Letteratura scientifica selezionata sul tema "Zooplankton – Antarctic Ocean"

Abstract Acoustic surveys for biomass estimation require accurate identification of echoes from the target species. In one objective technique for identifying Antarctic krill, the difference between mean volume-backscattering strength at two frequencies is used, but can misclassify small krill and other plankton. Here, we investigate ways to improve target identification by including characteristics of backscattering energy and morphology of aggregations. To do this, multi-frequency acoustic data were collected concurrently with target fishing of Antarctic krill and other euphausiid and salp aggregations. Parameter sets for these known aggregations were collated and used to develop empirical classifications. Both linear discriminant-function analysis (DFA) and the artificial neural network technique were employed. In both cases, acoustic-backscattering energy parameters were most important for discriminating between Antarctic krill and other zooplankton. However, swarm morphology and other parameters improved the discrimination, particularly between krill and salps. Our study suggests that for krill-biomass estimates, a simple DFA based on acoustic-energy parameters is a substantial improvement over current dB-difference acoustic methods; but studies requiring the discrimination of zooplankton other than krill must still be supported by target fishing.

Abstract. The Southern Ocean provides a vital service by absorbing about one-sixth of humankind's annual emissions of CO2. This comes with a cost – an increase in ocean acidity that is expected to have negative impacts on ocean ecosystems. The reduced ability of phytoplankton and zooplankton to precipitate carbonate shells is a clearly identified risk. The impact depends on the significance of these organisms in Southern Ocean ecosystems, but there is very little information on their abundance or distribution. To quantify their presence, we used coulometric measurement of particulate inorganic carbonate (PIC) on particles filtered from surface seawater into two size fractions: 50–1000 µm to capture foraminifera (the most important biogenic carbonate-forming zooplankton) and 1–50 µm to capture coccolithophores (the most important biogenic carbonate-forming phytoplankton). Ancillary measurements of biogenic silica (BSi) and particulate organic carbon (POC) provided context, as estimates of the biomass of diatoms (the highest biomass phytoplankton in polar waters) and total microbial biomass, respectively. Results for nine transects from Australia to Antarctica in 2008–2015 showed low levels of PIC compared to Northern Hemisphere polar waters. Coccolithophores slightly exceeded the biomass of diatoms in subantarctic waters, but their abundance decreased more than 30-fold poleward, while diatom abundances increased, so that on a molar basis PIC was only 1 % of BSi in Antarctic waters. This limited importance of coccolithophores in the Southern Ocean is further emphasized in terms of their associated POC, representing less than 1 % of total POC in Antarctic waters and less than 10 % in subantarctic waters. NASA satellite ocean-colour-based PIC estimates were in reasonable agreement with the shipboard results in subantarctic waters but greatly overestimated PIC in Antarctic waters. Contrastingly, the NASA Ocean Biogeochemical Model (NOBM) shows coccolithophores as overly restricted to subtropical and northern subantarctic waters. The cause of the strong southward decrease in PIC abundance in the Southern Ocean is not yet clear. The poleward decrease in pH is small, and while calcite saturation decreases strongly southward, it remains well above saturation ( > 2). Nitrate and phosphate variations would predict a poleward increase. Temperature and competition with diatoms for limiting iron appear likely to be important. While the future trajectory of coccolithophore distributions remains uncertain, their current low abundances suggest small impacts on overall Southern Ocean pelagic ecology.

AbstractSeasonal ice of the Southern Ocean, occupying some 15 x 106 km2, supports a distinctive biota based on algae that live on, within and immediately beneath the ice floes. How this annually-forming habitat recruits its biota, and the fate of the biota after the ice thaws in late summer, are little-known. Studies in the Weddell Sea in 1984–88 have shown that the seasonal ice is important as the wintering substrate of krill Euphausia superba which, together with other zooplankton and fish, supports a large breeding population of seals and penguins. Clearly a key habitat in the economy of the Southern Ocean, this seasonal ice is likely to be vulnerable to small climatic changes.

Preliminary results of the pilot study of the zooplankton in the region between the Ross and Scotia Seas from November 2017 to April 2018 are presented. In total, 53 zooplankton samples were collected in the top 100 m water layer using vertical tows of a 0.1 m2 Juday net from four Ukrainian longliners operating during the Antarctic toothfish fishery. Total zooplankton abundance ranged from 3 to 2836 ind m−3 with a global mean of 360 ± 550 (±1 SD) ind m−3. The highest abundances were recorded at the northeastern Ross Sea. At those stations, small copepods (mainly Oithona spp., Oncaea spp., Ctenocalanus spp. and copepod nauplii) numerically dominated the samples. Total biomass ranged from 0.3 to 85 mg DW m−3 with a mean of 10.9 ± 14.5 mg DW m−3. The highest biomasses were recorded at the eastern Ross Sea, where pelagic tunicates Salpa thompsoni, siphonophores and ctenophora Callianira sp. accounted for >90% of total zooplankton biomass. At other stations, zooplankton biomass generally ranged from 5 to 20 mg DW m−3 with no clear pattern in distribution. The community composition was driven by the sampling latitude and/or season rather than longitudinally. This pilot study emphasized the unique opportunity to investigate zooplankton dynamics in the regions traditionally not sampled during the oceanographic surveys. It also created unprecedented opportunities to increase the seasonal and geographical zooplankton sampling coverage using ships of opportunity at a fraction of a dedicated oceanographic survey costs. The potential of such surveys are enormous in both providing invaluable information, contributing to existing long-term databases and enhancing an international collaboration in the Southern Ocean, particularly in light of recent modeling initiatives of the whole Antarctic system undertaken by the Commission for the Conservation of Antarctic Marine Living Resources.

