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.

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.

Rapid biogeographical plankton shifts in the North Atlantic Ocean

Large-scale biogeographical changes in the biodiversity of a key zooplankton group (calanoid copepods) were detected in the north-eastern part of the North Atlantic Ocean and its adjacent seas over the period 1960–1999. These findings provided key empirical evidence for climate change impacts on marine ecosystems at the regional to oceanic scale. Since 1999, global temperatures have continued to rise in the region. Here, we extend the analysis to the period 1958–2005 using all calanoid copepod species assemblages (nine species assemblages based on an analysis including a total of 108 calanoid species or taxa) and show that this phenomenon has been reinforced in all regions. Our study reveals that the biodiversity of calanoid copepods are responding quickly to sea surface temperature (SST) rise by moving geographically northward at a rapid rate up to about 23.16 km yr−1. Our analysis suggests that nearly half of the increase in sea temperature in the northeast Atlantic and adjacent seas is related to global temperature rises (46.35% of the total variance of temperature) while changes in both natural modes of atmospheric and oceanic circulation explain 26.45% of the total variance of temperature. Although some SST isotherms have moved northwards by an average rate of up to 21.75 km yr−1 (e.g. the North Sea), their movement cannot fully quantify all species assemblage shifts. Furthermore, the observed rates of biogeographical movements are far greater than those observed in the terrestrial realm. Here, we discuss the processes that may explain such a discrepancy and suggest that the differences are mainly explained by the fluid nature of the pelagic domain, the life cycle of the zooplankton and the lesser anthropogenic influence (e.g. exploitation, habitat fragmentation) on these organisms. We also hypothesize that despite changes in the path and intensity of the oceanic currents that may modify quickly and greatly pelagic zooplankton species, these organisms may reflect better the current impact of climate warming on ecosystems as terrestrial organisms are likely to significantly lag the current impact of climate change.

Causes and prediction of abrupt climate-driven ecosystem shifts in the North Atlantic

Warming of the global climate is now unequivocal and its impact on Earth’ functional units has become more apparent. Here, we show that marine ecosystems are not equally sensitive to climate change and reveal a critical thermal boundary where a small increase in temperature triggers abrupt ecosystem shifts seen across multiple trophic levels. This large-scale boundary is located in regions where abrupt ecosystem shifts have been reported in the North Atlantic sector and thereby allows us to link these shifts by a global common phenomenon. We show that these changes alter the biodiversity and carrying capacity of ecosystems and may, combined with fishing, precipitate the reduction of some stocks of Atlantic cod already severely impacted by exploitation. These findings offer a way to anticipate major ecosystem changes and to propose adaptive strategies for marine exploited resources such as cod in order to minimize social and economic consequences.

On the processes linking climate to ecosystem changes

While documentation of climate effects on marine ecosystems has a long history, the underlying processes have often been elusive. In this paper we review some of the ecosystem responses to climate variability and discuss the possible mechanisms through which climate acts. Effects of climatological and oceanographic variables, such as temperature, sea ice, turbulence, and advection, on marine organisms are discussed in terms of their influence on growth, distribution, reproduction, activity rates, recruitment and mortality. Organisms tend to be limited to specific thermal ranges with experimental findings showing that sufficient oxygen supply by ventilation and circulation only occurs within these ranges. Indirect effects of climate forcing through effects on the food web are also discussed. Research and data needs required to improve our knowledge of the processes linking climate to ecosystem changes are presented along with our assessment of our ability to predict ecosystem responses to future climate change scenarios. (C) 2009 Elsevier B.V. All rights reserved.

Predator–prey reversal: A possible mechanism for ecosystem hysteresis in the North Sea?

