An overview of plankton ecology in the North Sea

The purpose of this report is to give an overview of plankton ecology in the North Sea, and the processes that effect it, as derived from current research. The Sir Alister Hardy Foundation has extensive data for the North Sea area, and other sources have also been used to provide information for this report. Shortfalls in current research have also been highlighted. The information contained herein is to be contributed towards an information base for the Strategic Environmental Assessment. The North Sea is an extension of the North Atlantic that has an area of 574,980 km2. The deepest area is off the coast of Norway (660m), with a number of shallow areas, such as the Dogger Bank (15m). The North Sea represents a large source of hydrocarbons that have been exploited since the early 1970s. The aim of this study is to provide the Department of Trade and Industry with biological data on the planktonic community of the North Sea, as a contribution towards the Strategic Environmental Assessment (SEA 2). An overview of phyto- and zoo- plankton community composition, plankton blooms, Calanus, mero-, pico- and megaplankton, sensitivity to disturbance / contamination, phytodetritus and vertical fluxes and the resting stages of phytoplankton is made using the results of the survey database. Additional published literature has also been used, and gaps in available data have been highlighted. 1.3 The Continuous Plankton Recorder (CPR) survey provides a unique long-term dataset of plankton abundance in the North Atlantic and North Sea (Warner and Hays 1994). The survey has been running for almost 70 years, using ‘ships of opportunity’ to tow CPRs on regular, and incidental routes, sampling at a depth of 10 m. Each sample represents 18 km of tow and approximately 3 m3 of filtered seawater. Over 400 taxa of plankton are routinely identified by a team of taxonomists. The samples are also compared to colour charts to give an indication of ‘greenness’, which provides a visual index of chlorophyll value. CPRs have been towed for over 4 million nautical miles, accumulating almost 200,000 samples. The design of the CPR has remained virtually unchanged since sampling started, thus providing a consistency of sampling that provides good historical comparisons. By systematically monitoring the plankton over a period, changes in abundance and long term trends can be distinguished. From this baseline data, inferences can be made, particularly concerning climate change and potentialanthropogenic impacts.

Temporal changes in plankton of the North Sea: community shifts and environmental drivers

This paper analyses long-term and seasonal changes in the North Sea plankton community during the period 1970 to 2008. Based on Continuous Plankton Recorder (CPR) data covering 38 yr, major changes in both phytoplankton and zooplankton abundance and community structure were identified. Regime changes were detected around 1978, 1989 and 1998. The first 2 changes have been discussed in the literature and are defined as a cold episodic event (1978) and a regime shift towards a warm dynamic regime (1989). The effect of these 2 regime changes on plankton indicators was assessed and checked against previous studies. The 1998 change represents a shift in the abundance and seasonal patterns of dinoflagellates and the dominant zooplankton group, the neritic copepods. Furthermore, environmental factors such as air temperature, wind speed and the North Atlantic water inflow were identified as potential drivers of change in seasonal patterns, and the most-likely environmental causes for detected changes were assessed. We suggest that a change in the balance of dissolved nutrients driven by these environmental factors was the cause of the latest change in plankton community structure, which in turn could have affected the North Sea fish community.

Long-term variability and environmental preferences of calycophoran siphonophores in the Bay of Villefranche (North-Western Mediterranean)

