Spatial distribution patterns of demersal and epibenthic communities on the Galician continental shelf (NW Spain)

Spatial distribution patterns of epibenthic communities on the Galician continental shelf were studied using multivariate methods. Data came from 5 surveys carried out in 2002, 2003 and 2004. Beam trawls and otter trawls were used to study epibenthic communities along 8 transects perpendicular to the coastline. The role of depth, season, latitude and sediment characteristics was examined. Seven habitats were described according to bathymetry and sediment characteristics. There were weak linear relationships between environmental variables and species richness, biomass and species diversity. However, the canonical analysis showed that depth and sediment characteristics greatly influence smaller epibenthic communities sampled by beam trawl. Six assemblages were obtained for beam trawls: inner shelf mud, very fine sands, and fine sands, middle shelf sands, and outer shelf very fine sands, and fine sands. Five assemblages were identified for larger-sized and swimming epibenthos sampled with otter trawls. These assemblages were also determined according to depth and sediment type but sediment characteristics were less important. Otter trawl assemblages were the same as the beam trawl ones, except for on the outer shelf where no differences between sediment type were detected. For both gears, inner and outer shelf assemblages displayed a higher biotic variability than the middle shelf, as a consequence of a higher environmental heterogeneity. Typifying species were mainly eurytopic in the middle shelf, whereas eurytopic and stenotopic species characterised the inner and outer shelves.     

The role of pelagic fish as forage for the demersal fish community in the southern Bay of Biscay

The small pelagic fish represents the link between nekton and demersal communities, and they are an important food source for some demersal fish species. In the autumns between 1993 and 2002, 74,869 stomach contents of 25 demersal fish species were analysed during the scientific surveys organized by the Instituto Español de Oceanografía. These species represent the demersal fish community of the southern Bay of Biscay. An important part of their diet (39% by volume) was composed of pelagic fish species (e.g. Engraulis encrasicolus, Gadiculus argenteus, Micromesistius poutassou, Trachurus trachurus, Sardina pilchardus, Scomber scombrus). Among all these prey species, the relevance of M. poutassou and G. argenteus stood out because of their high abundance both in the diets and during the scientific surveys. The relevance of pelagic fish as prey increased with predator size, reaching more than 60% of the diet by volume in the length range 25–29 cm. However, fish predators larger than 50 cm depended less on pelagic fish (33%), since they were also able to feed on other sources such as megafaunal invertebrates, mainly cephalopods. The pelagic fish resource was primarily exploited by 12 demersal fish species, with Merluccius merluccius and Zeus faber being the main ones feeding on small pelagics, which was related to the vertical movement of predators and prey through the water column. Survey abundance indices were used as indicators of prey abundance in the ecosystem, both for biomass and number. There was evidence for density-dependant feeding by predators on E. encrasicolus, G. argenteus and T. trachurus, while the main discrepancies between abundance in the stomachs and in the surveys were due to differential availability of prey length classes in the environment. Small prey species and individuals were exploited by most demersal fish while large prey species were less accessible to predators.     

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.     

Spatio-temporal variations in deep-sea demersal communities off the Balearic Islands (western Mediterranean)

The spatial and temporal variations of deep-sea megafaunal assemblages from the western Mediterranean are analysed in the present paper. The assemblages from two locations of the Balearic Islands situated 120 km apart were compared using data collected seasonally on a bathymetric stratum covering the 150–750 m depth range during six bottom-trawl surveys. The assemblage structure, in terms of species composition, species dominance and population sizes, was differentially affected by the spatio-temporal variables analysed (depth, location and fishing period). Although depth was the main factor determining the assemblage composition, the differences obtained between the two locations were also relevant. On the upper slope these between-location differences in the dynamics of megafaunal assemblages were found to be related to the effect of fishing exploitation. Population size-based metrics and biomass spectra were good predictors of meso-scale fishing effects, and were mainly reflected by elasmobranchs and demersal teleosts. Nevertheless, the effects of fishing depended on the species considered. Two dominant large-sized fish species found on the upper slope in both localities, Galeus melastomus and Phycis blennoides, had higher biomass values associated with lower fishing effort. Although the mean body weight (MBW) of both species and also the mean maximum body weight (MMBW) of G. melastomus agreed with this pattern, the P. blennoides MMBW did not. This last case could be indicative of natural size-trends such as the bigger–deeper phenomenon which refers to the displacement of large individuals towards the deeper limit of their bathymetric distribution, beyond the maximum depth sampled in this study for this species. By contrast, the target species of the upper slope fishery, the red shrimp Aristeus antennatus, was not negatively affected by the direct impact of fishing activity and other environmental factors, such as the presence of specific water masses could also be important.     

