Spatial Heterogeneity and Genetic Variation in the Copepod Neocalanus Cristatus Along Two Transects in the North Pacific Sampled By the Continuous Plankton Recorder

We present a macrogeographic study of spatial heterogeneity in an important subarctic Pacific copepod and describe the first genetic analysis of population structure using Continuous Plankton Recorder (CPR) samples. Samples of Neocalanus cristatus were collected at a constant depth of similar to 7 m from two CPR tow-routes, (i) an east-west similar to 6500-km transect from Vancouver Island, Canada to Hokkaido Island, Japan, and (ii) a north-south transect of similar to 2250 km from Anchorage, Alaska to Tacoma, Washington. Analysis of these samples revealed three features of the biology of N. cristatus. First, N. cristatus undergoes small-scale diel vertical migration that is larger among stages CV- adult (3-6 times more abundant at 7 m at night), than stages CI-CIV (only 2-4 times higher at night). Secondly, while there were no regions where N. cristatus did not appear, each transect sampled a few large-scale macrogeographic patches. Thirdly, an analysis of molecular variation, using a partial sequence of the N. cristatus cytochrome oxidase I gene, revealed that 7.3% (P < 0.0001) of the total genetic variation among N. cristatus sampled from macrogeographic patches by the CPR could be explained by spatial heterogeneity. We suggest that spatial heterogeneity at macrogeographic scales may be important in plankton evolution.

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.

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.

Fish data from the Continuous Plankton Recorder survey

The Continuous Plankton Recorder (CPR) survey has been sampling plankton in the North Sea since 1931. However the identification of the larval and juvenile fish taken in the survey has not been a part of the routine analysis of the samples. Specialist analysis of the fish was carried out between 1948 and the early 1980s but the data were available as hard copy only. As part of MarBEF, data on >60 taxa from 1948 to 1972 have been entered on a database which is now linked to EUROBIS to show the biogeographical information and the data are available for general research. Examples of the data are shown. These data provide a background on the variability of fish stocks before the recent period of rapid warming and in some cases before significant fisheries developed. Data for subsequent years will be made available as possible and work is now underway to bring the analysis up to date. This will provide time series over six decades.

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.

Variation among northeast Atlantic regions in the responses of zooplankton to climate change: Not all areas follow the same path

Broad scale climate forcing can interact with local environmental processes to affect the observed ecological phenomena. This causes potential problems of over-extrapolation for results from a limited number of sites or the averaging out of region-specific responses if data from too wide an area are combined. In this study, an area similar in extent to the Celtic Biscay Large Marine Ecosystem, but including off-shelf areas, was partitioned using clustering of satellite chlorophyll (chl-a) measurements. The resulting clusters were used to define areas over which to combine copepod data from the Continuous Plankton Recorder. Following filtering due to data limitations, nine regions were defined with sufficient records for analysis. These regions were consistent with known oceanographic structure in the study area. Off-shelf regions showed a progressively later timing in the seasonal peak of chl-a measurements moving northwards. Generalised additive models were used to estimate seasonal and multiannual signals in the adult and juvenile stages of Calanus finmarchicus, C. helgolandicus and the Paracalanus–Pseudocalanus group. Associations between variables (sea surface temperature (SST), phenology and annual abundance) differed among taxonomic groups, but even within taxonomic groups, relationships were not consistent across regions. For example, in the deep waters off Spain and Portugal the annual abundance of Calanus finmarchicus has a weak positive association with SST, in contrast to the pattern in most other regions. The regions defined in this study provide an objective basis for investigations into the long term dynamics of plankton populations and suggest suitable sub regions for deriving pelagic system indicators.

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.

Using the Continuous Plankton Recorder to investigate the absence of North Atlantic right whales (Eubalaena glacialis) from the Roseway Basin foraging ground

North Atlantic right whales (Eubalaena glacialis) were absent from Roseway Basin, located off southeastern Nova Scotia, for a 7-year period (1993–1999). The objective of this study was to examine the availability of the right whale's main prey, Calanus finmarchicus, in Roseway Basin during those 7 years to determine if the whales’ absence was due to inadequate prey resources. Since we had no historical data on zooplankton abundances at depth on the Scotian Shelf, near-surface zooplankton abundance data from the Continuous Plankton Recorder were used to infer water-column abundances. In addition, environmental parameters that are often correlated with high zooplankton concentrations were examined. The hypotheses tested were that changes in these parameters would be detectable between three time periods: pre-1993, 1993–1999 and post-1999. Calanus finmarchicus abundance was found to be lowest during 1993–1999, suggesting that right whales were not foraging in Roseway Basin because of the near-absence of their main prey species. Decreased in situ salinity and density proved to be indicators of the changes in circulation in the 1990s that may have affected the advection of C. finmarchicus onto the Scotian Shelf.

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.