Geographical Variation of Body Size of Neocalanus cristatus, N. plumchrus and N. flemingeri in the Subarctic Pacific and Its Marginal Seas: Implications for the Origin of Large Form of N. flemingeri in the Oyashio Area

Size distributions of Neocalanus cristatus, N. flemingeri and N. plumchrus were investigated in the eastern and the western subarctic gyres and three marginal seas of the North Pacific during the diapause period to examine the geographical variation in body size of Neocalanus species and to clarify the origin of the large biennial N. flemingeri which has been observed in the Oyashio region. There were significant among region variations in body sizes for all three species of Neocalanus. Generally, the body sizes of the copepods were larger in the marginal seas and marginal areas of the open ocean. In the open ocean, the body sizes increased westward. These patterns of variation in the body sizes roughly correlated with local food availability. Distribution of biennial N. flemingeri was restricted to the Sea of Japan, the Okhotsk Sea and the Oyashio region. The large-sized biennial N. flemingeri were abundantly observed in the Okhotsk Sea, and the medium-sized biennial individuals were observed in the Sea of Japan. These facts strongly suggest that the large biennial N. flemingeri in the Oyashio region are advected from the Okhotsk Sea.     

Geographical Variations in Prosome Length and Body Weight of Neocalanus Copepods in the North Pacific

Geographical variations in prosome length and body weight of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were investigated on samples from North-South and East-West transects in the North Pacific during spring to early summer in 1998 and 1999. Southward and eastward increasing patterns were pronounced for water temperature, although no significant pattern was observed for chlorophyll a concentrations. All Neocalanus species showed large geographical variations in prosome length and body weight, being smaller in the southern and eastern waters. Comparing the relationship between prosome length and body weight, large deviations (lower body weight at a given prosome length) were evident for the eastern specimens of N. cristatus and N. plumchrus. In stepwise regression analysis, the geographical variations of prosome length and body weight revealed a significantly negative correlation with temperature variations. These results suggest that temperature is a more important environmental factor than chlorophyll a concentration in its effect on geographical variations in prosome length and body weight of Neocalanus copepods in the North Pacific. This revised version was published online in July 2006 with corrections to the Cover Date.     

Grazing impact of the copepod community in the Oyashio region of the western subarctic Pacific Ocean

The role of copepod grazing on the ecosystem dynamics in the Oyashio region, western subarctic Pacific was investigated during six cruises from June 2001 to June 2002. In situ grazing rates of the copepod community (CGR) were measured by the gut fluorescence method in respect to developmental stages of dominant species. In terms of biomass, more than 80% of the copepod community was dominated by six large calanoid species (Neocalanus cristatus, Neocalanus flemingeri, Neocalanus plumchrus, Eucalanus bungii, Metridia pacifica and Metridia okhotensis) throughout the year. Resulting from the observed pattern of the interzonal migrating copepods, the CGR in the Oyashio region was divided into three phases, i.e. spring (bloom), summer (post-bloom) and autumn-winter phase. During the spring bloom, late copepodites of the interzonal migrating species, N. cristatus, N. flemingeri and E. bungii appeared in the surface layer (0-50 m) to consume the production of the bloom, resulting in a high grazing rate of the copepod community (7.9 mg Chl m⁻² d⁻¹), though its impact on phytoplankton community was low due to the high primary productivity. During the post-bloom period, although the copepod community which was dominated by N. cristatus, N. plumchrus, M. pacifica and newly recruited E. bungii still maintained a high biomass, the CGR was generally lower (1.8-2.6 mg Chl m⁻² d⁻¹ for June and August 2001), probably due to the lower availability of phytoplankton. Nevertheless, the highest CGR was also observed during this period (10.5 mg Chl m⁻² d⁻¹ in June 2002). The high CGR on autotrophic carbon accounted for 69% of the primary production, suggesting that the copepod community in the Oyashio region potentially terminates the phytoplankton bloom. Abundant occurrence of young E. bungii, which is a characteristic phenomenon in the Oyashio region, was largely responsible for the high grazing pressure in June 2002 suggesting that success of reproduction, growth, and survival in E. bungii during the spring bloom is an important factor in controlling phytoplankton abundance during the post-bloom season. During autumn and winter, CGR was the lowest in the year (0.29-0.38 mg Chl. m⁻² d⁻¹) due to the disappearance of the interzonal migrating copepods from the surface layer. Diel migrant M. pacifica was the most important grazer during this period. The annual ingestion of the copepod community is estimated as 37.7 gC m⁻² on autotrophic carbon (converted using C:Chl ratio of 30) or 137.9 gC m⁻² on suspended particles (using C:Chl ratio of in situ value, 58-191), accounting for 13% and 46% of annual primary production, respectively. This study confirms that copepod grazing is an important pathway in carbon flow in the Oyashio region and in particular their role in the phytoplankton dynamics is significant for the termination of the spring bloom.     

