Linking growth to environmental histories in central Baltic young-of-the-year sprat, Sprattus sprattus: an approach based on otolith microstructure analysis and hydrodynamic modelling

Otolith microstructure analysis and hydrodynamic modelling were combined to study growth patterns in young‐of‐the‐year (YoY) sprat, Sprattus sprattus, which were sampled in October 2002 in the central Baltic Sea. The observed ‘window of survival’, approximated by the distribution of back‐calculated days of first feeding (DFF), was narrow compared to the extended spawning season of sprat in the Baltic Sea (mean± SD = 22 June ± 14.1 days) and indicated that only individuals born in summer survived until October 2002. Within the group of survivors, individuals born later in the season exhibited faster larval, but more rapidly decreasing juvenile growth rates than earlier born conspecifics. Back‐calculated larval growth rates of survivors (0.48–0.69 mm day^?1) were notably higher than those previously reported for average larval sprat populations, suggesting that the YoY population was predominantly comprised of individuals which grew faster during the larval stage. Daily mean temperatures, experienced across the entire YoY population, were derived from Lagrangian particle simulations and correlated with (1) detrended otolith growth and (2) back‐calculated, daily somatic growth rates of survivors. The results showed that abrupt changes in ambient temperature can be detected in the seasonal pattern of otolith growth, and that higher temperatures led to significantly faster growth throughout the entire age range of YoY sprat.     

Predicting the dispersion of sprat larvae (Sprattus sprattus (L.)) in the German Bight

Sprat (Sprattus sprattus (L.)) larval dispersal in the German Bight is predicted with a model system consisting of two three-dimensional circulation models and a three-dimensional transport model. Horizontal distribution data for specific length classes of larvae, obtained from field surveys carried out in the German Bight during summer 1991, are used as initial conditions for the starting points of the tracers in the transport model. Sprat larval distributions are predicted and are compared with distributions of the appropriate length class of larvae observed during a subsequent cruise about 3 weeks later. The predicted larval (tracer) distributions compare favourably to observed larvae distributions in the inner model areas but not in the areas close to the coast. Factors contributing to inaccurate predictions are uncertainties in horizontal and vertical larval distributions as well as the model grid resolution, which is too coarse in coastal areas.     

An example of meso‐scale hydrographic features in the central Baltic Sea and their influence on the distribution and vertical migration of sprat,Sprattus sprattus balticus (Schn.)

A wind‐driven meso‐scale pattern of temperature, salinity and oxygen was found along a transect in the northern Bornholm Basin (southern Baltic Sea). Strong winds caused currents along this transect, which shifted cold intermediate water (minimum: 3.6°C) towards the south. The transect was surveyed with a towed CTD‐system and hydroacoustics in parallel to investigate the distribution of sprat, Sprattus sprattus balticus (Schn.) in relation to the observed meso‐scale pattern. In those parts of the transect where the cold intermediate water was observed, sprat were restricted to water layers below the halocline. In other parts of the transect, sprat moved into higher water layers and occupied a wider depth range. The important factor was temperature, which set an upper limit to the vertical sprat distribution. The development of hydrography, as measured in the field, was evaluated with a hydrodynamic model.     

Temporal and spatial variability in growth and condition of dab (Limanda limanda) and sprat (Sprattus sprattus) larvae in the Irish Sea

Variability in the high mortality rate during early life stages is considered to be one of the principal determinants of year‐class variability in fish stocks. The influence of water column stability on the spatial distribution of fish larvae and their prey is widely acknowledged. Water column stability may also impact growth through the early life history of fishes, and consequently alter the probability of survival to maturity by limiting susceptibility to predation and starvation. As a test of this concept, the variability in condition and growth of dab (Limanda limanda) and sprat (Sprattus sprattus) larvae was investigated in relation to seasonal stratification of the water column in the north‐western Irish Sea. RNA/DNA ratios and otolith microincrement analysis were used to estimate nutritional status and recent growth rates of larvae captured on four cruises in May and June of 1998 and 1999. Dab and sprat larvae were less abundant in 1999 and were in poorer condition with lower growth rates than in 1998. Dab larvae of <13 mm also exhibited spatial variability with higher RNA/DNA ratios at the seasonal tidal‐mixing front compared with stratified and mixed water masses. However, the growth and nutritional status of sprat larvae was uncorrelated to water column stability, meaning the more favourable feeding conditions generally associated with the stratified pool and tidal‐mixing front in the Irish Sea were not reflected in the growth and condition of these larvae. This suggests that the link between stability, production and larval growth is more complicated than inferred by some previous studies. The existence of spatio‐temporal heterogeneity in the growth and condition of these larvae has implications for larval survival and the recruitment success of these species in the Irish Sea.     

