Pteropods in Southern Ocean ecosystems

To date, little research has been carried out on pelagic gastropod molluscs (pteropods) in Southern Ocean ecosystems. However, recent predictions are that, due to acidification resulting from a business as usual approach to CO₂ emissions (IS92a), Southern Ocean surface waters may begin to become uninhabitable for aragonite shelled thecosome pteropods by 2050. To gain insight into the potential impact that this would have on Southern Ocean ecosystems, we have here synthesized available data on pteropod distributions and densities, assessed current knowledge of pteropod ecology, and highlighted knowledge gaps and directions for future research on this zooplankton group.Six species of pteropod are typical of the Southern Ocean south of the Sub-Tropical Convergence, including the four Thecosomes Limacina helicina antarctica, Limacina retroversa australis, Clio pyramidata, and Clio piatkowskii, and two Gymnosomes Clione limacina antarctica and Spongiobranchaea australis. Limacina retroversa australis dominated pteropod densities north of the Polar Front (PF), averaging 60 ind m⁻³ (max = 800 ind m⁻³) and 11% of total zooplankton at the Prince Edward Islands. South of the PF L. helicina antarctica predominated, averaging 165 ind m⁻³ (max = 2681 ind m⁻³) and up to >35% of total zooplankton at South Georgia, and up to 1397 ind m⁻³ and 63% of total zooplankton in the Ross Sea. Combined pteropods contributed <5% to total zooplankton in the Lazarev Sea, but 15% (max = 93%) to macrozooplankton in the East Antarctic. In addition to regional density distributions we have synthesized data on vertical distributions, seasonal cycles, and inter-annual density variation.Trophically, gymnosome are specialist predators on thecosomes, while thecosomes are considered predominantly herbivorous, capturing food with a mucous web. The ingestion rates of L. retroversa australis are in the upper range for sub-Antarctic mesozooplankton (31.2-4196.9 ng pig ind⁻¹ d⁻¹), while those of L. helicina antarctica and C. pyramidata are in the upper range for all Southern Ocean zooplankton, in the latter species reaching 27,757 ng pig ind⁻¹ d⁻¹ and >40% of community grazing impact. Further research is required to quantify diet selectivity, the effect of phytoplankton composition on growth and reproductive success, and the role of carnivory in thecosomes.Life histories are a significant knowledge gap for Southern Ocean pteropods, a single study having been completed for L. retroversa australis, making population studies a priority for this group. Pteropods appear to be important in biogeochemical cycling, thecosome shells contributing >50% to carbonate flux in the deep ocean south of the PF. Pteropods may also contribute significantly to organic carbon flux through the production of fast sinking faecal pellets and mucous flocs, and rapid sinking of dead animals ballasted by their aragonite shells. Quantification of these contributions requires data on mucous web production rates, egestion rates, assimilation efficiencies, metabolic rates, and faecal pellet morphology for application to sediment trap studies.Based on the available data, pteropods are regionally significant components of the Southern Ocean pelagic ecosystem. However, there is an urgent need for focused research on this group in order to quantify how a decline in pteropod densities may impact on Southern Ocean ecosystems.     

Response of Nereis diversicolor population (Polychaeta,Nereididae) to the pollution impact – Authie and Seine estuaries (France)

A survey within the French National Programme of Ecotoxicology was carried out in 2002, 2003 and 2004 to study the response of Nereis diversicolor populations (Polychaeta, Nereididae) to the impact of pollution in the Authie estuary (non-contaminated site) and in the Seine estuary (contaminated site). In the period studied, the density varied from 672 ind. m⁻² to 3584 ind. m⁻² in the Authie estuary and from 80 ind. m⁻² to 920 ind. m⁻² in the Seine estuary. Biomass varied from 3.94 g m⁻² (dry weight) in February 2004 to 38.0 g m⁻² in August 2003 in the Authie estuary and from 3.4 g m⁻² in February 2002 to 0.6 g m⁻² in February 2004 in the Seine estuary. Density and biomass of the populations of N. diversicolor were consistently lower in the Seine estuary than in the Authie estuary. Size frequency histograms permit the analysis of the cohorts as well as the elaboration of the growth curves. For the individuals from the Authie estuary, the relation between dry weight (DW) and length L3 (prostomium, peristomium and chaetiger 1) was DW = 4.2205 L3^2.9832. For those from the Seine estuary, the relation between dry weight and L3 was DW = 0.4697e^1.7209L3. The individuals of N. diversicolor should belong to eight cohorts in Authie estuary (two cohorts each year) instead of six cohorts for those from the Seine estuary. These differences can be attributed to the effect of pollution on the population of N. diversicolor.     