Vertical distributions of large ontogenetically migrating copepods in the Oyashio region during their growing season

Neocalanus flemingeri, Neocalanus plumchrus, Neocalanus cristatus and Eucalanus bungii are large and dominant mesozooplankton occurring throughout the subarctic Pacific. They are an important trophic link and transporter of organic matter to the mesopelagic zone. Vertical distributions of these copepods were investigated from March to October 2000 in the Oyashio region of the western subarctic Pacific. Neocalanus plumchrus and N. flemingeri were distributed in the surface layer (0–50 m) and N. cristatus and E. bungii in the subsurface layer (50–100 m). However, when examined in detail, clear seasonal and vertical differences were observed. Neocalanus plumchrus was concentrated in the top 20 m from late April to the end of July, and N. flemingeri showed a little deeper distribution from May to July. Neocalanus cristatus showed a deeper distribution than that of grazing individuals of E. bungii from April to early July, but grazing individuals of E. bungii (C3–C6) showed a deeper distribution than that of N. cristatus from the end of July to October. Early copepodites of E. bungii were distributed much shallower than late copepodite stages and overlapped with copepodites of N. plumchrus and N. flemingeri. These results suggest that the four species of large copepods have established habitat segregation by season, vertical distribution and food resource partitioning in the Oyashio region as well as other regions of the North Pacific.     

Egg production and early development of the subarctic copepods Neocalanus cristatus,N. plumchrus and N. flemingeri

Reproduction and early development of the large subarctic copepods Neocalanus cristatus, N. plumchrus and N. flemingeri were investigated in the laboratory. All three species produced eggs at intervals of 5.0 to 14.7 days, depending on species and temperature. The females of N. flemingeri released the fewest clutches of the three species, but with the largest clutch size. Clutch size of N. cristatus was smallest with longest intervals between clutches. Mean±1 S.E. of total fecundities were 386±116 eggs for N. cristatus, 840±214 eggs for N. plumchrus, and 924±346 eggs for N. flemingeri. The egg laying period of a female at 2°C was estimated to be 91 days for N. cristatus, 60 days for N. plumchrus and 47 days for N. flemingeri. The color and outer characteristics of eggs and nauplii of these species were quite different. C : N ratios of the eggs were 9.3–10.5, which were slightly higher than that of females or CV. Egg hatching times for each species were 4.6–5.7 days at 2°C and decreased with increasing temperature at a Q₁₀ of 2.8–3.0. N. cristatus nauplii developed to copepodid stage I (CI) without feeding, and the developmental time in days from hatching to CI was expressed as a Bělehrádek equation, D_CI=17068×(T+14.7)^−2.05. Reproduction strategies of the three species of Neocalanus are discussed with reference to their life history strategies.     

High feeding rates on large particles by Neocalanus flemingeri and N. plumchrus,and consequences for phytoplankton community structure in the subarctic Pacific Ocean

The open subarctic Pacific Ocean is a high nitrate low chlorophyll (HNLC) system characterized by low concentrations of phytoplankton, a community dominated by small cells, and iron-limited growth of, especially, the larger phytoplankton. In such systems the main energy and material flow is through the microbial web, with large copepods considered primarily to be grazers on the larger microzooplankton occupying the top of this web. Consistent with this is the recognition that much of the nutrition of the dominant copepods in this system, Neocalanus flemingeri, N. plumchrus and N. cristatus, is derived from microzooplankton. Also, these copepods consume only a small fraction of the total phytoplankton production. In this paper, we show that the contribution made by N. flemingeri and N. plumchrus to establishing and maintaining the community structure of this ecosystem should be re-evaluated. Our experiments indicate these grazers have high clearance rates on large particles, including both large phytoplankton and microzooplankton, and this selective removal contributes to establishment and maintenance of the observed foodweb structure in the Gulf of Alaska. These high feeding rates combined with large populations of these two Neocalanus species concentrated in the upper layer of the ocean, result in population-based feeding rates approximately equal to the growth rates of large phytoplankton under iron-limited conditions. We conclude that N. flemingeri and N. plumchrus populations (a) directly prevent the accumulation of large phytoplankton cells by selectively feeding on them at high rates, and (b) indirectly stimulate the accumulation of the smaller phytoplankton by consumption of their major grazers, the microzooplankton.     

