Identification of the copepodite developmental stages of twenty-six North Atlantic copepods. Occasional Publication of the Marine Biological Association 21.

In zooplankton copepod studies there is often the requirement to be able to identify the six copepodite developmental stages of different species, or to know their body dimensions. However, this information is not available for many species, or is dispersed through the literature. This guide gathers together both original and previously published information on morphology and measurements for the stages of twenty-six common North Atlantic copepod species and tabulates them in a standard format. For each species additional notes useful in their identification are also given.

Case history and persistence of the non-indigenous diatom Coscinodiscus wailesii in the north-east Atlantic

The introduction of non-indigenous marine plankton species can have a considerable ecological and economic effect on regional systems. Their presence, however, can go unnoticed until they reach nuisance status and as a consequence few case histories exist containing information on their initial appearance and their spatio-temporal patterns. Here we report on the occurrence of the non-indigenous diatom Coscinodiscus wailesii in 1977 in the English Channel, its subsequent geographical spread into European shelf seas, and its persistence as a significant member of the diatom community in the north-east Atlantic from 1977-1995.

Long-term and regional variability of phytoplankton biomass in the Northeast Atlantic (1960-1995)

Long-term regional changes in phytoplankton biomass in the Northeast Atlantic and North Sea are investigated using data from the Continuous Plankton Recorder survey. During the last decade there have been large changes in the long-term variation in phytoplankton biomass in the Northeast Atlantic and North Sea. Most regions, particularly in the North Sea, have shown a considerable increase in phytoplankton biomass while the opposite pattern was seen in the northern oceanic region of the Northeast Atlantic. These different spatial responses show similar patterns of change to the decadal variability in sea surface temperature influenced by the North Atlantic Oscillation index. Two rare oceanographic events and their relationship to the interannual changes in phytoplankton biomass are discussed. The results highlight the importance of maintaining long-term biological monitoring programmes to assess the biological responses to slow oceanic/atmospheric processes and to rare or episodic physical events.

Decadal basin-scale changes in diatoms, dinoflagellates, and phytoplankton color across the North Atlantic

The Continuous Plankton Recorder (CPR) survey has been used to characterize phytoplankton and zooplankton space-time dynamics in the North Sea since 1931 and in the North Atlantic since 1939. Phytoplankton biomass is assessed from these samples by visual assessment of the green color of the silk mesh, the Phytoplankton Color Index (PCI), and the total count of diatoms and dinoflagellates. Species with a frequency of occurrence greater than 1% in the samples are used as indicator species of the community. We investigated (1) long-term fluctuations of phytoplankton biomass, total diatoms, and total dinoflagellates; (2) geographical variation of patterns; (3) the relationship between phytoplankton and climate forcing in the North Atlantic CPR samples; (4) the relative contribution of diatoms and dinoflagellates to the PCI; and (5) the fluctuations of the dominant species over the period of survey to provide more information on the processes linking climate to changes in the phytoplankton community. As a result of the differences in microscopic analysis methods prior to 1958, our analyses were conducted for the period ranging from 1958 to 2002. The North Atlantic was divided into six regions identified through bathymetric criteria and separated along a North-South axis. Based on 12 monthly time series, we demonstrate increasing trends in PCI and total dinoflagellates and a decrease in total diatoms.

Regional climate change and harmful algal blooms in the northeast Atlantic

We investigated long-term spatial variability in a number of Harmful Algal Blooms (HABs) in the northeast Atlantic and North Sea using data from the Continuous Plankton Recorder. Over the last four decades, some dinoflagellate taxa showed pronounced variation in the south and east of the North Sea, with the most significant increases being restricted to the adjacent waters off Norway. There was also a general decrease along the eastern coast of the United Kingdom. The most prominent feature in the interannual bloom frequencies over the last four decades was the anomalously high values recorded in the late 1980s in the northern and central North Sea areas. The only mesoscale area in the northeast Atlantic to show a significant increase in bloom formation over the last decade was the Norwegian coastal region. The changing spatial patterns of HAB taxa and the frequency of bloom formation are discussed in relation to regional climate change, in particular, changes in temperature, salinity, and the North Atlantic Oscillation (NAO). Areas highly vulnerable to the effects of regional climate change on HABs are Norwegian coastal waters and the Skagerrak. Other vulnerable areas include Danish coastal waters, and to a lesser extent, the German and Dutch Bight and the northern Irish Sea. Quite apart from eutrophication, our results give a preview of what might happen to certain HAB genera under changing climatic conditions in temperate environments and their responses to variability of climate oscillations such as the NAO.

