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.

The North Pacific Continuous Plankton Recorder Survey

The Continuous Plankton Recorder (CPR) survey of the North Pacific is a PICES project now in its ninth year and facing an uncertain future. CPRs have been towed behind commercial ships along two (north–south and east–west) transects for a total of ~ nine times per year. Samples are collected with a filtering mesh and are then microscopically processed for plankton abundance in the laboratory. The survey, so far, has accumulated 3,648 processed samples (with approximately three times as many archived without processing), each representing 18 km of the transect (Fig. 1) and containing an abundance of data on over 290 phytoplankton and zooplankton taxa. A CTD with a fluorometer has been attached to the CPR sampling at the east–west transect in more recent years to provide supplementary environmental data.

Plankton populations at the bifurcation of the North Pacific Current

As the eastward-flowing North Pacific Current approaches the North American continent it bifurcates into the southward-flowing California Current and the northward-flowing Alaska Current. This bifurcation occurs in the south-eastern Gulf of Alaska and can vary in position. Dynamic height data from Project Argo floats have recently enabled the creation of surface circulation maps which show the likely position of the bifurcation; during 2002 it was relatively far north at 53 degrees N then, during early 2003, it moved southwards to a more normal position at 45 degrees N. Two ship-of-opportunity transects collecting plankton samples with a Continuous Plankton Recorder across the Gulf of Alaska were sampled seasonally during 2002 and 2003. Their position was dependent on the commercial ship's operations; however, most transects sampled across the bifurcation. We show that the oceanic plankton differed in community composition according to the current system they occurred in during spring and fall of 2002 and 2003, although winter populations were more mixed. Displacement of the plankton communities could have impacts on the plankton's reproduction and development if they use cues such as day length, and also on foraging of higher trophic-level organisms that use particular regions of the ocean if the nutritional value of the communities is different. Although we identify some indicator taxa for the Alaska and California currents, functional differences in the plankton communities on either side of the bifurcation need to be better established to determine the impacts of bifurcation movement on the ecosystems of the north-east Pacific.

Changes in development timing and cohort width of Neocalanus plumchrus degree ' flemingeri copepods in the eastern North Pacific

Neocalanus plumchrus/flemingeri copepods make up a large proportion of spring mesozooplankton biomass and are a valuable nutritional source for many higher trophic levels. Copepodites through to sub-adult stage are present in surface waters for a relatively short period of time each spring, and the date of maximum biomass has been calculated as the date when 50% of the population were at the sub-adult, CV stage. This index allows quite a precise date to be calculated from relatively infrequent sampling and interannual comparisons between 1957 and 2004 have demonstrated that the timing of peak abundance is significantly advanced in warmer years. However, recent data from the Continuous Plankton Recorder survey, which samples the surface NE Pacific more frequently during spring, has found that maximum numbers of CV copepodites occur after the 50% point is reached so that maximum biomass occurs some weeks later than predicted by this index (although comparisons between years show that the magnitude of the timing shift is similar). Comparisons with depth-stratified profiles from the BIONESS show that this is not just due to single-depth near-surface sampling by the CPR. We speculate on the cause of this change which could be related to the width of the cohort (which appears to now be narrower, at least in warm years) or the length of time that the CV stage needs to spend in the surface accumulating lipid before beginning diapause. A narrower cohort has implications for predators who will have less time to take advantage of this food source.

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.

Decadal changes in climate and ecosystems in the North Atlantic Ocean and adjacent seas

Climate change is unambiguous and its effects are clearly detected in all functional units of the Earth system. This study presents new analyses of sea-surface temperature changes and show that climate change is affecting ecosystems of the North Atlantic. Changes are seen from phytoplankton to zooplankton to fish and are modifying the dominance of species and the structure, the diversity and the functioning of marine ecosystems. Changes also range from phenological to biogeographical shifts and have involved in some regions of the Atlantic abrupt ecosystem shifts. These alterations reflect a response of pelagic ecosystems to a warmer temperature regime. Mechanisms are complex because they are nonlinear exhibiting tipping points and varying in space and time. Sensitivity of organisms to temperature changes is high, implicating that a small temperature modification can have sustained ecosystem effects. Implications of these changes for biogeochemical cycles are discussed. Two observed changes detected in the North Sea that could have opposite effects on carbon cycle are discussed. Increase in phytoplankton, as inferred from the phytoplankton colour index derived from the Continuous Plankton Recorder (CPR) survey, has been detected in the North Sea. This pattern has been accompanied by a reduction in the abundance of the herbivorous species Calanus finmarchicus. This might have reduced the grazing pressure and increase diatomaceous ‘fluff’, therefore carbon export in the North Sea. Therefore, it could be argued that the biological carbon pump might increase in this region with sea warming. In the meantime, however, the mean size of organisms (calanoid copepods) has dropped. Such changes have implications for the turnover time of biogenic carbon in plankton organisms and the mean residence time of particulate carbon they produce. The system characterising the warmer period is more based on recycling and less on export. The increase in the minimum turnover time indicates an increase in the ecosystem metabolism, which can be considered as a response of the pelagic ecosystems to climate warming. This phenomenon could reduce carbon export. These two opposite patterns of change are examples of the diversity of mechanisms and pathways the ecosystems may exhibit with climate change. Oversimplification of current biogeochemical models, often due to lack of data and biological understanding, could lead to wrong projection on the direction ecosystems and therefore some biogeochemical cycles might take in a warmer world.

