Distribution of phytoplankton in the southern Black Sea in summer 1996, spring and autumn 1998

The species composition, abundance, and biomass of micro- (>15 μm) and nano- (<15 μm) phytoplankton were studied along the southern Black Sea during June–July 1996 and March–April and September 1998. A total of 150 species were identified, 50% of them being dinoflagellates. The average total phytoplankton abundance changed from 77×10³ cells l⁻¹ in spring to 110×10³ cells l⁻¹ in autumn and biomass from 250 μg l⁻¹ in summer to 1370 μg l⁻¹ in spring. Based on the extensive sampling grid from June–July 1996, phytoplankton seemed to have a rather homogeneous biomass distribution in the southern Black Sea. In all periods, the coccolithophorid Emiliania huxleyi was the most abundant species, its contribution to the total abundance ranging from 73% in autumn to 43% in spring. However, in terms of biomass, diatoms made up the bulk of phytoplankton in spring (97%, majority being Proboscia alata) and autumn (73%, majority being Pseudosolenia calcar-avis), and dinoflagellates in summer (74%, Gymnodinium sp.). There was a remarkable similarity in the dominant species between the western and eastern regions of the southern Black Sea, indicating transport of phytoplankton within the basin.     

Size-fractionated phytoplankton biomass and species composition in the Indian sector of the Southern Ocean during austral summer

During the late austral summer of 1994, Antarctic waters were characterized by low phytoplankton biomass. Along the 62°E meridian transect, between 49°S and 67°S, chlorophyll (Chl.) a concentration in the upper 150 m was on average 0.2 mg m⁻³. However, in the Seasonal Ice Zone (SIZ) chlorophyll a concentrations were higher, with a characteristic deep chlorophyll maximum. The highest value (0.6 mg Chl. a m⁻³) was measured at the Antarctic Divergence, 64°S, corresponding to the depth of the temperature minimum (100 m). This deep biomass maximum decreased from South to North, disappeared in the Permanently Open Ocean Zone (POOZ) and reappeared with less vigour in the vicinity of the Polar Front Zone (PFZ). In the SIZ, the upper mixed layer was shallow, biomass was higher and the >10 μm fraction was predominant. In this zone the >10 μm, 2–10 μm and <2 μm size fractions represented on the average 46%, 25.1% and 28.9% of the total integrated Chl. a stock in the upper 100 m, respectively. The phytoplankton assemblage was diverse, mainly composed of large diatoms and dinoflagellate cells which contributed 42.7% and 33.1% of the autotrophic carbon biomass, respectively. Moving northwards, in parallel with the decrease in biomass, the biomass of autotrophic pico- and nanoflagellates (mainly Cryptophytes) increased steadily. In the POOZ, the picoplanktonic size fraction contributed 47.4% of the total integrated Chl. a stock. A phytoplankton community structure with low biomass and picoplankton-dominated assemblage in the POOZ contrasted with the relatively rich, diverse and diatom-dominated assemblage in the SIZ. These differences reflect the spatial and temporal variations prevailing in the Southern Ocean pelagic ecosystem.     

Seasonal carbon distribution of copepods in the eastern Weddell Sea, Antarctica

Copepods were sampled by a multiple opening-closing net in the eastern Weddell Sea during various seasons (late winter/early spring, summer, autumn). Total copepod biomass integrated over the upper 1000 m varied seasonally between 1.7 mg C m⁻³ in late winter/early spring and 3.7 mg C m⁻³ in autumn. After the dark season the copepods were rather evenly distributed vertically and highest biomass levels were found in the mid-water layers between about 200 m and 500 m. By contrast, especially in summer but also in autumn copepod biomass concentrated in the uppermost water layer. A total of 64 calanoid species were identified in the upper 1000 m with maximum species numbers in the deepest layer. The large calanoids Calanus propinquus, Calanoides acutus, Metridia gerlachei, Euchaeta antarctica and the small calanoid Microcalanus pygmaeus prevailed and accounted for 60–70% of total copepod biomass, while the small poecilostomatoid Oncaea and the cyclopoid Oithona species comprised about 20%. Hence, the distribution pattern of the entire copepod biomass is strongly influenced by the life cycles of a few dominant species.     

