Biophysical modelling of larval drift, growth and survival for the prediction of anchovy (Engraulis encrasicolus) recruitment in the Bay of Biscay (NE Atlantic)

Fish recruitment is the result of the integration of small‐scale processes affecting larval survival over a season and large oceanic areas. A hydrodynamic model was used to explore and model these physical–biological interaction mechanisms and then to perform the integration from individual to population scales in order to provide recruitment predictions for fisheries management. This method was applied to the case of anchovy (Engraulis encrasicolus) in the Bay of Biscay (NE Atlantic). The main data available to investigate survival mechanisms were past growth (otolith) records of larvae and juveniles sampled at sea. The drift history of these individuals was reconstructed by a backtracking procedure using hydrodynamic simulations. The relationships between (real) growth variation and variations in physical parameters (estimated by hydrodynamic simulations) were explored along the individual trajectories obtained. These relationships were then used to build and adjust individual‐based growth and survival models. Thousands of virtual buoys were released in the hydrodynamic model in order to reproduce the space–time spawning dynamics. Along the buoy trajectories (representative of sub‐cohorts), the biophysical model was run to simulate growth and survival as a function of the environment encountered. The survival rate after 3 months of drift was estimated for each sub‐cohort. The sum of all these survival rates over the season constituted an annual recruitment index. This index was validated over a series of recruitment estimations. The modelling choices, model results and the potential use of the recruitment index for fisheries management are discussed.     

Relationships between anchovy (Engraulis encrasicolus) recruitment and environment in the Bay of Biscay (1967–1996)

Various hypotheses have been put forward to explain the mechanisms leading to recruitment variability in fish populations. These have been based on either physical (wind stress, upwelling) or biological (food and predation) processes. In the Bay of Biscay, the hypothesis of a physical influence on anchovy recruitment has been confirmed. Oceanographic conditions in the Bay of Biscay in the spring and summer, influenced by north-easterly winds of medium and low intensity, seem to induce good levels of recruitment to the anchovy ( Engraulis encrasicolus ) population. An index of upwelling was significantly correlated with annual recruitment of anchovy ( P  < 0.0001) for the period 1967–1996. This physical parameter explains about 59% of the variability in the recruitment of the Bay of Biscay anchovy. Two factors affecting productivity in the Bay of Biscay may be linked to north-easterly winds, namely weak upwelling and an extension of the area influenced by river outflows. Both of these factors, together with low turbulence and stability, may act to enhance survival of the early life-history stages of anchovy by increasing food availability. The potential use of this upwelling index to forecast the recruitment of the Bay of Biscay anchovy offers possibilities for improving the management of this population.     

Climate,oceanography, and recruitment: the case of the Bay of Biscay anchovy (Engraulis encrasicolus)

The Bay of Biscay anchovy has experienced, since 2001, a succession of low recruitments, resulting in the collapse of the stock in 2005; this has led to successive closures of the fishery. This study investigates the possible impact of different controlling factors [North Atlantic Oscillation (NAO), East Atlantic (EA) pattern, turbulence, upwelling, and river flow] upon anchovy recruitment and fishery catches. Fifty‐five percent of the recruitment variability of this fishery can be explained by upwelling over the spawning area; this is related, in turn, to the EA pattern. The conceptual understanding of the system proposed for the Bay of Biscay anchovy suggests that negative EA periods are associated with northeasterly wind circulation, which produces weak upwelling over the continental shelf. This pattern results in hydrodynamic stability over the area, leading, probably, to adequate food availability. A positive EA (which extends onwards, from 1998) is associated with southwesterly winds and downwelling over the continental shelf; this leads, probably, to the dispersion of anchovy food and larvae, together with increasing mortality.     

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.     

The influence of mesoscale ocean processes on anchovy (Engraulis encrasicolus) recruitment in the Bay of Biscay estimated with a three-dimensional hydrodynamic mode

The relationship between anchovy ( Engraulis encrasicolus ) recruitment in the Bay of Biscay and environmental variables during their planktonic phase (March to July) was investigated from 1986 to 1997. Meteorological variables (wind and temperature) are forcing effects on the sea, but they are not thought to be processes that govern larval survival directly. Food-web dynamics are believed to be more closely linked to larval survival and are related to the physical vertical water column structure. Therefore, we used a three-dimensional (3D) hydrodynamic model to characterize three major physical mesoscale processes affecting vertical structure in south-east Biscay: stratification, upwelling and river plume extent. Indices were estimated from the model outputs to characterize and quantify the space/time evolution of these structures during the period March to July. A multiple linear regression analysis was then used to analyse hierarchy in the explanatory power of the physical indices. Coastal upwelling and shelf stratification breakdown indices were the most significant explanatory variables, with positive and negative effect on recruitment, respectively. A model with these two indices explains 75% of the recruitment variability of anchovy observed in the period 1987–96.     