Long-term variability of the main calycophoran siphonophores was investigated between 1974 and 1999 in a coastal station in the north-western Mediterranean. The data were collected at weekly frequency using a macroplankton net (680 μm mesh size) adapted to quantitatively sample delicate gelatinous plankton. A 3-year collection (1967–1969) of siphonophores from offshore waters using the same methodology showed that the patterns of variability observed inshore were representative of siphonophores’ changes at a regional scale. The aims of the study were: (i) to investigate the patterns of variability that characterised the dominant calycophoran species and assemblages; (ii) to identify the environmental optima that were associated with a significant increase in the dominant siphonophore species and (iii) to verify the influence of hydroclimatic variability on long-term changes of siphonophores. Our results showed that during nearly 3 decades the standing stock of calycophoran siphonophores did not show any significant change, with the annual maximum usually recorded in spring as a result of high densities of the dominant species Lensia subtilis, Muggiaea kochi and Muggiaea atlantica. Nevertheless, major changes in community composition occurred within the calycophoran population. Since the middle 1980s, M. kochi, once the most dominant species, started to decrease allowing other species, the congeneric M. atlantica and Chelophyes appendiculata, to increasingly dominate in spring and summer–autumn, respectively. The comparison of environmental and biotic long-term trends suggests that the decrease of M. kochi was triggered by hydrological changes that occurred in the north-western Mediterranean under the forcing of large-scale climate oscillations. Salinity, water stratification and water temperature were the main hydroclimatic factors associated with a significant increase of siphonophores, different species showing different environmental preferences.

Effects of environmental conditions on the seasonal distribution of phytoplankton biomass in the North Sea

We study the spatial and seasonal variability of phytoplankton biomass (as phytoplankton color) in relation to the environmental conditions in the North Sea using data from the Continuous Plankton Recorder survey. By using only environmental fields and location as predictor variables we developed a nonparametric model (generalized additive model) to empirically explore how key environmental factors modulate the spatio-temporal patterns of the seasonal cycle of algal biomass as well as how these relate to the ,1988 North Sea regime shift. Solar radiation, as manifest through changes of sea surface temperature (SST), was a key factor not only in the seasonal cycle but also as a driver of the shift. The pronounced increase in SST and in wind speed after the 1980s resulted in an extension of the season favorable for phytoplankton growth. Nutrients appeared to be unimportant as explanatory variables for the observed spatio-temporal pattern, implying that they were not generally limiting factors. Under the new climatic regime the carrying capacity of the whole system has been increased and the southern North Sea, where the environmental changes have been more pronounced, reached a new maximum.

The influence of phytoplankton productivity, temperature and environmental stability on the control of copepod diversity in the North East Atlantic

The patterns of copepod species richness (S) and their relationship with phytoplankton productivity, temperature and environmental stability were investigated at climatological, seasonal and year-to-year time scales as well as scales along latitudinal and oceanic–neritic gradients using monthly time series of the Continuous Plankton Recorder (CPR) Survey collected in the North East Atlantic between 1958 and 2006. Time series analyses confirmed previously described geographic patterns. Equatorward and towards neritic environments, the climatological average of S increases and the variance explained by the seasonal cycle decreases. The bi-modal character of seasonality increases equatorward and the timing of the seasonal cycle takes place progressive earlier equatorward and towards neritic environments. In the long-term, the climatological average of S decreased significantly (p < 0.001) between 1958 and 2006 in the Bay of Biscay and North Iberian shelf at a rate of ca. 0.04 year−1, and increased at the same rate between 1991 and 2006 in the northernmost oceanic location. The climatological averages of S correlate positively with those of the index of seasonality of phytoplankton productivity (ratio between the minimum and maximum monthly values of surface chlorophyll) and sea surface temperature, and negatively with those of the proxy for environmental stability (monthly frequency of occurrence of daily averaged wind speed exceeding 10 m s−1). The seasonal cycles of S and phytoplankton productivity (surface chlorophyll as proxy) exhibit similar features in terms of shape, timing and explained variance, but the relationship between the climatological averages of both variables is non-significant. From year-to-year, the annual averages of S correlate negatively with those of phytoplankton productivity and positively with those of sea surface temperature along the latitudinal gradient, and negatively with those of environmental stability along the oceanic–neritic gradient. The annual anomalies of S (i.e. factoring out geographic variation) show a unimodal relationship with those of sea surface temperature and environmental stability, with S peaking at intermediate values of the anomalies of these variables. The results evidence the role of seasonality of phytoplankton productivity on the control of copepod species richness at seasonal and climatological scales, giving support to the species richness–productivity hypothesis. Although sea surface temperature (SST) is indeed a good predictor of richness along the latitudinal gradient, it is unable to predict the increase of richness form oceanic to neritic environments, thus lessening the generality of the species richness–energy hypothesis. Meteo-hydrographic disturbances (i.e. SST and wind speed anomalies as proxies), presumably through its role on mixed layer depth dynamics and turbulence and hence productivity, maximise local diversity when occurring at intermediate frequency and or intensity, thus providing support to the intermediate disturbance hypothesis on the control of copepod diversity.