Environmental contextPersistent organic pollutants (POPs) are potentially toxic chemicals capable of long distance transport and are often found far from their source. Little is known of their behaviour in Antarctica, where the marine plankton food web is driven by strong seasonal variations in solar radiation. Here the first dynamic coupled ecosystem–fugacity model to describe how POPs distribute through the Antarctic environment is presented. The model is used to identify the important processes that govern the presence of hexachlorobenzene in Antarctic plankton. AbstractPolar regions can be repositories for many persistent organic pollutants (POPs). However, comparatively little is known of the movement and behaviour of POPs in Antarctic ecosystems. These systems are characterised by strong seasonal effects of light on plankton dynamics. This work describes a mass-conserving, fugacity-based dynamic model to describe the movement of POPs in the Antarctic physical and plankton systems. The model includes dynamic corrections for changes in the population volumes and the temperature dependence of the fugacity capacities, and was developed by coupling a dynamic Nutrient–Phytoplankton–Zooplankton–Detritus (NPZD) ecosystem model to fugacity models of the chemistry and biology of the Southern Ocean. The model is applied to the movement of hexachlorobenzene, a POP found in the Antarctic environment. The model predicts that the burden of HCB in the plankton varies with the seasonal cycle in Antarctic waters, and induces a seasonal variation in the biomagnification factor of zooplankton. This suggests that time series of POP concentrations in Antarctic biotic and abiotic systems should be measured over complete seasonal cycles. Furthermore, detritus is shown to be a key contributor to the movement of POPs in polar environments, linking physical and biological components of the model.

The organic carbon particle export to the interior layers in the Southern Ocean in the New Zealand–Tasmania Sector was approximately 170 mmolC m−2 yr−1. The export of particulate inorganic carbon in CaCO3 was 110 mmolC m−2 yr−1 and was contributed mostly by pteropods shells in the Antarctic Zones. The Si flux from biogenic opal at the sub-Antarctic Zone was 67 mmolSi m−2 yr−1 and rapidly increased to the south up to nearly 1 molSi m−2 yr−1 in the Antarctic Zone. The Antarctic Polar Front clearly demarcated the area where the biological pump was driven by CaCO3 to the north and biogenic SiO2 particle export to the south. Summer stratification caused by the sub-zero winter water layer in the Seasonal Ice Zone (SIZ) curtails the zooplankton community and hinders the replenishment of Fe. This hypothesis explains the large organic carbon export with large f- and export ratios at the SIZ and extremely large opal production at the Antarctic Circumpolar Zone. Estimated regeneration rate of CO2 from the export production and settling particulate fluxes of organic carbon in the water column between 100 m to 1 km was about 13 mmolC m−2 d−1 in the Antarctic Zone and Polar Frontal Zone.

Samples of phyto- and zooplankton were collected in the Indian sector of the Southern Ocean (38–67°S, 18–84°E) and analysed for organochlorine residues (PCBs and pesticides). The PCB concentration in particulate matter (mainly phytoplankton) appeared to be high and similar to that of temperate zones: 0.7 μg g−1 dry weight. Contamination levels were more constant expressed per water volume than per dry weight, and seven times lower (1.2 μg m−3) than in northern temperate zones (8.8 μm−3 in the North Sea). The Antarctic ecosystems are thus less contaminated than temperate ones — as expected — but the very low phytoplankton biomass present cause high PCB levels per unit of biomass. These results confirm the necessity of using different systems of units in order to correctly express the contamination levels and to identify the main mechanisms responsible for the accumulation of stable pollutants. PCB levels in netplankton samples (mainly zooplankton) were comparable with phytoplankton on a dry weight basis (0.7 μm−3), lower on a lipid weight basis (5.8 μg g−1 lw for netplankton, 16.3 for particulate matter) and were much higher per seawater volume (27.2 μg m−3 for netplankton, 1.2 for particulate matter). Netplankton contamination is comparable in the Antarctic (0.35 μg g−1 dw) and the North Sea (0.70) since zooplankton feeding on phytoplankton has similar levels of contamination in both ecosystems. Lindane, heptachlor epoxide, dieldrin, DDE and DDT were observed in various samples at trace levels. The high DDE/DDT ratio reflects the more recent origin of Antarctic organochlorines.

