Past, present and future state of the biogeochemical Si cycle in the Baltic Sea

The Baltic Sea is one of many aquatic ecosystems that show long-term declines in dissolved silicate (DSi) concentrations due to anthropogenic alteration of the biogeochemical Si cycle. Reductions in DSi in aquatic ecosystems have been coupled to hydrological regulation reducing inputs, but also with eutrophication, although the relative significance of both processes remains unknown for the observed reductions in DSi concentrations. Here we combine present and historical data on water column DSi concentrations, together with estimates of present river DSi loads to the Baltic, the load prior to damming together with estimates of the long-term accumulation of BSi in sediments. In addition, a model has been used to evaluate the past, present and future state of the biogeochemical Si cycle in the Baltic Sea. The present day DSi load to the Baltic Sea is 855 ktons y^− 1. Hydrological regulation and eutrophication of inland waters can account for a reduction of 420 ktons y^− 1 less riverine DSi entering the Baltic Sea today. Using published data on basin-wide accumulation rates we estimate that 1074 ktons y^− 1 of biogenic silica (BSi) is accumulating in the sediments, which is 36% higher than earlier estimates from the literature (791 ktons y^− 1). The difference is largely due to the high reported sedimentation rates in the Bothnian Sea and the Bothnian Bay. Using river DSi loads and estimated BSi accumulation, our model was not able to estimate water column DSi concentrations as burial estimates exceeded DSi inputs. The model was then used to estimate the BSi burial from measured DSi concentrations and DSi load. The model estimate for the total burial of BSi in all three basins was 620 ktons y^− 1, 74% less than estimated from sedimentation rates and sediment BSi concentrations. The model predicted 20% less BSi accumulation in the Baltic Proper and 10% less in the Bothnian Bay than estimated, but with significantly less BSi accumulation in the Bothnian Sea by a factor of 3. The model suggests there is an overestimation of basin-wide sedimentation rates in the Bothnian Bay and the Bothnian Sea. In the Baltic Proper, modelling shows that historical DSi concentrations were 2.6 times higher at the turn of the last century (ca. 1900) than at present. Although the DSi decrease has leveled out and at present there are only restricted areas of the Baltic Sea with limiting DSi concentrations, further declines in DSi concentrations will lead to widespread DSi limitation of diatoms with severe implications for the food web.     

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.     

Diatom stratigraphy and long-term dissolved silica concentrations in the Baltic Sea

In many parts of the world coastal waters with anthropogenic eutrophication have experienced a gradual depletion of dissolved silica (DSi) stocks. This could put pressure on spring bloom diatom populations, e.g. by limiting the intensity of blooms or by causing shifts in species composition. In addition, eutrophication driven enhanced diatom growth is responsible for the redistribution of DSi from the water phase to the sediments, and changes in the growth conditions may be reflected in the sediment diatom stratigraphy.To test for changes in diatom communities we have analyzed four sediment cores from the Baltic Sea covering approximately the last 100 years. The sediment cores originate from the western Gulf of Finland, the Kattegat, the Baltic Proper and the Gulf of Riga. Three out of the four cores reveal only minor changes in composition of diatom assemblages, while the Gulf of Riga core contains major changes, occurring after the second World War. This area is set apart from the other Baltic Sea basins by a high frequency of low after spring bloom DSi concentrations (< 2 µmol L^− 1) during a relatively well defined time period from 1991–1998. In 1991 to 1993 a rapid decline of DSi spring concentrations and winter stocks (down to 5 µmol L^− 1) in the Gulf was preceded by exceptionally intense diatom spring blooms dominated by the heavily silicified species Thalassiosira baltica (1991–1992; up to 5.5 mg ww L^− 1). T. baltica has been the principal spring bloom diatom in the Gulf of Riga since records began in 1975. DSi consumption and biomass yield experiments with cultured T. baltica suggest that intense blooms can potentially exhaust the DSi stock of the water column and exceed the annual Si dissolution in the Gulf of Riga. The phytoplankton time series reveals another exceptional T. baltica bloom period in 1981–1983 (up to 8 mg L^− 1), which, however, took place before the regular DSi measurements. These periods may be reflected in the conspicuous accumulation of T. baltica frustules in the sediment core corresponding to ca. 1975–1985.     

