Distribution,size frequencies and maturity stages of krill,Euphausia superba,in relation to sea-ice in the Northern Weddell Sea

Summary The spring (November) and early summer (December) distribution pattern and demography of Euphausia superba in the seasonally ice covered areas of the north-western Weddell Sea are described. Sampling with a Rectangular Midwater Trawl (RMT) and during SCUBA diving were undertaken in the pack-ice during the EPOS expedition 1988/1989 of RV Polarstern. These data were compared with former cruises during late winter (October) and summer (February). The biomass values of krill (1–27 g/m²) in the water column within the ice covered areas were of the same order of magnitude as earlier acoustic estimates in the ice-edge region of the same area. No significant correlation between percentage ice cover and krill abundance in the water column was found. Cluster analyses of size frequency distributions and maturity stage composition revealed the demography of the animals sampled. Samples obtained by SCUBA and RMT are compatible and results show a geographic and demographic separation of the krill. It is proposed that krill in the northern zone are largely emigrants from the Bellingshausen Sea, while the krill in the southern zone are of eastern Weddell Sea origin. Further studies concentrating on the under-ice distribution of krill are recommended.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation. Publication no 62 of the Alfred-Wegener-Institut für Polar- und Meeresforschung     

Top predators as indicators for ecosystem events in the confluence zone and marginal ice zone of the Weddell and Scotia Seas,Antarctica, November 1988 to January 1989 (EPOS Leg 2)

Summary The pelagic summer distribution of Antarctic seabirds, seals and whales was studied in the marginal ice zone of the northwestern Weddell Sea from November 1988 to January 1989. In order to relate top predators to other components of the ecosystem studied simultaneously, their distribution is mainly described in terms of energy flow. Bird, seal, and probably also whale requirements were highest in ice-covered areas. There was no evidence of higher numbers of top predators along the ice edge: densities generally increased further into the ice. In the pack ice, combined energy requirements of top predators often amounted to about 200.000 kJ/day/km², or about 45 kg fresh food, indicating high abundance and availability of prey under the ice. There was a lack of conformity between top predator abundance on the ice and abundance of other life in the water column below. In open water, bird requirements were generally less than 25.000 kJ/day/km², seals were virtually absent and whales were distributed unevenly. Tubenosed birds concentrated along the outer ice edge in early summer but they moved north to open water during December, leaving the area of maximum phytoplankton biomass associated with the retreating ice edge. This pattern matched northward movements of krill swarms that may be related to changes in quality rather than quantity of phytoplankton stocks.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Elemental composition of suspended matter in the Scotia-Weddell Confluence area during spring and summer 1988 (EPOS leg 2)

Summary During austral spring and summer 1988 the upper 500 m of water column in the Scotia-Weddell Confluence was sampled for the elemental composition of total suspended matter. For particulate organic carbon surface water concentrations ranged between 2.5 and 15 mol/l, with an estimated 19 to 47% of this pool being detrital carbon. In late November, the highest surface water particulate organic carbon concentrations (15 mol/l) occurred in the Confluence area where they coincided with a maximum in particulate Si (1.7 mol/l). Later in the season particulate Si in the Confluence area decreased to 0.3 mol/l. In the Scotia Sea on the contrary, surface water particulate Si increased with time and reached 3 mol/l in late December. For particulate Ca and Sr in surface water, strong gradients are observed across the Scotia Front (e.g. Ca: from 230 to 10 nmol/l; Sr: from 1.0 to 0.1 nmol/l), with highest concentrations in the Scotia Sea. In general, these distributions are confirmed by the observations on plankton species composition, done by other participants. In the Scotia Sea heavily calcified coccolithophorids and diatoms occurred throughout the season, while in the Confluence area heavily calcified coccolithophorids were absent and a switch-over from diatom to naked flagellate dominance was observed following a krill event. In the surface waters, the lithogenic Si fraction represents on average only 4% of the total particulate Si content. However, this fraction reaches 60% below 100 m depth in the Confluence area, due mainly to the presence of a sub-surface maximum in the aluminosilicate load (particulate Al content up to 30 pmol/l), probably reflecting advection of resuspended shelf sediments. Subsurface Ba/barite concentrations are highest in the Scotia Sea (280 pmol/l) and decrease through the Scotia Front to reach values of 100 pmol/l and less in the Confluence area, the marginal ice zone and the closed pack ice zone.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Influence of physical and biological processes on the concentration of O2 and CO2 in the ice-covered Weddell Sea in the Spring of 1988