Historical fluctuations in spawning location of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during 1967–73 and 2000–2004

The spatial extent of small pelagic fish spawning habitat is influenced by environmental factors and by the state of the adult population. In return, the configuration of spawning habitat affects recruitment and therefore the future structure of the adult population. Interannual changes in spatial patterns of spawning reflect variations in adult population structures and their environment. The present study describes the historical changes in the spatial distribution of spawning of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) in the Bay of Biscay during two periods: 1967–72 and 2000–2004. Using data from egg surveys conducted in spring, the spatial distributions of anchovy and sardine eggs are characterized by means of geostatistics. For each survey, a map of probability of egg presence is constructed. The maps are then compared to define (1) recurrent spawning areas, (2) occasional spawning areas and (3) unfavourable spawning areas during each period. Sardine spawning habitat is generally fragmented and appears spatially limited by the presence of cold bottom water. It is confined to coastal or shelf break refuge areas in years of restricted spawning extent. For anchovy, recurrent spawning sites are found in Gironde and Adour estuaries whilst spawning can extend further offshore in years of more intense spawning. For both species, the mean pattern of spawning has changed between 1967–72 and 2000–2004. Noticeably, the spatial distribution of anchovy eggs in spring has expanded northward. This trend possibly results from changes in environmental conditions during the last four decades.     

Growth and movement patterns of early juvenile European anchovy (Engraulis encrasicolus L.) in the Bay of Biscay based on otolith microstructure and chemistry

Various hypotheses have been put forward to explain the mechanisms in the Bay of Biscay that result in a good recruitment of European anchovy. Anchovy larvae from the spawning area in the Gironde River plume are advected towards off-shelf waters, where juveniles are commonly observed. Otolith microstructural and chemical analysis were combined to assess the importance of this off-shelf transport and to determine the relative contribution of these areas for anchovy survival. Chemical analysis of otoliths showed that anchovy juveniles in the Bay of Biscay can be divided into two groups: a group that drifts towards off-shelf waters early in their life and returns later, and a group that remains in the low salinity waters of the coastal area. The first group presents significantly faster growth rates (0.88 mm day⁻¹) than those remaining in the coastal waters (0.32 mm day⁻¹). This may be due to off-shelf waters being warmer in spring/summer, and to the fact that the lower food concentration is compensated for by higher prey visibility. Furthermore, the group of juveniles that drifted off the spawning area and had faster growth rates represents 99% of the juvenile population. These findings support the hypothesis that anchovy in the Bay of Biscay may use off-shelf waters as a spatio-temporal loophole, suggesting that transport off the shelf may be favourable for recruitment.     

From egg to juvenile in the Bay of Biscay: spatial patterns of anchovy (Engraulis encrasicolus) recruitment in a non‐upwelling region

In this study the spatial distribution of eggs, larvae and juveniles of European anchovy (Engraulis encrasicolus) was followed in 2004 and 2005 during three consecutive cruises each year in spring–summer to test what the limits of retention are in a non‐upwelling area. Eggs, small larvae and large juveniles were mainly distributed over the shelf, whereas large larvae and small juveniles were found mainly off the shelf. Although overall distributions were similar, the 2 yr differed in that there was more of a coastal distribution of individuals in 2004, whereas in 2005 more individuals were found off the shelf. There were no significant differences in the length–weight relationships for individuals found on and off the shelf or between years. The correspondence in circulation patterns and the lack of difference in the length–weight relationships suggest that a single population is present, larvae drifting off the shelf due to currents and returning as mobile juveniles. Quantile regression analysis of the long‐term recruitment index suggests that transport off the shelf may favour good recruitments. This would suggest that in non‐upwelling regions the retention area resulting in good recruitment may not be restricted to the shelf.     

Size,permeability and buoyancy of anchovy (Engraulis Encrasicolus) and sardine (Sardina Pilchardus) eggs in relation to their physical environment in the Bay of Biscay