Biochemical composition of marine sediment from the eastern Weddell Sea (Antarctica): High nutritive value in a high benthic-biomass environment

Within the SCAR's international EASIZ programme, as part of the benthic–pelagic coupling experiment, grain size and organic matter contents in marine surface sediment were measured. Samples were taken during the austral autumn of 2000 from 3 regions in the eastern Weddell Sea: Kapp Norvegia, Four Seasons Bank, and Austasen.In general, sediments were fine sand with a grain size fraction < 200 μm representing more than 40% of the total weight. The sediments from Four Seasons Bank (64 to 107 m depth) were coarser than those from Austasen and Kapp Norvegia (209 to 480 m depth), presumably due to winnowing of fine sediment at shallow depths. Organic carbon (OC) content ranged from 0.25% to 1.2% and constituted 10% to 97% of the total carbon. The samples from Kapp Norvegia presented the highest OC values. Overall, protein (PRT), lipid (LPD), and carbohydrate (CHO) contents were similar to those in sediment from cold regions (e.g., the North Atlantic and the Ross Sea) but higher than those in sediment from other Antarctic and more septentrional regions (e.g., the Ross Sea and the Mediterranean). The difference within the Antarctic is explained through the local conditions in Terra Nova Bay and Kapp Norvegia. In the Antarctic, PRT and LPD carbon were the main contributors to the biopolymeric carbon (BPC). In the eastern Weddell Sea shelf, the BPC accounted for more than 90% of the OC in most of the samples. More than 82% of the total PRT, LPD, and CHO were present in the fraction < 200 μm. This work remarks the existence of sediments with a high nutritional value persistent several weeks after the spring–summer pulse of fresh organic matter. It is also highlighted the high potential availability of these sediments (due to its grain size) for the benthic communities inhabiting this high-latitude continental shelf.     

Thorium-234 derived information on particle residence times and sediment deposition in shallow waters of the south-western Baltic Sea

Activities of the naturally occurring, short-lived and highly particle-reactive radionuclide tracer ²³⁴Th in the dissolved and particulate phase were measured at three shallow-water stations (maximum water depths: 15.6, 22.7 and 30.1 m) in Mecklenburg Bay (south-western Baltic Sea) to constrain the time scales of the dynamics and the depositional fate of particulate matter. Activities of particle-associated (> 0.4 μm) and total (particulate + dissolved) ²³⁴Th were in the range of 0.08–0.11 dpm L^− 1 and 0.11–0.20 dpm L^− 1, respectively. The activity ratio of total ²³⁴Th and its long-lived and conservative parent nuclide ²³⁸U was well below unity (range: 0.09–0.19) indicating substantial radioactive disequilibria throughout the water column, very dynamic trace-metal scavenging and particle export from the water column at all three stations. For the discussion the ²³⁴Th data of this study were combined with previously published water-column ²³⁴Th and particulate-matter data from Mecklenburg Bay (Kersten et al., 1998. Applied Geochemistry 13, 339–347). The resulting average vertical distribution of total ²³⁴Th/²³⁸U disequilibria was used to estimate the depositional ²³⁴Th flux to the sediment. There was a virtually constant net downward flux of ²³⁴Th of about 28 dpm m^− 2 d^− 1 leaving each water layer of one meter thickness. Thorium-234-derived net residence times of particulate material regarding settling from a given layer in the water column were typically on the order of days, but with maximum values of up to a couple of weeks. Based on an average ratio of particulate matter (PM) to particle-associated ²³⁴Th a net flux of about 145 mg PM m^− 2 d^− 1 was estimated to leave each water layer of one meter thickness. The estimated cumulative water-column-derived particulate-matter fluxes at the seafloor are higher by a factor of about 2 than previously published sediment-derived estimates for Mecklenburg Bay. This suggests that about half of the settling particulate material is exported from the study area and/or subject to processes such as mechanical breakdown, remineralisation and dissolution. Lateral particulate-matter redistribution and particle breakdown in the water column (as opposed to the sediment) seem to be favoured by (repeated) particle resuspension from and resettling to the seafloor before ultimate sedimentary burial. The importance of net lateral redistribution of particulate material seems to increase towards the seafloor and be particularly high within the bottommost few meters of the water column.     