Summary In October and November 1988, measurements of oxygen and total dissolved inorganic carbon (TCO₂) concentrations were made in the northwestern Weddell Sea to the south and north of the marginal ice edge, in order to estimate the relative importance, regarding their variations, of both biological (photosynthesis and respiration) and physical (transport of O₂ and CO₂ by turbulent movements and by intrusion from the atmosphere) processes. In the ice-covered region, both respiration and upwelling determined the O₂ and TCO₂ variations, whilst in the open water just north of the marginal ice edge, photosynthetic activity was the most important factor controlling O₂ and TCO₂ levels. These findings underline the importance of the activity of the pelagic ecosystem in determining the concentration of O₂ and CO₂ not only in the ice-free but also in the ice-covered Antarctic Ocean.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Nitrogen uptake in the Weddell Sea during late winter and spring

Summary Uptake rates of ammonium, nitrate and urea were measured during the EPOS leg 1 cruise to the Weddell Sea in October–November 1988 using the isotope ¹⁵N. Nitrate was the most important nitrogen source both for ice algae (f-ratio 0.88) and for phytoplankton in the water column (f-ratio 0.85). Indications of a gradual decrease in % new production with time were found in the outer marginal ice zone. Nitrogen uptake rates in ice algae from the sub-ice assemblage were light-limited at in situ irradiances. Significant regeneration of ammonium was found in ice algal samples only.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Acoustic observations on krill spring-summer migration and patchiness in the northern Weddell Sea

Summary Continuous acoustic observations with a 30 kHz and a 150 kHz echo-sounder were made from November 1988 to January 1989 on one repeatedly sampled transect, running along 49°W from the open waters of the Scotia Sea into the Weddell Sea pack-ice. Swarm signals occurring on the echorecords were mainly found in the upper 100 m of the watercolumn, in size varying vertically from 1 to 70 m, and horizontally from less than 5 to over 3,000 m. Catches with a RMT 1+8 indicated that the observed swarm signals were most probably caused by krill, Euphausia superba. From late November to early January the swarms migrated northwards away from the ice-edge and towards greater depths, while simultaneously growing in size. The average number of swarms observed per 10 nautical miles along the transects remained fairly constant throughout the cruise, but the average swarmsize and total aggregation size increased during the period studied. The echo-data give evidence of the spring-summer migration of major parts of the krill stock from under the ice-cover towards open waters.Data presented here were collected during the European Polarstern Study (EPOS), funded by the European Science Foundation     

Organic carbon in the upper layer and its sedimentation during the ice-retreat period in the Scotia-Weddell Sea, 1988

Summary The maximum of POC in the surface layer (200–400 g·l^–1) followed the retreating ice from end November to end December. In the upper 150 m DOC amounted to 10–20 times the POC content. Free floating sediment traps at 150 m showed a daily sedimentation of 0.6–11.6% of the standing stock of POC, 0.1–2.6% of the chlorophyll-a and 5–190% of the gross primary production. Maximum sedimentation occurred during grazing of a krill swarm, indicating the important role of krill swarms in the downward flux. Also at most other stations krill faecal strings formed a large part of the downward flux.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Short-term variations in particulate matter sedimentation off Kapp Norvegia,Weddell Sea,Antarctica: relation to water mass advection,ice cover,plankton biomass and feeding activity