Seasonal variation of horizontal material transport through the eastern channel of the Tsushima Straits

We conducted hydrographic observations ten times in the Tsushima Strait to reveal seasonal variations of horizontal material transports such as of heat, freshwater, chlorophyll a, and dissolved inorganic nitrogen (DIN) and phosphorus (DIP) through the eastern channel of the Tsushima Strait (ECTS). The volume, freshwater, and heat transport results are of nearly the same order as results reported in previous studies. The annual mean DIN and DIP transports of 3.59 kmol/s and 0.29 kmol/s are large relative to those of the Changjiang and the Taiwan Strait and are horizontally transported through the ECTS. Nutrient transports are high in July–August and October and low in April and November. Increased nutrient transports in July–August and October are due to the appearance of a cold saline water mass in the bottom layer of the ECTS. Changes in DIN transports in summer and autumn, which account for two-thirds of the total annual DIN transport, would have a large effect on the nitrogen budget and biological productivity in the Tsushima Warm Current region.     

Mixed layer depth climatology of the North Pacific based on Argo observations

A monthly mean climatology of the mixed layer depth (MLD) in the North Pacific has been produced by using Argo observations. The optimum method and parameter for evaluating the MLD from the Argo data are statistically determined. The MLD and its properties from each density profile were calculated with the method and parameter. The monthly mean climatology of the MLD is computed on a 2° × 2° grid with more than 30 profiles for each grid. Two bands of deep mixed layer with more than 200 m depth are found to the north and south of the Kuroshio Extension in the winter climatology, which cannot be reproduced in some previous climatologies. Early shoaling of the winter mixed layer between 20–30°N, which has been pointed out by previous studies, is also well recognized. A notable feature suggested by our climatology is that the deepest mixed layer tends to occur about one month before the mixed layer density peaks in the middle latitudes, especially in the western region, while they tend to coincide with each other in higher latitudes.     

Reevaluation of historical ocean heat content variations with time-varying XBT and MBT depth bias corrections

As reported in former studies, temperature observations obtained by expendable bathythermographs (XBTs) and mechanical bathythermographs (MBTs) appear to have positive biases as much as they affect major climate signals. These biases have not been fully taken into account in previous ocean temperature analyses, which have been widely used to detect global warming signals in the oceans. This report proposes a methodology for directly eliminating the biases from the XBT and MBT observations. In the case of XBT observation, assuming that the positive temperature biases mainly originate from greater depths given by conventional XBT fall-rate equations than the truth, a depth bias equation is constructed by fitting depth differences between XBT data and more accurate oceanographic observations to a linear equation of elapsed time. Such depth bias equations are introduced separately for each year and for each probe type. Uncertainty in the gradient of the linear equation is evaluated using a non-parametric test. The typical depth bias is +10 m at 700 m depth on average, which is probably caused by various indeterminable sources of error in the XBT observations as well as a lack of representativeness in the fall-rate equations adopted so far. Depth biases in MBT are fitted to quadratic equations of depth in a similar manner to the XBT method. Correcting the historical XBT and MBT depth biases by these equations allows a historical ocean temperature analysis to be conducted. In comparison with the previous temperature analysis, large differences are found in the present analysis as follows: the duration of large ocean heat content in the 1970s shortens dramatically, and recent ocean cooling becomes insignificant. The result is also in better agreement with tide gauge observations. On leave from the Meteorological Research Institute of the Japan Meteorological Agency.     

Observations of the North Equatorial Current,Mindanao Current,and Kuroshio current system during the 2006/07 El Niño and 2007/08 La Niña

Two onboard observation campaigns were carried out in the western boundary region of the Philippine Sea in December 2006 and January 2008 during the 2006/07 El Niño and the 2007/08 La Niña to observe the North Equatorial Current (NEC), Mindanao Current (MC), and Kuroshio current system. The NEC and MC measured in late 2006 under El Niño conditions were stronger than those measured during early 2008 under La Niña conditions. The opposite was true for the current speed of the Kuroshio, which was stronger in early 2008 than in late 2006. The increase in dynamic height around 8°N, 130°E from December 2006 to January 2008 resulted in a weakening of the NEC and MC. Local wind variability in this region did not appear to contribute to changes in the current system.     

Large bio-geographical shifts in the north-eastern Atlantic Ocean: From the subpolar gyre, via plankton, to blue whiting and pilot whales

Pronounced changes in fauna, extending from the English Channel in the south to the Barents Sea in the north-east and off Greenland in the north-west, have occurred in the late 1920s, the late 1960s and again in the late 1990s. We attribute these events to exchanges of subarctic and subtropical water masses in the north-eastern North Atlantic Ocean, associated with changes in the strength and extent of the subpolar gyre. These exchanges lead to variations in the influence exerted by the subarctic or Lusitanian biomes on the intermediate faunistic zone in the north-eastern Atlantic. This strong and persistent bottom-up bio-physical link is demonstrated using a numerical ocean general circulation model and data on four trophically connected levels in the food chain – phytoplankton, zooplankton, blue whiting, and pilot whales. The plankton data give a unique basin-scale depiction of these changes, and a long pilot whale record from the Faroe Islands offers an exceptional temporal perspective over three centuries. Recent advances in simulating the dynamics of the subpolar gyre suggests a potential for predicting the distribution of the main faunistic zones in the north-eastern Atlantic a few years into the future, which might facilitate a more rational management of the commercially important fisheries in this region.     

