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.

Interregional Biological Responses in the North Atlantic to Hydrometeorological Forcing

Using data from the CPR survey seven case studies are described that document different spatial and temporal responses in the plankton to hydroclimatic events. Long-term trends in the plankton of the eastern Atlantic and the North Sea over the last five decades are examined. Two of the examples revisit correlations that have been described between copepod abundance in the eastern Atlantic and North Sea and indices of atmospheric variability, the North Atlantic Oscillation index and the Gulf Stream North Wall index. Evidence for an increase in levels of Phytoplankton Colour (a visual index of chlorophyll) on the eastern and western sides of the Atlantic is presented. Changes in three trophic levels and in the hydrodynamics and chemistry of the North Sea circa 1988 are outlined as a regime shift. Two of the case studies emphasise the importance of variability in oceanic advection into shelf seas and the role of western and eastern margin currents at the shelf edge. The plankton appear to be integrating hydrometeorological signals and reflecting basin scale changes in circulation of surface, intermediate and deep waters in part associated with the NAO. The extent to which climatic variability may be contributing to the observed changes in the plankton is discussed with a forecast of potential future ecosystem effects in a climate change scenario.

Long-term changes in the pelagos, benthos and fisheries of the North Sea

Pronounced changes have occurred in the fisheries, plankton and benthos of the North Sea over the last five decades. Attribution of the relative contribution of anthropogenic versus natural hydrometeorological modulation to these changes is still unclear. As a background a summary history of our understanding of the state of health of the North Sea is outlined. We then focus on two contrasting periods in the North Sea, one between 1978-82 (cold) and the other post 1987 (warm) when pronounced alterations in many ecosystem characteristics occurred. The scale of the changes in the second of these periods is sufficiently large and wide ranging for it to have been termed a regime shift. A combination of local, regional and far field hydrometeorological forcing, and in particular variability in oceanic inflow, is believed to be responsible for the observed changes. Finally attention is drawn to the poor status of North Sea fish stocks where 7 stocks are documented as being fished outside safe biological limits. This situation is primarily believed to be a consequence of overfishing, but may have been exacerbated by environmental change.

Climate-related increases in jellyfish frequency suggest a more gelatinous future for the North Sea

Data obtained since 1958 from the continuous plankton recorder show an increasing occurrence of jellyfish in the central North Sea that is positively related to the North Atlantic Oscillation (NAO) and Atlantic inflow to the northern North Sea. Since 1970, jellyfish frequency has been also significantly negatively correlated with mean annual pH, independent of NAO trends. Jellyfish frequency increased in the mid-1980s, coincident with the reported regime shift in the North Sea and tracking trends in phytoplankton color. As models produced under all climate-change scenarios indicate a move toward a positive NAO, and pH of the oceans is predicted to decrease with rising CO2, we suggest that jellyfish frequency will increase over the next 100 yr.

Climate effects and benthic-pelagic coupling in the North Sea

The North Sea is one of the most biologically productive ecosystems in the world and supports important fisheries. Climate-induced changes occurred in the pelagic ecosystems of the North Sea during the 1980s. These changes, which have been observed from phytoplankton to fish and among permanent (holoplankton) and temporary (meroplankton) plankton species, have resulted in alterations in plankton community composition and seasonality. Until now, the effects of climate-driven changes on biological linkages between pelagic and benthic ecosystems have not been examined. The present study indicates that changes in benthic organisms could have a profound effect on the trophodynamics of the pelagos. We demonstrate this by analyses of a long-term time series of North Sea plankton and sea surface temperature data. We discover that pronounced changes in the North Sea meroplankton, mainly related to an increased abundance and spatial distribution of the larvae of a benthic echinoderm, Echinocardium cordatum, result primarily from a stepwise increase in sea temperature after 1987 that has caused warmer conditions to occur earlier in the year than previously. Key stages of reproduction in E. cordatum, gametogenesis and spawning, appear to be influenced by winter and spring sea temperature and their larval development is affected by the quantity and quality of their phytoplankton food. Our analyses suggest that a new thermal regime in the North Sea in winter and spring may have benefited reproduction and survival in this benthic species. As a result, E. cordatum may be altering the trophodynamics of the summer pelagic ecosystem through competition between its larvae and holozooplankton taxa.

