Surface patterns of zooplankton spatial variability detected by high frequency sampling in the NW Mediterranean. Role of density fronts

High frequency sampling was performed in daylight hours along a 35 km transect in the Ligurian Sea to investigate the upper layer zooplankton distribution during the spring phytoplankton bloom. The results show detailed spatial structure and biomass of key zooplankton functional groups, copepods, salps and krill larvae, within the different water masses characterizing this region. Although observed values of total copepod biomass distribution were rather constant along the transect, species-specific patterns were observed in the copepod spatial distribution. The larger species Calanus helgolandicus, as well as Centropages typicus, Oithona spp., and Oncaea spp., were associated with the frontal zone. However, Acartia spp. had a scattered distribution, and Clausocalanus/Paracalanus did not have a clear pattern. In addition, krill larvae were concentrated in the frontal area and salps had a scattered pattern. The cross-shore zooplankton distribution appeared strongly influenced by both the Northern Ligurian current governing inshore waters, which acts as a major flushing forcing, and the Ligurian front, which governs offshore waters and may act as retention area for zooplankton.     

Morphological and biochemical differentiation in Antarctic krill

During the February 1981 cruise FIBEX MD-25 between 30–50°E and 61–64°S, hydrography showed the presence of two gyres, confirmed by the geostrophic circulation relative to 1000 m from Levitus climatology, at the borders of these gyres concentrations of highly morphologically differentiated krill were found. Gaussian component analysis of krill samples, pooled by sectors, showed three cohorts of Euphausia superba in the western sector and one in the eastern sector. Across the sampling area, Thysanoessa macrura and E. superba occurred at separate stations. Analysis of cohorts in T. macrura separated two size groups in both the western and the eastern sectors. The use of a Differentiation Index (D.I.) [Färber-Lorda, J., 1990. Somatic length relationships and ontogenetic morphometric differentiation of Euphausia superba and Thysanoessa macrura of the southwest Indian Ocean during summer (February 1981). Deep-Sea Res. 37, 1135–1143.], based on somatic lengths, allows studying certain morphological differences within the populations sampled. Morphologically different and bigger males II (D.I. from 2.8 to 3.5) were present only in the southern transect while smaller males I (D.I. from 3.5 to 5.0) were present over the entire area. Biochemical composition of both species showed significant differences among stations for protein, lipids, and carbohydrates. A significant difference in lipid content was found between males I, and males II. For T. macrura, percentage of lipid content in mature animals was much higher than that in E. superba. The D.I. size distribution showed that when populations of E. superba were highly differentiated (corresponding to mature animals) in morphology, lipid content was high, and they were located near a gyre. Differences in morphometry can influence distribution of the species, because different developing stages have different swimming capacities. It is shown that, together with hydrography and trophic conditions, lipid content and morphometry of krill populations, are different but complementary aspects that help to understand krill ecology and distribution.     

Succession of the ecosystems of the Aral Sea during its transition from oligohaline to polyhaline water body

