Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

Zooplankton associated with the oxygen minimum zone system in the northern upwelling region of Chile during March 2000

Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemical role by promoting carbon flux into the subsurface OMZ (oxygen minimum zone). This work identifies the dominant zooplankton species inhabiting the area influenced by the OMZ in March 2000 off Iquique (20°S, northern Chile). Abundance and vertical distribution studies revealed 17 copepod and 9 euphausiid species distributed between the surface and 600 m at four stations sampled both by day and by night. Some abundant species remained in the well-oxygenated upper layer (30 m), with no evidence of diel vertical migration, apparently restricted by a shallow (40–60 m) oxycline. Other species, however, were found closely associated with the OMZ. The large-sized copepod Eucalanus inermis was found below the oxycline and performed diel vertical migrations into the OMZ, whereas the very abundant Euphausia mucronata performed extensive diel vertical migrations between the surface waters and the core of the OMZ (200 m), even crossing it. A complete assessment of copepods and euphausiids revealed that the whole sampled water column (0–600 m) is occupied by distinct species having well-defined habitats, some of them within the OMZ. Ontogenetic migrations were evident in Eucalanidae and E. mucronata. Estimates of species biomass showed a substantial (>75% of total zooplankton biomass) daily exchange of C between the photic layer and the OMZ. Both E. inermis and E. mucronata can actively exchange about 37.8 g C m⁻² d⁻¹ between the upper well-oxygenated (0–60 m) layer and the deeper (60–600 m) OMZ layer. This migrant biomass may contribute about 7.2 g C m⁻² d⁻¹ to the OMZ system through respiration, mortality, and production of fecal pellets within the OMZ. This movement of zooplankton in and out of the OMZ, mainly as a result of the migratory behavior of E. mucronata, suggests a very efficient mechanism for introducing large amounts of freshly produced carbon into the OMZ system and should, therefore, be considered when establishing C budgets for coastal upwelling systems.     

Vertical changes in abundance,biomass and community structure of copepods down to 3000 m in the southern Bering Sea

Vertical changes in abundance, biomass and community structure of copepods down to 3000 m depth were studied at a single station of the Aleutian Basin of the Bering Sea (53°28′N, 177°00′W, depth 3779 m) on the 14th June 2006. Both abundance and biomass of copepods were greatest near the surface layer and decreased with increase in depth. Abundance and biomass of copepods integrated over 0–3000 m were 1,390,000 inds. m^?2 and 5056 mg C m^?2, respectively. Copepod carcasses occurred throughout the layer, and the carcass:living specimen ratio was the greatest in the oxygen minimum layer (750–100 m, the ratio was 2.3). A total of 72 calanoid copepod species belonging to 34 genera and 15 families occurred in the 0–3000 m water column (Cyclopoida, Harpacticoida and Poecilostomatoida were not identified to species level). Cluster analysis separated calanoid copepod communities into 5 groups (A–E). Each group was separated by depth, and the depth range of each group was at 0–75 m (A), 75–500 m (B), 500–750 m (C), 750–1500 m (D) and 1500–3000 m (E). Copepods were divided into four types based on the feeding pattern: suspension feeders, suspension feeders in diapause, detritivores and carnivores. In terms of abundance the most dominant group was suspension feeders (mainly Cyclopoida) in the epipelagic zone, and detritivores (mainly Poecilostomatoida) were dominant in the meso- and bathypelagic zones. In terms of biomass, suspension feeders in diapause (calanoid copepods Neocalanus spp. and Eucalanus bungii) were the major component (ca. 10–45%), especially in the 250–3000 m depth. These results are compared with the previous studies in the same region and that down to greater depths in the worldwide oceans.     

