High concentrations of exopolymeric substances in Arctic winter sea ice: implications for the polar ocean carbon cycle and cryoprotection of diatoms

Exopolymeric substances (EPS) produced by microorganisms play important roles in various aquatic, porous, and extreme environments. Only recently has their occurrence in sea ice been considered. We used macroscopic and microscopic approaches to study the content and possible ecological role of EPS in wintertime fast ice near Barrow, Alaska (71°20′ N, 156°40′ W). Using Alcian blue staining of melted ice samples, we observed high concentrations of EPS in all samples examined, ranging from 0.79 to 7.71 mg xanthan gum equivalents (XGEQV) l⁻¹. Areal conversions to carbon equivalents yielded 1.5−1.9 g C m⁻² ice in March and 3.3−4.0 g C m⁻² in May (when the ice was thicker). Although EPS did not correlate with macronutrient or pigment data, the latter analyses indicated ongoing or recent biological activity in the ice within temperature horizons of −11°C to −9°C and warmer. EPS correlated positively with bacterial abundance (although no functional relationship could be deduced) and with dissolved organic carbon (DOC) concentrations. Ratios of EPS/DOC decreased at colder temperatures within the core, arguing against physical conversion of DOC to EPS during freezing. When sea-ice segments were maintained at representative winter temperatures (−5°C,−15°C and −25°C) for 3−14 months, the total EPS content increased significantly at rates of 5−47 μg XGEQV l⁻¹ d⁻¹, similar to published rates of EPS production by diatoms. Microscopic images of ice-core sections at these very cold temperatures, using a recently developed non-invasive method, revealed diatoms sequestered in spacious brine pockets, intact autofluorescent chloroplasts in 47% of the (pennate) diatoms observed, and indications of mucus in diatom-containing pores. The high concentrations of EPS detected in these winter ice cores represent a previously unrecognized form of organic matter that may contribute significantly to polar ocean carbon cycles, not only within the ice but after springtime release into the water column. The EPS present in very high concentrations in the brine of these microhabitats appear to play important buffering and cryoprotectant roles for microorganisms, especially diatoms, against harsh winter conditions of high salinity and potential ice-crystal damage.     

Total oxyradical scavenging capacity and cell membrane stability of haemocytes of the Arctic scallop, Chlamys islandicus, following benzo(a)pyrene exposure

Industrial activities, notably oil and gas industries, are expanding in the Arctic. Most of the biomarkers were developed using temperate organisms living at temperatures above 10 °C. Little is known about the biomarker responses of organisms living between −1.88 and 5 °C. Therefore, assessment of the toxicity of chemicals to cold-water adapted species is required. In this study, the Arctic scallop, Chlamys islandicus, was selected as a key species for bio-monitoring because of wide distribution in Arctic waters and its commercial value. Test animals, stored in seawater at 2 °C, were injected with benzo(a)pyrene (diluted in cod liver oil 5 mg ml⁻¹) in the adductor muscle every 24 h for four days giving a final dose of 0, 74 and 90.6 mg kg⁻¹ wet weight for control, low and high dose, respectively. The biomarkers used were total oxyradical scavenging capacity (TOSC) in the digestive gland and cell membrane stability of haemocytes. TOSC values were significantly reduced (ca. 30%) in exposed groups (P<0.05), indicating a depletion in oxyradical molecular scavengers. The antioxidant defences appeared to be overwhelmed by the reactive oxygen species as the plasma membranes of haemocytes were destabilised (P<0.05) probably due to lipid peroxidation. These data indicate that reactive oxygen species (ROS) were produced by Arctic scallops via the metabolisation of benzo(a)pyrene at 2 °C.     

Grazing experiments and model simulations of the role of zooplankton in Phaeocystis food webs

