Covariation in climate, zooplankton biomass and mackerel recruitment in the southern Gulf of St Lawrence

Between 1982 and 1991, an annual survey of stage I egg production of Atlantic mackerel ( Scomber scombrus ) was conducted in June/early July in the southern Gulf of St Lawrence. We investigated the relationship between interannual variability in biomass of zooplankton, determined from the archived survey plankton samples, and mackerel recruitment, estimated from the proportion of three-year-olds in the catch of the commercial fishery. Zooplankton biomass varied by a factor of 2.5, primarily owing to fluctuations in the >1000 μm size fraction. The index of mackerel recruitment fluctuated by a factor of ≈20 and was positively related (linear regression: P  < 0.05; n  = 10) to variations in the zooplankton biomass. Both mackerel recruitment and zooplankton biomass were negatively related (linear regression: P  < 0.05) to RIVSUM, a measure of freshwater discharge from the St Lawrence River system and an index of variability in the region's climate. Three hypotheses are put forward to explain these observations: (1) there is a strong link between interannual variation in abundance of copepod females, which produce prey for mackerel larvae, and larval survival; the exceptional recruitment and subsequent year class resulted from an exceptional production of Calanus finmarchicus nauplii; (2) years of high zooplankton biomass provide better feeding conditions and consequently higher survival of mackerel juveniles; and (3) mackerel recruitment and zooplankton biomass are independently under the control of an underlying physical process, without strong trophic linkage. The first hypothesis is supported by a study of copepod species composition and female abundance conducted for four of the survey years. At the present time, none of these hypotheses can be ruled out.     

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.     

Production, Quality, and Low Temperature Incubation of Eggs of Atlantic Cod Gadus morhua and Haddock Melanogrammus aeglefinus in Captivity

Atlantic cod Gadus morhua and haddock Melanogrammus aeglefinus broodstock maintained under altered regimens of temperature and photoperiod spawned up to 8 mo per year. The cod broodstock produced viable embryos from October through June. The haddock broodstock produced viable embryos from December through May. Egg diameters were largest during the middle of the spawning season when water temperature was at a minimum, resulting in an inverse relationship between egg diameter and water temperature in both species. Egg quality was high, as evaluated by buoyancy, fertilization rate, regularity of early cleavage, and percent viable hatch. Low temperature incubation of cod and haddock eggs extended the embryonic period. Cod embryos tolerated a wider range of temperatures than haddock. High mortality (1 90%) was observed before hatching in haddock embryos incubated at 1 C. Atlantic cod embryos hatched at temperatures as low as —1 C, extending the embryonic period to 59 d. At 8 C Atlantic cod and haddock embryos hatched in 11–12 d. To determine if extending the embryo incubation time by using low temperatures had a detrimental effect, embryos were incubated through hatch at either 1 C or 6 C, and the larvae from both groups reared at 6 C. Growth and early survival of larvae were comparable in both treatments.     

Modelling the retention and survival of Browns Bank haddock larvae using an early life stage model

Browns Bank, the principal spawning ground for haddock on the south-west Nova Scotian shelf, is composed of two distinct production zones: the inshore Bay of Fundy (BoF) region and the offshore south-west Nova (SWN) bank region. Fish growing in the BoF are larger at age than those in the SWN region. Analysis of research vessel (RV) data shows that the majority of age-2 haddock have the size of south-west Nova Scotia fish, suggesting that the majority of surviving fish were retained in SWN. We used an early life stage (ELS) model to address two questions. First, we asked whether the length-at-age difference between Bof and SWN is evident at the larval stage. Using a temperature-based growth model, we found that predicted size differences for late larvae would be less than 0.5 mm. From consideration of the average growth curves to age-2 for the two regions, we showed that this difference was not the seed for the size difference in later life stage. The second question we addressed was how well the ELS model predicted partitioning of late larvae between SWN and the BoF corresponding to the ratio of SWN/BoF age-2 juveniles found in the RV data. We ran the model on a representative number of years between 1973 and 1992, and found a significant correlation between the model simulation and the data (cor=0.71, P =0.047). This indicates that the age-2 size distribution is a reflection of retention and survival occurring during the first two months of life. Model/data discrepancies are explained in terms of differential mortality between the two regions.     

