South Georgia, Antarctica: a productive, cold water, pelagic ecosystem

The South Georgia region is characterised by high biomass and productivity of phytoplankton, zooplankton and vertebrate predators. Important commercial fisheries have been based at the island since the late 1700s, initially exploiting seals and whales, and currently taking krill Euphausia superba and finfish. Despite studies dating from the beginning of the last century, the causes of the high productivity remain unclear. The island lies within the Antarctic Zone of the Antarctic Circumpolar Current, to the south of the Polar Front. The offshore waters to its north and east are affected by a northwards deflection of the Southern Antarctic Circumpolar Current Front, together with waters from the Weddell-Scotia Confluence. Despite a retentive circulation over the shelf, the high productivity of phytoplankton and copepods is widespread, occurring far downstream and possibly extending to the Polar Front. High phytoplankton concentrations (>20 mg chlorophyll a m -3 ) may be linked to enhanced supply of iron or reduced forms of nitrogen (up to ~3 mmol ammonium m -3 ). Although macronutrients are generally not limiting in the Antarctic Zone, silicate concentrations of 2 mg chl a m -3 occur for 4 to 5 mo. Biomass of krill plus net-caught zooplankton in summer is around 15 to 20 g dry mass m -2 , equally dominated by krill and copepods. This greatly exceeds typical values for Antarctica, and is high compared to productive northern shelves. Zooplankton, and in particular krill, appear to have a pivotal role in regulating energy flow in this food web, through selective grazing and possibly also through nutrient regeneration. Abundances of krill and copepods are negatively related across a wide range of scales, suggesting direct interaction through competition or predation. Evidence suggests that when phytoplankton stocks are low, energy flow through krill is maintained by their feeding on the large populations of small copepods. Metazoans and higher predators at South Georgia can feed across several trophic levels according to prey abundance, and they regenerate substantial quantities of reduced nitrogen. Therefore we suggest that these groups have a controlling influence on lower trophic levels, both stabilising population sizes and maintaining high rates of energy flow. Hydrography, nutrient concentrations, phytoplankton, copepod, and krill biomasses fluctuate between years. Periodically (once or twice a decade), shortages of krill in summer result in breeding failures among many of their predators. This appears to be a downstream effect from wider scale, Scotia Sea phenomena, although the processes involved are unclear. The elevated biomass and energy flows at South Georgia appear to be caused by locally enhanced growth rates; there is no evidence so far for any physical concentration mechanism. Even for krill, which do not breed there, local growth rates are probably of a similar order to the biomass removed by their main land-based predators in summer. Thus the transfer of energy to higher predators depends on local enhancement of fluxes through the food web as well as the supply of plankton to the island by the Antarctic Circumpolar Current.