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.

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.

Spatial variability of the plankton trophic interaction in the North Sea: a new feature after the early 1970s

Traditionally, marine ecosystem structure was thought to be bottom-up controlled. In recent years, a number of studies have highlighted the importance of top-down regulation. Evidence is accumulating that the type of trophic forcing varies temporally and spatially, and an integrated view – considering the interplay of both types of control – is emerging. Correlations between time series spanning several decades of the abundances of adjacent trophic levels are conventionally used to assess the type of control: bottom-up if positive or top-down if this is negative. This approach implies averaging periods which might show time-varying dynamics and therefore can hide part of this temporal variability. Using spatially referenced plankton information extracted from the Continuous Plankton Recorder, this study addresses the potential dynamic character of the trophic structure at the planktonic level in the North Sea by assessing its variation over both temporal and spatial scales. Our results show that until the early-1970s a bottom-up control characterized the base of the food web across the whole North Sea, with diatoms having a positive and homogeneous effect on zooplankton filter-feeders. Afterwards, different regional trophic dynamics were observed, in particular a negative relationship between total phytoplankton and zooplankton was detected off the west coast of Norway and the Skagerrak as opposed to a positive one in the southern reaches. Our results suggest that after the early 1970s diatoms remained the main food source for zooplankton filter-feeders east of Orkney–Shetland and off Scotland, while in the east, from the Norwegian Trench to the German Bight, filter-feeders were mainly sustained by dinoflagellates.

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.

Spatio-Temporal Variability of the North Sea Cod Recruitment in Relation to Temperature and Zooplankton

The North Sea cod (Gadus morhua, L.) stock has continuously declined over the past four decades linked with overfishing and climate change. Changes in stock structure due to overfishing have made the stock largely dependent on its recruitment success, which greatly relies on environmental conditions. Here we focus on the spatio-temporal variability of cod recruitment in an effort to detect changes during the critical early life stages. Using International Bottom Trawl Survey (IBTS) data from 1974 to 2011, a major spatio-temporal change in the distribution of cod recruits was identified in the late 1990s, characterized by a pronounced decrease in the central and southeastern North Sea stock. Other minor spatial changes were also recorded in the mid-1980s and early 1990s. We tested whether the observed changes in recruits distribution could be related with direct (i.e. temperature) and/or indirect (i.e. changes in the quantity and quality of zooplankton prey) effects of climate variability. The analyses were based on spatially-resolved time series, i.e. sea surface temperature (SST) from the Hadley Center and zooplankton records from the Continuous Plankton Recorder Survey. We showed that spring SST increase was the main driver for the most recent decrease in cod recruitment. The late 1990s were also characterized by relatively low total zooplankton biomass, particularly of energy-rich zooplankton such as the copepod Calanus finmarchicus, which have further contributed to the decline of North Sea cod recruitment. Long-term spatially-resolved observations were used to produce regional distribution models that could further be used to predict the abundance of North Sea cod recruits based on temperature and zooplankton food availability.

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.

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.