The Continuous Plankton Recorder survey and the North Atlantic Oscillation: Interannual- to Multidecadal-scale patterns of phytoplankton variability in the North Atlantic Ocean

At interannual to multidecadal time scales, much of the oceanographic and climatic variability in the North Atlantic Ocean can be associated with the North Atlantic Oscillation (NAO). While evidence suggests that there is a relationship between the NAO and zooplankton dynamics in the North Atlantic Ocean, the phytoplankton response to NAO-induced changes in the environment is less clear. Time series of monthly mean phytoplankton colour values, as compiled by the Continuous Plankton Recorder (CPR) survey, are analysed to infer relationships between the NAO and phytoplankton dynamics throughout the North Atlantic Ocean. While a few areas display highly significant (p < 0.05) trends in the CPR colour time series during the period 1948–2000, nominally significant (p < 0.20) positive trends are widespread across the basin, particularly on the continental shelves and in a transition zone stretching across the Central North Atlantic. When long-term trends are removed from both the NAO index and CPR colour time series, the correlation between them ceases to be significant. Several hypotheses are proposed to explain the observed variability in the CPR colour and its relationship with climate in the North Atlantic.     

Macroecological study of Centropages typicus in the North Atlantic Ocean

Centropages typicus is a temperate neritic-coastal species of the North Atlantic Oceans, generally found between the latitudes of the Mediterranean and the Norwegian Sea. Therefore, the species experiences a large number of environments and adjusts its life cycle in response to changes in key abiotic parameters such as temperature. Using data from the Continuous Plankton Recorder (CPR) survey, we review the macroecology of C. typicus and factors that influence its spatial distribution, phenology and year-to-year to decadal variability. The ecological preferences are identified and quantified. Mechanisms that allow the species to occur in such different environments are discussed and hypotheses are proposed as to how the species adapts to its environment. We show that temperature and both quantity and quality of phytoplankton are important factors explaining the space and time variability of C. typicus. These results show that C. typicus will not respond only to temperature increase in the region but also to changes in phytoplankton abundance, structure and composition and timing of occurrence. Methods such as a decision tree can help to forecast expected changes in the distribution of this species with hydro-climatic forcing.     

An overview of statistical methods applied to CPR data

Since the beginning of the Continuous Plankton Recorder (CPR) survey in 1931, information on the abundance of a large number of plankton species or taxa has been obtained on a monthly basis in the northern North Atlantic. The many different ecological issues in which the survey has been involved have led to the application of a range of statistical methods. In this paper, we review some of the methods applied to the CPR data by presenting new and up-to-date analyses. Special attention is devoted to multivariate analysis, which has been used extensively to extract information from the CPR database. Results obtained from recently applied geostatistical methods on CPR data are then considered. An example of a time series decomposition by the use of Eigenvector filtering is presented to illustrate time-series analysis.     

Summarising spatial and temporal information in CPR data

The Continuous Plankton Recorder survey provides pan-oceanic data on geographic distribution, species composition, seasonal cycles of abundance, and long-term change during the last 70 years. In this paper we compare and contrast some of the historic data-analytic protocols of the survey, focusing primarily on the various means by which spatio-temporal information in CPR data has been exposed. Relative strengths and limitations are assessed, followed by suggestions for future approaches to the visualisation and summarising of CPR data.     

Oceanic distribution and life cycle of Calanus species in the Norwegian Sea and adjacent waters

The distribution and demography of Calanus finmarchicus, C. glacialis and C. hyperboreus were studied throughout their growth season on a basin scale in the Norwegian Sea using ordination techniques and generalized additive models. The distribution and demographic data were related to the seasonal development of the phytoplankton bloom and physical characteristics of water masses. The resulting quantified relationships were related to knowledge on life cycle and adaptations of Calanus species. C. finmarchicus was the numerically dominant Calanus species in Coastal, Atlantic and Arctic waters, showing strong association with both Atlantic and Arctic waters. C. hyperboreus and C. glacialis were associated with Arctic water; however, C. glacialis was occasionally observed in the Norwegian Sea and is probably an expatriate advected into the area from various origins. Demography indicated one generation per year of C. finmarchicus, a two-year life cycle of C. hyperboreus, and both one- and two-year life cycles for C. glacialis in the water masses where they were most abundant. For the examined Calanus species, young copepodites of the new generation seemed to be tuned to the phytoplankton bloom in their main water mass. The development of C. finmarchicus was delayed in Arctic water, and mis-match between feeding stages and the phytoplankton bloom may reduce survival and reproductive success of C. finmarchicus in Arctic water. Based on low abundances of C. hyperboreus CI–III in Atlantic water and main recruitment to CI prior to the phytoplankton bloom, we suggest that reproduction of C. hyperboreus in Atlantic water is not successful.     

