Effects on zooplankton of a warmer ocean: Recent evidence from the Northeast Pacific

The consequences for pelagic communities of warming trends in mid and high latitude ocean regions could be substantial, but their magnitude and trajectory are not yet known. Environmental changes predicted by climate models (and beginning to be confirmed by observations) include warming and freshening of the upper ocean and reduction in the extent and duration of ice cover. One way to evaluate response scenarios is by comparing how "similar" zooplankton communities have differed among years and/or locations with differing temperature. The subarctic Pacific is a strong candidate for such comparisons, because the same mix of zooplankton species dominates over a wide range of temperature climatologies, and observations have spanned substantial temperature variability at interannual-to-decadal time scales. In this paper, we review and extend copepod abundance and phenology time series from net tow and Continuous Plankton Recorder surveys in the subarctic Northeast Pacific. The two strongest responses we have observed are latitudinal shifts in centers of abundance of many species (poleward under warm conditions), and changes in the life cycle timing of Neocalanus plumchrus in both oceanic and coastal regions (earlier by several weeks in warm years and at warmer locations). These zooplankton data, plus indices of higher trophic level responses such as reproduction, growth and survival of pelagic fish and seabirds, are all moderately-to-strongly intercorrelated (vertical bar r vertical bar = 0.25-0.8) with indices of local and basin-scale temperature anomalies. A principal components analysis of the normalized anomaly time series from 1979 to 2004 shows that a single "warm-and-low-productivity" vs. "cool-and-high-productivity" component axis accounts for over half of the variance/covariance. Prior to 1990, the scores for this component were negative ("cool" and "productive") or near zero except positive in the El Nino years 1983 and 1987. The scores were strongly and increasingly positive ("warm" and "low productivity") from 1992 to 1998; negative from 1999 to 2002; and again increasingly positive from 2003-present. We suggest that, in strongly seasonal environments, anomalously high temperature may provide misleading environmental cues that contribute to timing mismatch between life history events and the more-nearly-fixed seasonality of insolation, stratification, and food supply. Crown Copyright (c) 2007 Published by Elsevier Ltd. All rights reserved.

Challenges for implementing the Marine Strategy Framework Directive in a climate of macroecological change

Unprecedented basin-scale ecological changes are occurring in our seas. As temperature and carbon dioxide concentrations increase, the extent of sea ice is decreasing, stratification and nutrient regimes are changing and pH is decreasing. These unparalleled changes present new challenges for managing our seas, as we are only just beginning to understand the ecological manifestations of these climate alterations. The Marine Strategy Framework Directive requires all European Member States to achieve good environmental status (GES) in their seas by 2020; this means management towards GES will take place against a background of climate-driven macroecological change. Each Member State must set environmental targets to achieve GES; however, in order to do so, an understanding of large-scale ecological change in the marine ecosystem is necessary. Much of our knowledge of macroecological change in the North Atlantic is a result of research using data gathered by the Continuous Plankton Recorder (CPR) survey, a near-surface plankton monitoring programme that has been sampling in the North Atlantic since 1931. CPR data indicate that North Atlantic and North Sea plankton dynamics are responding to both climate and human-induced changes, presenting challenges to the development of pelagic targets for achievement of GES in European Seas. Thus, the continuation of long-term ecological time series such as the CPR survey is crucial for informing and supporting the sustainable management of European seas through policy mechanisms.

Why is it so hard to set MSFD indicators and targets? Messages from the plankton

Unprecedented basin-scale ecological changes are occurring in our seas. As temperature and carbon dioxide concentrations increase, the extent of sea ice is decreasing, stratification and nutrient regimes are changing, and pH is decreasing. These unparalleled changes present new challenges for managing our seas as we are only just beginning to understand the ecological manifestations of these climate alterations. The Marine Strategy Framework Directive requires all European Member States to achieve Good Environmental Status (GES) in their seas by 2020; this means management toward GES will take place against a background of climate-driven macroecological change. Each Member State must set environmental targets to achieve GES; however, in order to do so an understanding of large-scale ecological change in the marine ecosystem is necessary. Much of our knowledge of macroecological change in the North Atlantic is a result of research using data gathered by the Continuous Plankton Recorder (CPR) survey, a near-surface plankton monitoring program which has been sampling in the North Atlantic since 1931. CPR data indicate that North Atlantic and North Sea plankton dynamics are responding to both climate and human-induced changes, presenting challenges to the development of pelagic targets for achievement of GES in European seas. Thus the continuation of long-term ecological time-series such as the CPR is crucial for informing and supporting the sustainable management of European seas through policy mechanisms.