During 22 field trips from 1990 to 2002 (mainly the western basin of the Large Aral) data on salinity, phytoplankton, zooplankton, zoobenthos and fish fauna have been collected. In 2002, the salinity of the western basin reached 75 ppt, while that in the eastern basin, 150 ppt. In 1999–2002, 159 species of planktonic algae have been recorded. This is approximately twice as low as recorded before. The diversity of Cyanophyta, Pyrrhophyta and Chlorophyta in particular has dropped in the past few years. As before, currently Bacillariophyta is the most diverse plankton. However, the composition of dominants has changed. Once previously dominant species, Actinocyclus ehrenbergii, vanished from the plankton of the Aral Sea and was replaced by such diatoms as Amphora coffeaformis, A. coffeaformis var. acutiuscula and Synedra tabulata var. parva.Since 1970s, a gradual decrease in the diversity of zooplankton has been taking place. Since 1997, the formerly dominant Calanipeda aquaedulcis vanished, which apparently was the reason for the emergence of Moina salina and Artemia parthenogenetica. Since 2000, artemia has been dominant in the plankton of the Aral Sea, constituting 99% of the zooplankton biomass.In the 1970–1980s, a rapid decrease in the biodiversity of the zoobenthos was observed. In the 1990s, most aboriginal and introduced species became extinct. Currently, the bivalve mollusk Syndosmya segmentum, the ostracod Cyprideis torosa and larvae of the dipteran Chironomus salinarius can still be recorded in the western basin. In the eastern basin no benthos is observed.By 1998, in the Large Aral, only five fish species survived: baltic herring Clupea harengus membras, flounder Platichthys flesus luscus, atherine Atherina boyeri caspia and bullheads Neogobius fluviatilis and Potamoschistus caucasicus. Since 2002, only flounder and atherina have been recorded in the western basin of the Large Aral. No fish have been recorded in the eastern part of the Aral Sea in 2002.With increasing salinity and transition of the Aral Sea from an oligohaline to a polyhaline water body, its biota is becoming drastically poorer. Almost all local species became extinct in the Aral; however, some still survive (including some endemics) in some lakes around the Aral Sea. In the near future, artemia will be the only animal in the Aral Sea.     

Error quantification of a high resolution coupled hydrodynamic-ecosystem coastal-ocean model: Part3, validation with Continuous Plankton Recorder data

Data collected during the Continuous Plankton Recorder (CPR) survey has been used to validate a three-dimensional hydrodynamic ecosystem model simulation of the North-west European Shelf for the years 1988–89. The CPR time series is unique to the North Atlantic region as a validation tool. Data were extracted from the model to correspond with those collected by the CPR survey, and both the model and survey plankton data were standardised to allow the comparison of model biomass with survey counts. Simple linear regression and absolute error maps provide a qualitative evaluation of spatio-temporal model performance of simulated diatoms, flagellates, total phytoplankton and omnivorous mesozooplankton. Comparisons of z-scores indicate that the model reproduces the main pelagic seasonal features, and there is good correlation between magnitudes of these features with respect to standard deviations from a long-term mean. The model is replicating up to 62% of the mesozooplankton seasonality across the domain, with variable results for the phytoplankton. There are, however, differences in the timing of patterns in plankton seasonality. The validation exercise has highlighted that the spring diatom bloom in the model is too early, suggesting the need to reparameterise the response of phytoplankton to changing light levels in the model. Errors in the north and west of the domain imply that model turbulence and vertical density structure need to be improved to more accurately capture plankton dynamics.     

Validation of the 3D biogeochemical model MIRO&CO with field nutrient and phytoplankton data and MERIS-derived surface chlorophyll a images

This paper presents results obtained with MIRO&CO-3D, a biogeochemical model dedicated to the study of eutrophication and applied to the Channel and Southern Bight of the North Sea (48.5°N–52.5°N). The model results from coupling of the COHERENS-3D hydrodynamic model and the biogeochemical model MIRO, which was previously calibrated in a multi-box implementation. MIRO&CO-3D is run to simulate the annual cycle of inorganic and organic carbon and nutrients (nitrogen, phosphorus and silica), phytoplankton (diatoms, nanoflagellates and Phaeocystis), bacteria and zooplankton (microzooplankton and copepods) with realistic forcing (meteorological conditions and river loads) for the period 1991–2003. Model validation is first shown by comparing time series of model concentrations of nutrients, chlorophyll a, diatom and Phaeocystis with in situ data from station 330 (51°26.00′N, 2°48.50′E) located in the centre of the Belgian coastal zone. This comparison shows the model's ability to represent the seasonal dynamics of nutrients and phytoplankton in Belgian waters. However the model fails to simulate correctly the dissolved silica cycle, especially during the beginning of spring, due to the late onset (in the model) of the early spring diatom bloom. As a general trend the chlorophyll a spring maximum is underestimated in simulations. A comparison between the seasonal average of surface winter nutrients and spring chlorophyll a concentrations simulated with in situ data for different stations is used to assess the accuracy of the simulated spatial distribution. At a seasonal scale, the spatial distribution of surface winter nutrients is in general well reproduced by the model with nevertheless a small overestimation for a few stations close to the Rhine/Meuse mouth and a tendency to underestimation in the coastal zone from Belgium to France. PO₄ was simulated best; silica was simulated with less success. Spring chlorophyll a concentration is in general underestimated by the model. The accuracy of the simulated phytoplankton spatial distribution is further evaluated by comparing simulated surface chlorophyll a with that derived from the satellite sensor MERIS for the year 2003. Reasonable agreement is found between simulated and satellite-derived regions of high chlorophyll a with nevertheless discrepancies close to the boundaries.     