A combined empirical and modelling study was conducted to further examine the potential importance of grazing by zooplankton in pelagic food webs in which Phaeocystis is a significant or dominant component. Laboratory experiments were designed to measure ingestion of Phaeocystis and other potential prey items which co-occur with Phaeocystis. Grazers included copepods and ciliates, and prey included Phaeocystis colonies and solitary cells, diatoms, ciliates, bacteria, and detritus. These data were expressed in the model currency of nitrogen units, and fit to hyperbolic tangent equations which included minimum prey thresholds. These equations and literature data were used to constrain a food web model whose purpose was to investigate trophic interactions rather than to mimic actual events. Nevertheless, the model output was similar to the general pattern and magnitude of development of Phaeocystis–diatom communities in some environments where they occur, e.g. north Norwegian waters. The model included three forms of nitrogen, three phytoplankton groups, bacteria, two zooplankton groups, and detritus, with detailed flows between compartments. An important component of the model was inclusion of variable prey preferences for zooplankton. The experiments and model simulations suggest several salient conclusions. Phaeocystis globosa colonies were eaten by a medium-sized copepod species, but ingestion appeared to be strongly dependent upon a proper size match between grazer and prey. If not, colonies were eaten little if at all. Phaeocystis solitary cells were ingested rapidly by ciliate microzooplankton, in agreement with prior literature observations. In contrast, detritus was eaten comparatively slowly by both ciliates and copepods. Both types of zooplankton exhibited apparent minimum prey thresholds below which grazing did not occur or was inconsequential. Model simulations implied that transitions between life cycle stages of Phaeocystis may potentially be important to phytoplankton–zooplankton interactions, and that relative rates of ingestion of Phaeocystis by various zooplankton may have significant impacts upon material fluxes through and out of Phaeocystis–diatom ecosystems. Indirect effects of trophic interactions appear to be equally significant as direct effects.     

Uncoupled distributions of transparent exopolymer particles (TEP) and dissolved carbohydrates in the Southern Ocean

Transparent exopolymer particles (TEP) are formed by the assembly of dissolved precursors, mainly mono and polysaccharides (DMCHO and DPCHO) that are released by microorganisms. Although TEP formation plays a significant role in carbon export to deep waters and can affect gas exchange at the sea surface, simultaneous measurements of TEP and their precursors in natural waters have been scantly reported. In this study, we described the spatial (vertical and regional) distribution of TEP, DMCHO and DPCHO in a region located around the Antarctic Peninsula, assessed their contribution to the total organic carbon pool, and explored their relationships with phytoplankton (with chlorophyll a (chl a) as a proxy) and bacteria. TEP concentration ranged from undetectable values to 48.9 µg XG eq L^− 1 with a mean value of 15.4 µg XG eq L^− 1 (11.6 µg TEP-C L^− 1). DMCHO and DPCHO showed average values of 4.3 µmol C L^− 1 and 8.6 µmol C L^− 1, respectively. We did not find simple relationships between the concentrations of TEP and dissolved carbohydrates, but a negative correlation between DMCHO and DPCHO was observed. Chl a was the best regressor of TEP concentration in waters within the upper mixed layer, while bacterial production was the best regressor of TEP concentration below the mixed layer, underlining the direct link between these particles and bacterial activity in deep waters.     

Biological components of Ross Sea short-term particle fluxes in the austral summer of 1995–1996

The Ross Sea, a region of high seasonal production in the Southern Ocean, is characterized by blooms of the haptophyte Phaeocystis antarctica and of diatoms. The different morphology, structural composition and consumption of these two phytoplankton by grazing zooplankton may result in different carbon cycling dynamics and carbon flux from the euphotic zone. We sampled short-term (2 days) particle flux at 5 sites from 177.6°W to 165°E along a transect at 76.5°S with traps placed below the euphotic zone at 200 m during December 1995–January 1996. We estimated carbon flux of as many eucaryotic organisms and fecal pellets as possible using microscopy for counts and measurements and applying volume:carbon conversions from the literature. Eucaryotic organisms contributed about 20–40% of the total organic carbon flux in both the central Ross Sea polynya and in the western polynya, and groups of organisms differed in contribution to the carbon flux at the different sites. Algal carbon flux ranged from 4.5 to 21.1 mg C m⁻² day⁻¹ and consisted primarily of P. antarctica (cell plus mucus) and diatom carbon at all sites. Different diatom species dominated the diatom flux at different sites. Carbon fluxes of small pennate diatoms may have been enhanced by scavenging, by sinking senescent P. antarctica colonies. Heterotrophic carbon flux ranged from 9.2 to 37.6 mg C m⁻² day⁻¹ and was dominated by athecate heterotrophic dinoflagellate carbon in general and by carbon flux of a particular large athecate dinoflagellate at two sites. Fecal pellet carbon flux ranged from 4.6 to 54.5 mg C m⁻² day⁻¹ and was dominated by carbon from ovoid/angular pellets at most sites. Analysis of fecal pellet contents suggested that large protozoans identified by light microscopy contributed to ovoid/angular fecal pellet fluxes. Carbon flux as a percentage of daily primary production was lowest at sites where P. antarctica predominated in the water column and was highest at sites where fecal pellet flux was highest. This indicates the importance of grazers in carbon export.     

