Continuous plankton records: Geographical variations in numerical abundance,biomass and production of euphausiids in the North Atlantic Ocean and the North Sea

Geographical variations in the numbers, biomass and production of euphausiids and the contribution of common species to the total are described from samples taken during 1966 and 1967 in the North Atlantic Ocean and the North Sea by the Continuous Plankton Recorder at 10 m depth. Euphausiids were most abundant in the central and western North Atlantic Ocean and the Norwegian Sea. Thysanoessa longicaudata (Krøyer) was numerically dominant. Biomass was greatest in the Norwegian Sea and the north-eastern North Sea where Meganyctiphanes norvegica (M. Sars) accounted for 81 and 59%, respectively, of the total biomass. Production was highest off Nova Scotia and in Iberian coastal waters; the dominant species were T. raschi (M. Sars) in the former area and Nyctiphanes couchi (Bell) in the latter. The mean P:B ratios were correlated with temperature.     

Continuous plankton records: Changes in the composition and abundance of the phytoplankton of the north-eastern Atlantic ocean and North Sea, 1958–1974

In most areas of the north-eastern Atlantic Ocean, diatoms have declined drastically in abundance in the last decade. Additionally, in areas to the north of 59°N Ceratium species and an index of total phytoplankton have also declined. South of 59°N the phytoplankton index has increased, diatoms have declined and Ceratium species have remained at a constant level of abundance. A possible explanation of the increase in the phytoplankton index at a time when the diatoms were declinig south of 59°N is the development of unidentified phytoplankton organisms such as microflagellates. As many of the variables influencing phytoplankton standing crop are governed in turn by the prevailing weather, the phytoplankton changes may well be a consequence of the general deterioration, since 1940, of North Atlantic weather. Changes in phytoplankton which may be attributed to an amelioration of climate since 1971 are evident as yet only in the southern North Sea.     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

The role of ciliates,heterotrophic dinoflagellates and copepods in structuring spring plankton communities at Helgoland Roads,North Sea

Mesocosm experiments coupled with dilution grazing experiments were carried out during the phytoplankton spring bloom 2009. The interactions between phytoplankton, microzooplankton and copepods were investigated using natural plankton communities obtained from Helgoland Roads (54°11.3′N; 7°54.0′E), North Sea. In the absence of mesozooplankton grazers, the microzooplankton rapidly responded to different prey availabilities; this was most pronounced for ciliates such as strombidiids and strobilids. The occurrence of ciliates was strongly dependent on specific prey and abrupt losses in their relative importance with the disappearance of their prey were observed. Thecate and athecate dinoflagellates had a broader food spectrum and slower reaction times compared with ciliates. In general, high microzooplankton potential grazing impacts with an average consumption of 120% of the phytoplankton production (P _p ) were measured. Thus, the decline in phytoplankton biomass could be mainly attributed to an intense grazing by microzooplankton. Copepods were less important phytoplankton grazers consuming on average only 47% of P _p . Microzooplankton in turn contributed a substantial part to the copepods’ diets especially with decreasing quality of phytoplankton food due to nutrient limitation over the course of the bloom. Copepod grazing rates exceeded microzooplankton growth, suggesting their strong top-down control potential on microzooplankton in the field. Selective grazing by microzooplankton was an important factor for stabilising a bloom of less-preferred diatom species in our mesocosms with specific species (Thalassiosira spp., Rhizosolenia spp. and Chaetoceros spp.) dominating the bloom. This study demonstrates the importance of microzooplankton grazers for structuring and controlling phytoplankton spring blooms in temperate waters and the important role of copepods as top-down regulators of microzooplankton.     

Continuous Plankton Records: Seasonal cycles of phytoplankton and copepods in the North Atlantic ocean and the North Sea

Data from the Continuous Plankton Recorder survey of the North Atlantic Ocean and the North Sea are used to study geographical variations in the amplitude, duration and timing of the seasonal cycles of total phytoplankton and total copepods. It is shown that the distribution of overwintering stocks influences the distributions throughout the year. There is a relationship between the timing of the spring increase of phytoplankton and the amplitude of the seasonal variation in sea surface temperature. In the open ocean, the timing of the spring increase of phytoplankton corresponds with the spring warming of the surface waters. In the North Sea the spring increase occurs earlier, associated, perhaps, with transient periods of vertical stability, resulting in a relatively slower rate of increase. It is suggested that in the open ocean the higher rate of increase is under-exploited by copepods due to low overwintering stocks and longer generation times. Exceptionally early spring increases of phytoplankton off the west coast of Greenland and over the Norwegian shelf are probably associated with permanent haloclines. A high and late autumn peak of phytoplankton off the coast of Portugal may be associated with coastal upwelling.     

