Analysis of the parasitic copepod species richness among Mediterranean fish

The Mediterranean ichthyofauna is composed of 652 species belonging to 405 genera and 117 families. Among these, 182 were studied for their parasitic copepods. The analysis of all the works conducted on these crustacea yielded 226 species distributed in 88 genera and 20 families. For each fish species we have established a file providing the species name of the fish, its family, its geographical distribution within the Mediterranean and some of its bio-ecological characteristics. Within each file, all the parasitic copepod species reported on each host species were listed. This allowed to know the species richness (SR) of these hosts. We thus produced 182 files within which 226 copepod species are distributed. A program was created under the Hypercard software, in order to analyse our data. Two parameters were studied. The first one is the mean species richness (MSR), which corresponds to the mean of the different SR found on the different host species. The second is the parasite–host ratio (P/H), which is the ratio of the number of copepod species by the number of host species. These parameters are calculated by our program for all the 182 species of Mediterranean fishes retained in our investigation, on the first hand, and, on the second hand, for one particular group of fish species. We used the following variables to investigate their correlations with copepod species richness: taxonomy—fish families, genera and species; biometry—maximal size of the adult fish; eco-ethology—mode of life (benthic, pelagic or nectonic), displacements (sedentary, migratory with environmental change, or migratory without environmental change), behaviour (solitary or gregarious). Other variables (colour, food, reproduction, abundance, distribution area) were also analysed but did not reveal any clear correlation. Providing that our study does not rely on quantitative (prevalence, intensity) but qualitative basis our aim was only to reveal some tendencies. These tendencies are as follows: (1) In many cases, parasite and host phylogeny seem to play an important role. There are fish families with copepods and families with few species of these parasites. The phyletic constraints could be due to the morphological characteristics of the habitat (e.g. structure of the gills) or biological/ecological characteristics that we were unable to identify. (2) It appears that the presence in a same environment of related fish species (e.g. several species of the same genus, or numerous genera of the same family) is correlated with high parasite richness. A likely explanation is that such situations favours alternated processes of lateral transfers and speciation. (3) Some eco-ethological criteria seem to favour the establishment of a large parasite species richness. It should be noted for instance that Mediterranean fishes the most often infected with copepods are generally nectonic or pelagic, migratory, and gregarious species.     

Evidence of episodic changes in deep-sea mesozooplankton abundance and composition in the Levantine Sea (Eastern Mediterranean)

Mesozooplankton samples from the 4270-m deep Ierapetra Basin in the oligotrophic Levantine Sea have revealed a strong temporal variability in abundance and composition hitherto unknown for the deep sea pelagic zone. The phenomenon reflected by a survey in June 1993 as compared to January 1987 is assessed by (i) the increase of the mesozooplankton standing crop throughout the water column, (ii) the absence of a decline in mesozooplankton abundance with increasing depth below 1000 m, (iii) the outstanding dominance of two interzonal calanoid copepods at bathypelagic and abyssopelagic depths and (iv) a substantial faunal change due to the codominance of Calanus helgolandicus, which was found for the first time in the area. The underlying hydrographical factors are addressed in the light of the Eastern Mediterranean Transient (EMT). Apart from a possible long-term faunal change, the EMT may exaggerate episodic plankton blooms and surface-abyssopelagic coupling in space and time.     

Life cycle of Epischura baicalensis Sars (Copepoda, Calanoida) in Lake Baikal

Epischura baicalensis Sars is a dominant pelagic species of Lake Baikal zooplankton. This is endemic to Lake Baikal and inhabits the entire water column. It produces two generations per year: the winter–spring and the summer. These copepods develop under different ecological conditions and vary in the duration of life stages, reproduction time, maturation of sex products and adult males and females lifespan. The total life period of the animals from each generation is one year. One female can produce 10 egg sacks every 10–20 days during its life time. The ratio of males and females is 1:1. One of the most essential features of the ecology of E. baicalensis is the alteration of its mass inhabited areas during a year, as well as in day time. This is due to the need for various conditions for gonad maturation, reproduction, nourishing and protection from being consumed by planktivors.     

