An overview of plankton ecology in the North Sea

The purpose of this report is to give an overview of plankton ecology in the North Sea, and the processes that effect it, as derived from current research. The Sir Alister Hardy Foundation has extensive data for the North Sea area, and other sources have also been used to provide information for this report. Shortfalls in current research have also been highlighted. The information contained herein is to be contributed towards an information base for the Strategic Environmental Assessment. The North Sea is an extension of the North Atlantic that has an area of 574,980 km2. The deepest area is off the coast of Norway (660m), with a number of shallow areas, such as the Dogger Bank (15m). The North Sea represents a large source of hydrocarbons that have been exploited since the early 1970s. The aim of this study is to provide the Department of Trade and Industry with biological data on the planktonic community of the North Sea, as a contribution towards the Strategic Environmental Assessment (SEA 2). An overview of phyto- and zoo- plankton community composition, plankton blooms, Calanus, mero-, pico- and megaplankton, sensitivity to disturbance / contamination, phytodetritus and vertical fluxes and the resting stages of phytoplankton is made using the results of the survey database. Additional published literature has also been used, and gaps in available data have been highlighted. 1.3 The Continuous Plankton Recorder (CPR) survey provides a unique long-term dataset of plankton abundance in the North Atlantic and North Sea (Warner and Hays 1994). The survey has been running for almost 70 years, using ‘ships of opportunity’ to tow CPRs on regular, and incidental routes, sampling at a depth of 10 m. Each sample represents 18 km of tow and approximately 3 m3 of filtered seawater. Over 400 taxa of plankton are routinely identified by a team of taxonomists. The samples are also compared to colour charts to give an indication of ‘greenness’, which provides a visual index of chlorophyll value. CPRs have been towed for over 4 million nautical miles, accumulating almost 200,000 samples. The design of the CPR has remained virtually unchanged since sampling started, thus providing a consistency of sampling that provides good historical comparisons. By systematically monitoring the plankton over a period, changes in abundance and long term trends can be distinguished. From this baseline data, inferences can be made, particularly concerning climate change and potentialanthropogenic impacts.

Variability In Abundance Vertical-Distribution And Ontogenetic Migrations Of Thysanoessa-Longicaudata (Crustacea Euphausiacea) In The Northeastern Atlantic-Ocean

The vertical distribution, seasonal and ontogenetic migrations and seasonal variability in abundance of Thysanoessa longicaudata (Krøyer) were investigated using the Longhurst-Hardy Plankton Recorder for a 4 yr period (March, 1971 to May, 1975) at Ocean Weather Station “I” (59°00′N; 19°00′W) in the north-eastern Atlantic Ocean. Of 8 species of euphausiids identified at this position, the vast majority were T. longicaudata (for example, 99.5% of the total euphausiids in 1972 belonged to this species). From March to October the majority of calyptopes, furciliae and adults of T. longicaudata were found in the upper 100 m. The major spawning occurred in spring at a water temperature of 9° to 10°C and calyptopes and furciliae appeared in late April, reaching their maximum abundance in May. There was no evidence of large-scale diurnal migrations, although an extensive ontogenetic migration of young developmental stages was observed. The eggs were found from 100 m down to 800 m, the maximum depth of sampling, and the vertical distribution of the three naupliar stages showed a “developmental ascent” as they matured. During the main reproductive period in May, over 70% of all nauplii were below 500 m while more than 94% of Calyptopis Stage I were above 500 m with their maximum abundance in the euphotic zone (0 to 50 m). Calyptopis Stage I is the first feeding stage and it is this stage which shows the largest ontogenetic migration. Brief descriptions of the egg and nauplii are given.

Plankton Of The Fladen Ground During Flex-76 .3. Vertical-Distribution Population-Dynamics And Production Of Calanus-Finmarchicus (Crustacea Copepoda)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR), the Longhurst Hardy Plankton Recorder (LHPR) and by our colleagues from other participating Institutes during the Fladen Ground Experiment (FLEX 76) were used to describe the vertical distribution and population dynamics of Calanus finmarchicus (Gunnerus) and to provide estimates of the production and carbon budget of the population from 19 March to 3 June, 1976. Total production of the 19 March to 3 June, 1976. Total production of the nauplii and copepodite stages (including adults), during the exponential growth phase in May, was estimated to be in the range of 0.49 to 0.91 g C m-2 d-1 or 29.0 to 55 g dry wt m-2 (14.5 to 27.8 g C m-2) for the three successive 10 d periods in May. Two gross growth efficiencies (K 1) (20 and 34%), together with the lower value of C. finmarchicus production, were used to calculate the gross ingestion levels of algae as 2.45 and 1.44 g C m-2 d-1 (73.5 and 43.2 g C m-2 over the May period). These ingestion levels, together with the algae ingested by other zooplankton species, are greater than the estimated total phytoplankton production of 45.9 g C m-2 over the FLEX period. A number of factors are discussed which could explain the discrepancies between the production estimates. One suggestion is that the vertical distribution of the development stages of this herbivorous copepod and their diel and ontogenetic migration patterns enable it to efficiently exploit its food source. Data from the FLEX experiment indicated that the depletion of nutrients limited the size of the spring bloom, but that it was the grazing pressure exerted by C. finmarchicus which was responsible for the control and depletion of the phytoplankton in the spring of 1976 in the northern North Sea.