Distribution of plankton lipids and their role in the biological transformation of Antarctic primary production

Production and transfer of lipid through the Antarctic food web is reviewed for the Indian Ocean sector. The slow settling fine particles showed a marked inter-annual variability in biochemical composition with an increase in lipid content as % organic carbon. Comparison of the fatty acid spectra of different size categories of organic particles indicated that fine particles are dominated by saturated, monoenoic and branched acids, while larger material (50–100 μm, 200–500 μm net collected fractions) displayed a signature dominated by polyunsaturated acids. Zooplankton taxa displayed different strategies of lipid accumulation. Lipid content was highest in Thysanoessa macrura females and copepodite stages of Calanus propinquus. Relatively low levels were recorded for juveniles and male stages of euphausiids. Reserve lipids varied with species: C. propinquus showed equal content of triglycerides and wax esters, T. macrura showed a dominance of wax esters and Euphausia superba and Themisto gaudichaudii accumulated only triglycerides. Computed as carbon equivalent and integrated over 200 m, lipids in slow settling particles represented 22.6% of annual primary production. Similar computation with mesozooplankton and E. superba data on biomass and population structure from several summer cruises indicated values of carbon accumulation as lipid reserves and egg production of 4.2 and 0.1% of annual primary production for copepods and 4.4 and 3.8% for E. superba. When all trophic levels are considered, the overall mean exceeded 30% of annual primary production.     

Spatial and temporal variability of size-fractionated biomass and primary production in the Ross Sea (Antarctica) during austral spring and summer

The results of a study on the spatial and temporal dynamics of size-fractionated biomass and production of phytoplankton in the Ross Sea during the austral spring and summer are reported. The spring cruise took place in the offshore Ross Sea from 14 November to 14 December 1994. Sampling was carried out on a transect of 27 stations distributed from 76.5 to 72.0°S along 175°E, and covered the three main Antarctic environments of the polynya open waters, the marginal ice zone and the pack ice area. Three subsystems were identified. The subsystem of the polynya was characterised by the predominance of the micro- and nano-planktonic fractions, chlorophyll (Chl a) concentrations from 69.6 to 164.7 mg m⁻² and production rates from 0.68 to 1.14 g C m⁻² day⁻¹. The second subsystem, the marginal ice zone, showed a relative increase of the micro-planktonic fraction, high biomass levels (from 99.64 to 220 mg Chl m⁻²) and production rates from 0.99 to 2.7 g C m⁻² day⁻¹. The subsystem of the pack ice area had a phytoplankton community dominated by the pico-planktonic fraction and showed low biomasses (from 19.4 to 37.7 mg Chl m⁻²) and production rates (0.28 to 0.60 g C m⁻² day⁻¹). Selective grazing by krill is considered an important factor in determining the size structure of the phytoplankton communities. The summer study consisted of a time series carried out in inshore waters of Terra Nova Bay from 12 January to 8 February 1990. In a well stabilised water column and with high levels of PAR always available, the primary production rates of a community dominated by micro-plankton varied from 0.52 to 1.2 g C m⁻² day⁻¹ (average 0.84). A high P/B ratio, up to 3, and a remarkably elevated mean phaeopigment (Phaeo)/Chl a ratio of 2.4 indicated an active removal of biomass by grazing, confirmed by the presence of faecal pellets in quantities reaching 6000 m⁻³ in the upper 50 m. The peculiarities of the inshore versus offshore environments in terms of community size structure, production processes and their implications as regards the food web are discussed.     

