Meso- and microzooplankton responses to an in situ iron fertilization experiment (SEEDS II) in the northwest subarctic Pacific

A mesoscale iron fertilization experiment was carried out in the western subarctic Pacific during summer 2004. The iron-patch was traced for 26 days after the enrichment, and the abundance and behavior of meso- and microzooplankton was compared with those outside of the patch. The surface chlorophyll-a concentration in the patch was high between days 10 and 13 (2.5 mg m⁻³) and decreased to the initial level after day 20. Microzooplankton grazing rates, estimated by a dilution method, was mostly balanced with phytoplankton growth rates throughout the observed period. Dominant mesozooplankton species in the upper 200 m were copepods: dominated by Eucalanus bungii, Neocalanus plumchrus and Metridia pacifica. Species composition did not change in the patch over the observation period. The copepod biomass was 3–5 times higher than in Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study (SEEDS), the previous iron-enrichment experiment in the same area, before the bloom, and exponentially increased both inside and outside the patch, which was mainly brought by the development of N. plumchrus. The development rates of N. plumchrus were not significantly different between inside and outside the patch. Estimated grazing rate suggest that the copepod grazing was main cause of the low accumulation of phytoplankton biomass, and dominance of grazing-resistant organisms such as large ciliates, large diatoms and diatoms with extremely long setae. “Arrested migration” for M. pacifica and upward shift of vertical distribution by E. bungii were observed during the bloom period, even if the accumulation of phytoplankton biomass was very low compared to other iron-enrichment experiments. These results indicate that the copepod grazing shaped the food-web structure of the lower trophic levels (biomass and species composition) in SEEDS II.     

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.     

Seasonal and spatial variations of zooplankton in the central and southeastern Bering Sea during the mid-1990s

Hydrographic and plankton surveys were conducted over the basin and slope of the southeastern Bering Sea during April, June/July and September of 1994 and in June/July 1995, and seasonal and spatial variations of zooplankton community were investigated in relation to the oceanographic conditions. In July 1994, sea surface temperature (SST) ranged 5.3–8.7 °C, and the thermocline was between 30 and 50 m. In July 1995, however, SST was warmer (7.3–12.4 °C), and the thermocline was shallower (20–30 m). The thermal front at the shelf was also stronger in July 1995 than in July 1994. Surface salinity was higher in 1994 than 1995. A total of 17 taxonomic groups of zooplankton were identified from the plankton samples. In 1994, the highest density was observed in September. Copepods were the major taxon during all surveys. While some taxa such as euphausiids, ostracods, and Neocalanus spp. were most abundant in spring, others such as Calanus spp., Metridia pacifica, chaetognaths, and pteropods were most abundant in September. Adults and late-stage copepodites of Eucalanus bungii were abundant in spring, and were replaced by 1st–3rd stages of copepodites in summer. Zooplankton density was ca. 4 times higher in 1995 than in 1994, in part because of warm water temperature.     

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.     

Mesozooplankton community characteristics in the NE subarctic Pacific

Mesozooplankton biomass, species composition, abundance, and vertical distribution were determined along a transect from the continental slope off the west coast of Canada to Ocean Station Papa (OSP) in the open-ocean waters of the NE subarctic Pacific as part of the Canadian Joint Global Ocean Flux Study of this area. All of these measurements had distinct seasonal patterns. At OSP biomass peaked in spring, coincident with the annual biomass maximum of large copepods of the genus Neocalanus. Early copepodites of these copepods were present in surface waters at all stations along the transect in winter, but N. plumchrus and N. flemingeri copepodites were only at the offshore stations in spring. This indicated that these large copepods had completed the growth phase of their life cycle slower in the open ocean than closer to shore where they had already descended to deep water by May or June. Summer biomass was low compared to the spring peak. The summer mesozooplankton abundance was similar to the springtime abundance, but the composition had changed from large-bodied copepods in the spring to small copepods and fewer non-copepod taxa in the summer, which accounts for the reduction in total biomass. Winter biomass was the lowest of the year. Winter species composition was similar to summer except for the appearance of juvenile stages of the genera Neocalanus and Calanus. Diel changes in biomass in the upper 150 m were found in summer but not in winter or spring. Vertical distributions of copepods were often distinct, with closely related species occupying different depth strata. Measurements of wet weight at OSP were higher than the long-term mean wet weight during winter and spring, and lower during summer.     