The size and specific gravity of eggs of marine pelagic fish partly determine their dispersal and survival. Using an original dataset of anchovy and sardine eggs, sampled in spring over the last decade in the Bay of Biscay, we provide a parameterization of these properties on ambient water temperature and salinity. We used the density gradient column for measurement of egg specific gravity. The column was also filled with homogeneous water for sinking velocity experiments. For anchovy, these experiments confirm that the effect of egg permeability through the chorion could be neglected when modelling sinking, while it has to be considered for sardine, its perivitelline space representing 78.6% (±6.2%) of the total egg volume, as opposed to 5–10% for most teleosts species. We estimated a coefficient of permeability of the chorion of 0.0038 mm s^?1. However, permeability should not affect the measurement of sardine egg specific gravity in a gradient column, provided a minimum duration before reading is respected for equilibrium to be reached. In relation to their environment, we found that the egg specific gravity is largely determined by sea surface salinity for both species, whereas egg size is weakly but significantly impacted by temperature, for sardine only. On average, the estimated difference in specific gravity between egg and surface water is ?0.92 σ_T for anchovy and ?1.06 σ_T for sardine. The detailed parameterization of the relationship between eggs and water properties should prove useful, in particular to modellers dealing with the dispersal of fish early life stages.     

A Bayesian model for anchovy (Engraulis encrasicolus): the combined forcing of man and environment

Fishery collapses frequently result from combined pressures of the environment and man, which are difficult to discern because of the complexities involved and our limited knowledge. Models to resolve this complexity often become too sophisticated, with too many assumptions and, consequently, with little capacity to predict beyond calibration data. In this paper we implement a different procedure where the model is kept simple and uncertainty accounts for the equation imperfectness to reproduce ecological complexity. Human and environmental forcing on an anchovy ( Engraulis encrasicolus ) stock are simulated with only six parameters plus their error terms, and the uncertainty is computed with Bayesian methods. The simple structure is able to reproduce the major dynamical features of this species in the Gulf of Cádiz, including data on life stages and age structure that had no contact with the model. This is a distinct performance for a frugal approach working on a mid-trophic species and a positive instance where parsimony can simulate the interaction of man, fish and the environment, provided uncertainty is accounted for in the process.     

Effect of habitat conditions on reproduction of the European anchovy (Engraulis encrasicolus) in the Strait of Sicily

The aims of this study were to describe the reproductive cycle of the European anchovy (Engraulis encrasicolus) off the south coast of Sicily and determine whether intra‐ and inter‐annual reproductive trait variations, if any, are adaptive responses which maximize reproductive success under environmental fluctuations. Biological data were collected from purse seine and mid‐water pelagic trawl commercial catches landed in Sciacca (Sicily) over 6 yr (1997–2002) at fortnightly intervals, analysing a total of 84 581 individuals. No inter‐annual changes in length at first reproduction were observed, with a mean pooled value of 11.26 cm for both sexes being found. Spawning intensity, indicated by gonadosomatic index, condition factor and length–weight relationships, seem to be governed by food availability prior to spawning. Anchovy reproductive investment was limited by the area's low primary production. There was a synchrony between reproductive cycle and temperature. Water warming marks the onset of a period of high water stability in the area, and its later cooling marks the onset of a period with low water stability. The relationship between reproductive cycle and temperature is therefore probably a reproductive strategy having evolved to ensure that spawning takes place during the period of the year when water column stability is higher, favouring food concentration and egg and larval retention in the spawning areas.     

Spawning site selection by European anchovy (Engraulis encrasicolus) in relation to oceanographic conditions in the Strait of Sicily

European anchovy egg occurrence and density data from summer surveys (1998–2007) and oceanographic data were examined to study the mechanisms that control the spatial distribution of anchovy spawning habitat in the Strait of Sicily. Quotient analysis indicated habitat preference for temperature (18–19°C), bottom depth (50–100 m), water column stability (13–14 cycle h^?1), fluorescence (0.10–0.15 μg m^?3 Chl a), salinity (37.5–37.6 PSU), current speed (0.20–0.25 m s^?1) and density (26.7–26.8 kg m^?3, σ_t). Canonical discriminant analysis identified temperature, column stability and fluorescence as major drivers of anchovy spawning habitat. Three of the 4 years which had lower egg abundance were warmer years, with low values of primary productivity. A geostrophic current flowing through the Strait (the Atlantic Ionic Stream, AIS) was confirmed as the main source of environmental variability in structuring the anchovy spawning ground by its influence on both the oceanography and distribution of anchovy eggs. This 10‐yr data series demonstrates recurrent but also variable patterns of oceanographic flows and egg distribution. A lack of freshwater flow in this area appears to depress productivity in the region, but certain and variable combinations of environmental conditions can elevate production in some sub‐areas in most years or other sub‐areas in fewer years. These temporal and spatial patterns are consistent with an ocean triad theory postulating that processes of oceanographic enrichment, concentration, and retention may help predict fishery yields.     