Formation and export of deep water in the Labrador and Irminger Seas in a GCM

The influence of changes in the rate of deep water formation in the North Atlantic subpolar gyre on the variability of the transport in the Deep Western Boundary Current is investigated in a realistic hind cast simulation of the North Atlantic during the 1953–2003 period. In the simulation, deep water formation takes place in the Irminger Sea, in the interior of the Labrador Sea and in the Labrador Current. In the Irminger Sea, deep water is formed close to the boundary currents. It is rapidly exported out of the Irminger Sea via an intensified East Greenland Current, and out of the Labrador Sea via increased southeastward transports. The newly formed deep water, which is advected to Flemish Cap in approximately one year, is preceded by fast propagating topographic waves. Deep water formed in the Labrador Sea interior tends to accumulate and recirculate within the basin, with a residence time of a few years in the Labrador Sea. Hence, it is only slowly exported northeastward to the Irminger Sea and southeastward to the subtropical North Atlantic, reaching Flemish Cap in 1–5 years. As a result, the transport in the Deep Western Boundary Current is mostly correlated with convection in the Irminger Sea. Finally, the deep water produced in the Labrador Current is lighter and is rapidly exported out of the Labrador Basin, reaching Flemish Cap in a few months. As the production of deep-water along the western periphery of the Labrador Sea is maximum when convection in the interior is minimum, there is some compensation between the deep water formed along the boundary and in the interior of the basin, which reduces the variability of its net transport. These mechanisms which have been suggested from hydrographic and tracer observations, help one to understand the variability of the transport in the Deep Western Boundary Current at the exit of the subpolar gyre.     

Use of Continuous Plankton Recorder information in support of marine management: applications in fisheries, environmental protection, and in the study of ecosystem response to environmental change

The Continuous Plankton Recorder (CPR) survey was conceived from the outset as a programme of applied research designed to assist the fishing industry. Its survival and continuing vigour after 70 years is a testament to its utility, which has been achieved in spite of great changes in our understanding of the marine environment and in our concerns over how to manage it. The CPR has been superseded in several respects by other technologies, such as acoustics and remote sensing, but it continues to provide unrivalled seasonal and geographic information about a wide range of zooplankton and phytoplankton taxa. The value of this coverage increases with time and provides the basis for placing recent observations into the context of long-term, large-scale variability and thus suggesting what the causes are likely to be. Information from the CPR is used extensively in judging environmental impacts and producing quality status reports (QSR); it has shown the distributions of fish stocks, which had not previously been exploited; it has pointed to the extent of ungrazed phytoplankton production in the North Atlantic, which was a vital element in establishing the importance of carbon sequestration by phytoplankton.The CPR continues to be the principal source of large-scale, long-term information about the plankton ecosystem of the North Atlantic. It has recently provided extensive information about the biodiversity of the plankton and about the distribution of introduced species. It serves as a valuable example for the design of future monitoring of the marine environment and it has been essential to the design and implementation of most North Atlantic plankton research.     

Continuous Plankton Recorder sampling off the east coast of North America: history and status

The Continuous Plankton Recorder (CPR) survey has sampled plankton on 14 routes off the coasts of the northeast United States and Canada between 1959 and 2000. Six of these routes are still operating and are sampled on a monthly basis. Some 2047 CPR tows have been made to the end of 2000 and the resulting database represents the most extensive time series of marine plankton available anywhere in the northwest Atlantic. The location and time span of coverage of each route is presented. In addition selected information is presented on:

1. zooplankton abundance as departures from baselines for the northeast US continental shelf;

2. interannual variation in seasonality of Gulf of Maine phytoplankton;

3. zooplankton relationships to local hydrography of the Gulf of Maine and to the North Atlantic Oscillation;

4. time-space matrices of zooplankton abundance and anomalies southeast of New York City;

5. time series of phyto- and zooplankton on the Scotian Shelf;

6. seasonal cycles of Phytoplankton Colour and of zooplankton on the Scotian Shelf and Georges Bank, and in the Gulf of Maine; and

7. monthly abundance of zooplankton in Narragansett Bay, Rhode Island.

