An overview of Calanus helgolandicus ecology in European waters

We review current knowledge and understanding of the biology and ecology of the calanoid copepod Calanus helgolandicus in European waters, as well as provide a collaborative synthesis of data from 18 laboratories and 26 sampling stations in areas distributed from the northern North Sea to the Aegean and Levantine Seas. This network of zooplankton time-series stations has enabled us to collect and synthesise seasonal and multi-annual data on abundance, body size, fecundity, hatching success and vertical distribution of C. helgolandicus. An aim was to enable comparison with its congener Calanus finmarchicus, which has been studied intensively as a key component of European and north east Atlantic marine ecosystems. C. finmarchicus is known to over-winter at depth, whereas the life-cycle of C. helgolandicus is less well understood. Overwintering populations of C. helgolandicus have been observed off the Atlantic coast between 400 and 800 m, while in the Mediterranean there is evidence of significant deep-water populations at depths as great as 4200 m. The biogeographical distribution of C. helgolandicus in European coastal waters covers a wide range of habitats, from open ocean to coastal environments, and its contribution to mesozooplankton biomass ranges from 6% to 93%. Highest abundances were recorded in the Adriatic and off the west coast of Spain. C. helgolandicus is generally found in 9-20 C water, with maximum abundances from 13-17 C. In contrast, C. finmarchicus is found in cooler water between 0 and 15 C, with peak abundances from 0 to 9 C. As water has warmed in the North Atlantic over recent decades, the range of C. helgolandicus and its abundance on the fringes of its expanding range have increased. This review will facilitate development of population models of C. helgolandicus. This will not only help answer remaining questions but will improve our ability to forecast future changes, in response to a warming climate, in the abundance and distribution of this important species.

Ocean structure and climate (Eastern North Atlantic): in situ measurement and remote sensing

Structure and climate of the east North Atlantic are appraised within a framework of in situ measurement and altimeter remote sensing from 0 degree - 60 degree N. Long zonal expendable bathythermograph /conductivity-temperature-depth probe sections show repeating internal structure in the North Atlantic Ocean. Drogued buoys and subsurface floats give westward speeds for eddies and wavelike structure. Records from longterm current meter deployments give the periodicity of the repeating structure. Eddy and wave characteristics of period, size or wavelength, westward propagation speed, and mean currents are derived at 20 degree N, 26 degree N, 32.5 degree N, 36 degree N and 48 degree N from in situ measurements in the Atlantic Ocean. It is shown that ocean wave and eddy-like features measured in situ correlate with altimeter structure. Interior ocean wave crests or cold dome-like temperature structures are cyclonic and have negative surface altimeter anomalies; mesoscale internal wave troughs or warm structures are anticyclonic and have positive surface height anomalies. Along the Eastern Boundary, flows and temperature climate are examined in terms of sla and North Atlantic Oscillation (NAO) Index. Longterm changes in ocean climate and circulation are derived from sla data. It is shown that longterm changes from 1992 to 2002 in the North Atlantic Current and the Subtropical Gyre transport determined from sla data correlate with winter NAO Index such that maximum flow conditions occurred in 1995 and 2000. Minimum circulation conditions occurred between 1996-1998. Years of extreme negative winter NAO Index result in enhanced poleward flow along the Eastern Boundary and anomalous winter warming along the West European Continental Slope as was measured in 1990, 1996, 1998 and 2001.

Validation of the NEMO-ERSEM operational ecosystem model for the North West European Continental Shelf

This paper details updates to the Met Office's operational coupled hydrodynamic-ecosystem model from the 7 km Medium-Resolution Continental Shelf – POLCOMS-ERSEM (MRCS-PE) system (Siddorn et al., 2007) to the 7 km Atlantic Margin Model NEMO-ERSEM (AMM7-NE) system. We also provide a validation of the ecosystem component of the new operational system. Comparisons have been made between the model variables and available in situ, satellite and climatological data. The AMM7-NE system has also been benchmarked against the MRCS-PE system. The transition to the new AMM7-NE system was successful and it has been running operationally since March 2012 and has been providing products through MyOcean (http://www.myocean.eu.org) since that time. The results presented herein show the AMM7-NE system performs better than the MRCS-PE system with the most improvement in the model nutrient fields. The problem of nutrient accumulation in the MRCS-PE system appears to be solved in the new AMM7-NE system with nutrient fields improved throughout the domain as discussed in Sect. 4. Improvements in model chlorophyll are also seen but are more modest.

