Phytoplankton biomass and primary production in the marginal ice zone of the northwestern Weddell Sea during austral summer

During the austral summer of 1995, distributions of phytoplankton biomass (as chlorophyll a), primary production, and nutrient concentrations along two north-south transects in the marginal ice zone of the northwestern Weddell Sea were examined as part of the 8th Korean Antarctic Research Program. An extensive phytoplankton bloom, ranging from 1.6 to 11.2 mg m⁻³ in surface chlorophyll a concentration, was encountered along the eastern transect and extended ca. 180 km north of the ice edge. The spatial extent of the bloom was closely related to the density field induced by the input of meltwater from the retreating sea ice. However, the extent (ca. 200 km) of the phytoplankton bloom along the western transect exceeded the meltwater-influenced zone (ca. 18 km). The extensive bloom along the western transect was more closely related to local hydrography than to the proximity of the ice edge and the resulting meltwater-induced stability of the upper water column. In addition, the marginal ice zone on the western transect was characterized by a deep, high phytoplankton biomass (up to 8 mg Chl a m⁻³) extending to 100-m depth, and the decreased nutrient concentration, which was probably caused by passive sinking from the upper euphotic zone and in situ growth. Despite the low bloom intensity relative to the marginal ice zone in both of the transects, mean primary productivity (2.6 g C m⁻² day⁻¹) in shelf waters corresponding to the northern side of the western transect was as high as in the marginal ice zone (2.1 g C m⁻² day⁻¹), and was 4.8 times greater than that in open waters, suggesting that shelf waters are as highly productive as the marginal ice zone. A comparison between the historical productivity data and our data also shows that the most productive regions in the Southern Ocean are shelf waters and the marginal ice zone, with emerging evidence of frontal regions as another major productive site. Accepted: 27 September 1998     

Phytoplankton biomass,P-I relationships and primary production in the Weddell Sea,Antarctica, during the austral autumn

An investigation into the changing phytoplankton biomass and total water column production during autumn sea ice formation in the eastern Weddell Sea, Antarctica showed reduced biomass concentrations and extremely low daily primary production. Mean chlorophyll-a concentration for the entire study period was extremely low, 0.15±0.01 mg.m⁻³ with a maximum of 0.35 mg.m⁻³ found along the first transect to the east of the grid. Areas of low biomass were identified as those either associated with heavy grazing or with deep mixing and corresponding low light levels. In most cases phytoplankton in the <20-μm size classes dominated. Integrated biomass to 100 m ranged from 7.1 to 28.0 mg.m⁻² and correlated positively with surface chlorophyll-a concentrations. Mean P^Bmax (photosynthetic capacity) and α^B (initial slope of the photosynthesis-irradiance curve) were 1.25±0.19 mgC. mgChla ⁻¹.h⁻¹ and 0.042±0.009 mgC.mgChla ⁻¹.h⁻¹.(μmol.m⁻².s⁻¹)⁻¹ respectively. The mean index of photoadaptation,I _k, was 32.2±4.0 μmol.m⁻².s⁻¹ and photoinhibition was found in all cases. Primary production was integrated to the critical depth (Z _cr) at each production station and ranged from 15.6 to 41.5 mgC.m⁻².d⁻¹. It appears that, other than grazing intensity, the relationship between the critical depth and the mixing depth (Z _mix) is an important factor as, ultimately, light availability due both to the late season and growing sea ice cover severely limits production during the austral autumn.     