Changes in the shelf macrobenthic community over large temporal and spatial scales in the Bohai Sea, China

Over the past 20 years, the Bohai Sea has been subjected to a considerable human impact through over-fishing and pollution. Together with the influence of the Yellow River cut-off, the ecosystem experienced a dramatic change. In order to integrate available information to detect any change in macrobenthic community structure and diversity over space and time, data collected during the 1980s and the 1990s from 3 regions of the Bohai Sea (Laizhou Bay, 16 stations, 37–38°N, 119–120.5°E; central Bohai Sea, 25 stations, 38–39°N, 119–121°E; eastern Bohai Bay, 12 stations, 38–39°N, 118.5–119°E) were reanalyzed in a comparative way by means of a variety of statistical techniques. A considerable change in community structure between the 1980s and the 1990s and over the geographical regions at both the species and family level were revealed. After 10 years, there was a considerable increase in abundance of small polychaetes, bivalves and crustaceans but decreased number of echinoderms. Once abundant in Laizhou Bay in the 1980s, a large echinoderm Echinocardium cordatum and a small mussel Musculista senhousia almost disappeared from the surveying area in the 1990s. Coupled with the increased abundance was the increased species richness in general whereas evenness was getting lower in central Bohai Sea and Bohai Bay but increased in Laizhou Bay. K-dominance plot showed the same trend as evenness J′. After 10 years, the macrobenthic diversity in the Bohai Sea as a whole was slightly reduced and a diversity ranking of central Bohai Sea > Laizhou Bay > eastern Bohai Bay over space was also suggested. Sediment granulometry and organic content were the two major agents behind the observed changes.     

Community structure of mesopelagic copepods (>500 μm) in the Ligurian Sea (Western Mediterranean)

The distribution of mesopelagic copepods between 250 and 1000 m depth was studied along a transect off the coast of Villefranche-sur-Mer in June 1991 by means of a BIONESS multiple-net sampler. Among the most abundant species at all stations were Pleuromamma gracilis, Paraeuchaeta acuta and overwintering Calanus helgolandicus CV copepodites, each species inhabiting different depth layers with maximum abundances in the 250–350 m, 450–550 m and the 700–850 m layer, respectively. Cluster analysis of the vertical distribution of all species caught revealed three distinct strata (‘TOP', ‘INTERMEDIATE', ‘BOTTOM') consisting of characteristic species that occurred in the same stratum at all stations. Differences in total abundances of the species assemblage occupying each layer are discussed with regard to predation pressure by mesopelagic macroplanktonic crustaceans and fish. We show that each stratum is characterized by distinct trophic interactions and life strategies, which play an important role in the structuring of mesopelagic zooplankton communities.     

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.     

Influence of the heavy fuel spill from the Prestige tanker wreckage in the overlying seawater column levels of copper, nickel and vanadium (NE Atlantic ocean)

The water column above the Prestige wreckage was sampled during two consecutive campaigns: Prestinaut (December 2002) two weeks after the tanker sunk and HidroPrestige0303 (March 2003) one month after the sealing of the main fuel leaks. Samples of the original cargo fuel and the emulsified fuel in the surface of the ocean were also collected. Analysis of the fuel indicated the release of 135 kg of Cu, 1700 kg of Ni and 5300 kg of V from the original fuel to the water column, remaining 35 kg of Cu, 3100 kg of Ni and 13,800 kg of V in the emulsified fuel. The metal partitioning between the water column and the emulsioned floating fuel, Cu > Ni ~ V, are in accordance with the stability index for the metal–nitrogen bond in metalloporphyrins. This release had an impact on dissolved trace metal concentrations in the water column. An increase on dissolved copper (2.8–4.7 nM) and nickel (2.2–8.0 nM) with respect to natural values (1–3 nM for Cu and 1.6–5 nM for Ni) was observed. Values for vanadium (28–35 nM) were in the range of pristine North Atlantic waters (30–36 nM). This contamination was especially observed in the upper water column (0–50 m), associated with the mixing of seawater with the fuel moving upwards, and in deep waters, where the residence time of fuel is higher. Future research in this field should focus on the environmental variables and the processes that control the release of contaminants from fuels for a better assessment of the contamination in oil-spill events.     