Vertical profiles of particulate organic matter and its relationship with chlorophyll-a in the upper layer of the NE Mediterranean Sea

Particulate organic matter (POM), nutrients, chlorophyll-a (CHL) and primary production measurements were performed in the upper layer of three different regions (cyclonic, anticyclonic and frontal+peripherial) of the NE Mediterranean Sea in 1991–1994. Depth profiles of bulk POM exhibited a subsurface maximum, coinciding with the deep chlorophyll maximum (DCM) established near the base of the euphotic zone of the Rhodes cyclone and its periphery, where the nutricline was situated just below the euphotic zone for most of the year. Moreover, the POM peaks were broader and situated at shallower depths in late winter–early spring as compared to its position in the summer–autumn period. Under prolonged winter conditions, as experienced in March 1992, the characteristic POM feature disappeared in the center of the Rhodes cyclone, where the upper layer was entirely occupied by nutrient-rich Levantine deep water. Deep convective processes in the cyclonic gyre led to the formation of vertically uniform POM profiles with low concentrations of particulate organic carbon (POC) (2.1 μM), nitrogen (0.21 μM), total particulate phosphorus (PP) (0.02 μM) and chlorophyll-a (0.5 μg/L) in the euphotic zone. Though the Levantine deep waters ascended up to the surface layer with the nitrate/phosphate molar ratios (28–29) in March 1992, the N/P molar ratio of bulk POM in the upper layer was low as 10–12, indicating luxury consumption of phosphate during algal production. Depth-integrated primary production in the euphotic zone ranged from 38.5 for oligotrophic autumn to 457 mg C m⁻² day⁻¹ for moderately mesotrophic cool winter conditions.     

Modelling the silica pump in the Permanently Open Ocean Zone of the Southern Ocean

A coupled 1D physical–biogeochemical model has been built to simulate the cycles of silicon and of nitrogen in the Indian sector of the Permanently Open Ocean Zone of the Southern Ocean. Based on a simplified trophic network, that includes two size classes of phytoplankton and of zooplankton, and a microbial loop, it has been calibrated by reference to surface physical, chemical and biological data sets collected at the KERFIX time-series station (50°40′S–68°25′E). The model correctly reproduces the high nutrient low chlorophyll features typical of the studied area. In a region where the spring–summer mixed layer depth is usually deeper than 60 m, the maximum of chlorophyll never exceeds 1.5 mg m⁻³, and the annual primary production is only 68 g C m⁻² year⁻¹. In the surface layer nitrate is never exhausted (range 27–23.5 mmoles m⁻³) while silicic acid shows strong seasonal variations (range 5–20 mmoles m⁻³). On an annual basis 71% of the primary production sustained by nanophytoplankton is grazed by microzooplankton. Compared to North Atlantic, siliceous microphytoplankton is mainly prevented from blooming because of an unfavourable spring–summer light-mixing regime. Silicic acid limitation (high half saturation constant for Si uptake: 8 mmoles m⁻³) also plays a major role on diatom growth. Mesozooplankton grazing pressure excerpts its influence especially in late spring. The model illustrates the efficiency of the silica pump in the Southern Ocean: up to 63% of the biogenic silica that has been synthetized in the photic layer is exported towards the deep ocean, while only 11% of the particulate organic nitrogen escapes recycling in the surface layer.     

Silicon dynamics in the Oder estuary, Baltic Sea

Studies on dissolved silicate (DSi) and biogenic silica (BSi) dynamics were carried out in the Oder estuary, Baltic Sea in 2000–2005. The Oder estuary proved to be an important component of the Oder River–Baltic Sea continuum where very intensive seasonal DSi uptake during spring and autumn, but also BSi regeneration during summer take place. Owing to the regeneration process annual DSi patterns in the river and the estuary distinctly differed; the annual patterns of DSi in the estuary showed two maxima and two minima in contrast to one maximum- and one minimum-pattern in the Oder River. DSi concentrations in the river and in the estuary were highest in winter (200–250 μmol dm^− 3) and lowest (often less than 1 μmol dm^− 3) in spring, concomitant with diatom growth; such low values are known to be limiting for new diatom growth. Secondary DSi summer peaks at the estuary exit exceeded 100 μmol dm^− 3, and these maxima were followed by autumn minima coinciding with the autumn diatom bloom. Seasonal peaks in BSi concentrations (ca. 100 μmol dm^− 3) occurred during the spring diatom bloom in the Oder River. Mass balance calculations of DSi and BSi showed that DSi + BSi import to the estuary over a two year period was 103.2 kt and that can be compared with the DSi export of 98.5 kt. The difference between these numbers gives room for ca. 2.5 kt BSi to be annually exported to the Baltic Sea. Sediment cores studies point to BSi annual accumulation on the level of 2.5 kt BSi. BSi import to the estuary is on the level of ca. 10.5 kt, thus ca. 5 kt of BSi is annually converted into the DSi, increasing the pool of DSi that leaves the system. BSi concentrations being ca. 2 times higher at the estuary entrance than at its exit remain in a good agreement with the DSi and BSi budgeting presented in the paper.     