Impacts of the Oceans on Climate Change

The oceans play a key role in climate regulation especially in part buffering (neutralising) the effects of increasing levels of greenhouse gases in the atmosphere and rising global temperatures. This chapter examines how the regulatory processes performed by the oceans alter as a response to climate change and assesses the extent to which positive feedbacks from the ocean may exacerbate climate change. There is clear evidence for rapid change in the oceans. As the main heat store for the world there has been an accelerating change in sea temperatures over the last few decades, which has contributed to rising sea‐level. The oceans are also the main store of carbon dioxide (CO2), and are estimated to have taken up ∼40% of anthropogenic-sourced CO2 from the atmosphere since the beginning of the industrial revolution. A proportion of the carbon uptake is exported via the four ocean ‘carbon pumps’ (Solubility, Biological, Continental Shelf and Carbonate Counter) to the deep ocean reservoir. Increases in sea temperature and changing planktonic systems and ocean currents may lead to a reduction in the uptake of CO2 by the ocean; some evidence suggests a suppression of parts of the marine carbon sink is already underway. While the oceans have buffered climate change through the uptake of CO2 produced by fossil fuel burning this has already had an impact on ocean chemistry through ocean acidification and will continue to do so. Feedbacks to climate change from acidification may result from expected impacts on marine organisms (especially corals and calcareous plankton), ecosystems and biogeochemical cycles. The polar regions of the world are showing the most rapid responses to climate change. As a result of a strong ice–ocean influence, small changes in temperature, salinity and ice cover may trigger large and sudden changes in regional climate with potential downstream feedbacks to the climate of the rest of the world. A warming Arctic Ocean may lead to further releases of the potent greenhouse gas methane from hydrates and permafrost. The Southern Ocean plays a critical role in driving, modifying and regulating global climate change via the carbon cycle and through its impact on adjacent Antarctica. The Antarctic Peninsula has shown some of the most rapid rises in atmospheric and oceanic temperature in the world, with an associated retreat of the majority of glaciers. Parts of the West Antarctic ice sheet are deflating rapidly, very likely due to a change in the flux of oceanic heat to the undersides of the floating ice shelves. The final section on modelling feedbacks from the ocean to climate change identifies limitations and priorities for model development and associated observations. Considering the importance of the oceans to climate change and our limited understanding of climate-related ocean processes, our ability to measure the changes that are taking place are conspicuously inadequate. The chapter highlights the need for a comprehensive, adequately funded and globally extensive ocean observing system to be implemented and sustained as a high priority. Unless feedbacks from the oceans to climate change are adequately included in climate change models, it is possible that the mitigation actions needed to stabilise CO2 and limit temperature rise over the next century will be underestimated.

A biological consequence of reducing Arctic ice cover: arrival of the Pacific diatom Neodenticula seminae in the North Atlantic for the first time in 800,000 years

The Continuous Plankton Recorder survey has monitored plankton in the Northwest Atlantic at monthly intervals since 1962, with an interegnum between 1978 and 1990. In May 1999, large numbers of the Pacific diatom Neodenticula seminae were found in Continuous Plankton Recorder (CPR) samples in the Labrador Sea as the first record in the North Atlantic for more than 800 000 years. The event coincided with modifications in Arctic hydrography and circulation, increased flows of Pacific water into the Northwest Atlantic and in the previous year the exceptional occurrence of extensive ice-free water to the North of Canada. These observations indicate that N. seminae was carried in a pulse of Pacific water in 1998/early 1999 via the Canadian Arctic Archipelago and/or Fram Strait. The species occurred previously in the North Atlantic during the Pleistocene from similar to 1.2 to similar to 0.8 Ma as recorded in deep sea sediment cores. The reappearance of N. seminae in the North Atlantic is an indicator of the scale and speed of changes that are taking place in the Arctic and North Atlantic oceans as a consequence of regional climate warming. Because of the unusual nature of the event it appears that a threshold has been passed, marking a change in the circulation between the North Pacific and North Atlantic Oceans via the Arctic. Trans-Arctic migrations from the Pacific into the Atlantic are likely to occur increasingly over the next 100 years as Arctic ice continues to melt affecting Atlantic biodiversity and the biological pump with consequent feedbacks to the carbon cycle.