Depth profiles of volatile iodine and bromine-containing halocarbons in coastal Antarctic waters

Measurements of bromoform (CHBr₃), diiodomethane (CH₂I₂), chloroiodomethane (CH₂ICl) and bromoiodomethane (CH₂IBr) were made in the water column (5–100 m depth) of the Southern Ocean within 0–40 km of the Antarctic sea ice during the ANTXX1/2 transect of the German R/V Polarstern, at five locations between 70–72°S and 9–11°W in the Antarctic spring/summer of 2003–2004. Some of the profiles exhibited a very pronounced layer of surface sea-ice meltwater, as evidenced by salinity minima and temperature maxima, along with surface maxima in concentrations of CHBr₃, CH₂I₂, CH₂ICl and CH₂IBr. These results are consistent with in situ surface halocarbon production by ice algae liberated from the sea ice, although production within the sea ice followed by transport cannot be entirely ruled out. Additional sub-surface maxima in halocarbons occurred between 20 and 80 m. At a station further from shore and not affected by surface sea-ice meltwater, surface concentrations of CH₂I₂ were decreased whereas CH₂ICl concentrations were increased compared to the stations influenced by meltwater, consistent with photochemical conversion of CH₂I₂ to CH₂ICl, perhaps during upward mixing from a layer at  70 m enhanced in iodocarbons. Mean surface (5–10 m) water concentrations of halocarbons in these coastal Antarctic waters were 57 pmol l^− 1 CHBr₃ (range 44–78 pmol l^− 1), 4.2 pmol l^− 1 CH₂I₂ (range 1.7–8.2 pmol l^− 1), 0.8 pmol l^− 1 CH₂IBr (range 0.2–1.4 pmol l^− 1), and 0.7 pmol l^− 1 CH₂ICl (range 0.2–2.4 pmol l^− 1). Concurrent measurements in air suggested a sea-air flux of bromoform near the Antarctic coast of between 1 and 100 (mean 32.3, median 10.4) nmol m^− 2 day^− 1 and saturation anomalies of 557–1082% (mean 783%, median 733%), similar in magnitude to global shelf values. In surface samples affected by meltwater, CH₂I₂ fluxes ranged from 0.02 to 6.1 nmol m^− 2 day^− 1, with mean and median values of 1.9 and 1.1 nmol m^− 2 day^− 1, respectively.     

The impact of groundwater discharges on mercury partitioning, speciation and bioavailability to mussels in a coastal zone

The Mussel Watch program conducted along the French coasts for the last 20 years indicates that the highest mercury concentrations in the soft tissue of the blue mussel (Mytilus edulis) occur in animals from the eastern part of Seine Bay on the south coast of the English Channel, the “Pays de Caux”. This region is characterized by the presence of intertidal and submarine groundwater discharges, and no particular mercury effluent has been reported in its vicinity. Two groundwater emergence systems in the karstic coastal zone of the Pays de Caux (Etretat and Yport with slow and fast water percolation pathways respectively) were seasonally sampled to study mercury distribution, partitioning and speciation in water. Samples were also collected in the freshwater–seawater mixing zones in order to compare mercury concentrations and speciation between these “subterranean” or “groundwater” estuaries and the adjacent macrotidal Seine estuary, characterized by a high turbidity zone (HTZ). The mercury concentrations in the soft tissue of mussels from the same areas were monitored at the same time.The means of the “dissolved” (< 0.45 μm) mercury concentrations (HgT_D) in the groundwater springs were 0.99 ± 0.15 ng l^− 1 (n = 18) and 0.44 ± 0.17 ng l^− 1 (n = 17) at Etretat and Yport respectively. High HgT_D concentrations were associated with strong runoff over short water pathways during storm periods, while low concentrations were associated with long groundwater pathways. Mean particulate mercury concentrations were 0.22 ± 0.05 ng mg^− 1 (n = 16) and 0.16 ± 0.10 ng mg^− 1 (n = 17) at Etretat and Yport respectively, and decreased with increasing particle concentration probably as a result of dilution by particles from soil erosion. Groundwater mercury speciation was characterized by high reactive-to-total mercury ratios in the dissolved phase (HgR_D/HgT_D: 44–95%), and very low total monomethylmercury concentrations (MMHg < 8 pg l^− 1). The HgT_D distributions in the Yport and Etretat mixing zones were similar (overall mean concentration of 0.73 ± 0.21 ng l^− 1, n = 43), but higher than those measured in the adjacent industrialized Seine estuary (mean: 0.31 ± 0.11 ng l^− 1, n = 67). In the coastal waters along the Pays de Caux dissolved monomethylmercury (MMHg_D) concentrations varied from 9.5 to 13.5 pg l^− 1 (2 to 8% of the HgT_D). Comparable levels were measured in the Seine estuary (range: 12.2– 21.1 pg l⁻¹; 6–12% of the HgT_D). These groundwater karstic estuaries seem to be mostly characterized by the higher HgT_D and HgR_D concentrations than in the adjacent HTZ Seine estuary. While the HTZ of the Seine estuary acts as a dissolved mercury removal system, the low turbid mixing zone of the Pays de Caux receives the dissolved mercury inputs from the groundwater seepage with an apparent Hg transfer from the particulate phase to the “dissolved” phase (< 0.45 μm). In parallel, the soft tissue of mussels collected near the groundwater discharges, at Etretat and Yport, exhibited significantly higher values than those found in the mussel from the mouth of the Seine estuary. We observe that this difference mimics the differences found in the mercury distribution in the water, and argue that the dissolved phase of the groundwater estuaries and coastal particles are significant sources of bioavailable mercury for mussels.     