100-years-changes in the phytoplankton community of Kiel Bight (Baltic Sea)

Literature data from 1905/06, 1912/13 and 1949/50 were compared with recent data (2001–2003) from Kiel Bight in order to investigate changes in phytoplankton composition and biomass, which may serve as indicators of environmental changes. In terms of biomass, diatomophyceae and dinophyceae are by far the most important groups. Their ratio is still close to unity. The share of diatomophyceae increased strongly in years with exceptionally high summer blooms (2001) or exceptionally early spring blooms (2003). The summer and autumn blooms of Chaetoceros and Skeletonema, detected in the early 20th century, are replaced by other diatoms (Cerataulina pelagica, Dactyliosolen fragilissimus, Proboscia alata, Pseudo-nitzschia spp.). Chaetoceros and Skeletonema are still important components of the spring blooms. Now as before, the autumn blooms are dominated by Ceratium spp., sometimes also by diatoms. Newly appearing bloom-forming species are mostly potentially toxic (Dictyocha speculum, Prorocentrum minimum, Pseudo-nitzschia spp.). The total phytoplankton biomass has roughly doubled in the course of the last century. The reference condition for phytoplankton biomass in Kiel Bight in the sense of the Water Framework Directive was defined at 55 mg C m^− 3 (± 10%, annual mean). The mean annual biomass of diatomophyceae and dinophyceae was 25 mg C m^− 3 (± 40%) for each, indicating that the sum of their carbon biomass amounted to 90% (± 10%) of the total phytoplankton biomass on an annual average. Diatomophyceae represented at least 80% of carbon biomass in the spring bloom peak at the beginning of the 20th century.     

Particle flux, and composition of sedimenting matter, in the Greenland Sea

Vertical flux of particulate material was recorded with moored sediment traps during 1988/1989 in the Greenland Sea at 72°N, 10°W. This region exhibits pronounced seasonal variability in ice cover. Annual fluxes at 500 m water depth were 22. 79, 8.55, 2.39, 3.81 and 0.51 g m⁻² for total flux (dry weight), carbonate particulate biogenic silicate, particulate organic carbon and nitrogen, respectively. Fluxes increased in April, maximum rates of all compounds occurred in May–June, and consistently high total flux rates of around 100 mg m⁻²d⁻¹ prevailed the summer. The increasing flux of biogenic particles measured in April is indicative of an early onset of algal growth in spring. Small pennate diatoms dominated in the trap collections during April, and were still numerous during the high flux period when Thalassiosira species were the most abundant diatoms. During May–June, up to 22% of the Thalassiosira cells collected were viable-looking cells. The faecal pellet flux increased after the May–June event. Therefore we conclude that the diatoms settled as phytodetritus, most likely in rapidly sinking aggregates. From seasonal nutrient profiles it is concluded that diatoms contribute 25% to new production during spring and 50% on an annual basis. More than 50% of newly produced silicate particles are dissolved above the 500 m horizon. High new production during spring does not lead to a pronounced sedimentation pulse of organic matter during spring but elevated vertical export is observed during the entire growth period.     