Continuous plankton records: overwintering and annual fluctuations in the abundance of zooplankton

A study of seasonal variations in the pattern of year-to-year changes, for 1948 to 1982, in the abundance of four zooplankton taxa (Pseudocalanus elongatus Acartia clausi, Calanus finmarchicus and Hyperiidea) for the North Sea suggests that an element of the variations, represented at least in part by a quasi-linear downward trend, has its origins in winter and its occurrence through the year is a function of inherent persistence. A downward linear trend is the dominant pattern in the annual fluctuations in abundance of many species of zooplankton throughout the north-east Atlantic and the North Sea. It is argued that it is reasonable to extrapolate from the limited data set included in the detailed study and to assume a winter origin for the pattern wherever it occurs. Some implications for ideas about the dynamics of the plankton ecosystem are considered.     

Distribution of nutrients and AOU in the Mediterranean Water (North Atlantic Ocean)

Water samples were collected in the Northeast Atlantic Ocean (33-37^° N and 11-16^° W) in May 1983, and vertical profiles of chemical and physical parameters allowed us to identify the different water masses. A significant difference in salinity and its maximum can be found at different places and depths, as a consequence of the spreading of the Mediterranean Water (MW) from the Strait of Gibraltar. The apparent oxygen utilization shows a correlation with nutrients in North Atlantic Central Water. The relative percentages of water masses were assessed in order to examine the depth of maximum influence of MW as a function of latitude and to apply this knowledge to the black scabbard fish long-line strategy. The long-line fishing depth location was adapted to the specific MW distribution of the different banks with improvements in the mean catch efficiency. The thickness of the MW prevalence layer has a pronounced decrease to the south. MW influence clearly sinks and decrease to south-west. The highest percentage of the MW was found in the Gorringe Bank area and the lowest in the Madeira area.     

Deep distributions of oceanic cirripede larvae in the Sargasso Sea and surrounding North Atlantic Ocean

From deep ( 1000 m), multi-depth zooplankton samples taken in the Azores frontal region from November 1980 to June 1981, high numbers of the two larval forms ofLepas pectinata, the commonest lepadid cirripede in this area, were sampled. The nauplii and cyprid larvae are large and long-lived. Nauplii were sampled where they feed, in the upper 150 m, closely associated with the fluorescence maximum. The cyprids were mainly sampled between 300 and 400 m, with a sharp cut-off in numbers just below the distribution peak. This deep distribution is intriguing, as the cyprids, which were not migrating diurnally, are non-feeding. To complete their life cycle they must settle on floating debris at the surface, where very few were sampled. They are negatively buoyant and there is no apparent physical reason for this deep distribution. There was no obvious pattern of changes in individual length or length/dry weight distribution with depth, the population appeared homogeneous. There was also no accumulation of other zooplankton at the same depth, or evidence from gut contents of large carnivores that the distribution pattern was caused by selective predation. A close association exists betweenL. pectinata andSargassum spp. weed, the cyprids preferentially settling on small fragments of weathered weed, the supply of which may be enhanced after winter. During winter, the harsh environment at the surface of the northern Sargasso Sea may discourage cyprid settlement. The deep distribution may be an ontogenetic migration conferring some survival advantage and saving energy due to the lower temperature and oxygen levels at depth. If it is a behavioural mechanism, duration of the cyprid stage will be extended until a more clement time of year, allowing them to settle over a longer period. The cyprids of other species were also sampled at depth, so the phenomenon appears to be widespread. If subsequent research confirms these observations as an ontogenetic migration, our current understanding of the fouling behaviour of oceanic lepadid cirripedes would be considerably modified.     

The occurrence of wax esters in crustaceans from the North-east Atlantic Ocean

The lipids of 2 species and 1 group of copepods, 4 species of euphausiids, 2 species of mysids, and 6 species of decapods, taken from different depths in the North-eastern Atlantic Ocean and Western Mediterranean Sea, have been analysed specifically for their wax ester content. The mid-water animals living below 500 m contained large amounts of wax ester (30 to 80% total lipid), while those animals living in the upper 600 to 700 m or on the sea floor had only small wax ester fractions (5 to 10% total lipid). It is suggested that the wax esters are laid down by those animals living in areas of low food density, where energy conservation is important, partly to achieve a more neutral buoyancy. In addition, it would appear that wax esters can function as energy reserves.     

Continuous plankton records: relationships between species of phytoplankton and zooplankton in the seasonal cycle

Seasonal fluctuations in abundance and their geographical variations in the north-east Atlantic Ocean and the North Sea for 49 taxa of both phytoplankton and zooplankton have been studied by multivariate techniques. Most of the observed variability in both seasonal and geographical fluctuations in abundance can be attributed to differences between the taxa with respect to population growth rates modulated by temperature coupled with the distribution of overwintering stocks, with the overall pattern also influenced by the timing of the establishment and breakdown of vertical stability in the water column. The observations indicate that for any species, the similarity between its seasonal cycles in the various areas, which represent a wide range of hydrographic regimes, is appreciably greater than any differences induced by interaction between the species in any particular area.     