Vertical habitat partitioning by large calanoid copepods in the oceanic subarctic Pacific during Spring

The copepods Neocalanus plumchrus, N. flemingeri, N. cristatus, and Eucalanus bungii dominate the net zooplankton throughout the subarctic Pacific Ocean. All four species have an extensive seasonal ontogenetic vertical migration, completing most or all of their feeding and somatic growth in spring and early summer. We used stratified tows with MOCNESS and BIONESS instrumented net systems to resolve their upper ocean vertical distributions in May and June of 1984, 1987 and 1988. In each year the feeding copepodite stages of all four species were concentrated above the permanent halocline (roughly from 0 to 150m). However, the four species showed strong vertical species zonation and segregation within this layer. We consistently found a near-surface pair (N. plumchrus and N. flemingeri) and a subsurface pair (N. cristatus and E. bungii). The boundary between these groups shifts vertically, but was sharply defined and was very often coincident with a weak and transient thermocline marking the base of the layer actively mixed by surface wind and wave energy. Diel vertical migration was very limited during our sampling periods.The data suggest that the vertical distribution patterns of the copepods could be set by responses to the local intensity of turbulent mixing in the watercolumn. N. plumchrus and N. flemingeri occupied a stratum characterized by strong turbulence. N. cristatus and E. bungii occupied a stratum that was a local minimum in turbulence profiles. The depth of the boundary between the species pairs was deeper when winds and surface energy inputs were strong. The vertical partition pattern may also be determined by a difference in feeding strategy between the species pairs. N. plumchrus and N. flemingeri may feed on the enhanced protozoan population of the mixed layer, while N. cristatus and E. bungii feed on particle aggregates settling from above.     

Protozoa in the diets of Neocalanus spp. in the oceanic subarctic Pacific Ocean

Copepod species of the genus Neocalanus dominate the zooplankton biomass of the oceanic subarctic Pacific Ocean. Neocalanus spp. populations in the subarctic Pacific environment are successful: they feed, accumulate lipid, and persist from year to year. Prior experimental observations derived from a variety of methods indicated that, although their functional morphology is such that they clear the small phytoplankton cells characteristic of the oceanic subarctic Pacific environment efficiently, Neocalanus spp. do not consume sufficient phytoplankton to meet even basic metabolic requirements in that environment. Hence, their success in the subarctic Pacific must depend on their ability to obtain nutrition from other sources. As part of the SUPER (SUbarctic Pacific Ecosystem Research) program, experiments were performed to test the hypothesis that N. plumchrus and N. cristatus obtain a significant portion of their nutrition from planktonic Protozoa. The experiments demonstrate that Protozoa alone do not provide sufficient nutrition for N. cristatus to meet its basic metabolic needs. Protozoa constitute the major dietary component of N. plumchrus however, in agreement with the predictions of Frost's (1987) model of the subarctic Pacific ecosystem. At a minimum this diet permits N. plumchrus to meet basic metabolic requirements. Copepod grazing activities appear to be sufficient to control protozoan stocks in the oceanic subarctic Pacific during late spring and early summer when Neocalanus spp. inhabit the upper water column.     

Horizontal transport of the calanoid copepod Neocalanus in the North Pacific: The influences of the current system and the life history