Build-up and decline of summer phytoplankton biomass in the eastern Weddell Sea,Antarctica

The seasonal development and decline of phytoplankton was investigated in the eastern Weddell Sea during summer and fall 1991. During the first half of the study (15 Jan–13 Feb) in an area off Vestkapp, favourable irradiance/mixing regimes initiated net phytoplankton growth in ice-free waters on the shelf and in stretches of open water over the partially ice-covered deep ocean. Chi a concentrations in the upper water column were moderate (0.2–0.8 g l^–1), but significantly above winter values. Later in the season (16 Feb–11 March), a phytoplankton bloom with surface Chl a concentrations ranging from 1.6–2.3 g l^–1 was encountered in an area further to the east. We suggest that the upper water column must have been stratified in this region for time scales of weeks to faciliate bloom development. Bacterial biomass and productivity generally paralleled the seasonal development of the phytoplankton. Nitrate concentrations in the upper mixed layer were substantially lower than would be expected from the existing phytoplankton standing stock, suggesting that heterotrophic consumption of organic matter by bacteria and zooplankton removed a large fraction of the primary production. The shallow seasonal pycnocline was eventually eroded by the passage of a storm, resulting in a homogeneous distribution of phytoplankton biomass over the entire water column, followed by sedimentation and deposition of phytodetritus on the sea floor. After the storm induced destratification, bacterial productivity was particularly high, amounting to more than half of the primary production (range: 10%–120%) in the upper water column. Subsequently, phytoplankton biomass in the upper water column decreased to values <1 g Chl a l^–1. The combination of low incident irradiances and incessant deep mixing prevented the phytoplankton biomass to increase again. During the last week of the investigation, extensive new-ice formation was observed. A major fraction of the residual surface plankton was incorporated into new sea ice, thus terminating the pelagic growth season of the phytoplankton in the eastern Weddell Sea.     

Diatom composition and biomass variability in nearshore waters of Maxwell Bay, Antarctica, during the 1992/1993 austral summer

 Diatom composition and biomass were investigated in the nearshore water (<30 m in depth) of Maxwell Bay, Antarctica during the 1992/1993 austral summer. Epiphytic or epilithic diatoms such as Fragilaria striatula, Achnanthes brevipes var. angustata and Licmophora spp. dominated the water column microalgal populations. Within the bay, diatom biomass in surface water was several times higher at the nearshore (2.4–14 μg C l^-1) than at the offshore stations (>100 m) (1.2–3.2 μg C l^-1) with a dramatic decrease towards the bay mouth. Benthic forms accounted for >90% of diatom carbon in all nearshore stations, while in the offshore stations planktonic forms such as Thalassiosira antarctica predominated (50–>90%). Microscopic examination revealed that many of these diatoms have become detached from a variety of macroalgae growing in the intertidal and shallow subtidal bottoms. Epiphytic diatoms persistently dominated during a 19-day period in the water column at a fixed nearshore station, and the biomass of these diatoms fluctuated from 0.86 to 53 μg C l^-1. A positive correlation between diatom biomass and wind speed strongly suggests that wind-driven resuspension of benthic forms is the major mechanism increasing diatom biomass in the water column. Received: 28 April 1995/Accepted: 1 April 1996     

Biology of Salpa thompsoni in waters adjacent to the Ross Sea, Southern Ocean, during austral summer 2008

Depth-stratified vertical sampling was carried out during the New Zealand International Polar Year cruise to the Ross Sea on board the RV Tangaroa in February–March 2008. The distribution (horizontal and vertical), density and population biology of Salpa thompsoni were investigated. Salps were found at two of the four major sampling locations, e.g. near the continental slope of the Ross Sea and in the vicinity of seamounts to the north of the Ross Sea. Both abundance and biomass of S. thompsoni were highest near the seamounts in the Antarctic Circumpolar Current reaching ~2,500 ind 1,000 m⁻³ and 8.2 g dry wt 1,000 m⁻³ in the water column sampled. The data showed that S. thompsoni populations were able to utilize horizontal and vertical discontinuities in water column structure, in particular the warm Circumpolar Deep Water (CDW), to persist in the high Antarctic. Although salps appeared to continue migrating to the surface colder layers to feed, both aggregate chain and young embryo release seem to be restricted to the CDW. This study for the first time has provided evidence that low Antarctic salp species may successfully reproduce in the hostile high Antarctic realm.     