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.     

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.     

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.     

Mesozooplankton responses to iron-fertilization in the western subarctic Pacific (SEEDS2001)

A mesoscale iron-fertilization experiment was carried out in the western subarctic Pacific during summer 2001. The iron-patch was traced for 14 days after the fertilization, and the abundance and behavior of mesozooplankton were compared with those outside of the patch. The phytoplankton biomass in the patch rapidly increased to over 15 times the initial level by the later half of the observation period, and was composed of large-sized (>10 mm), centric diatoms. Dominant zooplankton species in the upper 200-m depth were large copepods: Neocalanus plumchrus, Neocalanus cristatus, Eucalanus bungii and Metridia pacifica. Mesozoplankton biomass as well as species composition did not change significantly in the patch over the observation period. Furthermore, no changes of vertical distribution or diel vertical migration were observed for any species or stages of mesozooplankton throughout the observation period. However, the abundance of the first copepodite stages of N. plumchrus and E. bungii increased several fold in the patch after the diatom bloom formation compared to the densities outside the patch. The increases of both species are considered to be due to lowered mortality during the egg and nauplius stages. Spawning of N. plumchrus takes place at depth using lipid storage, while spawning of E. bungii takes place in the surface layer supported by grazing. These facts suggest that the relative importance of nauplii in the diets of the large copepods was decreased in the patch by the diatom bloom. Gut-pigment contents of dominant copepods in the patch increased 4–18 times, and the maximum values were observed during the bloom peak. However, the grazing impact on phytoplankton was low throughout the experiment, especially during the bloom period (<6% of the primary production).     

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.     

Zooplankton inside an Arctic Ocean cold-core eddy: Probable origin and fate

In September 2004, an extensive survey of a cold-core eddy in the Canada Basin, western Arctic was carried out with high-horizontal-resolution physical and chemical sampling and lower-horizontal-resolution biological sampling. The eddy was located over the continental slope north of the Chukchi Shelf and had a radius of ∼8 km. Its core was centered at a depth of ∼160 m. Water mass characteristics and the presence of copepods from the North Pacific Ocean (Neocalanus flemingeri and Metridia pacifica) demonstrated that the core contained water of Pacific origin. Vertical distributions of zooplankton were associated with the physical structure of the water column. For most taxa, concentrations in the eddy core were elevated compared with those in similar density water in the surrounding Basin. Based on tracer-age estimates and previous observations of eddy formation, the eddy is believed to have been formed during the previous spring/summer from the Chukchi shelfbreak jet. Surprisingly, the eddy also contained elevated abundances of Arctic-origin copepods (Metridia longa and Calanus glacialis). Analysis of a shelf–basin transect occupied in the region in August 2004 showed that these species were present in high abundances in relatively shallow water (50 m) inshore of the shelfbreak due to upwelling of deeper basin water, and copepods, onto the shelf in response to easterly winds. If the formation of the observed eddy occurred during, or shortly after, a period of such winds, upwelled Arctic-origin copepods on the shelf might have been entrained into the feature. Our observations suggest that formation and subsequent migration of such eddies may provide a mechanism for transporting zooplankton from the Chukchi Shelf into the interior Canada Basin. The periodic input of high abundances of zooplankton from productive shelf areas could affect food webs in the less productive basin.     

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.     

Trophic relationships at intrannual spatial and temporal scales of macro and megafauna around a submarine canyon off the Catalonian coast (western Mediterranean)