The horizontal transport of calanoid copepod Neocalanus flemingeri, N. plumchrus and N. cristatus in the subarctic North Pacific has been investigated by particle tracking experiments using an ocean circulation model. In our physical numerical model, the current and frontal systems in the subarctic Pacific are reproduced realistically, and fine-resolved (not smoothed) and vertically sheared western boundary currents in the model enable us to assess the differences in horizontal transports among Neocalanus species. In the experiments, seasonal vertical migration and life cycle of Neocalanus species is included, and this attempt is a novel approach in examining the transports of the zooplanktons. The maximal depths of the vertical migrations and the lengths of the surface and mesopelagic dwelling duration are the essential factors in characterizing the trajectory of the horizontal transport. Both small and large forms of N. flemingeri (hereafter NF-SF and NF-LF) whose mesopelagic-inhabited depths are the shallowest among the three Neocalanus species are transported to more distant regions from initial positions compared with the others. The longest transport distance is over 5000 km for NF-SF. On the other hand, the transport distance of the deepest inhabitant N. cristatus (hereafter NC) is the shortest. The differences in the transport distance are due to the current speed of the western boundary currents at the inhabited depth at which they spend most of their lifetime (ca. 3/4 of their lifetime). More than 82% of NF-LF that originated from the Okhotsk Sea completes its life cycle in the Okhotsk Sea, and all the animals out to the Pacific are transported to the south of the subarctic front where their survivals are unexpected due to high temperatures. This is consistent with the observed limited distribution of NF-LF in the Okhotsk Sea and its surrounding waters. Geographical genetic variations of Neocalanus species are reported as very low. This implies frequent genetic exchange all over the Pacific and its adjacent seas. Being consistent with the above, the present study suggests that the trans-Pacific transport of Neocalanus species can be accomplished within a few generations.     

Downward carbon transport by diel vertical migration of the copepods Metridia pacifica and Metridia okhotensis in the Oyashio region of the western subarctic Pacific Ocean

Seasonal change in the downward carbon transport due to respiration and mortality through diel vertical migration (DVM) of the calanoid copepods Metridia pacifica and Metridia okhotensis was estimated in the Oyashio region, western subarctic Pacific during six cruises from June 2001 to June 2002. M. pacifica (C4, C5 and adult females) was an active migratory species throughout the year though its DVM amplitude varied among seasons and stages. The mean distribution depths of adult females during the daytime were positively related with the illumination level in the water column, being shallowest in April and deepest in January. M. okhotensis generally showed less-extensive migrations than M. pacifica. Therefore, together with their lower abundance, this species is considered to be a less-important mechanism of downward transport of carbon except for April when their DVM was more active and descended deeper than M. pacifica, which remained in the upper 150 m even during the daytime. The mean migrating biomass of the two Metridia species was 558 mg C m⁻² d⁻¹ and was high during summer to winter (263–1676 mg C m⁻² d⁻¹) and low during spring (59–63 mg C m⁻² d⁻¹). Total downward flux through DVM fluctuated between 1.0 and 20.0 mg C m⁻² d⁻¹ with an annual mean of 8.0 mg C m⁻² d⁻¹. Contribution of the respiratory flux was greater than the mortality flux and accounted for 64–98% of total migratory flux throughout the year except for January when contribution of both fluxes was equal. Overall the annual carbon transport by DVM of Metridia spp. was estimated as 3.0 g C m⁻² year⁻¹, corresponding to 15% of the annual total POC flux at 150 m at the study site, suggesting that DVM is a significant process for carbon export in the subarctic region as well as that in tropical and subtropical oceanic regions. Since DVM in M. pacifica is more active during the non-bloom season when the gravitational flux of particulate matter is low, this species plays an important role in driving the biological pump in the subarctic Pacific during summer to winter.     

Production of <Emphasis Type="Italic">Acartia steueri</Emphasis> (Copepoda: Calanoida) in Ilkwang Bay,Southeastern Coast of Korea

Production of the marine calanoid copepod Acartia steueri was measured from 2 October 1991 to 8 October 1992 at a station in Ilkwang Bay, on the southeastern coast of Korea. Phytoplankton standing stock ranged over 1.0 to 9.3 mg chl.a m⁻³, and annual primary productivity (by the C-14 method) at three stations was estimated at 200 gC m⁻² yr⁻¹. Acartia steueri (nauplii + copepodids + adults) were present in the plankton throughout the year, with seasonal variation in abundance. Biomass of A. steueri, excluding the NI stage, was 0.01–4.55 mgC m⁻³ (mean: 0.68 mgC m⁻³) with peaks in November, February, May and July-early August, and relatively low biomass in September– January. Instantaneous growth rates of the nauplius stages were higher than the copepodid stages. Annual production of A. steueri was 25.1 mgC m⁻³ yr⁻¹ (or 166 mgC m⁻² yr⁻¹), showing peaks in November, May and July–August with a small peak in February, and low production in December–April and September–October. There were no significant relationships between the daily production rate of A. steueri and temperature or chlorophyll a concentration, indicating that unknown other factors might be related to the variation of the production rate.     