Microphytoplankton assemblages in the waters surrounding South Georgia, Antarctica during austral summer 1994

The composition and distribution of microphytoplankton assemblages in the vicinity of South Georgia were examined during a krill survey conducted during voyage 119 the RV Africana in austral summer (January/February) 1994. Microphytoplankton distribution was studied at 83 stations along a zig-zag transect in the waters surrounding South Georgia. Oceanographic and zooplankton data suggest that conditions were anomalous during the investigation, in that sub-Antarctic surface waters had probably been advected southwards and were present around the island. The two most widely distributed species were Corethron criophilum and Eucampia antarctica, which were recorded at all stations sampled. Also well-represented amongst the microphytoplankton assemblages were temperate, neritic species such as Chaetoceros atlanticus, Proboscia alata and Odontella weissflogii. This can probably be related to the intrusion of sub-Antarctic surface waters unusually far south. Cluster and ordination analysis identified three distinct groupings of stations in the waters surrounding the islands. These were found to the east, west and north of the islands. The diatom groupings identified during this investigation broadly correspond to three water masses previously described in the vicinity of South Georgia. Despite the anomalous hydrographical conditions that existed during the investigation, the differences in phytoplankton were negligible when compared to normal years. Small mesoscale features are, however, important in imposing biogeographic patterns on microphytoplankton. Received: 21 February 1996 / Accepted: 3 November 1996     

Microzooplankton grazing in the surface water of the Southern Ocean during an austral summer

Microzooplankton grazing was investigated in surface waters of the Indo-Pacific and Atlantic sectors of the Southern Ocean by the dilution method. Phytoplankton growth varied mainly between 0.1 and 0.4 day⁻¹, and microzooplankton grazing between 0.0 and 0.3 day⁻¹. Great fluctuations in phytoplankton growth rate were observed at one station within 3 weeks and between closely spaced stations. Microzooplankton grazing rates were similar to phytoplankton growth rate despite the variation of phytoplankton growth rates, although in some cases, phytoplankton growth overwhelmed microzooplankton grazing. These observations suggest that microzooplankton are the main consumers of primary producers, and that steady state between phytoplankton growth and microzooplankton grazing is usually established in the Southern Ocean in austral summer. Received: 5 November 1996 / Accepted: 4 March 1997     

Physiological characteristics of the antarctic copepod Calanoides acutus during late summer in the Weddell Sea

Ingestion, respiration and excretion rates as well as lipid and protein body content of the dominant Antarctic copepod Calanoides acutus (CIV to adult females) were studied during the period covering the end of phytoplankton bloom (February) to the beginning of transition to overwintering (March-April). Daily rations measured with gut fluorescence varied from 2.2 to 2.7% in surface C. acutus. Weight-specific respiration and excretion rates in deep C. acutus decreased by a factor of 11 and 3.5–3.8 compared to their surface counterparts. High lipid (up to 455 µg ind^–1) and protein (198 µg ind^–l) content was observed in surface C. acutus CV in February; a month later the animals with similar lipid and protein content were found in the depth (500–1000 m layer). Their lipid reserves were enough to overwinter and probably to ascend, molt and reproduce. At the same time some of the deep CVs had much lower protein and lipid content and could survive only for 4–5 months. Our own and literature data led to the conclusion that females of C. acutus reach adulthood at the age of more than one year while development of males could be completed in one year.     

Light and productivity of Antarctic phytoplankton during austral summer in an ice edge region in the Weddell-Scotia Sea

The photosynthesis-irradiance relationships (P-I curves) ofnatural plankton samples were studied in the Weddell Sea ice-edgezone, between Elephant Island and South Orkney Islands, duringthe austral summer of 1988–89. Three water bodies weredistinguished in the region: Bellingshausen Sea waters modifiedafter flowing through Drake Passage and Bransfield Strait, WeddellSea waters and Weddell Sea waters modified by melting. The stationssituated in modified Bellingshausen waters showed a net phytoplanktoncomposition which was different from that of the other two waterbodies. Weddell Sea waters and Weddell Sea waters modified bymelting of sea ice had the same net phytoplankton composition.In the area of modified Weddell Sea waters, there was an accumulationof phytoplankton in the upper 40 m (>4 mg Chl m^–1).p^B, and     