Simulation of the impact of climate change on the egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island,the western coast of Japan

In order to investigate the impact of climate change on egg and larval transport of Japanese anchovy (Engraulis japonicus) off Kyushu Island western Japan, we conducted particle‐tracking simulations on transport success/failure to fishing grounds from 1960 to 2007. The modeled transport success since the mid‐1990s increased and decreased in the offshore and coastal zones, respectively, compared with the 1960s and 1970s. The estimated northward shift of the spawning ground and weakened Tsushima Warm Current contributed to increase in modeled transport success to the offshore zone. Conversely, the weakening trend of the modeled onshore current in the Goto‐Nada Sea combined with the northward shift of the spawning ground resulted in unsuccessful larval transport. These results suggest that fluctuations in juvenile and subadult anchovy catches in this area may be attributable to changes in the physical environment. The present study showed that changes in transport success induced by oceanographic fluctuations related to climate change, have the potential to affect anchovy recruitment off the western coast of Japan.     

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.     

山东半岛南部癆鱼产卵场癆鱼仔、稚鱼摄食的研究

研究了山东半岛南部Ti鱼产卵场体长2．5－17．9mm的Ti鱼仔、稚鱼的摄食。Ti鱼仔、稚鱼主要饵料为原生动物的鼎形虫、单角铠甲虫、三角铠甲虫。小型桡足类的卵、桡足类的无节幼体和桡足幼体。桡足类种类为墨氏胸刺水蚤、小拟哲水蚤、克氏纺锤水蚤、双刺纺锤水蚤和拟长腹剑蚤等小型类桡足类。各种饵料所占的个数的百分比为：鼎形虫2．2％，单角铠甲虫7．6％，三角铠甲虫3．3％，桡足类的卵42．2％，无节幼体30．4％，桡足幼体14．1％，Ti鱼仔、稚鱼的摄食率为16．2％，摄食活动主要在白天，在水的中下层进行。由于摄食率低，肠道饱满度差，饥锇可能是引起Ti鱼仔、稚鱼死亡的主要原因。     

Transport and environmental temperature variability of eggs and larvae of the Japanese anchovy (Engraulis japonicus) and Japanese sardine (Sardinops melanostictus) in the western North Pacific estimated via numerical particle-tracking experiments

Numerical particle-tracking experiments were performed to investigate the transport and variability in environmental temperature experienced by eggs and larvae of Pacific stocks of the Japanese anchovy ( Engraulis japonicus ) and Japanese sardine ( Sardinops melanostictus ) using high-resolution outputs of the Ocean General Circulation Model for the Earth Simulator (OFES) and the observed distributions of eggs collected from 1978 to 2004. The modeled anchovy individuals tend to be trapped in coastal waters or transported to the Kuroshio–Oyashio transition region. In contrast, a large proportion of the sardines are transported to the Kuroshio Extension. The egg density-weighted mean environmental temperature until day 30 of the experiment was 20–24°C for the anchovy and 17–20°C for the sardine, which can be explained by spawning areas and seasons, and interannual oceanic variability. Regression analyses revealed that the contribution of environmental temperature to the logarithm of recruitment per spawning (expected to have a negative relationship with the mean mortality coefficient) was significant for both the anchovy and sardine, especially until day 30, which can be regarded as the initial stages of their life cycles. The relationship was quadratic for the anchovy, with an optimal temperature of 21–22°C, and linear for the sardine, with a negative coefficient. Differences in habitat areas and temperature responses between the sardine and anchovy are suggested to be important factors in controlling the dramatic out-of-phase fluctuations of these species.     

Interannual differences in growth and hatch‐date distributions of early juvenile European anchovy in the Bay of Biscay: implications for recruitment

In order to understand better the recruitment variability in European anchovy in the Bay of Biscay, it is important to investigate the processes that affect survival during the early life stages. Anchovy juvenile growth trajectories and hatch‐date distributions were inferred over a 3‐year period based on otolith microstructure analysis. Otolith growth trajectories showed a characteristic shape depending on their hatch‐date timing. Earlier‐born juveniles had notably broader maximum increments than later born conspecifics, resulting in higher growth rates. This observation suggests that early hatching would be beneficial for larval and juvenile growth, and, therefore, survival. The estimated juvenile hatch‐date distributions were relatively narrow compared with the extended anchovy spawning season (March–August) in the Bay of Biscay and indicated that only individuals originated mainly from the summer months (June–August) survived until autumn. Hatch‐date distributions were markedly different among years and seemed to influence the interannual recruitment strength of anchovy. We conclude that years characterized by juvenile survivors originating from the peak spawning period (May and June) would lead to considerable recruitment success. Downwelling events during the peak spawning period seem to affect larval survival. Furthermore, size‐dependent overwinter mortality would be an additional process that regulates recruitment strength in the anchovy population in the Bay of Biscay.