Long-term changes in the meroplankton of the North Sea

Data from the continuous plankton recorder (CPR) survey collected in the late-1940s to early-1960s indicated that the abundance of decapod larvae was low and the seasonal peak of abundance was late following cold winters. The phenological effect of temperature was shown to be consistent with relationships between both geographical and interannual patterns of variation. Analyses of CPR data collected from the 1940s to the present day reveal large-scale long-term changes in the abundance and phenology of the North Sea meroplankton. Echinoderm larvae, whose peak abundance has advanced by 47 days, show the greatest shift in timing. Echinoderm larvae have also increased in abundance to become the most abundant taxon in North Sea CPR samples. Genetic and morphological analyses of CPR samples show that the variations in echinoderm larvae are mainly attributable to an increasing abundance and earlier occurrence of the larvae of a resident species, Echinocardium cordatum, rather than a change in species composition. The remarkable scale of the changes in abundance and phenology of the meroplankton, which are greater than those seen in the holoplankton, has stimulated the development of further research into the causes and effects of these changes.

The mid-eighties regime shift and appendicularian abundance

There is accumulating evidence suggesting that a regime shift occurred in the North Atlantic during the mid-eighties. This shift has been reported primarily from Continuous Plankton Recorder (CPR) data as a stepwise change in plankton abundance and copepod community structure. Here we analyse the CPR data for appendicularian abundance to show that a similar stepwise increase occurred in the abundance of appendicularians during the mid-eighties. Furthermore, we compare these results to data recorded by other zooplankton time series programmes to show that a similar abrupt increase in appendicularian abundance during the mid-early eighties has also been recorded in other areas. The fact that such a change occurred at locations so distant apart as Helgoland Roads in the North Sea or the White Sea in the Arctic suggests that these changes have a global origin. The strong dependence of appendicularian phenology with temperature points out to direct links to global climate change.

Spatial patterns and trends in abundance of larval sandeels in the North Sea: 1950-2005

Early recruitment indices based on larval fish data from the Continuous Plankton Recorder (CPR) have the potential to inform stock assessments of Ammodytes marinus in the North Sea. We evaluate whether the CPR data are reliable for sandeel larvae. Spatially, CPR larval data were comparable with catches by dedicated larval samplers (Gulf and bongo nets) during ICES coordinated surveys in 2004 and 2009. ICES data are also used to explore environmental influences on sandeel distributions. Temporally, CPR data correlate with larval data from plankton surveys off Stonehaven (1999–2005), with sandeel 0-group trawl data at the east Fair Isle ground (since 1984), and with recruitment data (since 1983) for the Dogger Banks stock assessment area. Therefore, CPR data may provide an early recruit index of relative abundance for the Dogger Banks assessment area, where the majority of the commercial catch of A. marinus is taken, and the Wee Bankie area that is particularly important for seabird foraging. While warm conditions may stimulate the production of sandeel larvae, their natural mortality is typically greater, in the Dogger Banks and Wadden Sea areas, when the larvae are hatched in warm years and/or with abundant 1-year-old sandeel that are likely to be cannibalistic.

Challenges for implementing the Marine Strategy Framework Directive in a climate of macroecological change

Unprecedented basin-scale ecological changes are occurring in our seas. As temperature and carbon dioxide concentrations increase, the extent of sea ice is decreasing, stratification and nutrient regimes are changing and pH is decreasing. These unparalleled changes present new challenges for managing our seas, as we are only just beginning to understand the ecological manifestations of these climate alterations. The Marine Strategy Framework Directive requires all European Member States to achieve good environmental status (GES) in their seas by 2020; this means management towards GES will take place against a background of climate-driven macroecological change. Each Member State must set environmental targets to achieve GES; however, in order to do so, an understanding of large-scale ecological change in the marine ecosystem is necessary. Much of our knowledge of macroecological change in the North Atlantic is a result of research using data gathered by the Continuous Plankton Recorder (CPR) survey, a near-surface plankton monitoring programme that has been sampling in the North Atlantic since 1931. CPR data indicate that North Atlantic and North Sea plankton dynamics are responding to both climate and human-induced changes, presenting challenges to the development of pelagic targets for achievement of GES in European Seas. Thus, the continuation of long-term ecological time series such as the CPR survey is crucial for informing and supporting the sustainable management of European seas through policy mechanisms.