     

Distribution of Calanus finmarchicus in the northern North Atlantic and Arctic Ocean—Expatriation and potential colonization

The distribution of Calanus finmarchicus was studied on a transect across the central Greenland Sea, and on five transects from the Eurasian shelves across the Atlantic Inflow in the Arctic Ocean. Stage composition was used as an indicator for successful growth; gonad maturity and egg production were taken as indicators for reproductive activity. On the Arctic Ocean transects, these parameters were measured simultaneously from the sibling species Calanus glacialis. Response of egg production rate to different temperatures at optimal food conditions was very similar between both species in the laboratory. C. finmarchicus was present at all stations studied, but young developmental stages were only present close to the regions of submergence of Atlantic water under the Polar water. This together with a decreasing abundance and biomass from west to east along the Atlantic Inflow in the Arctic Ocean and reproductive failure indicates that C. finmarchicus is expatriated in the Arctic Ocean. We hypothesize that the late availability of food in the Arctic Ocean, rather than low temperature per se, limits reproductive success. Better reproductive success in the very low temperature regions of the Return Atlantic Current and the marginal ice zone in the Greenland Sea supports this hypothesis. The possibility for a replacement of C. glacialis by C. finmarchicus and consequences for the ecosystem after increasing warming of the Arctic are discussed.     

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea,English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C. typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas.     

The Continuous Plankton Recorder: concepts and history, from Plankton Indicator to undulating recorders

Alister Hardy conceived the Continuous Plankton Recorder (CPR) survey in the 1920s as a means of mapping near-surface plankton in space and time, interpreting the changing fortunes of the fisheries and relating plankton changes to hydrometeorology and climatic change. The seed he planted has grown to become the most extensive long-term survey of marine organisms in the world and the breadth of his vision becomes ever more apparent. The survey has now run for over 70 years and its value increases with every passing decade. Operating from ‘ships of opportunity’ the machines used are robust, reliable and easy to handle. Wherever possible, all the sampling and analytical methods have not been changed to maintain the consistency of the time series. Computerisation and the development of new statistical approaches have increased our ability to handle the large quantities of information generated and enhance the sensitivity of the data analyses. This overview, based on almost 900 papers, recounts the various phases in the history of the survey. It starts with the Indicator Survey (1921–1934), the deployment of the first CPR on the Discovery Expedition (1924–1927) and the early CPR survey in the North Sea (1931–1939). The survey reopened in 1946 after the Second World War and expanded across the North Atlantic to North America from 1959. Taxonomic studies were initiated and an emphasis was placed on patterns of distribution, which were seen to reflect the varying oceanographic conditions. The years 1968–1976 saw further expansion with operations even in the American Great Lakes, publication of a Plankton Atlas and initial evidence for a downward trend in plankton biomass. At about this time electronic instrumentation was attached to CPRs to make additional measurements and work was started on the development of a new generation of undulating Continuous Plankton and Environmental Recorders (CPERs). In 1976 the survey moved to Plymouth. Scientific priorities in the UK changed in the subsequent decade and funding became more difficult to secure even though some of the CPR papers being published at the time are now regarded as classics in plankton ecology. In 1988 the UK Natural Environment Research Council (NERC) decided to close the survey. An international rescue operation led to the creation of the Sir Alister Hardy Foundation for Ocean Science (SAHFOS) in 1990, which has continued with consortium funding from a number of countries, and from 1999 again included NERC. The scientific rationale of the survey has gained credibility as concern over climate change and other anthropogenic effects has grown and as the key role that plankton plays as an indicator of large-scale environmental conditions becomes ever more apparent. Recently, the survey became an integral component of the Global Ocean Observation System (GOOS) and expanded into the North Pacific. It plays a complementary role in many large international and multidisciplinary projects and is providing inspiration, advice and support to daughter surveys elsewhere in the world. At the start of a new millennium, Hardy’s vision from the 1920s is a powerful driving force not just in international biological oceanography, but in global environmental science.     