Review on the seasonal variation of the surface circulation in the Japan/East Sea

The seasonal variation of the surface circulation in the Japan/East Sea (JES) and the Tsushima/Korea Straits (TKS) is reviewed and discussed, focusing on mesoscale and submesoscale variabilities.The monsoon modified by coastal geographical features near Vladivostok generates a dipole of vortex off Vladivostok which induces dramatic changes in the surface circulation in the northwest JES, splitting the Subpolar Gyre into two smaller gyres by generating the Vladivostok Dome. Between these two smaller gyres, the Northwest Thermal Front is formed and current reversal is induced along the North Korean coast. The winter monsoon also induces a current reversal along the Sakhalin coast. The volume transport of the surface Subpolar Gyre has two maxima in January and August. The maximum in August is induced by the summer intensification of the Liman-North Korean Cold Current and the shallow and narrow surface coastal jet generated by the sea ice and snow melting. The maximum in January is induced by the northwest monsoon and associated cooling.Salient features in the TKS are the submesoscale variabilities. In the western channel, submesoscale eddies with length scale of about 80 km and time scale of 5–6 days develop in the cold period. On the lee side of the Tsushima Islands, Karman-like vortex pairs are generated in the warm period. Anticyclonic vortices generated at the northern tip of the Tsushima Islands have a time scale of 5 to 8 days, length scale of 35 to 60 km, and propagate toward the JES with a phase speed of 8 cm/s. Cyclonic vortices south of the anticyclonic counter part of the vortex pairs are rather stationary with intermittent occasional propagation toward the east. The development of stratification seems to be necessary for the development of Karman-like vortex pairs.Summarizing the results above, a schematic surface circulation with seasonal change is proposed.     

Convection and the seeding of the North Atlantic bloom

Observations of vertical velocities in deep wintertime mixed layers using neutrally buoyant floats show that the convectively driven vertical velocities, roughly 1000 m per day, greatly exceed the sinking velocities of phytoplankton, 10 m or less per day. These velocities mix plankton effectively and uniformly across the convective layer and are therefore capable of returning those that have sunk to depth back into the euphotic zone. This mechanism cycles cells through the surface layer during the winter and provides a seed population for the spring bloom. A simple model of this mechanism applied to immortal phytoplankton in the subpolar Labrador Sea predicts that the seed population in early spring will be a few percent of the fall concentration if the plankton sink more slowly than the mean rate at which the surface well-mixed layer grows over the winter. Plankton that sink faster than this will mostly sink into the abyss with only a minute fraction remaining by spring. The shallower mixed layers of mid-latitudes are predicted to be much less effective at maintaining a seed population over the winter, limiting the ability of rapidly sinking cells to survive the winter.     