Feeding and overwintering of Antarctic krill across its major habitats: The role of sea ice cover, water depth, and phytoplankton abundance

Antarctic krill (Euphausia superba) were sampled in contrasting habitats: a seasonally ice-covered deep ocean (Lazarev Sea), ice-free shelves at their northern range (South Georgia) and the Antarctic Peninsula (Bransfield Strait), and shelf and oceanic sites in the Scotia Sea. Across 92 stations, representing a year-round average, the food volume in krill stomachs comprised 71 +/- 29% algae, 17 +/- 21% protozoans, and 12 +/- 25% metazoans. Fatty acid trophic markers showed that copepods were consistently part of krill diet, not a switch food. In open waters, both diatom and copepod consumption increased with phytoplankton abundance. Under sea ice, ingestion of diatoms became rare, whereas feeding on copepods remained constant. During winter, larvae contained high but variable proportions of diatom markers, whereas in postlarvae the role of copepods increased with krill body length. Overwintering differed according to habitat. Krill from South Georgia had lower lipid stores than those from the Bransfield Strait or Lazarev Sea. Feeding effort was much reduced in Lazarev Sea krill, whereas most individuals from the Bransfield Strait and South Georgia contained phytoplankton and seabed detritus in their stomachs. Their retention of essential body reserves indicates that krill experienced most winter hardship in the Lazarev Sea, followed by South Georgia and then Bransfield Strait. This was reflected in the delayed development from juveniles to adults in the Lazarev Sea. Circumpolar comparisons of length frequencies suggest that krill growth conditions are more favorable in the southwest Atlantic than in the Lazarev Sea or off East Antarctica because of longer phytoplankton bloom periods and rewarding access to benthic food.

Late winter under ice pelagic microbial communities in the high Arctic Ocean and the impact of short-term exposure to elevated CO2 levels

Polar Oceans are natural CO2 sinks because of the enhanced solubility of CO2 in cold water. The Arctic Ocean is at additional risk of accelerated ocean acidification (OA) because of freshwater inputs from sea ice and rivers, which influence the carbonate system. Winter conditions in the Arctic are of interest because of both cold temperatures and limited CO2 venting to the atmosphere when sea ice is present. Earlier OA experiments on Arctic microbial communities conducted in the absence of ice cover, hinted at shifts in taxa dominance and diversity under lowered pH. The Catlin Arctic Survey provided an opportunity to conduct in situ, under-ice, OA experiments during late Arctic winter. Seawater was collected from under the sea ice off Ellef Ringnes Island, and communities were exposed to three CO2 levels for 6 days. Phylogenetic diversity was greater in the attached fraction compared to the free-living fraction in situ, in the controls and in the treatments. The dominant taxa in all cases were Gammaproteobacteria but acidification had little effect compared to the effects of containment. Phylogenetic net relatedness indices suggested that acidification may have decreased the diversity within some bacterial orders, but overall there was no clear trend. Within the experimental communities, alkalinity best explained the variance among samples and replicates, suggesting subtle changes in the carbonate system need to be considered in such experiments. We conclude that under ice communities have the capacity to respond either by selection or phenotypic plasticity to heightened CO2 levels over the short term.

The potential use of mapped extent and distribution of habitats as indicators of Good Environmental Status (GES)