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.

Increased blooms of a dinoflagellate in the NW Atlantic

Sampling by the Continuous Plankton Recorder (CPR) over the NW Atlantic from 1960 to 2000 has enabled long-term studies of the larger components of the phytoplankton community, highlighting various changes, particularly during the 1990s. Analysis of an index of phytoplankton biomass, the Phytoplankton Colour Index (PCI) has revealed an increase over the past decade, most marked during the winter (December to February) months. Examination of the structure of the community using multiple linear-regression models indicates that the winter phytoplankton community composition has changed markedly in the 1990s compared to the 1960s. One phytoplankter, the dinoflagellate Ceratium arcticum (Cleve), has undergone dramatic changes in abundance during this period, with pronounced large winter blooms and decreased autumnal levels, and its contribution to the Phytoplankton Colour index values has increased significantly. Other dominant species in the phytoplankton community, both diatoms and dinoflagellates, did not show the same variations over the examined time period. It is suggested that the response of C. arcticum is probably a result of previously reported changes in stratification in the NW Atlantic, due to dynamic hydro-climatic (freshening and cooling) events.

Differential contribution of diatoms and dinoflagellates to phytoplankton biomass in the NE Atlantic Ocean and the North Sea

Sampling by the continuous plankton recorder (CPR) survey over the North Atlantic Ocean and the North Sea has enabled long-term studies of phytoplankton biomass. Analysis of an index of phytoplankton biomass, the phytoplankton colour index (PCI), has previously shown an increase in phytoplankton biomass in the NE Atlantic. In the current study, further investigations were conducted to determine the contribution of diatom and dinoflagellate cell counts to the PCI, their fluctuations over the last 45 yr and their geographical variations in the eastern North Atlantic and the North Sea. An increased contribution of dinoflagellates to the PCI was revealed over the south NE Atlantic and the northern North Sea. In contrast, the contribution of diatoms decreased in the north NE Atlantic and the northern North Sea. No discernible trends were found in the other regions of the North Sea. The relative contributions of diatoms and dinoflagellates to the PCI led to the identification of 3 geographically distinct dynamic regimes in the diatom/dinoflagellate dynamics in the NE Atlantic and the North Sea. Finally, it is stressed that the discrepancy observed in the patterns of PCI and diatom and dinoflagellate cell counts suggests that changes in PCI do not reflect changes in the community structure and that the exclusive use of PCI is not adequate to investigate the long-term trends in the trophic link between phytoplankton and herbivorous zooplankton.

An individual-based model of the early life history of mackerel (Scomber scombrus) in the eastern North Atlantic, simulating transport, growth and mortality

The main purpose of this paper is to provide the core description of the modelling exercise within the Shelf Edge Advection Mortality And Recruitment (SEAMAR) programme. An individual-based model (IBM) was developed for the prediction of year-to-year survival of the early life-history stages of mackerel (Scomber scombrus) in the eastern North Atlantic. The IBM is one of two components of the model system. The first component is a circulation model to provide physical input data for the IBM. The circulation model is a geographical variant of the HAMburg Shelf Ocean Model (HAMSOM). The second component is the IBM, which is an i-space configuration model in which large numbers of individuals are followed as discrete entities to simulate the transport, growth and mortality of mackerel eggs, larvae and post-larvae. Larval and post-larval growth is modelled as a function of length, temperature and food distribution; mortality is modelled as a function of length and absolute growth rate. Each particle is considered as a super-individual representing 10 super(6) eggs at the outset of the simulation, and then declining according to the mortality function. Simulations were carried out for the years 1998-2000. Results showed concentrations of particles at Porcupine Bank and the adjacent Irish shelf, along the Celtic Sea shelf-edge, and in the southern Bay of Biscay. High survival was observed only at Porcupine and the adjacent shelf areas, and, more patchily, around the coastal margin of Biscay. The low survival along the shelf-edge of the Celtic Sea was due to the consistently low estimates of food availability in that area.

