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

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.

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.

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.

North Atlantic climatic signals and the plankton of the European Continental Shelf

Climatic variability on the European Continental Shelf is dominated by events over the North Atlantic Ocean, and in particular by the North Atlantic Oscillation (NAO). The NAO is essentially a winter phenomenon, and its effects will be felt most strongly by populations for which winter conditions are critical. One example is the copepod Calanus finmarchicus, whose northern North Sea populations overwinter at depth in the North Atlantic. Its annual abundance in this region is strongly dependent on water transports at the end of the winter, and hence on the NAO index. Variations in the NAO give rise to changes in the circulation of the North Atlantic Ocean, with additional perturbations arising from El Ni (n) over tildeo - Southern Oscillation (ENSO) events in the Pacific, and these changes can be delayed by several years because of the adjustment time of the ocean circulation. One measure of the circulation is the latitude of the north wall of the Gulf Stream (GSNW index). Interannual variations in the plankton of the Shelf Seas show strong correlations with the fluctuations of the GSNW index, which are the result of Atlantic-wide atmospheric processes. These associations imply that the interannual variations are climatically induced rather than due to natural fluctuations of the marine ecosystem, and that the zooplankton populations have not been significantly affected by anthropogenic processes such as nutrient enrichment or fishing pressure. While the GSNW index represents a response to atmospheric changes over two or more years, the zooplankton populations correlated with it have generation times of a few weeks. The simplest explanation for the associations between the zooplankton and the GSNW index is that the plankton are responding to weather patterns propagating downstream from the Gulf Stream system. It seems that these meteorological processes operate in the spring. Although it has been suggested that there was a regime shift in the North Sea in the late 1980s, examination of the time-series by the cumulative sum (CUSUM) technique shows that any changes in the zooplankton of the central and northern North Sea are consistent with the background climatic variability. The abundance of total copepods increased during this period but this change does not represent a dramatic change in ecosystem processes. It is possible some change may have occurred at the end of the time-series in the years 1997 and 1998.

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.

Relationships between North Atlantic salmon, plankton and hydroclimatic change in the Northeast Atlantic

The abundance of wild salmon (Salmo salar) in the North Atlantic has declined markedly since the late 1980s as a result of increased marine mortality that coincided with a marked rise in sea temperature in oceanic foraging areas. There is substantial evidence to show that temperature governs the growth, survival, and maturation of salmon during their marine migrations through either direct or indirect effects. In an earlier study (2003), long-term changes in three trophic levels (salmon, zooplankton, and phytoplankton) were shown to be correlated significantly with sea surface temperature (SST) and northern hemisphere temperature (NHT). A sequence of trophic changes ending with a stepwise decline in the total nominal catch of North Atlantic salmon (regime shift in ∼1986/1987) was superimposed on a trend to a warmer dynamic regime. Here, the earlier study is updated with catch and abundance data to 2010, confirming earlier results and detecting a new abrupt shift in ∼1996/1997. Although correlations between changes in salmon, plankton, and temperature are reinforced, the significance of the correlations is reduced because the temporal autocorrelation of time-series substantially increased due to a monotonic trend in the time-series, probably related to global warming. This effect may complicate future detection of effects of climate change on natural systems.

Monitoring marine plankton ecosystems: Identification of the most relevant indicators of the state of an ecosystem

The number of variables involved in the monitoring of an ecosystem can be high and often one of the first stages in the analysis is to reduce the number of variables. We describe a method developed for geological purposes, using the information theory, that enables selection of the most relevant variables. This technique also allows the examination of the asymmetrical relationships between variables. Applied to a set of physical and biological variables (plankton assemblages in four areas of the North Sea), the method shows that biological variables are more informative than physical variables although the controlling factors are mainly physical (sea surface temperature in winter and spring). Among biological variables, diversity measures and warm-water species assemblages are informative for the state of the North Sea pelagic ecosystems while among physical variables sea surface temperature in late winter and early spring are highly informative. Although often used in bioclimatology, the utilisation of the North Atlantic Oscillation (NAO) index does not seem to provide a lot of information. The method reveals that only the extreme states of this index has an influence on North Sea pelagic ecosystems. The substantial and persistent changes that were detected in the dynamic regime of the North Sea ecosystems and called regime shift are detected by the method and corresponds to the timing of other shifts described in the literature for some European Systems such as the Baltic and the Mediterranean Sea when both physical and biological variables are considered.

Climate-induced changes in the North Sea Decapoda over the last 60 years

In the mid-1980s the North Sea ecosystem experienced a climate-induced regime shift that has favoured decapods and detritivores in the benthos and jellyfish in the plankton over commercial fisheries. Here, we investigate changes among the Decapoda in the North Sea plankton over the last 60 yr. Decapods are important predators in the plankton and the benthos where they can influence productivity and structure communities. In the North Sea it has been suggested that a climate-driven increase in decapod abundance has been important in propagating the climate signal through the North Sea food web. We show that climate-induced changes in the Decapoda in the central and southern North Sea include the presence of new warm-water taxa, changes in the abundance and proportions of commercial species of shrimp, and an earlier occurrence of decapod larvae in the plankton compared with the period 1981–1983. Notable amongst the warm-water taxa appearing in the North Sea is the predatory swimming crab Polybius henslowii that can swarm in large numbers when conditions are favourable and that is known to exhibit range shifts in response to fluctuations in hydroclimatic forcing. We suggest that climate-induced changes among North Sea decapods have played an important role in the trophic amplification of a climate signal and the development of the new North Sea dynamic regime. Understanding these changes is likely to be imperative for a successful ecosystem-based approach to the future management of North Sea fisheries at a time of climate change.