Th-derived particulate organic carbon fluxes in the northern Barents Sea with comparison to drifting sediment trap fluxes

Large-volume sampling of ²³⁴Th was conducted to estimate particulate organic carbon (POC) export in conjunction with drifting sediment trap deployments in the northern Barents Sea in July 2003 and May 2005. ²³⁴Th-derived POC fluxes averaged 42.3 ± 39.7 mmol C m^− 2 d^− 1 in 2003 and 47.1 ± 30.6 mmol C m^− 2 d^− 1 in 2005. Sediment trap POC fluxes averaged 13.1 ± 8.2 mmol C m^− 2 d^− 1 in 2003 and 17.3 ± 11.4 mmol C m^− 2 d^− 1 in 2005, but better reflected the transient bloom conditions that were observed at each station within a season. Although ²³⁴Th fluxes agreed within a factor 2 at most stations and depths sampled, sediment trap POC fluxes were lower than large-volume POC flux estimates at almost every station. This may represent an under-collection of POC by the drifting sediment traps or, conversely, an over-collection of POC by the large-volume sampling of ²³⁴Th. It is hypothesized that the offset between the two methods is partly due to the presence of the prymnesiophyte Phaeocystis pouchetii, which potentially causes a large variation in > 53-μm POC/²³⁴Th ratios. Due to the large proportion of dissolved carbon or mucilage released by P. pouchetii, and because it is thought that P. pouchetii does not contribute significantly to the vertical export of biogenic matter in the Barents Sea, the application of large-volume sampling of ²³⁴Th may yield relatively high, and possibly inaccurate POC/²³⁴Th ratios. Hence, POC fluxes derived from ²³⁴Th sampling may be inappropriate and drifting sediment traps might be a more reliable method to measure the vertical export of biogenic matter in regions that have recurrent P. pouchetii blooms, such as the Barents Sea.     

Present-day biosiliceous sedimentation in the northwestern Ross Sea, Antarctica

Sedimentological and oceanographic inferences have been obtained for the NW Ross Sea using sedimentary ²¹⁰Pb as a tracer together with determinations of biogenic silica and organic carbon. ²¹⁰Pb chronologies give apparent accumulation rates ranging between 14 and 80 mg cm⁻² yr⁻¹ (0.02–0.12 cm yr⁻¹) in the shelf basins. Even if a profile of ²¹⁰Pb is present in sediments from the top of the banks, here sediment accumulation rate is practically null, and physical mixing is responsible for the downward transport of fine particles and associated ²¹⁰Pb. The accuracy of ²¹⁰Pb-derived accumulation rates is discussed with respect to ¹⁴C dates. The annual rate of biogenic accumulation from ²¹⁰Pb appears to be ca. 8 times higher than the value derived using radiocarbon. Bioturbation is probably responsible for the discrepancy but also temporal and spatial variations in opal accumulation play a key-role. Contrasting measured and expected inventories of ²¹⁰Pb, a residence time of about 50 years has been tentatively estimated for the water in the NW Ross Sea. Furthermore, the data suggest that the pattern of present-day biosiliceous sediment accumulation in the Ross Sea is mainly driven by biogenic silica production in the water column, the SW area being the most productive part of the Ross Sea, by high sediment accumulation rate which enhances the seabed preservation, and by hydrodynamics, which is so effective to resuspend fine biogenic particles from the topographic highs. Resuspended particles are then deposited onto the flanks. The material which accumulates in the central part of the basins derives basically from primary production and settling through the water column.     

