Drifter observations in the summer time Bay of Biscay slope current

During the summer of 2012, 20 surface drifters drogued at 50 m depth were deployed on the continental slope to the north of the Bay of Biscay. Initially after release the drifters all crossed the slope, with 14 continuing equatorward, parallel to the slope following an absolute dynamic topography feature and 6 returning to the slope, in an eddy, visible in chlorophyll-a maps. Lagrangian statistics show an anisotropic flow field that becomes less tied to the absolute dynamic topography and increasingly dominated by diffusion and eddy processes. A weaker tie to the absolute dynamic topography allowed for total of 8 of the drifters crossed from the deep water onto the shelf, showing pathways for flow across the slope. A combination of drifter trajectories, absolute dynamic topography and chlorophyll-a concentration maps have been used to show that small anticyclonic eddies, tied to the complex slope topography provide a mechanism for on shelf transport. During the summer, the presence of these eddies can be seen in surface chlorophyll-a maps.

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.

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.

Variations in the distributions of the Centropages chierchiae and Temora stylifera (Copepoda: Calanoida) in the north-eastern Atlantic Ocean and western european shelf waters.

Centropages chierchiae and Temora stylifera occurred rarely in the Continuous Plankton Recorder (CPR) survey in the Bay of Biscay, Celtic Sea, and English Channel before 1988. By 2000 they were found frequently and in abundance. The seasonal cycles of abundance of these species differ, C. chierchiae occurring mainly in the summer while T. stylifera was found most frequently in late autumn or winter towards the northern limits of its distribution. The increase in abundance of both species is related to temperature. However, in the years when it was found in the samples, the frequency of occurrence of C. chierchiae was correlated positively with the strength of the shelf edge current and negatively with the North Atlantic Oscillation (NAO) while the reverse was true for T. stylifera.

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.

Particle export during a bloom of Emiliania huxleyi in the North-West European continental margin

Coccolithophores, the dominant pelagic calcifiers in the oceans, play a key role in the marine carbon cycle through calcification, primary production and carbon export, the main drivers of the biological CO2 pump. In May 2002 a cruise was conducted on the outer shelf of the North-West European continental margin, from the north Bay of Biscay to the Celtic Sea (47.0 degrees-50.5 degrees N, 5.0 degrees-11.0 degrees W), an area where massive blooms of Emiliania huxleyi are observed annually. Biogeochemical variables including primary production, calcification, partial pressure of CO2 (pCO(2)), chlorophyll-a (Chl-a), particle load, particulate organic and inorganic carbon (POC, PIC) and Th-234, were measured in surface waters to assess particle dynamic and carbon export in relation to the development of a coccolithophore bloom. We observed a marked northward decrease in Chl-a concentration and calcification rates: the bloom exhibited lower values and may be less well developed in the Goban Spur area. The export fluxes of POC and PIC from the top 80 m, determined using the ratios of POC and PIC to Th-234 of particles, ranged from 81 to 323 mg C m(-2) d(-1) and from 30 to 84 mg C m(-2) d(-1), respectively. The highest fluxes were observed in waters presenting a well-developed coccolithophore bloom, as shown by high reflectance of surface waters. This experiment confirms that the occurrence of coccolithophores promotes efficient export of organic and inorganic carbon on the North-West European margin.

Comparing copepod time-series in the north of Spain: Spatial autocorrelation of community composition

Four time-series of copepod species biomass in the north of Spain were contrasted to demonstrate spatial autocorrelation of local communities and their responses to short-term local and regional variability in oceanographic conditions. The series represented coastal and oceanic environments along a marked gradient of influence of seasonal upwelling from Galicia to the Mar Cantábrico (S Bay of Biscay), and each one included at least 10 years of continuous data collected at monthly frequency. Community composition (i.e. species number and diversity) was very consistent through the region, but local variations in the presence of new species and the relative proportions of common species allowed for the characterisation of the response to the environment at each site. Small-sized species were more frequent near the coast. A few species, however, captured the main patterns of variability in all series. Calanus helgolandicus and Acartia (mainly Acartia clausi) were generally the main contributors to total biomass, while other species as Paracalanus parvus and Clausocalanus spp. were important only at some locations. Most copepod indices were positively correlated with upwelling, either considering the whole community (biomass, species richness and diversity) or individual species, but only in the coastal series analysed since 1991. Copepods in the nearby ocean, however, showed negative correlations with upwelling in the period 1960–1986. The effects of upwelling may have been modulated by local factors, as showed by the increases in biomass, number of species and diversity in associations with increases in sea surface temperature in Galicia, while in the Mar Cantábrico only the warming-tolerant species increased and those typical of upwelling decreased. Density stratification of the water column was associated with decreases in total copepod biomass in Galicia, while it favoured the increase in species richness in the Mar Cantábrico. Nearly all significant responses of copepods to environmental variability were delayed by up to 5 months, showing the importance of considering time-lags in the analysis of temporal responses of zooplankton.

