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.

Acquisition of continuous plankton recorder data

The Continuous Plankton Recorder has been deployed on a seasonal basis in the north Pacific since 2000, accumulating a database of abundance measurements for over 290 planktonic taxa in over 3,500 processed samples. There is an additional archive of over 10,000 samples available for further analyses. Exxon Valdez Oil Spill Trustee Council financial support has contributed to about half of this tally, through four projects funded since 2002. Time series of zooplankton variables for sub-regions of the survey area are presented together with abstracts of eight papers published using data from these projects. The time series covers a period when the dominant climate signal in the north Pacific, the Pacific Decadal Oscillation (PDO), switched with unusual frequency between warm/positive states (pre-1999 and 2003-2006) and cool/negative states (1999-2002 and 2007). The CPR data suggest that cool negative years show higher biomass on the shelf and lower biomass in the open ocean, while the reverse is true in warm (PDO positive) years with lower shelf biomass (except 2005) and higher oceanic biomass. In addition, there was a delay in plankton increase on the Alaskan shelf in the colder spring of 2007, compared to the warmer springs of the preceding years. In warm years, smaller species of copepods which lack lipid reserves are also more common. Availability of the zooplankton prey to higher trophic levels (including those that society values highly) is therefore dependent on the timing of increase and peak abundance, ease of capture and nutritional value. Previously published studies using these data highlight the wide-ranging applicability of CPR data and include collaborative studies on; phenology in the key copepod species Neocalanus plumchrus, descriptions of distributions of decapod larvae and euphausiid species, the effects of hydrographic features such as mesoscale eddies and the North Pacific Current on plankton populations and a molecularbased investigation of macro-scale population structure in N. cristatus. The future funding situation is uncertain but the value of the data and studies so far accumulated is considerable and sets a strong foundation for further studies on plankton dynamics and interactions with higher trophic levels in the northern Gulf of Alaska.

Recent trends in plankton and upwelling intensity off Galicia (NW Spain)

The mean intensity of the NE Atlantic upwelling system at its northern limit (Galicia, NW Spain) decreased during the last 40 years. At the same time, warming of surface waters was detected. Plankton biomass and composition are expected to reflect such changes when integrated over large time and space scales. In this study, biomass, abundance and species composition of phyto- and zooplankton were analysed to search for significant patterns of annual change and relations with upwelling intensity. Regionally integrated, mostly offshore, data were obtained from the Continuous Plankton Recorder (since 1958) whereas coastal data from Vigo and A Coruña came from the Radiales program (since 1987). No significant trends were found in phytoplankton biomass at either regional or local scales. However, there was a significant decrease in diatom abundance at regional scales and also of large species at local scales. Zooplankton abundance (mainly copepods) significantly decreased offshore but increased near the coast. Biomass of zooplankton also increased near the coast, with the fastest rates in the south. Warm-water species, like Temora stylifera, were increasingly abundant at both regional and local scales. Significant correlations between upwelling intensity and plankton suggest that climatic effects were delayed for several years. Our results indicate that the effects of large scale climatic trends on plankton communities are being effectively modulated within the pelagic ecosystem in this upwelling region.

Influence of the Pacific Decadal Oscillation on phytoplankton phenology and community structure in the western North Pacific

Phytoplankton phenology and community structure in the western North Pacific were investigated for 2001–2009, based on satellite ocean colour data and the Continuous Plankton Recorder survey. We estimated the timing of the spring bloom based on the cumulative sum satellite chlorophyll adata, and found that the Pacific Decadal Oscillation (PDO)-related interannual SST anomaly in spring significantly affected phytoplankton phenology. The bloom occurred either later or earlier in years of positive or negative PDO (indicating cold and warm conditions, respectively). Phytoplankton composition in the early summer varied depending on the magnitude of seasonal SST increases, rather than the SST value itself. Interannual variations in diatom abundance and the relative abundance of non-diatoms were positively correlated with SST increases for March–April and May–July, respectively, suggesting that mixed layer environmental factors, such as light availability and nutrient stoichiometry, determine shifts in phytoplankton community structure. Our study emphasised the importance of the interannual variation in climate-induced warm–cool cycles as one of the key mechanisms linking climatic forcing and lower trophic level ecosystems.

