Effects of formalin preservation on stable carbon and nitrogen isotope signatures in Calanoid copepods: implications for the use of Continuous Plankton Recorder Survey samples in stable isotope analyses

Preserved and archived organic material offers huge potential for the conduct of retrospective and long-term historical ecosystem reconstructions using stable isotope analyses, but because of isotopic exchange with preservatives the obtained values require validation. The Continuous Plankton Recorder (CPR) Survey is the most extensive long-term monitoring program for plankton communities worldwide and has utilised ships of opportunity to collect samples since 1931. To keep the samples intact for subsequent analysis, they are collected and preserved in formalin; however, previous studies have found that this may alter stable carbon and nitrogen isotope ratios in zooplankton. A maximum ~0.9‰ increase of δ15N and a time dependent maximum ~1.0‰ decrease of δ13C were observed when the copepod, Calanus helgolandicus, was experimentally exposed to two formalin preservatives for 12 months. Applying specific correction factors to δ15N and δ13C values for similarly preserved Calanoid species collected by the CPR Survey within 12 months of analysis may be appropriate to enable their use in stable isotope studies. The isotope values of samples stored frozen did not differ significantly from those of controls. Although the impact of formalin preservation was relatively small in this and other studies of marine zooplankton, changes in isotope signatures are not consistent across taxa, especially for δ15N, indicating that species-specific studies may be required. Copyright © 2011 John Wiley & Sons, Ltd.

Comparing copepod time-series in the north of Spain: Spatial autocorrelation of community composition

Four time-series of copepod species biomass in the north of Spain were contrasted to demonstrate spatial autocorrelation of local communities and their responses to short-term local and regional variability in oceanographic conditions. The series represented coastal and oceanic environments along a marked gradient of influence of seasonal upwelling from Galicia to the Mar Cantábrico (S Bay of Biscay), and each one included at least 10 years of continuous data collected at monthly frequency. Community composition (i.e. species number and diversity) was very consistent through the region, but local variations in the presence of new species and the relative proportions of common species allowed for the characterisation of the response to the environment at each site. Small-sized species were more frequent near the coast. A few species, however, captured the main patterns of variability in all series. Calanus helgolandicus and Acartia (mainly Acartia clausi) were generally the main contributors to total biomass, while other species as Paracalanus parvus and Clausocalanus spp. were important only at some locations. Most copepod indices were positively correlated with upwelling, either considering the whole community (biomass, species richness and diversity) or individual species, but only in the coastal series analysed since 1991. Copepods in the nearby ocean, however, showed negative correlations with upwelling in the period 1960–1986. The effects of upwelling may have been modulated by local factors, as showed by the increases in biomass, number of species and diversity in associations with increases in sea surface temperature in Galicia, while in the Mar Cantábrico only the warming-tolerant species increased and those typical of upwelling decreased. Density stratification of the water column was associated with decreases in total copepod biomass in Galicia, while it favoured the increase in species richness in the Mar Cantábrico. Nearly all significant responses of copepods to environmental variability were delayed by up to 5 months, showing the importance of considering time-lags in the analysis of temporal responses of zooplankton.

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.

Long-Term Retrospective Analysis of Mackerel Spawning in the North Sea: A New Time Series and Modeling Approach to CPR Data

We present a unique view of mackerel (Scomber scombrus) in the North Sea based on a new time series of larvae caught by the Continuous Plankton Recorder (CPR) survey from 1948-2005, covering the period both before and after the collapse of the North Sea stock. Hydrographic backtrack modelling suggested that the effect of advection is very limited between spawning and larvae capture in the CPR survey. Using a statistical technique not previously applied to CPR data, we then generated a larval index that accounts for both catchability as well as spatial and temporal autocorrelation. The resulting time series documents the significant decrease of spawning from before 1970 to recent depleted levels. Spatial distributions of the larvae, and thus the spawning area, showed a shift from early to recent decades, suggesting that the central North Sea is no longer as important as the areas further west and south. These results provide a consistent and unique perspective on the dynamics of mackerel in this region and can potentially resolve many of the unresolved questions about this stock