Pore water phosphate and ammonia below the permanent halocline in the south-eastern Baltic Sea and their benthic fluxes under anoxic conditions

In this paper the results of a study on the distribution of pore water phosphates and ammonia, and their fluxes under anoxic condition in a deep (> 70 m) accumulation-type bottom of the south-eastern Baltic Sea, namely in the Gdańsk Deep and the adjacent areas, are presented. All measurements were taken during the growth period, i.e. in September 2000, April 2001 and June 2002. Benthic phosphate and ammonia fluxes were estimated using Fick's First Law. Phosphate and ammonia concentrations ranged from 7.5 to 266.3 μmol dm^− 3 and from 53.6 to 1248.3 μmol dm^− 3, respectively. The values recorded in the central part of the Gdańsk Deep were lower than those found both on its slopes and on the SW slope of the Gotland Deep. The lowest phosphate contents were typical of the Oblique Sill which separates the Gdańsk and Gotland Deeps.In 1993–2002, as a result of anoxia the sediments in the Gdańsk Deep released about 5.1 × 10³ t P and 22.8 × 10³ t N. These loads supplied on average 1.5% and 0.9% of phytoplankton's demand for P and N, respectively. In comparison to the total external load of nutrients discharged to the Gulf of Gdańsk (i.e. 8.79 × 10³ t year^− 1 P_tot and 130.79 × 10³ t year^− 1 N_tot; [Witek, Z., Humborg, Ch., Savchuk, O., Grelowski, A. and Łysiak-Pastuszak, E., 2003. Nitrogen and phosphorus budgets of the Gulf of Gdańsk (Baltic Sea). Est. Coast. Shelf Sci., 57:239–248.]), the return flux of P and N from the anoxic sediments to the water column in the Gdańsk Deep was a minor source of these elements.     

The contribution of total suspended solids to the Bay of Biscay by Cantabrian Rivers (northern coast of the Iberian Peninsula)

Information of suspended sediments fluxes of small rivers to the coastal zone is sparse, and this is particularly so for the Iberian Rivers. To help address this shortage of information, the relationship between fluvial discharge and total suspended solids (TSS) for the main 28 Cantabrian Rivers using data from 22 years monitoring by the COCA network has been analysed, and their particulate material fluxes to the Bay of Biscay coasts have been quantified. The Cantabrian Fluvial System (drainage basin area of 20,333 km²) may be considered as a quasi-homogeneous fluvial system with an average discharge of 561 m^− 3 s^− 1 and average loads of 35 kg_TSS s^− 1 with rivers showing similar average yields of 56 t km^− 2 a^− 1. The average TSS contribution is 1.2 ± 0.2 10⁹ kg a^− 1. This seaward flux of sediment is dispersed along the entire North Iberian coast and is rather modest (25% of the total supply) in comparison with the output from the French Rivers to the Bay of Biscay. The TSS loads of Cantabrian Rivers indicate they are similar to world upland rivers and those of other parts of Northern Europe according to Milliman and Syvistki [Milliman and Syvistki, 1992. Geomorphic/tectonic control of sediment discharge to the ocean: the importance of small mountainous rivers. Journal of Geology, 100: 525–544] and Milliman [Milliman, 2001. Delivery and fate of fluvial water and sediment to the sea: a marine geologist's view of European rivers. Scientia Marina, 65: 121–132]. Although their TSS flux is practically negligible (13,000 times lower) when compared to the world average flux, they provide a good example of the role of small Atlantic temperate rivers.     

Biogeochemistry of sulfur in a sediment core from the west-central Baltic Sea: Evidence from stable isotopes and pyrite textures