The North Atlantic subpolar gyre regulates the spawning distribution of blue whiting (Micromesistius poutassou)

The spawning stock of blue whiting (Micromesistius poutassou), an economically important pelagic gadoid in the North Atlantic Ocean, increased threefold after 1995. The reproductive success of the stock is largely determined during the very early stages of life, but little is known about the spawning dynamics of this species. Here we show that the spawning distribution of blue whiting is variable, regulated by the hydrography west of the British Isles. When the North Atlantic subpolar gyre is strong and spreads its cold, fresh water masses east over Rockall Plateau, the spawning is constrained along the European continental slope and in a southerly position near Porcupine Bank. When the gyre is weak and conditions are relatively saline and warm, the spawning distribution moves northwards along the slope and especially westwards covering Rockall Plateau. The apparent link between the spawning distribution and the subpolar gyre is the first step towards understanding the reproduction variability, which currently is the main challenge for appropriate management of the blue whiting stock.

Monitoring marine plankton ecosystems: Identification of the most relevant indicators of the state of an ecosystem

The number of variables involved in the monitoring of an ecosystem can be high and often one of the first stages in the analysis is to reduce the number of variables. We describe a method developed for geological purposes, using the information theory, that enables selection of the most relevant variables. This technique also allows the examination of the asymmetrical relationships between variables. Applied to a set of physical and biological variables (plankton assemblages in four areas of the North Sea), the method shows that biological variables are more informative than physical variables although the controlling factors are mainly physical (sea surface temperature in winter and spring). Among biological variables, diversity measures and warm-water species assemblages are informative for the state of the North Sea pelagic ecosystems while among physical variables sea surface temperature in late winter and early spring are highly informative. Although often used in bioclimatology, the utilisation of the North Atlantic Oscillation (NAO) index does not seem to provide a lot of information. The method reveals that only the extreme states of this index has an influence on North Sea pelagic ecosystems. The substantial and persistent changes that were detected in the dynamic regime of the North Sea ecosystems and called regime shift are detected by the method and corresponds to the timing of other shifts described in the literature for some European Systems such as the Baltic and the Mediterranean Sea when both physical and biological variables are considered.

A blooming jellyfish in the northeast Atlantic and Mediterranean

A long-term time series of plankton records collected by the continuous plankton recorder (CPR) Survey in the northeast Atlantic indicates an increased occurrence of Cnidaria since 2002. In the years 2007 and 2008, outbreaks of the warm-temperate scyphomedusa, Pelagia noctiluca, appeared in CPR samples between 45° N to 58° N and 1° W to 26° W. Knowing the biology of this species and its occurrence in the adjacent Mediterranean Sea, we suggest that P. noctiluca may be exploiting recent hydroclimatic changes in the northeast Atlantic to increase its extent and intensity of outbreaks. In pelagic ecosystems, Cnidaria can affect fish recruitment negatively. Since P. noctiluca is a highly venomous species, outbreaks can also be detrimental to aquaculture and make bathing waters unusable, thus having profound ecological and socio-economic consequences.

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.

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.

Effects on zooplankton of a warmer ocean: Recent evidence from the Northeast Pacific