The influence of biotic and abiotic factors on the growth of sprat (Sprattus sprattus) in the Baltic Sea

We analysed spatial and temporal patterns of condition factor (CF) of sprat (Sprattus sprattus) between 1986 and 2000 in different areas of the Baltic proper. Results showed a moderate increase in CF between 1986 and 1989. Afterwards, an abrupt decrease in CF, between 22% and 29%, occurred in all the areas of the Baltic proper. However, from 1998 and onwards, CF increased compared to the previous years. Weight-at-age and CF of Baltic herring showed a similar pattern in the same period. Several factors affecting variability of sprat CF were evaluated. Data showed that sprat CF was density-dependent and possibly related to the pelagic fish abundance and individual food intake. Temperature did not influence sprat CF while salinity seemed to affect variability of sprat CF in the northern Baltic proper. Changes in salinity levels may shape pelagic fish growth rates both indirectly changing the zooplankton community structure and abundance and/or directly via fish physiology and metabolisms.     

Variability of larval Baltic sprat (Sprattus sprattus L.) otolith growth: a modeling approach combining spatially and temporally resolved biotic and abiotic environmental key variables

Plankton sampling was conducted in the Baltic to obtain sprat larvae. Their individual drift patterns were back‐calculated using a hydrodynamic model. The modelled positions along the individual drift trajectories were subsequently used to provide insight into the environmental conditions experienced by the larvae. Autocorrelation analysis revealed that successive otolith increment widths of individual larvae were not independent. Otolith increment width was then modelled using two different generalized additive model (GAM) analyses (with and without autocorrelation), using environmental variables determined for each modelled individual larval position as explanatory variables. The results indicate that otolith growth was not only influenced by the density of potential prey but was controlled by a number of simultaneously acting environmental factors. The final model, not considering autocorrelation, explained more than 80% of the variance of otolith growth, with larval age as a factor variable showing the strongest significant impact on otolith growth. Otolith growth was further explained by statistically significant ambient environmental factors such as temperature, bottom depth, prey density and turbulence. The GAM analysis, taking autocorrelation into account, explained almost 98% of the variability, with the previous otolith increment showing the strongest significant effect. Larval age as well as ambient temperature and prey abundance also had a significant effect. An alternative approach applied individual‐based model (IBM) simulations on larval drift, feeding, growth and survival starting as exogenously feeding larvae at the back‐calculated positions. The IBM results revealed optimal growth conditions for more than 97% of the larvae, with a tendency for our IBM to slightly overestimate larval growth.     

Coupling ecosystem and individual‐based models to simulate the influence of environmental variability on potential growth and survival of larval sprat (Sprattus sprattus L.) in the North Sea

To investigate the impact of changing environmental conditions in the North Sea on the distribution and survival of early life stages of a marine fish species, we employed a suite of coupled model components: (i) an Eulerian coupled hydrodynamic/ecosystem (Nutrients, Phyto‐, Zooplankton, Detritus) model to provide both 3‐D fields of hydrographical properties, and spatially and temporally variable prey fields; (ii) a Lagrangian transport model to simulate temporal changes in cohort distribution; and (iii) an individual‐based model (IBM) to depict foraging, growth and survival of fish early life stages. In this application, the IBM was parameterized for sprat (Sprattus sprattus L.) and included non‐feeding (egg and yolk‐sac larval) stages as well as foraging and growth subroutines for feeding (post‐yolk sac) larvae. Sensitivity analyses indicated that the angle of visual acuity, assimilation efficiency and the maximum food consumption rate were the most critical intrinsic model parameters. As an example, we applied this model system for 1990 in the North Sea. Results included not only information concerning the interplay of temperature and prey availability on larval fish survival and growth but also information on mechanisms underlying larval fish aggregation within frontal zones. The good agreement between modelled and in situ estimates of sprat distribution and growth rates in the German Bight suggested that interconnecting these different models provided an expedient tool to scrutinize basic processes in fish population dynamics.     