Spatial patterns of zooplankton and nekton in a hydrothermally active axial valley on Juan de Fuca Ridge

Zooplankton and nekton at 2000 m depth in the axial valley of Endeavour Segment, Juan de Fuca Ridge, show marked variability in abundances in a plane at 20 m above bottom. A remotely operated vehicle flew a gridded rectangle 3.2×0.5 km that included two large high-temperature and two small low-temperature vent fields. Numbers of zooplankton, jellyfish, shrimp and fish were recorded with a video camera, and the abundance patterns were examined with the program SADIE^©. Each organism group displayed a distinctive distribution pattern. Abundance gaps over the high-temperature fields were significant and, for the more abundant copepods, were related to the locations of individual smokers. Pelagic shrimp and macrourid fish abundances were correlated and concentrated around the northern high temperature field. Distinct aggregations of zooplankton and nekton were correlated with the fluid indicators from both the low temperature diffuse effluent and the focused high temperature vents. Patterns were likely established by organism choice that forms aggregations and gaps, and by physical processes that entrain passive particles near vigorous smoker plumes. While enhanced plankton and nekton numbers were not observed over the vent fields, overall abundances in the axial valley may be sustained by production transported from the vent fields on the seafloor.     

Mesoscale physical variability affects zooplankton production in the Labrador Sea

Surface distribution (0–100 m) of zooplankton biomass and specific aminoacyl-tRNA synthetases (AARS) activity, as a proxy of structural growth, were assessed during winter 2002 and spring 2004 in the Labrador Sea. Two fronts formed by strong boundary currents, several anticyclonic eddies and a cyclonic eddy were studied. The spatial contrasts observed in seawater temperature, salinity and fluorescence, associated with those mesoscale structures, affected the distributions of both zooplankton biomass and specific AARS activity, particularly those of the smaller individuals. Production rates of large organisms (200–1000 μm) were significantly related to microzooplankton biomass (63–200 μm), suggesting a cascade effect from hydrography through microzooplankton to large zooplankton. Water masses defined the biomass distribution of the three dominant species: Calanus glacialis was restricted to cold waters on the shelves while Calanus hyperboreus and Calanus finmarchicus were widespread from Canada to Greenland. Zooplankton production was up to ten-fold higher inside anticyclonic eddies than in the surrounding waters. The recent warming tendency observed in the Labrador Sea will likely generate weaker convection and less energetic mesoscale eddies. This may lead to a decrease in zooplankton growth and production in the Labrador basin.     

Zooplankton metabolism and carbon demand at two seamounts in the NE Atlantic

Zooplankton metabolic rates, determined from electron transfer system (ETS) activity, were studied at two seamounts (Seine: 34°N, 14°W, summit depth ∼170 m; Sedlo: 40°N, 27°W, summit depth ∼750 m) in the northeast (NE) Atlantic during three cruises in November 2003, April 2004 and July 2004. ETS activity and respiratory carbon demand were measured for samples taken at seamount and open-ocean locations in order to probe the hypothesis of locally enhanced seamount productivity. ETS activity and biomass revealed no consistent diel patterns of feeding activity and vertical migration at Seine and Sedlo Seamounts. Spatial differences of biomass-specific ETS activity were observed at both seamounts and coincided with differences in food abundance and quality. At Seine Seamount in April 2004, biomass-specific ETS activity was on average higher at the seamount locations compared to the open ocean, though the enhancement was of a lower magnitude than spatial and temporal variability and had no apparent influence on zooplankton respiratory carbon demand or biomass. A persistent pattern of reduced zooplankton biomass above the summit location at Seine Seamount in April 2004 and July 2004 resulted in a local reduction of respiratory carbon demand. At Sedlo Seamount in November 2003, large spatial differences in biomass-specific ETS activity observed at the seamount locations resulted in a large range of respiratory carbon demand at the seamount, but were not reflected in zooplankton biomass. The depth-integrated (0–150 m) median respiratory carbon demand of the zooplankton community estimated from day and night hauls was 2.1 mg C m⁻² d⁻¹ at Seine Seamount (range: 0.3–6.3) and 2.9 mg C m⁻² d⁻¹ at Sedlo Seamount (range: 1.6–12.0). The sporadic nature and low magnitude of locally higher zooplankton respiration rates at the seamounts, which did not result in locally higher zooplankton standing stock biomass, lead us to reject the hypothesis that locally enhanced seamount productivity provides an autochthonous food supply to the resident faunas at Seine and Sedlo Seamounts. Instead, we conclude that the faunas at both seamounts are more likely supported by advection of food from the surrounding ocean.     