The spatial and temporal changes of near-bottom macrofauna (suprabenthos and macroplankton) and the trophic relationships of megabenthic decapod crustaceans were analyzed off the Catalonian coasts (western Mediterranean) around Berenguera submarine canyon in four periods (April and December 1991, March and July 1992) and four zones (within Berenguera Canyon at ca. 450 m, and on adjacent slope at ca. 400, 600 m and 1200 m). In March 1992, we found the highest macrofauna abundance and the smallest sizes in the canyon, suggesting a positive effect of river discharges on suprabenthos recruitment. By contrast, macroplankton (decapods, fishes and euphausiids) did not show higher recruitment into canyons. After analyzing the diet of 23 decapod crustaceans, we found a significant segregation between guilds feeding on zooplankton and on benthos. Zooplankton (euphausiids and Pasiphaeidae) and infauna (polychaetes, Calocaris macandreae and ophiuoroids) were consistently the main prey exploited by decapod crustaceans around Berenguera Canyon. We also found some macrophyte (Posidonia oceanica) consumption, which was higher in periods of water column homogeneity (winter–spring and late autumn). Positive correlations between decapods' gut fullness (F) and decapod abundance indicate feeding aggregations, while positive correlations were also found between F and Llobregat River (situated ca. 18 km from Berenguera head) flow 1 to 2 months before sampling. Increases in F were delayed only 1 month when zooplankton feeders were analyzed alone, while benthos feeders did not show significant relationships with any environmental variables. That indicates that the response of megabenthic decapods feeding on benthos to environmental shifts is slower than that of zooplankton feeders. The importance of river flows in enhancing food supply of macro- and megabenthos dwelling close to submarine canyons was apparent, with a delay in the fauna response of 0–2 months after river flow peaks.     

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.     

Distribution and relative importance of jellyfish in a region of hydrothermal venting

Net sampling to 3000 m depth at Endeavour Ridge in the northeast Pacific in July 1991–1994 shows that medusae in the immediate vicinity of the hydrothermal vent fields often make up a larger proportion of the total zooplankton abundance and biomass from mesopelagic to bathypelagic depths than in the surrounding waters. This was particularly evident in the dominant Trachymedusae, and least evident in the siphonophores. In addition, the large red Scyphomedusa Stygiomedusa gigantea was a major biomass component in the region of the deep (1000–1800 m depth) migrating scattering layers at the vent field, but was not found in any net tows greater than 10 km away from vents. There is no concurrent increase in relative or percent biomass of fish or chaetognaths, which are the other major predators in the community. We hypothesize that predaceous medusae respond opportunistically to the enhanced zooplankton biomass throughout the water column around vents in spring to early summer, in a way that other predators do not.     

Estimating copepods' abundance with paired nets: Implications of mesh size for population studies

Three representative copepods occurring in the southern Patagonian shelf, i.e., Calanus australis, Drepanopus forcipatus and Oithona helgolandica were sampled by the first time with paired nets of 66 and 150 µm mesh size. The stage-specific estimates of abundance of their populations were statistically analyzed to assess differences in the catchability by both plankton nets. Differences between nets were significant only for smaller developmental stages and species, with higher catchabilities by the 66 µm net, while no differences were detected for most of the stages of the medium and large size species. A significant effect of the spatial distribution on the estimates of abundance was detected for the majority of the species and stages but the interaction between mesh size and spatial distribution was not significant. This means that differences between both nets were maintained across the latitudinal gradient. Our results strongly suggest the convenience of using the 66 µm plankton net to estimate the abundance of the three copepods' populations. Furthermore, we propose a correction factor to adjust past estimates of abundances from 150 µm mesh collections.     

Fragmentation of marine snow by swimming macrozooplankton: A new process impacting carbon cycling in the sea

Comparisons of the abundances and size distributions of marine snow (aggregated particles >0.5 mm in diameter) in the upper 100 m of the water column at ten stations off Southern California in the late afternoon with those in the same parcel of water the following morning, after nocturnal vertical migration by zooplankton had occurred, revealed the existence of a previously undescribed process affecting marine particle dynamics. Aggregate abundances increased overnight and changes were positively and significantly correlated only with the abundance of the common euphausiid, Euphausia pacifica, and with no other biological or physical factor. Moreover, mean aggregate size decreased and aggregate size distributions shifted toward smaller size classes when euphausiids were abundant. The only conclusion consistent with these findings was that euphausiids were physically disaggregating marine snow into smaller, more numerous aggregates through shear stresses generated while swimming. Video-recording of both tethered and free-swimming E. pacifica in the laboratory dramatically confirmed that aggregates passing within 8–10 mm of the animal's abdomen were fragmented either by entrainment and direct impact with the beating pleopods or by eddies generated during swimming. At the abundances observed in this study, swimming E. pacifica would have sufficiently disturbed 3–33% of the water column each night to disrupt the aggregates contained therein. This is the first evidence for the fragmentation of large particles by the swimming activities of zooplankton and suggests that macrozooplankton and micronekton play a significant role in the particle dynamics of the water column regardless of whether they consume particles or not. Disaggregation of marine snow by swimming and migrating animals may alter the sizes of particles available to grazers and microbial colonizers and reduce the flux of particulate carbon by generating smaller particles, which potentially sink more slowly and reside longer in the water column. This newly discovered process reduces carbon flux while simultaneously conserving carbon and provides a previously unconsidered link between animal behavior and the biogeochemistry of the sea. It may help explain the exponential reduction in particle flux with depth observed in parts of the ocean and help balance oceanic carbon models.     