The Mediterranean Sea Regime Shift at the End of the 1980s, and Intriguing Parallelisms with Other European Basins

Regime shifts are abrupt changes encompassing a multitude of physical properties and ecosystem variables, which lead to new regime conditions. Recent investigations focus on the changes in ecosystem diversity and functioning associated to such shifts. Of particular interest, because of the implication on climate drivers, are shifts that occur synchronously in separated basins. In this work we analyze and review long-term records of Mediterranean ecological and hydro-climate variables and find that all point to a synchronous change in the late 1980s. A quantitative synthesis of the literature (including observed oceanic data, models and satellite analyses) shows that these years mark a major change in Mediterranean hydrographic properties, surface circulation, and deep water convection (the Eastern Mediterranean Transient). We provide novel analyses that link local, regional and basin scale hydrological properties with two major indicators of large scale climate, the North Atlantic Oscillation index and the Northern Hemisphere Temperature index, suggesting that the Mediterranean shift is part of a large scale change in the Northern Hemisphere. We provide a simplified scheme of the different effects of climate vs. temperature on pelagic ecosystems.

Transregional linkages in the North-Eastern Atlantic - An ‘end-to-end ’ analysis of pelagic ecosystems

This review examines interregional linkages and gives an overview perspective on marine ecosystem functioning in the north-eastern Atlantic. It is based on three of the 'systems' considered by the European Network of Excellence for Ocean Ecosystems Analysis (EUR-OC EANS was established in 2004 under the European Framework VI funding programme to promote integration of marine ecological research within Europe), the Arctic and Nordic Seas, North Atlantic shelf seas and North Atlantic. The three systems share common open boundaries and the transport of water, heat, nutrients and particulates across these boundaries modifies local processes. Consistent with the EUR-OC EANS concept of 'end-to-end' analyses of marine food webs, the review takes an integrated approach linking ocean physics, lower trophic levels and working up the food web to top predators such as marine mammals. We begin with an overview of the regions focusing on the major physical patterns and their implications for the microbial community, phytoplankton, zooplankton, fish and top predators. Human-induced links between the regional systems are then considered and finally possible changes in the regional linkages over the next century are discussed. Because of the scale of potential impacts of climate change, this issue is considered in a separate section. The review demonstrates that the functioning of the ecosystems in each of the regions cannot be considered in isolation and the role of the atmosphere and ocean currents in linking the North Atlantic Ocean, North Atlantic shelf seas and the Arctic and Nordic Seas must be taken into account. Studying the North Atlantic and associated shelf seas as an integrated 'basin-scale' system will be a key challenge for the early twenty-first century. This requires a multinational approach that should lead to improved ecosystem-based approaches to conservation of natural resources, the maintenance of biodiversity, and a better understanding of the key role of the north-eastern Atlantic in the global carbon cycle.

The North Atlantic subpolar gyre regulates the spawning distribution of blue whiting (Micromesistius poutassou)

The spawning stock of blue whiting (Micromesistius poutassou), an economically important pelagic gadoid in the North Atlantic Ocean, increased threefold after 1995. The reproductive success of the stock is largely determined during the very early stages of life, but little is known about the spawning dynamics of this species. Here we show that the spawning distribution of blue whiting is variable, regulated by the hydrography west of the British Isles. When the North Atlantic subpolar gyre is strong and spreads its cold, fresh water masses east over Rockall Plateau, the spawning is constrained along the European continental slope and in a southerly position near Porcupine Bank. When the gyre is weak and conditions are relatively saline and warm, the spawning distribution moves northwards along the slope and especially westwards covering Rockall Plateau. The apparent link between the spawning distribution and the subpolar gyre is the first step towards understanding the reproduction variability, which currently is the main challenge for appropriate management of the blue whiting stock.

Decadal fluctuations in North Atlantic water inflow in the North Sea between 1958-2003: impacts on temperature and phytoplankton populations

The circulation of Atlantic water along the European continental slope, in particular the inflow into the North Sea, influences North Sea water characteristics with consequent changes in the environment affecting plankton community dynamics. The long-term effect of fluctuating oceanographic conditions oil the North Sea, pelagic ecosystem is assessed. It is shown that (i) there are similar regime shifts in the inflow through the northern North Sea and in Sea, Surface Temperature, (ii) long-term phytoplankton trends are influenced by the inflow only in some North Sea regions, and (iii) the spatial variability in chemicophysical and biological parameters highlight the influence of smaller scale processes.

Comparative analysis European wide marine ecosystem shifts - a large scale approach for developing the basis for ecosystem-based management

Abrupt and rapid ecosystem shifts (where major reorganizations of food-web and community structures occur), commonly termed regime shifts, are changes between contrasting and persisting states of ecosystem structure and function. These shifts have been increasingly reported for exploited marine ecosystems around the world from the North Pacific to the North Atlantic. Understanding the drivers and mechanisms leading to marine ecosystem shifts is crucial in developing adaptive management strategies to achieve sustainable exploitation of marine ecosystems. An international workshop on a comparative approach to analysing these marine ecosystem shifts was held at Hamburg University, Institute for Hydrobiology and Fisheries Science, Germany on 1-3 November 2010. Twenty-seven scientists from 14 countries attended the meeting, representing specialists from seven marine regions, including the Baltic Sea, the North Sea, the Barents Sea, the Black Sea, the Mediterranean Sea, the Bay of Biscay and the Scotian Shelf off the Canadian East coast. The goal of the workshop was to conduct the first large-scale comparison of marine ecosystem regime shifts across multiple regional areas, in order to support the development of ecosystem-based management strategies.