Greenland halibut diet in the Northwest Atlantic from 1978 to 2003 as an indicator of ecosystem change

The Northwest Atlantic marine community underwent dramatic changes during the last 30 years, including the collapse of many groundfish stocks and an increase in shrimp populations. Greenland halibut Reinhardtius hippoglossoides is an important commercial species and one of the top fish predators in this system. It is a large, wide-ranging flatfish that is found at depths up to 2200 m and it has an opportunistic diet which makes it a potential candidate for an ecosystem indicator. Analysis of stomach contents of Greenland halibut between 1978 and 2003 indicates that diet composition reflects the major changes in community structure. Over the entire period there was a clear increase in the importance of invertebrates, particularly after 1992. This change was associated with a higher importance of Pandalus shrimp and Gonatus squid and a protracted reliance on zooplankton by predators under 25 cm length. Capelin Mallotus villosus was the dominant prey between 1978 and 1992 for predators in the 12–63 cm range, but its importance dropped off drastically in the mid 1990s. Levels of main prey in the diet of Greenland halibut correlated well with fishery-independent surveys. Greenland halibut sample capelin well, compared to bottom trawl surveys and acoustic surveys. Greenland halibut consumed small shrimp which are not routinely caught by surveys and may be important in deriving information on year classes and growth of shrimp. Our results suggest that Greenland halibut's diet is a useful tracker of ecosystem change.     

Environmental drivers of megafaunal assemblage composition and biomass distribution over mainland and insular slopes of the Balearic Basin (Western Mediterranean)

The influence of mesoscale physical and trophic variables on deep-sea megafauna, a scale of variation often neglected in deep-sea studies, is crucial for understanding their role in the ecosystem. Drivers of megafaunal assemblage composition and biomass distribution have been investigated in two contrasting areas of the Balearic basin in the NW Mediterranean: on the mainland slope (Catalonian coasts) and on the insular slope (North of Mallorca, Balearic Islands). An experimental bottom trawl survey was carried out during summer 2010, at stations in both sub-areas located between 450 and 2200 m water depth. Environmental data were collected simultaneously: near-bottom physical parameters, and the elemental and isotopic composition of sediments. Initially, data were analysed along the whole depth gradient, and then assemblages from the two areas were compared. Analysis of the trawls showed the existence of one group associated with the upper slope (US=450–690 m), another with the middle slope (MS=1000–1300 m) and a third with the lower slope (LS=1400–2200 m). Also, significant differences in the assemblage composition were found between mainland and insular slopes at MS. Dominance by different species was evident when the two areas were compared by SIMPER analysis. The greatest fish biomass was recorded in both areas at 1000–1300 m, a zone linked to minimum temperature and maximum O₂ concentration on the bottom. Near the mainland, fish assemblages were best explained (43% of total variance, DISTLM analysis) by prey availability (gelatinous zooplankton biomass). On the insular slope, trophic webs seemed less complex and were based on vertical input of surface primary production. Decapods, which reached their highest biomass values on the upper slope, were correlated with salinity and temperature in both the areas. However, while hydrographic conditions (temperature and salinity) seemed to be the most important variables over the insular slope, resource availability (gelatinous zooplankton and Calocaris macandreae) predominated and explained 59% of decapod assemblage variation over the mainland slope. Both fish and decapods were linked to net primary production recorded over the mainland 3 months before sampling, while the delay between the input of food from the surface and fish abundance was only 1 month on the insular slope. Our results suggest that trophic relationships over insular slopes probably involve a shorter food chain than over mainland slopes and one that is likely more efficient in terms of energy transfer.     

Zooplankton distribution across Fram Strait in autumn: Are small copepods and protozooplankton important?

We investigated zooplankton distribution in September 2006/2007 at eight stations across Fram Strait in contrasting water masses ranging from cold Polar water to warm Atlantic water. Our main objectives were: (1) to describe the plankton community in the upper 200 m during autumn, and (2) to investigate the importance of small-sized copepods and protozooplankton in an arctic ecosystem when the majority of the large Calanus species had entered diapause. We sampled both with a WP-2 net and Go-Flo bottle and show that small copepods <1 mm are significantly undersampled using a WP-2 net with 90 μm mesh.Small copepods and protozooplankton made a significant contribution both in terms of abundance and total zooplankton biomass at all stations in September, when the large calanoid copepods had left the upper 200 m. The dominating group in the upper 60 m at all stations was Oithona spp. nauplii and their daily estimated grazing potential on the <10 μm phytoplankton ranged from 0.1% to 82% of the standing stock. Both Oithona copepodites and nauplii biomass showed a significantly positive relation with temperature, but not with potential food. Heterotrophic protozooplankton, on the other hand, were most likely bottom-up regulated by the availability of phytoplankton <10 μm. We hypothesise that Oithona nauplii and protozooplankton compete for food and conclude that there was a strong link between the zooplankton community and the microbial food web in Fram Strait.     