Simulating the dispersion of vertically migrating sprat larvae (Sprattus sprattus (L.)) in the German Bight with a circulation and transport model system

A model system consisting of two three-dimensional circulation models and a three-dimensional transport model is used to simulate the dispersal of sprat larvae (Sprattus sprattus (L.)) in the German Bight. A Northwest European shelf sea model simulates the currents of the North Sea and provides sea surface elevation data for the open boundaries of the German Bight circulation model. The German Bight circulation model has a horizontal grid resolution of 2.75 km and a vertical resolution of 5 m except for the bottom layer. The driving forces of the circulation models are the M2-tide, six-hourly time-dependent wind stress and air pressure fields as well as monthly climatological density fields. The simulated currents serve as input to the transport model. The model area, grid size and the vertical resolution of the transport model are in accordance with the German Bight circulation model. The transport model simulates advection and diffusion using a Monte Carlo method. Furthermore it incorporates a simulation of active vertical movement of the larvae. The vertical migration pattern is dependent on the time of day and size of the larvae and is based on field data from the German Bight. Horizontal larvae distribution data, gained from two out of five field surveys carried out in the German Bight during spring and summer 1991, are used as initial values for the starting points of the tracers in the transport model. Simulated tracer distributions are compared with larval distributions observed during subsequent cruises about 3 weeks later. Discrepancies and similarities between tracer and larval distributions are discussed.     

Variations in the drift of larval cod (Gadus morhua L.) in the Baltic Sea: combining field observations and modelling

Coupled three-dimensional (3-D) physical oceanographic modelling and field sampling programmes were carried out in May 1988 and August 1991 to investigate the potential drift of larval cod ( Gadus morhua L.) in the Bornholm Basin of the Baltic Sea. The goals were to predict the transport of cod larvae, thus aiding the identification of physical processes influencing larval retention/dispersal. Numerical simulations were performed using a 3-D eddy-resolving baroclinic model based on the Bryan–Cox–Semtner code adapted for the Baltic Sea. Within the Bornholm Basin, the model was initialized with ground truth data of physical parameters obtained on the research cruises, and all simulations were forced with actual wind data. Outside the basin, generalized hydrographic features of the Baltic Sea were utilized by incorporation of simulated hydrographic fields from previous model runs typical for the time periods considered. Larval drift was simulated either by incorporation of passive drifters, or as the initial horizontal distribution of larvae implemented into the model. Drift model simulations of larval transport agreed relatively well with field observations. The influence of variations in the vertical distribution on a smaller scale, i.e. vertical deviations of ± 6 m from the observed mean centre of mass, on the drift was examined, revealing no significant differences in the drift of larvae depending on their vertical distribution. The different wind forcing during the investigated time periods was linked to a retention situation in May 1988 and to a dispersal situation in August 1991. Finally, observed spatial distribution patterns of 1-group cod based on Baltic Young Fish Surveys (BYFS) were compared with their predicted transport in the larval phase and examined with respect to recruitment.     

Oceanographic influences on the distribution of Baltic cod, Gadus morhua, during spawning in the Bornholm Basin of the Baltic Sea

The Baltic Sea is a stratified, semienclosed sea typified by a low-salinity surface layer and a deep saline layer of varying volume, salinity, temperature and oxygen concentration. The relationships between these oceanographic factors and the distribution of Baltic cod are presented, utilizing results from a survey carried out during the 1995 spawning period in the Bornholm Basin, at present the main spawning area of this stock. Cod distribution, abundance and population structure were estimated from hydroacoustic and trawl data and related to hydrographic parameters as well as to bottom depth. In the central basin, cod were aggregated in an intermediate layer about 15 m thick. This area of peak abundance was defined at its upper limit by the halocline and at the lower limit by oxygen content. The majority of individuals caught in the basin centre were in spawning or pre-spawning condition with a high proportion of males to females. On the basin slopes, aggregations of cod were found near the bottom. These individuals were mainly immature and maturing stages with an increasing proportion of females to males with size. Salinity and oxygen conditions were found to be the major factors influencing the vertical and horizontal distribution of adult cod. Abundance of immature cod was also positively related to decreasing bottom depths. The effect of temperature was minor. The observed size- and sex-dependent spawning aggregation patterns, in association with habitat volume and stock size, may influence cod catchability and thereby the assessment and exploitation patterns of this stock.     