Long-term variability and environmental preferences of calycophoran siphonophores in the Bay of Villefranche (North-Western Mediterranean)

Long-term variability of the main calycophoran siphonophores was investigated between 1974 and 1999 in a coastal station in the north-western Mediterranean. The data were collected at weekly frequency using a macroplankton net (680 μm mesh size) adapted to quantitatively sample delicate gelatinous plankton. A 3-year collection (1967–1969) of siphonophores from offshore waters using the same methodology showed that the patterns of variability observed inshore were representative of siphonophores’ changes at a regional scale. The aims of the study were: (i) to investigate the patterns of variability that characterised the dominant calycophoran species and assemblages; (ii) to identify the environmental optima that were associated with a significant increase in the dominant siphonophore species and (iii) to verify the influence of hydroclimatic variability on long-term changes of siphonophores. Our results showed that during nearly 3 decades the standing stock of calycophoran siphonophores did not show any significant change, with the annual maximum usually recorded in spring as a result of high densities of the dominant species Lensia subtilis, Muggiaea kochi and Muggiaea atlantica. Nevertheless, major changes in community composition occurred within the calycophoran population. Since the middle 1980s, M. kochi, once the most dominant species, started to decrease allowing other species, the congeneric M. atlantica and Chelophyes appendiculata, to increasingly dominate in spring and summer–autumn, respectively. The comparison of environmental and biotic long-term trends suggests that the decrease of M. kochi was triggered by hydrological changes that occurred in the north-western Mediterranean under the forcing of large-scale climate oscillations. Salinity, water stratification and water temperature were the main hydroclimatic factors associated with a significant increase of siphonophores, different species showing different environmental preferences.

The influence of phytoplankton productivity, temperature and environmental stability on the control of copepod diversity in the North East Atlantic

The patterns of copepod species richness (S) and their relationship with phytoplankton productivity, temperature and environmental stability were investigated at climatological, seasonal and year-to-year time scales as well as scales along latitudinal and oceanic–neritic gradients using monthly time series of the Continuous Plankton Recorder (CPR) Survey collected in the North East Atlantic between 1958 and 2006. Time series analyses confirmed previously described geographic patterns. Equatorward and towards neritic environments, the climatological average of S increases and the variance explained by the seasonal cycle decreases. The bi-modal character of seasonality increases equatorward and the timing of the seasonal cycle takes place progressive earlier equatorward and towards neritic environments. In the long-term, the climatological average of S decreased significantly (p < 0.001) between 1958 and 2006 in the Bay of Biscay and North Iberian shelf at a rate of ca. 0.04 year−1, and increased at the same rate between 1991 and 2006 in the northernmost oceanic location. The climatological averages of S correlate positively with those of the index of seasonality of phytoplankton productivity (ratio between the minimum and maximum monthly values of surface chlorophyll) and sea surface temperature, and negatively with those of the proxy for environmental stability (monthly frequency of occurrence of daily averaged wind speed exceeding 10 m s−1). The seasonal cycles of S and phytoplankton productivity (surface chlorophyll as proxy) exhibit similar features in terms of shape, timing and explained variance, but the relationship between the climatological averages of both variables is non-significant. From year-to-year, the annual averages of S correlate negatively with those of phytoplankton productivity and positively with those of sea surface temperature along the latitudinal gradient, and negatively with those of environmental stability along the oceanic–neritic gradient. The annual anomalies of S (i.e. factoring out geographic variation) show a unimodal relationship with those of sea surface temperature and environmental stability, with S peaking at intermediate values of the anomalies of these variables. The results evidence the role of seasonality of phytoplankton productivity on the control of copepod species richness at seasonal and climatological scales, giving support to the species richness–productivity hypothesis. Although sea surface temperature (SST) is indeed a good predictor of richness along the latitudinal gradient, it is unable to predict the increase of richness form oceanic to neritic environments, thus lessening the generality of the species richness–energy hypothesis. Meteo-hydrographic disturbances (i.e. SST and wind speed anomalies as proxies), presumably through its role on mixed layer depth dynamics and turbulence and hence productivity, maximise local diversity when occurring at intermediate frequency and or intensity, thus providing support to the intermediate disturbance hypothesis on the control of copepod diversity.

Biomass changes and trophic amplification of plankton in a warmer ocean

Ocean warming can modify the ecophysiology and distribution of marine organisms, and relationships between species, with nonlinear interactions between ecosystem components potentially resulting in trophic amplification. Trophic amplification (or attenuation) describe the propagation of a hydroclimatic signal up the food web, causing magnification (or depression) of biomass values along one or more trophic pathways. We have employed 3-D coupled physical-biogeochemical models to explore ecosystem responses to climate change with a focus on trophic amplification. The response of phytoplankton and zooplankton to global climate-change projections, carried out with the IPSL Earth System Model by the end of the century, is analysed at global and regional basis, including European seas (NE Atlantic, Barents Sea, Baltic Sea, Black Sea, Bay of Biscay, Adriatic Sea, Aegean Sea) and the Eastern Boundary Upwelling System (Benguela). Results indicate that globally and in Atlantic Margin and North Sea, increased ocean stratification causes primary production and zooplankton biomass to decrease in response to a warming climate, whilst in the Barents, Baltic and Black Seas, primary production and zooplankton biomass increase. Projected warming characterized by an increase in sea surface temperature of 2.29 ± 0.05 °C leads to a reduction in zooplankton and phytoplankton biomasses of 11% and 6%, respectively. This suggests negative amplification of climate driven modifications of trophic level biomass through bottom-up control, leading to a reduced capacity of oceans to regulate climate through the biological carbon pump. Simulations suggest negative amplification is the dominant response across 47% of the ocean surface and prevails in the tropical oceans; whilst positive trophic amplification prevails in the Arctic and Antarctic oceans. Trophic attenuation is projected in temperate seas. Uncertainties in ocean plankton projections, associated to the use of single global and regional models, imply the need for caution when extending these considerations into higher trophic levels.