Plankton distribution and the impact of copepod grazing on primary production in Fram Strait, Greenland Sea

Two hydrobiological transects across the East Greenland Shelf and the open waters of Fram Strait in summer were chosen to illustrate the distribution and production of phyto- and zooplankton in relation to water masses and ice cover. The parameters used were temperature and salinity, inorganic nutrients, chlorophyll a, primary production, phytoplankton species composition, abundance of the dominant herbivorous copepods Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa and egg production of C. finmarchicus and C. glacialis. Grazing impact of copepodites and adults of these four species was modelled for each station by using egg production rates as an index of growth. Seasonal development of plankton communities was closely associated with the extent of the ice cover, hydrographic conditions and the water masses typical of the different hydrographic domains. Four regions were identified from their biological activities and physical environment: The Northeast Water polynya on the East Greenland Shelf, with a springbloom of diatoms and active reproduction of herbivorous copepods. The pack ice region, dominated by small flagellates and negligible grazing activities. The marginal ice zone, with high variability and strong gradients of autotroph production related to eddies and ice tongues, an active microbial loop and low egg production. The open water, with high station-to-station variability of most of the parameters, probably related to hydrographic mesoscale activities. Here, Phaeocystis pouchetii was a prominent species in the phytoplankton communities. Its presence may at least partly be responsible for the generally low egg production in the open waters. Grazing impact on primary production was always small, due to low zooplankton biomass in the polynya and due to low ingestion in the remaining regions.     

On the relation between organic and inorganic carbon in the Weddell Sea

Carbon cycling in the Weddell Sea was investigated during the ANT X/7 cruise with `FS Polarstern' December 1992–January 1993. Samples were taken on a cross section from Kapp Norvegia to Joinville Island, and on a section from the Larsen Ice Shelf to the northeast. The following quantities were measured: total carbon dioxide (TCO₂), fluorescence from humic substances and total organic carbon. The distribution of TCO₂ was strongly positively correlated to the time elapsed since the various water masses were last ventilated. In general, humic substance fluorescence was positively correlated with TCO₂, with the exception of the productive part of the western Weddell Sea, where the correlation was negative in the surface mixed layer. The increased fluorescence at the surface is suggested to be a result of biological production. The distribution of total organic carbon showed less structure, since this quantity includes a particulate component, which is subject to dispersion processes different from those of the dissolved components TCO₂ and humic substances. The mean total organic carbon concentration below the surface mixed layer was 50 μmol l⁻¹. At some stations, a steep TOC maximum around 2000 m depth was observed. This was interpreted to result from mass sinking of phytoplankton blooms. Total organic carbon had a maximum in surface water, and at some stations also a second subsurface maximum. In the Warm Deep Water (WDW), TCO₂ and fluorescence had their maximum values, while total organic carbon tended to be low. In low productivity surface water in the eastern part of the Kapp Norvegia–Joinville Island section, the lowest flourescence was found. Surface water is eventually formed from Warm Deep Water, which had the highest fluorescence values, and therefore it is concluded that humic substances were removed in situ from surface water. In the central area of the Weddell Sea, TCO₂ and fluorescence showed the highest Warm Deep Water maxima, while total organic carbon was low. The Warm Deep Water in this area is part of the so-called Central Intermediate Water which circulates for a long time within the Weddell Gyre. Reduced total organic carbon, which coincides with the most pronounced Central Intermediate Water characteristics, and high TCO₂ can thus both be accounted for by continued degradation of organic matter in this water mass. The associated fluorescence maximum implies that humic substances are also produced during mineralisation. Recently formed bottom water, by contrast, could be seen as patches of low TCO₂, low fluorescence and high total organic carbon along the western slope of the Weddell Sea.     

Distribution, reproduction and development of Calanus species in the Northeast water in relation to environmental conditions