Plankton Of The Fladen Ground During Flex-76 .2. Population-Dynamics And Production Of Thysanoessa-Inermis (Crustacea Euphausiacea)

Samples taken in the northern North Sea with the Continuous Plankton Recorder (CPR), the Undulating Oceanographic Recorder (UOR) and the Longhurst-Hardy Plankton Recorder (LHPR) during the Fladen Ground Experiment in 1976 (FLEX 76) are used to describe the vertical distribution and population dynamics of Thysanoessa inermis (Krøyer) and to provide estimates of the production and carbon budget of the population from 19th March to 3 June 1976. Spawning occurred in late April and early May, in near synchronisation with the start of the spring bloom of phytoplankton. Eggs, nauplii and calyptopes reached maximum abundance in succession, and furciliae were numerous when sampling ceased in early June. Adults increased in length from a mean of 12.1 mm in mid-March to 17.5 mm in early June and the estimated production was 2.40 mg m-3 over the 74 d period. Total carbon ingested by the population of T. inermis was estimated to be 10 mg C m-2 d-1 in the upper 100m which was only 1.5% of the daily primary production of 0.68 gC m-2 measured over the FLEX period 26 March to 4 June 1976. The grazing by T. inermis on the phytoplankton population was assumed to have little effect on the control and depletion of the spring phytoplankton bloom during FLEX 77.

Diel vertical migration of decapod larvae in the Portuguese coastal upwelling ecosystem: implications for offshore transport

The vertical distribution of decapod larvae off the northwest Portuguese coast was analysed in relation to associated environmental conditions from sampling during a 69 h period around a current meter mooring located on the shelf, approximately 21 km off the coast. Plankton samples were collected every 2 h at the surface with a neuston net and through the water column with a Longhurst Hardy Plankton Recorder (Pro-LHPR), allowing a very detailed resolution of larval vertical distribution. Environmental data (temperature, salinity, and chlorophyll a) were obtained every hour. To investigate the horizontal distribution of decapod larvae in relation to the coast, a plankton-sampling grid was carried out before the 69 h fixed station. Larvae of shelf decapod species were widely distributed over the shelf, while those of inshore species were found much closer to the coast. Decapod larvae (zoeae and megalopae) showed clear diel vertical migrations, only appearing in the upper 20 m at night, a migration that did not appear to be affected by physical conditions in the water column. Larval densities were highly variable, 0.01 to 215 ind. m super(-3) for zoeae and 0 to 93 ind. m super(-3) for megalopae, the zoeae being generally more abundant. The results indicated that during the day larvae accumulate very close to the bottom. The diel vertical migration behaviour is discussed as one of the contributing mechanisms for larval retention over the shelf, even with offshore transport conditions promoted by coastal upwelling, and is hence of major relevance for the recruitment success of decapod species that inhabit inshore and shelf zones of coastal upwelling systems.

US NOAA Fisheries and UK SAHFOS CPR surveys: comparison of methods and data

The US National Oceanic and Atmospheric Administration (NOAA) Fisheries Continuous Plankton Recorder (CPR) Survey has sampled four routes: Boston–Nova Scotia (1961–present), New York toward Bermuda (1976–present), Narragansett Bay–Mount Hope Bay–Rhode Island Sound (1998–present) and eastward of Chesapeake Bay (1974–1980). NOAA involvement began in 1974 when it assumed responsibility for the existing Boston–Nova Scotia route from what is now the UK's Sir Alister Hardy Foundation for Ocean Science (SAHFOS). Training, equipment and computer software were provided by SAHFOS to ensure continuity for this and standard protocols for any new routes. Data for the first 14 years of this route were provided to NOAA by SAHFOS. Comparison of collection methods; sample processing; and sample identification, staging and counting techniques revealed near-consistency between NOAA and SAHFOS. One departure involved phytoplankton counting standards. This has since been addressed and the data corrected. Within- and between-survey taxonomic and life-stage names and their consistency through time were, and continue to be, an issue. For this, a cross-reference table has been generated that contains the SAHFOS taxonomic code, NOAA taxonomic code, NOAA life-stage code, National Oceanographic Data Center (NODC) taxonomic code, Integrated Taxonomic Information System (ITIS) serial number and authority and consistent use/route. This table is available for review/use by other CPR surveys. Details of the NOAA and SAHFOS comparison and analytical techniques unique to NOAA are presented.

