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.

Simulation of mackerel (Scomber scombrus) recruitment with an individual-based model and comparison with field data

An individual-based model (IBM) for the simulation of year-to-year survival during the early life-history stages of the north-east Atlantic stock of mackerel (Scomber scombrus) was developed within the EU funded Shelf-Edge Advection, Mortality and Recruitment (SEAMAR) programme. The IBM included transport, growth and survival and was used to track the passive movement of mackerel eggs, larvae and post-larvae and determine their distribution and abundance after approximately 2 months of drift. One of the main outputs from the IBM, namely distributions and numbers of surviving post-larvae, are compared with field data as recruit (age-0/age-1 juveniles) distribution and abundance for the years 1998, 1999 and 2000. The juvenile distributions show more inter-annual and spatial variability than the modelled distributions of survivors; this may be due to the restriction of using the same initial egg distribution for all 3 yr of simulation. The IBM simulations indicate two main recruitment areas for the north-east Atlantic stock of mackerel, these being Porcupine Bank and the south-eastern Bay of Biscay. These areas correspond to areas of high juvenile catches, although the juveniles generally have a more widespread distribution than the model simulations. The best agreement between modelled data and field data for distribution (juveniles and model survivors) is for the year 1998. The juvenile catches in different representative nursery areas are totalled to give a field abundance index (FAI). This index is compared with a model survivor index (MSI) which is calculated from the total of survivors for the whole spawning season. The MSI compares favourably with the FAI for 1998 and 1999 but not for 2000; in this year, juvenile catches dropped sharply compared with the previous years but there was no equivalent drop in modelled survivors.

Is observed variability in the long-term results of the Continuous Plankton Recorder survey a response to climate change?

In the more than 50 years that the Continuous Plankton Recorder (CPR) survey has operated on a regular monthly basis in the north-east Atlantic and North Sea, large changes have been witnessed in the planktonic ecosystem. These changes have taken the form of long-term trends in abundance for certain species or stepwise changes for others, and in many cases are correlated with a mode of climatic variability in the North Atlantic, either: (1) the North Atlantic Oscillation (NAO), a basin-scale atmospheric alteration of the pressure field between the Azores high pressure cell and the Icelandic Low; or (2) the Gulf Stream Index (GSI), which measures the latitudinal position of the north wall of the Gulf Stream. Recent work has shown that the changes in the GSI are coupled with the NAO and Pacific Southern Oscillation with a 2 year lag. The plankton variability is also possibly linked to changes observed in the distribution and flux of water masses in the surface, intermediate and deep waters of the North Atlantic. For example, in the last two decades, the extent and location of the formation of North Atlantic Deep Water, Labrador Sea Intermediate Water and Norwegian Sea intermediate and upper-layer water has altered considerably. This paper discusses the extent to which observed changes in plankton abundance and distribution may be linked to this basin-scale variability in hydrodynamics. The results are also placed within the context of global climate warming and the possible effects of the observed melting of Arctic permafrost and sea ice on the subpolar North Atlantic.

North Sea Ecosystem: Status Report

The North Sea is a dynamic large marine ecosystem which is bordered by a dense coastal population, contains a productive oil and gas province, has a dense shipping network and has one of the most productive fisheries in the world. An assessment of the state of health of the North Sea was initiated in 1987 as part of a developing series of international initiatives at Ministerial level to address concerns over the impact of these activities on the marine ecosystem. Four North Sea Ministerial Conferences (1984, 1987, 1990, 1995) and an Intermediate Ministerial Meeting (1993) have been held to date to develop a harmonized approach to the sustainable management of the North Sea. In 1988 at the request of Ministers a North Sea Task Force was established to co-ordinate work leading to the production of a Quality Status Report (QSR) on the North Sea in December 1993. In recognition of the large geographical and ecological diversity exhibited, a sub-regional approach was adopted and a total of 13 sub-regional assessment reports were produced to a common protocol. The Task Force established a five-year plan to co-ordinate research, monitoring and modelling and other special topics in the preparations for the QSR. As part of this exercise a ‘Monitoring Master Plan’ was drawn up to provide for the first time reliable spatial information on the distribution of chemical contaminants and biological effects throughout the North Sea. The Task Force was a unique structure in international collaboration with a fixed remit that ended in December 1993. It was successful in bringing together many diverse organisations with interests in the North Sea and co-ordinated to a tight timetable the production of the QSR. The experiences gained are now being applied to the whole north east Atlantic under a new OSPAR Convention and have wide application to other Large Marine Ecosystems.