The vertical and horizontal distribution of mesozooplankton biomass and its composition, together with the reproduction and development of the three dominant herbivorous copepods Calanus finmarchicus, C. glacialis and C. hyperboreus, were studied in the Northeast Water between the end of May and end of July, 1993. Biomass varied between 0.15 and 10.6 g m⁻² and was lower on the shallow banks. Highest biomass was found on the shelf slope and in the entrances to Belgica and Westwind Troughs. There was no seasonal trend during the study period. Among the zooplankton taxa, copepods were dominant, with 84% biomass of all other taxa, followed by chaetognaths with 14%. The large Calanus species made up 91% of copepod biomass. The most abundant species, C. glacialis, inhabited areas of low current speeds on Belgica and Ob Bank, C. hyperboreus dominated shelf slope and trough stations, while C. finmarchicus was most abundant in the Return Atlantic Current along the shelf slope and also eastern Belgica Trough. C. glacialis was the only Calanus species spawning during this period, but young copepodites of the other species were also abundant. Egg production of C. glacialis was at a maximum by our arrival and continued to at least mid August. According to the results from starvation experiments, its egg production was fuelled by food uptake, but was decoupled from phytoplankton chlorophyll until July, indicating ice-algae and microzooplankton as an alternative food source. Only when the polynya approached its maximum extent was a close relationship to phytoplankton established. Due to both spatial and temporal heterogeneity of the occurrence of young copepodite stages it was impossible to follow the growth of cohorts of developmental stages. Herbivorous carbon requirements estimated from egg production rates were mostly less than one third of the phytoplankton stock. From egg production and the distribution of young stages, the outer Westwing Trough seems to be the centre of biological activities. This may be related to the formation of young ice in winter in this area, which was found to carry a large mass of under-ice algae. High biomass but low production in Belgica Trough indicate this as an advective and expatriated community of C. finmarchicus and C. hyperboreus, where grazing is negligible, at least during early summer.     

One-dimensional modelling of the plankton ecosystem of the north-western Corsican coastal area in relation to meteorological constraints

In order to study the influence of wind mixing on the spring variability of the plankton production of the north western Corsican coastal area, a one-dimensional (1D), vertical, coupled hydrodynamic/biological model (ECOHYDROMV) is used. A hydrodynamic 1D model of the water column with a k–l turbulent closure is applied. The biological model comprises six state variables, representing the plankton ecosystem in the spring period: phytoplankton, copepods, nitrate, ammonium, particulate organic matter of phytoplanktonic origin and particulate organic matter of zooplanktonic origin. The system is influenced by turbulence (expressed by the vertical eddy diffusivity), temperature and irradiance. The model takes into account momentum and heat surface fluxes computed from meteorological data in order to simulate a typical spring atmospheric forcing for the considered area. Results show that primary production vertical structure is characterised by a subsurface maximum which deepens with time and is regulated by the opposite gradients of nitrate concentration and irradiance. Surface plankton productivity is mainly controlled by turbulent vertical transport of nutrients into the mixed layer. The short time scale variability of turbulent mixing generated by the wind appears to be responsible for the plurimodal shape of plankton blooms, observed in the considered area. Furthermore, the model is applied to the study of the spring evolution of the plankton communities off the bay of Calvi (Corsica) for the years 1986 and 1988. In order to initiate and validate the model, time series of hydrological, chemical and biological data have been used. The model reproduces accurately the spring evolution of the phytoplankton biomass measured in situ and illustrates that its strong variability in those years was in close relation to the variability of the wind intensity.     

Dynamics of suprabenthos-zooplankton communities around the Balearic Islands (western Mediterranean): Influence of environmental variables and effects on the biological cycle of Aristeus antennatus