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.

Challenges and difficulties in assessing the Environmental Status under the requirements of the Ecosystem Approach in North-African countries, illustrated by eutrophication assessment

Marine ecosystems provide many ecosystem goods and services. However, these ecosystems and the benefits they create for humans are subject to competing uses and increasing pressures. As a consequence of the increasing threats to the marine environment, several regulations require applying an ecosystem-based approach for managing the marine environment. Within the Mediterranean Sea, in 2008, the Contracting Parties of the Mediterranean Action Plan decided to progressively apply the Ecosystem Approach (EcAp) with the objective of achieving Good Environmental Status (GES) for 2018. To assess the Environmental Status, the EcAp proposes 11 Ecological Objectives, each of which requires a set of relevant indicators to be integrated. Progress towards the EcAp entails a gradual and important challenge for North-African countries, and efforts have to be initiated to propose and discuss methods. Accordingly, to enhance the capacity of North-African countries to implement EcAp and particularly to propose and discuss indicators and methods to assess GES, the aim of this manuscript is to identify the practical problems and gaps found at each stage of the Environmental Status assessment process. For this purpose, a stepwise method has been proposed to assess the Environmental Status using Ecologic Objective 5-Eutrophication as example.

Challenges and difficulties in assessing the Environmental Status under the requirements of the Ecosystem Approach in North-African countries, illustrated by eutrophication assessment

Marine ecosystems provide many ecosystem goods and services. However, these ecosystems and the benefits they create for humans are subject to competing uses and increasing pressures. As a consequence of the increasing threats to the marine environment, several regulations require applying an ecosystem-based approach for managing the marine environment. Within the Mediterranean Sea, in 2008, the Contracting Parties of the Mediterranean Action Plan decided to progressively apply the Ecosystem Approach (EcAp) with the objective of achieving Good Environmental Status (GES) for 2018. To assess the Environmental Status, the EcAp proposes 11 Ecological Objectives, each of which requires a set of relevant indicators to be integrated. Progress towards the EcAp entails a gradual and important challenge for North-African countries, and efforts have to be initiated to propose and discuss methods. Accordingly, to enhance the capacity of North-African countries to implement EcAp and particularly to propose and discuss indicators and methods to assess GES, the aim of this manuscript is to identify the practical problems and gaps found at each stage of the Environmental Status assessment process. For this purpose, a stepwise method has been proposed to assess the Environmental Status using Ecologic Objective 5-Eutrophication as example.

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.

Visions for the North Sea: The Societal Dilemma Behind Specifying Good Environmental Status.

We augment discussions about the Good Environmental Status of the North Sea by developing two extreme visions and assessing their societal benefits. One vision (‘Then’) assumes restoration of benthic functioning; we contend that trawling had already degraded the southern North Sea a century ago. Available information is used to speculate about benthic functioning in a relatively undisturbed southern North Sea. The second vision (‘Now’) draws on recent benthic functioning. The supply of five ecosystem services, supported by benthic functioning, is discussed. ‘Then’ offers confidence in the sustainable supply of diverse services but restoration of past function is uncertain and likely to be paired with costs, notably trawling restraints. ‘Now’ delivers known and valued services but sustained delivery is threatened by, for example, climate change. We do not advocate either vision. Our purpose is to stimulate debate about what society wants, and might receive, from the future southern North Sea.