Global synchrony of an accelerating risein sea surface temperature

The oceans have shown a recent rapid and accelerating rise in temperature with, given the close link between temperature and marine organisms, pronounced effects on ecosystems. Here we describe for the first time a globally synchronous pattern of pulsed short period (�1 year long) emanations of warm sea surface temperature anomalies from tropical seas towards the poles on the shelf/slope with an intensification of the warming after the 1976/1977, 1986/1987 and 1997/1998 El Nin˜os. On the eastern margins of continents the anomalies propagate towards the poles in part by largely baroclinic boundary currents, reinforced by regional atmospheric warming. The processes contributing to the less continuous warm anomalies on western margins are linked to the transfer of warmth from adjacent western boundary currents. These climate induced events show a close parallelism with the timing of ecosystem changes in shelf seas, important for fisheries and ecosystem services, and melting of sea-ice.

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.

Past, present and future nutrient loads of the North Sea: causes and consequences

Comprehensive, aggregate nutrient budgets were established for two compartments of the North Sea, the shallow coastal and deeper open regions, and for three different periods, representing pre-eutrophication (∼1950), eutrophication (∼1990) and contemporary (∼2000) phases. The aim was to quantify the major budget components, to identify their sources of variability, to specify the anthropogenic components, and to draw implications for past and future policy. For all three periods, open North Sea budgets were dominated (75%) by fluxes from and to the North-East Atlantic; sediment exchange was of secondary importance (18%). For the coastal North Sea, fluxes during the eutrophication period were dominated by sediment exchange (49% of all inputs), followed by exchange with the open sea (21%), and anthropogenic inputs (19%). Between 1950 and 1990, N-loading of coastal waters increased by a factor of 1.62 and P-loading by 1.45. These loads declined after 1990. Interannual variability in Atlantic inflow was found to correspond to a variability of 11% in nutrient load to the open North Sea. Area-specific external loads of both the open and coastal North Sea were below Vollenweider-type critical loads when expressed relative to depth and flushing. External area-specific load of the coastal North Sea has declined since 1990 from 1.8 to about 1.4 g P m−2 y−1 in 2000, which is close to the estimate of 1.3 for 1950. N-load declined less, leading to an increase in N/P ratio.

Black-legged kittiwakes as indicators of environmental change in the North Sea; evidence from long-term studies

Top predators, particularly seabirds, have repeatedly been suggested as indicators of marine ecosystem status. One region currently under pressure from human fisheries and climate change is the North Sea. Standardized seabird monitoring data have been collected on the Isle of May, an important seabird colony in the northwestern North Sea, over the last 10–20 years. Over this period oceanographic conditions have varied markedly, and between 1990 and 1999 a major industrial fishery for sandlance (Ammodytes marinus), the main prey of most seabird species, was prosecuted nearby. Sandlance fishing grounds close to seabird colonies down the east coast of the UK were closed in 2000 in an attempt to improve foraging opportunities for breeding seabirds, particularly black-legged kittiwakes (Rissa tridactyla). Initially this closure seemed to be beneficial for kittiwakes with breeding success recovering to pre-fishery levels. However, despite the ban continuing, kittiwakes and many other seabird species in the North Sea suffered severe breeding failures in 2004. In this paper, we test the predictive power of four previously established correlations between kittiwake breeding success and climatic/trophic variables to explain the observed breeding success at the Isle of May in 2004. During the breeding season, kittiwakes at this colony switch from feeding on 1+ group to 0 group sandlance, and results up until 2003 indicated that availability of both age classes had a positive effect on kittiwake breeding success. The low breeding success of kittiwakes in 2004 was consistent with the late appearance and small body size of 0 group sandlance, but at odds with the two variables likely to operate via 1 group availability (lagged winter sea surface temperature and larval sandlance cohort strength in 2003). The reason for the discrepancy is currently unknown, but analysis of 1 group sandlance body composition indicated that lipid content in 2004 was extremely low, and thus fish eaten by kittiwakes during pre-breeding and early incubation were likely to be of poor quality. Monitoring of reproductive success of kittiwakes, although useful, was clearly not sufficient to tease apart the complex causation underlying the 2004 event. Monitoring programs such as this, therefore, need to be complemented by detailed research to identify the mechanisms involved, and to attribute and predict the effects of natural and human-induced environmental change.