Why is it so hard to set MSFD indicators and targets? Messages from the plankton

Unprecedented basin-scale ecological changes are occurring in our seas. As temperature and carbon dioxide concentrations increase, the extent of sea ice is decreasing, stratification and nutrient regimes are changing, and pH is decreasing. These unparalleled changes present new challenges for managing our seas as we are only just beginning to understand the ecological manifestations of these climate alterations. The Marine Strategy Framework Directive requires all European Member States to achieve Good Environmental Status (GES) in their seas by 2020; this means management toward GES will take place against a background of climate-driven macroecological change. Each Member State must set environmental targets to achieve GES; however, in order to do so an understanding of large-scale ecological change in the marine ecosystem is necessary. Much of our knowledge of macroecological change in the North Atlantic is a result of research using data gathered by the Continuous Plankton Recorder (CPR) survey, a near-surface plankton monitoring program which has been sampling in the North Atlantic since 1931. CPR data indicate that North Atlantic and North Sea plankton dynamics are responding to both climate and human-induced changes, presenting challenges to the development of pelagic targets for achievement of GES in European seas. Thus the continuation of long-term ecological time-series such as the CPR is crucial for informing and supporting the sustainable management of European seas through policy mechanisms.

Is there a decline in marine phytoplankton?

Phytoplankton account for approximately 50% of global primary production, form the trophic base of nearly all marine ecosystems, are fundamental in trophic energy transfer and have key roles in climate regulation, carbon sequestration and oxygen production. Boyce et al.1 compiled a chlorophyll index by combining in situ chlorophyll and Secchi disk depth measurements that spanned a more than 100-year time period and showed a decrease in marine phytoplankton biomass of approximately 1% of the global median per year over the past century. Eight decades of data on phytoplankton biomass collected in the North Atlantic by the Continuous Plankton Recorder (CPR) survey2, however, show an increase in an index of chlorophyll (Phytoplankton Colour Index) in both the Northeast and Northwest Atlantic basins3, 4, 5, 6, 7 (Fig. 1), and other long-term time series, including the Hawaii Ocean Time-series (HOT)8, the Bermuda Atlantic Time Series (BATS)8 and the California Cooperative Oceanic Fisheries Investigations (CalCOFI)9 also indicate increased phytoplankton biomass over the last 20–50 years. These findings, which were not discussed by Boyce et al.1, are not in accordance with their conclusions and illustrate the importance of using consistent observations when estimating long-term trends.

The influence of phytoplankton productivity, temperature and environmental stability on the control of copepod diversity in the North East Atlantic

The patterns of copepod species richness (S) and their relationship with phytoplankton productivity, temperature and environmental stability were investigated at climatological, seasonal and year-to-year time scales as well as scales along latitudinal and oceanic–neritic gradients using monthly time series of the Continuous Plankton Recorder (CPR) Survey collected in the North East Atlantic between 1958 and 2006. Time series analyses confirmed previously described geographic patterns. Equatorward and towards neritic environments, the climatological average of S increases and the variance explained by the seasonal cycle decreases. The bi-modal character of seasonality increases equatorward and the timing of the seasonal cycle takes place progressive earlier equatorward and towards neritic environments. In the long-term, the climatological average of S decreased significantly (p < 0.001) between 1958 and 2006 in the Bay of Biscay and North Iberian shelf at a rate of ca. 0.04 year−1, and increased at the same rate between 1991 and 2006 in the northernmost oceanic location. The climatological averages of S correlate positively with those of the index of seasonality of phytoplankton productivity (ratio between the minimum and maximum monthly values of surface chlorophyll) and sea surface temperature, and negatively with those of the proxy for environmental stability (monthly frequency of occurrence of daily averaged wind speed exceeding 10 m s−1). The seasonal cycles of S and phytoplankton productivity (surface chlorophyll as proxy) exhibit similar features in terms of shape, timing and explained variance, but the relationship between the climatological averages of both variables is non-significant. From year-to-year, the annual averages of S correlate negatively with those of phytoplankton productivity and positively with those of sea surface temperature along the latitudinal gradient, and negatively with those of environmental stability along the oceanic–neritic gradient. The annual anomalies of S (i.e. factoring out geographic variation) show a unimodal relationship with those of sea surface temperature and environmental stability, with S peaking at intermediate values of the anomalies of these variables. The results evidence the role of seasonality of phytoplankton productivity on the control of copepod species richness at seasonal and climatological scales, giving support to the species richness–productivity hypothesis. Although sea surface temperature (SST) is indeed a good predictor of richness along the latitudinal gradient, it is unable to predict the increase of richness form oceanic to neritic environments, thus lessening the generality of the species richness–energy hypothesis. Meteo-hydrographic disturbances (i.e. SST and wind speed anomalies as proxies), presumably through its role on mixed layer depth dynamics and turbulence and hence productivity, maximise local diversity when occurring at intermediate frequency and or intensity, thus providing support to the intermediate disturbance hypothesis on the control of copepod diversity.