Summary A multi-cup sediment trap was deployed at 250m in the shelf area off Kapp Norvegia, Weddell Sea (630 m water depth) to determine the relative importance of water mass advection, sea ice movement, phytoplankton biomass and plankton feeding. Short-term fluctuations in sedimentation were determined using a sampling frequency of 2.7 days over 54 days during January and February 1988. Three periods of enhanced sedimentation were associated with water mass exchange, settling of diatoms following break-up of ice cover and release of fecal matter by krill feeding on particulate matter derived from phytoplankton and ice algae. An initial sedimentation pulse (28 Jan) was mainly due to sinking pelagic diatoms and krill fecal strings containing algae released from sea ice passing over the trap position. The ¹³C-composition of the sedimented organic carbon was about-24. The isotope ratio decreased sharply by about 5.5 at the end of the first pulse indicating the source of sinking matter becoming pelagic diatoms of the retreating ice-edge. At this time the diatom Corethron criophilum contributed a very high proportion of the organic flux causing an increase of the opal/C_org ratios. The second pulse (6 Feb) was due to empty diatom frustules, minipellets and small planktonic aggregates. Much of the organic carbon was transported by round fecal pellets. During the third pulse (14 Feb), round fecal pellets transported even more; the percentage of C. criophilum to the diatom organic carbon flux was more than 80% (>2mg C m^–2 day^–1).Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Effects of Antarctic sea ice biota on seeding as studied in aquarium experiments

Summary The potential seeding impact of sea ice microbial communities was studied during late austral winter early spring 1988 in the Weddell Sea, Antarctica. Experiments were performed in seawater aquariums with natural seawater and seawater enriched with crushed ice. Algal, protozoan and bacterial cell numbers were followed, as well as nutrients and DOC levels. The results showed a potential seeding effect of sea ice communities to the water column. However, the type of ice communities differed greatly from each other and the effect of such seeding will be patchy. In our experiments seeding of seawater by ice rich in algae, flagellates and/or particulate organic carbon lead to the development of communities dominated either by diatoms or bacteria.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Abundance of cryptophyceae and chlorophyll b-containing organisms in the Weddell-Scotia Confluence area in the spring of 1988

Summary During a cruise in the Weddell-Scotia Confluence area (EPOS Leg 2: November–January 1988/1989) nanophytoplankton composition was determined by employing taxon-specific pigment measurements with HPLC. The biomass of the most important components was estimated by using specific pigment ratios measured in cultures of two cryptomonads and a prasinophyte. Highest cryptophyte biomass was found along the retreating ice-edge; the contribution of cryptophytes to total phytoplankton crop increased with time, reaching monospecific bloom conditions at the end of the cruise. Chlorophyll b-containing organisms and Prymnesiophyceae were present everywhere and dominated in the ice-covered part of the survey area. Cryptophyte-specific pigment measurements were in reasonable agreement with cryptophyte cell numbers. Prasinophyte cell counts, however, did not match with measured chlorophyll b concentrations. The quantitative importance of the nanophytoplankton groups reported here underlines the diversity of the plankton in the Southern Ocean's marginal ice zone system which may have implications for food chain dynamics.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Community structure of the epipelagic zooplankton community under the sea-ice of the northern Weddell Sea

Summary The present paper describes the composition, abundance, biomass and diversity of the meso- and macrozooplankton in the epipelagic zone of the open water and under the ice of the northern Weddell Sea. Samples were collected in October/November 1988 with a multiple RMT1+8 net during the European Polarstern Study (EPOS). Multivariate analysis resulted in two distinct site clusters, a northern one mainly located in the open water/marginal ice zone and a southern one extending from the marginal ice zone into the consolidated pack-ice. Clusters were, however, faunistically coherent with a high degree in positive covariation of species. There was no basis for the separation into communities, but differences occurred on the population level in numerical abundances, biomass (wet weight) and in a shift in species dominance. Different ice zones and vertical layers were tested among each other with regard to their relative species abundance. Significant differences were found between the upper 60 m layer of the open sea, the upper 60 m layer of the closed pack-ice and the so called transitional zone. Species richness and diversity was lowest directly under the closed pack-ice. Abundance and biomass was highest in the surface layer of the open water, while both variablès decreased dramatically under the ice. Copepods dominated numerically in open water, while salps dominated in biomass. Euphausia superba and Thysanoessa macrura were the dominant species in the upper water column of the closed pack-ice zone. Krill was the only species with increasing abundance in the sub-ice area and a dominance in biomass of more than 91% demonstrated its unique importance for the sub-ice habitat.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Aggregation of algae released from melting sea ice: implications for seeding and sedimentation