Heterogeneity of impacts of high CO2 on the North Western European Shelf

The increase in atmospheric CO2 is a dual threat to the marine environment: from one side it drives climate change, leading to modifications in water temperature, circulation patterns and stratification intensity; on the other side it causes a decrease in marine pH (ocean acidification, or OA) due to the increase in dissolved CO2. Assessing the combined impact of climate change and OA on marine ecosystems is a challenging task. The response of the ecosystem to a single driver can be highly variable and remains still uncertain; additionally the interaction between these can be either synergistic or antagonistic. In this work we use the coupled oceanographic–ecosystem model POLCOMS-ERSEM driven by climate forcing to study the interaction between climate change and OA. We focus in particular on carbonate chemistry, primary and secondary production. The model has been run in three different configurations in order to assess separately the impacts of climate change on net primary production and of OA on the carbonate chemistry, which have been strongly supported by scientific literature, from the impact of biological feedbacks of OA on the ecosystem, whose uncertainty still has to be well constrained. The global mean of the projected decrease of pH at the end of the century is about 0.27 pH units, but the model shows significant interaction among the drivers and high variability in the temporal and spatial response. As a result of this high variability, critical tipping point can be locally and/or temporally reached: e.g. undersaturation with respect to aragonite is projected to occur in the deeper part of the central North Sea during summer. Impacts of climate change and of OA on primary and secondary production may have similar magnitude, compensating in some area and exacerbating in others.

Ecosystem function and services provided by the deep sea

The deep sea is often viewed as a vast, dark, remote, and inhospitable environment, yet the deep ocean and seafloor are crucial to our lives through the services that they provide. Our understanding of how the deep sea functions remains limited, but when treated synoptically, a diversity of supporting, provisioning, regulating and cultural services becomes apparent. The biological pump transports carbon from the atmosphere into deep-ocean water masses that are separated over prolonged periods, reducing the impact of anthropogenic carbon release. Microbial oxidation of methane keeps another potent greenhouse gas out of the atmosphere while trapping carbon in authigenic carbonates. Nutrient regeneration by all faunal size classes provides the elements necessary for fueling surface productivity and fisheries, and microbial processes detoxify a diversity of compounds. Each of these processes occur on a very small scale, yet considering the vast area over which they occur they become important for the global functioning of the ocean. The deep sea also provides a wealth of resources, including fish stocks, enormous bioprospecting potential, and elements and energy reserves that are currently being extracted and will be increasingly important in the near future. Society benefits from the intrigue and mystery, the strange life forms, and the great unknown that has acted as a muse for inspiration and imagination since near the beginning of civilization. While many functions occur on the scale of microns to meters and timescales up to years, the derived services that result are only useful after centuries of integrated activity. This vast dark habitat, which covers the majority of the globe, harbour processes that directly impact humans in a variety of ways; however, the same traits that differentiate it from terrestrial or shallow marine systems also result in a greater need for integrated spatial and temporal understanding as it experiences increased use by society. In this manuscript we aim to provide a foundation for informed conservation and management of the deep sea by summarizing the important role of the deep sea in society.

Genotyping an Emiliania huxleyi (Prymnesiophyceae) bloom event in the North Sea reveals evidence of asexual reproduction

Due to the unprecedented rate at which our climate is changing, the ultimate consequence for many species is likely to be either extinction or migration to an alternate habitat. Certain species might, however, evolve at a rate that could make them resilient to the effects of a rapidly changing environment. This scenario is most likely to apply to species that have large population sizes and rapid generation times, such that the genetic variation required for adaptive evolution can be readily supplied. Emiliania huxleyi (Lohm.) Hay and Mohler (Prymnesiophyceae) is likely to be such a species as it is the most conspicuous extant calcareous phytoplankton species in our oceans with generation times of 1 day−1. Here we report on a validated set of microsatellites, in conjunction with the coccolithophore morphology motif genetic marker, to genotype 93 clonal isolates collected from across the world. Of these, 52 came from a single bloom event in the North Sea collected on the D366 UK Ocean Acidification cruise in June-July 2011. There were 26 multilocus genotypes (MLGs) encountered only once in the North Sea bloom and 8 MLGs encountered twice or up to six times. Each of these repeated MLGs exhibited Psex values of less than 0.05 indicating each repeated MLG was the product of asexual reproduction and not separate meiotic events. In addition, we show that the two most polymorphic microsatellite loci, EHMS37 and P01E05, are reporting on regions likely undergoing rapid genetic drift during asexual reproduction. Despite the small sample size, there were many more repeated genotypes than previously reported for other bloom-forming phytoplankton species, including a previously genotyped E. huxleyi bloom event. This study challenges our current assumption that sex is the predominant mode of reproduction during bloom events. Whilst genetic diversity is high amongst extant populations of E. huxleyi, the root cause for this diversity and ultimate fate of these populations still requires further examination. Nonetheless, we show that certain CMM genotypes are found everywhere; while others appear to have a regional bias.