Pattern and scale of variability among Northwest Atlantic Shelf plankton communities

Continuous plankton recorders (CPRs) have been used in the Northwest Atlantic for almost 50 years. While data collected by these surveys have provided valuable information on long-term variability in plankton populations, all previous analyses have been limited to only a portion of the geographic range of the available data. Here we present an analysis of the CPR data from the Mid Atlantic Bight to the Labrador Sea. Across this wide geographic range, we found many common associations among the taxa. In particular, the changes in most regions were strongly size structured, with small and medium copepods varying together and often positively related to indicators of phytoplankton abundance. The time series from nearby regions were strongly correlated; however, after 1990, the spatial pattern became more complex. During this period, several of the copepod taxa, noticeably Calanus finmarchicus and Centropages typicus, experienced a series of anomalies that appeared to propagate from northeast to southwest. Although the direction of propagation was consistent with the shelf circulation, the anomalies propagated at a rate much slower than typical current speeds. The timing of the copepod anomalies and their phase speed were similar in character to observed changes in salinity and the position of the Shelf Slope Front. The correspondence between the changes in the plankton community and changes in the physical environmental suggests that physical conditions are a strong driver of interannual variability in Northwest Atlantic Shelf ecosystems.

Bottom-up effects of climate on fish populations: data from the Continuous Plankton Recorder

The Continuous Plankton Recorder (CPR) dataset on fish larvae has an extensive spatio-temporal coverage that allows the responses of fish populations to past changes in climate variability, including abrupt changes such as regime shifts, to be investigated. The newly available dataset offers a unique opportunity to investigate long-term changes over decadal scales in the abundance and distribution of fish larvae in relation to physical and biological factors. A principal component analysis (PCA) using 7 biotic and abiotic parameters is applied to investigate the impact of environmental changes in the North Sea on 5 selected taxa of fish larvae during the period 1960 to 2004. The analysis revealed 4 periods of time (1960–1976; 1977–1982; 1983–1996; 1997–2004) reflecting 3 different ecosystem states. The larvae of clupeids, sandeels, dab and gadoids seemed to be affected mainly by changes in the plankton ecosystem, while the larvae of migratory species such as Atlantic mackerel responded more to hydrographic changes. Climate variability seems more likely to influence fish populations through bottom-up control via a cascading effect from changes in the North Atlantic Oscillation (NAO) impacting on the hydro dynamic features of the North Sea, in turn impacting on the plankton available as prey for fish larvae. The responses and adaptability of fish larvae to changing environmental conditions, parti cularly to changes in prey availability, are complex and species-specific. This complexity is enhanced with fishing effects interacting with climate effects and this study supports furthering our under - standing of such interactions before attempting to predict how fish populations respond to climate variability

A 60-year ocean colour data set from the continuous plankton recorder

The phytoplankton colour index (PCI) of the Continuous Plankton Recorder (CPR) survey is an in situ measure of ocean colour, which is considered a proxy of the phytoplankton biomass. PCI has been extensively used to describe the major spatiotemporal patterns of phytoplankton in the North Atlantic Ocean and North Sea since 1931. Regardless of its wide application, the lack of an adequate evaluation to test the PCI's quantitative nature is an important limitation. To address this concern, a field trial over the main production season has been undertaken to assess the numerical values assigned by previous investigations for each category of the greenness of the PCI. CPRs were towed across the English Channel from Roscoff to Plymouth consecutively for each of 8 months producing 76 standard CPR samples, each representing 10 nautical miles of tow. The results of this experiment test and update the PCI methodology, and confirm the validity of this long-term in situ ocean colour data set. In addition, using a 60-year time series of the PCI of the western English Channel, a comparison is made between the previous and the current revised experimental calculations of PCI.

Macroscale factors affecting diatom abundance: a synergistic use of Continuous Plankton Recorder and satellite remote sensing data

Diatoms exist in almost every aquatic regime; they are responsible for 20% of global carbon fixation and 25% of global primary production, and are regarded as a key food for copepods, which are subsequently consumed by larger predators such as fish and marine mammals. A decreasing abundance and a vulnerability to climatic change in the North Atlantic Ocean have been reported in the literature. In the present work, a data matrix composed of concurrent satellite remote sensing and Continuous Plankton Recorder (CPR) in situ measurements was collated for the same spatial and temporal coverage in the Northeast Atlantic. Artificial neural networks (ANNs) were applied to recognize and learn the complex non-monotonic and non-linear relationships between diatom abundance and spatiotemporal environmental factors. Because of their ability to mimic non-linear systems, ANNs proved far more effective in modelling the diatom distribution in the marine ecosystem. The results of this study reveal that diatoms have a regular seasonal cycle, with their abundance most strongly influenced by sea surface temperature (SST) and light intensity. The models indicate that extreme positive SSTs decrease diatom abundances regardless of other climatic conditions. These results provide information on the ecology of diatoms that may advance our understanding of the potential response of diatoms to climatic change.