Annual cycle of distribution of three ice-associated copepods along the coast near Dumont d'Urville, Terre Adélie (Antarctica)

In polar regions sea ice is a site of enhanced primary production during winter and provides important habitat for small grazers, such as copepods. We sampled zooplankton from the sea ice and water column throughout 2005, near Dumont d'Urville station (Terre Adélie, Antarctica). Three species of ice-associated copepods were found: two calanoid copepods Paralabidocera antarctica and Stephos longipes and the harpacticoid copepod Drescheriella glacialis. P. antarctica was the most abundant of the three and was closely associated with the sea ice during most of the year. This species had a one year life cycle with a probable over-wintering period in the sea ice as nauplii and a short copepodite phase in spring. Reproduction and spawning occurred in early summer. A comparison with two other populations (near Syowa and Davis stations) along the east coast of Antarctica showed that there was a temporal shift in the life cycles of the three populations, which was linked to variability in sea ice conditions. D. glacialis was the second most abundant copepod and was more common during the winter than during summer, indicating its preference for the sea ice habitat. In autumn, the presence of D. glacialis in the deeper part of the water column suggested that this species colonised the sea ice from the benthos. S. longipes was found only in the water column near Dumont d'Urville and was not very abundant. In Terre Adélie particular environmental conditions, such as the absence of a permanent sea ice zone throughout the year, a longer time of open water, strong katabatic winds and the presence of polynyas, have influenced both the abundance and distribution of the three common ice-associated copepods.     

Trophic flows,ecosystem structure and fishing impacts in the South Catalan Sea,Northwestern Mediterranean

An exploited ecosystem from the continental shelf and upper slope of the Northwestern Mediterranean Sea was described by means of an Ecopath mass-balance model with the aim of characterising its functioning and structure and describing the ecosystem impacts of fishing. This application included some complexities added to the general modelling methodology due to the high biodiversity of the Mediterranean Sea and the multispecific nature of the fishery, and to the difficulties of working with fishing data which are usually irregularly or imprecisely collected. The model comprised 40 functional groups including primary producers, the main species of benthic, demersal and pelagic invertebrates, fishes and non-fish vertebrates and three detritus groups. In addition, trawling, purse seine, longline and troll bait fishing fleets were included.Results showed that the functional groups were organized into four trophic levels with the highest levels corresponding to anglerfish, dolphins, large pelagic fishes and adult hake. The system was dominated by the pelagic fraction, where sardine and anchovy prevailed in terms of fish biomasses and catches. Detritus and detritivorous groups also played key roles in the ecosystem and important coupled pelagic-demersal interactions were described. Considering Odum's theory of ecosystem development, the ecosystem was placed on an intermediate-low developmental stage due, at least partially, to the impact of fishing activity. This highlighted the high intensity of fishing in the ecosystem, in accordance with the general assessment of western Mediterranean marine resources, and fishing fleets were ranked as top predators of the system. The low trophic level of the catch was in line with the long history of exploitation in the area. However, the steady decline of pelagic landings between 1994 and 2003, coupled with a decrease of the pelagic biomass within the system, underlined the low resistance of the system in front of perturbations. This decline was reproduced under Ecosim dynamic simulations combining different scenarios of moderate increase of fishing effort and an environmental forcing affecting the availability of preys to small and medium-sized pelagic fishes under wasp-waist flow control.     

Trace element levels in some commercially valuable fish species from coastal waters of Mediterranean Sea, Egypt

Concentrations of 5 trace elements (Cd, Pb, Cu, Cr, Zn) were determined in muscle of fishes collected from two coastal areas of the Egyptian coast of the Mediterranean Sea west of Alexandria (El-Mex Bay and Eastern Harbour). Sardinella aurita, Alepes djedaba, Siganus luridus, Siganus rivulatus, Sphyraena chrysotoenia, and Scomberomorus commerson were collected from El-Mex Bay. While Boops boops, Lithognathus mormyrus, Sparisoma cretense, Serranus cabrilla and Synodus saurus were collected from the E.H. In all fishes zinc was highest (up to 57 mg/kg) followed by Cr, Cu, Pb and Cd. The concentrations of several metals in fishes were significantly different among the species in each area. For all trace element examined, the concentrations decreased significantly with body weight of some fishes. In contrast, a positive correlation with body weight was found especially for Zn and Cu concentrations in 5 fish species and for Pb in 4 fish species. The concentrations of Cd, Pb and Zn were higher in fishes from El-Mex Bay (3.76, 6.49 and 57.21 mg/kg) than those from Eastern Harbour, whereas the opposite trend was observed for Cu and Cr, revealing the direct sources of trace metal pollution present in El-Mex coastal area. Levels of Cr surpassed the Maximum Permissible Concentration in most fish tissues, followed by Pb and Cd in some species. Cu and Zn presented concentrations below the maximum permissible levels in fish tissues.     

