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We investigated physiological parameters (elemental and biochemical composition, metabolic rates, feeding activity and growth) of adult Antarctic krill in the Lazarev Sea in late spring (December), mid autumn (April) and mid winter (July and August) to evaluate proposed hypotheses of overwintering mechanisms. Our major observations are: (1) respiration rates were reduced by 30 to 50% in autumn and winter, compared to values in late spring; (2) feeding activity was reduced by 80 to 86% in autumn and winter, compared to late spring, at similar food concentrations; (3) feeding was omnivorous during winter; (4) with each moult, krill grew by 0.5 to 3.8% in length; (5) body lipids and, to a small extent, body proteins were consumed during winter. Adult Euphausia superba thus adopt metabolic slowdown and omnivorous feeding activity at low rates to survive the winter season in the Lazarev Sea. By mid autumn, metabolic activity is reduced, most likely being influenced by the Antarctic light regime, which is accompanied by a reduction in feeding activity and growth. Although at a low level, the feeding activity during winter seems to provide an important energy input.

A ntarctic krill Euphausia superba, a key species in Southern Ocean food webs 1 , plays a central role in ecosystem processes and community dynamics of apex predators, and is the target of a commercial fishery 2 . Krill spawn in late spring and their larvae develop during summer, autumn and under the ice in winter to emerge as juveniles in the following spring. The newly spawned eggs sink from the surface to up to 1,000 m depth where they hatch and the developing larvae actively swim upwards to feed in the upper water column 1 . Larval Antarctic krill have low lipid reserves, which are insufficient to support long periods of starvation. Therefore, winter, when primary production is minimal, is assumed to be a critical bottleneck for larval krill development and hence recruitment to the adult population 3,4 . Present hypotheses suggest that high algal biomass in winter sea ice enhances larval krill winter-feeding conditions and growth [5][6][7][8] . This implies that larvae have access to this high algal biomass within the sea ice. However, recent observations 9,10 indicate that the linkage between sea ice and krill recruitment success is not as direct as has been suggested. The timing of ice-edge advance and annual ice-season duration is highly variable, and does not necessarily show a clear link to krill recruitment in the following year ( Supplementary Figs. 1 and 2). Along the Antarctic Peninsula, adult krill have a five to six year population cycle with oscillations in biomass exceeding an order of magnitude 9 . According to bioenergetics models, part of the variability is due to interannual variation in reproductive output 11 as well as autumn blooms that may govern the possible overwinter survival rate of larvae 11,12 . In the Bransfield Strait, three krill winter surveys have shown that krill abundance is an order of magnitude higher than in summer, regardless of concurrent sea-ice conditions 10 A dominant Antarctic ecological paradigm suggests that winter sea ice is generally the main feeding ground for krill larvae. Observations from our winter cruise to the southwest Atlantic sector of the Southern Ocean contradict this view and present the first evidence that the pack-ice zone is a food-poor habitat for larval development. In contrast, the more open marginal ice zone provides a more favourable food environment for high larval krill growth rates. We found that complex under-ice habitats are, however, vital for larval krill when water column productivity is limited by light, by providing structures that offer protection from predators and to collect organic material released from the ice. The larvae feed on this sparse ice-associated food during the day. After sunset, they migrate into the water below the ice (upper 20 m) and drift away from the ice areas where they have previously fed. Model analyses indicate that this behaviour increases both food uptake in a patchy food environment and the likelihood of overwinter transport to areas where feeding conditions are more favourable in spring.

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