Ocean climate anomalies and the ecology of the North Sea

Long-term changes in the plankton of the North Sea are investigated using data from the continuous plankton recorder (CPR) survey. During the last 4 decades, there appears to have been 2 large anomalous periods within the plankton data set, one that occurred in the late 1970s and the other in the late 1980s. These anomalous periods seem to be largely synchronous with unusual ocean climate conditions that have occurred episodically over a timescale of decades. The unusual ocean climate conditions prevailing at these 2 time periods appear to contain important hydrographical elements that involve oceanic incursions into the North Sea. This paper, using data from the CPR survey and providing evidence from other studies, focuses on the relationship between the long-term changes in the biology of the North Sea and these 2 exceptional hydro-climatic events. Here, we suggest that while atmospheric variability and associated changes in regional temperatures have a dominant effect on the marine environment, oceanic influences on the ecology of a semi-closed environment such as the North Sea may have been underestimated in the past.

Decadal basin-scale changes in diatoms, dinoflagellates, and phytoplankton color across the North Atlantic

The Continuous Plankton Recorder (CPR) survey has been used to characterize phytoplankton and zooplankton space-time dynamics in the North Sea since 1931 and in the North Atlantic since 1939. Phytoplankton biomass is assessed from these samples by visual assessment of the green color of the silk mesh, the Phytoplankton Color Index (PCI), and the total count of diatoms and dinoflagellates. Species with a frequency of occurrence greater than 1% in the samples are used as indicator species of the community. We investigated (1) long-term fluctuations of phytoplankton biomass, total diatoms, and total dinoflagellates; (2) geographical variation of patterns; (3) the relationship between phytoplankton and climate forcing in the North Atlantic CPR samples; (4) the relative contribution of diatoms and dinoflagellates to the PCI; and (5) the fluctuations of the dominant species over the period of survey to provide more information on the processes linking climate to changes in the phytoplankton community. As a result of the differences in microscopic analysis methods prior to 1958, our analyses were conducted for the period ranging from 1958 to 2002. The North Atlantic was divided into six regions identified through bathymetric criteria and separated along a North-South axis. Based on 12 monthly time series, we demonstrate increasing trends in PCI and total dinoflagellates and a decrease in total diatoms.

Differential contribution of diatoms and dinoflagellates to phytoplankton biomass in the NE Atlantic Ocean and the North Sea

Sampling by the continuous plankton recorder (CPR) survey over the North Atlantic Ocean and the North Sea has enabled long-term studies of phytoplankton biomass. Analysis of an index of phytoplankton biomass, the phytoplankton colour index (PCI), has previously shown an increase in phytoplankton biomass in the NE Atlantic. In the current study, further investigations were conducted to determine the contribution of diatom and dinoflagellate cell counts to the PCI, their fluctuations over the last 45 yr and their geographical variations in the eastern North Atlantic and the North Sea. An increased contribution of dinoflagellates to the PCI was revealed over the south NE Atlantic and the northern North Sea. In contrast, the contribution of diatoms decreased in the north NE Atlantic and the northern North Sea. No discernible trends were found in the other regions of the North Sea. The relative contributions of diatoms and dinoflagellates to the PCI led to the identification of 3 geographically distinct dynamic regimes in the diatom/dinoflagellate dynamics in the NE Atlantic and the North Sea. Finally, it is stressed that the discrepancy observed in the patterns of PCI and diatom and dinoflagellate cell counts suggests that changes in PCI do not reflect changes in the community structure and that the exclusive use of PCI is not adequate to investigate the long-term trends in the trophic link between phytoplankton and herbivorous zooplankton.

An individual-based model of the early life history of mackerel (Scomber scombrus) in the eastern North Atlantic, simulating transport, growth and mortality

The main purpose of this paper is to provide the core description of the modelling exercise within the Shelf Edge Advection Mortality And Recruitment (SEAMAR) programme. An individual-based model (IBM) was developed for the prediction of year-to-year survival of the early life-history stages of mackerel (Scomber scombrus) in the eastern North Atlantic. The IBM is one of two components of the model system. The first component is a circulation model to provide physical input data for the IBM. The circulation model is a geographical variant of the HAMburg Shelf Ocean Model (HAMSOM). The second component is the IBM, which is an i-space configuration model in which large numbers of individuals are followed as discrete entities to simulate the transport, growth and mortality of mackerel eggs, larvae and post-larvae. Larval and post-larval growth is modelled as a function of length, temperature and food distribution; mortality is modelled as a function of length and absolute growth rate. Each particle is considered as a super-individual representing 10 super(6) eggs at the outset of the simulation, and then declining according to the mortality function. Simulations were carried out for the years 1998-2000. Results showed concentrations of particles at Porcupine Bank and the adjacent Irish shelf, along the Celtic Sea shelf-edge, and in the southern Bay of Biscay. High survival was observed only at Porcupine and the adjacent shelf areas, and, more patchily, around the coastal margin of Biscay. The low survival along the shelf-edge of the Celtic Sea was due to the consistently low estimates of food availability in that area.