Distribution of bacterial abundance and cell-specific nucleic acid content in the Northeast Pacific Ocean

We tested the idea that bacterial cells with high nucleic acid content (HNA cells) are the active component of marine bacterioplankton assemblages, while bacteria with low nucleic acid content (LNA cells) are inactive, with a large data set (>1700 discrete samples) based on flow cytometric analysis of bacterioplankton in the Northeast Pacific Ocean off the coast of Oregon and northern California, USA. Samples were collected in the upper 150 m of the water column from the coast to 250 km offshore during 14 cruises from March 2001 to September 2003. During this period, a wide range of trophic states was encountered, from dense diatom blooms (chlorophyll-a concentrations up to 43 μg l⁻¹) at shelf stations during upwelling season (March–September) to lower chlorophyll-a concentrations (0.1–5 μg l⁻¹) during winter (November–February) and at basin stations (>1700 m depth). We found only weakly positive relations of log total bacterial abundance to log chlorophyll-a concentration (as a proxy for availability of organic substrate), and of HNA bacteria as a fraction of total bacteria to log chlorophyll-a. Abundance of HNA and LNA bacteria co-varied positively in all regions, although HNA bacteria were more responsive to high phytoplankton biomass in shelf waters than in slope and basin waters. Since LNA cell abundance in general showed responses similar to those of HNA cell abundance to changes in phytoplankton biomass, our data do not support the hypothesis that HNA cells are the sole active component of marine bacterioplankton.     

The seasonal succession of zooplankton in the Southern Ocean south of Australia,part I: The seasonal ice zone

Between November 2001 and March 2002 an Australian/Japanese collaborative study completed six passes of a transect line in the Seasonal-Ice Zone (south of 62°S) along 140°E. Zooplankton samples were collected with a NORPAC net on 22–28 November, and a Continuous Plankton Recorder on 10–15 January, 11–12 February, 19–22 February, 25–26 February, and 10–11 March. Zooplankton densities were lowest on 22–28 November (ave=61 individuals (ind) m⁻³), when almost the entire transect was covered by sea ice. By 10–15 January sea surface temperature had increased by ∼2 °C across the transect line, and the study area was ice-free. Total zooplankton abundance had increased to maximum levels for the season (ave=1301 ind m⁻³; max=1979 ind m⁻³), dominated by a “Peak Community” comprising Oithona similis, Ctenocalanus citer, Clausocalanus laticeps, foraminiferans, Limacina spp., appendicularians, Rhincalanus gigas and large calanoid copepodites (C1–3). Total densities declined on each subsequent transect, returning to an average of 169 ind m⁻³ on 10–11 March. The seasonal density decline was due to the decline in densities of “Peak Community” taxa, but coincided with the rise of Euphausia superba larvae into the surface waters, increased densities of Salpa thompsoni, and an increased contribution of C4 to adult stages to the populations of Calanoides acutus, Calanus propinquus and Calanus simillimus. The seasonal community succession appeared to be influenced by the low sea ice extent and southward projection of the ACC in this region. The relatively warm ACC waters, together with low krill biomass, favoured high densities of small grazers during the January/February bloom conditions. The persistence of relatively warm surface waters in March and the seasonal decrease in chlorophyll a biomass provided favorable conditions for salps, which were able to penetrate south of the Southern Boundary.