Modeling larval Calanus finmarchicus on Georges Bank: time-varying mortality rates and a cannibalism hypothesis

Throughout the North Atlantic, the copepod Calanus finmarchicus dominates the zooplankton biomass, linking primary production and higher trophic levels. On Georges Bank, the peak abundance of larval (naupliar) stages occurs in March–April and represents a potential source of prey for cod and haddock larvae. Following this maximum, naupliar abundance declines dramatically, reaching a minimum in May and increasing again in June. Explaining the naupliar seasonal cycle is critical for predicting climate effects on C. finmarchicus dynamics, including whether environmental variability may lead to a mismatch with larval fish. Here, an age-within-stage population dynamics model is used to investigate the factors controlling the temporal variation of C. finmarchicus nauplii in three Georges Bank sub-regions. The model incorporates temperature- and food-dependent development and egg production, as well as female abundance derived from the US Global Ocean Ecosystem Dynamics (GLOBEC) program. Use of field-estimated constant mortality rates overestimates May abundances by as much as an order of magnitude. These data/model discrepancies can not be explained by temperature or food-limitation effects on physiological rates. Instead, accurate simulation requires use of time-varying early stage mortalities, which differ from published estimates in both magnitude and trend. These mortality rates are correlated with C. finmarchicus female abundance, implying cannibalism as a possible regulatory factor. Thus, the biological control of predation (including cannibalism) must be considered to predict the effects of climate on C. finmarchicus and associated larval fish populations.     

Interannual differences in larval haddock survival: hypothesis testing with a 3D biophysical model of Georges Bank

The ultimate goal of early life studies of fish over the past century has been to better understand recruitment variability. As evident in the Georges Bank haddock (Melanogrammus aeglefinus) population, there is a strong relationship between recruitment success and processes occurring during the planktonic larval stage. This research sought new insights into the mechanisms controlling the recruitment process in fish populations using biological–physical modeling methods together with laboratory and field data sets. We created the first three‐dimensional model of larval haddock on Georges Bank by coupling models of hydrodynamics, lower trophic levels, a single copepod species, and larval haddock. Interactions between feeding, metabolism, growth, vertical behavior, advection, predation, and the physical environment of larval haddock were quantitatively investigated using the coupled models. Particularly, the model was used to compare survival over the larval period and the sources of mortality in 1995 and 1998, 2 years of disparate haddock recruitment. The results of model simulations suggest that the increased egg hatching rates and higher food availability, which reduced starvation and predation, in 1998 contributed to its larger year‐class. Additionally, the inclusion of temperature‐dependent predation rates produced model results that better agreed with observations of the mean hatch date of survivors. The results from this biophysical model imply that food limitation and its related losses to starvation and predation, especially from hatch to 7 mm, may be responsible for interannual variability in recruitment and larval survival outside of the years studied.     

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.     

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.     

Climate‐induced environmental conditions influencing interannual variability of Mediterranean bluefin (Thunnus thynnus) larval growth

Daily growth variability of bluefin (Thunnus thynnus) larvae sampled in their Balearic Sea spawning grounds during the 2003–2005 spawning seasons was examined. Multi‐factorial ANOVA was applied to study the effects of environmental variables, such as temperature at 10 m depth (T10), microzooplankton dry weight (MDW) and protein/dry weight ratio (PROT/MDW) on larval growth. The 2003 bluefin tuna (BFT) larval cohort showed the fastest growth, recognizable from enhanced otolith and somatic mass increment compared to the 2004–2005 larval cohorts. The 2003 BFT larvae showed greater recent growth than the 2004–2005 BFT cohorts, which decreased in the last stages of development. Growth differences between the 2004 and 2005 larval cohorts were not significant. The environmental conditions between 2003 and 2004–2005 were highly contrasting as a result of the 2003 warming anomaly. Somatic and otolith growth rates (OGR) were significantly related to T10 and MDW, as well as to the PROT/MDW ratios. Nonetheless, the effect of T10 on OGR depended on the relative high (H) or low (L) levels of MDW and PROT/DW. Higher OGR was observed when T10 was high, MDW was low and PROT/DW was high. This environmental scenario conditions were met during 2003, which recorded the highest surface temperature and low planktonic biomass. Somatic growth, expressed as larval DW growth increase (DWGR), showed three‐factor significant interactions with T10*MDW*PROT/MDW, in which the two‐way interactions of MDW*PROT/MDW showed differences in the function of T10 levels.     