Summary Factors influencing the fate of ice algae released from melting sea ice were studied during a R V Polarstern cruise (EPOS Leg 2) to the northwestern Weddell Sea. The large-scale phytoplankton distribution patterns across the receding ice edge and small-scale profiling of the water column adjacent to melting ice floes indicated marked patchiness on both scales. The contribution of typical ice algae to the phytoplankton was not significant. In experiments simulating the conditions during sea ice melting, ice algae revealed a strong propensity to form aggregates. Differences in the aggregation potential were found for algal assemblages collected from the ice interior and the infiltration layer. Although all algal species collected from the ice were also found in aggregates, the species composition of dispersed and aggregated algae differed significantly. Aggregates were of a characteristic structure consisting of monospecific microaggregates which are likely to have formed in the minute brine pockets and channels within the ice. Sinking rates of aggregates were three orders of magnitude higher than those of dispersed ice algae. These observations, combined with the negligible seeding effect of ice algae found during this study, suggest that ice algae released from the melting sea ice are subject to rapid sedimentation. High grazing pressure at the ice edge of the investigation area is another factor eliminating ice algae released during melting.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

The distribution and abundance of krill faecal material and oval pellets in the Scotia and Weddell Seas (Antarctica) and their role in particle flux

Summary The abundance and depth distribution of zooplankton faeces in spring to early summer were investigated along meridional transects (47°W and 49°W) that extended from the Scotia Sea (57°S) across the Weddell-Scotia Confluence and into the Weddell Gyre (62°S). The sea ice edge retreated from 59°30S to 61°S during the study. Faeces were sampled with nets, Niskin bottles and sediment traps and subsequently analysed by light and electron (SEM) microscopy. Krill faecal strings and oval faecal pellets of unknown origin were by far the most important zooplankton faeces and highest concentrations were always found in the Confluence often close to the ice border. Krill faeces were usually more abundant in the uppermost layer (0–50m) where they contributed an average of 130 g dry weight m^–3. There was an exponential decrease with depth, with a minimum of 0.6 g dry weight m^–3 in the 500–1000 m stratum. Oval pellets were more evenly distributed in the upper 1000 m of the water column, with an average of 9 g dry weight m ^–3, although there was a small peak (20 g dry weight m^–3) in the subsurface layer (50–150 m depth). Consecutive collections (day-night) of krill faeces using drifting sediment traps showed that only the larger strings sank from 50 to 150 m depth. Peritrophic membranes appeared to deteriorate during sinking. Diatoms (in particular Nitzschia and Thalassiosira spp.) contributed by far the bulk of material in krill and oval faeces. In samples collected near or under the pack ice, remains of crustaceans in both krill- and oval faeces were also found.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Size and stage composition of age class 0 Antarctic krill (<Emphasis Type="Italic">Euphausia superba</Emphasis>) in the ice–water interface layer during winter/early spring

The condition and survival of Antarctic krill (Euphausia superba) strongly depends on sea ice conditions during winter. How krill utilize sea ice depends on several factors such as region and developmental stage. A comprehensive understanding of sea ice habitat use by krill, however, remains largely unknown. The aim of this study was to improve the understanding of the krill’s interaction with the sea ice habitat during winter/early spring by conducting large-scale sampling of the ice–water interface (0–2 m) and comparing the size and developmental stage composition of krill with the pelagic population (0–500 m). Results show that the population in the northern Weddell Sea consisted mainly of krill that were <1 year old (age class 0; AC0), and that it was comprised of multiple cohorts. Size per developmental stage differed spatially, indicating that the krill likely were advected from various origins. The size distribution of krill differed between the two depth strata sampled. Larval stages with a relatively small size (mean 7–8 mm) dominated the upper two metre layer of the water column, while larger larvae and AC0 juveniles (mean 14–15 mm) were proportionally more abundant in the 0- to 500-m stratum. Our results show that, as krill mature, their vertical distribution and utilization of the sea ice appear to change gradually. This could be the result of changes in physiology and/or behaviour, as, e.g., the krill’s energy demand and swimming capacity increase with size and age. The degree of sea ice association will have an effect on large-scale spatial distribution patterns of AC0 krill and on predictions of the consequences of sea ice decline on their survival over winter.     