The role of zooplankton grazing on dimethylsulfide (DMS) and dissolved dimethylsulfoniopropionate (DMSPd) production was investigated in the Antarctic Ocean in January and February 2002. Dominant herbivorous macrozooplankton of this region, the Antarctic krill Euphausia superba and the tunicate Salpa thompsoni, were used in shipboard incubation experiments. The concentration of DMSPd + DMS increased in the water during incubation with krill. The production rate was 2.96 ± 2.78 nmol DMSPd + DMS·krill–1·h–1 (mean ± SD). In addition, the DMSPd + DMS production rate was linearly related to the ingestion rate of krill (r2 = 0.664, p ≤ 0.01). Addition of salps to natural surface water, however, did not change the DMSPd + DMS concentrations. During the experiments, both animals fed on phytoplankton cells. The fecal pellets of krill contained broken phytoplankton cells, whereas those of salps contained unbroken cells. These results suggest that sloppy feeding by krill is a more likely mechanism for producing DMS and DMSPd than the direct ingestion of phytoplankton cells by salps. The decrease of DMS concentrations in the upper 200 m of the water column from January to February may be explained, in part, by changes in the composition of the macrozooplankton community.

AbstractThe community composition of zooplankton with an emphasis on copepods was assessed in the frontal zones of the Indian sector of the Southern Ocean (SO) during summer 2013. Copepods were the dominant group in both the bongo net and multiple plankton sampler across the entire region. High zooplankton abundance was recorded along each transect in the Polar Front (PF). Community structure in this front was dominated by common taxa, including Ctenocalanus citer, Clausocalanus spp., Calanoides acutus, Calanus propinquus, Calanus australis and Rhincalanus gigas, which together accounted for > 62% of the total abundance. Calocalanus spp., Neocalanus tonsus and C. propinquus were indicator species in the Sub-Tropical Front (STF), Sub-Antarctic Front and PF, respectively. A strong contrast in population structure and biovolume was observed between then PF and the STF. The community structure of smaller copepods was associated with the high-temperature region, whereas communities of larger copepods were associated with the low-temperature region. Thus, it seems probable that physical and biological characteristics of the SO frontal regions are controlling the abundance and distribution of zooplankton community structure by restricting some species to the warmer stratified zones and some species to the well-mixed zone.

The role of fish in the Antarctic food web in inshore and offshore waters is analysed, taking as an example the coastal marine communities of the southern Scotia Arc (South Orkney Islands and South Shetland Islands) and the west Antarctic Peninsula. Inshore, the ecological role of demersal fish is more important than that of krill. There, demersal fish are major consumers of benthos and also feed on zooplankton (mainly krill in summer). They are links between lower and upper levels of the food web and are common prey of other fish, birds and seals. Offshore, demersal fish depend less on benthos and feed more on zooplankton (mainly krill) and nekton, and are less accessible as prey of birds and seals. There, pelagic fish (especially lantern fish) are more abundant than inshore and play an important role in the energy flow from macrozooplankton to higher trophic levels (seabirds and seals). Through the higher fish predators, energy is transferred to land in the form of fish remains, pellets (birds), regurgitation and faeces (birds and seals). However, in the general context of the Antarctic marine ecosystem, krill (Euphausia superba) plays the central role in the food web because it is the main food source in terms of biomass for most of the high level predators from demersal fish up to whales. This has no obvious equivalent in other marine ecosystems. In Antarctic offshore coastal and oceanic waters the greatest proportion of energy from the ecosystem is transferred to land directly through krill consumers, such as flying birds, penguins, and seals. Beside krill, the populations of fish in the Antarctic Ocean are the second most important element for higher predators, in particular the energy-rich pelagic Myctophidae in open waters and the pelagic Antarctic silver fish (Pleuragramma antarcticum) in the high Antarctic zone. Although the occurrence of these pelagic fish inshore has been poorly documented, their abundance in neritic waters could be higher than previously believed.