温度、盐度对条石鲷胚胎发育影响及初孵仔鱼饥饿耐受力

研究了盐度、温度对条石鲷胚胎发育的影响和初孵仔鱼对饥饿的耐受力。结果表明,胚胎发育的适宜盐度范围为20～35,最适盐度范围为25～30,孵化率都在89%以上,盐度低于20或高于35,孵化率都降低。盐度对孵化时间有一定影响,盐度高于35或者低于15条件下孵化时间都延长。胚胎发育的适宜温度范围为21～27℃,孵化率都在77%以上,最适温度范围为23～25℃,孵化率达89%以上,低于19℃或者高于27℃时孵化率降低且初孵仔鱼畸形率升高。胚胎孵化时间与温度成负相关关系,胚胎发育速率与温度成正相关关系。条石鲷胚胎发育的生物学零度（阈温度）为10.06℃,有效积温值为405.3℃·h。对胚胎发育的Q10值进行了计算。饥饿条件下,条石鲷初孵仔鱼8日龄全部死亡,半数死亡时间出现在第6天,饥饿对初孵仔鱼生长具有明显的抑制作用。     

Hatching time effect on the intra‐spawning larval morphology and growth in Northern pike (Esox luciusL.)

Based on the analysis of 11 morphometric variables of body (total length, body area and perimeter, myotome height and eye diameter) and yolk sac (length, height, area, perimeter and volume) of pike larvae, the aim of this study was to evaluate how larval size at hatching and growth of larvae hatched from single egg batches vary according to three hatching times: early, mid and late. Hatching time structures strongly pike larval morphometrics. Early hatched larvae have smaller body sizes at hatch, faster growth and higher yolk use efficiency than late hatched ones. Early hatched larvae seem to be premature and hatch at precocious developmental stage whereas late hatched individuals continue their growth within the egg shell and hatch at larger size but with lower reserves (yolk). A compensatory growth phase was observed for the early hatching pike larvae particularly during the first 5 days post hatch. Consequently, no significant difference in body parameters was recorded from day 10 post hatching whatever the hatching time. The higher growth accomplished by early hatched larvae may be related to a particular metabolic activity that converts more efficiently yolk into body tissues.     

Modelling the transport of larval sandeels on the north-west European shelf

This study considers the interannual variability in the advection of sandeel ( Ammodytes marinus ) larvae on the north-west European shelf. We use recently collated data on spawning location, age of larvae and the time of hatching (between 1955 and 1993) as input to a particle-tracking model. Utilizing hourly currents from a 39 year run of a two-dimensional sea circulation model, cohorts of sandeel larvae were represented as a distribution of particles in space and time, and particle drift was calculated for the larval life stage. The final distributions of particles were compared with actual distributions of 0-group sandeels in the North Sea, as determined from surveys conducted at around the end of the larval phase. Preliminary examination of the data between 1973 and 1993 indicates that differences in predicted drift could explain some of the large interannual variability in 0-group distributions. This approach may help to improve our understanding of the frequent colonization events that are typical of this species.     

The growth of larval cod and haddock in the Irish Sea: a model with temperature,prey size and turbulence forcing

We applied a physiological individual‐based model for the foraging and growth of cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) larvae, using observed temperature and prey fields data from the Irish Sea, collected during the 2006 spawning season. We used the model to estimate larval growth and survival and explore the different productivities of the cod and haddock stocks encountered in the Irish Sea. The larvae of both species showed similar responses to changes in environmental conditions (temperature, wind, prey availability, daylight hours) and better survival was predicted in the western Irish Sea, covering the spawning ground for haddock and about half of that for cod. Larval growth was predicted to be mostly prey‐limited, but exploration of stock recruitment data suggests that other factors are important to ensure successful recruitment. We suggest that the presence of a cyclonic gyre in the western Irish Sea, influencing the retention and/or dispersal of larvae from their spawning grounds, and the increasing abundance of clupeids adding predatory pressure on the eggs and larvae; both may play a key role. These two processes deserve more attention if we want to understand the mechanisms behind the recruitment of cod and haddock in the Irish Sea. For the ecosystem‐based management approach, there is a need to achieve a greater understanding of the interactions between species on the scale a fish stock is managed, and to work toward integrated fisheries management in particular when considering the effects of advection from spawning grounds and prey–predator reversal on the recovery of depleted stocks.     