Going Global: planning the next 80 years of the Continuous Plankton Recorder Survey

Going Global: planning the next 80 years of the Continuous Plankton Recorder Survey. Operated by the Sir Alister Hardy Foundation for Ocean Science (SAHFOS), the Continuous Plankton Recorder (CPR) survey is the world’s largest, sampling 4 ocean basins, and longest running (since 1931) plankton biodiversity monitoring programme. Having sampled enough miles to circumnavigate the globe over 200 times, the CPR database houses over 2.5 million entries, describing the distribution of 500 phytoplankton and zooplankton taxa. Routinely sampling in the Arctic, Atlantic, Pacific and Southern Oceans, the survey analyses 4000 samples yearly. Data collected from these samples are made freely available for bona fide scientific purposes. The CPR survey data is used to generate a better understanding of changes in the plankton and to date some 1000 papers have been published on plankton biodiversity. This year sees the 80th anniversary of the CPR survey and to celebrate and build upon this unique monitoring programme, SAHFOS intends to further develop its global plankton perspective. Work will be extended into the South Atlantic and Indian Ocean and an international partnership with complementary surveys in Australia, Canada, America, Japan and South Africa will be implemented. The Digital Object will describe the CPR survey using compilations made by Plymouth Art College and BBC film footage.

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 Copepoda of the Southern North Sea. 1932-37.

I. The monthly changes in the distribution and abundance of the Copepoda in the southern North Sea have been investigated from June 1932 to December 1937 by using the Continuous Plankton Recorder. This was towed at a standard depth of 10 metres by ships sailing on regular lines from Hull to Rotterdam, to Bremen and towards the Skagerrak, and later from London to Esbjerg. 2. The methods are described and those limitations which apply more particularly to the Copepoda are discussed (pp. 175 to 186 and 198 to 203). 3. The first part of the report deals with the Copepoda as a whole, i.e. the total population. The difference between the summer and winter distributions is stressed. The variations in numbers from year to year are found to be considerable and it is suggested that they are sufficiently large to be reflected in the success or failure of the broods of those fish which are at some period of their development dependent upon the Copepoda for food. 4. The second part deals with the data concerning the constituent species or groups of allied species ; a list of these is given on p. 197. 5. The group Paracalanus + Pseudocalanus was by far the most abundant and together with the genera Temora and Acartia was found to be responsible for most of the fluctuations in the population (pp. 205 to 208). 6. The distributions, seasonal and spatial, of the other common forms are described, with the exception of that of Oalantts finmarchicus which is to be the subject of a later report. 7. The recorder results are compared with the findings of the International Council survey from 1902 to 1908; some marked disagreements are discussed (pp. 227 to 232). 8. The appearance of the northern forms Oandacia armata and Metridia lucens during the winters of 1932-33, 1933-34 and 1937 are recorded (pp. 222 to 223) 9. A summarised account of the main seasonal changes in the area is given (pp. 232 to 234) and followed by a brief comparison of the 5½ years investigated.

Ecological Relations Between the Herring and the Plankton off The North-East Coast of England

I. 430 plankton samples, which were taken by several herring drifters using the Continuous Plankton Recorder in the Shields fishing area during the summer seasons of 1931 to 1933, are analysed to show the main changes in the plankton during those seasons. 2. A comparison is made between the proportions of the different zooplankton organisms found in the plankton and the proportions of these recorded by Savage (1937) in the stomachs of herring obtained from drifters working in the same area and during the same time. The comparisons are made for 29 ten-day periods in the seasons 1931 to 1933, and in addition, for 6 ten-day periods relating to a single drifter which obtained both plankton and stomach samples at the same time in 1932. 3. The comparisons in 2 provide evidence that the herring feeds by selecting certain organisms by individual acts of capture and not by swimming open-mouthed to strain out the plankton indiscriminately: (a) Calanus and Temora in the stomachs either correspond fairly closely to the proportions in the plankton or they may be in very much higher proportions. The latter is always true regarding Anomalocera. (b) Acartia, Oithona, Cladocera and Lamellibranch larvae are always in larger proportions in the plankton than in the stomachs; this applies also to Centropages with two insignificant exceptions. (c) There is a close correspondence between the numbers of Limacina and Sagitta in the plankton and stomachs in the latter half of the 1931 season, but not during 1932 and 1933, when the numbers in the stomachs were insignificant ; during the former period there was a great scarcity of Calanus in the plankton.