The biogeochemistry of the sulfur cycle in a ca. 5-m-long sediment core from the eastern slope (221 m water depth) of the Landsort Deep in the west-central Baltic Sea was investigated by analyzing the solid phase records of sulfur isotopes and pyrite textures, besides selected main and minor elements. The sediments were deposited during post-glacial history of the Baltic Sea when the basin experienced alteration of brackish (Yoldia Sea, Littorina Sea) and freshwater (Baltic Ice Lake, Ancylus Lake) conditions. The stable isotopic composition of total sulfur was analyzed as a function of depth. In selected samples pyrite (FeS₂), greigite (Fe₃S₄), and barite (BaSO₄) fractions were separated for isotope analyses. Pyrite textures were analyzed by SEM and optical microscopy.Microbial reactions associated with the oxidation of organic matter resulted in assemblages of authigenic sulfide minerals which for the post-Ancylus Lake brackish water environment are dominated by pyrite and for freshwater environments by Fe-monosulfides. The sulfur isotopic composition of the brackish water Littorina Sea sediments (δ³⁴S between −40 and −27‰ vs. V-CDT) is believed to be determined by cellular sulfate reduction rates and reactions involving intermediate sulfur species. The availability of reactive iron and decomposable organic matter as well as sedimentation rate and the chemocline position are important variables upon the δ³⁴S values of sulfides in brackish water environment. The syn-depositional abundance of sulfur and organic matter, and transport of dissolved sulfur species vs. rates of microbial reactions determine δ³⁴S in the freshwater sediments. The upper part of the Ancylus Lake sediments is sulfidized by downward diffusing H₂S and/or sulfate from overlying brackish water sediments. Minor concretionary barite formation in the freshwater sediments is most likely due to the reaction of pore water sulfate diffusing downward from brackish water sediments with barium desorbed from freshwater sediments. The size distribution of pyrite framboids in the brackish sediments indicates that the formation mainly occurred from anoxic pore waters, although some pyrite formation in an anoxic water column cannot be excluded.     

Organic carbon budget for the Gulf of Bothnia

We calculated input of organic carbon to the unproductive, brackish water basin of the Gulf of Bothnia from rivers, point sources and the atmosphere. We also calculated the net exchange of organic carbon between the Gulf of Bothnia and the adjacent marine system, the Baltic Proper. We compared the input with sinks for organic carbon; permanent incorporation in sediments and mineralization and subsequent evasion of CO₂ to the atmosphere. The major fluxes were riverine input (1500 Gg C year^− 1), exchange with the Baltic Proper (depending on which of several possible DOC concentration differences between the basins that was used in the calculation, the flux varied between an outflow of 466 and an input of 950 Gg C year^− 1), sediment burial (1100 Gg C year^− 1) and evasion to the atmosphere (3610 Gg C year^− 1). The largest single net flux was the emission of CO₂ to the atmosphere, mainly caused by bacterial mineralization of organic carbon. Input and output did not match in our budget which we ascribe uncertainties in the calculation of the exchange of organic carbon between the Gulf of Bothnia and the Baltic Proper, and the fact that CO₂ emission, which in our calculation represented 1 year (2002) may have been overestimated in comparison with long-term means. We conclude that net heterotrophy of the Gulf of Bothnia was due to input of organic carbon from both the catchment and from the Baltic Proper and that the future degree of net heterotrophy will be sensible to both catchment export of organic carbon and to the ongoing eutrophication of the Baltic Proper.     

Decadal-scale variations of trophic levels at high trophic levels in the Yellow Sea and the Bohai Sea ecosystem

A total of 2759 stomachs collected from a bottom trawl survey carried out by R/V “Bei Dou” in the Yellow Sea between 32°00 and 36°30N in autumn 2000 and spring 2001 were examined. The trophic levels (TL) of eight dominant fish species were calculated based on stomach contents, and trophic levels of 17 dominant species in the Yellow Sea and the Bohai Sea reported in later 1950s and mid-1980s were estimated so as to be comparable. The results indicated that the mean trophic level at high trophic levels declined from 4.06 in 1959–1960 to 3.41 in 1998–1999, or 0.16–0.19·decade^− 1 (mean 0.17·decade^− 1) in the Bohai Sea, and from 3.61 in 1985–1986 to 3.40 in 2000–2001, or 0.14·decade^− 1 in the Yellow Sea; all higher than global trend. The dominant species composition in the Yellow Sea and the Bohai Sea changed, with the percentage of planktivorous species increases and piscivorous or omnivorous species decreases, and this was one of the main reasons for the decline in mean trophic level at high tropic levels. Another main reason was intraspecific changes in TL. Similarly, many factors caused decline of trophic levels in the dominant fish species in the Yellow Sea and the Bohai Sea. Firstly, TL of the same prey got lower, and anchovy (Engraulis japonicus) as prey was most representative. Secondly, TLs of diet composition getting lower resulted in not only decline of trophic levels but also changed feeding habits of some species, such as spotted velvetfish (Erisphex pottii) and Trichiurus muticus in the Yellow Sea. Thirdly, species size getting smaller also resulted in not only decline of trophic levels but also changed feeding habits of some species, such as Bambay duck (Harpodon nehereus) and largehead hairtail (Trichiurus haumela). Furthermore, fishing pressure and climate change may be interfering to cause fishing down the food web in the China coastal ocean.     