Larval group differentiation in Atlantic cod (Gadus morhua) inside and outside the Gullmar Fjord

The spatial and temporal occurrence of pelagic fish stages and their biological variability may affect their dispersal and survival, and ultimately fish recruitment. We collected Atlantic cod larvae at one station inside and at one station outside the Gullmar Fjord, eastern Skagerrak, in order to investigate small-scale larval group differentiation. Rectangular midwater trawl hauls were taken every 6 h (during 24 h) from three separate depth intervals between the surface and 70 m depth. About 80% and 20% of all larvae occurred above the halocline at the Fjord station and the Coastal station, respectively. Hatching (at both stations) occurred from the 3rd week in February to the 1st week in May, indicating that cod larvae were present for at least 5 months (from late February to early August). The length and hatch date frequency distributions of larvae from the surface layer were unimodal inside the fjord but bimodal outside the fjord. Analyses of seven microsatellite DNA loci indicated that larvae collected inside the fjord (where local spawning occurs) were genetically distinct from larvae sampled on the outside (F_ST = 0.0026). The two age cohorts outside the fjord were not, however, genetically different, nor were larvae collected at different depths. We conclude that small-scale variability of vertical concentration and larval life history variability should have consequences for interpreting models of larval dispersal and survival, and subpopulation structure analyses.     

Effect of different diets on larval production, quality and fatty acid profile of the marine ornamental shrimp Lysmata amboinensis (de Man, 1888), using wild larvae as a standard

The present work evaluates the influence of broodstock diets [Marine Cuisine^®– MC, MC supplemented with highly unsaturated fatty acid- (HUFA) enriched Artemia biomass – MC + AB, and MC supplemented with squid – MC + S] on larval production, newly hatched and early zoeal stage survival and fatty acid profile of newly hatched larvae of Lysmata amboinensis . These parameters are compared with those from larvae hatched from embryos spawned in the wild. The number (±SE) of larvae produced with MC and MC + S (1077 ± 219 and 1103 ± 184, respectively) was similar to that in broodstock carrying embryos spawned in the wild (1224 ± 111), while those fed MC + AB displayed significantly lower values (1044 ± 161). Larvae produced with MC + AB displayed lower survival for all starvation periods, while larvae spawned in the wild displayed the highest survival. No larvae resisted 144 h of starvation and none moulted to zoea II. The fatty acid comparison revealed that larvae from embryos spawned in the wild displayed the highest levels of DHA, as well as higher DHA/EPA and n -3/ n -6 ratios. These results suggest that broodstock diets commonly used to promote ornamental shrimp's maturation (based on mixed frozen components) are far from being optimal.     

Effect of Bacillus subtilis on growth development and survival of larvae Macrobrachium rosenbergii (de Man)

A feeding experiment was conducted to investigate the effect of Bacillus subtilis bacterium, on larval growth and development rate of Macrobrachium rosenbergii (de Man) during February 28 to April 8, 2005 in University Putra Malaysia hatchery. Newly hatched larvae of M. rosenbergii were reared with two dietary treatments consisting of newly hatched Artemia salina nauplii with B. subtilis (10⁸ cells ml⁻¹), and newly hatched A. salina nauplii without B. subtilis carried out in triplicate in 60‐L aquarium (50 L⁻¹). After trial, the larvae that fed B. subtilis‐treated Artemia naupli were found to have higher survival and a faster rate of metamorphosis than larvae that were fed with nontreated Artemia naupli. There were significant differences between B. subtilis‐treated Artemia naupli and nontreated Artemia diet in larval growth and development rate of metamorphosis (P < 0.05). Larval survival after 40 days was significantly greater (P < 0.05) in the B. subtilis‐treated groups (55.3 ± 1.02) compared with the nontreated groups (36.2 ± 5.02%).     