Factors influencing the abundance of deep pelagic bioluminescent zooplankton in the Mediterranean Sea

Measurements of the density of deep pelagic bioluminescent zooplankton (BL) were made with the Intensified Silicon Intensifier Target (ISIT) profiler in the Ligurian, Tyrrhenian, Adriatic, Ionian Seas and the Strait of Sicily from ～300 m to near seafloor. Mean BL densities ranged from 2.61 m^?3 at 500–1000 m depth in the Adriatic Sea to 0.01 m^?3 at 4000–5000 m depth in the E Ionian Sea. We investigated drivers of spatial variation of deep pelagic BL density. Linear regression was applied between surface chlorophyll a (Chl a) concentration and underlying BL density. Chl a values were determined from satellite derived 100 km radius composites (six 10-day means per ISIT deployment, over preceding 60 days). At 500–1000 m depth we found a significant positive relationship between mean BL density and mean Chl a in the period prior to 0–10 days (at 1% level) and prior to 10–40 days (at 5% level). Beyond 40 days no relationship between BL density and Chl a was found at this depth. At depths 1000–1500 m BL density values were low and no significant relationship with Chl a was detected. Generalised additive modelling (GAM) was used to assess the influence of benthic hotspots (seamount; cold water coral mound; mud volcano) on overlying BL density. A reduction in BL density was found downstream of the Palinuro seamount from 300 to 600 m. No effect on BL density in the overlying water column was detected from the presence of cold water corals. Higher BL densities were detected over the W Madonna dello Ionio mud volcano than at other sites sampled in the NW Ionian Sea. We find surface Chl a to be a good predictor of BL density in the mesopelagic zone; below this depth we hypothesise that processes affecting the efficiency of particle export to deep water may exert greater influence on BL density.     

Spring distribution of larger (>64 μm) protozoans in the Atlantic sector of the Southern Ocean

Distribution of larger protozoans (armoured dinoflagellates, tintinnids, heliozoans, radiolarians and foraminiferans >64 μm) is presented for three major water masses of the Southern Ocean: the Polar Front region (PFr), the southern Antarctic Circumpolar Current (southern ACC) and the northern Weddell Gyre. Sampling took place during the SO-JGOFS cruise ANT X/6 of R/V Polarstern (October–November 1992) along a meridional transect at 6°W between 48°00′S and 59°30′S. Multinet samples (64 μm mesh size) were taken at six stations from the surface down to 500 m depth at five different depth intervals. In the upper 100 m of the water column abundances of larger protozoans varied between 94 and 10,930 ind. m^–3, with highest abundances in the PFr, where phytoplankton blooms occurred, and lowest values in the Antarctic Circumpolar Current–Weddell Gyre Boundary (AWB). Foraminiferans and polycystine and smaller (<300 μm) phaeodarian radiolarians dominated larger protozoan assemblages in the PFr. In open water of the southern ACC, tintinnids, armoured dinoflagellates, foraminiferans and smaller (<300 μm) phaeodarian radiolarians were equally important. The heliozoans Sticholonche spp. and nassellarian radiolarians dominated assemblages in the Weddell Gyre and AWB. Larger protozoan biomasses ranged between 2 and 674 μg C m⁻³ and were always dominated by larger (>300 μm) phaeodarians. Highest biomasses were found in the AWB between 200 and 500 m depth. Standing stocks of larger protozoans constituted a negligible fraction of zooplankton biomass in the upper 200 m of the water column. In deeper layers of the ice-covered Weddell Gyre and AWB their biomasses, dominated by larger (>300 μm) phaeodarians, was significant contributing up to 45% to total larger protozoan and metazoan biomass. Analysis of correlation between distribution patterns and environmental conditions at the stations sampled indicate that spring distribution patterns of heterotrophic armoured dinoflagellates, polycystine radiolarians and foraminiferans follow productivity in the water column. Of the protozoan groups studied the smaller (<300 μm) phaeodarian radiolarians also showed a significant correlation with productivity during spring, however, results from previous studies do not suggest a consistent pattern. Spring distribution patterns of other larger protozoans were not related to differences in productivity in the water column, and effects such as ice-cover, grazing or silica limitation might be determining. Dead radiolarian skeletons constituted on average 27, 8 and 11% of the population of nassellarians, spumellarians and smaller (<300 μm) phaeodarians, respectively. The contribution of dead radiolarian skeletons to total radiolarian stocks varied with depth and water mass. Differences between live and skeleton assemblages composition were observed. These differences should be taken into consideration when interpreting the geological record.     