Long-term changes in Krill biomass and distribution in the Barents Sea: are the changes mainly related to capelin stock size and temperature conditions?

Krill plays a significant role in the Barents Sea ecosystem, providing energy transport between different trophic levels. The current paper presents the results of a long-term study (1980–2009) based on pelagic trawl catches from August to September. Our investigations show that the krill species were distributed widely in the Barents Sea and that the largest krill concentrations were restricted to the west-central and eastern parts of the Barents Sea. The current paper presents the relative biomass indices, and the estimates must be interpreted as minimum biomass. The mean annual krill biomass was estimated to be 22 million tonnes in wet weight, with the highest values being as much as 48 million tonnes. Capelin is the largest pelagic stock, and in some years, their biomass can amount to 4–7 million tonnes, which can impose high predation pressure on krill. When their biomass is high, capelin may consume close to 26 million tonnes annually. The predation from pelagic (herring and blue whiting) and bottom (cod and haddock) fish species was much lower, being 9 and 1 million tonnes, respectively. A negative relationship between krill biomass and capelin stock size above 74°N was observed during the study period. However, during the last decade, the krill biomass has increased despite heavy predation from capelin in some years. A positive significant linear relationship between the mean annual Kola temperature and the krill biomass seems to indicate that the recent warming conditions have favourable impacts on the krill populations in the Barents Sea.     

Spring distribution and ecological role of seabirds and marine mammals in the Weddell Sea,Antarctica

Summary The early spring distribution of seabirds, pinnipeds and cetaceans was quantitatively determined in the northern Weddell Sea during the EPOS 1 cruise of the icebreaking RV Polarstern. Two hundred and ninety-one half hour counts were made in the Antarctic region between October 18 and November 16, 1988 (+94 counts in sub-antarctic and sub-tropical water). The bird populations were dominated by the Adélie Penguin Pygoscelis adeliae, which represented 90% of the birds counted in the closed pack ice (CPI), with a mean density of 31 penguins per km². Crabeater seal Lobodon carcinophagus, the most common pinniped, had a mean density of 1.2/km² in the CPI with local concentrations reaching 14/km². Expressed as biomass, mean values of about 90 kg/km² were found in the whole Antarctic zone for seabirds, 50 for seals, and 35 for baleen cetaceans. Densities were higher in the CPI: 140, 180 and 115 kg/km² respectively, were found for birds, pinnipeds and baleen whales, and lowest in open water (8 kg/km² for the birds, 0.2 for the pinnipeds and no whales). A minimum value of food intake by seabirds and marine mammals was estimated to be 2.7 mg C/m²/day for the Antarctic zone (4.1 in the CPI). Taking into account that pinnipeds density and cetacean food uptake are underestimated, and that Antarctic seabirds consume much more fish than generally suspected, a minimum krill production of 16 mg C/m²/day is proposed for the Antarctic zone: 22.5 in the CPI, 3 in open water, and intermediate values in the marginal ice zones.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Biological activity in the antarctic zooplankton community

Summary Zooplankton biomass and respiratory ETS activity were studied along a transect at 49°W, running from open Scotia Sea water (57°S) into the Weddell Sea pack ice (62°S) in November–December 1988. Zooplankton biomass and respiratory activity were relatively high in the ice edge region and in the frontal zone separating the Scotia Sea from waters in the south; low biomasses and activities were encountered under the pack ice. The higher activities in the frontal zone were mainly attributed to locally higher water temperatures, while in the ice edge region they were probably related to changes of the Zooplankton population into more active developmental stages. This development in life stages is possibly a response to ice edge generated phytoplankton blooms. A comparison of the most abundant Zooplankton taxa indicated that amphipods and euphausiids had relatively the highest weight specific ETS activities. The developmental stages of Euphausia superba, from juvenile to sexually mature adults, showed a large variability in ETS activity; only the most inactive stages were found under the pack ice. The difference in respiration between the most active and inactive stages was at least a factor of 6 and is of importance to the overwintering of the species in the under ice habitat.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