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.

Macroecological study of Centropages typicus in the North Atlantic Ocean

Centropages typicus is a temperate neritic-coastal species of the North Atlantic Oceans, generally found between the latitudes of the Mediterranean and the Norwegian Sea. Therefore, the species experiences a large number of environments and adjusts its life cycle in response to changes in key abiotic parameters such as temperature. Using data from the Continuous Plankton Recorder (CPR) Survey, we review the macroecology of C. typicus and factors that influence its spatial distribution, phenology and year-to-year to decadal variability. The ecological preferences are identified and quantified. Mechanisms that allow the species to occur in such different environments are discussed and hypotheses are proposed as to how the species adapts to its environment. We show that temperature and both quantity and quality of phytoplankton are important factors explaining the space and time variability of C. typicus. These results show that C. typicus will not respond only to temperature increase in the region but also to changes in phytoplankton abundance, structure and composition and timing of occurrence. Methods such as a decision tree can help to forecast expected changes in the distribution of this species with hydro-climatic forcing. (C) 2007 Elsevier Ltd. All rights reserved.

Long-term responses of North Atlantic calcifying plankton to climate change

The global increase in atmospheric carbon dioxide concentration is potentially threatening marine biodiversity in two ways. First, carbon dioxide and other greenhouse gases accumulating in the atmosphere are causing global warming1. Second, carbon dioxide is altering sea water chemistry, making the ocean more acidic2. Although temperature has a cardinal influence on all biological processes from the molecular to the ecosystem level3, acidification might impair the process of calcification or exacerbate dissolution of calcifying organisms4. Here, we show however that North Atlantic calcifying plankton primarily responded to climate-induced changes in temperatures during the period 1960–2009, overriding the signal from the effects of ocean acidification. We provide evidence that foraminifers, coccolithophores, both pteropod and nonpteropod molluscs and echinoderms exhibited an abrupt shift circa 1996 at a time of a substantial increase in temperature5 and that some taxa exhibited a poleward movement in agreement with expected biogeographical changes under sea temperature warming6,7. Although acidification may become a serious threat to marine calcifying organisms, our results suggest that over the study period the primary driver of North Atlantic calcifying planktonwas oceanic temperature.

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.

Comparing copepod time-series in the north of Spain: Spatial autocorrelation of community composition

Four time-series of copepod species biomass in the north of Spain were contrasted to demonstrate spatial autocorrelation of local communities and their responses to short-term local and regional variability in oceanographic conditions. The series represented coastal and oceanic environments along a marked gradient of influence of seasonal upwelling from Galicia to the Mar Cantábrico (S Bay of Biscay), and each one included at least 10 years of continuous data collected at monthly frequency. Community composition (i.e. species number and diversity) was very consistent through the region, but local variations in the presence of new species and the relative proportions of common species allowed for the characterisation of the response to the environment at each site. Small-sized species were more frequent near the coast. A few species, however, captured the main patterns of variability in all series. Calanus helgolandicus and Acartia (mainly Acartia clausi) were generally the main contributors to total biomass, while other species as Paracalanus parvus and Clausocalanus spp. were important only at some locations. Most copepod indices were positively correlated with upwelling, either considering the whole community (biomass, species richness and diversity) or individual species, but only in the coastal series analysed since 1991. Copepods in the nearby ocean, however, showed negative correlations with upwelling in the period 1960–1986. The effects of upwelling may have been modulated by local factors, as showed by the increases in biomass, number of species and diversity in associations with increases in sea surface temperature in Galicia, while in the Mar Cantábrico only the warming-tolerant species increased and those typical of upwelling decreased. Density stratification of the water column was associated with decreases in total copepod biomass in Galicia, while it favoured the increase in species richness in the Mar Cantábrico. Nearly all significant responses of copepods to environmental variability were delayed by up to 5 months, showing the importance of considering time-lags in the analysis of temporal responses of zooplankton.

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea, English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas. (C) 2007 Elsevier Ltd. All rights reserved.