Distribution and feeding of chaetognaths in the epipelagic zone of the Lazarev Sea (Antarctica) during austral summer

This study seeks to determine the effects of local hydrography on the distribution, abundance and feeding of chaetognaths in the Lazarev Sea, an area strongly controlled by physical processes which has been held responsible for initiating the Weddell Polynya. Zooplankton samples were taken at 39 stations on four transects located between 6°W and 3°E and from 60°S to 70°S between surface and 350 m. The dominant species, Eukrohnia hamata, accounted for 86.5% of all chaetognaths, followed by Sagitta gazellae (8.1%) and Sagitta marri (5.4%). These three species showed distinct vertical and horizontal distribution patterns. While E. hamata and S. marri had maximum abundances below 250 m depth, S. gazellae showed a narrow distribution band in the upper 150 m depth. The distribution pattern was strongly modified at the Greenwich meridian with an upward transport of a high abundance of deep dwelling organisms (S. marri and E. hamata) and a displacement of S. gazellae to the surface, likely coupled with the rise of the warm, saline halo around the Maud Rise. Small copepods were the main prey of all three chaetognath species. Feeding rates (FR) varied among species and depth. Sagitta marri showed the highest FR with 0.38 prey d⁻¹, followed by S. gazellae and E. hamata (0.22 and 0.07 prey d⁻¹). Feeding rates were usually highest in the 25–80-m stratum. Size distribution and maturity of E. hamata revealed a dominance of small and immature organisms along all depths and stations, suggesting that this area might be acting as an important source of recently spawned organisms to the surface.     

Copepods in summer platelet ice in the eastern Weddell Sea,Antarctica

Copepods in platelet-ice layers underlying fast ice and in the water column below were studied at Drescher Inlet, eastern Weddell Sea in February 1998. Three copepod species were found: Drescheriella glacialis and Paralabidocera antarctica occurred in platelet-ice layers, while Stephos longipes was only present in the water column. The distribution of all species varied considerably between station and depth. D. glacialis dominated the platelet-ice community and occurred at all five platelet-ice sampling sites, except one, with numbers of up to 26 ind. l^–1. In contrast, P. antarctica was only found in low numbers (up to 2 ind. l^–1) at one site. The total copepod abundance in the platelet ice was not associated with algal biomass, although it was strongly correlated with high ammonium concentrations (up to 9 M) in the interstitial water between the platelets. This is the first indirect evidence to support the hypothesis that zooplankton excretion can partly account for the high ammonium values often found in platelet-ice layers.     

Phytoplankton biomass and production during late austral spring (1991) and summer (1993) in the Bransfield Strait

Phytoplankton biomass and productivity were measured during two cruises in the Bransfield Strait in December 1991 (D91) and January/February 1993 (J93). Strong seasonal variability in productivity values was observed due to differences in the physiological response of phytoplankton. However, although the photosynthetic capacity of phytoplankton was markedly lower in D91 [P _m ^B =0.61 ± 0.25 mg C (mg Chla)⁻¹ h⁻¹] than in J93 [P _m ^B =2.18 ± 0.91 mg C (mg Chla)⁻¹ h⁻¹], average water column chlorophyll values in different areas of the strait were approximately similar in D91 (49–78 mg Chla m⁻²) and J93 (22–76 mg Chla m⁻²). The spatial distribution of chlorophyll was patchy and generally associated with the influence of the different water masses that meet together in the Bransfield Strait. No correlation was found between the mixed layer depth and either the integrated chlorophyll or the productivity. Our results suggest that major phytoplankton blooms in the Bransfield Strait are advected from the nearby Gerlache Strait or Bellingshausen Sea following the main eastward surface currents. Accepted: 5 July 1998     