The association of Antarctic krill Euphausia superba with the under-ice habitat

The association of Antarctic krill Euphausia superba with the under-ice habitat was investigated in the Lazarev Sea (Southern Ocean) during austral summer, autumn and winter. Data were obtained using novel Surface and Under Ice Trawls (SUIT), which sampled the 0-2 m surface layer both under sea ice and in open water. Average surface layer densities ranged between 0.8 individuals m(-2) in summer and autumn, and 2.7 individuals m(-2) in winter. In summer, under-ice densities of Antarctic krill were significantly higher than in open waters. In autumn, the opposite pattern was observed. Under winter sea ice, densities were often low, but repeatedly far exceeded summer and autumn maxima. Statistical models showed that during summer high densities of Antarctic krill in the 0-2 m layer were associated with high ice coverage and shallow mixed layer depths, among other factors. In autumn and winter, density was related to hydrographical parameters. Average under-ice densities from the 0-2 m layer were higher than corresponding values from the 0-200 m layer collected with Rectangular Midwater Trawls (RMT) in summer. In winter, under-ice densities far surpassed maximum 0-200 m densities on several occasions. This indicates that the importance of the ice-water interface layer may be under-estimated by the pelagic nets and sonars commonly used to estimate the population size of Antarctic krill for management purposes, due to their limited ability to sample this habitat. Our results provide evidence for an almost year-round association of Antarctic krill with the under-ice habitat, hundreds of kilometres into the ice-covered area of the Lazarev Sea. Local concentrations of postlarval Antarctic krill under winter sea ice suggest that sea ice biota are important for their winter survival. These findings emphasise the susceptibility of an ecological key species to changing sea ice habitats, suggesting potential ramifications on Antarctic ecosystems induced by climate change.     

Late winter microbial communities in the western Weddell Sea (Antarctica)

Microbial communities in the water column and sea ice were studied during the EPOS-cruise on R/V Polarstern in the western Weddell Sea in late winter (October–November 1988). Samples were taken from four transects from heavy pack-ice to open water. The results indicated the important role of protozoans especially in the ice-edge area. Heterotrophic nanofiagellates, dinoflagellates, ciliates and sarcodines showed significant positive correlations with chlorophyll a. Autotrophic picoplankton and autotrophic flagellates, which were probably motile zooids of Phaeocystis pouchetii (up to 3×10⁶ cells 1^–1), were most abundant in the areas of low or medium chlorophyll a concentration. Sea ice contained high numbers of heterotrophic organisms, and the distribution of the different groups showed distinct vertical zonation. At two sites, the microbial assembly beneath the ice was clearly influenced by communities from the melting ice.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

Hydrography and biochemical indicators of microplankton biomass in the Bransfield Strait (Antarctica) during January 1994

 The relationships between hydrography and spatial distribution of several biochemical indicators of microplankton biomass (chlorophyll, protein and ATP) were studied in an area covering the eastern part of the Bransfield Strait and the northern part of the Weddell Sea, during Antarctic summer (January 1994). Four hydrographic zones were identified: (a) the northern part of the Bransfield Strait, covered by waters of Bellings- hausen Sea origin; (b) a Weddell Sea water mass that affected most of the study area; (c) the Weddell-Scotia Confluence waters, observed north of Elephant Island; and (d) waters influenced by ice melting, found towards the southeastern part of the sampled area. The highest values of biomass indicators (chlorophyll a, ATP and protein) were found in the zones affected by ice-melting processes and in waters from the Bellingshausen Sea. The lowest values of all biochemical parameters were found in the Weddell Sea and in the Weddell-Scotia Confluence waters. A high variability in the hydrographic structure and the distribution of biochemical indicators was observed. The degree of stabilization of the water column, the depth of the upper mixed layer and the grazing pressure of herbivorous zooplankton played a major role in the development, accumulation and spatial variability of microplankton biomass. Received: 15 August 1995/Accepted: 18 February 1996