Spawning season and temperature relationships for sardine (Sardina pilchardus) in the eastern North Atlantic

Spawning temperature preferences for sardine (Sardina pilchardus) in the eastern North Atlantic were determined from field data. These were compared with climatological temperature cycles (1986-2002) derived from satellite data by geographical region, to predict spawning seasons. Optimum spawning temperatures were determined as 14.0-15.0oC from the English Channel to Portugal and 16.0–18.0oC for all north-west African regions. Spawning seasons were closely related to the general latitudinal trend of the annual temperature cycle, with modification by upwelling in the western Iberian and north-west African regions. Some differences between temperature-based spawning season predictions and field observations were related to variations in seasonal plankton production. Correlations in the annual time-series of favourable spawning temperatures suggested relatively strong linkages between the southern areas from Portugal to Senegal. There was no consistent relationship between annual variations in duration of temperature-predicted spawning seasons and observed field abundance of eggs.

Is there a connection between high transport of water through the Rockall Trough and ecological changes in the North Sea?

Changes in the ecosystem of the North Sea may occur as pronounced inter-annual and step-wise shifts as well as gradual trends. Marked inter-annual shifts have occurred at least twice in the last two decades, the late 1980s and the late 1990s, that appear to reflect an increased inflow of oceanic water and species. Numerical modelling has demonstrated a link between altered rates of inflow of oceanic water into the northern North Sea and a regime shift after 1988. In 1989 and 1997 oceanic species not normally found in the North Sea were observed there, suggesting pulses of oceanic water had entered the basin and triggered the subsequent ecosystem change. The oceanic water has origins mainly west of Britain in the Rockall Trough, where the long-term mean volume transport is around 3.7Sv northwards (1Sv=10 super(6)m super(3)s super(1)), but in early 1989 and early 1998 was observed to be more than twice the mean value, reaching over 7Sv. These periods of high transport coinciding with the inferred pulses of oceanic water into the North Sea suggest a connection through the continental shelf edge current. Copyright 2001 International Council for the Exploration of the Sea

The CPR Survey (1948-1997): a gridded database browser of plankton abundance in the North Sea.

The Continuous Plankton Recorder (CPR) survey provides a unique multi- decadal dataset on the abundance of plankton in the North Sea and North Atlantic and is one of only a few monitoring programmes operating at a large spatio- temporal scale. The results of all samples analysed from the survey since 1946 are stored on an Access Database at the Sir Alister Hardy Foundation for Ocean Science (SAHFOS) in Plymouth. The database is large, containing more than two million records (~80 million data points, if zero results are added) for more than 450 taxonomic entities. An open data policy is operated by SAHFOS. However, the data are not on-line and so access by scientists and others wishing to use the results is not interactive. Requests for data are dealt with by the Database Manager. To facilitate access to the data from the North Sea, which is an area of high research interest, a selected set of data for key phytoplankton and zooplankton species has been processed in a form that makes them readily available on CD for research and other applications. A set of MATLAB tools has been developed to provide an interpolated spatio-temporal description of plankton sampled by the CPR in the North Sea, as well as easy and fast access to users in the form of a browser. Using geostatistical techniques, plankton abundance values have been interpolated on a regular grid covering the North Sea. The grid is established on centres of 1 degree longitude x 0.5 degree latitude (~32 x 30 nautical miles). Based on a monthly temporal resolution over a fifty-year period (1948-1997), 600 distribution maps have been produced for 54 zooplankton species, and 480 distribution maps for 57 phytoplankton species over the shorter period 1958-1997. The gridded database has been developed in a user-friendly form and incorporates, as a package on a CD, a set of options for visualisation and interpretation, including the facility to plot maps for selected species by month, year, groups of months or years, long-term means or as time series and contour plots. This study constitutes the first application of an easily accessed and interactive gridded database of plankton abundance in the North Sea. As a further development the MATLAB browser is being converted to a user- friendly Windows-compatible format (WinCPR) for release on CD and via the Web in 2003.

Congruent trends in long-term zooplankton decline in the north-east Atlantic and basking shark (Cetorhinus maximus ) fishery catches off west Ireland

Trends in basking shark (Cetorhinus maximus) fishery catches off Achill Island, west Ireland between 1949 and 1975 were examined in relation to zooplankton (total copepod) abundance in four adjacent sea areas over a 27-year period. The numbers of basking sharks caught and copepod abundance showed downward trends and were positively correlated (r-value range, 0.44–0.74). A possible explanation for the downward trend in shark catches was that progressively fewer basking sharks occurred there between 1956 and 1975 because fewer copepods, their food resource, occurred near the surface off west Ireland over the same period. We suggest that the decline in basking sharks may have been due to a distributional shift of sharks to more productive areas, rather than a highly philopatric, localized stock that was over-exploited.

