How well does the Continuous Plankton Recorder (CPR) sample zooplankton? A comparison with the Longhurst Hardy Plankton Recorder (LHPR) in the northeast Atlantic

The Continuous Plankton Recorder (CPR) survey has collected data on basin- scale zooplankton abundance in the North Atlantic since the 1930s. These data have been used in many studies to elucidate seasonal patterns and long-term change in plankton populations, as well as more recently to validate ecosystem models. There has, however, been relatively little comparison of the data from the CPR with that from other samplers. In this study we compare zooplankton abundance estimated from the CPR in the northeast Atlantic with near-surface samples collected by a Longhurst-Hardy Plankton Recorder (LHPR) at Ocean Weather Station India (59 degree N, 19 degree W) between 1971 and 1975. Comparisons were made for six common copepods in the region: Acartia clausi, Calanus finmarchicus, Euchaeta norvegica, Metridia lucens, Oithona sp. and Pleuromamma robusta. Seasonal cycles based on CPR data were similar to those recorded by the LHPR. Differences in absolute abundances were apparent, however, with the CPR underestimating abundances by a factor of between 5 and 40, with the exception of A. clausi. Active avoidance by zooplankton is thought to be responsible. This avoidance is species specific, so that care must be taken describing communities, as the CPR emphasises those species that are preferentially caught, a problem common to many plankton samplers.

Can the Continuous Plankton Recorder (CPR) be used to infer sardine diet?

Knowledge on the impact of climate variability in the diet of planktivorous fish is limited by the laborious work involved in stomach content analysis, impractical for large scale studies. Routine measurements of plankton such as the Continuous Plankton Recorder (CPR) survey provide valuable information of the temporal variation of phyto- and zooplankton prey availability for higher trophic levels. Sardines are a world-wide distributed and commercially important planktivorous fish, at the basis of the pelagic marine food web. Being predominantly non-selective filter-feeders, their diets closely correspond to the water plankton species and a significant relationship was recently found between Sardina pilchardus feeding intensity and remotely sensed chlorophyll alpha . Data of sardine stomach prey composition and CPR were obtained during 2003 for the same location off the west coast of Portugal, an area characterised by strong seasonality of plankton abundance and composition, mainly governed by upwelling events. Phyto- and zooplankton prey in sardine stomachs were identified to the lowest possible taxa and their numerical and volumetric abundance was registered, as well as their contribution to the prey carbon content. The seasonal variation of the abundance and composition of sardine diet was then compared to the abundance and composition of the water plankton obtained with the CPR at the same time and for the same area where the fish were collected, in order to evaluate if CPR data can be used to proxy sardine prey availability and diet composition at large temporal scales.

Genetic analysis of Continuous Plankton Recorder (CPR) samples

Genetic analysis of Continuous Plankton Recorder (CPR) samples is enabling greater taxonomic resolution and the study of plankton population structure. Here, we present some results from the genetic analysis of CPR samples collected in the North Sea and north-eastern Atlantic that reveal the impacts of climate on benthic-pelagic coupling and the food web. We show that pronounced changes in the North Sea meroplankton are related to an increased abundance and spatial distribution of the larvae of the benthic echinoderm, Echinocardium cordatum. Key stages of reproduction in E. cordatum, gametogenesis and spawning, are influenced by winter and spring sea temperature (January-May). A stepwise increase in sea temperature after 1987, which has created warmer conditions earlier in the year, together with increased summer phytoplankton, may benefit the reproduction and survival of this benthic species. Competition between the larvae of E. cordatum and other holozooplanlcton taxa may now be altering the trophodynamics of the summer pelagic ecosystem. In the north-eastern Atlantic the genetic analysis of fish larvae sampled by the CPR has revealed an unprecedented increase in the abundance of juvenile snake pipefish, Entelurus aequoreiis since 2002. We argue that increased sea surface temperatures in winter and spring when the eggs of E. aqueoreus, which are brooded by the male, are developing and the young larvae are growing in the plankton are a likely cause. The increased abundance of this species in Atlantic and adjacent European seas already appears to be influencing the marine food web.

Spatial distribution of key zooplankton species using continuous plankton recorder (CPR) data from the North Atlantic (2000-2009)