Removal of large predatory fishes from marine ecosystems has resulted in persistent ecosystem shifts, with collapsed predator populations and super-abundant prey populations. One explanation for these shifts is reversals of predator–prey roles that generate internal feedbacks in the ecosystems. Pelagic forage fish are often predators and competitors to the young life stages of their larger fish predators. I show that cod recruitment in the North Sea has been negatively related to the spawning-stock biomass of herring for the last 44 years. Herring, together with the abundance of Calanus finmarchicus, the major food for cod larvae, were the main predictors of cod recruitment. These predictors were of equivalent importance, worked additively, and explained different parts of the dynamics in cod recruitment. I suggest that intensive harvesting of cod has released herring from predator control, and that a large population of herring suppresses cod recruitment through predation on eggs and larvae. This feedback mechanism can promote alternative stable states and therefore cause hysteresis to occur under changing conditions; however, harvesting of herring might at present prevent a shift in the ecosystem to a herring-dominated state.

Regional and annual variation in black-legged kittiwake breeding productivity is related to sea surface temperature

Spatiotemporal variation in seabird demographic parameters is often pronounced and may be an important source of information on the state of marine ecosystems. Black-legged kittiwakes Rissa tridactyla in Britain and Ireland show strong regional structure in breeding productivity, and both temporal and spatial variation are probably related to abundance of the principal prey of breeding kittiwakes, the lesser sandeel Ammodytes marinus. Annual regional estimates of sandeel abundance do not exist, prohibiting direct tests of this hypothesis. We examined relationships between kittiwake breeding productivity and 2 potential proxies of sandeel abundance, winter sea surface temperature (SST) and abundance of Calanus copepods, within and among 6 regions in Britain and Ireland from 1986 to 2004. Means and trends in winter SST differed among regions, with higher means and less pronounced increasing trends in western (Atlantic) regions than in eastern (North Sea) regions. A negative relationship between breeding productivity and winter SST in the previous year was found within 2 regions (East Scotland and Orkney), as well as in a cross-regional analysis. Results were inconclusive for Calanus abundance, with a positive relationship in East Scotland and negative in Orkney. These results demonstrate that although a single environmental driver (SST) is related to both within- and between-region variation in a key demographic parameter, regional heterogeneity in SST trends as well as the importance of other factors may lead to highly variable responses. Understanding this heterogeneity is critical for predicting long-term effects of climate change or other anthropogenic drivers on marine ecosystems.

Spatial and temporal variability in Scotian Shelf zooplankton communities

Zooplankton are indicators of the response of marine ecosystems to environmental variability. The relationships between zooplankton communities on the Scotian Shelf and hydrographic and geographic regions of the Scotian Shelf in the 1990s and 2000s were described using complementary data sets, each resolving different space and time scales. The Atlantic Zone Monitoring Program (AZMP) sampled Scotian Shelf zooplankton from the whole water column twice per year at stations along three cross-shelf transects and semi-monthly at a fixed station on the inshore central shelf, while Continuous Plankton Recorder (CPR) samples were collected from near-surface waters approximately monthly on an along-shelf transect. Variability patterns were compared among these three data sets to identify robust spatial and interannual trends. Stations were clustered based on taxonomic composition, and spatial clusters were compared to hydrographic boundaries and bathymetry to determine whether temporal changes in community composition were driven by changes in water mass distributions on the shelf. This project identifies zooplankton community and abundance shifts that may affect fish recruitment in the northwest Atlantic and contributes to development of ecosystem-based fisheries management on the Scotian Shelf.

Trophic amplification of climate warming

Ecosystems can alternate suddenly between contrasting persistent states due to internal processes or external drivers. It is important to understand the mechanisms by which these shifts occur, especially in exploited ecosystems. There have been several abrupt marine ecosystem shifts attributed either to fishing, recent climate change or a combination of these two drivers. We show that temperature has been an important driver of the trophodynamics of the North Sea, a heavily fished marine ecosystem, for nearly 50 years and that a recent pronounced change in temperature established a new ecosystem dynamic regime through a series of internal mechanisms. Using an end-to-end ecosystem approach that included primary producers, primary, secondary and tertiary consumers, and detritivores, we found that temperature modified the relationships among species through nonlinearities in the ecosystem involving ecological thresholds and trophic amplifications. Trophic amplification provides an alternative mechanism to positive feedback to drive an ecosystem towards a new dynamic regime, which in this case favours jellyfish in the plankton and decapods and detritivores in the benthos. Although overfishing is often held responsible for marine ecosystem degeneration, temperature can clearly bring about similar effects. Our results are relevant to ecosystem-based fisheries management (EBFM), seen as the way forward to manage exploited marine ecosystems.