Spatio‐temporal trends of sailfish,Istiophorus platypterus catch rates in relation to spawning ground and environmental factors in the equatorial and southwestern Atlantic Ocean

Spatial and temporal trends of sailfish catch rates in the southwestern and equatorial Atlantic Ocean in relation to environmental variables were investigated using generalized additive models and fishery‐dependent data. Two generalized additive models were fit: (i) ‘spatio‐temporal’, including only latitude, longitude, month, and year; and (ii) ‘oceanographic’, including sea surface temperature (SST), chlorophyll‐a concentration, wind velocity, bottom depth, and depth of mixed layer and year. The spatio‐temporal model explained more (average ~40%) of the variability in catch rates than the oceanographic model (average ~30%). Modeled catch rate predictions showed that sailfish tend to aggregate off the southeast coast of Brazil during the peak of the spawning season (November to February). Sailfish also seem to aggregate for feeding in two different areas, one located in the mid‐west Atlantic to the south of ~15°S and another area off the north coast of Brazil. The oceanographic model revealed that wind velocity and chlorophyll‐a concentration were the most important variables describing catch rate variability. The results presented herein may help to understand sailfish movements in the Atlantic Ocean and the relationship of these movements with environmental effects.     

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.     

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.     

Variation in the distribution of yellowfin sole Limanda aspera larvae in warm and cold years in the eastern Bering Sea

Multiyear periods of relatively cold temperatures (2007–2013) and warm temperatures (2001–2005 and 2014–2018) altered the eastern Bering Sea ecosystem, affecting ocean currents and wind patterns, plankton community, and spatial distribution of fishes. Yellowfin sole Limanda aspera larvae were collected from the inner domain (≤50 m depth) of the eastern Bering Sea among four warm years (2002, 2004, 2005, 2016), an average year (2006), and three cold years (2007, 2010, 2012). Spatial distribution and density of larvae among those years was analyzed using generalized additive models that included timing of sea-ice retreat, areal coverage of water ≤0°C, and water temperature as covariates. Analyses indicated a combination of temperature effects on the location and timing of spawning, and on egg and larval survival, may explain the variation in larval density and distribution among years. During warm years, higher density and wider spatial distribution of larvae may be due to earlier spawning, an expansion of the spawning area, and higher egg and larvae survival due to favorable temperatures. Larval distribution contracted shoreward, and density was lower during cold conditions and was likely due to fish spawning closer to shore to remain in preferred temperatures, later spawning, and increased mortality. Predicted drift trajectories from spawning areas showed that larvae would reach nursery grounds in most years. Years when the drift period was longer than the pelagic phase of the larvae occurred during both warm and cold conditions indicating that settlement outside of nursery areas could happen during either temperature condition.     

基于GAM的长江口鱼类资源时空分布及影响因素

根据2006至2017年长江口及其邻近海域鱼类资源调查,运用广义加性模型研究长江口鱼类资源密度与环境因子之间的关系,并对2017年鱼类资源密度的时空分布进行预测。结果显示,春夏秋冬四个季节最佳GAM偏差解释率分别为69.6%、55.9%、51.4%和47.4%,交叉验证回归线斜率的平均效应为0.62～0.88。盐度、水温和溶解氧是影响长江口鱼类资源密度的主要环境影响因子且在不同季节对鱼类资源密度有不同的影响机制。总体上,在春、夏、秋季,盐度与鱼类资源密度之间存在正向相关性;在夏、秋、冬季,水温对鱼类资源密度有显著影响,在秋季与鱼类资源密度之间存在正向相关性;在春、秋、冬季,溶解氧对鱼类资源密度有显著影响,在冬季与鱼类资源密度之间存在正向线性相关。研究表明,2017年夏季鱼类资源密度较高;在长江口南支的自然延伸水域存在鱼类资源密度的相对低值,在崇明岛向海自然延伸方向水域存在鱼类资源密度的相对高值。后续研究将对长江口鱼类资源进行不同生态类型区分,以期更加准确地掌握影响各生态类型鱼类时空分布的环境因素及其时空分布信息。     

Habitat suitability modelling for sardine juveniles (Sardina pilchardus) in the Mediterranean Sea