Linking mercury exposure to habitat and feeding behaviour in Beaufort Sea beluga whales

Mercury (Hg) levels in the Beaufort Sea beluga population have been increasing since the 1990's. Ultimately, it is the Hg content of prey that determines beluga Hg levels. However, the Beaufort Sea beluga diet is not understood, and little is known about the diet Hg sources in their summer habitat. During the summer, they segregate into social groups based on habitat use leading to the hypothesis that they may feed in different food webs explaining Hg dietary sources. Methyl mercury (MeHg) and total mercury (THg) levels were measured in the estuarine-shelf, Amundsen Gulf and epibenthic food webs in the western Canadian Arctic collected during the Canadian Arctic Shelf Exchange Study (CASES) to assess their dietary Hg contribution. To our knowledge, this is the first study to report MeHg levels in estuarine fish and epibenthic invertebrates from the Arctic Ocean. Although the Mackenzie River is a large source of Hg, the estuarine-shelf prey items had the lowest MeHg levels, ranging from 0.1 to 0.27 μg/g dry weight (dw) in arctic cisco (Coregonus autumnalis) and saffron cod (Eleginus gracilis) respectively. Highest MeHg levels occurred in fourhorn sculpin (Myoxocephalus quadricornis) (0.5 μg/g dw) from the epibenthic food web. Beluga hypothesized to feed in the epibenthic and Amundsen Gulf food webs had the highest Hg levels matching with high Hg levels in associated food webs, and estuarine-shelf belugas had the lowest Hg levels (2.6 μg/g dw), corresponding with the low food web Hg levels, supporting the variation in dietary Hg uptake. The trophic level transfer of Hg was similar among the food webs, highlighting the importance of Hg sources at the bottom of the food web as well as food web length. We propose that future biomagnification studies incorporate predator behaviour with food web structure to assist in the evaluation of dietary Hg sources.     

Hydrographic structure and zooplankton abundance and diversity off Paita, northern Peru (1994 to 2004) — ENSO effects, trends and changes

The objective of the present study was to verify possible spatial, seasonal, and inter-annual changes in the zooplankton off Paita (northern Peru), an upwelling area located closely to the limits of cold Humboldt Current and warm Equatorial Surface Waters. Zooplankton was sampled at subsurface on 53 occasions from August 1994 to December 2004 at four stations located 2 to 30 km offshore with a WP-2 net (300 µm). Extremely high surface water temperatures combined with low salinities were observed during the 1997/98 El Niño up to 29.0 °C) and in April 2002 (up to 25.0 °C). Temperatures more than 2 °C above monthly average were also observed in October 1994, in April 2000, and in November 2004. Significant trends were observed for oxygen concentration (increase) and several horizontal and vertical gradients. Among the copepods (72% of all individuals), the most abundant species were Paracalanus parvus (28%), Acartia tonsa (26%), and Calanus sp. (10%). The strong 1997–98 El Niño (EN) event led to drastic changes in species composition that were reversed during the 1998–99 La Niña (LN) event. Community parameters such as total abundance, diversity, equitability and species richness displayed marked variations associated with the 1997–98 EN and long-term trends. Long-term trends were significant for several vertical and horizontal temperature and oxygen gradients, indicating an increase in upwelling intensity at the shelf during the study period. 10-year-trends were also significant for total zooplankton abundance (increase) and community evenness (J, decline). Our data confirmed the importance of the weak EN in 2002/03 for the study region. Within the trend of increasing zooplankton abundance, a sharp step or shift was observed from 1999 to 2000. When using sequential t-tests to detect shifts in (x + 1) transformed abundance data, a significant rupture was found between the last sampling in 1999 and the first sampling in 2000. Also, a substantial decrease in diel variability occurred after 1999, probably due to changes in vertical migration patterns. The considerable increase in zooplankton abundance over the study period, the ENSO effect, and the 1999–2000 transition are discussed with regard to synchronicity with other zooplankton time series. The present study contributes with the first evidence from an important area located in the Humboldt Current for synchronous trends and changes that were previously observed elsewhere in the Pacific. Our results demonstrated the importance of long-term zooplankton monitoring studies in upwelling areas, and confirms the idea of dramatic changes in pelagic ecosystem structure occurring in the East Pacific.     