Influence of climate change and trophic coupling across four trophic levels in the Celtic Sea

Climate change has had profound effects upon marine ecosystems, impacting across all trophic levels from plankton to apex predators. Determining the impacts of climate change on marine ecosystems requires understanding the direct effects on all trophic levels as well as indirect effects mediated by trophic coupling. The aim of this study was to investigate the effects of climate change on the pelagic food web in the Celtic Sea, a productive shelf region in the Northeast Atlantic. Using long-term data, we examined possible direct and indirect ‘bottom-up’ climate effects across four trophic levels: phytoplankton, zooplankton, mid-trophic level fish and seabirds. During the period 1986–2007, although there was no temporal trend in the North Atlantic Oscillation index (NAO), the decadal mean Sea Surface Temperature (SST) in the Celtic Sea increased by 0.66±0.02°C. Despite this, there was only a weak signal of climate change in the Celtic Sea food web. Changes in plankton community structure were found, however this was not related to SST or NAO. A negative relationship occurred between herring abundance (0- and 1-group) and spring SST (0-group: p = 0.02, slope = −0.305±0.125; 1-group: p = 0.04, slope = −0.410±0.193). Seabird demographics showed complex species–specific responses. There was evidence of direct effects of spring NAO (on black-legged kittiwake population growth rate: p = 0.03, slope = 0.0314±0.014) as well as indirect bottom-up effects of lagged spring SST (on razorbill breeding success: p = 0.01, slope = −0.144±0.05). Negative relationships between breeding success and population growth rate of razorbills and common guillemots may be explained by interactions between mid-trophic level fish. Our findings show that the impacts of climate change on the Celtic Sea ecosystem is not as marked as in nearby regions (e.g. the North Sea), emphasizing the need for more research at regional scales.

Comparisons of zooplankton time series

Evidence for climate-correlated low frequency variability of various components of marine ecosystems has accumulated rapidly over the past 2 decades. There has also been a growing recognition that society needs to learn how the fluctuations of these various components are linked, and to predict the likely amplitude and steepness of future changes. Demographic characteristics of marine zooplankton make them especially suitable for examining variability of marine ecosystems at interannual to decadal time scales. Their life cycle duration is short enough that there is little carryover of population membership from year to year, but long enough that variability can be tracked with monthly-to-seasonal sampling. Because zooplankton are rarely fished, comparative analysis of changes in their abundance can greatly enhance our ability to evaluate the importance of and interaction between physical environment, food web, and fishery harvest as causal mechanisms driving ecosystem level changes. A number of valuable within-region analyses of zooplankton time series have been published in the past decade, covering a variety of modes of variability including changes in total biomass, changes in size structure and species composition, changes in spatial distribution, and changes in seasonal timing. But because most zooplankton time series are relatively short compared to the time scales of interest, the statistical power of local analyses is often low, and between-region and between-variable comparisons are also needed. In this paper, we review the results of recent within- and between-region analyses, and suggest some priorities for future work.

High dispersal potential has maintained long-term population stability in the North Atlantic copepod Calanus finmarchicus

The cool-water copepod Calanus finmarchicus is a key species in North Atlantic marine ecosystems since it represents an important food resource for the developmental stages of several fish of major economic value. Over the last 40 years, however, data from the Continuous Plankton Recorder survey have highlighted a 70 per cent reduction in C. finmarchicus biomass, coupled with a gradual northward shift in the species's distribution, which have both been linked with climate change. To determine the potential for C. finmarchicus to track changes in habitat availability and maintain stable effective population sizes, we have assessed levels of gene flow and dispersal in current populations, as well as using a coalescent approach together with palaeodistribution modelling to elucidate the historical population demography of the species over previous changes in Earth's climate. Our findings indicate high levels of dispersal and a constant effective population size over the period 359 000–566 000 BP and suggest that C. finmarchicus possesses the capacity to track changes in available habitat, a feature that may be of crucial importance to the species's ability to cope with the current period of global climate change.