Sea surface distribution of coccolithophores in the eastern Pacific sector of the Southern Ocean (Bellingshausen and Amundsen Seas) during the late austral summer of 2001

Horizontal distributions of coccolithophores were observed in sea surface water samples collected on the RV Polarstern between 27 February and 10 April, 2001, in the Pacific sector of the Southern Ocean (Bellingshausen and Amundsen Seas). These samples were analyzed to gain information about the distribution of coccolithophores in relation to the oceanic fronts of the Southern Ocean. A total of fifteen species of coccolithophores were identified, showing cell abundances of up to 67 × 10³ cells/l down to 63°S. Emiliania huxleyi was the most abundant taxon, always accounting for more than 85% of the assemblage. The second most abundant species was Calcidiscus leptoporus, with values lower than 7%. Cell density increases significantly in both the Subantarctic and Polar Fronts (155 and 151 × 10³ cells/l, respectively), decreasing abruptly in the intervening Polar Frontal Zone and to the south of the Polar Front. Although temperature at high latitudes is the main factor controlling the biogeographical distribution of coccolithophores, at the regional level (Southern Ocean) the frontal systems, and consequently nutrient distribution, play a crucial role.     

Surface phytoplankton composition and carbon biomass distribution in the Crozet Basin during austral summer of 1999: variability across frontal zones

Surface phytoplankton assemblages were studied in January/February 1999 in the Crozet Basin (43°50S–45°20S; 61°E–64°30E) between the northern Polar Zone and the Agulhas Front. Cell concentrations increased several fold northwards from the SubAntarctic Zone (SAZ) and reached peak numbers (average 2×10⁶ cells l^–l ) in the central and western Subtropical Zone (STZ). The most spectacular increase in cell numbers occurred at the Subtropical Front (STF) and was attributed to dinoflagellates and diatoms. Nanoflagellates and picoplankton were dominant in the entire area (average 2.8×10⁵–1.6×10⁶ cells l^–l). In the SAZ they were followed by coccolithophorids, dinoflagellates and diatoms. In the STZ coccolithophorids were often outnumbered by dinoflagellates. Diatoms were dominated by Pseudonitzschia delicatissima and were generally the least abundant algae, but reached peak densities of 1.2–4×10⁵ cells l^–l at, and north of the STF. Coccolithophorids contained mainly Emiliania huxleyi, but in the SAZ and STF Gephyrocapsa oceanica was a co-dominant species. Dinoflagellates were dominated by nano-sized species of Gymnodinium, Gyrodinium and Prorocentrum. The numbers of dinoflagellate and coccolithophorid species increased considerably in the convergence zone (STZ), which suggests their in-situ development. Heterotrophic dinoflagellates and ciliates were mainly present in the subtropics. Cell carbon biomass was attributed chiefly to auto- and heterotrophic dinoflagellates (av. 23–72 g C l^–l; 68–87%), showing their important contribution to the carbon flow. Variations in cell concentrations across the fronts and water masses, and the distribution of major species were most likely controlled by the combined effect of such factors as nutrient renewal in the convergence zone, availability of iron, increased water-column stability at fronts, and high horizontal gradients in surface-water temperature.     

Algal and bacterial processes in platelet ice during late austral summer

 The biota inhabiting layers of platelet ice were investigated in the Weddell Sea during late austral summer. Due to meltwater release, the salinity of the interstitial water between platelets was reduced. Algae and bacteria accumulated within this ice environment attaining concentrations of up to 500 μg in total pigments (chlorophyll a plus phaeopigments) and 2 mg in bacterial biomass per liter. Pennate diatoms of the genus Fragilariopsis were most common in the platelet layer, while ice-free water was dominated by autotrophic nanoflagellates. Protozoa contributed only 5% or less to the total protistan (microalgae plus protozoa) cell concentration in the ice, compared to about 10% in open water, thus suggesting a low grazing pressure within the platelet habitat. The bulk of bacterial biomass occurred within the dense assemblages of pennate diatoms that grew attached to the ice platelets. Algal and bacterial concentrations in the interstitial water between platelets were much lower. Measurements of bacterial growth showed that substantial heterotrophic potential can be established within assemblages inhabiting late summer platelet ice. Small-scale analyses of bacterial activity patterns revealed that those bacteria that were closely associated with ice and/or algae showed considerably less biomass-specific substrate uptake than cells that occurred freely suspended in the interstitial water, indicating that their physiological state differed. Received: 21 October 1995/Accepted: 27 January 1996     