Effects of mesoscale phytoplankton variability on the copepods Neocalanus flemingeri and N. plumchrus in the coastal Gulf of Alaska

The copepods Neocalanus flemingeri and N. plumchrus are major components of the mesozooplankton on the shelf of the Gulf of Alaska, where they feed, grow and develop during April–June, the period encompassing the spring phytoplankton bloom. Satellite imagery indicates high mesoscale variability in phytoplankton concentration during this time. Because copepod ingestion is related to food concentration, we hypothesized that phytoplankton ingestion by N. flemingeri and N. plumchrus would vary in response to mesoscale variability of phytoplankton. We proposed that copepods on the inner shelf, where the phytoplankton bloom is most pronounced, would be larger and have more lipid stores than animals collected from the outer shelf, where phytoplankton concentrations are typically low. Shipboard feeding experiments with both copepods were done in spring of 2001 and 2003 using natural water as food medium. Chlorophyll concentration ranged widely, between 0.32 and 11.44 μg l⁻¹ and ingestion rates varied accordingly, between 6.0 and 627.0 ng chl cop⁻¹ d⁻¹. At chlorophyll concentrations<0.50 μg l⁻¹, ingestion is always low, <40 ng cop⁻¹ d⁻¹. Intermediate ingestion rates were observed at chlorophyll concentrations between 0.5 and 1.5 μg l⁻¹, and maximum rates at chlorophyll concentrations>1.5 μg l⁻¹. Application of these feeding rates to the phytoplankton distribution on the shelf allowed locations and time periods of low, intermediate and high daily feeding to be calculated for 2001 and 2003. A detailed cross-shelf survey of body size and lipid store in these copepods, however, indicated they were indistinguishable regardless of collection site. Although the daily ingestion of phytoplankton by N. flemingeri and N. plumchrus varied widely because of mesoscale variability in phytoplankton, these daily differences did not result in differences in final body size or lipid storage of these copepods. These copepods efficiently dealt with small and mesoscale variations in their food environment such that mesoscale structure in phytoplankton did not affect their final body size.     

Does transported seagrass provide an important trophic link in unvegetated, nearshore areas?

The contribution of detritus from seagrass and other primary producers to faunal production in unvegetated nearshore areas was examined primarily using stable isotopes. Fish, macroinvertebrates, meiofauna and primary producers (seagrasses, macroalgae, seston and benthic microalgae) were sampled from sites in south-western Australia. All samples were analysed for δ¹³C and δ¹⁵N values and fish gut contents were determined. δ¹³C values for seagrasses in the region were high compared to other macrophytes, ranging from 49.9 to −8.2‰ compared to −19.8 to −12.6‰ for macroalgae. The δ¹⁵N values ranged between 4.0 and 7.7‰ for the red, brown and green algae, and between 3.2 and 5.9‰ for seagrasses. Seston and benthic microalgae samples had a mean δ¹³C value of −12.8 and −14.0‰, respectively, and their δ¹⁵N values were comparable to the macroalgae. All invertebrate fauna had mean δ¹³C values considerably lower than seagrasses. However, individual samples harpacticoid copepods and polychaetes had a value as high as −11.7‰. δ¹⁵N values for consumers were higher than those of the primary producers, except for copepods and amphipods. The δ¹³C values for fish had a relatively small range, between −16.6 and −13.1‰, and the δ¹⁵N values of fish were elevated compared to the invertebrates and primary producers, ranging mostly between 10.0 and 12.6‰. Mixing model analysis based on δ¹³C values indicated that seagrass ranked low as a likely carbon source for all invertebrates other than harpacticoid copepods at a single site and some samples of polychaetes. The δ¹³C values for fish were similar to those of a combination of harpacticoid and calanoid copepods, amphipods and polychaetes. The consumption of harpacticoid copepods by some fish species indicates that Amphibolis and Posidonia species in south-western Australia can contribute to the food web of unvegetated nearshore areas as detritus, but brown algae is likely to make a greater contribution. At least for the time of year that was sampled, the flow of detrital seagrass material into the foodweb may be mediated by specific detrivores, in this case harpactacoid copepods, rather than by all detritivores.     

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.     