Experiments on the survival of six brackish macro-invertebrates from the Baltic Sea after dredged spoil coverage and its implications for the field

Physical disturbance by disposal of dredged materials in estuarine and coastal waters may result in burial of benthic fauna. Survival rates depend on a variety of factors including the type and amount of disposed materials and the lifestyle of the organisms. Laboratory burial experiments using six common macrobenthic invertebrates from a brackish habitat of the western Baltic Sea were performed to test the organisms' escape reaction to dredged material disposal. Experimental lab-results were then extrapolated to a field situation with corresponding bottom topography and covering layer thicknesses at experimental field disposal study sites. Resulted survival rates were then verified by comparison with results of an earlier field study at the same disposal sites.Our experimental design in the lab included the disposal of two types of dredged material (i.e. ‘till’ and ‘sand/till mixture’) and two covering layer depths (i.e. 10–20 cm and 14–40 cm). All three bivalves Arctica islandica (Linnaeus), Macoma balthica (Linnaeus), Mya arenaria (Linnaeus) and the polychaete Nephtys hombergii (Savigny) successfully burrowed to the surface of a 32–41 cm deposited sediment layer of till or sand/till mixture and restored contact with the overlying water. These high escape potentials could partly be explained by the heterogeneous texture of the till and sand/till mixture with ‘voids’. The polychaete Bylgides (Harmothoe) sarsi (Malmgren) successfully burrowed through a 16 cm covering layer whereas the polychaete Lagis koreni (Malmgren) showed almost no escaping reaction. No general differences in escape behaviour after burial were detected between our test species from the brackish habitat and those reported in the literature for the same species in marine environments. However, a size-dependence in mobility of motile polychaetes and M. arenaria was apparent within our study. In comparison to a thick coverage, thin covering layers (i.e. 15–16 cm and 20 cm) increased the chance of the organisms (N. hombergii and M. arenaria) to reach the sediment surface after burial. This was not observed for the other test species. While crawling upward to the new sediment surfaces burrowing velocities of up to 8 cm d^− 1 were observed for the bivalves and up to 20 cm d^− 1 for N. hombergii. Between 17 and 79% of the test organisms showed burrowing activity after experimental burial. The survival rate (defined as the ability to regained contact with the sediment surface) ranged from 0 to 33%, depending on species and on burial depth. The organisms reached the sediment surface by burrowing (polychaetes and bivalves) and/or by extending their siphons to the new sediment surface (bivalves). The extrapolation of laboratory survival rates to the two disposal sites was obtained based on the in situ thicknesses of the dredged spoil layers measured by multi-beam echo sounder. This resulted in total average survival rate estimates for the test species of 45 and 43% for the two disposal sites. The results obtained during the laboratory tests and the following extrapolation to the field were verified by the range of results from a previous field study, using grab sampling shortly before and after a disposal event in June 2001. The effect of dredged material disposal on the tested Baltic Sea benthic macrofauna was assessed by extrapolating the verified laboratory results to the field.     

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.     