Seasonal variability in sea surface oceanographic conditions in the Aegean Sea (Eastern Mediterranean): an overview

Seasonal variability and the spatial distribution of sea surface temperatures (SST) and salinities (SSS) are reviewed, in relation to the prevailing climatological conditions, heat fluxes, water budget and general water circulation patterns. Within this context, consideration is given to: sea surface temperatures; air temperatures; precipitation; evaporation; wind speeds and directions; freshwater (mainly riverine) discharges throughout the Aegean; and the exchange of water masses with the Black Sea and eastern Mediterranean Sea. The investigation of satellite images, covering a 6-yr period (1988–1994), has enabled a synthesis of the monthly sea surface thermal distribution to be established.The climate of the Aegean Sea is characterised by annual air temperatures of 16–19.5°C, precipitation of about 500 mm yr⁻¹ and evaporation of some 4 mm d⁻¹. The Aegean has a negative heat budget (approximately −25 W m⁻²) and positive water balance (+ 1.0 m yr⁻¹), when inflow from the Black Sea is considered. During the summer, the (northerly) Etesians are the dominant winds over the Sea.Mean monthly sea surface temperatures (SST) vary from 8°C in the north during winter, up to 26°C in the south during summer. SST depends mainly upon air temperature; there is a month's delay between the former and latter maxima. The sea surface salinity (SSS) varies also spatially and seasonally, ranging from less than 31 psu, in the north, to more than 39 psu, in the southeast; lower values (< 25 psu) occur adjacent to the river mouths. SSSs present their maximum differences during summer, whilst during winter and autumn the distribution of SSS is more uniform. The overall spatial SST and SSS distribution pattern is controlled by: distribution of the (colder) Black Sea Waters; advection of the (warmer) Levantine Waters, from the southeastern part of the Aegean; upwelling and downwelling; and, to a lesser extent, but locally important, freshwater riverine inflows.     

Physical and biological correlates of virus dynamics in the southern Beaufort Sea and Amundsen Gulf

As part of the Canadian Arctic Shelf Exchange Study (CASES), we investigated the spatial and seasonal distributions of viruses in relation to biotic (bacteria, chlorophyll-a (chl a)) and abiotic variables (temperature, salinity and depth). Sampling occurred in the southern Beaufort Sea Shelf in the region of the Amundsen Gulf and Mackenzie Shelf, between November 2003 and August 2004. Bacterial and viral abundances estimated by epifluorescence microscopy (EFM) and flow cytometry (FC) were highly correlated (r² = 0.89 and r² = 0.87, respectively), although estimates by EFM were slightly higher (FC = 1.08 × EFM + 0.12 and FC = 1.07 × EFM + 0.43, respectively). Viral abundances ranged from 0.13 × 10⁶ to 23 × 10⁶ ml^− 1, and in surface waters were ~ 2-fold higher during the spring bloom in May and June and ~ 1.5-fold higher during July and August, relative to winter abundances. These increases were coincident with a ~ 6-fold increase in chl a during spring and a ~ 4-fold increase in bacteria during summer. Surface viral abundances near the Mackenzie River were ~ 2-fold higher than in the Mackenzie Shelf and Amundsen Gulf regions during the peak summer discharge, concomitant with a ~ 5.5-fold increase in chl a (up to 2.4 μg l^− 1) and a ~ 2-fold increase in bacterial abundance (up to 22 × 10⁵ ml^− 1). Using FC, two subgroups of viruses and heterotrophic bacteria were defined. A low SYBR-green fluorescence virus subgroup (V2) representing ~ 71% of the total viral abundance, was linked to the abundance of high nucleic acid fluorescence (HNA) bacteria (a proxy for bacterial activity), which represented 42 to 72% of the bacteria in surface layers. A high SYBR-green fluorescence viral subgroup (V1) was more related to high chl a concentrations that occurred in surface waters during spring and at stations near the Mackenzie River plume during the summer discharge. These results suggest that V1 infect phytoplankton, while most V2 are bacteriophages. On the Beaufort Sea shelf, viral abundance displayed seasonal and spatial variations in conjunction with chl a concentration, bacterial abundance and composition, temperature, salinity and depth. The highly dynamic nature of viral abundance and its correlation with increases in chl a concentration and bacterial abundance implies that viruses are important agents of microbial mortality in Arctic shelf waters.     