The continuous plankton recorder (CPR) survey is an upper layer plankton monitoring program that has regularly collected samples, at monthly intervals, in the North Atlantic and adjacent seas since 1946. Water from approximately 6 m depth enters the CPR through a small aperture at the front of the sampler and travels down a tunnel where it passes through a silk filtering mesh of 270 µm before exiting at the back of the CPR. The plankton filtered on the silk is analyzed in sections corresponding to 10 nautical miles (approx. 3 m**3 of seawater filtered) and the plankton microscopically identified (Richardson et al., 2006 and reference therein). In the present study we used the CPR data to investigate the current basin scale distribution of C. finmarchicus (C5-C6), C. helgolandicus (C5-C6), C. hyperboreus (C5-C6), Pseudocalanus spp. (C6), Oithona spp. (C1-C6), total Euphausiida, total Thecosomata and the presence/absence of Cnidaria and the Phytoplankton Colour Index (PCI). The PCI, which is a visual assessment of the greenness of the silk, is used as an indicator of the distribution of total phytoplankton biomass across the Atlantic basin (Batten et al., 2003). Monthly data collected between 2000 and 2009 were gridded using the inverse-distance interpolation method, in which the interpolated values were the nodes of a 2 degree by 2 degree grid. The resulting twelve monthly matrices were then averaged within the year and in the case of the zooplankton the data were log-transformed (i.e. log10 (x+1).

Molecular analysis of Continuous Plankton Recorder samples, an examination of echinoderm larvae in the North Sea

Analysis of the biological time series of plankton samples collected by the Continuous Plankton Recorder (CPR) in the North Atlantic and North Sea has shown a regime shift in the plankton in this region. Both the distributions of planktonic organisms and their timing of occurrence in the seasonal cycle have changed and these changes appear to ref lect global warming. In the North Sea the planktonic larvae of echinoderms have shown a recent dramatic increase in both relative and absolute abundance and their seasonal peak of occurrence has advanced by 47 days. The identity of the echinoderm larvae involved in this change has, however, remained equivocal. The small size of many organisms like echinoderm larvae combined with incomplete taxonomic keys hinders their visual identification and their fragility often means that useful morphological features are damaged during sampling by the CPR. Here, using new molecular methods applied to CPR samples, we show that planktonic larvae of the benthic Echinocardium cordatum dominate the North Sea plankton. We argue that since this species benefits from mild winters and warmer waters their numerical increase in the plankton is consistent with recent climatic changes that appear to be affecting the wider ecology of this region.

Use of Continuous Plankton Recorder information in support of marine management: applications in fisheries, environmental protection, and in the study of ecosystem response to environmental change

The Continuous Plankton Recorder (CPR) survey was conceived from the outset as a programme of applied research designed to assist the fishing industry. Its survival and continuing vigour after 70 years is a testament to its utility, which has been achieved in spite of great changes in our understanding of the marine environment and in our concerns over how to manage it. The CPR has been superseded in several respects by other technologies, such as acoustics and remote sensing, but it continues to provide unrivalled seasonal and geographic information about a wide range of zooplankton and phytoplankton taxa. The value of this coverage increases with time and provides the basis for placing recent observations into the context of long-term, large-scale variability and thus suggesting what the causes are likely to be. Information from the CPR is used extensively in judging environmental impacts and producing quality status reports (QSR); it has shown the distributions of fish stocks, which had not previously been exploited; it has pointed to the extent of ungrazed phytoplankton production in the North Atlantic, which was a vital element in establishing the importance of carbon sequestration by phytoplankton. The CPR continues to be the principal source of large-scale, long-term information about the plankton ecosystem of the North Atlantic. It has recently provided extensive information about the biodiversity of the plankton and about the distribution of introduced species. It serves as a valuable example for the design of future monitoring of the marine environment and it has been essential to the design and implementation of most North Atlantic plankton research.

Using continuous plankton recorder data

The continuous plankton recorder (CPR) survey is the largest multi-decadal plankton monitoring programme in the world. It was initiated in 1931 and by the end of 2004 had counted 207,619 samples and identified 437 phyto- and zooplankton taxa throughout the North Atlantic. CPR data are used extensively by the research community and in recent years have been used increasingly to underpin marine management. Here, we take a critical look at how best to use CPR data. We first describe the CPR itself, CPR sampling, and plankton counting procedures. We discuss the spatial and temporal biases in the Survey, summarise environmental data that have not previously been available, and describe the new data access policy. We supply information essential to using CPR data, including descriptions of each CPR taxonomic entity, the idiosyncrasies associated with counting many of the taxa, the logic behind taxonomic changes in the Survey, the semi-quantitative nature of CPR sampling, and recommendations on choosing the spatial and temporal scale of study. This forms the basis for a broader discussion on how to use CPR data for deriving ecologically meaningful indices based on size, functional groups and biomass that can be used to support research and management. This contribution should be useful for plankton ecologists, modellers and policy makers that actively use CPR data.

Distribution and abundance of sardine (Sardina pilchardus) eggs in the English Channel from Continuous Plankton Recorder sampling

Continuous Plankton Recorder (CPR) samples from the English Channel and adjacent Celtic shelf, taken over the period 1958-1980, were analysed for sardine (Sardina pilchardus) eggs. Results showed the progression of sardine spawning along the English Channel from west to east from March to August and a return from east to west from September to November. This corresponds with the two seasonal peaks of sardine egg abundance in the western Channel: the main summer peak being in May/June, with a smaller autumn peak in October/November. Long-term changes in sardine egg abundance in CPR samples showed a decline in summer spawning from the late 1960s, but no clear trend in autumn-spawned egg abundance. Similar patterns were observed in the numbers of sardine eggs sampled by conventional plankton net tows at the time-series Station L5 off Plymouth. This supports the use of the longer time-series of sardine egg data at L5 as being representative of a wider area and emphasizes the importance in continuation of the L5 time-series.