Spawning season and temperature relationships for sardine (Sardina pilchardus) in the eastern North Atlantic

Spawning temperature preferences for sardine (Sardina pilchardus) in the eastern North Atlantic were determined from field data. These were compared with climatological temperature cycles (1986-2002) derived from satellite data by geographical region, to predict spawning seasons. Optimum spawning temperatures were determined as 14.0-15.0oC from the English Channel to Portugal and 16.0–18.0oC for all north-west African regions. Spawning seasons were closely related to the general latitudinal trend of the annual temperature cycle, with modification by upwelling in the western Iberian and north-west African regions. Some differences between temperature-based spawning season predictions and field observations were related to variations in seasonal plankton production. Correlations in the annual time-series of favourable spawning temperatures suggested relatively strong linkages between the southern areas from Portugal to Senegal. There was no consistent relationship between annual variations in duration of temperature-predicted spawning seasons and observed field abundance of eggs.

Congruent trends in long-term zooplankton decline in the north-east Atlantic and basking shark (Cetorhinus maximus ) fishery catches off west Ireland

Trends in basking shark (Cetorhinus maximus) fishery catches off Achill Island, west Ireland between 1949 and 1975 were examined in relation to zooplankton (total copepod) abundance in four adjacent sea areas over a 27-year period. The numbers of basking sharks caught and copepod abundance showed downward trends and were positively correlated (r-value range, 0.44–0.74). A possible explanation for the downward trend in shark catches was that progressively fewer basking sharks occurred there between 1956 and 1975 because fewer copepods, their food resource, occurred near the surface off west Ireland over the same period. We suggest that the decline in basking sharks may have been due to a distributional shift of sharks to more productive areas, rather than a highly philopatric, localized stock that was over-exploited.

Ecological effects of the North Atlantic oscillation

Climatic oscillations as reflected in atmospheric modes such as the North Atlantic Oscillation (NAO) may be seen as a proxy for regulating forces in aquatic and terrestrial ecosystems. Our review highlights the variety of climate processes related to the NAO and the diversity in the type of ecological responses that different biological groups can display. Available evidence suggests that the NAO influences ecological dynamics in both marine and terrestrial systems, and its effects may be seen in variation at the individual, population and community levels. The ecological responses to the NAO encompass changes in timing of reproduction, population dynamics, abundance, spatial distribution and interspecific relationships such as competition and predator-prey relationships. This indicates that local responses to large-scale changes may be more subtle than previously suggested. We propose that the NAO effects may be classified as three types: direct, indirect and integrated. Such a classification will help the design and interpretation of analyses attempting to relate ecological changes to the NAO and, possibly, to climate in general.

Biodiversity of North Atlantic and North Sea calanoid copepods

Spatial patterns in pelagic biodiversity are the result of factors acting from a global to a local scale. The global patterns have been studied intensively using taxa such as foraminifera and euphausiids. However, these studies do not allow direct comparisons of neritic and oceanic regions or examination of relationships between local and regional patterns of pelagic diversity. Here we present a map of the diversity of calanoid copepods, a key planktonic group, summarising 40 yr of continuous monthly investigations in the North Atlantic and North Sea. The high number of samples (168 162) allowed mesoscale patterns in diversity to be detected for the first time at an ocean-basin level. Our results demonstrate pronounced local spatial variability in planktonic diversity and refine previous global studies at a lower resolution. They form a baseline at which long-term changes in planktonic diversity can be better assessed and ecosystem management plans implemented.