The trophodynamics of the numerically dominant mesozooplankton (200-2000 m) in the vicinity of the Subtropical Convergence (STC) in the Indian sector of the Southern Ocean during austral autumn (April / May) 2007 were investigated as part of the Southern Ocean Ecosystem Variability Study. The survey consisted of six north-south transects each bisecting the STC between 38º to 43ºS and 38º to 41º45’E. In total, 48 stations situated at 30 nautical mile intervals were occupied over a period of ten days. Hydrographic data revealed a well defined surface and sub-surface expression of the STC, which appeared to meander considerably between 41ºS and 41º15’S. Surface chlorophyll-a (chla) concentrations were low, ranging between 0.08 and 0.68 mg chl-a.m-3 and were generally dominated by the picophytoplankton (<2 m) which made up 66.6% (SD±17.6) of the total pigment. Chl-a concentrations integrated over the top 150m of the water column ranged between 11.97 and 40.07 mg chl-a.m-2 and showed no significant spatial patterns (p>0.05). Total integrated mesozooplankton abundance and biomass during the study ranged between 3934.9 and 308521.4 ind.m-2 (mean = 47198.19; SD±62411.4 ind.m-2) and between 239.8 and 4614.3 mg Dwt.m-2 (mean = 1338.58; SD ±1060.5), respectively. Again, there were no significant spatial patterns in the total mesozooplankton abundance or biomass within the region of study (p>0.05). No significant correlations were found between biological (chlorophyll-a concentrations and zooplankton abundance) and physico-chemical variables (temperature and salinity) (p>0.05). The total mesozooplankton community was numerically dominated by copepods of the genera Pleuromamma, Calanus, Oncaea and Oithona. Other important representatives of the mesozooplankton community included the tunicate, Salpa thompsoni, and the pteropod, Limacina retroversa. At the 40% similarity level, numerical analysis identified five distinct mesozooplankton groupings within the survey area. Differences between the groupings were associated with changes in the relative contribution of numerically dominant species rather than the presence or absence of individual species. No groupings were associated with any specific feature of the front within the survey area. The feeding rates of the six most numerically abundant mesozooplankton species (Calanus simillimus, Limacina retroversa, Pleuromamma abdominalis, Clausocalanus breviceps, Oncaea conifera, Salpa thompsoni) accounting for on average 39% of the total mesozooplankton counts, were investigated using the gut fluorescence technique. For all species, the total gut pigment contents during the night time were significantly higher than the daytime values (p<0.05 for all species). The gut evacuation rates (k) for selected mesozooplankton ranged between 0.14 and 0.81 h-1. The ingestion rates ranged between 147.8 and 5495.4 ng(pigm)ind-1.day-1 which corresponded to a daily ration of between 2.4 and 10.9% body carbon. The combined grazing impact of the selected species on the daily phytoplankton standing stock was highly variable and ranged between 1.2 and 174.1% with an average of 27.3% (SD±38.78%) within the survey area. The highest grazing impact (>60%) was typically associated with those stations where the pteropod, L. retroversa, and the tunicate, S. thompsoni, contributed more than 5% of the total mesozooplankton counts. No significant differences were found in the grazing impact of any or all selected species situated either north, south or in the immediate vicinity of the front (p>0.05 in all cases). The lack of defined spatial patterns in the mesozooplankton abundance and community structure suggests that the STC did not act as a significant biogeographic barrier to the distribution of mesozooplankton during the study. It is presumed that the large scale mixing event caused by a storm prior to this study was responsible for the observed lack of elevated biological activity within the region of the STC.

The population structure and feeding dynamics of the hyperiid amphipod, Themisto gaudichaudii, was investigated during two cruises of the South African National Antarctic Programme conducted in the Indian sector of the Polar Frontal Zone during austral autumn (April) 2004 and 2005. During the 2004 cruise the frontal features that delimit the PFZ converged to form a single distinctive feature. In 2005, the research cruise was conducted in the vicinity of a cold-core eddy which was spawned from the Antarctic Polar Front. Total mesozooplankton abundance and biomass during the 2004 study ranged from 55.19 to 860.57 ind. m⁻³, and from 2.60 to 38.42 mg dwt m⁻³, respectively. In 2005 the abundance and biomass ranged from 23.1 to 2160.64 ind. m⁻³, and from 0.76 to 35.16 mg dwt m⁻³, respectively. The mesozooplankton community was numerically dominated by copepods, pteropods, and ostracods during both surveys. The abundance and biomass of Themisto gaudichaudii in the region of investigation was < 0.2 ind. m⁻³ (range 0.01 to 0.15 ind. m⁻³) and < 0.06 mg dwt m⁻³ (range 0.02 to 0.06 mg dwt m⁻³) during 2004, while in 2005 the abundance and biomass of the amphipod ranged from < 0.01 to 0.2 ind. m⁻³ and < 0.01 to 0.04 mg dwt m⁻³, respectively. These values correspond to < 1% of the total mesozooplankton abundance and biomass during both surveys. T. gaudichaudii exhibited no significant spatial patterns in abundance, biomass and total length during both 2004 and 2005 (p > 0.05 in all cases). A key feature of the two investigations was the virtual absence of juveniles (total length < 15 mm) among the amphipod population, supporting the suggestion that they exhibit strong seasonal patterns in reproduction. Gut content analysis during both years indicated that for both the male and female amphipods’, copepods were the most prevalent prey species found in stomachs, followed by chaetognaths and pteropods. Results of electivity studies indicate that T. gaudichaudii is an opportunistic predator, generally feeding on the most abundant mesozooplankton prey. Results of in vitro incubations indicated that the total daily feeding rate of T. gaudichaudii during 2004 ranged from 11.45 to 20.90 ind. m⁻³ d⁻¹, which corresponds to between 0.12 and 1.64% of the total mesozooplankton standing stock. In 2005, the feeding rate ranged between 0.1 and 1.73% of the total mesozooplankton standing stock. The low predation impact of T. gaudichaudii during this study can be related to their low abundances and high interannual variability throughout the region of investigation.