Continuous Plankton Recorder Database: evolution, current uses and future directions.

The Continuous Plankton Recorder (CPR) survey, operated by the Sir Alister Hardy Foundation for Ocean Science (SAHFOS), is the largest plankton monitoring programme in the world and has spanned >70 yr. The dataset contains information from ~200 000 samples, with over 2.3 million records of individual taxa. Here we outline the evolution of the CPR database through changes in technology, and how this has increased data access. Recent high-impact publications and the expanded role of CPR data in marine management demonstrate the usefulness of the dataset. We argue that solely supplying data to the research community is not sufficient in the current research climate; to promote wider use, additional tools need to be developed to provide visual representation and summary statistics. We outline 2 software visualisation tools, SAHFOS WinCPR and the digital CPR Atlas, which provide access to CPR data for both researchers and non-plankton specialists. We also describe future directions of the database, data policy and the development of visualisation tools. We believe that the approach at SAHFOS to increase data accessibility and provide new visualisation tools has enhanced awareness of the data and led to the financial security of the organisation; it also provides a good model of how long-term monitoring programmes can evolve to help secure their future.

Is observed variability in the long-term results of the Continuous Plankton Recorder survey a response to climate change?

In the more than 50 years that the Continuous Plankton Recorder (CPR) survey has operated on a regular monthly basis in the north-east Atlantic and North Sea, large changes have been witnessed in the planktonic ecosystem. These changes have taken the form of long-term trends in abundance for certain species or stepwise changes for others, and in many cases are correlated with a mode of climatic variability in the North Atlantic, either: (1) the North Atlantic Oscillation (NAO), a basin-scale atmospheric alteration of the pressure field between the Azores high pressure cell and the Icelandic Low; or (2) the Gulf Stream Index (GSI), which measures the latitudinal position of the north wall of the Gulf Stream. Recent work has shown that the changes in the GSI are coupled with the NAO and Pacific Southern Oscillation with a 2 year lag. The plankton variability is also possibly linked to changes observed in the distribution and flux of water masses in the surface, intermediate and deep waters of the North Atlantic. For example, in the last two decades, the extent and location of the formation of North Atlantic Deep Water, Labrador Sea Intermediate Water and Norwegian Sea intermediate and upper-layer water has altered considerably. This paper discusses the extent to which observed changes in plankton abundance and distribution may be linked to this basin-scale variability in hydrodynamics. The results are also placed within the context of global climate warming and the possible effects of the observed melting of Arctic permafrost and sea ice on the subpolar North Atlantic.

Structure of phytoplankton (Continuous Plankton Recorder and SeaWiFS) and impact of climate in the Northwest Atlantic Shelves

All marine organisms are affected to some extent by the movement and thermal properties of oceanic currents. However phytoplankton, because of its small size is most directly coupled to the physical environment. The intense hydrodynamic activity observed in the Northwest Atlantic Shelves Province makes this region especially intriguing from the point of view of physical-biological interactions. In the present work, remote sensed data of Sea Surface Height (SSH) anomalies, Sea-surface chlorophyll a concentrations (SeaWiFS), and Sea Surface Temperature (SST) are used to complement the Continuous Plankton Recorder (CPR) survey that continuously sampled a route between Norfolk (Virginia, USA; 39° N, 71° W) and Argentia (Newfoundland; 47° N, 54° W) over the period 1995–1998. Over this period, we examined physical structures (i.e. SST and SSH) and climatic forcing associated with space-time phytoplankton structure. Along this route, the phytoplankton structures were mainly impacted by the changes in surface flow along the Scotian Shelf rather than significantly influenced by the mesoscale features of the Gulf Stream. These changes in water mass circulation caused a drop in temperature and salinity along the Scotian Shelf that induced changes in phytoplankton and zooplankton abundance.

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.

The Continuous Plankton Recorder: towards a global perspective

The Continuous Plankton Recorder (CPR) began its first routine deployment to collect plankton samples in the North Sea in 1931. The better part of a century after that event, the Recorder is used for sampling plankton widely in many oceans. The CPR is designed to be towed behind ships of opportunity (such as commercial and passenger vessels) and to collect plankton samples along the ship’s path. The samples provide information on the broad scale spatial distributions of larger hard-shelled phytoplankton and robust mesozooplankton organisms (Richardson et al., 2006). Changes in these distributions provide indices for the biological health of the ocean including those required for fisheries and for assessment of climate change impacts.