An overview of Calanus helgolandicus ecology in European waters

We review current knowledge and understanding of the biology and ecology of the calanoid copepod Calanus helgolandicus in European waters, as well as provide a collaborative synthesis of data from 18 laboratories and 26 sampling stations in areas distributed from the northern North Sea to the Aegean and Levantine Seas. This network of zooplankton time-series stations has enabled us to collect and synthesise seasonal and multi-annual data on abundance, body size, fecundity, hatching success and vertical distribution of C. helgolandicus. An aim was to enable comparison with its congener Calanus finmarchicus, which has been studied intensively as a key component of European and north east Atlantic marine ecosystems. C. finmarchicus is known to over-winter at depth, whereas the life-cycle of C. helgolandicus is less well understood. Overwintering populations of C. helgolandicus have been observed off the Atlantic coast between 400 and 800 m, while in the Mediterranean there is evidence of significant deep-water populations at depths as great as 4200 m. The biogeographical distribution of C. helgolandicus in European coastal waters covers a wide range of habitats, from open ocean to coastal environments, and its contribution to mesozooplankton biomass ranges from 6% to 93%. Highest abundances were recorded in the Adriatic and off the west coast of Spain. C. helgolandicus is generally found in 9-20 C water, with maximum abundances from 13-17 C. In contrast, C. finmarchicus is found in cooler water between 0 and 15 C, with peak abundances from 0 to 9 C. As water has warmed in the North Atlantic over recent decades, the range of C. helgolandicus and its abundance on the fringes of its expanding range have increased. This review will facilitate development of population models of C. helgolandicus. This will not only help answer remaining questions but will improve our ability to forecast future changes, in response to a warming climate, in the abundance and distribution of this important species.

Pulses in the eastern margin current and warmer water off the north west European shelf linked to North Sea ecosystem changes

The North Sea ecosystem has recently undergone dramatic changes, observed as altered biomass of individual species spanning a range of life forms from algae to birds, with evidence for an approximate doubling in the abundance of both phytoplankton and benthos as part of a regime shift after 1987. Remarkably, these changes, in part recorded in the Phytoplankton Colour Index of the Continuous Plankton Recorder (CPR) survey, are notable as episodic shifts occurring in 1988/89 and 1998 imposed on a gradual decadal trend. These biological events are shown to be a response to coincident changes in oceanic input and water temperature. Geostrophic transports have been calculated from a hydrographic section across the Rockall Trough, and a time series of seasurface temperature derived from satellite observations. The 2 pulses of oceanic incursion into the North Sea in circa 1988 and 1998 coincided with strong northward advection of anomalously warm water at the edge of the continental shelf.

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.

Spatial, seasonal and long-term fluctuations of plankton in relation to hydroclimatic features in the English Channel, Celtic Sea and Bay of Biscay

Plankton collected by the Continuous Plankton Recorder (CPR) survey were investigated for the English Channel, Celtic Sea and Bay of Biscay from 1979 to 1995. The main goal was to study the relationship between climate and plankton and to understand the factors influencing it. In order to take into account the spatial and temporal structure of biological data, a three-mode principal component analysis (PCA) was developed. It not only identified 5 zones characterised by their similar biological composition and by the seasonal and inter-annual evolution of the plankton, it also made species associations based on their location and year-to-year change. The studied species have stronger year-to-year fluctuations in abundance over the English Channel and Celtic Sea than the species offshore in the Bay of Biscay. The changes in abundance of plankton in the English Channel are negatively related to inter-annual changes of climatic conditions from December to March (North Atlantic Oscillation [NAO] index and air temperature). Thus, the negative relationship shown by Fromentin and Planque (1996; Mar Ecol Prog Ser 134:111-118) between year-to-year changes of Calanus finmarchicus abundance in the northern North Atlantic and North Sea and NAO was also found for the most abundant copepods in the Channel. However, the hypothesis proposed to explain the plankton/NAO relationship is different for this region and a new hypothesis is proposed. In the Celtic Sea, a relationship between the planktonic assemblage and the air temperature was detected, but it is weaker than for the English Channel. No relationship was found for the Bay of Biscay. Thus, the local physical environment and the biological composition of these zones appear to modify the relationship between winter climatic conditions and the year-to-year fluctuations of the studied planktonic species. This shows, therefore, that the relationship between climate and plankton is difficult to generalise.

North Sea Ecosystem: Status Report

The North Sea is a dynamic large marine ecosystem which is bordered by a dense coastal population, contains a productive oil and gas province, has a dense shipping network and has one of the most productive fisheries in the world. An assessment of the state of health of the North Sea was initiated in 1987 as part of a developing series of international initiatives at Ministerial level to address concerns over the impact of these activities on the marine ecosystem. Four North Sea Ministerial Conferences (1984, 1987, 1990, 1995) and an Intermediate Ministerial Meeting (1993) have been held to date to develop a harmonized approach to the sustainable management of the North Sea. In 1988 at the request of Ministers a North Sea Task Force was established to co-ordinate work leading to the production of a Quality Status Report (QSR) on the North Sea in December 1993. In recognition of the large geographical and ecological diversity exhibited, a sub-regional approach was adopted and a total of 13 sub-regional assessment reports were produced to a common protocol. The Task Force established a five-year plan to co-ordinate research, monitoring and modelling and other special topics in the preparations for the QSR. As part of this exercise a ‘Monitoring Master Plan’ was drawn up to provide for the first time reliable spatial information on the distribution of chemical contaminants and biological effects throughout the North Sea. The Task Force was a unique structure in international collaboration with a fixed remit that ended in December 1993. It was successful in bringing together many diverse organisations with interests in the North Sea and co-ordinated to a tight timetable the production of the QSR. The experiences gained are now being applied to the whole north east Atlantic under a new OSPAR Convention and have wide application to other Large Marine Ecosystems.