Continuous Plankton Records - Persistence In Time-Series And The Population-Dynamics Of Pseudocalanus-Elongatus And Acartia-Clausi

Rates of population increase in early spring and the sizes of overwintering stocks were calculated for the planktonic copepods Pseudocalanus elongatus and Acartia clausi for a set of areas covering the open waters of the north-east Atlantic Ocean and the North Sea for the period 1948 to 1979. For both species, the rates of population increase were higher in the open ocean than in the North Sea and appear to be related to temperature. The overwintering stocks in the North Sea were larger than those in the open ocean and are probably related to phytoplanton concentration. P. elongatus shows higher overwintering stocks and lower rates of population increase than A. clausi, resulting in different levels of persistence in the stocks of the two species. It is suggested that this difference in persistence is responsible for differences between the two species with respect to geographical distribution in summer and different patterns of year-to-year fluctuations in abundance.

Continuous Plankton Records - Persistence In Time-Series Of Annual Means Of Abundance Of Zooplankton

Time-series of annual means of abundance of zooplankton of the north-east Atlantic Ocean and the North Sea, for the period 1948 to 1977, show considerable associations between successive years. The seasonal dynamics of the stocks appear to be consistent with at least a proportion of this being due to inherent persistence from year-to-year. Experiments with a simple model suggest that the observed properties of the time-series cannot be reproduced as a response to simple random forcing. The extent of trends and long wavelength variations can be simulated by introducing fairly extensive persistence into the perturbations, but this underestimates the extent of shorter wavelength variability in the observed time-series. The effect of persistence is to increase the proportion of trend and long wavelength variability in time-series of annual means, but stocks can respond to short wavelength perturbations provided these have a clearly defined frequency.

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.

The Mechanics of QSR 2000

Assessment of the quality of the marine environment forms an important part of the new 1992 OSPAR Convention for the Protection of the Marine Environment of the North-East Atlantic that was ratified and entered into force on 25 March 1998. In the ministerial statement at the signing of the Convention it was agreed that the first assessment (Quality Status Report, QSR) for all Convention waters should be produced for the year 2000. To oversee this charge a new Environmental Assessment and Monitoring Committee (ASMO) was established and a junior group under this committee, to implement necessary actions, the Assessment Co-ordination Group (ACG). Because of the wide geographical diversity and varying levels of information available in different parts of the Convention area it was decided to produce five regional reports for: I The Arctic; II The North Sea; III The Celtic seas; IV The Bay of Biscay and Iberian Coast; V The Wider Atlantic, which will be synthesised in a holistic QSR for the year 2000. The report for the North Sea will largely be an update of QSR 1993 and forms the third cycle of a developing management system for the North Sea. This paper will present the procedures that have been adopted to implement the QSRs, and outlines the guidelines that have been developed for their structure, format, design and publication.

Effect of phytoplankton size classes on bio-optical properties of phytoplankton in the Western Iberian coast: Application of models

Chlorophyll-a satellite products are routinely used in oceanography, providing a synoptic and global view of phytoplankton abundance. However, these products lack information on the community structure of the phytoplankton, which is crucial for ecological modelling and ecosystem studies. To assess the usefulness of existing methods to differentiate phytoplankton functional types (PFT) or phytoplankton size classes from satellite data, in-situ phytoplankton samples collected in the Western Iberian coast, on the North-East Atlantic, were analysed for pigments and absorption spectra. Water samples were collected in five different locations, four of which were located near the shore and another in an open-ocean, seamount region. Three different modelling approaches for deriving phytoplankton size classes were applied to the in situ data. Approaches tested provide phytoplankton size class information based on the input of pigments data (Brewin et al., 2010), absorption spectra data (Ciotti et al., 2002) or both (Uitz et al., 2008). Following Uitz et al. (2008), results revealed high variability in microphytoplankton chlorophyll-specific absorption coefficients, ranging from 0.01 to 0.09 m2 (mg chl)− 1 between 400 and 500 nm. This spectral analysis suggested, in one of the regions, the existence of small cells (< 20 μm) in the fraction of phytoplankton presumed to be microphytoplankton (based on diagnostic pigments). Ciotti et al. (2002) approach yielded the highest differences between modelled and measured absorption spectra for the locations where samples had high variability in community structure and cell size. The Brewin et al. (2010) pigment-based model was adjusted and a set of model coefficients are presented and recommended for future studies in offshore water of the Western Iberian coast.