A novel method for estimating vertical eddy diffusivities using diurnal signals with application to western Long Island Sound

We present an approach that allows the estimation of vertical eddy diffusivity coefficients from buoy measurements made at two or more depths. By measuring the attenuation and phase lag of a scalar signal generated periodically at the surface as it propagates downwards, the vertical eddy diffusivity coefficients can be calculated as K_v = ωΔz²/2ln²(α₂/α₁), where α₂/α₁ is the ratio of the real amplitudes at frequency ω at the two depths separated by Δz = z₂ − z₁; as K_V = ωΔz²/2φ², where φ is the phase lag at the frequency ω; or as K_v = iωΔz²/ln²(U₂/U₁), where U₂/U₁ is the ratio of the complex signal amplitudes at the two depths. The method requires that horizontal fluxes be small at the ω frequency and that the signal-to-noise ratios at the two depths allow the determination of the amplitude and phase of ω.Application of this method to summertime 2004 western Long Island Sound oxygen and temperature buoy measurements at two depths provides a time-series of two-day average vertical eddy diffusivity estimates. Using these eddy diffusivities in conjunction with measured vertical concentration gradients, we obtain a time-series of vertical transport rates for oxygen and heat and estimate mean downward fluxes for June and July as 150–260 mMol m^− 2 day^− 1 and 100–400 W m^− 2 respectively. These estimates are of a similar magnitude to sub-pycnocline O₂ and heat demands of 240 ± 200 mMol m^− 2 day^− 1 and 180 ± 60 W m^− 2 that we infer from simple budgets, implying that vertical transport is significant to both budgets.The eddy coefficients obtained from the independent O₂ and temperature measurements have a 68% correlation, and the O₂ flux estimates show a correlation of 41% to measured rates of change in bottom dissolved oxygen levels. Our results indicate that extended time-series of eddy diffusivity coefficients can be obtained from in situ buoy measurements and the method shows promise as a way to constrain the vertical transport variability in budgets of dissolved materials in estuaries.     

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.     

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.     

Protist entrapment in newly formed sea ice in the Coastal Arctic Ocean

Protist abundance and taxonomic composition were determined in four development stages of newly formed sea ice (new ice, nilas, young ice and thin first-year ice) and in the underlying surface waters of the Canadian Beaufort Sea from 30 September to 19 November 2003. Pico- and nanoalgae were counted by flow cytometry whereas photosynthetic and heterotrophic protists ≥ 4 µm were identified and counted by inverted microscopy. Protists were always present in sea ice and surface water samples throughout the study period. The most abundant protists in sea ice and surface waters were cells < 4 µm. They were less abundant in sea ice (418–3051 × 10³ cells L^− 1) than in surface waters (1393–5373 × 10³ cells L^− 1). In contrast, larger protists (≥ 4 µm) were more abundant in sea ice (59–821 × 10³ cells L^− 1) than in surface waters (22–256 × 10³ cells L^− 1). These results suggest a selective incorporation of larger cells into sea ice. The ≥ 4 µm protist assemblage was composed of a total number of 73 taxa, including 12 centric diatom species, 7 pennate diatoms, 11 dinoflagellates and 16 flagellates. The taxonomic composition in the early stage of ice formation (i.e., new ice) was very similar to that observed in surface waters and was composed of a mixed population of nanoflagellates (Prasinophyceae and Prymnesiophyceae), diatoms (mainly Chaetoceros species) and dinoflagellates. In older stages of sea ice (i.e., young ice and thin first-year ice), the taxonomic composition became markedly different from that of the surface waters. These older ice samples contained relatively fewer Prasinophyceae and more unidentified nanoflagellates than the younger ice. Diatom resting spores and dinoflagellate cysts were generally more abundant in sea ice than in surface waters. However, further studies are needed to determine the importance of this winter survival strategy in Arctic sea ice. This study clearly shows the selective incorporation of large cells (≥ 4 µm) in newly formed sea ice and the change in the taxonomic composition of protists between sea ice and surface waters as the fall season progresses.     