Fluxes of biogenic carbon in the Southern Ocean: roles of large microphagous zooplankton

The Southern Ocean is an extreme environment, where waters are permanently cold, a seasonal ice cover extends over large areas, and the solar energy available for photosynthesis is severely restricted, either by vertical mixing to considerable depths or, especially south of the Antarctic Circle, by prolonged seasonal periods of low or no irradiance. Such conditions would normally lead to low productivity and a water column dominated by recycling processes involving microbial components of pelagic communities but this does not seem to be the case in the Southern Ocean, where there is efficient export to large apex predators and deep waters. This paper investigates the role of large microphagous zooplankton (salps, krill, and some large copepods) in the partitioning of biogenic carbon among the pools of short- and long-lived organic carbon and sequestered biogenic carbon. Large microphagous zooplankton are able to ingest microbial-sized particles and thus repackage small, non-sinking particles into both metazoan biomass and large, rapidly sinking faeces. Given the wide spatio-temporal extent of microbial trophic pathways in the Southern Ocean, large zooplankton that are omnivorous or able to ingest small food particles have a competitive advantage over herbivorous zooplankton. Krill efficiently transfer carbon to a wide array of apex predators and their faecal pellets are exported to depth during occasional brief sedimentation episodes in spring time. Salps may be a significant link towards some fish (directly) and other apex predators (indirectly) and, at some locations (especially in offshore waters) and time, they may account for most of the downward flux of biogenic carbon. Large copepods are a trophic link towards fish and at least one whale species, and their grazing activity generally impedes the export of organic particles to depth. As a result, biogenic carbon is channelled mainly towards apex predators and episodically into the deep ocean. Without these original interactions, Antarctic waters might well be dominated by microbial components and recycling processes instead of active export from the generally small primary producers towards large apex predators.     

Colonization of a new man-made river (Marchfeldcanal, Lower Austria) by benthic copepods

The role of copepods in the colonization and emigration processes of benthos and drift of an irrigation canal were studied. During the 3 first years of existence of the canal, fifteen species of copepods were recorded (1 diaptomid, 11 cyclopoids, 3 harpacticoids). Copepod succession in the canal was initially represented by lentic species, then by predatory species like Macrocyclops albidus that developed with the proliferation of filamentous algae and epibionts during the first two years. Primary consumers appeared from the beginning of the flowing conditions but started to dominate only from the third year. Smaller species associated with the bed sediments like the harpacticoid species Nitocra hibernica, were late colonizers. The latter as well as Paracyclops fimbriatus and Eucyclops serrulatus were the dominant copepod species in the canal. Total copepod density in the benthos ranged from 15,800 to 68,000 ind./m², with a mean of 7.6% of the total zoobenthos. Copepods constituted up to 25% of the total faunal drift density, ranging from 50 to 140 ind./m³. Their highest abundance was observed around midnight. A three-fold increase of the discharge temporarily affected the drift density, but did not change the diel rhythm of the drift. Cyclopoid copepods namely P. fimbriatus (54.77%), copepodid cyclopoids (20.76%) and E. serrulatus (16.11%) formed the main part of the drift samples. The harpacticoid Nitocra hibernica was a minor participant in the drift processes.     

On the relation of structure, perception and activity in marine planktonic copepods

The goal of this paper is to illustrate how in juvenile and adult subtropical marine planktonic copepods various structures or morphological features function in concert to detect prey and predators. Without motion by either food (e.g. flagellate, ciliate) or feeder (e.g. feeding current) or both (e.g. Acartia spp. and ciliate) few feeding activities will occur. Through motion a food particle is either perceived mechanically or chemically to be followed by appendage activities. A combination of mechano- and chemosensors on their cephalic appendages (and probably on other extremities) serve juvenile and adult copepods to perceive signals. Perception is followed by alternation of motion and sensing by these appendages, or by no motion at all (e.g. behavior by Eucalanus pileatus when perceiving a weak hydrodynamic signal). Non-moving and extended sensors (setae) are best suited for mechanical/hydrodynamic perceptions in those copepods which lack a feeding current and hardly move. Numerous mechanosensors arranged in three dimensions on the first antennae (A1) are required to perceive the precise location of moving prey at a distance (e.g. Oithona feeding on ciliates but also sinking particles). Those copepods which create a weak or intermittent feeding current can supplement nutrition with carnivory, which requires perception by the A1 (e.g. Centropages velificatus adults). These two groups require, in addition to perception of prey motion/location, rapid motion by their appendages (A1, second maxillae M2, etc.) to capture the prey. Nauplii, which satiate at far lower food levels than adults, have one of several means of food acquisition: encounter through forward motion, perception through feeding current, or perception of a moving food particle. The nearly continuous motion of most calanoid nauplii makes them vulnerable to predation because all three pairs of appendages are usually moving. Opposite are nauplii of cyclopoid and a few calanoid species which move only occasionally. Copepodid stages and adults use non-moving and often extended setae on the tips of their A1 to perceive predators at a distance. This structure and their pronounced escape motion may reduce their vulnerability to predation as compared to nauplii.     