Uptake and metabolism of arsenate by anexic cultures of the microalgae Dunaliella tertiolecta and Phaeodactylum tricornutum

Axenic cultures of the microalgae species, Dunaliella tertiolecta and Phaeodactylum tricornutum were grown at arsenic (As) concentrations typically found in uncontaminated marine environments ( 2 µg L^− 1) under different phosphorus concentrations. D. tertiolecta accumulated higher arsenic concentrations (mean: 13.7 ± 0.7 µg g^− 1 dry mass) than P. tricornutum (mean: 1.9 ± 0.2 µg g−1 dry mass). Media phosphorus concentrations (0.6–3 mg/L) had little influence on microalgae growth rates or arsenic accumulation. Arsenic was present as lipid bound (29–38%; 4.2–9.5%), water-soluble (20–29%; 26–34%) and residue bound (41–45%; 57–69%) arsenic species in D. tertiolecta and P. tricornutum respectively. Hydrolysed lipids contained mostly glycerol arsenoribose (OH- ribose), dimethylarsinate (DMA) and inorganic arsenic (As(V)) moieties. Water-soluble species of microalgae were very different. D. tertiolecta contained inorganic arsenic (54–86%) with variable amounts of DMA (7.4–20%), arsenoriboses (5–25%) and traces of methylarsonate (MA) ( 1%). P. tricornutum contained mostly DMA (32–56%) and phosphate arsenoribose (PO₄-ribose, 23–49%) and small amounts of OH-ribose (3.8–6.5%) and As(V) (9–16%). Both microalgae contained an unknown cationic arsenic species. The residue fractions of both microalgae contained predominately inorganic arsenic (99–100%). These results show that at natural seawater arsenic concentrations, both algae take up substantial amounts of inorganic arsenic that is complexed with structural elements or sequestered in vacuoles as stable complexes. A significant portion is also incorporated into lipids. Arsenic is metabolised to simple methylated species and arsenoriboses.     

Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific

Grazing experiments and production estimation based on life-history analysis of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were carried out in the Oyashio region to understand the carbon flows associated with the interzonal migrating copepods. These copepods, and also Eucalanus bungii, fed on nano- and micro-sized organisms non-selectively throughout the season. However, diatoms were the dominant food resource until May and organisms, such as ciliates were the major resource after May. Daily growth rate was estimated from the Ikeda–Motoda, Huntley–Lopez and Hirst–Sheader models. Since the growth rates were considered to be overestimates for the Huntley–Lopez model and underestimates for the other two models, we applied the weight-specific growth rates previously reported for these species in the Bering Shelf. Surface biomass of Neocalanus increased rapidly in June during the appearance of C5, and a successive increase of overwintering stock was evident in the deeper layer. The deep biomass decreased gradually from September to May during the dormant and reproduction period. N. cristatus has the largest annual mean biomass (2.3 gC m⁻²), followed by N. plumchrus (1.1) and N. flemingeri (0.4). Daily production rate of Neocalanus varied from 0.4 to 363.4 mgC m⁻² day⁻¹, to which N. cristatus was the largest contributor. Annual production was estimated as 11.5 gC m⁻² year⁻¹ for N. cristatus, 5.7 for N. plumchrus and 2.1 for N. flemingeri, yielding annual P/B ratio of 5 for each species. The annual production of Neocalanus accounted for 13.2% of the primary production in the Oyashio region. Their fecal pellets were estimated to account for 14.9% (0.7 gC m⁻² year⁻¹) of sinking flux of organic carbon at 1000-m depth. Moreover, their export flux by ontogenetic vertical migration, which is not measured by sediment trap observations, is estimated to be 91.5% (4.3 gC m⁻² year⁻¹) of carbon flux of sinking particles at 1000-m depth. These results suggest the important role of interzonal migrating copepods in the export flux of carbon.     