Biodiversity as a dynamic variable in the Gulf of Maine continuous plankton recorder transect

Studies relating biodiversity to ecosystem processes typically do not take into account changes in biodiversity through time. Marine systems are highly dynamic, with biodiversity changing at diel, seasonal and inter-decadal timescales. We examined the dynamics of biodiversity in the Gulf of Maine pelagic zooplankton community. Taxonomic data came from the Gulf of Maine continuous plankton recorder (CPR) transect, spanning the years 1961–2006. The CPR transect also contains coincident information on temperature and phytoplankton biomass (measured by the phytoplankton color index). Taxonomic richness varied at all timescales considered. The relationships between temperature and richness, and between phytoplankton and richness, also depended on temporal scale. The temperature–richness relationship was monotonic at the multi-decadal scale, and tended to be hump-shaped at finer scales; the productivity–richness relationship was hump-shaped at the multi-decadal scale, and tended to be monotonic at finer scales. Seasonal biodiversity dynamics were linked to temperature; inter-decadal biodiversity dynamics were linked to phytoplankton.

The importance of zooplankton in reducing levels of atmospheric CO sub(2) via the biological pump

Zooplankton play a key role in climate change through the transfer of large quantities of CO sub(2) to the deep ocean by a process known as the biological pump. Plankton composition is crucial as associated mineral material facilitates sinking of carbon rich debris and some taxa package faecal and detrital material. Ocean acidification may impact calcareous groups. Zooplankton have also been shown to be highly sensitive indicators of environmental change. Results will be presented to show that ocean temperature, circulation and planktonic ecosystems (using data from the Continuous Plankton Recorder, CPR survey) in the North Atlantic are changing rapidly in concert and that there is evidence to suggest that the changes are an ocean wide response to global warming with potential feedback effects. Given the importance of the oceans to the carbon cycle, even a minor change in the flux of carbon to the deep ocean would have a big impact increasing growth of atmospheric CO sub(2). We have virtually no understanding of the spatial and temporal variability in the efficiency of the biological pump for most of the world's ocean. Establishing new plankton monitoring programmes backed up by appropriate research to help understand processes is needed to address this gap in knowledge. There is little doubt within a global change context and the future of mankind that a potential acceleration in the growth of atmospheric carbon due to a reduction in the efficiency of the biological pump is a key issue for future research in zooplankton ecology.

A biological consequence of reducing Arctic ice cover: arrival of the Pacific diatom Neodenticula seminae in the North Atlantic for the first time in 800,000 years

The Continuous Plankton Recorder survey has monitored plankton in the Northwest Atlantic at monthly intervals since 1962, with an interegnum between 1978 and 1990. In May 1999, large numbers of the Pacific diatom Neodenticula seminae were found in Continuous Plankton Recorder (CPR) samples in the Labrador Sea as the first record in the North Atlantic for more than 800 000 years. The event coincided with modifications in Arctic hydrography and circulation, increased flows of Pacific water into the Northwest Atlantic and in the previous year the exceptional occurrence of extensive ice-free water to the North of Canada. These observations indicate that N. seminae was carried in a pulse of Pacific water in 1998/early 1999 via the Canadian Arctic Archipelago and/or Fram Strait. The species occurred previously in the North Atlantic during the Pleistocene from similar to 1.2 to similar to 0.8 Ma as recorded in deep sea sediment cores. The reappearance of N. seminae in the North Atlantic is an indicator of the scale and speed of changes that are taking place in the Arctic and North Atlantic oceans as a consequence of regional climate warming. Because of the unusual nature of the event it appears that a threshold has been passed, marking a change in the circulation between the North Pacific and North Atlantic Oceans via the Arctic. Trans-Arctic migrations from the Pacific into the Atlantic are likely to occur increasingly over the next 100 years as Arctic ice continues to melt affecting Atlantic biodiversity and the biological pump with consequent feedbacks to the carbon cycle.

Assessing marine plankton community structure from long-term monitoring data with multivariate autoregressive (MAR) models: a comparison of fixed station versus spatially distributed sampling data

We examined how marine plankton interaction networks, as inferred by multivariate autoregressive (MAR) analysis of time-series, differ based on data collected at a fixed sampling location (L4 station in the Western English Channel) and four similar time-series prepared by averaging Continuous Plankton Recorder (CPR) datapoints in the region surrounding the fixed station. None of the plankton community structures suggested by the MAR models generated from the CPR datasets were well correlated with the MAR model for L4, but of the four CPR models, the one most closely resembling the L4 model was that for the CPR region nearest to L4. We infer that observation error and spatial variation in plankton community dynamics influenced the model performance for the CPR datasets. A modified MAR framework in which observation error and spatial variation are explicitly incorporated could allow the analysis to better handle the diverse time-series data collected in marine environments.