The habitat of mesopelagic scyphomedusae in Monterey Bay,California

In the mesopelagic zone, at depths of 200–1000 m in the Monterey Submarine Canyon, CA, medusae in three genera of scyphozoa, Atolla, Periphylla and Poralia, were observed, videotaped and collected over a 9-year period (1990–1998). Environmental data were obtained simultaneously using a remotely operated vehicle (ROV) with sensors for depth, temperature, salinity and dissolved oxygen. Shipboard measurements of these same properties at two reference stations in the region defined the local water masses and helped identify species niches using the metric of spiciness and oxygen levels of the waters in which medusae were visually “captured”. The most abundant genus of mesopelagic scyphomedusae was Atolla, found associated most strongly with the spicy (warm, salty) waters of the California Undercurrent, usually above the core of the oxygen minimum zone (OMZ; O₂>0.5 ml/l). The least abundant mesopelagic scyphomedusa was Periphylla, which occurred in more variable waters, including those with a greater contribution of fresher, colder (less spicy), subarctic water and, hence, most like those at the offshore California Current station in the most depleted oxygen zone (averaging O₂ <0.3 ml/l). Poralia was mostly confined to the densest, coldest water, with peak abundance at the lower boundary of the OMZ (i.e., 0.3< O₂<0.5 ml/l). These spiciness measures on local isopycnal surfaces within the mesopelagic zone, supported by data on dissolved oxygen concentrations, indicate highly significant but fine-scale habitat differences in species habitats in Central California waters. This in situ investigation appears to be one of only a few studies to document fine-scale, water mass affinities of mesopelagic zooplankton.     

High-frequency fluxes of labile compounds in the central Ligurian Sea,northwestern Mediterranean

Sinking particles were collected every 4 h with drifting sediment traps deployed at 200 m depth in May 1995 in a 1-D vertical system during the DYNAPROC observations in the northwestern Mediterranean sea. POC, proteins, glucosamine and lipid classes were used as indicators of the intensity and quality of the particle flux. The roles of day/night cycle and wind on the particle flux were examined. The transient regime of production from late spring bloom to pre-oligotrophy determined the flux intensity and quality. POC fluxes decreased from, on average, 34 to 11 mg m⁻² d⁻¹, representing 6–14% of the primary production under late spring bloom conditions to 1–2% under pre-oligotrophic conditions. Total protein and chloroplast lipid fluxes correlated with POC and reflected the input of algal biomass into the traps. As the season proceeded, changes in the biochemical composition of the exported material were observed. The C/N ratio rose from 7.8 to 12. Increases of serine (10–28% of total proteins), total lipids (7–9 to 14–28% of POC) and reserve lipids (1–5 to 5–22% of total lipids) were noticeable, whereas total protein content in POC decreased (20–27 to 18–7%). N-acetyl glucosamine, a tracer of fecal pellet flux, showed that zooplankton grazing was a major vector of downward export during the decaying bloom. Against this background pattern, episodic events specifically increased the flux, modifying the quality and the settling velocity of particles. Day/night signals in biotracers (POC, N-acetyl glucosamine, protein and chloroplast lipids) showed that zooplankton migrations were responsible for sedimentation of fresh material through fast sinking particles (V=170–180 m d⁻¹) at night. Periodic signatures of re-processed material (high lipolysis and bacterial biomass indices) suggested that other zooplankton fecal pellets or small aggregates, probably of lower settling velocities (V<170 m d⁻¹), contributed to the flux during calm periods. At the beginning of the experiment, during the development of a prymnesiophyte bloom in the upper layers, the sterol signal with no periodicity enabled us to estimate high particle settling velocities (⩾600 m d⁻¹) likely related to large aggregate formation. A wind event increased biotracer fluxes (POC, protein, chloroplast lipids). The rapid transmission of surface signals through extremely fast sinking particles could be a general feature of particle fluxes in marine areas unaffected by horizontal advection.     