Historical data reveal power-law dispersal patterns of invasive aquatic species.

Understanding how invasive species spread is of particular concern in the current era of globalisation and rapid environmental change. The occurrence of super-diffusive movements within the context of Lévy flights has been discussed with respect to particle physics, human movements, microzooplankton, disease spread in global epidemiology and animal foraging behaviour. Super-diffusive movements provide a theoretical explanation for the rapid spread of organisms and disease, but their applicability to empirical data on the historic spread of organisms has rarely been tested. This study focuses on the role of long-distance dispersal in the invasion dynamics of aquatic invasive species across three contrasting areas and spatial scales: open ocean (north-east Atlantic), enclosed sea (Mediterranean) and an island environment (Ireland). Study species included five freshwater plant species, Azolla filiculoides, Elodea canadensis, Lagarosiphon major, Elodea nuttallii and Lemna minuta; and ten species of marine algae, Asparagopsis armata, Antithamnionella elegans, Antithamnionella ternifolia, Codium fragile, Colpomenia peregrina, Caulerpa taxifolia, Dasysiphonia sp., Sargassum muticum, Undaria pinnatifida and Womersleyella setacea. A simulation model is constructed to show the validity of using historical data to reconstruct dispersal kernels. Lévy movement patterns similar to those previously observed in humans and wild animals are evident in the re-constructed dispersal pattern of invasive aquatic species. Such patterns may be widespread among invasive species and could be exacerbated by further development of trade networks, human travel and environmental change. These findings have implications for our ability to predict and manage future invasions, and improve our understanding of the potential for spread of organisms including infectious diseases, plant pests and genetically modified organisms.

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.

Identifying species and ecosystem sensitivities.

Executive Summary The programme of work was commissioned in September 1998 to supply information to underpin the UK’s commitments to protection and conservation of the ecosystems and biodiversity of the marine environment under the 1992 OSPAR Convention on the Protection of the Marine Environment of the North East Atlantic. The programme also provided support for the implementation of the Biodiversity Convention and the EU Habitats Directive. The MarLIN programme initiated a new approach to assessing sensitivity and recoverability characteristics of seabed species and biotopes based on structures (such as the seabed biotopes classification) and criteria (such as for assessing rarity and defining ‘sensitivity’) developed since 1997. It also developed tools to disseminate the information on the Internet. The species researched were those that were listed in conventions and directives, included in Biodiversity Action Plans, or were nationally rare or scarce. In addition, species were researched if they maintained community composition or structure and/or provided a distinctive habitat or were special to or especially abundant in a particular situation or biotope At its conclusion in August 2001, the work carried out under the contract with DETR/DEFRA had: · Developed protocols, criteria and structures for identifying ‘sensitivity’ and ‘recoverability’, which were tested by a programme management group. · Developed a database to hold research data on biology and sensitivity of species and biotopes. · Defined the link between human activities and the environmental factors likely to be affected by those activities. · Developed a user-friendly Web site to access information from the database, on the sensitivity and recoverability characteristics of over 100 species and basic information on over 200 species. Additionally, the project team have: · Brought together and facilitated discussion between current developers and users of electronic resources for environmental management, protection and education in the conference ‘Using Marine Biological Information in the Electronic Age’ (19-21 July 1999). · Contributed to the development of Ecological Quality Objectives for the North Sea (Scheveningen, 11- 3 September 1999 and subsequent papers). · Provided detailed information on species as a supplement to the National Biodiversity Network Gateway demonstration www.searchnbn.net. · Developed a peer-reviewed approach to electronic publication of updateable information. · Promoted the contract results and the MarLIN approach to the support of marine environmental management and protection at European research fora and, through the web site, internationally. The information available through the Web site is now being used by consultants and Government agencies. The DEFRA contract has been of critical importance in establishing the Marine Life Information Network (MarLIN) programme and has encouraged support from other organisations. Other related work in the MarLIN programme is on-going, especially to identify sensitivity of biotopes to support management of SACs (contract from English Nature in collaboration with Scottish Natural Heritage), to access data sources (in collaboration with the National Biodiversity Network) and to establish volunteer recording schemes for marine life. The results of the programme are best viewed on the Web site (www.marlin.ac.uk). Three reports have been produced during the project. A final report detailing the work undertaken, a brochure ‘Identifying the sensitivity of seabed ecosystems’ and a CD-ROM describing the programme and demonstrating the Web site have been delivered as final products in addition to the Web site.