Dynamics of suprabenthos and zooplankton were analyzed in two areas located in the NW (off Sóller harbour) and S (off Cabrera Archipelago) of Mallorca (Balearic Islands, western Mediterranean) at depths ranging between 135–780 m. Four stations situated respectively at 150 m (shelf-slope break), and at bathyal depths of 350, 650 and 750 m were sampled at bi-monthly intervals during six cruises performed between August 2003 and June 2004. Suprabenthos showed maximum biomass in both areas from late spring to summer (April to August), while minimum biomass was found in autumn (September–November). Though variable, temporal dynamics of zooplankton showed peaks of biomass in late winter and summer (February and June), while minimals occurred in autumn (August–September) and, at bathyal depths, in April. Suprabenthos (abundance; MDS analyses) showed a sample aggregation as a function of depth (3 groups corresponding to the shelf-slope break, upper slope — over 350 m; and the middle, deeper part of the slope — over 650–750 m), without any separation of hauls by season. By contrast, zooplankton samples were separated by season and not by depth. There was evidence of three seasonal groups corresponding to summer (June 2004–August 2003), autumn–winter (September and November 2003, February 2004), and spring (April 2004), being especially well established off Sóller. In general, suprabenthos was significantly correlated with the sediment variables (e.g. total organic matter content (% OM), potential REDOX), whereas zooplankton was almost exclusively dependent on Chl a at the surface, which suggests two different food sources for suprabenthos and zooplankton. The increase of suprabenthos abundance in April–June was paralleled by a sharp increase (ca. 2.8 times) in the %OM on sediment during the same period, coupled ca. 1–2 months of delay with the peak of surface Chl a recorded in February–March (from satellite imagery data). Suprabenthos biomass was also correlated with salinity close to the bottom, suggesting a link between suprabenthos abundance and changes in the oceanographic condition of water masses close to the bottom. It is suggested that a higher suprabenthos biomass recorded off Sóller in comparison to that off Cabrera in June could, in turn, be related to a seasonal inflow of Levantine Intermediate Water (LIW) in April–June in this area at mid bathyal depths (350–650 m). This trend would be based on: 1) it was evident only at mid-slope depths between 350–750 m, coinciding with the LIW distribution, and 2) it was not recorded among zooplankton (collected throughout the water column). The possible effect of the fluctuations of suprabenthos and zooplankton on higher trophic levels has been explored studying the diet and food consumption rates of the red shrimp Aristeus antennatus, as indicator species by its dominance in bathyal communities. A. antennatus increased its food consumption from February to April–June 2004 off Sóller, which in the case of large (CL > 40 mm) specimens was found in both areas. In addition, there was a shift of diet from winter to spring–early summer. In this last period, A. antennatus preyed upon euphausiids and mesopelagic decapods and fish, while benthos (e.g. polychaetes and bivalves) decreased in the diet. This indicates an increase in the food consumption and probably in the caloric content of the diet in pre-spawning females in April–June 2004, which is synchronized with the period when gonad development begins in A. antennatus females (May–June). Anyway, macrozooplankton, and not suprabenthos, is crucial as a high energetic food source in the coupling between food intake and reproduction in the red shrimp.     

Seasonal and interannual variation of physical and biological processes during 1994–2001 in the Sea of Japan/East Sea: A three-dimensional physical–biogeochemical modeling study

A Pacific basin-wide physical–biogeochemical model has been used to investigate the seasonal and interannual variation of physical and biological fields with analyses focusing on the Sea of Japan/East Sea (JES). The physical model is based on the Regional Ocean Model System (ROMS), and the biogeochemical model is based on the Carbon, Si(OH)₄, Nitrogen Ecosystem (CoSiNE) model. The coupled ROMS–CoSiNE model is forced with the daily air–sea fluxes derived from the National Centers for Environmental Prediction (NCEP) and the National Center for Atmospheric Research (NCAR) reanalysis for the period of 1994 to 2001, and the model results are used to evaluate climate impact on nutrient transport in Mixed Layer Depth (MLD) and phytoplankton spring bloom dynamics in the JES.The model reproduces several key features of sea surface temperature (SST) and surface currents, which are consistent with the previous modeling and observational results in the JES. The calculated volume transports through the three major straits show that the Korea Strait (KS) dominates the inflow to the JES with 2.46 Sv annually, and the Tsugaru Strait (TS) and the Soya Strait (SS) are major outflows with 1.85 Sv and 0.64 Sv, respectively. Domain-averaged phytoplankton biomass in the JES reaches its spring peak 1.8 mmol N m^− 3 in May and shows a relatively weak autumn increase in November. Strong summer stratification and intense consumption of nitrate by phytoplankton during the spring result in very low nitrate concentration at the upper layer, which limits phytoplankton growth in the JES during the summer. On the other hand, the higher grazer abundance likely contributes to the strong suppression of phytoplankton biomass after the spring bloom in the JES. The model results show strong interannual variability of SST, nutrients, and phytoplankton biomass with sudden changes in 1998, which correspond to large-scale changes of the Pacific Decadal Oscillation (PDO). Regional comparisons of interannual variations in springtime were made for the southern and northern JES. Variations of nutrients and phytoplankton biomass related to the PDO warm/cold phase changes were detected in both the southern and northern JES, and there were regional differences with respect to the mechanisms and timing. During the warm PDO, the nutrients integrated in the MLD increased in the south and decreased in the north in winter. Conversely, during the cold PDO, the nutrients integrated in the MLD decreased in the south and increased in the north. Wind divergence/convergence likely drives the differences in the southern and northern regions when northerly and northwesterly monsoon dominates in winter in the JES. Subjected to the nutrient change, the growth of phytoplankton biomass appears to be limited neither by nutrient nor by light consistently both in the southern and northern regions. Namely, the JES is at the transition zone of the lower trophic-level ecosystem between light-limited and nutrient-limited zones.     