Diel variations of the bathymetric distribution of zooplankton groups and biomass in Cap-Ferret Canyon,France

The bathymetric distribution, abundance and diel vertical migrations (DVM) of zooplankton were investigated along the axis of the Cap-Ferret Canyon (Bay of Biscay, French Atlantic coast) by a consecutive series of synchronous net hauls that sampled the whole water column (0–2000 m in depth) during a diel cycle. The distribution of appendicularians (maximum 189 individuals m⁻³), cladocerans (maximum 287 individuals m⁻³), copepods (copepods<4 mm, maximum 773 individuals m⁻³, copepods>4 mm, maximum 13 individuals m⁻³), ostracods (maximum 8 individuals m⁻³), siphonophores (maximum >2 individuals m⁻³) and peracarids (maximum >600 individuals 1000 m⁻³) were analysed and represented by isoline diagrams. The biomass of total zooplankton (maximum 18419 μg C m⁻³, 3780 μg N m⁻³) and large copepods (>4 mm maximum 2256 μg C m⁻³, 425 μg N m⁻³) also were determined. Vertical migration was absent or affected only the epipelagic zone for appendicularians, cladocerans, small copepods and siphonophores. Average amplitude of vertical migration was about 400–500 m for ostracods, some hyperiids and mysids, and large copepods, which were often present in the epipelagic, mesopelagic, and bathypelagic zones. Large copepods can constitute more than 80% of the biomass corresponding to total zooplankton. They may play an important role in the active vertical transfer of carbon and nitrogen.     

Ascending and descending particle flux from hydrothermal plumes at Endeavour Segment,Juan de Fuca Ridge

Bio-acoustic surveys and associated zooplankton net tows have documented anomalously high concentrations of zooplankton within a 100 m layer above the hydrothermal plumes at Endeavour Segment, Juan de Fuca Ridge. These and other data suggest that congregating epi-plume zooplankton are exploiting a food substrate associated with the hydrothermal plume. Ascending, organic-rich particles could provide a connection. Consequently, two paired sequentially sampling ascending and descending particle flux traps and a current meter were deployed on each of three moorings from July 1994 to May 1995. Mooring sites included an on-axis site (OAS; 47°57.0′N, 129°05.7′W) near the main Endeavour vent field, a “down-current” site 3 km west of the main vent field (WS), and a third background station 43 km northeast of the vent field (ES). Significant ascending and descending particle fluxes were measured at all sites and depths. Lipid analyses indicated that ascending POC was derived from mid-depth and deep zooplankton whereas descending POC also contained a component of photosynthetically derived products from the sea surface. Highest ascending POC fluxes were found at the hydrothermal plume-swept sites (OAS and WS). The limited data available, however, precludes an unequivocal conclusion that hydrothermal processes contribute to the ascending flux of organic carbon at each site. Highest ascending to descending POC flux ratios were also found at WS. Observed trends in POC, PMn/PTi, and PFe/PTi clearly support a hydrothermal component to the descending flux at the plume-swept WS site (no descending data was recovered at OAS) but not at the background ES site. Alternative explanations for ascending particle data are discussed. First-order calculations for the organic carbon input (5–22 mg C m⁻² d⁻¹) required to sustain observed epi-plume zooplankton anomalies at Endeavour are comparable both to measured total POC flux to epi-plume depths (2–5 mg C m⁻² d⁻¹: combined hydrothermal and surface derived organic carbon) and to estimates of the total potential in situ organic carbon production (2–9 mg C m⁻² d⁻¹) from microbial oxidation of hydrothermal plume H₂, CH₄ and NH₄⁺.     