Microbial community structure and biomass in surface waters during a Polar Front summer bloom along 170°W

As part of the US Joint Global Ocean Flux Study (US JGOFS) Southern Ocean Program, flow cytometry and epifluorescent microscopy were utilized to determine abundance, distribution and size structure of the microbial community in the Polar Front region during the summer biomass maximum. Surface samples were collected approximately every 10 km along 170°W during two N–S transects, separated in time by two weeks. Phytoplankton abundance and size structure varied with distinct latitudinal trends. Autotrophic biomass was lowest north of the Polar Front reflecting the dominance of small cells. The highest biomass (170 μg C l⁻¹) occurred at 65°S where the composition was strongly influenced by large centric diatoms. Farther south, the diatom community shifted to the dominance of smaller pennate diatoms. Total grazer biomass and size distributions followed similar patterns, ranging from 4 μg C l⁻¹ in the north to 52 μg C l⁻¹ in the south where larger (>20 μm) grazers were more abundant. Heterotrophic bacteria varied over an order of magnitude in abundance across the study site, with size generally increasing from north to south. In the second transect, phytoplankton biomass at 65°S was 50% lower, and grazer biomass and bacterial populations were slightly greater, indicating the decline of the bloom. The changes in biomass and community structure along 170°W and the reduction of phytoplankton standing stock at 65°S over time suggests adjacent, yet different, microbial systems in terms of carbon flux, spanning from primarily recycling to export-dominated.     

Mesozooplankton grazing during the Phaeocystis globosa bloom in the southern bight of the North Sea

During spring blooms 1998 and 1999, three complementary methods were used to evaluate the in situ feeding activities of the dominant copepod species of the Belgian coastal zone: gut pigment content analysis using HPLC, the ¹⁴C tracer method, and cell count experiments. The results obtained by all three methods consistently showed that Phaeocystis globosa is not an adequate food source for the spring copepods in the Belgian coastal zone. Our results demonstrated that, among the potential prey, copepods strongly selected diatoms and microzooplankton, and that these types of prey accounted for the major part of the ingested carbon. However, diatoms and microzooplankton ingestion did not always seem sufficient in terms of carbon to avoid food limitation. Comparison of clearance rates exerted on different potential prey types during the P. globosa peak with those before and after the P. globosa peak showed that the copepods' feeding pressure on diatoms was reduced during the P. globosa peak while that on microzooplankton was not. The low grazing pressure on P. globosa, together with the preferential grazing on diatoms, which reduces the competition for nutrients, and the predation on microzooplankton organisms, which reduces the microzooplankton grazing pressure on P. globosa cells, are likely to favour the P. globosa bloom in the Southern Bight of the North Sea.     

Plankton dynamics and carbon flux in an area of upwelling off the coast of Morocco

A carbon flux study was carried out off the coast of Morocco, at 31°N, in a region characterized by the presence of a persistent cyclonic eddy. Two short-term (4 and 3 day) deployments of free-floating sediment traps were combined with water column sampling and rate process measurements as the ship followed the traps. For a period of 36 h between trap deployments, a hydrographic section was run along 31°30'N as part of a larger scale survey being carried out simultaneously on the R.V. A. von Humboldt. The first trap deployment was near the eastern margin of the eddy and the traps moved to the north and west in a frontal jet associated with its northern boundary. After the second deployment, which was at the recovery point of the first, the traps moved to the west and then to the southwest. Throughout the study, chlorophyll concentrations varied between 27 and 125 mg m⁻² (0–100 m), with highest concentrations in the upwelled water nearest the coast and in upwelled water generated within the cyclonic eddy. Particulate organic carbon (POC) and particulate organic nitrogen (PON) concentrations were relatively uniform (13.6±1.8 and 1.63±28 g m⁻² with phytoplankton carbon accounting for 16–85% of total POC. Bacterial carbon was 5% of total POC and mesozooplankton carbon concentrations were equivalent to 9% of total POC. Microzooplankton biomass was not assessed but POC:PON ratios in the water column were often high, suggesting there was sometimes a large detrital component in the POC. Primary production rates varied between 1.0 and 2.5 g C m⁻² day⁻¹. Bacterial consumption accounted for 50% of primary production. Metabolic rates suggested that copepods were ingesting more than 0.4 g C m⁻² day⁻¹. while filtration rates suggested that ingestion of phytoplankton carbon was only 0.2 g C m⁻²day⁻¹, even when phytoplankton constituted 85% of the POC. f-ratios (based on uptake rates for ¹⁵N-nitrate and ammonia) were between 0.1 and 0.4, and excretion by mesozooplankton could account for 40% of the daily ammonium uptake by phytoplankton. HPLC pigment analysis showed that when chlorophyll biomass was high, diatoms were dominant, whereas when it was low, small prymnesiophytes, chlorophytes and diatoms were all important. The composition of the fluoresecent pigments in material in the sediment traps indicated that intact phytoplankton and copepod faecal pellets were the main sources but the relative rates of sedimentation of pigment, POC and PON for the two trapping periods did not reflect differences that were observed in the overlying water column. This was likely to be the result of spatial heterogeneity and strong horizontal currents heterogeneity and strong horizontal currents within the euphotic zone. Thus, material collected at 100 m probably did not originate in the water column immediately overlying the traps and trapping efficiencies might also have been variable.     