A comparison of two zooplankton time series data collected in the Gulf of Maine

Interannual and seasonal trends of zooplankton abundance and species composition were compared between the Bongo net and Continuous Plankton Recorder (CPR) time series in the Gulf of Maine. Data from 5799 Bongo and 3118 CPR samples were compared from the years 1978–2006. The two programs use different sampling methods, with the Bongo time series composed of bimonthly vertically integrated samples from locations throughout the region, while the CPR was towed monthly at 10 m depth on a transect that bisects the region. It was found that there was a significant correlation between the interannual (r = 0.67, P < 0.01) and seasonal (r = 0.95, P < 0.01) variability of total zooplankton counts. Abundance rankings of individual taxa were highly correlated and temporal trends of dominant copepods were similar between samplers. Multivariate analysis also showed that both time series equally detected major shifts in community structure through time. However, absolute abundance levels were higher in the Bongo and temporal patterns for many of the less abundant taxa groups were not similar between the two devices. The different mesh sizes of the samplers probably caused some of the discrepancies; but diel migration patterns, damage to soft bodied animals and avoidance of the small CPR aperture by some taxa likely contributed to the catch differences between the two devices. Nonetheless, Bongo data presented here confirm the previously published patterns found in the CPR data set, and both show that the abundance increase of the 1990s has been followed by average to below average levels from 2002 to 06.

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.

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.

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.

A biological consequence of reducing Arctic ice cover: arrival of the Pacific diatom Neodenticula seminae in the North Atlantic for the first time in 800,000 years

The Continuous Plankton Recorder survey has monitored plankton in the Northwest Atlantic at monthly intervals since 1962, with an interegnum between 1978 and 1990. In May 1999, large numbers of the Pacific diatom Neodenticula seminae were found in Continuous Plankton Recorder (CPR) samples in the Labrador Sea as the first record in the North Atlantic for more than 800 000 years. The event coincided with modifications in Arctic hydrography and circulation, increased flows of Pacific water into the Northwest Atlantic and in the previous year the exceptional occurrence of extensive ice-free water to the North of Canada. These observations indicate that N. seminae was carried in a pulse of Pacific water in 1998/early 1999 via the Canadian Arctic Archipelago and/or Fram Strait. The species occurred previously in the North Atlantic during the Pleistocene from similar to 1.2 to similar to 0.8 Ma as recorded in deep sea sediment cores. The reappearance of N. seminae in the North Atlantic is an indicator of the scale and speed of changes that are taking place in the Arctic and North Atlantic oceans as a consequence of regional climate warming. Because of the unusual nature of the event it appears that a threshold has been passed, marking a change in the circulation between the North Pacific and North Atlantic Oceans via the Arctic. Trans-Arctic migrations from the Pacific into the Atlantic are likely to occur increasingly over the next 100 years as Arctic ice continues to melt affecting Atlantic biodiversity and the biological pump with consequent feedbacks to the carbon cycle.

Ecological Investigations with the Continuous Plankton Recorder: The Phytoplankton in the Southern North Sea. 1932-37.