Microphytobenthos biomass at Kerguelen's Land (Subantarctic Indian Ocean): repartition and variability during austral summers

Microphytobenthos biomass has been measured at several coastal sites on the SE of the main island of the Kerguelen Archipelago (Indian Ocean), during several austral summers (1985–1992), using a conventional fluorometric method. Heterogeneity tests, conducted on two different intertidal sites (Port-Aux-Français, PAF; and Port-Raymond, PRA), showed low standard deviations, whereas the mean concentrations were highly different. Pigment concentrations showed a high variability related to the characteristics of the sediments: from low biomass in coarse intertidal sand, submitted to intense scouring (0.32±0.31 μg Chl a g⁻¹ dw, 0.29±0.14 μg Phaeo g⁻¹ dw) to high biomass in intertidal muddy sand in sheltered areas, particularly along estuaries (54 μg Chl a, 15 μg Phaeo g⁻¹ dw at Korrigan). The subtidal muddy sediments under a Macrocystis pyrifera (Linné) and Durvillaea antarctica (Chamisso in Choris) belt exhibited high concentrations in phaeopigment (Phaeo) (up to 136±83 μg g⁻¹ dw; PRA), while the concentration of chlorophyll a (Chl a) was relatively low. The dense macroalgal canopy supports an important epiphytic diatom biomass (mainly the genera Cocconeis Ehrenberg and Grammatophora Ehrenberg), which is sedimenting after degradation and is in part responsible of the high levels of Phaeo in all sediments. Macroalgal debris were observed, but diatom frustules were dominant in most surficial subtidal sediments. A circatidal mud, in the Morbihan Bay, made of a sponge spicule mat (50 m deep; 4.96 μg Chl a g⁻¹ dw), showed a very low Chl a/Phaeo ratio (0.1), while it reached up to 6 in intertidal sand. Surprisingly, a penguin rookery beach, at the east side of Courbet Peninsula, was characterized by a very low biomass (0.07±0.04 μg Chl a g⁻¹ dw), while it was nutrient enriched, particularly with nitrates.In comparison with the data at the similar latitude, but in temperate regions from the Northern Hemisphere, the microphytobenthos biomass, recorded at Kerguelen's Land, exhibited relatively high pigment concentrations, particularly the Phaeo, and supported a dense and diversified subtidal macrofauna composed of polychaetes (particularly Thelepus extensus Hutchings and Glasby), sea urchins, mytillids and gammarids. The exuberant macroalgal canopy, coastal indentations and low tidal amplitude must be in part responsible of these large benthic primary and secondary biomasses.     

Seasonality of picophytoplankton chlorophyll a and biomass in the central Cantabrian Sea, southern Bay of Biscay

Seasonal changes in the abundance and biomass of cyanobacteria (Synechococcus and Prochlorococcus) and picoeukaryotes were studied by flow cytometry in the upper layers of the central Cantabrian Sea continental shelf, from April 2002 to April 2006. The study area displayed the typical hydrographic conditions of temperate coastal zones. A marked seasonality of the relative contribution of prokaryotes and eukaryotes was found. While cyanobacteria were generally more abundant for most of the year (up to 2.4 10⁵ cells mL^− 1), picoeukaryotes dominated the community (up to 10⁴ cells mL^− 1) from February to May. The disappearance of Prochlorococcus from spring through summer is likely related to shifts in the prevailing current regime. The maximum total abundance of picophytoplankton was consistently found in late summer–early autumn. Mean photic-layer picoplanktonic chlorophyll a ranged from 0.06 to 0.53 µg L^− 1 with a relatively high mean contribution to total values (33 ± 2% SE), showing maxima around autumn and minima in spring. Biomass (range 0.58–40.16 mg C m^− 3) was generally dominated by picoeukaryotes (mean ± SE, 4.28 ± 0.27 mg C m^− 3) with an average contribution of cyanobacteria of 30 ± 2%. Different seasonality of pigment and biomass values resulted in a clear temporal pattern of picophytoplanktonic carbon to chlorophyll a ratio, which ranged from 10 (winter) to 140 (summer). This study highlights the important contribution of picoplanktonic chlorophyll a and carbon biomass in this coastal ecosystem.     