Distributions of Calanus spp. and other mesozooplankton in the Labrador Sea in relation to hydrography in spring and summer (1995-2000)

We collected mesozooplankton samples in the upper 100 m in spring or early summer each year between 1995 and 2000 along a section from Hamilton Bank (Labrador) to Cape Desolation (Greenland), and along additional sections in spring 1997 and early summer 1995. The North Atlantic waters of the central basin were characterised by the presence of the copepods Calanus finmarchicus, Euchaeta norvegica and Scolecithrocella minor and euphausiids. Calanus glacialis, Calanus hyperboreus and Pseudocalanus spp. were associated with the Arctic waters over the shelves. Amongst the other enumerated groups larvaceans were concentrated over the shelves and around the margins. Amphipods, pteropods and the copepods Oithona spp. and Oncaea spp. showed no definable relationships with water masses or bathymetry, while the diel migrant ostracods and chaetognaths were confined to deep water. Metrida longa, also a strong diel migrant, and Microcalanus spp., a mainly deep water species and possible diel migrant, were both sometimes quite abundant on the shelves as well as in the central basin, consistent with their likely Arctic origins.Analysis of community structure along the section across the Labrador Sea indicated that stations could be grouped into five different zones corresponding to: the Labrador Shelf; the Labrador Slope; the western and central Labrador Sea; the eastern Labrador Sea and Greenland Slope; and, the Greenland Shelf. The boundaries between zones varied spatially between years, but community composition was relatively consistent within a given zone and a given season (spring versus early summer). The relationship between community composition and water masses was not entirely straightforward. For example, Labrador Shelf water was generally confined to the shelf, but in spring 2000 when it also dominated the adjacent slope zone, the community in the Labrador Slope zone was similar to those found in other years. Conversely, in spring 1997, when Arctic organisms were unusually abundant in the Labrador Slope zone, there was no increased contribution of shelf water. In addition, North Atlantic organisms were often found on the shelves when no slope or central basin water was present.Although other organisms were sometimes very abundant, the mesozooplankton preserved dry weight biomass was dominated everywhere by the three species of Calanus, which together always accounted for ≥70%. One species, C. finmarchicus, comprised >60% of the total mesozooplankton biomass and >80% of the abundance of large copepods in spring and summer throughout the central Labrador Sea. In western and central regions of the central basin average C. finmarchicus biomass was ca 4 g dry weight m⁻² and average abundance, ca 17?000 m⁻² over both seasons. Highest levels (ca 7 g dry weight m⁻², >100?000 m⁻²) occurred in the northern Labrador Sea in spring and in eastern and southwest regions in early summer. C. hyperboreus contributed ca 20% of the total mesozooplankton biomass in the central basin in spring and <5% in early summer, while C. glacialis accounted for <1%. Over the shelves, C. hyperboreus contributed a maximum of 54% and 3.6 g dry weight m⁻², and C. glacialis, a maximum of 29% and 1 g dry weight m⁻², to the total mesozooplankton biomass.     

Trans-Atlantic responses of Calanus finmarchicus populations to basin-scale forcing associated with the North Atlantic Oscillation

Populations of the copepod species Calanus finmarchicus often dominate the springtime biomass and secondary production of shelf ecosystems throughout the North Atlantic Ocean. Recently, it has been hypothesised that interannual to interdecadal fluctuations observed in such populations are driven primarily by climate-associated changes in ocean circulation. Here, we compare evidence from the North Sea and Gulf of Maine/Western Scotian Shelf (GoM/WSS) linking fluctuations in C. finmarchicus abundance to changes in ocean circulation associated with the North Atlantic Oscillation (NAO). A particularly striking contrast emerges from this Trans-Atlantic comparison: whereas the North Sea C. finmarchicus population exhibits a negative correlation with the NAO index, the GoM/WSS population exhibits a more complex, positive association with the index. The physical processes underlying these contrasting population responses are discussed in the context of regional- to basin-scale circulation changes associated with the NAO.     

Macrophysiology of Calanus finmarchicus in the North Atlantic Ocean

Copepods represent the major part of the dry weight of the mesozooplankton in pelagic ecosystems and therefore have a central role in the secondary production of the North Atlantic Ocean. The calanoid copepod species Calanus finmarchicus is the main large copepod in subarctic waters of the North Atlantic, dominating the dry weight of the mesozooplankton in regions such as the northern North Sea and the Norwegian Sea. The objective of this work was to investigate the relationships between both the fundamental and realised niches of C. finmarchicus in order to better understand the future influence of global climate change on the abundance, the spatial distribution and the phenology of this key-structural species. Based on standardised Principal Component Analyses (PCAs), a macroecological approach was applied to determine factors affecting the spatial distribution of C. finmarchicus and to characterise its realised niche. Second, an ecophysiological model was used to calculate the Potential Egg Production Rate (PEPR) of C. finmarchicus and the centre of its fundamental niche. Relationships between the two niches were then investigated by correlation analysis. We found a close relationship between the fundamental and realised niches of C. finmarchicus at spatial, monthly and decadal scales. While the species is at the centre of its niche in the subarctic gyre, our joint macroecological and macrophysiological analyses show that it is at the edge of its niche in the North Sea, making the species in this region more vulnerable to temperature changes.     