Carbon flows in Baltic Sea food webs — a re-evaluation using a mass balance approach

The brackish Baltic Sea has been seen as particularly suitable for studies of food webs. Compared to fully marine ecosystems, it has low species diversity, which means fewer trophic linkages to analyse. The Baltic Sea is also one of the best-studied areas of the world, suggesting that most data requirements for food web models should be fulfilled. Nevertheless, the influence of physical and biological factors on trophic interactions and biogeochemical patterns varies spatially in the Baltic Sea, adding considerable complexity to food web studies. Food web structure and processes can be described and compared quantitatively between areas by estimating the flow of matter or energy through the organisms. Most such models have been based on carbon, though studies of complementary flows of other elements limiting production, such as nitrogen and phosphorus would be desirable. However, since ratios between carbon and other elements are used in calculating these flows, it is crucial, as a first step, to quantify the flows of carbon as accurately as possible.In this study, we used the EcopathII software (ver 3.1) to analyse models of carbon flow through the food webs in the three main areas of the Baltic Sea; the Baltic proper, Bothnian Sea and Bothnian Bay. A previously published study on carbon flow in the Baltic Sea [Elmgren, R. 1984. Trophic dynamics in the enclosed, brackish Baltic Sea. Rapp. P.-V. Reun. — Cons. Int. Explor. Mer. (183) 152–169.] was complemented with the data on respiration and flow to detritus [Wulff, F., Ulanowicz, R. 1989. A comparative anatomy of the Baltic Sea and Chesapeeake Bay ecosystems. In: F. Wulff, J.G. Field, K.H. Mann (Eds.), Flow Analysis of Marine Ecosystems: Theory and Practice. New York: Springer-Verlag.] in order to present complete mass balance models of carbon. The purpose of re-evaluating previous models with new analytic tools was to check how well their carbon flows balance, and to provide a basis for improved mass balance models using more recent data, including nutrients other than carbon.The resulting mass balance networks for the Baltic proper, Bothnian Sea and the Bothnian Bay were shown to deviate from steady state. There was an organic carbon surplus of 45, 25 and 18 g C m⁻² year⁻¹ in the pelagic zones of the Baltic proper, Bothnian Sea and Bothnian Bay, respectively. The Ecopath network analysis confirmed that the overall carbon flow was highest in the Baltic proper, somewhat lower in the Bothnian Sea and much lower in the Bothnian Bay. The only clear differences in food web structure between the basins was that the average trophic level was lower for demersal fish in the Bothnian Sea and higher for macrofauna in the Bothnian Bay, compared to the other basins. The analysis showed weakness in our current understanding in Baltic Sea food webs and highlighted areas where improvements could be made with more recent data.     

Estimation of TKE dissipation rates in dense bottom plumes using a Pulse Coherent Acoustic Doppler Profiler (PC-ADP) — Structure function approach