Composition of the phytoplankton pigments in the surface waters along a transect from the Northwestern to the southeastern Pacific Ocean

Multidisciplinary oceanic investigation was undertaken in Aug–Sep. 2003 along a transect from Northwestern (Busan, Korea) to Southeastern Pacific (Talcahuano, Chile) to understand the physical, chemical and biological features in the surface water, and to depict their interaction with the atmosphere. Among the twenty parameters measured, we describe the physical, chemical and biological features. Physico-chemical data were analyzed in conjunction with the geographic position and yielded 7 peculiar surface water masses. The first water mass (28.4°N, 130.8°E to 21.5°N, 139.5°E) was warm and low in phosphate and nitrate content, and high in silicate. The concentration of phytoplankton pigment was one of the lowest. The second (20.4°N, 140.7°E to 2.2°S, 162.9°E) was the warmest and the least saline. Nitrate and phosphate concentration were one of the lowest. Chlorophyll a (Chl a) concentration was the lowest among the surface waters. The third (3.4°S, 164.0°E to 14.5°S, 173.3°E) was warm. Nitrate concentration was the lowest. CHL-a, peridinin (Perid), violaxanthin (Viola), zeaxanthin (Zea), chlorophyll-b (Chl b) and β-CAR were abundant. The fourth (18.6°S, 177.5°E to 31.8°S, 123.9°W) was saline and poor in nutrient concentration. The contributions of 19′-butanoyloxyfucoxanthin (But-fuco), 19′-hexanoyloxyfucoxanthin (Hex-fuco), and CHL b to CHL a were non-negligible. The fifth (32.4°S, 122.1°W to 33.8°S, 117.2°W) was relatively cold and well oxygenated. Concentration of Fuco, But-fuco, Hex-fuco and Chl b was high. The sixth (34.2°S, 115.4°W to 37.4°S, 92.1°W) was cold, well oxygenated and enriched with phosphate and nitrate. Concentration of phytoplankton pigment was, however, one of the lowest. The seventh, located off the Chilean coast, from 37.2°S, 87.2°W to 36.1°S, 74.1°W was well oxygenated and highly enriched with nitrate and phosphate. Phytoplankton pigments such as Fuco, Perid, But-fico, and Hex-fuco were rich. The 7 surface water masses are partially attributed to Kuroshio Current, North Equatorial Current and North Equatorial Countercurrent, South Equatorial current, South Pacific Subtropical Gyre, South Pacific Current, Subtropical Front and Chilean coastal water. The differences in physicochemical characteristics and the history of the surface water resulted in difference in quantity and composition of the phytoplankton pigment.     

Long-term bottom water warming in the north Ross Sea

We measured potential temperature, salinity, and dissolved oxygen profiles from the surface to the bottom at two locations in the north Ross Sea (65.2°S, 174.2°E and 67.2°S, 172.7°W) in December 2004. Comparison of our data with previous results from the same region reveals an increase in potential temperature and decreases in salinity and dissolved oxygen concentration in the bottom layer (deeper than 3000 m) over the past four decades. The changes were significantly different from the analytical precisions. Detailed investigation of the temperature, salinity, dissolved oxygen and σ ₃ value distributions and the bottom water flow in the north Ross Sea suggests a long-term change in water mass mixing balance. That is to say, it is speculated that the influence of cool, saline, high-oxygen bottom water (high-salinity Ross Sea Bottom Water) formed in the southwestern Ross Sea has possibly been decreased, while the influences of relatively warmer and fresher bottom water (low-salinity Ross Sea Bottom Water) and the Adélie Land Bottom Water coming from the Australia-Antarctic Basin have increased. The possible impact of global warming on ocean circulation needs much more investigation.     

Possibility of recent changes in vertical distribution and size composition of chlorophyll-a in the western North Pacific region

Our analysis of the last three decades of retrospective data of vertical distributions and size composition of chlorophyll-a (Chl-a) over the western North Pacific has revealed significant changes of three indices related to Chl-a during summer season, as follows: (1) decreasing linear trend of the proportion of Chl-a in surface layer to that of the whole water column by 0.4 and 2.3% year⁻¹ in the subtropical area along 137°E (STA₁₃₇) during 1972 to 1997 and in the Kuroshio Extension area along 175°E (KEA₁₇₅) during 1990 to 2001; (2) increasing linear trend of the depth of subsurface Chl-a maximum (DCM) by 0.4 and 2.6 m year⁻¹ in STA₁₃₇ and KEA₁₇₅; and (3) decreasing linear trend of larger-size Chl-a (>3 μm) by 0.1 and 2.5% year⁻¹ in STA₁₃₇ and KEA₁₇₅, respectively. Water density (σ _θ ) at 75 m depth had also decreased by 0.006 and 0.05 year⁻¹ in STA₁₃₇ and KEA₁₇₅, respectively. The ratio of biogenic opal to biogenic CaCO₃ in the sinking flux decreased by 0.015 year⁻¹ in the subtropical region from 1997 to 2005. These findings may indicate that the subsurface chlorophyll maximum is deepening and larger phytoplankton such as diatoms has been decreasing during the past decade, associated with the decreasing density of surface water caused by warming in the western North Pacific, especially in the summer.