Pulses in the eastern margin current and warmer water off the north west European shelf linked to North Sea ecosystem changes

The North Sea ecosystem has recently undergone dramatic changes, observed as altered biomass of individual species spanning a range of life forms from algae to birds, with evidence for an approximate doubling in the abundance of both phytoplankton and benthos as part of a regime shift after 1987. Remarkably, these changes, in part recorded in the Phytoplankton Colour Index of the Continuous Plankton Recorder (CPR) survey, are notable as episodic shifts occurring in 1988/89 and 1998 imposed on a gradual decadal trend. These biological events are shown to be a response to coincident changes in oceanic input and water temperature. Geostrophic transports have been calculated from a hydrographic section across the Rockall Trough, and a time series of seasurface temperature derived from satellite observations. The 2 pulses of oceanic incursion into the North Sea in circa 1988 and 1998 coincided with strong northward advection of anomalously warm water at the edge of the continental shelf.

A regime shift in the North Sea circa 1988 linked to changes in the North Sea horse mackerel fishery

After 1987, Phytoplankton Colour (a visual estimate of chlorophyll) measured on samples taken by the continuous plankton recorder (CPR) in the North Sea increased substantially, both in level and seasonal extent, compared to earlier years since 1946. Many species of phytoplankton and zooplankton showed marked changes in abundance at about the same time. These events coincided with a large increase in catches of the western stock of the horse mackerel (Trachurus trachurus L.) in the northern North Sea reflecting a northerly expansion of the stock along the shelf edge from the Bay of Biscay to the North Sea after 1987. Using a 3D hydrodynamic model, with input from measured wind parameters, monthly transport of oceanic water into the North Sea has been calculated for the period 1976–1994, integrated for a section from Orkney to Shetland to Norway. A substantial increase in oceanic inflow occurred in the winter months, December to March, from 1988. Higher sea surface temperatures were also measured after 1987 especially in spring and summer months. These biological and physical events may be a response to observed changes in pressure distribution over the North Atlantic. From 1988 onwards, the North Atlantic Oscillation (NAO) index, the pressure difference between Iceland and the Azores, increased to the highest positive level observed in this century. Positive NAO anomalies are associated with stronger and more southerly tracks of the westerly winds and higher temperatures in western Europe. These changing wind distributions may have led to an increase in the northerly advection of water along the western edge of the European shelf and may have assisted the migration of the horse mackerel. This study is possibly a unique demonstration of a correlation between three different trophic levels of a marine ecosystem and hydrographic and atmospheric events at decadal and regional scales. The results emphasise the importance of maintaining into the future long term programmes such as the CPR.

Ecological effects of the North Atlantic oscillation

Climatic oscillations as reflected in atmospheric modes such as the North Atlantic Oscillation (NAO) may be seen as a proxy for regulating forces in aquatic and terrestrial ecosystems. Our review highlights the variety of climate processes related to the NAO and the diversity in the type of ecological responses that different biological groups can display. Available evidence suggests that the NAO influences ecological dynamics in both marine and terrestrial systems, and its effects may be seen in variation at the individual, population and community levels. The ecological responses to the NAO encompass changes in timing of reproduction, population dynamics, abundance, spatial distribution and interspecific relationships such as competition and predator-prey relationships. This indicates that local responses to large-scale changes may be more subtle than previously suggested. We propose that the NAO effects may be classified as three types: direct, indirect and integrated. Such a classification will help the design and interpretation of analyses attempting to relate ecological changes to the NAO and, possibly, to climate in general.

Biodiversity of North Atlantic and North Sea calanoid copepods

Spatial patterns in pelagic biodiversity are the result of factors acting from a global to a local scale. The global patterns have been studied intensively using taxa such as foraminifera and euphausiids. However, these studies do not allow direct comparisons of neritic and oceanic regions or examination of relationships between local and regional patterns of pelagic diversity. Here we present a map of the diversity of calanoid copepods, a key planktonic group, summarising 40 yr of continuous monthly investigations in the North Atlantic and North Sea. The high number of samples (168 162) allowed mesoscale patterns in diversity to be detected for the first time at an ocean-basin level. Our results demonstrate pronounced local spatial variability in planktonic diversity and refine previous global studies at a lower resolution. They form a baseline at which long-term changes in planktonic diversity can be better assessed and ecosystem management plans implemented.