Reconstruction of sea surface dimethylsulfide in the North Pacific during 1970s to 2000s

We proposed an empirical equation of sea surface dimethylsulfide (DMS, nM) using sea surface temperature (SST, K), sea surface nitrate (SSN, μM) and latitude (L, °N) to reconstruct the sea surface flux of DMS over the North Pacific between 25°N and 55°N: ln DMS = 0.06346 · SST − 0.1210 · SSN − 14.11 · cos(L) − 6.278 (R² = 0.63, p < 0.0001). Applying our algorithm to climatological hydrographic data in the North Pacific, we reconstructed the climatological distributions of DMS and its flux between 25 °N and 55 °N. DMS generally increased eastward and northward, and DMS in the northeastern region became to 2–5 times as large as that in the southwestern region. DMS in the later half of the year was 2–4 times as large as that in the first half of the year. Moreover, applying our algorithm to hydrographic time series datasets in the western North Pacific from 1971 to 2000, we found that DMS in the last three decades has shown linear increasing trends of 0.03 ± 0.01 nM year^− 1 in the subpolar region, and 0.01 ± 0.001 nM year^− 1 in the subtropical region, indicating that the annual flux of DMS from sea to air has increased by 1.9–4.8 μmol m^− 2 year^− 1. The linear increase was consistent with the annual rate of increase of 1% of the climatological averaged flux in the western North Pacific in the last three decades.     

Feeding selectivity of Calanus finmarchicus in the Trondheimsfjord

The feeding selectivity of Calanus finmarchicus was studied by carrying out three incubation experiments; two experiments with natural seawater sampled during spring bloom (Exp. 1) and post-bloom conditions (Exp. 2) and a third experiment with cultured dinoflagellates and ciliates (Exp. 3). In the first two experiments a gradient in ciliate concentration was created to investigate the potential for prey density dependent selective feeding of C. finmarchicus. Results of microplankton counts indicated C. finmarchicus to be omnivorous. Diatoms contributed chiefly to the diet during spring bloom conditions. Despite the high microphytoplankton biomass during the spring bloom (Exp. 1), ciliates were selected positively by C. finmarchicus when the ciliate biomass exceeded 6.5 μg C L^− 1. A selection in favor of large conic ciliates such as Laboea sp. and Strombidium conicum was indicated by positive selectivity indices. Ciliates were throughout positively selected by C. finmarchicus during Exp. 2, and selectivity indices indicated a negative selection of diatoms. The results from Exp. 3 showed that C. finmarchicus has the ability to switch from dinoflagellates to ciliates as sole food source, even if the dinoflagellate was offered in surplus. This suggests that other factors, such as nutrition may be of significance for the feeding selectivity of C. finmarchicus.     

Molluscan assemblages of seagrass-covered and bare intertidal flats on the Banc d'Arguin, Mauritania, in relation to characteristics of sediment and organic matter