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.     

fCO2 variability at the CARIACO tropical coastal upwelling time series station

Monthly seawater pH and alkalinity measurements were collected between January 1996 and December 2000 at 10°30′N, 64°40′W as part of the CARIACO (CArbon Retention In A Colored Ocean) oceanographic time series. One key objective of CARIACO is to study temporal variability in Total CO₂ (TCO₂) concentrations and CO₂ fugacity (fCO₂) at this tropical coastal wind-driven upwelling site. Between 1996 and 2000, the difference between atmospheric and surface ocean CO₂ concentrations ranged from about − 64.3 to + 62.3 μatm. Physical and biochemical factors, specifically upwelling, temperature, primary production, and TCO₂ concentrations interacted to control temporal variations in fCO₂. Air–sea CO₂ fluxes were typically depressed (0 to + 10 mmol C m^{− 2} day^{− 1}) in the first few months of the year during upwelling. Fluxes were higher during June–November (+ 10 to 20 mmol C m^{− 2} day^{− 1}). Fluxes were generally independent of the slight changes in salinity normally seen at the station, but low positive flux values were seen in the second half of 1999 during a period of anomalously heavy rains and land-derived runoff. During the 5 years of monthly data examined, only two episodes of negative air–sea CO₂ flux were observed. These occurred during short but intense upwelling events in March 1997 (−10 mmol C m^{− 2} day^{− 1}) and March 1998 (− 50 mmol C m^{− 2} day^{− 1}). Therefore, the Cariaco Basin generally acted as a source of CO₂ to the atmosphere in spite of primary productivity in excess of between 300 and 600 g C m^{− 2} year^{− 1}.     

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.     

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).     

Ammonia Nitrogen at the Water–Sediment Interface in Puck Bay (Baltic Sea)

The magnitude of the exchange flux at the water–sediment interface was determined on the basis of the ammonia concentration gradient at the near-bottom water–interstitial interface and Fick's first law. It was established that in Puck Bay, ammonia almost always passes from the sediment to water. Ammonia flux varied from 5 to 1434 μmol NH₄-N m⁻² day⁻¹. In total,c. 138·2 tonneammonia year⁻¹pass from sediments of Internal Puck Bay to near-bottom water, the equivalent value for External Puck Bay being 686·9 tonne year⁻¹. In total, about 825 tonne ammonia year⁻¹passes from the sediment to near-bottom water of Puck Bay. In interstitial waters, ammonia occurred in concentrations varying over a wide range (3–1084 μmol NH₄-N dm⁻³).The basic factors affecting the magnitude of ammonia concentration in interstitial waters included: oxidation of organic matter, type of sediment, and inflow of fresh underground waters to the region examined.This paper involves preliminary studies only and constitutes a continuation of the studies on ionic macrocomponents and phosphorus in interstitial waters of Puck Bay undertaken previously.     

Uptake of atmospheric carbon dioxide in Arctic shelf seas: evaluation of the relative importance of processes that influence pCO2 in water transported over the Bering–Chukchi Sea shelf

The uptake of atmospheric carbon dioxide in the water transported over the Bering–Chukchi shelves has been assessed from the change in carbon-related chemical constituents. The calculated uptake of atmospheric CO₂ from the time that the water enters the Bering Sea shelf until it reaches the northern Chukchi Sea shelf slope (1 year) was estimated to be 86±22 g C m⁻² in the upper 100 m. Combining the average uptake per m³ with a volume flow of 0.83×10⁶ m³ s⁻¹ through the Bering Strait yields a flux of 22×10¹² g C year⁻¹. We have also estimated the relative contribution from cooling, biology, freshening, CaCO₃ dissolution, and denitrification for the modification of the seawater pCO₂ over the shelf. The latter three had negligible impact on pCO₂ compared to biology and cooling. Biology was found to be almost twice as important as cooling for lowering the pCO₂ in the water on the Bering–Chukchi shelves. Those results were compared with earlier surveys made in the Barents Sea, where the uptake of atmospheric CO₂ was about half that estimated in the Bering–Chukchi Seas. Cooling and biology were of nearly equal significance in the Barents Sea in driving the flux of CO₂ into the ocean. The differences between the two regions are discussed. The loss of inorganic carbon due to primary production was estimated from the change in phosphate concentration in the water column. A larger loss of nitrate relative to phosphate compared to the classical ΔN/ΔP ratio of 16 was found. This excess loss was about 30% of the initial nitrate concentration and could possibly be explained by denitrification in the sediment of the Bering and Chukchi Seas.