Labrador Sea convection and subpolar North Atlantic Deep Water export in the SODA assimilation model

Labrador Sea convection was most intense and reached the greatest depths in the early 1990s, followed by weaker, shallower, and more variable convection after 1995. The Simple Ocean Data Assimilation (SODA) version 2.0.2/2.0.4 assimilation model is used to explore convective activity in the North Atlantic Ocean for the period from 1992 to 2007. Hydrographic conditions, which are relatively well observed during this period, are used to compare modeled and observed winter mixed-layer depths and water mass anomalies in relation to Deep Western Boundary Current transports and meridional overturning circulation (MOC) changes at the exit of the subpolar basin. The assimilation differs markedly from local observations in the March mixed-layer depth, which represents deep convection and water mass transformation. However, mean MOC rates at the exit of the subpolar gyre, forced by stratification in the mid-latitudes, are similar to estimates based on observations and show no significant decrease during the 1992–2007 period. SODA reproduces the deep Labrador Sea Water formation in the western North Atlantic without any clear indication of significant formation in the Irminger Sea while the lighter upper Labrador Sea Water density range is reached in the Irminger Sea in the 1990s, in agreement with existing assumptions of deep convection in the Irminger Sea and also supported by computed lag correlations with the Labrador Sea. Deep Water transformation mainly takes place in the eastern North Atlantic. The introduction of CFC-11 into the SODA model as a tracer reproduces the mean and multiyear variations of observed distributions.     

Seasonal development of Calanus finmarchicus in relation to phytoplankton bloom dynamics in the Norwegian Sea

Seasonal development of Calanus finmarchicus was studied in relation to the physical environment and phytoplankton bloom dynamics in the Norwegian Sea during eight basin-scale surveys from March to August 1995. Our main objective was to gain new knowledge about the life cycle of C. finmarchicus and its adaptation to the physical and biological environment of the Norwegian Sea. Time of spawning, estimated by temperature-dependent back-calculations from the occurrences of copepodite stage 1 (CIs), varied by water mass and occurred mainly during the phytoplankton pre-bloom and bloom periods. Recruitment to CI of the year's first generation (G1) generally occurred during the bloom and late bloom. The seasonal development of C. finmarchicus was progressively delayed from Coastal to Atlantic and to Arctic water, and from south to north within Atlantic and Arctic waters. This delay was partly linked to the phytoplankton bloom development that followed the same pattern, but development of C. finmarchicus also showed an increasing tendency to lag behind the phytoplankton development in colder waters. This may explain why C. finmarchicus are less successful in colder water. The consumption of nitrate was used as proxy for the seasonal history of phytoplankton development to aid interpretation of the lifecycle of C. finmarchicus. This approach allows us to align phytoplankton bloom and copepod development sequences despite temporal and geographical variation in bloom development, which otherwise tend to cause variability in quasi-synoptic and large-scale data. Two generations of C. finmarchicus were found in southern and northern regions of Coastal Water, and in southern Atlantic Water. In northern Atlantic Water and in Arctic Water, one generation was observed.     

Inter-decadal variability in zooplankton and phytoplankton abundance on the Newfoundland and Scotian shelves