The Banc d'Arguin, a non-estuarine area of shallows and intertidal flats off the tropical Saharan coast of Mauritania, is characterised by extensive intertidal and subtidal seagrass beds. We examined the characteristics of intertidal seagrass (Zostera noltii) meadows and bare areas in terms of the presence and abundance of molluscs (gastropods and bivalves). To explain observed differences between molluscan assemblages in seagrass and bare patches, some aspects of the feeding habitat (top-5 mm of the sediment) and of food (organic materials) of molluscs were examined. The novelty of this study is that phytopigments were measured and identified to assess source and level of decay (freshness) of organic material in the sediment and to study their importance as an explanatory variable for the distribution of molluscs. Over an area of 36 km² of intertidal flats, at 12 sites, paired comparisons were made between seagrass-covered and nearby bare patches. Within seagrass meadows, dry mass of living seagrass was large and amounted to 180 ±10 g AFDM m^− 2 (range 75–240). Containing twice the amount of silt per unit dry sediment mass, seagrass sediments were muddier than bare areas; the relative amount of organic material was also larger. The total number of species of bivalves and gastropods amounted to 27, 14 of which were found only in seagrass areas, 4 only in bare and 9 in both types of habitat. Among the three numerically most abundant species, the bivalves Anadara senilis, Dosinia hepatica and Loripes lacteus, the first was numerically most abundant in bare and the other two in seagrass-covered areas. Bare intertidal areas had greater mean total biomass of molluscs (80.5 g AFDM m^− 2) than seagrass meadows (30.0 g AFDM m^− 2). In both habitats, the bulk of the biomass was made up by A. senilis. Excluding this species, bare mudflats contained on average only 3.1 g AFDM m^− 2 and seagrass meadows 6.9 g AFDM m^− 2. As compared to previous surveys in 1980–1986, the biomass of A. senilis had increased almost 10-fold and D. hepatica, previously found in very small numbers, had become the most numerous species. However, the total biomass excluding that of A. senilis was similar. Concentrations of phytopigments were similar to those observed at temperate mudflats, indicating that the Banc d'Arguin might not be as oligotrophic as previously thought. Per unit of dry sediment mass, smaller amounts of phytopigments were found in bare than in seagrass areas. Per unit of dry organic material, bare sediments contained most (fresh) phytopigments. This suggests that in seagrass-covered meadows the organic material is more degraded than in bare sediments. Overall, the composition of phytopigments, quite surprisingly, indicated a benthic-diatom-dominated trophic system. Multivariate statistics revealed that patterns of zoobenthic assemblages were correlated with patterns of a combination of four environmental parameters: grain size of the sediment, amount of fresh phytopigments and amounts of leaves and roots of seagrass.     

Planktonic ciliates as food for the scyphozoan <Emphasis Type="Italic">Aurelia coerulea</Emphasis>: feeding and growth responses of ephyra and metephyra stages

Blooms of the moon jellyfish, Aurelia coerulea, often occur in eutrophic embayments where microzooplankton are abundant. Hence, microzooplankton may play a role in A. coerulea population dynamics. In this study, feeding and growth responses of the ephyra and metephyra stages of A. coerulea were examined on a diet of planktonic ciliates, the main components of microzooplankton. Laboratory experiments indicated that ephyrae actively fed on ciliates and that the central disc diameter increased over the 5-day incubation period. Carbon-specific growth rates estimated from daily increases of central disc diameter increased with carbon-specific prey consumption rates until about 4 µg C ind day^?1 and then became saturated. By fitting these results to rectangular hyperbolic equations, the maximum specific growth rates were estimated to be 0.32 day^?1 for ciliate prey. However, the central disc diameter of the metephyrae did not increase on a diet of ciliates during the 5-day incubation, even though consumption of prey ciliates was observed. These results suggest that planktonic ciliates serve as a food source for growth at the ephyra stage of A. coerulea and then the contribution to growth decreases at and after the metephyra stage. This prey switch-dependency on the development stage of A. coerulea may be associated with the frequent occurrence of jellyfish blooms in eutrophic embayments.     

Growth and mortality of larval and juvenile Japanese seaperch Lateolabrax japonicus in relation to seasonal changes in temperature and prey abundance in the Chikugo estuary

Japanese seaperch Lateolabrax japonicus migrate from Ariake Bay to the estuarine turbidity maximum (ETM) zone of the Chikugo River and inhabit there through the post-migration period (15–20 mm in standard length). The feeding, growth and mortality during the post-migration period of Japanese seaperch were analyzed in relation to seasonal changes in temperature and prey concentration. Larvae and juveniles were collected from ten sampling stations at 4–7 day intervals from 24 February to 24 April 2005 in the Chikugo estuary. Based on the otolith microstructure analysis the sampled fish were divided into nine cohorts, each cohort covering a 5 day hatch date period (22 December 2004 to 4 February 2005). The growth coefficient (G, day⁻¹) was higher and the mortality coefficient (M, day⁻¹) was lower in the later cohorts. The ratio of G to M as an index of stage-specific survival during the post-migration period significantly increased as the season progressed and exceeded 1.0 in the last cohort examined. Variability in abundance of the major prey organism, Sinocalanus sinensis, had a significant effect on the Japanese seaperch ingestion rate. Increase in temperature and spring bloom of S. sinensis is concluded to provide the later cohorts with a higher survival probability through increasing ingestion and growth rates during their post-migration period in spring 2005.     