Identification of potential juvenile grounds of short‐lived species such as European sardine (Sardina pilchardus) in relation to the environment is a crucial issue for effective management. In the current work, habitat suitability modelling was applied to acoustic data derived from both the western and eastern part of the Mediterranean Sea. Early summer acoustic data of sardine juveniles were modelled using generalized additive models along with satellite environmental and bathymetry data. Selected models were used to construct maps that exhibit the probability of presence in the study areas, as well as throughout the entire Mediterranean basin, as a measure of habitat adequacy. Areas with high probability of supporting sardine juvenile presence persistently within the study period were identified throughout the Mediterranean Sea. Furthermore, within the study period, a positive relationship was found between suitable habitat extent and the changes in abundance of sardine juveniles in each study area.     

黄海大头鳕0龄幼体分布及其与环境因子的关系

大头鳕是广泛分布于太平洋北部沿岸海域的重要经济和生态种,研究其幼体的生物学特性和分布有助于了解大头鳕的种群动态。本实验根据2016年6月、8月、10月、12月在黄海进行的渔业资源和环境调查数据,研究了黄海大头鳕种群幼体的生长规律;并运用广义可加模型(GAM)方法,分析了该海域大头鳕幼体分布及其与环境因子的关系。结果显示,黄海大头鳕幼体为正异速增长,体质量增长较快,b值为3.316 1。黄海大头鳕0龄幼体主要在底层盐度31.7~33.3、底层水温6.6~12.1°C、深度35.8~87.2 m、底质为细粉砂或黏土质软泥底质的海域中生活。研究表明,大头鳕幼体分布的季节变化与黄海冷水团的季节演变过程具有同步性。对大头鳕幼体密度有显著影响的环境因子依次为底层水温、底质、底层盐度,而深度对大头鳕幼体密度的影响不显著。     

Food organisms and feeding habits of larval and juvenile Japanese flounder Paralichthys olivaceus at Ohama Beach in Hiuchi-Nada, the central Seto Inland Sea, Japan

ABSTRACT:   The abundance of food organisms and feeding habits of larval and juvenile Japanese flounder were examined during the period from May to August in 1999, 2000 and 2001 at the sandy Ohama beach, the central Seto Inland Sea. The food organisms collected with a sledge net consisted of 40 families from 18 orders, dominated by mysids, crangonids and gammarids. The mean densities of mysids, crangonid shrimp ( Crangon spp.), gammarids and fish were 2.74, 6.74, 2.91 and 0.15 individuals/m², respectively. The main prey of the flounder ( n  = 202; range of total length 9.80–75.95 mm) was mysids and small crangonid shrimp (<14 mm in body length). Prey fish availability was low, as the density of fish was low. The small crangonid shrimp was abundant, and the large crangonid shrimp, which could prey on larval flounder, was not abundant. The crangonid shrimp was important not as a predator for the flounder but as prey. The flounder preferred epifaunal mysids, Nipponomysis ornata and Anisomysis ijimai , to sand-burrowing mysids, Iiella oshimai , and avoided crangonid shrimp.     

Patterns in the spawning of cod (Gadus morhua L.), sole (Solea solea L.) and plaice (Pleuronectes platessa L.) in the Irish Sea as determined by generalized additive modelling

Eleven ichthyoplankton cruises were undertaken covering most of the Irish Sea during the period February to June, 1995. To identify spawning localities and investigate temporal trends in egg production, the data on stage 1 A egg distributions of cod ( Gadus morhua ), plaice ( Pleuronectes platessa ) and sole ( Solea solea ) have been modelled using generalized additive models (GAMs). A two-stage approach was adopted where presence/absence was firstly modelled as a binary process and a GAM surface subsequently fitted to egg production (conditional on presence). We demonstrate that this approach can be used to model egg production both in space and in time. The spawning sites for cod, plaice and sole in the Irish Sea were defined in terms of the probability of egg occurrence. For cod, we demonstrate that by integrating under predicted egg production surfaces, a cumulative production curve can be generated and used to define percentiles of production and thus delimit the extent of the spawning season. However, for plaice and sole, the surveys did not fully cover the spawning season and the limitations that this imposes on GAM modelling of these data are discussed. Comparison of the spawning sites in 1995 with historical data suggests that the locations of cod, plaice and sole egg production in the Irish Sea have probably remained relatively constant over the last 30 years.