Long-term annual primary production in the Ulleung Basin as a biological hot spot in the East/Japan Sea

Although the Ulleung Basin is an important biological hot spot in East/Japan Sea (hereafter the East Sea), very limited knowledge for seasonal and annual variations in the primary productivity exists. In this study, a recent decadal trend of primary production in the Ulleung Basin was analyzed based on MODIS-derived monthly primary production for a better annual production budget. Based on the MODIS-derived primary production, the mean daily primary productivity was 766.8 mg C m-2 d-1 (SD=+/- 196.7 mg C m-2 d-1) and the annual primary productivity was 280.2 g C m-2 yr-1 (SD=+/- 14.9 g C m-2 yr-1) in the Ulleung Basin during the study period. The monthly contributions of primary production were not largely variable among different months, and a relatively small interannual production variability was also observed in the Ulleung Basin, which indicates that the Ulleung Basin is a sustaining biologically productive region called as hot spot in the East Sea. However, a significant recent decline in the annual primary production was observed in the Ulleung Basin after 2006. Although no strong possibilities were found in this study, the current warming sea surface temperature and a negative phase PDO index were suggested for the recent declining primary production. For a better understanding of subsequent effects on marine ecosystems, more intensive interdisciplinary field studies will be required in the Ulleung Basin.

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.

Trait-based representation of diatom functional diversity in a plankton functional type model of the eutrophied southern North Sea

We introduce a trait-based description of diatom functional diversity to an existing plankton functional type (PFT) model, implemented for the eutrophied coastal ecosystem in the Southern Bight of the North Sea. The trait-based description represents a continuum of diatom species, each characterized by a distinct cell volume, and includes size dependence of four diatom traits: the maximum growth rate, the half-saturation constants for nutrient uptake, the photosynthetic efficiency, and the relative affinity of copepods for diatoms. Through competition under seasonally varying forcing, the fitness of each diatom varies throughout time, and the outcome of competition results in a changing community structure. The predicted seasonal change in mean cell volume of the community is supported by field observations: smaller diatoms, which are more competitive in terms of resource acquisition, prevail during the first spring bloom, whereas the summer bloom is dominated by larger species which better resist grazing. The size-based model is used to determine the ecological niche of diatoms in the area and identifies a range of viable sizes that matches observations. The general trade-off between small, competitive diatoms and large, grazing-resistant species is a convenient framework to study patterns in diatom functional diversity. PFT models and trait-based approaches constitute promising complementary tools to study community structure in marine ecosystems.

Annual study of oxygenated volatile organic compounds in UK shelf waters

We performed an annual study of oxygenated volatile organic compound (OVOC) seawater concentrations at a site off Plymouth, UK in the Western English Channel over the period of February 2011–March 2012. Acetone concentrations ranged from 2–10 nM (nanomole/L) in surface waters with a maximum observed in summer. Concentrations correlated positively with net shortwave radiation and UV light, suggestive of photochemically linked acetone production. We observed a clear decline in acetone concentrations below the mixed layer. Acetaldehyde varied between 4–37 nM in surface waters with higher values observed in autumn and winter. Surface concentrations of methanol ranged from 16–78 nM, but no clear annual cycle was observed. Methanol concentrations exhibited considerable inter-annual variability. We estimate consistent deposition to the sea surface for acetone and methanol but that the direction of the acetaldehyde flux varies during the year.

Dynamics of air–sea CO2 fluxes in the northwestern European shelf based on voluntary observing ship and satellite observations

From January 2011 to December 2013, we constructed a comprehensive pCO2 data set based on voluntary observing ship (VOS) measurements in the western English Channel (WEC). We subsequently estimated surface pCO2 and air–sea CO2 fluxes in northwestern European continental shelf waters using multiple linear regressions (MLRs) from remotely sensed sea surface temperature (SST), chlorophyll a concentration (Chl a), wind speed (WND), photosynthetically active radiation (PAR) and modeled mixed layer depth (MLD). We developed specific MLRs for the seasonally stratified northern WEC (nWEC) and the permanently well-mixed southern WEC (sWEC) and calculated surface pCO2 with uncertainties of 17 and 16 μatm, respectively. We extrapolated the relationships obtained for the WEC based on the 2011–2013 data set (1) temporally over a decade and (2) spatially in the adjacent Celtic and Irish seas (CS and IS), two regions which exhibit hydrographical and biogeochemical characteristics similar to those of WEC waters. We validated these extrapolations with pCO2 data from the SOCAT and LDEO databases and obtained good agreement between modeled and observed data. On an annual scale, seasonally stratified systems acted as a sink of CO2 from the atmosphere of −0.6 ± 0.3, −0.9 ± 0.3 and −0.5 ± 0.3 mol C m−2 yr−1 in the northern Celtic Sea, southern Celtic sea and nWEC, respectively, whereas permanently well-mixed systems acted as source of CO2 to the atmosphere of 0.2 ± 0.2 and 0.3 ± 0.2 mol C m−2 yr−1 in the sWEC and IS, respectively. Air–sea CO2 fluxes showed important inter-annual variability resulting in significant differences in the intensity and/or direction of annual fluxes. We scaled the mean annual fluxes over these provinces for the last decade and obtained the first annual average uptake of −1.11 ± 0.32 Tg C yr−1 for this part of the northwestern European continental shelf. Our study showed that combining VOS data with satellite observations can be a powerful tool to estimate and extrapolate air–sea CO2 fluxes in sparsely sampled area.