The influence of oceanographic scenarios on the population dynamics of demersal resources in the western Mediterranean: Hypothesis for hake and red shrimp off Balearic Islands

The aim of the present paper is to study the relationships between some climatic indices and parental stock, recruitment and accessibility to trawl fishery of hake (Merluccius merluccius) and red shrimp (Aristeus antennatus) off Balearic Islands (western Mediterranean). Available annual catch per unit effort, recruitment and spawning stock biomass have been used as biological data. As environmental data, the meso-scale IDEA index and the large-scale North Atlantic Oscillation (NAO) and Mediterranean Oscillation (MO) indices have been used. To analyze possible links between these indices with the population dynamics of demersal resources, two non-linear approaches have been applied: (i) stock–recruitment relationships from Ricker and Beverton–Holt models, by sequentially incorporating environment factors; (ii) generalized additive modelling, both classical general and threshold non-additive models were considered. The latter simulate an abrupt change in explicative variables across different phases (time periods or climatic index values). The results have shown that two oceanographic scenarios around the Balearic Islands, associated with macro and meso-scale climate regimes, can influence the population dynamics of hake and red shrimp. This is especially true for recruitment, which seems to be enhanced during low NAO and IDEA indices periods. During these periods, colder-than-normal winters generate high amounts of cold Western Mediterranean Intermediate Waters (WIW) in the Gulf of Lions, which flow southwards and reach the Balearic Islands channels in spring, increasing the productivity in the area. This oceanographic scenario could also be favourable to the distribution of hake on the fishing grounds where the trawl fleet targets this species, increasing its accessibility to the fishery. Both spawning stock and abundance of red shrimp seems to be also enhanced by high MO index periods, which could reflect the increased presence of the saline and warm Levantine Intermediate Waters (LIW) in the study area, extending over the fishing grounds of this species. The proposed interactions can be useful to assess and manage these important demersal resources.     

Picophytoplankton and nanophytoplankton abundance and distribution in the southeastern Beaufort Sea (Mackenzie Shelf and Amundsen Gulf) during Fall 2002

The distribution of picophytoplankton (0.2–2 µm) and nanophytoplankton (2–20 µm) in the Beaufort Sea–Mackenzie Shelf and Amundsen Gulf regions during autumn, 2002 is examined relative to their ambient water mass properties (salinity, temperature and nutrients: nitrate + nitrite, phosphate, and silicate) and to the ratio of variable to maximum fluorescence, Fv/Fm. Total phytoplankton and cell abundances (< 20 µm) were mainly correlated with salinity. Significant differences in picophytoplankton cell numbers were found among waters near the mouth of the Mackenzie River, ice melt waters and the underlying halocline water masses of Pacific origin. Picophytoplankton was the most abundant phytoplankton fraction during the autumnal season, probably reflecting low nitrate concentrations (surface waters average ~ 0.65 µM). The ratio Fv/Fm averaged 0.44, indicating that cells were still physiologically active, even though their concentrations were low (max Chl a = 0.9 mg m^− 3). No significant differences in Fv/Fm were evident in the different water masses, indicating that rate limiting conditions for photosynthesis and growth were uniform across the whole system, which was in a pre-winter stage, and was probably already experiencing light limitation as a result of shortening day lengths.     

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.     

Bacterial production and microbial food web structure in a large arctic river and the coastal Arctic Ocean

Globally significant quantities of organic carbon are stored in northern permafrost soils, but little is known about how this carbon is processed by microbial communities once it enters rivers and is transported to the coastal Arctic Ocean. As part of the Arctic River-Delta Experiment (ARDEX), we measured environmental and microbiological variables along a 300 km transect in the Mackenzie River and coastal Beaufort Sea, in July–August 2004. Surface bacterial concentrations averaged 6.7 × 10⁵ cells mL^− 1 with no significant differences between sampling zones. Picocyanobacteria were abundant in the river, and mostly observed as cell colonies. Their concentrations in the surface waters decreased across the salinity gradient, dropping from 51,000 (river) to 30 (sea) cells mL^− 1. There were accompanying shifts in protist community structure, from diatoms, cryptophytes, heterotrophic protists and chrysophytes in the river, to dinoflagellates, prymnesiophytes, chrysophytes, prasinophytes, diatoms and heterotrophic protists in the Beaufort Sea.Size-fractionated bacterial production, as measured by ³H–leucine uptake, varied from 76 to 416 ng C L^− 1 h^− 1. The contribution of particle-attached bacteria (> 3 µm fraction) to total bacterial production decreased from > 90% at the Mackenzie River stations to < 20% at an offshore marine site, and the relative importance of this particle-based fraction was inversely correlated with salinity and positively correlated with particulate organic carbon concentrations. Glucose enrichment experiments indicated that bacterial metabolism was carbon limited in the Mackenzie River but not in the coastal ocean. Prior exposure of water samples to full sunlight increased the biolability of dissolved organic carbon (DOC) in the Mackenzie River but decreased it in the Beaufort Sea.Estimated depth-integrated bacterial respiration rates in the Mackenzie River were higher than depth-integrated primary production rates, while at the marine stations bacterial respiration rates were near or below the integrated primary production rates. Consistent with these results, P_CO2 measurements showed surface water supersaturation in the river (mean of 146% of air equilibrium values) and subsaturation or near-saturation in the coastal sea. These results show a well-developed microbial food web in the Mackenzie River system that will likely convert tundra carbon to atmospheric CO₂ at increasing rates as the arctic climate continues to warm.     