Prey landscapes help identify potential foraging habitats for leatherback turtles in the NE Atlantic

Identifying key marine megavertebrate habitats has become ever more important as concern increases regarding global fisheries bycatch and accelerated climate change. This will be aided by a greater understanding of the patterns and processes determining the spatiotemporal distribution of species of conservation concern. We identify probable foraging grounds for leatherback turtles in the NE Atlantic using monthly landscapes of gelatinous organism distribution constructed from Continuous Plankton Recorder Survey data. Using sightings data (n = 2013 records, 1954 to 2003) from 9 countries (UK, Ireland, France, Belgium, The Netherlands, Denmark, Germany, Norway and Sweden), we show sea surface temperatures of approximately 10 to 12 degree C most likely indicate the lower thermal threshold for accessible habitats during seasonal foraging migrations to high latitudes. Integrating maps of gelatinous plankton as a possible indicator of prey distribution with thermal tolerance parameters demonstrates the dynamic (spatial and temporal) nature of NE Atlantic foraging habitats. We highlight the importance of body size- related thermal constraints in structuring leatherback foraging populations and demonstrate a latitudinal gradient in body size (Bergmann's rule) where smaller animals are excluded from higher latitude foraging areas. We highlight the marine area of the European continental shelf edge as being both thermally accessible and prey rich, and therefore potentially supporting appreciable densities of foraging leatherbacks, with some suitable areas not yet extensively surveyed.

Reply to comment: is climate change the most likely driver of range expansion of a critically endangered top predator in northeast Atlantic waters?

The comment by Votier et al. (2008) on our recently published article (Wynn et al. 2007) contains two main criticisms: (i) that our analytical approach is inappropriate and (ii) that we have failed to acknowledge other factors that may have contributed to the change in Balearic Shearwater numbers recorded throughout northwest European waters. We strongly disagree with both these criticisms.

Air–sea fluxes of oxygenated volatile organic compounds across the Atlantic Ocean

We present air-sea fluxes of oxygenated volatile organics compounds (OVOCs) quantified by eddy covariance (EC) during the Atlantic Meridional Transect cruise in 2012. Measurements of acetone, acetaldehyde, and methanol in air as well as in water were made in several different oceanic provinces and over a wide range of wind speeds (1-18 m s(-1)). The ocean appears to be a net sink for acetone in the higher latitudes of the North Atlantic but a source in the subtropics. In the South Atlantic, seawater acetone was near saturation relative to the atmosphere, resulting in essentially zero net flux. For acetaldehyde, the two-layer model predicts a small oceanic emission, which was not well resolved by the EC method. Chemical enhancement of air-sea acetaldehyde exchange due to aqueous hydration appears to be minor. The deposition velocity of methanol correlates linearly with the transfer velocity of sensible heat, confirming predominant airside control. We examine the relationships between the OVOC concentrations in air as well as in water, and quantify the gross emission and deposition fluxes of these gases.

Seasonality and interannual variability of the European Slope Current from 20 years of altimeter data compared with in situ measurements

The European Slope Current (SC) is a major section of the warm poleward flow from the Atlantic to the Arctic, which also moderates the exchange of heat, salt, nutrients and carbon between the deep ocean and the European shelf seas. The mean structure of the geostrophic flow, seasonality, interannual variability and long-term trend of SC are appraised with an unprecedented continuous 20-year satellite altimeter dataset. Comparisons with long term in situ data showed a maximum correlation of r2=0.51 between altimeter and Acoustic Doppler Current Profilers (ADCP), with similar results for drogued buoy data. Mean geostrophic currents were appraised more comprehensively than previous attempts, and the paths of 4 branches of the North Atlantic Current (NAC) and positions of 5 eddies in the region were derived quantitatively. A consistent seasonal cycle in the flow of the SC was found at all 8 sections along the European shelf slope, with maximum poleward flow in the winter and minimum in the summer. The seasonal difference in the altimetry current speed amounted to ~8-10 cm s-1 at the northern sections, but only ~5 cm s-1 on the Bay of Biscay slopes. This extended altimeter dataset indicates significant regional and seasonal variations, and has revealed new insights into the interannual variability of the SC. It is shown that there is a peak poleward flow at most positions along a ~2000 km stretch of the continental slope from Portugal to Scotland during 1995-1997, but this did not clearly relate to the extreme negative North Atlantic Oscillation (NAO) in the winter of 1995-1996. The speed of the SC exhibited a long term decreasing trend of ~1% per year. By contrast the NAC showed no significant trend over the 20-year period. Major changes in the NAC occurred three times, and these changes followed decreases in the NAO index.