Daily and seasonal variations of the partial pressure of CO2 in surface seawater along Belgian and southern Dutch coastal areas

The variations of the partial pressure of CO₂ (pCO₂) and related parameters were determined in surface seawater along the Belgian coast, from January 1995 to June 1996, at both daily and seasonal time scales. The distribution of pCO₂ in this area is regulated by river input from the Scheldt, biological activity and hydrodynamics. The contribution of each of these processes varies as a function of the considered time scale: (i) the daily variation of pCO₂ depends on the tide although modulated by the biological diel cycle; (ii) the seasonal variation of pCO₂ depends on the input from the Scheldt and the seasonal variations of phytoplanktonic biomass. During winter, the plume of the river Scheldt is oversaturated in pCO₂ with respect to the atmosphere. During spring and summer, phytoplankton blooms occur both in the lower Scheldt estuary and in the river plume and may lead to undersaturation of pCO₂ in the easternmost area of the river plume. However, the degradation of phytoplankton induces oversaturation of pCO₂, in the westernmost area of the plume. Furthermore, the inter-annual variation of pCO₂ depends partly on the fluctuations of the discharge of the Scheldt. Our preliminary results strongly suggest that, on an annual basis, the Scheldt plume behaves as a net source of CO₂ to the atmosphere.     

An analysis of the characteristics of chlorophyll in the Sulu Sea

The seasonal and spatial variations of chlorophyll concentrations, Sea Surface Temperature (SST), wind fields and wind-induced Ekman pumping in the Sulu Sea are investigated using a set of new remote sensing measurements from October 1997 to December 2004. The results show the seasonality of chlorophyll, wind fields and SST and reveal the phytoplankton blooming events in the Sulu Sea basin during the northeast monsoon season. In summer, chlorophyll concentrations were relatively low (< 0.2 mg/m³) and distributed uniformly throughout the basin with a narrow belt of high chlorophyll concentrations along the coastal waters, particularly the coasts of Borneo and of the Sulu Archipelago. In winter, chlorophyll concentrations increased (> 0.2 mg/m³) throughout the entire basin, and phytoplankton bloomed southward to the central basin, while chlorophyll concentrations reached high levels (1 mg/m³) in the center of the blooms. One peak was observed during the northeast monsoon season each year. SSTs have significant negative correlations with chlorophyll concentrations; i.e., high and uniformly distributed in summer but lower with an obvious tongue of cold waters southward to the central basin in winter. The seasonal variation of chlorophyll concentrations and SST distribution were associated with the seasonally reversing monsoon. The winter phytoplankton blooming and the tongue of the cold waters were correlated to the vertical upwelling cold and nutrient-rich waters drawn by the northeast wind, with the center of the blooms and the location of cold tongues coinciding with the maximum of the wind speeds and the Ekman pumping velocities.     