The consequences for pelagic communities of warming trends in mid and high latitude ocean regions could be substantial, but their magnitude and trajectory are not yet known. Environmental changes predicted by climate models (and beginning to be confirmed by observations) include warming and freshening of the upper ocean and reduction in the extent and duration of ice cover. One way to evaluate response scenarios is by comparing how "similar" zooplankton communities have differed among years and/or locations with differing temperature. The subarctic Pacific is a strong candidate for such comparisons, because the same mix of zooplankton species dominates over a wide range of temperature climatologies, and observations have spanned substantial temperature variability at interannual-to-decadal time scales. In this paper, we review and extend copepod abundance and phenology time series from net tow and Continuous Plankton Recorder surveys in the subarctic Northeast Pacific. The two strongest responses we have observed are latitudinal shifts in centers of abundance of many species (poleward under warm conditions), and changes in the life cycle timing of Neocalanus plumchrus in both oceanic and coastal regions (earlier by several weeks in warm years and at warmer locations). These zooplankton data, plus indices of higher trophic level responses such as reproduction, growth and survival of pelagic fish and seabirds, are all moderately-to-strongly intercorrelated (vertical bar r vertical bar = 0.25-0.8) with indices of local and basin-scale temperature anomalies. A principal components analysis of the normalized anomaly time series from 1979 to 2004 shows that a single "warm-and-low-productivity" vs. "cool-and-high-productivity" component axis accounts for over half of the variance/covariance. Prior to 1990, the scores for this component were negative ("cool" and "productive") or near zero except positive in the El Nino years 1983 and 1987. The scores were strongly and increasingly positive ("warm" and "low productivity") from 1992 to 1998; negative from 1999 to 2002; and again increasingly positive from 2003-present. We suggest that, in strongly seasonal environments, anomalously high temperature may provide misleading environmental cues that contribute to timing mismatch between life history events and the more-nearly-fixed seasonality of insolation, stratification, and food supply. Crown Copyright (c) 2007 Published by Elsevier Ltd. All rights reserved.

Zooplankton response to a warmer northern Wadden Sea

Weekly measurements of mesozooplankton (>76 mu m) and hydrographic parameters have been carried out since 1984 in the List Tidal Basin (northern Wadden Sea). Monthly water temperature significantly increased by 0.04 degrees C year. The largest increase by 3 degrees C in 22 years occurred in September, implying, an extension of the warm summer period. Mean annual copepod abundance and length of copepod season correlated significantly with mean temperature from January to May. Except for an increasing Acartia sp. abundance during spring (April-May), no longterm trends in copepod abundance were observed. The percentage of carnivorous zooplankton increased significantly since 1984 mainly due to a sudden increase in the cyclopoid copepod Oithona similis in 1997. We expect that global warming will lead to a longer copepod season and higher copepod abundances in the northern Wadden Sea.

Non-linearities, regime shifts and recovery: The recent influence of climate on Black Sea chlorophyll

The Black Sea ecosystem experienced severe eutrophication-related degradation during the 1970s and 1980s. However, in recent years the Black Sea has shown some signs of recovery which are often attributed to a reduction in nutrient loading. Here, SeaWiFS chlorophyll a (chl a), a proxy for phytoplankton biomass, is used to investigate spatio-temporal patterns in Black Sea phytoplankton dynamics and to explore the potential role of climate in the Black Sea's recovery. Maps of chl a anomalies, calculated relative to the 8 year mean, emphasize spatial and temporal variability of phytoplankton biomass in the Black Sea, particularly between the riverine-influenced Northwest Shelf and the open Black Sea. Evolution of phytoplankton biomass has shown significant spatial variability of persistence of optimal bloom conditions between three major regions of the Black Sea. With the exception of 2001, chl a has generally decreased during our 8 year time-series. However, the winter of 2000–2001 was anomalously warm with low wind stress, resulting in reduced vertical mixing of the water column and retention of nutrients in the photic zone. These conditions were associated with anomalously high levels of chl a throughout much of the open Black Sea during the following spring and summer. The unusual climatic conditions occurring in 2001 may have triggered a shift in the Black Sea's chl a regime. The long-term significance of this recent shift is still uncertain but illustrates a non-linear response to climate forcing that makes future ecosystem changes in the pelagic Black Sea ecosystem difficult to predict.

