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Interpreting the vulnerability of pelagic calcifiers to ocean acidification (OA) is enhanced by an understanding of their critical thresholds and how these thresholds are modified by other climate change stressors (e.g., warming). To address this need, we undertook a three-part data synthesis for pteropods, one of the calcifying zooplankton group. We conducted the first meta-analysis and threshold analysis of literature characterizing pteropod responses to OA and warming by synthetizing dataset comprising of 2,097 datapoints. Meta-analysis revealed the extent to which responses among studies conducted on differing life stages and disparate geographies could be integrated into a common analysis. The results demonstrated reduced calcification, growth, development, and survival to OA with increased magnitude of sensitivity in the early life stages, under prolonged duration, and with the concurrent exposure of OA and warming, but not species-specific sensitivity. Second, breakpoint analyses identified OA thresholds for several endpoints: dissolution (mild and severe), calcification, egg development, shell growth, and survival. Finally, consensus by a panel of pteropod experts was used to verify thresholds and assign confidence scores for five endpoints with a sufficient signal: noise ratio to develop life-stage specific, duration-dependent thresholds. The range of aragonite saturation state from 1.5-0.9 provides a risk range from early warning to lethal impacts, thus providing a rigorous basis for vulnerability assessments to guide climate change management responses, including an evaluation of the efficacy of local pollution management. In addition, meta-analyses with OA, and warming shows increased vulnerability in two pteropod processes, i.e., shell dissolution and survival, and thus pointing toward increased threshold sensitivity under combined stressor effect.

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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As part of the Canadian contribution to the International Polar Year (IPY), several major international research programs have focused on offshore arctic marine ecosystems. The general goal of these projects was to improve our understanding of how the response of arctic marine ecosystems to climate warming will alter food web structure and ecosystem services provided to Northerners. At least four key findings from these projects relating to arctic heterotrophic food web, pelagic-benthic coupling and biodiversity have emerged: (1) Contrary to a long-standing paradigm of dormant ecosystems during the long arctic winter, major food web components showed relatively high level of winter activity, well before the spring release of ice algae and subsequent phytoplankton bloom. Such phenological plasticity among key secondary producers like zooplankton may thus narrow the risks of Climatic Change

Abstract. In oligotrophic tropical and subtropical oceans, where strong stratification can limit the replenishment of surface nitrate, dinitrogen (N 2 ) fixation by diazotrophs can represent a significant source of nitrogen (N) for primary production. The VAHINE (VAriability of vertical and tropHIc transfer of fixed N 2 in the south-wEst Pacific) experiment was designed to examine the fate of diazotroph-derived nitrogen (DDN) in such ecosystems. In austral summer 2013, three large (∼ 50 m 3 ) in situ mesocosms were deployed for 23 days in the New Caledonia lagoon, an ecosystem that typifies the low-nutrient, low-chlorophyll environment, to stimulate diazotroph production. The zooplankton component of the study aimed to measure the incorporation of DDN into zooplankton biomass, and assess the role of direct diazotroph grazing by zooplankton as a DDN uptake pathway. Inside the mesocosms, the diatom-diazotroph association (DDA) het-1 predominated during days 5-15 while the unicellular diazotrophic cyanobacteria UCYN-C predominated during days 15-23. A Trichodesmium bloom was observed in the lagoon (outside the mesocosms) towards the end of the experiment. The zooplankton community was dominated by copepods (63 % of total abundance) for the duration of the experiment. Using two-source N isotope mixing models we estimated a mean ∼ 28 % contribution of DDN to zooplankton nitrogen biomass at the start of the experiment, indicating that the natural summer peak of N 2 fixation in the lagoon was already contributing significantly to the zooplankton. Stimulation of N 2 fixation in the mesocosms corresponded with a generally low-level enhancement of DDN contribution to zooplankton nitrogen biomass, but with a peak of ∼ 73 % in mesocosm 1 following the UCYN-C bloom. qPCR analysis targeting four of the common diazotroph groups present in the mesocosms (Trichodesmium, het-1, het-2, UCYN-C) demonstrated that all four were ingested by copepod grazers, and that their abundance in copepod stomachs generally corresponded with their in situ abundance. 15 N 2 labelled grazing experiments therefore provided evidence for direct ingestion and assimilation of UCYN-C-derived N by the zooplankton, but not for het-1 and Trichodesmium, supporting an important role of secondary pathways of DDN to the zooplankton for the latter groups, i.e. DDN contributions to the dissolved N pool and uptake by nondiazotrophs. This study appears to provide the first evidence of direct UCYN-C grazing by zooplankton, and indicates that UCYN-C-derived N contributesPublished by Copernicus Publications on behalf of the European Geosciences Union.

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