Effects of temperature, density and early weaning on the survival and growth of Atlantic ditch shrimp Palaemonetes varians larvae

This investigation examined the effects of temperature, density and early weaning on the survival and growth of Palaemonetes varians larvae. Survival of larvae raised at 17.5 °C was not significantly different (average + standard deviation) (94 ± 5%) from the survival of those raised at 19.5 °C (95 ± 5%) and at 21.5 °C (94 ± 4%). However, the duration of the larval stage was significantly longer for shrimp reared at 17.5 °C (17.3 ± 0.8 days) compared with shrimp reared at 19.5 °C (14.3 ± 0.7 days) and at 21.5 °C (11.3 ± 0.6 days). No significant differences ( P >0.05) were found in the survival rate, final weight and length of larvae reared at the densities of 5, 10, 20 and 50 larvae L⁻¹. The survival of P. varians larvae fed solely on Artemia was significantly higher ( P <0.05) than larvae weaned with an artificial practical diet from Zoea II stage (94 ± 4% and 82 ± 1%, respectively, for Artemia and artificial diet-fed larvae), but no significant differences ( P >0.05) were observed in the final larval weight or length between these two treatments. The survival and growth of the larvae fed with the practical diet tested is a promising step ahead in the development of the culture of this species as it eliminates both the need for Artemia throughout all larval stages, and the need for more expensive artificial diets.     

Effects of temperature and lipid droplet adherence on mortality of hatchery-reared southern hake Merluccius australis larvae

Effects of exogenous (water temperature) and endogenous (lipid droplet adherence) factors were experimentally tested on early survival of southern hake Merluccius australis reared under controlled conditions. Experiments to determine the effect of temperature (10, 12 and 14 °C) on larval growth rates and yolk-sac absorption rates of unfed southern hake were carried out under laboratory conditions. There was no significant differences in growth rates at the temperature range tested (ANCOVA, F = 0.164, p > 0.25), but yolk-sac absorption rates and mortality increased with temperature (ANCOVA, F = 53.84, p < 0.001). A high percentage (between 31 and 81%) of hake eggs showed a lipid droplet not adhered (i.e., freely moving in the yolk, and not located in the posteriormost portion of the yolk-sac). In a second experiment, fed southern hake larvae with the lipid droplet not adhered during embryonic development did not survive after yolk-sac absorption. This study provides the first data on the influence of the lipid droplet absorption on larval survival of cultured hake, and can be used as an early indication of the quality of the batch.     

Inhibition of sexual maturation in tank reared haddock (Melanogrammus aeglefinus) through the use of constant light photoperiods

The haddock (Melanogrammus aeglefinus) is believed to be a potential candidate for aquaculture in the Atlantic coastal countries including the UK, Norway and Canada. However, under culture conditions, haddock will sexually mature prior to the attainment of a suitable harvest weight. Therefore, a long term tank based experiment was performed where three populations of haddock (hatched spring 2002, approximately 150 fish per population) were exposed to either a simulated natural photoperiod (SNP) or SNP until January or July 2003 and thereafter continuous illumination (Jan LL or July LL respectively) with individual growth rate and maturation status being recorded until July 2004 (27 months post-hatch). While the SNP treated population matured in the spring of 2004 (two years post-hatch) with 88% of the population being observed to release gametes, no mature individuals were observed in either of the LL treated populations and furthermore there was no evidence of gonadal development or elevation in sex steroids (testosterone, estradiol-17β or calcium). The application of LL appeared to directly stimulate the growth rate of haddock (e.g. 14 to 27% increase in weight thermal growth coefficient) and improve food conversion rates. At the end of the trial there was an approximate 50% increase in wet weight (SNP: 647 ± 53 g, Jan LL: 982 ± 34 g and July LL 985 ± 33 g), however no significant difference in weight in relation to the length of LL exposure was observed. Plasma melatonin analysis revealed that the natural diel rhythm had been inhibited in both LL treated populations. Along with highlighting the similarity in reproductive entrainment between haddock and other gadoids, these results demonstrate how photoperiod manipulation could be used as a management strategy to improve growth performance in farmed haddock stocks.