000 showed that there were no survivors from the early spring spawns, indicating a massive loss of early spawning effort. Moreover, the abundance of 1‐year‐old mackerel estimated from an acoustic survey carried out in 2001 was the lowest observed within the 1999–2008 time series. This low or null survival from the early spawns in 2000 could be due to the meteorological and oceanographic conditions of that spring, in particular two storm events in April after a relatively calm March. The first storm event from the north caused strong local wind in the southern Bay of Biscay but a weak oceanographic response. The second storm event from the southwest was mainly felt west of Galicia and caused a notable increase in shelf currents and a shift of the hydrographical structure along the shelf. Detailed analysis of strong wind pulses in early spring within the historical recruitment record suggests that strong local turbulence generated by high wind speeds and advection of larvae caused by the enhancement of shelf currents can contribute to reduced recruitment. Our observations indicate that, in 2000, both mechanisms were present.     

Investigations into the development of the pituitary gland-thyroid tissue axis and distribution of tissue thyroid hormone content in embryonic coho salmon (Oncorhynchus kisutch) from Lake Ontario

Total organism content of L-thyroxine (T₄) and triiodo-L-thyronine (T₃) were measured in the early developmental stages of a stock of Lake Ontario coho salmon from the egg to the yolk absorption stage. Whole organism T₄ levels were constant between the egg and pre-hatch embryo stages, but fell progressively during yolk absorption. T₃ levels were low from egg to eye-pigment appearance, but then increased prior to hatch and fell again during the post-hatch yolk absorption period. When expressed as ng/tissue, T₄ content of the body compartment rose progressively between days 67 and 87 post-fertilization, whilst T₄ content of the yolk compartment fell progressively during the same period; the pattern was not evident for tissue T₃ content. When expressed as ng/g dry weight of tissue, the inverse relationship was found for T₄, and T₃ content of the body and yolk compartments decreased progressively and increased progressively, respectively during the same period, suggesting that thyroid hormones were selectively retained in the yolk compartment. Intensely “immunostained” (using anti-human β-TSH antibody) thyrotropic cells were present in small numbers in the pars distalis of the embryonic pituitary at the eye-pigment appearance stage, and the numbers increased markedly until the pre-hatch period. Administration of either bovine thyrotropic hormone (bTSH) or ovine growth hormone (oGH) had no effect on thyroid hormone content of larvae challenged during the yolk absorption period, suggesting that the thyroid tissue was not responsive to exogenous bTSH challenge at this time, and that oGH-sensitive 5′-monodeiodination was either not present or at levels that were too low to cause an elevation in total T₃ content, or that the substrate levels were insufficient to permit a measureable increase in whole body T₃ content.     

Estimation of seasonal spawning ground locations and ambient sea surface temperatures for eggs and larvae of Pacific saury (Cololabis saira) in the western North Pacific

Since there have been practically no surveys of the eggs of Pacific saury (Cololabis saira) in the western North Pacific (WNP), its spawning ground (SG) distribution has been poorly resolved, based mainly on the larval distribution. This means of estimating SG distribution is imprecise because saury eggs drift for more than a week before they hatch, in a region with intense western boundary currents and their extensions. To improve our understanding of the immature saury, a large number of larvae (body length <25 mm) collected in the WNP during 1993–96 were numerically backtracked to take into account the advection by geostrophic and wind‐forced Ekman currents, and the SG locations and ambient sea surface temperatures (SSTs) for the eggs and larvae on the backtracking trajectories were estimated. The resulting seasonal distributions of SGs indicated that both the locations and the intensities of spawning change from season to season. Moreover, the ambient SSTs for eggs just after fertilization ranged from a high of around 21.5°C in early autumn (September to October) to a low of around 15.0°C in late spring (May to June) with an intermediate of around 20.0°C in winter (January to February). The ambient SSTs showed seasonally different gradients while the individuals developed from eggs to early larvae: the SSTs decreased throughout the autumn (September to December), stayed rather constant in winter (January to February), and increased throughout the spring (March to June). The ambient SSTs for the early larvae were at around 19.0°C in autumn and winter (September to February) and around 16.5°C throughout the spring (March to June).