The distribution, abundance, chemical composition, metabolism, and feeding ecology of the tentaculate ctenophore, Callianira antarctica (Chun 1897), were investigated during austral winter 2001and autumn & winter 2002, in the vicinity of Marguerite Bay west of the Antarctic Peninsula. Callianira antarctica had a widespread distribution during autumn and winter, and variable abundance (0.02 to 2.6 ind. m-2) during winter 2001 associated with specific circulation features. Size frequency distributions for autumn and winter suggest that more than half of the C. antarctica population may have experienced 'degrowth' during winter due to low food availability. Callianira antarctica is a fairly robust ctenophore with geometric mean (geomean) carbon (C) and nitrogen (N) values of 8.41 and 1.83% dry weight (DW), respectively. Winter oxygen consumption and ammonium excretion rates ranged from 0.059 to 0.410 micro l O2 [mg DW]-1 h-1 and 0.60 to 31.1 µg-at N [g DW]-1 h-1, respectively, at 0oC. Daily minimum maintenance rations based on respiration experiments were 2.7% to 3.6% of the total body carbon (TBC) for small ctenophores, and 1.4% to 1.9% TBC for larger ctenophores. Calanoid copepods and larval and juvenile Antarctic krill were offered to ctenophores in incubation experiments. Digestion times were variable, lasting 8 to 20 h, and were independent of ctenophore size and dependent on number and type of prey. Gut content analysis from one autumn and two winter seasons indicated C. antarctica preyed on both copepods and krill in situ, with an increased dependence on larval krill during winter. Lipid biomarker analysis on C. antarctica and their potential prey confirmed these results. Divers observed aggregations of C. antarctica passively drifting with tentacles extended near dense concentrations of larval Euphausia superba during winter. These observations along with gut content and lipid biomarker analysis suggest that larval krill is an important prey item for C. antarctica during winter.

The community structure and predation impact of carnivorous macrozooplankton (>2 cm; chaetognaths, medusae, ctenophores and mysids), with particular emphasis on the chaetognaths Eukrohnia hamata and Sagitta gazellae, were investigated during three surveys conducted in late austral summer (April/May) of 2001, 2004 and 2005 in the Polar Frontal Zone in the vicinity of the Prince Edward Islands (46º45’S, 37º50’E), Southern Ocean. The 2001 survey formed part of the Marion Offshore Variability Ecosystem Study (MOVES II), while the 2004 and 2005 surveys formed part of the Dynamics of Eddy Impacts on Marion’s Ecosystem study (DEIMEC III and IV respectively). Macrozooplankton samples were collected using WP-2, RMT-8 and Bongo nets. Results of the hydrographic survey indicated that the region of investigation, the Polar Frontal Zone (PFZ), is an area of high mesoscale variability. During the 2004 survey the Antarctic Polar Front (APF) and the Subantarctic Front (SAF) merged to form an intense frontal feature with subsurface temperature and salinity ranging from 8.5-7.5ºC and 34.15-33.88, respectively. A cyclonic cold core eddy, believed to have been spawned from the APF, was observed during the 2005 survey. Macrozooplankton abundance and biomass ranged from 0 to 43.731 ind. m⁻³, and from 0 to 41.55 mg wwt m⁻³ respectively, during the three surveys. Among the carnivorous macrozooplankton, chaetognaths (Eukrohnia hamata and Sagitta gazellae) were most prominent, contributing up to 85% of the total biomass during all three surveys. Elevated biomass values were found near and within the frontal feature during the 2004 survey, and also along the eddy edge during the 2005 survey. However, hierarchical cluster analysis did not reveal the presence of distinct zooplankton groupings associated with the various water masses encountered during the surveys and this is probably due to the high mesoscale variability in oceanographic conditions that are characteristic of the PFZ. The total average predation impact of the selected carnivorous macrozooplankton during the 2001, 2004 and 2005 surveys accounted for 4.93 ± 6.76%, 0.55 ± 0.51% and 4.88 ± 4.45 of the mesozooplankton standing stock, respectively. S. gazellae had the highest consumption rate in all three surveys, consuming up to 800 g Dwt 1000m⁻³d⁻¹ during the study. Of the two chaetognaths, E. hamata dominated the chaetognath standing stock. The combined abundance and biomass values of E. hamata and S. gazellae ranged from 0 to 43.73 ind. m⁻³ and from 0 to 41.551 mg wwt m⁻³ respectively, during the three surveys. Inter-annual variability in the chaetognath densities was apparent. Highest abundances and biomasses tended to be associated with specific water masses, confirming the existence of a relationship between zooplankton community structure and hydrographic conditions. Generally, about 90% of the chaetognaths contained no food in their guts. S. gazellae consumed a wider variety of prey. Oil droplets occurred in the guts of ≈ 51% of E. hamata. Cannibalism was low in both species, but greater in S. gazellae than E. hamata. During the three surveys, the feeding rate values of E. hamata and S. gazellae went up to 0.48 and 2.099 prey d⁻¹ respectively. S. gazellae also had a greater predation impact on the mesozooplankton standing stock than E. hamata. The mean predation impact of the chaetognaths combined was 0.31 ± 0.291%, 0.52 ± 0.28% and 0.53 ± 0.56% of the mesozooplankton standing stock during the 2001, 2004 and 2005 surveys, respectively. During all three surveys, the majority of individuals (≈ 76%) of the chaetognaths were at stage I maturity, suggesting that during the time of study the chaetognaths were not reproducing. In both species a significant difference (log-linear analysis, p < 0.05) in maturities between the years investigated was observed. In general, there were no differences in lengths and maturities between the different water masses encountered during the surveys. The lengths of E. hamata and S. gazellae ranged from 5 to 24 mm and from 9.4 to 63.6 mm, respectively.