1. The changes in the composition and distribution of the plankton of the southern North Sea have been investigated month by month, from June 1932 to December 1937; the present report deals with the phytoplankton. The survey was carried out by the Continuous Plankton Recorder, towed at a standard depth of 10 metres, by ships on regular steamship lines across the North Sea from Hull towards the Skagerrak, to Bremen and to Rotterdam, and later between London and Esbjerg. 2. The material and methods are described, together with a discussion on the validity of this type of survey and some comparison of its results with those obtained by other methods (pp. 76-86). 3. Particular attention has been paid to Rhizosolenia styliformis (pp. 92- 107), Biddulphia sinensis (pp. 108-115), Phaeocystis (pp. 149-153), and the Dinoflagellates (pp. 134-149); of these the first three are known to be of particular importance in relation to the herring fisheries. More generalised data are available for the principal diatoms other than R. styliformis and B. sinensis (pp. 116-134). 4. The main part of the work is an ecological study of the phytoplankton changes in time and space over the 5½ years. Each year is marked by some distinct variations in the abundance and the times of increase, maximum numbers and decline as recorded in the different forms. These variations in the annual cycles are compared on the different lines by a series of graphs arranged against a time scale of months, a set for each year being placed side by side (Plates I-XXI). More detailed studies by more frequent records were made in the autumns of 1934, 1935, 1936 and 1937 (cf. Figs. 3 and 4). The changes in spatial distribution are shown by a series of monthly maps arranged in a similar manner for each year (Plates XXII-LXIV). These intensive studies of the changes in time and space are also intended to form the basis for correlations with other features in the general ecology of the area (e. g. the zooplankton, hydrology, meteorology and fisheries) to be made in later publications. 5. Whilst each form has shown its own peculiar features, a trend towards a general increase in the phytoplankton as a whole has been observed during the period, although the years 1934 and 1936 have in some respects shown deviations and regressive features, and not all organisms have revealed the same trend. The possible relation of this gradual trend to other events observed in recent years in these and neighbouring waters is discussed (pp. 162-167). 6. The application of these results to the study of patchiness (pp. 154-158), inter-relationships in the plankton (pp. 159-160) and to water movements (pp. 160-162) is briefly discussed.

Ecological Investigations with the Continuous Plankton Recorder: The Zooplankton (other than Copepoda and Young Fish) in the Southern North Sea, 1932-37.

I. The report describes the main monthly changes in the distribution and abundance of the zooplankton, other than Copepoda and young fish (dealt with in separate reports), over the southern part of the North Sea from 1932 to 1937. The work is part of the survey carried out by Continuous Plankton Recorders towed at a depth of 10 metres on regular steamship lines between England and the Continent. 2. The limitations to the sampling method are discussed, and it is shown to be unsuitable for recording Mysidacea and Euphausiacea on account of their marked diurnal variation due presumably to vertical migration; they are omitted from the report. 3. The changing distribution of Sagitta, Limacina, Clione, Lamellibranch larvae, Cladocera, Caprellid Amphipoda, Decapod larvae, Echinoderm larvae and Oikopleura are shown in a series of monthly charts while their seasonal fluctuations are compared in time-chart histograms. 4. The Alima larvae of Squilla are recorded on a few occasions in the regions where the Channel opens into the North Sea. 5. The distributional characteristics of the different forms, i.e. their tendencies to even or " patchy " production, are compared.

Continuous Plankton Records: General Introduction to the 1938-39 Survey

The Bulletins in this volume, except the last, deal entirely with the results of this expanded survey of 1938 and 1939. They show for the first time, month by month, the main changes in the plankton over practically the whole of the North Sea for a year and eight months. They form an important basis for comparison with the results of the post-war survey, revived on an even more extensive scale and to be described in later volumes.

Comparing whole-link travel time models used in DTA

In a model commonly used in dynamic traffic assignment the link travel time for a vehicle entering a link at time t is taken as a function of the number of vehicles on the link at time t. In an alternative recently introduced model, the travel time for a vehicle entering a link at time t is taken as a function of an estimate of the flow in the immediate neighbourhood of the vehicle, averaged over the time the vehicle is traversing the link. Here we compare the solutions obtained from these two models when applied to various inflow profiles. We also divide the link into segments, apply each model sequentially to the segments and again compare the results. As the number of segments is increased, the discretisation refined to the continuous limit, the solutions from the two models converge to the same solution, which is the solution of the Lighthill, Whitham, Richards (LWR) model for traffic flow. We illustrate the results for different travel time functions and patterns of inflows to the link. In the numerical examples the solutions from the second of the two models are closer to the limit solutions. We also show that the models converge even when the link segments are not homogeneous, and introduce a correction scheme in the second model to compensate for an approximation error, hence improving the approximation to the LWR model.