The Healthy and Biologically Diverse Seas Evidence Group (HBDSEG) has been tasked with providing the technical advice for the implementation of the Marine Strategy Framework Directive (MSFD) with respect to descriptors linked to biodiversity. A workshop was held in London to address one of the Research and Development (R&D) proposals entitled: ‘Mapping the extent and distribution of habitats using acoustic and remote techniques, relevant to indicators for area/extent/habitat loss.’ The aim of the workshop was to identify, define and assess the feasibility of potential indicators of benthic habitat distribution and extent, and identify the R&D work which could be required to fully develop these indicators. The main points that came out of the workshop were: (i) There are many technical aspects of marine habitat mapping that still need to be resolved if cost-effective spatial indicators are to be developed. Many of the technical aspects that need addressing surround issues of consistency, confidence and repeatability. These areas should be tackled by the JNCC Habitat Mapping and Classification Working Group and the HBDSEG Seabed Mapping Working Group. (ii) There is a need for benthic ecologists (through the HBDSEG Benthic Habitats Subgroup and the JNCC Marine Indicators Group) to finalise the list of habitats for which extent and/or distribution indicators should be considered for development, building upon the recommendations from this report. When reviewing the list of indicators, benthic habitats could also be distinguished into those habitats that are defined/determined primarily by physical parameters (although including biological assemblages) (e.g. subtidal shallow sand) and those defined primarily by their biological assemblage (e.g. seagrass beds). This distinction is important as some anthropogenic pressures may influence the biological component of the ecosystem despite not having a quantifiable effect on the physical habitat distribution/extent. (iii) The scale and variety of UK benthic habitats makes any attempt to undertake comprehensive direct mapping exercises prohibitively expensive (especially where there is a need for repeat surveys for assessment). There is a clear need therefore to develop a risk-based approach that uses indirect indicators (e.g. modelling), such as habitats at risk from pressures caused by current human activities, to develop priorities for information gathering. The next steps that came out of the workshop were: (i) A combined approach should be developed by the JNCC Marine Indicators Group together with the HBDSEG Benthic Habitats Subgroup, which will compile and ultimately synthesise all the criteria used by the three different groups from the workshop. The agreed combined approach will be used to undertake a final review of the habitats considered during the workshop, and to evaluate any remaining habitats in order to produce a list of habitats for indicator development for which extent and/or distribution indicators could be appropriate. (ii) The points of advice raised at this workshop, alongside the combined approach aforementioned, and the final list of habitats for extent and/or distribution indicator development will be used to develop a prioritised list of actions to inform the next round of R&D proposals for benthic habitat indicator development in 2014. This will be done through technical discussions within JNCC and the relevant HBDSEG Subgroups. The preparation of recommendations by these groups should take into account existing work programmes, and consider the limited resources available to undertake any further R&D work.

Ocean-wide tracking of pelagic sharks reveals extent of overlap with longline fishing hotspots

Overfishing is arguably the greatest ecological threat facing the oceans, yet catches of many highly migratory fishes including oceanic sharks remain largely unregulated with poor monitoring and data reporting. Oceanic shark conservation is hampered by basic knowledge gaps about where sharks aggregate across population ranges and precisely where they overlap with fishers. Using satellite tracking data from six shark species across the North Atlantic, we show that pelagic sharks occupy predictable habitat ‘hotspots’ of high space use. Movement modelling showed sharks preferred habitats characterised by strong sea-surface-temperature gradients (fronts) over other available habitats. However, simultaneous Global Positioning System (GPS) tracking of the entire Spanish and Portuguese longline-vessel fishing fleets show an 80% overlap of fished areas with hotspots, potentially increasing shark susceptibility to fishing exploitation. Regions of high overlap between oceanic tagged sharks and longliners included the North Atlantic Current/Labrador Current convergence zone and the Mid-Atlantic Ridge south-west of the Azores. In these main regions, and sub-areas within them, shark/vessel co-occurrence was spatially and temporally persistent between years, highlighting how broadly the fishing exploitation efficiently ‘tracks’ oceanic sharks within their space-use hotspots year-round. Given this intense focus of longliners on shark hotspots our study argues the need for international catch limits for pelagic sharks and identifies a future role of combining fine-scale fish and vessel telemetry to inform the ocean-scale management of fisheries.

Progress in satellite remote sensing for studying physical processes at the ocean surface and its borders with the atmosphere and sea-ice

Physical oceanography is the study of physical conditions, processes and variables within the ocean, including temperature-salinity distributions, mixing of the water column, waves, tides, currents, and air-sea interaction processes. Here we provide a critical review of how satellite sensors are being used to study physical oceanography processes at the ocean surface and its borders with the atmosphere and sea-ice. The paper begins by describing the main sensor types that are used to observe the oceans (visible, thermal infrared and microwave) and the specific observations that each of these sensor types can provide. We then present a critical review of how these sensors and observations are being used to study i) ocean surface currents, ii) storm surges, iii) sea-ice, iv) atmosphere-ocean gas exchange and v) surface heat fluxes via phytoplankton. Exciting advances include the use of multiple sensors in synergy to observe temporally varying Arctic sea-ice volume, atmosphere- ocean gas fluxes, and the potential for 4 dimensional water circulation observations. For each of these applications we explain their relevance to society, review recent advances and capability, and provide a forward look at future prospects and opportunities. We then more generally discuss future opportunities for oceanography-focussed remote-sensing, which includes the unique European Union Copernicus programme, the potential of the International Space Station and commercial miniature satellites. The increasing availability of global satellite remote-sensing observations means that we are now entering an exciting period for oceanography. The easy access to these high quality data and the continued development of novel platforms is likely to drive further advances in remote sensing of the ocean and atmospheric systems.