Continuous plankton records: Persistence in time-series of annual means of abundance of zooplankton

Time-series of annual means of abundance of zooplankton of the north-east Atlantic Ocean and the North Sea, for the period 1948 to 1977, show considerable associations between successive years. The seasonal dynamics of the stocks appear to be consistent with at least a proportion of this being due to inherent persistence from year-to-year. Experiments with a simple model suggest that the observed properties of the time-series cannot be reproduced as a response to simple random forcing. The extent of trends and long wavelength variations can be simulated by introducing fairly extensive persistence into the perturbations, but this underestimates the extent of shorter wavelength variability in the observed time-series. The effect of persistence is to increase the proportion of trend and long wavelength variability in time-series of annual means, but stocks can respond to short wavelength perturbations provided these have a clearly defined frequency.     

Larval distribution and reproductive ecology of Thysanoessa macrura (Crustacea: Euphausiacea) in the Scotia Sea

In the Scotia Sea, the larvae of Thysanoessa macrura G.O. Sars, 1885 were scarce in spring (September 27 to October 6, 1971) and mainly present at the Calyptopis I stage, while in summer (December 17, 1974 to January 3, 1975) they were much more abundant and much further developed (up to Furcilia VI). The spawning season commences as early as September and, as indicated by the presence of numerous nauplii and metanauplii in summer, is rather long. T. macrura begins to spawn much earlier in West-Wind-Drift waters than in waters of the Weddell Sea. It starts to spawn simultaneously with Euphausia frigida and E. triacantha, but significantly earlier than E. superba. These timings of the breeding process are related to differential biogeographical characteristics of these species. Plankton collections made at depths of 500 or 1000 m have, for T. macrura, revealed a developmental ascent analogous to that of E. superba. Ontogenetical deep-water migrations are, consequently, characteristic not only for E. superba, but also for other Antarctic euphausiids (in particular E. frigida, but possibly not for inshore E. crystallorophias) and, as correctly postulated by several other investigators, for all oceanic euphausiid species.     

Seasonal distribution of modern coccolithophores in the western North Atlantic Ocean

The seasonal successions of standing crop and floral assemblage were studied for oceanic coccolithophores at 5 weather stations in the western North Atlantic Ocean. The coccolithophores were less populous during winter at the northern stations, whereas they became scarce during summer at the southern stations. The average standing crop in the surface water was quite consistent throughout all climatic regions, being a few tens of thousand cells per liter. The average standing crop at the 100 m level showed a clear trend of increasing southwards. The population at this depth was only one-tenth as large as that of the surface level at the northernmost station, whereas it was almost two-thirds as large as the surface levels at the southernmost station. Many of the major species exhibited seasonally biased occurrences. The trend was mostly repeated in consecutive years, although the magnitude of their abundances fluctuated from year to year in some case. The average species composition was obtained for the 5 weather stations at the surfacewater (surface and 20 m levels combined) and 100 m levels; Emiliania huxleyi dominated the flora at all stations. The relationship between occurrences of major species and ambient water temperature was also studied. Although some species were stenothermal, the majority exhibited eurythermal characteristics; this accounts for the high specific diversity of the coccolithophore community throughout the wide range of waters studied.Contribution No. 2899 of the Lamont-Doherty Geological Observatory of Columbia University.     

Latitudinal variation in aspects of the biology of Cyclothone braueri and C. microdon (Pisces: Gonostomatidae) in the eastern North Atlantic Ocean

Samples from a series of RMT-8 tows along 20°W in spring and autumn of various years were used to study the variability in aspects of the breeding biology of two species of Cyclothone: C. braueri and C. microdon in the eastern North Atlantic Ocean. Variables compared within each species included length-frequency distribution, maximum size, size at maturity, sex ratio, egg size and fecundity. Populations of both species at lower latitudes were generally smaller, reached maturity at a smaller size, and probably spawned earlier in the year and over a longer period than populations at higher latitudes. In both species, fecundity was highest at 40°N in the spring. The results suggested that C. microdon may not spawn at the northern extent of its range.     

