Use of satellite data for modelling food availability and survival of marine fish larvae

A modelling scheme is described which uses satellite retrieved sea-surface temperature and chlorophyll-a to derive monthly zooplankton biomass estimates in the eastern North Atlantic; this forms part of a bio-physical model of inter-annual variations in the growth and survival of larvae and post-larvae of mackerel (Scomber scombrus). The temperature and chlorophyll data are incorporated first to model copepod (Calanus) egg production rates. Egg production is then converted to available food using distribution data from the Continuous Plankton Recorder (CPR) Survey, observed population biomass per unit daily egg production and the proportion of the larval mackerel diet comprising Calanus. Results are validated in comparison with field observations of zooplankton biomass. The principal benefit of the modelling scheme is the ability to use the combination of broad scale coverage and fine scale temporal and spatial variability of satellite data as driving forces in the model; weaknesses are the simplicity of the egg production model and the broad-scale generalizations assumed in the raising factors to convert egg production to biomass.

Wind mixing, food availability and mortality of anchovy larvae Engraulis encrasicolus in the northern Adriatic Sea

A study was carried out in June/July 1996 in the River Po outflow in the northern Adriatic to investigate spawning of anchovy Engraulis encrasicolus and survival of larvae in relation to food availability and wind mixing. Hydrographic- and bongo net sampling was carried out on 2 grid surveys; one after a period of low winds and settled weather, and the other after an intervening period of strong winds, which resulted in a decrease in water column stratification. The spawning areas of anchovy and the larval distributions were associated with the river outflow plume (most clearly on the second survey grid, after the period of higher winds). Potential food particles for anchovy larvae, primarily copepod nauplii and copepodite stages, were also concentrated in the area influenced by the river outflow. Although there was a nearly 50% reduction in the mean water column abundance of potential food particles between the 2 survey grids, mostly due to a decline in abundance outside the immediate river plume area, there was no significant change in mortality of anchovy larvae between the 2 grids; the exponential decline in numbers of eggs and larvae to 10 mm in length being equivalent to overall mortality rates of 43.2%/d on the first survey and 44.7%/d on the second. The resilience of larval survival under potentially less favourable feeding conditions, following the period of wind mixing, was ascribed, in part, to the maintenance of local water column stratification by the superficial low salinity input from the River Po. This stratification in the immediate outflow area was associated with the presence of concentrated layers of potential food particles (typically >50 particles/L and 1.5 to 2.8 times the mean water column abundance) in the upper 10 m of the water column, coincident with peak numbers of anchovy larvae. However, since there was no evidence for lower larval survival in areas, less influenced by the immediate river outflow plume, a simple direct relationship between enhanced water column stability, improved feeding conditions and larval survival was not supported.

Distribution and abundance of sardine (Sardina pilchardus) eggs in the English Channel from Continuous Plankton Recorder sampling

Continuous Plankton Recorder (CPR) samples from the English Channel and adjacent Celtic shelf, taken over the period 1958-1980, were analysed for sardine (Sardina pilchardus) eggs. Results showed the progression of sardine spawning along the English Channel from west to east from March to August and a return from east to west from September to November. This corresponds with the two seasonal peaks of sardine egg abundance in the western Channel: the main summer peak being in May/June, with a smaller autumn peak in October/November. Long-term changes in sardine egg abundance in CPR samples showed a decline in summer spawning from the late 1960s, but no clear trend in autumn-spawned egg abundance. Similar patterns were observed in the numbers of sardine eggs sampled by conventional plankton net tows at the time-series Station L5 off Plymouth. This supports the use of the longer time-series of sardine egg data at L5 as being representative of a wider area and emphasizes the importance in continuation of the L5 time-series.

An individual-based model of the early life history of mackerel (Scomber scombrus) in the eastern North Atlantic, simulating transport, growth and mortality

The main purpose of this paper is to provide the core description of the modelling exercise within the Shelf Edge Advection Mortality And Recruitment (SEAMAR) programme. An individual-based model (IBM) was developed for the prediction of year-to-year survival of the early life-history stages of mackerel (Scomber scombrus) in the eastern North Atlantic. The IBM is one of two components of the model system. The first component is a circulation model to provide physical input data for the IBM. The circulation model is a geographical variant of the HAMburg Shelf Ocean Model (HAMSOM). The second component is the IBM, which is an i-space configuration model in which large numbers of individuals are followed as discrete entities to simulate the transport, growth and mortality of mackerel eggs, larvae and post-larvae. Larval and post-larval growth is modelled as a function of length, temperature and food distribution; mortality is modelled as a function of length and absolute growth rate. Each particle is considered as a super-individual representing 10 super(6) eggs at the outset of the simulation, and then declining according to the mortality function. Simulations were carried out for the years 1998-2000. Results showed concentrations of particles at Porcupine Bank and the adjacent Irish shelf, along the Celtic Sea shelf-edge, and in the southern Bay of Biscay. High survival was observed only at Porcupine and the adjacent shelf areas, and, more patchily, around the coastal margin of Biscay. The low survival along the shelf-edge of the Celtic Sea was due to the consistently low estimates of food availability in that area.