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     

The zooplankton community in the vicinity of the ice edge,western Weddell Sea,March 1986

Summary The zooplankton community in the vicinity of the ice edge in the west central Weddell Sea was investigated in the late austral summer (March 1986). Sampling was done with two ships operating concurrently, one in the pack ice and the other in the adjcent open sea. Metazoan microzooplankton (<1 mm) was most abundant in the epipelagic zone. It consisted mostly of copepod nauplii and copepods of the genera Oithona, Oncaea, Ctenocalanus and Microcalanus. While species composition was similar in both areas, vertical patterns differed in that the microzooplankton had sparse populations in the upper 50 m under the ice. This may have been related to water temperature which in the upper 50 m under the ice was more than 1°C cooler than in the open sea. Zooplankton in the 1–20 mm size range was dominated by the calanoid copepods Metridia gerlachei, Calanus propinquus and Calanoides acutus which constituted half the biomass in the upper 1000 m. Their populations had highest densities in the upper 150 m, though much of the C. acutus population resided below 300 m. Metridia gerlachei and C. propinquus underwent diel vertical migrations in both areas whereas C. acutus did not migrate. Species diversity in the epipelagic zone was moderate and the fauna was characterized by species typical of the oceanic east wind drift. Diversity increased with depth and was due primarily to the appearance of circumpolar mesopelagic copepods in Weddell Warm Deep Water. Biomass of 1–20 mm zooplankton in the 0–1000 m zone was low (1.1–1.3 gDWm^-2) compared to other Southern Ocean areas investigated with comparable methods. It is suggested that this is related to Weddell circulation patterns and the resulting low annual primary production in the central Weddell Sea.     

Mesozooplankton distribution patterns and grazing impacts of copepods and Euphausia crystallorophias in the Amundsen Sea, West Antarctica, during austral summer

The rapid melting of glaciers as well as the loss of sea ice in the Amundsen Sea makes it an ideal environmental setting for the investigation of the impacts of climate change in the Antarctic on the distribution and production of mesozooplankton. We examined the latitudinal distribution of mesozooplankton and their grazing impacts on phytoplankton in the Amundsen Sea during the early austral summer from December 27, 2010 to January 13, 2011. Mesozooplankton followed a latitudinal distribution in relation to hydrographic and environmental features, with copepods dominating in the oceanic area and euphausiids dominating in the polynya. Greater Euphausia crystallorophias biomass in the polynya was associated with lower salinity and higher food concentration (chlorophyll a, choanoflagellates, and heterotrophic dinoflagellates). The grazing impact of three copepods (Rhincalanus gigas, Calanoides acutus, and Metridia gerlachei) on phytoplankton was low, with the consumption of 3 % of phytoplankton standing stock and about 4 % of daily primary production. Estimated daily carbon rations for each of the three copepods were also relatively low (<10 %), barely enough to cover metabolic demands. This suggests that copepods may rely on food other than phytoplankton and that much of the primary production is channeled through microzooplankton. Daily carbon rations for E. crystallorophias were high (up to 49 %) with the grazing impact accounting for 17 % of the phytoplankton biomass and 84 % of primary production. The presence of E. crystallorophias appears to be a critical factor regulating phytoplankton blooms and determining the fate of fixed carbon in the coastal polynyas of the Amundsen Sea.