Mesozooplankton community structure and grazing impact in the Polar Frontal Zone (PFZ) of the Southern Ocean were investigated during two cruises of the South African National Antarctic Programme (SANAP), the Marion Offshore Ecosystem Variability Study I & II (MOEVS). During the first cruise (MOEVS I), a meso-scale oceanographic grid survey was conducted in the upstream region of the Prince Edward Islands (PEI) in austral autumn (April) 2001. Mesozooplankton samples, collected using a Bongo net (fitted with 200 and 300µm mesh nets) at depths between 200 and 300 m, were separated into three size fractions: 200-500 µm; 500-1000 µm; 1000-2000 µm by reverse filtration. Total surface (depth <5 m) chlorophyll-a (chl-a) concentration (measured fluorometrically) during the study ranged between 0.11 and 0.34 µg 1^(-1) and was always dominated by picophytoplankton (<2.0 µm). Total mesozooplankton abundance and biomass during the survey ranged between 49 and 1512 ind. m^(-3) and between 0.7 and 25 mg Dwt. m^(-3), respectively. Throughout the survey, the 200-500 µm class numerically dominated the mesozooplankton community, comprising an average of ~ 69% (SD = ± 12.3%). The dominant species in the 200-500 µm size fraction were the copepods Oithona similis, Calanus simillimus and Metridia lucens and the pteropod, Limacina retroversa. However, in terms of biomass, the 1000-2000 µm group was predominant, with dry weight values constituting an average of ~ 66% (SD = ± 10.2%). Biomass was dominated by carnivorous zooplankton, particularly the euphausiids, Euphausia vallentini and Thysanoessa vicina and the chaetognaths, Sagitta gazellae and Eukrohnia hamata. Three distinct groupings of stations were identified by multivariate analysis. The different station groupings identified reflect changes in the relative contributions of the rather than different species assemblages. During the second cruise (MOEVS II), conducted in April 2002 (austral autumn), mesozooplankton community structure and grazing impact were investigated at 13 stations in the west Indian sector of the PFZ. Total integrated chl-a biomass ranged between 11.17 and 28.34 mg chl-a m^(-2) and was always dominated by nano- and picophytoplankton (<20 µm). Throughout the study, small copepods, mainly Oithona similis and Ctenocalanus vanus, numerically dominated the mesozooplankton community comprising up to 85% (range 30 to 85%) of the total abundance. Grazing activity of the four most abundant copepods (O. similis, C. vanus, Calanus simillimus and Clausocalanus spp.), which comprised up to 93% of total mesozooplankton abundance, was investigated using the gut fluorescent technique. Results of gut fluorescence analyses indicated that C. simillimus, Clausocalanus spp. and Ctenocalanus vanus exhibited diel variability in gut pigments, with maximum values at various stages of the night. In contrast, O. similis did not demonstrate diel variation in gut pigment contents. Ingestion rates of the four copepods ranged from 23.23 to 1462.02 ng (pigm.) ind^(-1) day^(-1), depending on the species. The combined grazing impact of the four copepods, ranged between 1 and 36% of the phytoplankton standing stock per day, with the highest daily impact (~ 35.86%) occurring at stations in the vicinity of the Antarctic Polar Front. Among the copepods, O. similis and C. vanus were generally the most important consumers of phytoplankton biomass; together they were responsible for up to 89% (range 15 to 89%) of the total daily grazing impact. Carbon specific ingestion rates of the copepods varied between 42 and 320% body carbon per day, depending on the species. The study highlights the importance of small copepods in terms of both their significant contribution to total mesozooplankton numbers and their grazing impact on the phytoplankton standing stocks in the PFZ during austral autumn.