Circulation, hydrographic and nutrient characteristics of the Cilician Basin, Northeastern Mediterranean Sea

The water mass, circulation and chemical properties of the Cilician Basin, the northeastern Levantine Sea, are described on the basis of three hydrographic cruises performed during May 1997 (spring), July 1998 (summer) and October 2003 (autumn). The hydrographic data reveal the presence of Levantine Surface Water (LSW) and Modified Atlantic Water (MAW) within the upper 90 m layer, Levantine Intermediate Water (LIW) between 90 and 250 m, and Transitional Mediterranean Water (TMW) further below. The temporal variability of the circulation system is manifested by a change in shape, size and intensity of eddies as well as the pathways of the Lattakia Basin coastal current system. The nutrient concentrations varied between nitrate + nitrite = 0.16–0.31 μM, phosphate = 0.02–0.03 μM and silicate = 0.95–1.2 μM for the surface layer during sampling periods. Dissolved nutrient concentrations in the Transitional Mediterranean Water were: 2.1–5.3 μM for NO₃ + NO₂, 0.10–0.21 μM for PO₄ and 5.7–10 μM for Si. The molar ratios of nitrate to phosphate in the water column range between 5 and 20 in the surface layer and reach up to a value of 45 at the top of the nutricline at the depths of 29.05 kg/m³ isopycnal surface for most of the year. Below the nutricline the N / P ratios retain the values around 24–28.     

Microbial community structure in the marginal ice zone of the Bellingshausen Sea

Phytoplankton, bacteria and microzooplankton were investigated on a transect in the Bellingshausen Sea during the ice melt period in November–December 1992. The transect along the 85°W meridian comprised seven stations that progressed from solid pack-ice (70°S), through melting ice into open water (67°S). The abundance, biomass and taxonomic composition were determined for each component of the microbial community. The phytoplankton was mostly dominated by diatoms, particularly small (<20 μm) species. Diatom abundance ranged from 66 000 cells l⁻¹ under the ice to 410 000 cells l⁻¹ in open water. Phytoplankton biomass varied from <1 to 167 mg C m⁻³, with diatoms comprising 89–95% of the total biomass in open water and autotrophic nanoflagellates comprising 57% under the ice. The standing stocks of autotrophs in the mixed layer ranged from 95 mg C m⁻² under the pack-ice to 9478 mg C m⁻² in open waters. Bacterial abundance in ice-covered and open water stations varied from 1.1 to 5.5×10⁸ cells l⁻¹. Bacterial biomass ranged from 2.4 mg C m⁻³ under pack-ice to an average of 14 mg C m⁻³ in open water. The microzooplankton consisted mainly of aloricate oligotrich ciliates and heterotrophic dinoflagellates and these were most abundant in open waters. Their biomass varied between 0.2 and 54 mg C m⁻³ with a minimum at depth under the ice and maximum in open surface waters. Microheterotrophic standing stocks varied between 396 mg C m⁻² under pack-ice and 3677 mg C m⁻² in the open waters. The standing stocks of the total microbial community increased consistently from 491 mg C m⁻² at the ice station to 13 155 mg C m⁻² in open waters, reflecting the productive response of the community to ice-melt. The composition of the microbial community also shifted markedly from one dominated by heterotrophs (82% of microbial stocks) at the ice station to one dominated by autotrophs (73% of microbial stocks) in the open water. Our estimates suggest that the microbial community comprised >100% of the total particulate organic carbon (POC) under the ice and 62–66% of the measured POC in the open waters.     

Role of filtering and biodeposition by Adamussium colbecki in circulation of organic matter in Terra Nova Bay (Ross Sea, Antarctica)