Zooplankton structure and grazing in the Atlantic sector of the Southern Ocean in late austral summer 1993: Part 1. Ecological zonation

The composition, distribution, abundance, biomass and size structure of mesozooplankton, collected using Bongo nets in the top 300 m layer along a transect between the Antarctic continent and Cape Town, were investigated during the second South African Antarctic Marine Ecosystem Study (SAAMES II) in Jan.–Feb. 1993. Small (<10 mm) and medium (20–50 mm) size groups of zooplankton consistently dominated across the Southern Ocean. The highest zooplankton densities were recorded at the Antarctic Polar Front (APF) and at the Subtropical Convergence (STC). Minor peaks in zooplankton densities were observed in the southern vicinity of the Subantarctic Front (SAF) and APF. Elevated zooplankton stocks were also found within the Marginal Ice Zone (MIZ) and the Polar Frontal Zone. The lowest densities were recorded in the permanently open zone (MIZ–APF) and in the Subantarctic zone (SAF–STC). Copepods were generally important along the entire transect and formed the bulk of zooplankton stock within the MIZ and in the Polar Frontal Zone (APF–SAF), accounting for at least 40–95% of total abundance and biomass. Euphausiids were also a prominent group along the transect. Their contribution was highest (up to 80% of total biomass) between the MIZ and the APF, mainly because of the occurrence of swarms of the Antarctic krill Euphausia superba. Tunicates, Pyrosoma sp. and Salpa fusiformis, were found in great numbers only in the region of the STC and further north, while Salpa thompsoni was abundant at the southern boundary of the APF. Chaetognaths dominated samples numerically and by mass in the Subantarctic Zone. Results obtained from cluster and ordination analyses show that zooplankton community structure was well correlated with the position of various biogeographical zones separated by the main frontal systems of the Southern Ocean. Two major groupings of stations, separated by the SAF, were identified in these analyses. This front separated the Antarctic and the subantarctic/subtropical assemblages, confirming its important role as a biogeographical boundary.     

Microbial biomass and viral infections of heterotrophic prokaryotes in the sub-surface layer of the central Arctic Ocean

Seawater samples were collected for microbial analyses between 55 and 235 m depth across the Arctic Ocean during the SCICEX 97 expedition (03 September–02 October 1997) using a nuclear submarine as a research platform. Abundances of prokaryotes (range 0.043–0.47×10⁹ dm⁻³) and viruses (range 0.68–11×10⁹ dm⁻³) were correlated (r=0.66, n=150) with an average virus:prokaryote ratio of 26 (range 5–70). Biomass of prokaryotes integrated from 55 to 235 m ranged from 0.27 to 0.85 g C m⁻² exceeding that of phytoplankton (0.005–0.2 g C m⁻²) or viruses (0.02–0.05 g C m⁻²) over the same depth range by an order of magnitude on average. Using transmission electron microscopy (TEM), we estimated that 0.5% of the prokaryote community on average (range 0–1.4%) was visibly infected with viruses, which suggests that very little of prokaryotic secondary production was lost due to viral lysis. Intracellular viruses ranged from 5 to >200/cell, with an average apparent burst size of 45±38 (mean±s.d.; n=45). TEM also revealed the presence of putative metal-precipitating bacteria in 8 of 13 samples, which averaged 0.3% of the total prokaryote community (range 0–1%). If these prokaryotes are accessible to protistan grazers, the Fe and Mn associated with their capsules might be an important source of trace metals to the planktonic food web. After combining our abundance and mortality data with data from the literature, we conclude that the biomass of prokaryoplankton exceeds that of phytoplankton when averaged over the upper 250 m of the central Arctic Ocean and that the fate of this biomass is poorly understood.     