Simulating transport of 129I and idealized tracers in the northern North Atlantic Ocean

Transport of the radioactive tracer Iodine-129 (¹²⁹I, T _1/2 = 15.7 Myr) in the northern North Atlantic Ocean has been investigated using a global isopycnic Ocean General Circulation Model (OGCM) and observed data. ¹²⁹I originates mainly from the nuclear fuel reprocessing plants in Sellafield (UK) and La Hague (France), and is transported northwards along the Norwegian coast, and then into surface and intermediate layers in the Arctic Ocean through the Barents Sea and the Fram Strait, but also partly recirculating south along the eastern coast of Greenland. In the North Atlantic Subpolar Seas, ¹²⁹I is mainly found in dense overflow waters from the Nordic Seas being exported southwards in the Deep Western Boundary Current, and to a lesser extent in surface and intermediate layers circulating cyclonically within the Subpolar Gyre. Observed concentration of ¹²⁹I along a surface transect in the eastern Nordic Seas in 2001 is captured by the OGCM, while in the Nansen Basin of the Arctic Ocean the OGCM overestimates the observed values by a factor of two. The vertical profile of ¹²⁹I in the Labrador Sea, repeatedly observed since 1997 to present, is fairly realistically reproduced by the OGCM. This indicates that the applied model system has potential for predicting the magnitude and depth of overflow waters from the Nordic Seas into the North Atlantic Subpolar Seas. To supplement the information obtained from the ¹²⁹I distribution, we have conducted a number of idealized tracer experiments with the OGCM, including tracers mimicking pure water masses, and instantaneous pulse releases. New insight into time-scales of tracer transport in this region is obtained by utilizing a few recently developed methods based on the theory of Transit Time Distribution (TTD) and age of tracers. Implications for other types of “anomalies” in the northern North Atlantic Ocean, being anomalous hydrography or chemical tracers, and how they are interpreted, are discussed.     

Marine snow studies in the Northeast Atlantic Ocean: distribution,composition and role as a food source for migrating plankton

During a 25 d Lagrangian study in May and June 1990 in the Northeast Atlantic Ocean, marine snow aggregates were collected using a novel water bottle, and the composition was determined microscopically. The aggregates contained a characteristic signature of a matrix of bacteria, cyanobacteria and autotrophic picoplankton with inter alia inclusions of the tintiniid Dictyocysta elegans and large pennate diatoms. The concentration of bacteria and cyanobacteria was much greater on the aggregates than when free-living by factors of 100 to 6000 and 3000 to 2 500 000, respectively, depending on depth. Various species of crustacean plankton and micronekton were collected, and the faecal pellets produced after capture were examined. These often contained the marine snow signature, indicating that these organisms had been consuming marine snow. In some cases, marine snow material appeared to dominate the diet. This implies a food-chain short cut wherby material, normally too small to be consumed by the mesozooplankton, and considered to constitute the diet of the microplankton can become part of the diet of organisms higher in the food-chain. The micronekton was dominated by the amphipod Themisto compressa, whose pellets also contained the marine snow signature. Shipboard incubation experiments with this species indicated that (1) it does consume marine snow, and (2) its gut-passage time is sufficiently long for material it has eaten in the upper water to be defecated at its day-time depth of several hundred meters. Plankton and micronekton were collected with nets to examine their vertical distribution and diel migration and to put into context the significance of the flux of material in the guts of migrants. Gut flux for the T. compressa population was calculated to be up to 2% of the flux measured simultaneously by drifting sediment traps and <5% when all migrants are considered. The in situ abundance and distribution of marine snow aggregates (>0.6 mm) was examined photographically. A sharp concentration peak was usually encountered in the depth range 40 to 80 m which was not associated with peaks of in situ fluorescence or attenuation but was just below or at the base of the upper mixed layer. The feeding behaviour of zooplankton and nekton may influence these concentration gradients to a considerable extent, and hence affect the flux due to passive settling of marine snow aggregates.     

Survey of the occurrence and morphology of sub-cuticular bacteria in shelf echinoderms from the north-east Atlantic Ocean

Echinoderms from the shelf seas around the British Isles were examined by epifluorescence and transmission electron microscopy for the presence of sub-cuticular bacteria (SCB). Information was obtained on SCB distribution in 63 of the 88 species known from this area, and 40 of these contained SCB. The SCB were present in all five classes. Only 39% of the holothurian species had SCB. In each of the other four classes, 60% of the species were associated with SCB. No correlation was noted between host ecology and SCB distribution or morphology. SCB distribution appeared to be related to host phylogeny. Congeneric echinoderms almost always had or lacked SCB, and this was usually also true for co-familial species. SCB showed limited morphological diversity, with all of them fitting into a classification of three general types. There was no morphological evidence suggesting that they are chemoautotrophs or methylotrophs. Some species had two morphological types of symbiont. SCB load was quantified for four species, three of which had symbiont densities >10⁹ SCB g^-1 ash-free dry wt. SCB are probably heterotrophic and may be co-evolving with their hosts.