Simulation of mackerel (Scomber scombrus) recruitment with an individual-based model and comparison with field data

An individual-based model (IBM) for the simulation of year-to-year survival during the early life-history stages of the north-east Atlantic stock of mackerel (Scomber scombrus) was developed within the EU funded Shelf-Edge Advection, Mortality and Recruitment (SEAMAR) programme. The IBM included transport, growth and survival and was used to track the passive movement of mackerel eggs, larvae and post-larvae and determine their distribution and abundance after approximately 2 months of drift. One of the main outputs from the IBM, namely distributions and numbers of surviving post-larvae, are compared with field data as recruit (age-0/age-1 juveniles) distribution and abundance for the years 1998, 1999 and 2000. The juvenile distributions show more inter-annual and spatial variability than the modelled distributions of survivors; this may be due to the restriction of using the same initial egg distribution for all 3 yr of simulation. The IBM simulations indicate two main recruitment areas for the north-east Atlantic stock of mackerel, these being Porcupine Bank and the south-eastern Bay of Biscay. These areas correspond to areas of high juvenile catches, although the juveniles generally have a more widespread distribution than the model simulations. The best agreement between modelled data and field data for distribution (juveniles and model survivors) is for the year 1998. The juvenile catches in different representative nursery areas are totalled to give a field abundance index (FAI). This index is compared with a model survivor index (MSI) which is calculated from the total of survivors for the whole spawning season. The MSI compares favourably with the FAI for 1998 and 1999 but not for 2000; in this year, juvenile catches dropped sharply compared with the previous years but there was no equivalent drop in modelled survivors.

Modelling the vertical distribution of eggs of anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus)

Measurements were made of the density and settling velocity of eggs of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus), using a density-gradient column. These results were related to observed vertical distributions of eggs obtained from stratified vertical distribution sampling in the Bay of Biscay. Eggs of both species had slightly positive buoyancy in local seawater throughout most of their development until near hatching, when there was a marked increase in density and they became negatively buoyant. The settling velocity of anchovy eggs, which are shaped as prolate ellipsoids, was close to predictions for spherical particles of equivalent volume. An improved model was developed for prediction of the settling velocity of sardine eggs, which are spherical with a relatively large perivitelline volume; this incorporated permeability of the chorion and adjustment of the density of the perivitelline fluid to ambient seawater. Eggs of both species were located mostly in the top 20 m of the water column, in increasing abundance towards the surface. A sub-surface peak of egg abundance was sometimes observed at the pycnocline, particularly where this was pronounced and associated with a low-salinity surface layer. There was a progressive deepening of the depth distributions for successive stages of egg development. Results from this study can be applied for improved plankton sampling of sardine and anchovy eggs and in modelling studies of their vertical distribution.

Buoyancy measurements and vertical distribution of eggs of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus)

Measurements were made of the density and settling velocity of eggs of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus), using a density-gradient column. These results were related to observed vertical distributions of eggs obtained from stratified vertical distribution sampling in the Bay of Biscay. Eggs of both species had slightly positive buoyancy in local seawater throughout most of their development until near hatching, when there was a marked increase in density and they became negatively buoyant. The settling velocity of anchovy eggs, which are shaped as prolate ellipsoids, was close to predictions for spherical particles of equivalent volume. An improved model was developed for prediction of the settling velocity of sardine eggs, which are spherical with a relatively large perivitelline volume; this incorporated permeability of the chorion and adjustment of the density of the perivitelline fluid to ambient seawater. Eggs of both species were located mostly in the top 20 m of the water column, in increasing abundance towards the surface. A sub-surface peak of egg abundance was sometimes observed at the pycnocline, particularly where this was pronounced and associated with a low-salinity surface layer. There was a progressive deepening of the depth distributions for successive stages of egg development. Results from this study can be applied for improved plankton sampling of sardine and anchovy eggs and in modelling studies of their vertical distribution.

Spawning season and temperature relationships for sardine (Sardina pilchardus) in the eastern North Atlantic

Spawning temperature preferences for sardine (Sardina pilchardus) in the eastern North Atlantic were determined from field data. These were compared with climatological temperature cycles (1986-2002) derived from satellite data by geographical region, to predict spawning seasons. Optimum spawning temperatures were determined as 14.0-15.0oC from the English Channel to Portugal and 16.0–18.0oC for all north-west African regions. Spawning seasons were closely related to the general latitudinal trend of the annual temperature cycle, with modification by upwelling in the western Iberian and north-west African regions. Some differences between temperature-based spawning season predictions and field observations were related to variations in seasonal plankton production. Correlations in the annual time-series of favourable spawning temperatures suggested relatively strong linkages between the southern areas from Portugal to Senegal. There was no consistent relationship between annual variations in duration of temperature-predicted spawning seasons and observed field abundance of eggs.

