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.

An overview of the literature concerning the oceanography of the eastern North Atlantic region

An overview of the main oceanographic features of the eastern North Atlantic boundary, with emphasis toward the upper layers, is presented. The principal features discussed are: water mass boundaries; forcing by wind, density and tides; topographic features and effects; fronts; upwelling and downwelling; poleward flows; coastal currents; eddies. The occurrence and spatial and seasonal variability of these features is described in five regional sections: Celtic Sea and western English Channel; Bay of Biscay; western Iberia; Gulf of Cadiz; northwest Africa. This paper is intended to provide a base of physical oceanographic knowledge in support of research in fisheries, biological and chemical oceanography, and marine biology.

A change in the zooplankton of the central North Sea (55 degree to 58 degree N): A possible consequence of changes in the benthos

The mesozooplankton taken in continuous plankton recorder samples from the Central North Sea has changed from being numerically dominated by holoplanktonic calanoid copepod species from 1958 to the late 1970s to a situation where pluteus larvae of echinoid and ophiuroid echinoderms have been more abundant than any single holoplanktonic species in the 1980s and early 1990s. The abundance of the echinoderm larvae as a proportion of the zooplankton taken in the samples has followed a continuous increasing trend over the Dogger Bank, but off the eastern coast of northern England and southern Scotland the increase did not become obvious until the 1980s. This trend is consistent with reported increases in abundance of the macrobenthos. It is proposed that changes in the benthos have influenced the composition of the plankton.

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.

Coccolithophore bloom size variation in response to the regional environment of the subarctic North Atlantic

Several environmental/physical variables derived from satellite and in situ data sets were used to understand the variability of coccolithophore abundance in the subarctic North Atlantic. The 7-yr (1997–2004) time-series analysis showed that the combined effects of high solar radiation, shallow mixed layer depth (<20 m), and increased temperatures explained >89% of the coccolithophore variation. The June 1998 bloom, which was associated with high light intensity, unusually high sea-surface temperature, and a very shallow mixed layer, was found to be one of the most extensive (>995,000 km2) blooms ever recorded. There was a pronounced sea-surface temperature shift in the mid-1990s with a peak in 1998, suggesting that exceptionally large blooms are caused by pronounced environmental conditions and the variability of the physical environment strongly affects the spatial extent of these blooms. Consequently, if the physical environment varies, the effects of these blooms on the atmospheric and oceanic environment will vary as well.

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.