The annual cycle of nanoflagellates in the Central Cantabrian Sea (Bay of Biscay)

The annual cycle of nanoflagellates (NF) including autotrophic (ANF), heterotrophic (HNF) and mixotropic (MNF) flagellates carried out in a temperate sea (Central Cantabrian Sea, southern Bay of Biscay) is presented. Three stations with characteristics ranging from coastal to oceanic conditions were analysed in order to compare NF response to this gradient. Samples were monthly collected at each station at three different depths between February 2002 and December 2002. CTD profiles were also taken at each station. NF were grouped according to their trophic status into ANF, HNF and MNF. Abundance and biomass were determined for each group. The annual cycle showed a general pattern consisting in a maximum in July with secondary maxima in March and October and minimum values in May. ANF were the most important fraction, making a major contribution (nearly 75%) to total NF biomass in all stations. HNF represented over 20% along the cycle, except for a peak in spring found in every station. MNF reached less than 5%, showing low seasonability. Small flagellates (2–5 µm) dominated throughout the cycle. Microplankton community was also analysed in terms of abundance and biomass. A significant positive correlation (r² = 0.49) was obtained between 2–5 µm NF and 10–20 µm HNF–MNF biomasses, suggesting a possible trophic relationship between these groups which should be cautiously taken. No significant relationships were found between microplankton and NF or between nutrients and ANF, indicating that the regulation of NF numbers is complex and probably implicates other groups. In addition to this, the unexpected 2002 Chl a concentration pattern and the misplacing of upwelling events render necessary to perform additional studies to fully understand the precise behaviour of NF in the Cantabrian Sea. To the best of our knowledge, this is the first study of a NF cycle in a temperate sea that considers all functional groups.     

The biogeochemical cycling of methane in Ria de Vigo, NW Spain: Sediment processing and sea–air exchange

Methane (CH₄) concentrations were measured in the water column, in sediment porewaters, and in atmospheric air, in the Ría de Vigo, NW Spain, during both the onset (April 2003) and at the end of (September 2004) seasonal upwelling. In addition, CH₄ concentration and stable isotopic signatures (δ¹³CH₄) were measured in porewaters, and sediment methanogenesis and aerobic oxidation of CH₄ were determined in sediment incubations. Surface water column CH₄ (2 m depth) was in the range 3–180 nmol l^− 1 (110–8500% saturation) and followed a generally landward increase but with localised maxima in both the inner and middle Ría. These maxima were consistent with CH₄ inputs from underlying porewaters in which CH₄ concentrations were up to 3 orders of magnitude higher (maximum 350 μmol l^− 1). Surface water CH₄ concentrations were approximately three times higher in September than in April, consistent with a significant benthic CH₄ flux driven by enhanced sediment methanogenesis following the summer productivity maximum. CH₄ and δ¹³CH₄ in sediment porewaters and in incubated sediment slurries (20 °C) revealed significant sediment CH₄ oxidation, with an apparent isotopic fractionation factor (r_c) of  1.004. Using turbulent diffusion models of air–sea exchange we estimate an annual emission of atmospheric CH₄ from the Ría de Vigo of 18–44 × 10⁶ g (1.1–2.7 × 10⁶ mol). This estimate is approximately 1–2 orders of magnitude lower than a previous estimate based on a bubble transport model.