Pteropods In Southern Ocean Ecosystems

摘要

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 CO_2 emissions (lS92a), 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 floes, 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.