Potential impacts of climate change on the primary production of regional seas: A comparative analysis of five European seas

Regional seas are potentially highly vulnerable to climate change, yet are the most directly societally important regions of the marine environment. The combination of widely varying conditions of mixing, forcing, geography (coastline and bathymetry) and exposure to the open-ocean makes these seas subject to a wide range of physical processes that mediates how large scale climate change impacts on these seas’ ecosystems. In this paper we explore the response of five regional sea areas to potential future climate change, acting via atmospheric, oceanic and terrestrial vectors. These include the Barents Sea, Black Sea, Baltic Sea, North Sea, Celtic Seas, and are contrasted with a region of the Northeast Atlantic. Our aim is to elucidate the controlling dynamical processes and how these vary between and within these seas. We focus on primary production and consider the potential climatic impacts on: long term changes in elemental budgets, seasonal and mesoscale processes that control phytoplankton’s exposure to light and nutrients, and briefly direct temperature response. We draw examples from the MEECE FP7 project and five regional model systems each using a common global Earth System Model as forcing. We consider a common analysis approach, and additional sensitivity experiments. Comparing projections for the end of the 21st century with mean present day conditions, these simulations generally show an increase in seasonal and permanent stratification (where present). However, the first order (low- and mid-latitude) effect in the open ocean projections of increased permanent stratification leading to reduced nutrient levels, and so to reduced primary production, is largely absent, except in the NE Atlantic. Even in the two highly stratified, deep water seas we consider (Black and Baltic Seas) the increase in stratification is not seen as a first order control on primary production. Instead, results show a highly heterogeneous picture of positive and negative change arising from complex combinations of multiple physical drivers, including changes in mixing, circulation and temperature, which act both locally and non-locally through advection.

Quantitative pathways for Northeast Atlantic fisheries based on climate, ecological–economic and governance modelling scenarios

Here we present quantitative projections of potential futures for ecosystems in the North Atlantic basin generated from coupling a climate change-driven biophysical model (representing ecosystem and fish populations under climate change) and a scenario-driven ecological–economic model (representing fleets and industries under economic globalization). Four contrasting scenarios (Baseline, Fortress, Global Commons, Free Trade) were defined from the perspective of alternative regional management and governance of the oceanic basin, providing pathways for the future of ecosystems in the Northeast Atlantic basin by 2040. Results indicate that in the time frame considered: (1) the effects of governance and trade decisions are more significant in determining outcomes than the effects of climate change alone, (2) climate change is likely to result in a poleward latitudinal shift of species ranges and thus resources, with implications for exploitation patterns, (3) the level of fisheries regulation is the most important factor in determining the long term evolution of the fisheries system, (4) coupling climate change and governance impacts demonstrates the complex interaction between different components of this social–ecological system, (5) an important driver of change for the future of the North Atlantic and the European fishing fleets appears to be the interplay between wild fisheries and aquaculture development, and finally (6) scenarios demonstrate that the viability and profit of fisheries industries is highly volatile. This study highlights the need to explore basin-scale policy that combines medium to long-term environmental and socio-economic considerations, and the importance of defining alternative sustainable pathways.

Decadal reanalysis of biogeochemical indicators and fluxes in the North West European shelf-sea ecosystem

In this paper we present the first decadal reanalysis simulation of the biogeochemistry of the North West European shelf, along with a full evaluation of its skill and value. An error-characterized satellite product for chlorophyll was assimilated into a physical-biogeochemical model of the North East Atlantic, applying a localized Ensemble Kalman filter. The results showed that the reanalysis improved the model predictions of assimilated chlorophyll in 60% of the study region. Model validation metrics showed that the reanalysis had skill in matching a large dataset of in situ observations for ten ecosystem variables. Spearman rank correlations were significant and higher than 0.7 for physical-chemical variables (temperature, salinity, oxygen), ∼0.6 for chlorophyll and nutrients (phosphate, nitrate, silicate), and significant, though lower in value, for partial pressure of dissolved carbon dioxide (∼0.4). The reanalysis captured the magnitude of pH and ammonia observations, but not their variability. The value of the reanalysis for assessing environmental status and variability has been exemplified in two case studies. The first shows that between 340,000-380,000 km2 of shelf bottom waters were oxygen deficient potentially threatening bottom fishes and benthos. The second application confirmed that the shelf is a net sink of atmospheric carbon dioxide, but the total amount of uptake varies between 36-46 Tg C yr−1 at a 90% confidence level. These results indicate that the reanalysis output dataset can inform the management of the North West European shelf ecosystem, in relation to eutrophication, fishery, and variability of the carbon cycle.