A change in the zooplankton of the central North Sea (55 degree to 58 degree N): A possible consequence of changes in the benthos

The mesozooplankton taken in continuous plankton recorder samples from the Central North Sea has changed from being numerically dominated by holoplanktonic calanoid copepod species from 1958 to the late 1970s to a situation where pluteus larvae of echinoid and ophiuroid echinoderms have been more abundant than any single holoplanktonic species in the 1980s and early 1990s. The abundance of the echinoderm larvae as a proportion of the zooplankton taken in the samples has followed a continuous increasing trend over the Dogger Bank, but off the eastern coast of northern England and southern Scotland the increase did not become obvious until the 1980s. This trend is consistent with reported increases in abundance of the macrobenthos. It is proposed that changes in the benthos have influenced the composition of the plankton.

Feeding of anchovy Engraulis encrasicolus larvae in the northwestern Adriatic Sea in response to changing hydrobiological conditions

Results from depth integrated and vertically stratified plankton sampling in the northwestern Adriatic Sea were used for comparison of gut contents of larvae of European anchovy Engraulis encrasicolus with composition and concentration of potential prey in the plankton. Sampling was carried out over a grid of stations both before and after a period of increased wind mixing to investigate changes in food availability and larval feeding success. All larvae had empty guts soon after dusk, indicating daytime feeding and rapid gut clearance. With increasing larval length there was a greater percentage of specimens with empty guts, despite suitable food being available in the plankton for these larger larvae; this suggests differential gut evacuation during sampling-possibly related to the degree of gut development. Larval diet was principally the various developmental stages of copepods, especially calanoid and cyclopoid nauplii, which were preferentially selected by larvae, whereas selection was against harpacticoid nauplii. Lamellibranch larvae and Peridinium were generally abundant in the plankton, but were only present in the gut contents in any number when the preferred dietary organisms were present in the plankton at low concentrations. The number of food organisms in the gut contents increased with concentration of the preferred food organisms in the plankton up to a limit of similar to 50 organisms/l. Within the upper 18 m of the water column, there was a reduction in the proportion of larvae with food in their guts with increasing depth, irrespective of the vertical profile of food concentration. Following a period of wind mixing the composition of the plankton changed. This was reflected in the diet of anchovy larvae, which altered in parallel. There was also an overall 41% decrease in concentration of the preferred food particles of larvae in the plankton following the period of wind mixing, but larvae were still able to maintain their food intake. These results show that anchovy larvae can successfully adapt their diet to a changing prey field and suggest that in the conditions observed in the northern Adriatic, quite radical changes in the feeding environment were probably insufficient to affect overall larval mortality.

Climate-related increases in jellyfish frequency suggest a more gelatinous future for the North Sea

Data obtained since 1958 from the continuous plankton recorder show an increasing occurrence of jellyfish in the central North Sea that is positively related to the North Atlantic Oscillation (NAO) and Atlantic inflow to the northern North Sea. Since 1970, jellyfish frequency has been also significantly negatively correlated with mean annual pH, independent of NAO trends. Jellyfish frequency increased in the mid-1980s, coincident with the reported regime shift in the North Sea and tracking trends in phytoplankton color. As models produced under all climate-change scenarios indicate a move toward a positive NAO, and pH of the oceans is predicted to decrease with rising CO2, we suggest that jellyfish frequency will increase over the next 100 yr.

Population-Dynamics And Production Of Euphausiids. III Meganyctiphanes-Norvegica And Nyctiphanes-Couchi In The North-Atlantic Ocean And The North-Sea

Seasonal changes in abundance, size and aspects of the population structure of Meganyctiphanes norvegica (M. Sars) and Nyctiphanes couchi (Bell) are described from samples taken with the “Continuous Plankton Recorder” at 10 m depth over a 2 yr period (1966 and 1967) in the North Atlantic Ocean and the North Sea. M. norvegica lived for a maximum of just over 2 yr, and adults of both year-classes spawned during a limited breeding season in the spring or summer. N. couchi spawned over a prolonged breeding season, giving rise to a complex of cohorts with overlapping size ranges. It was concluded that 3 or 4 cohorts were spawned in each year and that the maximum life span was probably greater than 1 yr, although maturity may be attained in less than a year. Estimated annual production at 10 m depth for M. norvegica ranged from 0.80 to 18.74 mg m-3yr-1 and for N. couchi from 0.67 to 8.23 mg m-3yr-1. P:B ratios ranged from 1.3:1 to 6.3:1 for M. norvegica and 4.0:1 to 5.5:1 for N. couchi.