At Terra Nova Bay, the scallop Adamussium colbecki (Smith, 1902) characterises the soft and hard bottoms from 20 to 80 m depth, constituting large beds and reaching high values of density (50–60 individuals/m²) and biomass (120 g/m² DW soft tissues). To assess its role in the organic matter recycling in the coastal ecosystem, its filtering and biodeposition rates were evaluated in laboratory experiments during the austral summer 1993/94. Filtration rates, measured in a flow-through system, were calculated from the difference in particulate organic carbon (POC), nitrogen (PON) and chlorophyll-a (Chl-a) concentration in inflow and outflow water. Experiments were performed using natural sea water with POC, PON and Chl-a concentrations of about 450 μg/l, 90 μg/l and 2 μg/l, respectively. The biodeposition rate and the biochemical composition of the biodeposits were studied in order to detect how the organic matter is transformed through feeding activity of A. colbecki. At +1°C temperature, the average filtering rate was about 1 l h⁻¹ g⁻¹ (DW soft tissues) in specimens ranging in body mass from 2 to 3 g (DW soft tissues) and 6–7 cm long. The biodeposition rate in 3–8 cm long specimens, ranging from 0.4 to 5.7 g (DW soft tissues), was about 5.65 mg DW/g DW/day, leading to an estimate of Corg flux, through biodeposition by A. colbecki, of about 21 mg C m⁻² day⁻¹ at in situ conditions. Comparison between the biochemical composition of seston and biodeposits shows a decrease of the labile compounds, of the Chl-a/phaeopigments ratio in the biodeposits. The recorded C/N ratio decrease suggests a microbial colonisation in the biodeposits. This study suggests that Adamussium colbecki plays an important role in coupling the material fluxes from the water column to the sea bed, processing about 14% of total Carbon flux from the water column to the sediments, with an assimilation efficiency of 36%.     

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.     

Microbial food web responses to light and nutrients beneath the coastal Arctic Ocean sea ice during the winter–spring transition

We measured the abundance and biomass of phototrophic and heterotrophic microbes in the upper mixed layer of the water column in ice-covered Franklin Bay, Beaufort Sea, Canada, from December 2003 to May 2004, and evaluated the influence of light and nutrients on these communities by way of a shipboard enrichment experiment. Bacterial cell concentrations showed no consistent trends throughout the sampling period, averaging (± SD) 2.4 (0.9) × 10⁸ cells L^− 1; integrated bacterial biomass for the upper mixed layer ranged from 1.33 mg C m^− 3 to 3.60 mg C m^− 3. Small cells numerically dominated the heterotrophic protist community in both winter and spring, but in terms of biomass, protists with a diameter > 10 µm generally dominated the standing stocks. Heterotrophic protist biomass integrated over the upper mixed layer ranged from 1.23 mg C m^− 3 to 6.56 mg C m^− 3. Phytoplankton biomass was low and variable, but persisted during the winter period. The standing stock of pigment-containing protists ranged from a minimum value of 0.38 mg C m^− 3 in winter to a maximal value of 6.09 mg C m^− 3 in spring and the most abundant taxa were Micromonas-like cells. These picoprasinophytes began to increase under the ice in February and their population size was positively correlated with surface irradiance. Despite the continuing presence of sea ice, phytoplankton biomass rose by more than an order of magnitude in the upper mixed layer by May. The shipboard experiment in April showed that this phototrophic increase in the community was not responsive to pulsed nutrient enrichment, with all treatments showing a strong growth response to improved irradiance conditions. Molecular (DGGE) and microscopic analyses indicated that most components of the eukaryotic community responded positively to the light treatment. These results show the persistence of a phototrophic inoculum throughout winter darkness, and the strong seasonal response by arctic microbial food webs to sub-ice irradiance in early spring.     

Sediment community biomass and respiration in the Northeast water polynya, Greenland: a numerical simulation of benthic lander and spade core data