The aim of the present study was to assess the ecological role of the euthecosome pteropod, Limacina retroversa, in particular, and the mesozooplankton community, in general, in the pelagic ecosystem of the Polar Frontal Zone (PFZ), Southern Ocean. Data were collected from four oceanographic surveys to the Indian sector of the PFZ during austral autumn 2000, 2002, 2004 and 2005. Copepods, mainly Calanus simillimus, Oithona similis, Clausocalanus spp. and Ctenocalanus spp., typically dominated total mesozooplankton counts, accounting for, on average, between 75.5 % and 88.1 % (Mean = 77.4 %; SD = 13.4 %) of the total, during the present investigation. Results of the study indicate that L. retroversa may, at times, contribute substantially to total mesozooplankton abundances. During the study, L. retroversa contributed between 0.0 and 30.0 % (Mean = 5.3 %; SD = 7.1 %) to total mesozooplankton numbers. Significant small-scale variability in abundance and size structure of L. retroversa and abundance of copepods was minimal. Inter-annual variability, on the other hand, was significant between some years. Total pteropod numbers were greatest during April 2002 and 2004, while copepods exhibited greatest abundances during April 2004 only. Pearson’s Correlation analysis suggested that L. retroversa abundances were positively correlated to total surface chlorophyll-a (chl-a) concentrations. The significantly lower chl-a concentrations recorded during April 2005 may explain the reduced pteropod numbers observed during that survey. The size class structure of L. retroversa comprised mainly small and mediumsized individuals during all four surveys. This corresponds well with records from the northern hemisphere (sub-Arctic and Arctic waters) where Limacina spp. are reported to exhibit maximum spawning during mid to late-summer. Higher abundances of large individuals only occurred during April 2005, when chl-a concentrations were very low; possibly the result of delayed spawning, due to reduced food availability. Ingestion rates of the four most abundant copepods, determined using the gut fluorescence technique, ranged between 159.32 ng (pigm) ind⁻¹ day⁻¹ and 728.36 ng (pigm) ind⁻¹ day⁻¹ (Mean = 321.01 ng (pigm) ind⁻¹ day⁻¹; SD = 173.91 ng (pigm) ind⁻¹ day). Ingestion rates of L. retroversa were much higher, ranging from an average of 4 28.68 ng (pigm) ind⁻¹ day⁻¹ in April 2002 to 4 196.88 ng (pigm) ind⁻¹day⁻¹in April 2005 (Mean = 4157.36 ng (pigm) ind⁻¹ day⁻¹; SD = 35.37 ng (pigm) ind⁻¹day⁻¹). Average daily grazing rates for the pteropod varied between 0.39 mg (pigm) m⁻² day⁻¹ in April 2005 and 17.69 mg (pigm) m-2 day-1 in April 2004 (Mean = 6.13 mg (pigm) m⁻² day⁻¹; SD = 11.04 mg (pigm) m⁻² day⁻¹); corresponding average daily grazing impacts ranged between 8.4 % and 139.8 % of the phytoplankton standing stock in April 2005 and 2004, respectively (Mean = 48.5 %; SD = 84.5 %). Average daily grazing rates of the four copepods ranged from 4.58 mg (pigm) m⁻² day⁻¹ to 8.77 mg (pigm) m⁻² day⁻¹ -1, during April 2002 and 2004, respectively (Mean = 6.28 mg (pigm) m⁻² day⁻¹; SD = 5.94 mg (pigm) m⁻² day⁻¹). Collectively, the copepods removed an average of between 31.6 % and 89.8 % of the phytoplankton standing stock per day, during April 2002 and 2004, respectively (Mean = 70.8 %; SD = 86.7 %). The daily grazing impact of the copepods accounted for an average of between 40.4 % and 87.8 % of the total zooplankton grazing impact, during April 2004 and 2005, respectively (Mean = 75.0 %; SD = 65.5 %). L. retroversa was responsible for an average of 52.4 % and 59.5 % of the total zooplankton grazing impact, during April 2002 and 2004, respectively. However, during April 2005, when L. retroversa numbers were significantly lower than previous years, the pteropod contributed an average of only 7.5 % to the total zooplankton grazing impact. Thus, during the present investigation,the pteropod was responsible for removing a mean of 48.9 % of the available phytoplankton (SD = 74.9 %). The predation impact of the dominant carnivorous macrozooplankton and micronekton in the PFZ was determined during April 2004 and 2005 using daily ration estimates obtained from the literature. Additionally, gut content analysis was used to determine the contribution of L. retroversa to the diet of the dominant predators. Average predation impact ranged from 1.1 % and 5.7 % of the total mesozooplankton standing stock during April 2004 and 2005, respectively (Mean = 3.8 %; SD = 12.3 %). Chaetognaths and euphausiids dominated total carnivore numbers and made the greatest contributions to total predation impact during both years. Copepods appeared to be the main prey item of the dominant carnivorous macrozooplankton-micronekton in the region. L. retroversa was only detected in the gut contents of the amphipod, Themisto gaudichaudi, but not in either of the chaetognath species (Eukrohnia hamata and Sagitta gazellae) or the myctophid fish (Electrona spp.). The pteropod was found in 19 % of amphipod guts dissected. Pearson’s Correlation analyses showed that the four major predatory zooplankton groups found in the PFZ (chaetognaths, euphausiids, amphipods and myctophid fish) were positively correlated to abundances of L. retroversa, suggesting that the pteropod might be an important prey item for many of the carnivorous macrozooplankton/micronekton in the PFZ. To conclude, L. retroversa may play an important role in the pelagic ecosystem of the PFZ, in austral autumn. However, ocean acidification and calcium carbonate undersaturation (as a result of increased anthropogenic carbon dioxide emissions), that is predicted to occur within the next 50 – 100 years, will most likely have significant implications for the Sub-Antarctic pelagic ecosystem if L. retroversa cannot adapt quickly enough to the changes.