Modelling the combined impacts of climate change and direct anthropogenic drivers on the ecosystem of the northwest European continental shelf

The potential response of the marine ecosystem of the northwest European continental shelf to climate change under a medium emissions scenario (SRES A1B) is investigated using the coupled hydrodynamics-ecosystem model POLCOMS-ERSEM. Changes in the near future (2030–2040) and the far future (2082–2099) are compared to the recent past (1983–2000). The sensitivity of the ecosystem to potential changes in multiple anthropogenic drivers (river nutrient loads and benthic trawling) in the near future is compared to the impact of changes in climate. With the exception of the biomass of benthic organisms, the influence of the anthropogenic drivers only exceeds the impact of climate change in coastal regions. Increasing river nitrogen loads has a limited impact on the ecosystem whilst reducing river nitrogen and phosphate concentrations affects net primary production(netPP) and phytoplankton and zooplankton biomass. Direct anthropogenic forcing is seen to mitigate/amplify the effects of climate change. Increasing river nitrogen has the potential to amplify the effects of climate change at the coast by increasing netPP. Reducing river nitrogen and phosphate mitigates the effects of climate change for netPP and the biomass of small phytoplankton and large zooplankton species but amplifies changes in the biomass of large phytoplankton and small zooplankton.

Impacts of ocean acidification in a warming Mediterranean Sea: An overview

Mediterranean Sea fisheries supply significant local and international markets, based largely on small pelagic fish, artisanal fisheries and aquaculture of finfish (mainly seabass and seabream) and shellfish (mussels and oysters). Fisheries and aquaculture contribute to the economy of countries bordering this sea and provide food and employment to coastal communities employing ca 600,000 people. Increasing temperatures and heat wave frequency are causing stress and mortality in marine organisms and ocean acidification is expected to worsen these effects, especially for bivalves and coralligenous systems. Recruitment and seed production present possible bottlenecks for shellfish aquaculture in the future since early life stages are vulnerable to acidification and warming. Although adult finfish seem able to withstand the projected increases in seawater CO2, degradation of seabed habitats and increases in harmful blooms of algae and jellyfish might adversely affect fish stocks. Ocean acidification should therefore be factored into fisheries and aquaculture management plans. Rising CO2 levels are expected to reduce coastal biodiversity, altering ecosystem functioning and possibly impacting tourism being the Mediterranean the world’s most visited region. We recommend that ocean acidification is monitored in key areas of the Mediterranean Sea, with regular assessments of the likely socio-economic impacts to build adaptive strategies for the Mediterranean countries concerned.

Tracing early stages of species differentiation: Ecological, morphological and genetic divergence of Galapagos sea lion populations

Background: Oceans are high gene flow environments that are traditionally believed to hamper the build-up of genetic divergence. Despite this, divergence appears to occur occasionally at surprisingly small scales. The Galápagos archipelago provides an ideal opportunity to examine the evolutionary processes of local divergence in an isolated marine environment. Galápagos sea lions (Zalophus wollebaeki) are top predators in this unique setting and have an essentially unlimited dispersal capacity across the entire species range. In theory, this should oppose any genetic differentiation.
Results: We find significant ecological, morphological and genetic divergence between the western colonies and colonies from the central region of the archipelago that are exposed to different ecological conditions. Stable isotope analyses indicate that western animals use different food sources than those from the central area. This is likely due to niche partitioning with the second Galápagos eared seal species, the Galápagos fur seal (Arctocephalus galapagoensis) that exclusively dwells in the west. Stable isotope patterns correlate with significant differences in foraging-related skull morphology. Analyses of mitochondrial sequences as well as microsatellites reveal signs of initial genetic differentiation.
Conclusion: Our results suggest a key role of intra- as well as inter-specific niche segregation in the evolution of genetic structure among populations of a highly mobile species under conditions of free movement. Given the monophyletic arrival of the sea lions on the archipelago, our study challenges the view that geographical barriers are strictly needed for the build-up of genetic divergence. The study further raises the interesting prospect that in social, colonially breeding mammals additional forces, such as social structure or feeding traditions, might bear on the genetic partitioning of populations.