Increasing prevalence of the marine cladoceran Penilia avirostris (Dana, 1852) in the North Sea

Rising sea surface temperatures in the North Sea have had consequential effects on not only indigenous plankton species, but also on the possibility of successful colonisation of the area by invasive plankton species. Previous studies have noted the introduction and integration into the plankton community of various phytoplankton species, but establishment of zooplankton organisms in the North Sea is less well-documented. Examining continuous plankton recorder (CPR) survey data and zooplankton results from the Helgoland Roads study, the autumn of 1999 witnessed the occurrence of the marine cladoceran Penilia avirostris in large numbers in the North Sea. The rapid appearance of the species corresponded with exceptionally warm sea surface temperatures (SSTs). Since 1999, the species has become a regular feature of the autumnal zooplankton community of the North Sea. In 2002 and 2003, the species occurred in greater abundance than recorded before. It is suggested that increased autumn SSTs have proved favourable to P. avirostris, with warmer conditions contributing to the success of the species’ resting eggs and aiding colonisation.

Phaeocystis colony distribution in the North Atlantic Ocean since 1948, and interpretation of long-term changes in the Phaeocystis hotspot in the North Sea

Monitoring of Phaeocystis since 1948 during the Continuous Plankton Recorder survey indicates that over the last 5.5 decades the distribution of its colonies in the North Atlantic Ocean was not restricted to neritic waters: occurrence was also recorded in the open Atlantic regions sampled, most frequently in the spring. Apparently, environmental conditions in open ocean waters, also those far oVshore, are suitable for complete lifecycle development of colonies (the only stage recorded in the survey). In the North Sea the frequency of occurrence was also highest in spring. Its southeastern part was the Phaeocystis abundance hotspot of the whole area covered by the survey. Frequency was especially high before the 1960s and after the 1980s, i.e., in the periods when anthropogenic nutrient enrichment was relatively low. Changes in eutrophication have obviously not been a major cause of long-term Phaeocystis variation in the southeastern North Sea, where total phytoplankton biomass was related signiWcantly to river discharge. Evidence is presented for the suggestion that Phaeocystis abundance in the southern North Sea is to a large extent determined by the amount of Atlantic Ocean water Xushed in through the Dover Strait. Since Phaeocystis plays a key role in element Xuxes relevant to climate the results presented here have implications for biogeochemical models of cycling of carbon and sulphur. Sea-to-air exchange of CO2 and dimethyl sulphide (DMS) has been calculated on the basis of measurements during single-year cruises. The considerable annual variation in phytoplankton and in its Phaeocystis component reported here does not warrant extrapolation of such figures.

Macroecology of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and adjacent seas

Global climate change is expected to modify the spatial distribution of marine organisms. However, projections of future changes should be based on robust information on the ecological niche of species. This paper presents a macroecological study of the environmental tolerance and ecological niche (sensu Hutchinson 1957, i.e. the field of tolerance of a species to the principal factors of its environment) of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and adjacent seas. Biological data were collected by the Continuous Plankton Recorder (CPR) Survey, which samples plankton in the North Atlantic and adjacent seas at a standard depth of 7 m. Eleven parameters were chosen including bathymetry, temperature, salinity, nutrients, mixed-layer depth and an index of turbulence compiled from wind data and chlorophyll a concentrations (used herein as an index of available food). The environmental window and the optimum level were determined for both species and for each abiotic factor and chlorophyll concentration. The most important parameters that influenced abundance and spatial distribution were temperature and its correlates such as oxygen and nutrients. Bathymetry and other water-column-related parameters also played an important role. The ecological niche of C. finmarchicus was larger than that of C. helgolandicus and both niches were significantly separated. Our results have important implications in the context of global climate change. As temperature (and to some extent stratification) is predicted to continue to rise in the North Atlantic sector, changes in the spatial distribution of these 2 Calanus species can be expected. Application of this approach to the 1980s North Sea regime shift provides evidence that changes in sea temperature alone could have triggered the substantial and rapid changes identified in the dynamic regimes of these ecosystems. C. finmarchicus appears to be a good indicator of the Atlantic Polar Biome (mainly the Atlantic Subarctic and Arctic provinces) while C. helgolandicus is an indicator of more temperate waters (Atlantic Westerly Winds Biome) in regions characterised by more pronounced spatial changes in bathymetry.