Nutrient flux in the Rhode River: Tidal transport of microorganisms in brackish marshes

Concentrations of bacteria, chlorophyll a, and several dissolved organic compounds were determined during 11 tidal cycles throughout the year in a high and a low elevation marsh of a brackish tidal estuary. Mean bacterial concentrations were slightly higher in flooding (7·1 × 10⁶ cells ml⁻¹) than in ebbing waters (6·5 × 10⁶ cells ml⁻¹), and there were no differences between marshes. Mean chlorophyll a concentrations were 36·7 μg l⁻¹ in the low marsh and 20·4 μg l⁻¹ in the high marsh. Flux calculations, based on tidal records and measured concentrations, suggested a small net import of bacterial and algal biomass into both marshes. Over the course of individual tidal cycles, concentrations of all parameters were variable and not related to tidal stage. Heterotrophic activity measured by the uptake of ³H-thymidine, was found predominantly in the smallest particle size fractions (< 1·0 μm). Thymidine uptake was correlated with temperature (r = 0·48, P < 0·01), and bacterial productivity was estimated to be 7 to 42 μg Cl⁻¹ day⁻¹.     

Shelf spawning habitat of Emmelichthys nitidus in south-eastern Australia – Implications and suitability for egg-based biomass estimation

The spawning habitat of Emmelichthys nitidus (Emmelichthyidae) in south-eastern Australia is described from vertical ichthyoplankton samples collected along the shelf region off eastern through to south-western Tasmania during peak spawning in October 2005–06. Surveys covered eastern waters in 2005 (38.8–43.5°S), and both eastern and southern waters in 2006 (40.5°S around to 43.5°S off the south-west). Eggs (n = 10,393) and larvae (n = 378) occurred along eastern Tasmania in both years but were rare along southern waters south and westwards of 43.5°S in 2006. Peak egg abundances (1950–2640 per m⁻²) were obtained off north-eastern Tasmania (40.5–41.5°S) between the shelf break and 2.5 nm inshore from the break. Eggs were up to 5-days old, while nearly 95% of larvae were at the early preflexion stage, i.e. close to newly emerged. Average abundances of aged eggs pooled across each survey declined steadily from day-1 to day-5 eggs both in 2005 (97-18) and 2006 (175-34). Moreover, day-1 egg abundances were significantly greater 2.5 nm at either side of the break, including at the break, than in waters ≥5 nm both inshore and offshore from the break. These results, complemented with egg and larval data obtained in shelf waters off New South Wales (NSW; 35.0–37.7°S) in October 2002–03, indicate that the main spawning area of E. nitidus in south-eastern Australia lies between 35.5°S off southern NSW and 43.5°S off south-eastern Tasmania, and that spawning activity declines abruptly south and westwards of 43.5°S around to the south-west coast. In addition, quotient analyses of day-1 egg abundances point to a preferred spawning habitat contained predominantly within a 5 nm corridor along the shelf break, where waters are 125–325 m deep and median temperatures 13.5–14.0 °C. Spawning off eastern Tasmania is timed with the productivity outburst typical of the region during the austral spring, and the temperature increase from the mixing between the southwards advancing, warm East Australian Current and cooler subantarctic water over the shelf. Overall, ichthyoplankton data, coupled with reproductive information from adults trawled off Tasmania, indicate that E. nitidus constitutes a suitable species for the application of the daily egg production method (DEPM) to estimate spawning biomass. This finding, together with evidence in support of a discrete eastern spawning stock extending from southern NSW to southern Tasmania, strengthens the need for DEPM-based biomass estimates of E. nitidus prior to further fishery expansion.     

Predatory impact of an epiphytic hydrozoan in an upwelling area in the Bay of Coliumo (Dichato, Chile)

The natural diet of the epiphytic hydrozoan Obelia geniculata has been studied in an upwelling area in the Bay of Coliumo (Dichato, Chile) during two 24-h cycles. Number of prey per predator and predatory density have been measured. In both cycles more than 78% of the diet consisted of eggs of invertebrates and faecal pellets. Compared with other species of hydrozoans studied in a similar way, O. geniculata had a heterogeneous diet and a capture rate which was scarcely related to the peaks of abundance of its prey. This catch rate was between 632×10³ and 10 393×10³ prey m⁻² day⁻¹, which corresponds to a mean ingestion rate of 113% of the hydranth biomass per day. These results indicate the importance of small-sized benthic suspension feeders in upwelling systems.