Continuous Plankton Recorder (CPR) sampling on the Newfoundland and Scotian shelves covers three multi-year periods characterised by negative (1962–1971), positive (1992–2000) and negative/neutral (2001–2003) values of the NAO index. Water temperatures respond differently to changes in the NAO in different regions: a positive NAO index tends to lead to reduced temperatures on the Newfoundland shelf and to increased temperatures on the central/western Scotian shelf, and a negative NAO index to the reverse. Since the 1960s, the hydrographic changes due to the NAO have been superimposed on a freshening of the water column throughout the region, which is attributed to increased contribution of Arctic water outflow. Changes in plankton abundance measured by the CPR for the three time periods were generally, but not always, similar on the Newfoundland and eastern and western regions of the Scotian shelf, although Arctic species (e.g. Calanus glacialis, Calanus hyperboreus) were notably more abundant and warm water species (e.g. Metridia lucens, euphausiids) less abundant on the Newfoundland shelf than on the Scotian shelf. Three categories of phytoplankton (colour, diatoms, dinoflagellates) increased in abundance in the 1990s, and these increases generally persisted into 2001–2003. This is believed to be a response to the persistent freshening of the water column, probably due to increased stratification. The Arctic species C. glacialis and C. hyperboreus also showed persistent increases in abundance after 1992, perhaps due to increased transport from the Arctic, although the abundance of the Arctic slope water species Metridia longa decreased. Two groups, Calanus 1–4 and euphausiids, both thought to play important roles in the food chain, showed persistent decreases in abundance after 1992, especially on the Newfoundland shelf. In all regions, Calanus finmarchicus 5–6, Oithona spp. and Centropages hamatus abundance changed in association with variations in the NAO, although no common mechanism could be identified. C. finmarchicus 5–6 abundance decreased in the 1990s and increased after 2001, while the other two species showed the opposite pattern. Centropages typicus and M. lucens abundance on the Scotian shelf increased with rising temperature. This is attributed to increased production rates for the former and an increased influx of warm, M. lucens-rich, slope water on to the shelf for the latter. A comparison between ring net and CPR sampling on the Newfoundland shelf suggests that the Calanus 1–4 category is dominated by C. finmarchicus and that late stage C. glacialis and C. hyperboreus are grossly under-sampled compared to late stage C. finmarchicus.     

Regional variation in distribution pattern, population structure and growth rates of Meganyctiphanes norvegica and Thysanoessa longicaudata in the Irminger Sea, North Atlantic

Euphausiids are a key component of the northern North Atlantic marine ecosystem and Meganyctiphanes norvegica and Thysanoessa longicaudata are dominant both numerically and in terms of biomass. The Irminger Sea is remote and experiences often-hostile weather conditions. Consequently, few studies have been conducted there, and detailed information on the seasonal distribution, abundance and growth of euphausiids is limited. Here we explore patterns of abundance and spatial and temporal variation in length–frequency distribution in order to determine regional growth rates for both species in the Irminger Basin. Regional composite length–frequency distributions for spring, summer and winter were devised by aggregating discrete net haul data according to the results of a multivariate cluster analysis of length conducted on spring and summer net sample data. Three biologically distinct regions within the Irminger basin were apparent (Central Irminger Sea, Northern Irminger Sea and East of Greenland Shelf). These regions corresponded broadly with distinct physical zones within the basin. Modes in the composite length–frequency distributions were determined by fitting multiple normal distributions, and regional differences in growth were investigated by tracking modes between seasons. The results provide some evidence for regional variability in growth and population dynamics. The population structure and growth of M. norvegica was similar in the open ocean regions of the Northern and Central Irminger Basin, but different in the region around the East Greenland Shelf. There was a distinct absence of larger individuals (+I-group) in the open ocean regions compared to East Greenland Coast region, and growth rates were marginally higher. A similar pattern in population structure was also observed for T. longicaudata. Variability in growth and abundance are discussed in relation to prevailing environmental characteristics such as temperature and food availability.     

Zooplankton distribution across the Lomonosov Ridge,Arctic Ocean: species inventory,biomass and vertical structure

On a transect across the Lomonosov Ridge stratified zooplankton tows were made to the bottom at seven stations. A species inventory was established and compared with earlier observations in the Arctic Ocean. Differences between the Amundsen and Makarov basins are relatively small and correspond well with the general circulation patterns for Atlantic, Pacific, and neritic waters, suggesting slow mixing rates for the different basins. There were no remarkable differences in the species composition or their vertical distribution between the two sides of the Lomonosov Ridge. This indicates effective faunistic exchange across the ridge, although several bathy-pelagic species were almost or completely absent on top of the Ridge. Biomass showed a strong gradient along the transect, with a pronounced peak (9.5 g dry weight m⁻²) in the core of Atlantic water over the ridge, and minima over the deep basins. These differences were related to the effect of bottom topography for deep-living species, and the dynamics of the Atlantic layer for the meso- and epipelagic species. The maximum was formed mainly by the copepods Calanus hyperboreus and Metridia longa together with chaetognaths and ostracods. The presence of young developmental stages in some of the abundant species (C. hyperboreus, M. longa) suggests successful reproduction at all stations but C. finmarchicus was almost exclusively represented as old stages and adults. Comparison with earlier data on abundance and biomass from the Canada Basin (Russian Drift station “North Pole-22”) shows a pronounced difference with respect to both absolute quantities and relative composition. The copepod C. finmarchicus is completely absent in the central Canada Basin, and the portion of non-copepod zooplankton is dramatically decreased. This points to a reduced advection of Atlantic water or more severe food conditions in this basin.