Global synchrony of an accelerating risein sea surface temperature

The oceans have shown a recent rapid and accelerating rise in temperature with, given the close link between temperature and marine organisms, pronounced effects on ecosystems. Here we describe for the first time a globally synchronous pattern of pulsed short period (�1 year long) emanations of warm sea surface temperature anomalies from tropical seas towards the poles on the shelf/slope with an intensification of the warming after the 1976/1977, 1986/1987 and 1997/1998 El Nin˜os. On the eastern margins of continents the anomalies propagate towards the poles in part by largely baroclinic boundary currents, reinforced by regional atmospheric warming. The processes contributing to the less continuous warm anomalies on western margins are linked to the transfer of warmth from adjacent western boundary currents. These climate induced events show a close parallelism with the timing of ecosystem changes in shelf seas, important for fisheries and ecosystem services, and melting of sea-ice.

Monitoring of a quasi-stationary eddy in the Bay of Biscay by means of satellite, in situ and model results

The presence of a quasi-stationary anticyclonic eddy within the southeastern Bay of Biscay (centred around 44°30′N-4°W) has been reported on various occasions in the bibliography. The analysis made in this study for the period 2003–2010, by using in situ and remote sensing measurements and model results shows that this mesoscale coherent structure is present almost every year from the end of winter-beginning of spring, to the beginning of fall. During this period it remains in an area limited to the east by the Landes Plateau, to the west by Le Danois Bank and Torrelavega canyon and to the northwest by the Jovellanos seamount. All the observations and analysis made in this contribution, suggest that this structure is generated between Capbreton and Torrelavega canyons. Detailed monitoring from in situ and remote sensing data of an anticyclonic quasi-stationary eddy, in 2008, shows the origin of this structure from a warm water current located around 43°42′N-3°30′W in mid-January. This coherent structure is monitored until August around the same area, where it has a marked influence on the Sea Level Anomaly, Sea Surface Temperature and surface Chlorophyll-a concentration. An eddy tracking method, applied to the outputs of a numerical model, shows that the model is able to reproduce this type of eddy, with similar 2D characteristics and lifetimes to that suggested by the observations and previous works. This is the case, for instance, of the simulated MAY04 eddy, which was generated in May 2004 around Torrelavega canyon and remained quasi-stationary in the area for 4 months. The diameter of this eddy ranged from 40 to 60 km, its azimuthal velocity was less than 20 cm s−1, its vertical extension reached 3000–3500 m depth during April and May and it was observed to interact with other coherent structures.

Monitoring of a quasi-stationary eddy in the Bay of Biscay by means of satellite, in situ and model results

The presence of a quasi-stationary anticyclonic eddy within the southeastern Bay of Biscay (centred around 44°30′N-4°W) has been reported on various occasions in the bibliography. The analysis made in this study for the period 2003–2010, by using in situ and remote sensing measurements and model results shows that this mesoscale coherent structure is present almost every year from the end of winter-beginning of spring, to the beginning of fall. During this period it remains in an area limited to the east by the Landes Plateau, to the west by Le Danois Bank and Torrelavega canyon and to the northwest by the Jovellanos seamount. All the observations and analysis made in this contribution, suggest that this structure is generated between Capbreton and Torrelavega canyons. Detailed monitoring from in situ and remote sensing data of an anticyclonic quasi-stationary eddy, in 2008, shows the origin of this structure from a warm water current located around 43°42′N-3°30′W in mid-January. This coherent structure is monitored until August around the same area, where it has a marked influence on the Sea Level Anomaly, Sea Surface Temperature and surface Chlorophyll-a concentration. An eddy tracking method, applied to the outputs of a numerical model, shows that the model is able to reproduce this type of eddy, with similar 2D characteristics and lifetimes to that suggested by the observations and previous works. This is the case, for instance, of the simulated MAY04 eddy, which was generated in May 2004 around Torrelavega canyon and remained quasi-stationary in the area for 4 months. The diameter of this eddy ranged from 40 to 60 km, its azimuthal velocity was less than 20 cm s−1, its vertical extension reached 3000–3500 m depth during April and May and it was observed to interact with other coherent structures.