Long-term changes in North Atlantic zooplankton biomass

New measures of zooplankton biomass have been derived from CPR samples in the North Atlantic from 1958 to 2005. The final aim was to investigate how the zooplankton standing stock had varied throughout the last decades, knowing that in different areas of the North Atlantic significant changes in the distribution of the dominant zooplankton species as well as the plankton assemblage have been observed. During the forty-five years of monitoring the contribution of the different groups (e.g. copepods, euphausiids, meroplankton larvae) to the total zooplankton biomass has been evaluated. The changes in the phenology of the biomass were also considered. The relationship between quantity, quality and seasonal timing of plankton and the poor fish recruitment seen in recent years in the North Sea are also discussed.

Basin scale distribution of zooplankton in the Ligurian Sea (north-western Mediterranean) in late autumn

Mesozooplankton biomass and abundance were evaluated in epipelagic waters at 59 stations covering the Italian sector of the Ligurian Sea (north-western Mediterranean) in December 1990. This region is characterised by a cyclonic circulation which encloses a central divergence zone and is associated with a main thermohaline front offshore the western Ligurian coast. At the end of autumn, mesozooplankton biomass (range: 0.80–4.24 mg DW m−3) and the abundance (range: 83.8–932 ind. m−3) were lower in the divergence zone. On the contrary, in the Ligurian frontal zone at the periphery of the divergence and on the eastern continental shelf the greatest values of biomass and abundance were recorded. Copepods and appendicularians dominated the mesozooplankton community, the main taxa being the copepods Clausocalanus spp. (46% of total zooplankton) and Oithona spp. (15%) and the appendicularian Fritillaria spp. (12%). Three hydrological sub-regions, i.e. the divergence, the eastern continental shelf and the periphery of the divergence, were characterised by different zooplankton communities and characteristic species. Environmental differences between the three zones were mainly related to changes in bottom topography, sea surface temperatures and quantity of particulate organic matter. Vertical mesozooplankton abundance and taxa distribution from the surface to 1,900 m depth were also examined in one station. The results showed that the bulk of the community was concentrated in the upper 200 m, small copepods being dominant particularly in the upper 50 m. The copepod community was more diversified in sub-superficial waters, with a maximum observed in the 200–400 m layer. The distributions of main zooplankton taxa described in epipelagic waters in the eastern Ligurian Sea in autumn were compared with their distribution at surface in the north-western Mediterranean obtained by sampling performed with the Continuous Plankton Recorder in 1997–1999. The analysis of the zooplankton community in CPR samples confirms the dominance of small copepods (Paracalanus spp., Clausocalanus spp., Oithona spp.) and appendicularians in the north-western Mediterranean in late autumn-winter and shows that their distribution is mainly related to the main mesoscale hydrographic features characterising this basin.

Long-term variability and environmental preferences of calycophoran siphonophores in the Bay of Villefranche (North-Western Mediterranean)

Long-term variability of the main calycophoran siphonophores was investigated between 1974 and 1999 in a coastal station in the north-western Mediterranean. The data were collected at weekly frequency using a macroplankton net (680 μm mesh size) adapted to quantitatively sample delicate gelatinous plankton. A 3-year collection (1967–1969) of siphonophores from offshore waters using the same methodology showed that the patterns of variability observed inshore were representative of siphonophores’ changes at a regional scale. The aims of the study were: (i) to investigate the patterns of variability that characterised the dominant calycophoran species and assemblages; (ii) to identify the environmental optima that were associated with a significant increase in the dominant siphonophore species and (iii) to verify the influence of hydroclimatic variability on long-term changes of siphonophores. Our results showed that during nearly 3 decades the standing stock of calycophoran siphonophores did not show any significant change, with the annual maximum usually recorded in spring as a result of high densities of the dominant species Lensia subtilis, Muggiaea kochi and Muggiaea atlantica. Nevertheless, major changes in community composition occurred within the calycophoran population. Since the middle 1980s, M. kochi, once the most dominant species, started to decrease allowing other species, the congeneric M. atlantica and Chelophyes appendiculata, to increasingly dominate in spring and summer–autumn, respectively. The comparison of environmental and biotic long-term trends suggests that the decrease of M. kochi was triggered by hydrological changes that occurred in the north-western Mediterranean under the forcing of large-scale climate oscillations. Salinity, water stratification and water temperature were the main hydroclimatic factors associated with a significant increase of siphonophores, different species showing different environmental preferences.

Distribution and Seasonal Cycles of Decapod Crustacean Larvae in Continuous Plankton Records from the North Pacific Ocean

Decapoda taken in Continuous Plankton Recorder (CPR) samples from the Pacific in 1997 and 2000-2003 have been identified and measured. Some previously un-described larval stages were referred to species and characteristics of these are described. Distributions and seasonal occurrence of decapod taxa in the samples are described and discussed with particular emphasis on the dendrobranchiate shrimp Sergestes similis and the brachyurans Cancer spp. And Chionoecetes spp. There is a prolonged larval season at low levels of abundance off the Californian coast but in the more northern waters there is a shorter productive period but numbers of larvae per sample are high, particularly in June. Larvae of Chionoecetes and other Oregoninae were found only from May to July.

Warm-water decapods and the trophic amplification of climate in the North Sea

A long-term time series of plankton and benthic records in the North Sea indicates an increase in decapods and a decline in their prey species that include bivalves and flatfish recruits. Here, we show that in the southern North Sea the proportion of decapods to bivalves doubled following a temperature-driven, abrupt ecosystem shift during the 1980s. Analysis of decapod larvae in the plankton reveals a greater presence and spatial extent of warm-water species where the increase in decapods is greatest. These changes paralleled the arrival of new species such as the warm-water swimming crab Polybius henslowii now found in the southern North Sea. We suggest that climate-induced changes among North Sea decapods have played an important role in the trophic amplification of a climate signal and in the development of the new North Sea dynamic regime.

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.

North Sea ecosystem change from swimming crabs to seagulls

A recent increase in sea temperature has established a new ecosystem dynamic regime in the North Sea. Climate-induced changes in decapods have played an important role. Here, we reveal a coincident increase in the abundance of swimming crabs and lesser black-backed gull colonies in the North Sea, both in time and in space. Swimming crabs are an important food source for lesser black-backed gulls during the breeding season. Inhabiting the land, but feeding mainly at sea, lesser black-backed gulls provide a link between marine and terrestrial ecosystems, since the bottom-up influence of allochthonous nutrient input from seabirds to coastal soils can structure the terrestrial food web. We, therefore, suggest that climate-driven changes in trophic interactions in the marine food web may also have ensuing ramifications for the coastal ecology of the North Sea.

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.