Feeding ecology of mesopelagic zooplankton of the subtropical and subarctic North Pacific Ocean determined with fatty acid biomarkers

Mesopelagic zooplankton may meet their nutritional and metabolic requirements in a number of ways including consumption of sinking particles, carnivory, and vertical migration. How these feeding modes change with depth or location, however, is poorly known. We analyzed fatty acid (FA) profiles to characterize zooplankton diet and large particle (>51 μm) composition in the mesopelagic zone (base of euphotic zone ?1000 m) at two contrasting time-series sites in the subarctic (station K2) and subtropical (station ALOHA) Pacific Ocean. Total FA concentration was 15.5 times higher in zooplankton tissue at K2, largely due to FA storage by seasonal vertical migrators such as Neocalanus and Eucalanus. FA biomarkers specific to herbivory implied a higher plant-derived food source at mesotrophic K2 than at oligotrophic ALOHA. Zooplankton FA biomarkers specific to dinoflagellates and diatoms indicated that diatoms, and to a lesser extent, dinoflagellates were important food sources at K2. At ALOHA, dinoflagellate FAs were more prominent. Bacteria-specific FA biomarkers in zooplankton tissue were used as an indicator of particle feeding, and peaks were recorded at depths where known particle feeders were present at ALOHA (e.g., ostracods at 100–300 m). In contrast, depth profiles of bacterial FA were relatively constant with depth at K2. Diatom, dinoflagellate, and bacterial biomarkers were found in similar proportions in both zooplankton and particles with depth at both locations, providing additional evidence that mesopelagic zooplankton consume sinking particles. Carnivory indices were higher and increased significantly with depth at ALOHA, and exhibited distinct peaks at K2, representing an increase in dependence on other zooplankton for food in deep waters. Our results indicate that feeding ecology changes with depth as well as by location. These changes in zooplankton feeding ecology from the surface through the mesopelagic zone, and between contrasting environments, have important consequences for the quality and quantity of organic material available to deeper pelagic and benthic food webs, and for organic matter sequestration.     

Vertical distribution of zooplankton in Canary Island waters: implications for export flux

Zooplankton biomass, gut fluorescence and electron transfer system (ETS) activity were measured in vertical profiles (0–900 m) in two different size classes (<1 and >1 mm) in Canary Island waters. Both size fractions displayed a typical pattern of distribution with higher biomass, gut fluorescence and ETS in the shallower layers at night. By day, however, the vertical distribution varied between the size fractions, with higher biomass of the small fraction in the 0–200 m and a layer of large organisms at depth (∼500 m). For both size fractions, average ETS activity was higher by day than at night at depths between 200 and 600 m. Similarly, gut fluorescence was slightly higher by day below 200 m. The downward export of respiratory carbon was 1.92 and 4.29 mg C m⁻² d⁻¹ for samples obtained southwest of Gran Canaria Island and west of Tenerife Island respectively, being 2.68 mg C m⁻² d⁻¹ for the whole area. These values represented 16–45% (22–28% for the area) of the calculated passive particulate export production resulting from primary production. The estimated “gut flux” accounted for 0.35 (western zone) and 2.37 mg C m⁻² d⁻¹ (southwest of Gran Canaria), being 1.28 mg C m⁻² d⁻¹ for the whole area and represented between 3 and 25% (11–14% for the whole area) of the estimated passive particle export flux. These results agree with previous estimates and suggest that diel-migrant zooplankton can play an important role in the downward flux of carbon.     

Impact of submarine hydrothermal vents on the metal composition of krill and its excretion products

We have measured the metal composition and estimated the excretion rate of trace elements (Ag, Cd, Co, Cu, Ni, Pb, V and Zn) by Antarctic krill (Euphausia superba) in three locations (two located within a submarine hydrothermal vent field and one away from it) along the Antarctic Peninsula region of the Southern Ocean. Results indicated that krill excreted large amounts of Ag, Cu, Pb and Zn, (range: 1.9–41.2 pmol Ag g DW^? 1 h^? 1, 15.3–26.8 nmol Cu g DW^? 1 h^? 1, 308.7–1118.3 pmol Pb g DW^? 1 h^? 1 and 24.4–76.5 nmol Zn g DW^? 1 h^? 1), compared with the non-significant or undetectable release rates of Cd, Co, Ni and V. The metal composition of the excreted material from krill collected in the area of hydrothermal activity was similar to the metal composition reported for suspended particles emitted from those vents. Our results suggest that krill recycling of Ag, Cu, Pb and Zn could potentially influence trace metal concentrations in the water column of the Bransfield region of the Southern Ocean, and that the original source of metals to these waters may be hydrothermal vents.