Trophodynamics of carnivorous zooplankton in the region of the Subtropical Convergence (STC) in the Indian sector of the Southern Ocean was investigated during austral autumn (April 2007) as part of the first cruise of the Southern Ocean Ecosystem Variability Study. Within the region of the study, the STC was well defined by the 14°C surface isotherm which separated the Agulhas Return Current and Subtropical water in the north from Sub-Antarctic waters to the south. Total average abundance (3.89 ± 5.46ind 100m-3) and biomass (0.14 ± 0.27mg Dwt 100m-3) of carnivorous zooplankton south of the front were significantly higher than the total average abundance (1.33 ± 1.81ind 100m-3) and biomass (0.03 ± 0.05mg Dwt 100m-3) north of the front (p<0.001). There were no significant correlations between the selected physico-chemical (temperature and salinity) and the biological (mesozooplankton abundance and biomass) variables and the total abundance and biomass of the carnivorous zooplankton during the investigation (p>0.05 in all cases). There was no evidence of enhanced biomass and abundance values at stations occupied in the immediate vicinity of the front. Total average carnivorous zooplankton abundance was dominated by chaetognaths (Eukrohnia hamata Möbius 1875, Sagitta gazellae Ritler-Záhony 1909 and S. zetesios Fowler 1905) and euphausiids (Nematoscelis megalops Sars 1883, Euphausia longirostris Hansen 1908 and E. spinifera Sars 1883), which contributed up to 86.58 ± 32.91% of the total counts. The total average biomass was dominated by euphausiids and amphipods (Themisto gaudichaudii Guérin-Méneville 1825, Phronima sedentaria Forsskål 1775 and Vibilia armata Bovallius 1887) which contributed up to 71.45 ± 34.85% of the total counts. In general the populations of both the euphausiids and amphipods were dominated by females while the chaetognaths were dominated by juveniles. Numerical analysis identified two major zooplankton groupings within the survey area which did not coincide with the water masses within the survey area. The SIMPER procedure of the PRIMER package indicated differences between the groups were mainly attributed to changes in the abundance of the numerically dominant species rather than the presence or absence of individual species. The absence of any significant spatial patterns in the distribution of the carnivorous zooplankton suggests that the STC did not act as a biogeographical barrier during the present study. The mean feeding rates of the chaetognaths E. hamata, S. gazellae and S. zetesios were 1.82 ± 0.85prey d-1, 3.63 ± 2.08prey d-1 and 2.18 ± 0.59prey d-1, respectively. These rates correspond to a combined predation impact equivalent to <5% of the mesozooplankton standing stock or <10% of the mesozooplankton secondary production. Mesozooplankton, comprising mainly copepods was the dominant prey in the guts of the three chaetognath species. Total predation impact of the euphausiids, chaetognaths and amphipods, estimated using published daily ration data, on the mesozooplankton standing stock and secondary production ranged from 0.01% to 1.53% and from 0.03% to 30.54%, respectively. Among the carnivorous zooplankton, chaetognaths were generally identified as the dominant predators of mesozooplankton. Low predation impact of selected carnivorous zooplankton suggested that these organisms contributed little to the vertical carbon flux within the region of investigation during the study.

Thesis (Ph. D.)--Joint Program in Oceanography/Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Dept. of Biology; and the Woods Hole Oceanographic Institution), 2006.
Includes bibliographical references (p. 297-311).
The physical and biological forces that drive zooplankton distribution and patchiness in an antarctic continental shelf region were examined, with particular emphasis on the Antarctic krill, Euphausia superba. This was accomplished by the application of acoustic, video, and environmental sensors during surveys of the region in and around Marguerite Bay, west of the Antarctic Peninsula, in the falls and winters of 2001 and 2002. An important component of the research involved the development and verification of methods for extracting estimates of ecologically-meaningful quantities from measurements of scattered sound. The distribution of acoustic volume backscattering at the single frequency of 120 kHz was first examined as an index of the overall biomass of zooplankton. Distinct spatial and seasonal patterns were observed that coincided with advective features. Improved parameterization was then achieved for a theoretical model of Antarctic krill target strength, the quantity necessary in scaling measurements of scattered sound to estimates of abundance, through direct measurement of all necessary model parameters for krill sampled in the study region and survey period.
(cont.) Methods were developed for identifying and delineating krill aggregations, allowing the distribution of krill to be distinguished from that of the overall zooplankton community. Additional methods were developed and verified for estimating the length, abundance, and biomass of krill in each acoustically-identified aggregation. These methods were applied to multi-frequency acoustic survey data, demonstrating strong seasonal, inter-annual, and spatial variability in the distribution of krill biomass. Highest biomass was consistently associated with regions close to land where temperatures at depth were cool. Finally, the morphology, internal structure, and vertical position of individual krill aggregations were examined. The observed patterns of variability in aggregation characteristics between day and night, regions of high versus low food availability, and in the presence or absence of predators, together reinforced the conclusion that aggregation and diel vertical migration represent strategies to avoid visual predators, while also allowing the krill access to shallowly-distributed food resources. The various findings of this work have important implications to the fields of zooplankton acoustics and Antarctic krill ecology, especially in relation to the interactions of the krill with its predators.
by Gareth L. Lawson.
Ph.D.