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.

Covariance among North Sea ecosystem state indicators during the past 50 years - contrasts between coastal and open waters

Regime shift and principal component analysis of a spatially disaggregated database capturing time-series of climatic, nutrient and plankton variables in the North Sea revealed considerable covariance between groups of ecosystem indicators. Plankton and climate time-series span the period 1958–2003, those of nutrients start in 1980. In both regions, the period from 1989 to 2001 identified in principal component 1 had warmer surface waters, higher Atlantic inflow and stronger winds, than the periods before or after. However, it was preceded by a regime shift in both open (PC2) and coastal (PC3) waters during 1977 towards more hours of sunlight and higher water temperature, which lasted until 1997. The relative influence of nutrient availability and climatic forcing differed between open and coastal North Sea regions. Inter-annual variability in phytoplankton dynamics of the open North Sea was primarily regulated by climatic forcing, specifically by sea surface temperature, Atlantic inflow and co-varying wind stress and NAO. Coastal phytoplankton variability, however, was regulated by insolation and sea surface temperature, as well as Si availability, but not by N or P. Regime shifts in principal components of hydrographic and climatic variables (explaining 55 and 61% of the variance in coastal and open water variables) were detected using Rodionov's sequential t-test. These shifts in hydroclimatic variables which occurred around 1977, 1989, 1997 and 2001, were synchronized in open and coastal waters, and were tracked by open water chlorophyll and copepods, but not by coastal plankton. North–central–south or open-coastal spatial breakdowns of the North Sea explained similar amounts of variability in most ecosystem indicators with the exception of diatom abundance and chlorophyll concentration, which were clearly better explained using the open-coastal configuration.

Recruitment in a changing environment: the 2000s North Sea herring recruitment failure

Environmentally induced change appears to be impacting the recruitment of North Sea herring (Clupea harengus). Despite simultaneously having a large adult population, historically low exploitation, and Marine Stewardship Council accreditation (implying sustainability), there have been an unprecedented 6 sequential years of poor juvenile production (recruitment). Analysis suggests that the poor recruitment arises during the larval overwintering phase, with recent survival rates greatly reduced. Contemporary warming of the North Sea has caused significant changes in the plankton community, and a recently identified regime shift around 2000 shows close temporal agreement with the reduced larval survival. It is, therefore, possible that we are observing the first consequences of this planktonic change for higher trophic levels. There is no indication of a recovery in recruitment in the short term. Fishing mortality is currently outside the agreed management plan, and forecasts show a high risk of the stock moving outside safe biological limits soon, potentially precipitating another collapse of the stock. However, bringing the realized fishing mortality back in line with the management plan would likely alleviate the problem. This illustrates again that recruitment is influenced by more than just spawning-stock biomass, and that changes in other factors can be of equal, or even greater, importance. In such dynamically changing environments, recent management success does not necessarily guarantee future sustainability.

Modelling the future biogeography of North Atlantic zooplankton communities in response to climate change

Advances in habitat and climate modelling allow us to reduce uncertainties of climate change impacts on species distribution. We evaluated the impacts of future climate change on community structure, diversity, distribution and phenology of 14 copepod species in the North Atlantic. We developed and validated habitat models for key zooplankton species using continuous plankton recorder (CPR) survey data collected at mid latitudes of the North Atlantic. Generalized additive models (GAMs) were applied to relate the occurrence of species to environmental variables. Models were projected to future (2080–2099) environmental conditions using coupled hydroclimatix–biogeochemical models under the Intergovernmental Panel on Climate Change (IPCC) A1B climate scenario, and compared to present (2001–2020) conditions. Our projections indicated that the copepod community is expected to respond substantially to climate change: a mean poleward latitudinal shift of 8.7 km per decade for the overall community with an important species range variation (–15 to 18 km per decade); the species seasonal peak is expected to occur 12–13 d earlier for Calanus finmarchicus and C. hyperboreus; and important changes in community structure are also expected (high species turnover of 43–79% south of the Oceanic Polar Front). The impacts of the change expected by the end of the century under IPCC global warming scenarios on copepods highlight poleward shifts, earlier seasonal peak and changes in biodiversity spatial patterns that might lead to alterations of the future North Atlantic pelagic ecosystem. Our model and projections are supported by a temporal validation undertaken using the North Atlantic climate regime shift that occurred in the 1980s: the habitat model built in the cold period (1970–1986) has been validated in the warm period (1987–2004).