Simulations of biological production in the Rhodes and Ionian basins of the eastern Mediterranean

The biological production characteristics of the Rhodes and western Ionian basins of the eastern Mediterranean are studied by a one-dimensional, coupled physical–biological model. The biological model involves single aggregated compartments of phytoplankton, zooplankton, detritus as well as ammonium and nitrate forms of the inorganic nitrogen. It interacts with the physical model through the vertical eddy diffusivity which is calculated using the Mellor–Yamada level 2.5 turbulence parameterization. The model simulations demonstrate the importance of the contrasting physical oceanographic characteristics of these two basins on affecting their yearly planktonic structures. The annual primary production in the Rhodes basin is estimated as 97 g C m² yr⁻¹ which is comparable with the northwestern Mediterranean. The western Ionian basin, on the contrary, possesses only 10% of the Rhodes' productivity and therefore represent a most oligotrophic site in the eastern Mediterranean. The Rhodes basin reveals a strong bloom in early spring, typically in March, a weaker bloom in early winter, typically in January, and a subsurface production below the seasonal thermocline during summer. This structure is slightly modified in the western Ionian basin, and the early winter and early spring blooms are merged to cover the entire winter. These results are supported favorably by the available observations both in their magnitudes and timing.     

Seasonal variation of riverine nutrient inputs in the northern Bay of Biscay (France), and patterns of marine phytoplankton response

Seasonal variations in nutrient inputs are described for the main rivers (Loire and Vilaine) flowing into the northern Bay of Biscay. The river plumes are high in N/P ratio in late winter and spring, but not in the inner plume during the summer. Conservative behavior results in most nutrients entering the estuary and eventually reaching the coastal zone. Temporal and spatial aspects of phytoplankton growth and nutrient uptake in the northern Bay of Biscay distinguish the central area of salinity 34 from the plume area. The first diatom bloom appears offshore in late winter, at the edge of the river plumes, taking advantage of haline stratification and anticyclonic “weather windows.” In spring, when the central area of the northern shelf is phosphorus-limited, small cells predominate in the phytoplankton community and compete with bacteria for both mineral and organic phosphorus. At that period, river plumes are less extensive than in winter, but local nutrient enrichment at the river mouth allows diatom growth. In summer, phytoplankton become nitrogen-limited in the river plumes; the central area of the shelf is occupied by small forms of phytoplankton, which are located on the thermocline and use predominantly regenerated nutrients.     

Fluxes of biogenic carbon in the Southern Ocean: roles of large microphagous zooplankton

The Southern Ocean is an extreme environment, where waters are permanently cold, a seasonal ice cover extends over large areas, and the solar energy available for photosynthesis is severely restricted, either by vertical mixing to considerable depths or, especially south of the Antarctic Circle, by prolonged seasonal periods of low or no irradiance. Such conditions would normally lead to low productivity and a water column dominated by recycling processes involving microbial components of pelagic communities but this does not seem to be the case in the Southern Ocean, where there is efficient export to large apex predators and deep waters. This paper investigates the role of large microphagous zooplankton (salps, krill, and some large copepods) in the partitioning of biogenic carbon among the pools of short- and long-lived organic carbon and sequestered biogenic carbon. Large microphagous zooplankton are able to ingest microbial-sized particles and thus repackage small, non-sinking particles into both metazoan biomass and large, rapidly sinking faeces. Given the wide spatio-temporal extent of microbial trophic pathways in the Southern Ocean, large zooplankton that are omnivorous or able to ingest small food particles have a competitive advantage over herbivorous zooplankton. Krill efficiently transfer carbon to a wide array of apex predators and their faecal pellets are exported to depth during occasional brief sedimentation episodes in spring time. Salps may be a significant link towards some fish (directly) and other apex predators (indirectly) and, at some locations (especially in offshore waters) and time, they may account for most of the downward flux of biogenic carbon. Large copepods are a trophic link towards fish and at least one whale species, and their grazing activity generally impedes the export of organic particles to depth. As a result, biogenic carbon is channelled mainly towards apex predators and episodically into the deep ocean. Without these original interactions, Antarctic waters might well be dominated by microbial components and recycling processes instead of active export from the generally small primary producers towards large apex predators.