Sediment community metabolism (oxygen demand) was measured in the Northeast Water (NEW) polynya off Greenland employing two methods: in situ benthic chambers deployed with a benthic (GOMEX) lander and shipboard laboratory Batch Micro-Incubation Chambers (BMICs) utilizing ‘cores’ recovered from USNEL box cores. The mean benthic respiration rate measured with the lander was 0.057 mM O₂ m⁻² h⁻¹ (n = 5); whereas the mean measured with the BMICs was 0.11 mM O₂ m⁻² h⁻¹ (n = 21; p < 0.01 that the means were the same). In terms of carbon fluxes (14 and 27 mg C m⁻² d⁻¹), these respiration rates represent ca. 5–15% of the average net primary production measured in the euphotic zone in 1992. The biomass of the bacteria, meiofauna and macrofauna were measured at each location to quantify the relationship between total community respiration and total community biomass (mean 1.42 g C m⁻²). Average carbon residence time in the biota, calculated by dividing the biomass by the respiration, was on the order of 50–100 days, which is comparable to relatively oligotrophic continental margins at temperate latitudes.The biomass and respiration data for the aerobic heterotrophic bacteria, the infaunal invertebrates (meiofauna and macrofauna), and the epifaunal megabenthos (two species of brittle stars) are summarized in a ‘steady-state’ solution of a sediment food chain model, in terms of carbon. This carbon budget illustrates the relative importance of the sediment-dwelling invertebrates in the benthic subsystem, compared to the bacteria and the epibenthos, during the summer open-water period in mud-lined troughs at depths of about 300 m. The input needed to drive heterotrophic respiratory processes was within the range of the input of organic matter recorded in moored, time-sequencing sediment traps.A time-dependent numerical simulation of the model was run to investigate the potential responses of the three size groups of benthos to abrupt seasonal pulses of particulate organic matter. The model suggests that there is a time lag in the increase in bottom community biomass and respiration following the POC pulse, and provides hypothetical estimates for the potential carbon storage in the summer (open water), followed by catabolic losses during each ensuing winter (ice covered).This sequence of storage and respiration may contribute to the process of seasonal CO₂ ‘rectification’ (sensu Yager et al., 1995) in some Arctic ecosystems.     

Production of Acartia omorii (Copepoda: Calanoida) in Ilkwang Bay, southeastern coast of Korea

Production of the marine calanoid copepod Acartia omorii was measured from 2 October 1991 to 8 October 1992 at a station in Ilkwang Bay on the southeastern coast of Korea. A. omorii (nauplii + copepodites + adults) were present in the plankton throughout the year, with seasonal variation in abundance. Biomass of A. omorii was averaged at 0.44 mgC m^− 3, with peaks in February and July, and relatively low biomass in late summer and fall. Egg production rate ranged from 2.4 to 151.9 μgC m^− 3 day^− 1, which was equivalent to 95–6075 eggs m^− 3 day^− 1. Fecundity of an adult female was averaged at 38 eggs female^− 1 day^− 1. Instantaneous growth rates of copepodites were higher than those of nauplii stages. Annual production of A. omorii ranged from 33.5 mgC m^− 3 year^− 1 to 221 mgC m^− 2 year^− 1, showing a seasonal variation of daily production rate with peaks in February and July. The daily production rate of A. omorii was significantly correlated with chlorophyll a concentration. These results suggest that standing stocks and/or productivity of phytoplankton are the major influencing factors, rather than water temperature for the seasonal variation of production of A. omorii in Ilkwang Bay.     

The trophic role of the tunicate Salpa thompsoni in the Antarctic marine ecosystem

During a repeat grid survey and drogue study carried out in austral summer 1994/95, the abundance and feeding activity of salps were estimated in the Lazarev Sea region from net tows and in situ measurements of gut fluorescence. Throughout the survey area, Salpa thompsoni accounted for >95% of the total salp stock while Ihlea racovitzai was consistently represented in very low abundances. Maximum densities of S. thompsoni, with ≈4000 ind. 1000 m⁻³, were recorded in the Marginal Ice Zone (MIZ) in December when chlorophyll-a concentrations were well below 1 mg m⁻³. A dramatic decrease in salp stock was observed at the beginning of January, when S. thompsoni virtually disappeared from the most productive area of the MIZ where chlorophyll-a concentrations had by then reached bloom levels of 1.5–3 mg (Chl-a) m⁻³. In situ grazing measurements showed that throughout the cruise S. thompsoni exhibited the highest ingestion rates per individual of any of the most abundant components of the grazing pelagic community, with maxima of ≈160 μg (pigm) ind. ⁻¹ d⁻¹. These feeding rates are 3 to 5 times higher than those previously obtained using in vitro incubations. The total daily consumption of the population of S. thompsoni varied from 0.3 to 108% of daily primary production. We suggest that competitive removal of food by S. thompsoni, rather than direct predation, is responsible for the low krill abundances generally associated with salp swarms.