Genetic structure of whiting (Merlangius merlangus) in the north east Atlantic and adjacent waters

Accurate identification of stock boundaries is essential for efficient fisheries management, hence the present study focused on the genetic structure of whiting. To this aim, 488 individuals collected from the southern Bay of Biscay to the southern Norwegian coast were genotyped using seven microsatellites. A low level of genetic structuring was detected in Atlantic waters since only the Bay of Biscay differentiated from more northern samples. The lack of genetic structure along the western margin of the British Isles is consistent with a high level of passive transport of pelagic eggs and larvae due to the combined influence of the North Atlantic Current and the Shelf Edge Current. High levels of dispersal could also occur between the western British Isles and the North Sea through both the branching of the North Atlantic Current into the northern North Sea and from the residual current flowing from the English Channel to the Southern Bight. In contrast, a significant genetic structure was identified within the North Sea, and this may be associated with the complex oceanography of this basin and retention systems reducing larval dispersal. In addition, considering also genetic, phenotypic and tag-recapture data collected on whiting, a learned homing behaviour of adults toward spawning areas may be hypothesised.

An overview of the literature concerning the oceanography of the eastern North Atlantic region

An overview of the main oceanographic features of the eastern North Atlantic boundary, with emphasis toward the upper layers, is presented. The principal features discussed are: water mass boundaries; forcing by wind, density and tides; topographic features and effects; fronts; upwelling and downwelling; poleward flows; coastal currents; eddies. The occurrence and spatial and seasonal variability of these features is described in five regional sections: Celtic Sea and western English Channel; Bay of Biscay; western Iberia; Gulf of Cadiz; northwest Africa. This paper is intended to provide a base of physical oceanographic knowledge in support of research in fisheries, biological and chemical oceanography, and marine biology.

Feeding of anchovy Engraulis encrasicolus larvae in the northwestern Adriatic Sea in response to changing hydrobiological conditions

Results from depth integrated and vertically stratified plankton sampling in the northwestern Adriatic Sea were used for comparison of gut contents of larvae of European anchovy Engraulis encrasicolus with composition and concentration of potential prey in the plankton. Sampling was carried out over a grid of stations both before and after a period of increased wind mixing to investigate changes in food availability and larval feeding success. All larvae had empty guts soon after dusk, indicating daytime feeding and rapid gut clearance. With increasing larval length there was a greater percentage of specimens with empty guts, despite suitable food being available in the plankton for these larger larvae; this suggests differential gut evacuation during sampling-possibly related to the degree of gut development. Larval diet was principally the various developmental stages of copepods, especially calanoid and cyclopoid nauplii, which were preferentially selected by larvae, whereas selection was against harpacticoid nauplii. Lamellibranch larvae and Peridinium were generally abundant in the plankton, but were only present in the gut contents in any number when the preferred dietary organisms were present in the plankton at low concentrations. The number of food organisms in the gut contents increased with concentration of the preferred food organisms in the plankton up to a limit of similar to 50 organisms/l. Within the upper 18 m of the water column, there was a reduction in the proportion of larvae with food in their guts with increasing depth, irrespective of the vertical profile of food concentration. Following a period of wind mixing the composition of the plankton changed. This was reflected in the diet of anchovy larvae, which altered in parallel. There was also an overall 41% decrease in concentration of the preferred food particles of larvae in the plankton following the period of wind mixing, but larvae were still able to maintain their food intake. These results show that anchovy larvae can successfully adapt their diet to a changing prey field and suggest that in the conditions observed in the northern Adriatic, quite radical changes in the feeding environment were probably insufficient to affect overall larval mortality.

Digestion of natural food by larval and post-larval turbot Scophthalmus maximus

The digestion of natural, mainly crustacean zooplankton, by different age groups of turbot Scophthalmus maximus larvae was evaluated by comparisons of visual appearance, dry weight and carbon and nitrogen content of fresh food organisms with material recovered from faeces. Visually, the degree of digestion of food particles ranged from no discernible change of lamellibranch larvae, copepod eggs, intact copepod faecal pellets and some phytoplankton species, to varying degrees of removal of body constituents in copepods, cladocerans and decapod zoea. For crustaceans, the proportion of body constituents removed was related to the size and construction of their apparently indigestible exoskeleton. Uppon defaecation larger organisms showed the greatest percentage loss in dry weight and carbon. A high percentage of nitrogen was extracted from all organisms.