During a hydrographic survey in January 2006 the spreading of inflowing saline water was observed in the Arkona Basin (Western Baltic Sea). Two bottom mounted ‘pulse coherent’ acoustic Doppler profilers (PC-ADP) were used to measure the near-bottom current field of the dense plume with a high temporal (1 s) and spatial resolution (5 cm). In order to estimate the dissipation rate of turbulent kinetic energy () a structure function approach was applied to the beam velocity data. Simultaneous measurements with a microstructure shear profiler (MSS) and an acoustic Doppler velocimeter (ADV) supplied independent data for the verification of the structure function method. Additional measurements with standard CTD, near-bottom towed and vessel mounted acoustic Doppler current profilers (ADCP) completed the data set.The estimated dissipation rates from the structure function approach fit well with the values derived from the ADV and the MSS probe. It is shown that the structure function approach is a reliable and easily applicable method to derive estimates of TKE dissipation rates from PC-ADP beam velocities. The observed dissipation rates ranged between 5 · 10^− 6 and 1 · 10^− 8 W kg^− 1 depending on the hydrographic conditions. Inside the plume the dissipation rates exceeded that of the overlaying brackish water by two orders of magnitude. Since the noise level of velocity data in pulse coherent mode is considerably lower than in the Doppler mode the PC-ADP can also be used for estimates in marine environments with low turbulence level. Reynolds stresses estimated from the PC-ADP and the ADV agreed well at the same depth level. TKE production derived from PC-ADP measurements compared reasonably well with the dissipation rate of TKE in a varying environment.     

Methane release and coastal environment in the East Siberian Arctic shelf

In this paper we present 2 years of data obtained during the late summer period (September 2003 and September 2004) for the East Siberian Arctic shelf (ESAS). According to our data, the surface layer of shelf water was supersaturated up to 2500% relative to the present average atmospheric methane content of 1.85 ppm, pointing to the rivers as a strong source of dissolved methane which comes from watersheds which are underlain with permafrost. Anomalously high concentrations (up to 154 nM or 4400% supersaturation) of dissolved methane in the bottom layer of shelf water at a few sites suggest that the bottom layer is somehow affected by near-bottom sources. The net flux of methane from this area of the East Siberian Arctic shelf can reach up to 13.7 × 10⁴ g CH₄ km^− 2 from plume areas during the period of ice free water, and thus is in the upper range of the estimated global marine methane release. Ongoing environmental change might affect the methane marine cycle since significant changes in the thermal regime of bottom sediments within a few sites were registered. Correlation between calculated methane storage within the water column and both integrated salinity values (r = 0.61) and integrated values of dissolved inorganic carbon (DIC) (r = 0.62) suggest that higher concentrations of dissolved methane were mostly derived from the marine environment, likely due to in-situ production or release from decaying submarine gas hydrates deposits. The calculated late summer potential methane emissions tend to vary from year to year, reflecting most likely the effect of changing hydrological and meteorological conditions (temperature, wind) on the ESAS rather than riverine export of dissolved methane. We point out additional sources of methane in this region such as submarine taliks, ice complex retreat, submarine permafrost itself and decaying gas hydrates deposits.     

Sea ice and the onshore–offshore gradient in pre-winter zooplankton assemblages in southeastern Beaufort Sea

Zooplankton communities were studied in southeastern Beaufort Sea (Arctic Ocean) in September–October 2002. Cluster analysis and non-metric multidimensional scaling revealed three distinct mesozooplankton assemblages. A neritic assemblage occurred on the Mackenzie Shelf and in Franklin Bay, while distinct off-shelf assemblages prevailed in the Cape Bathurst Polynya and on the Beaufort Slope respectively. Over 95% of the mesozooplankton was comprised of eight copepod taxa. Pseudocalanus spp. contributed predominantly to the discrimination of the three assemblages and was the only significant indicator of the Shelf assemblage. Oithona similis, Oncaea borealis, Metridia longa and Calanus hyperboreus were indicators of the Polynya assemblage. Cyclopina sp. and Microcalanus pygmaeus were indicative of the overall off-shelf community (Polynya and Slope assemblages). The importance of omnivores and carnivores increased from the shelf to the polynya and the slope. Station depth and duration of reduced ice conditions during summer (< 50% ice concentration) underpinned the distribution of the assemblages (r² = 0.71 and 0.45 respectively). The abundance of Pseudocalanus spp. was independent of depth and increased with the duration of reduced ice conditions (r_s = 0.438). The abundance of Cyclopina sp., M. pygmaeus and other indicators of the offshore assemblages followed the opposite trend (r_s = − 0.467 and − 0.5 respectively). Under continued climate warming, a reduction of the ice cover will affect the biogeography of mesozooplankton on and around the Mackenzie Shelf, to the potential advantage of Pseudocalanus spp. and other calanoid herbivores.