The role of pelagic fish as forage for the demersal fish community in the southern Bay of Biscay

The small pelagic fish represents the link between nekton and demersal communities, and they are an important food source for some demersal fish species. In the autumns between 1993 and 2002, 74,869 stomach contents of 25 demersal fish species were analysed during the scientific surveys organized by the Instituto Español de Oceanografía. These species represent the demersal fish community of the southern Bay of Biscay. An important part of their diet (39% by volume) was composed of pelagic fish species (e.g. Engraulis encrasicolus, Gadiculus argenteus, Micromesistius poutassou, Trachurus trachurus, Sardina pilchardus, Scomber scombrus). Among all these prey species, the relevance of M. poutassou and G. argenteus stood out because of their high abundance both in the diets and during the scientific surveys. The relevance of pelagic fish as prey increased with predator size, reaching more than 60% of the diet by volume in the length range 25–29 cm. However, fish predators larger than 50 cm depended less on pelagic fish (33%), since they were also able to feed on other sources such as megafaunal invertebrates, mainly cephalopods. The pelagic fish resource was primarily exploited by 12 demersal fish species, with Merluccius merluccius and Zeus faber being the main ones feeding on small pelagics, which was related to the vertical movement of predators and prey through the water column. Survey abundance indices were used as indicators of prey abundance in the ecosystem, both for biomass and number. There was evidence for density-dependant feeding by predators on E. encrasicolus, G. argenteus and T. trachurus, while the main discrepancies between abundance in the stomachs and in the surveys were due to differential availability of prey length classes in the environment. Small prey species and individuals were exploited by most demersal fish while large prey species were less accessible to predators.     

Distribution of drifting seaweeds in eastern East China Sea

In offshore waters with relatively low primary production, drifting seaweeds composed of Sargassum species form an identical ecosystem such as an oasis in desert. Commercially important pelagic fishes such as jack mackerel (Trachurus japonicus) and yellow tail (Seriola quinqueradiata) spawn in East China Sea pass their juvenile period accompanying drifting seaweeds. Therefore drifting seaweeds are very important not only in offshore ecosystem but also fishery resources. However the distribution of drifting seaweeds in East China Sea has scarcely known. Then we conducted two research cruises of R/V Hakuho–Maru in May 2002 and in March 2004. During the cruises, drifting seaweeds were visually observed from the bridge and sampled with a towing net. The observation revealed that the drifting seaweeds were distributed along the front between the Kuroshio Current and coastal waters and mainly composed of one seaweed species, Sargassum horneri (Turner) C. Agardh from spring to early summer. There are no reports on geographical distribution of this species in the coasts south of southern Kyushu Island in Japan. Kuroshio Current flows northeastward there. Buoys with GPS attached to drifting seaweeds released off Zhejiang Province, China, in March 2005 to track their transport. Their positions monitored by ORBCOM satellite showed that they were transported to the area in East China Sea, where the drifting seaweeds were observed during the cruises, in 2 months. These facts suggest that S. horneri detached from Chinese coast in March or months earlier than March could be transported to fringe area of continental shelf and waters influenced by Kuroshio Current from March to May. Therefore the Sargassum forests, especially S. horneri, along the Chinese coast play a very important role in the ecosystem of the East China Sea as a source of drifting seaweeds.     

Copepod diversity in the water masses of the southern East China Sea north of Taiwan

The warm oceanic current Kuroshio and the continental shelf water of the East China Sea meet in the western North Pacific, north of Taiwan and form an upwelling when they converge. The intrusion of the Kuroshio westward over the East China Sea shelf thus results in complicated exchanges of waters between these two water masses. We studied the copepods in the plankton collection taken from an east–west transect crossing these waters in April 1995 when the intrusion of the Kuroshio over the East China Sea shelf was beginning to retreat. The taxonomy of copepods was carefully treated and erroneous species records reported in the literature were guarded against. We evaluated the copepod diversity, the association of copepod species, and the association of stations in these water masses.     

Quantifying copepod kinematics in a laboratory turbulence apparatus

We describe application of a new apparatus that permits simultaneous detailed observations of plankton behavior and turbulent velocities. We are able to acquire 3D trajectories amenable to statistical analyses for comparisons of copepod responses to well-quantified turbulence intensities that match those found in the coastal ocean environment. The turbulence characteristics consist of nearly isotropic and homogeneous velocity fluctuation statistics in the observation region. In the apparatus, three species of copepods, Acartia hudsonica, Temora longicornis, and Calanus finmarchicus were exposed separately to stagnant water plus four sequentially increasing levels of turbulence intensity. Copepod kinematics were quantified via several measures, including transport speed, motility number, net-to-gross displacement ratio, number of escape events, and number of animals phototactically aggregating per minute. The results suggest that these copepods could control their position and movements at low turbulence intensity. At higher turbulence intensity, the copepods movement was dominated by the water motion, although species-specific modifications due to size and swimming mode of the copepod influenced the results. Several trends support a dome-shaped variation of copepod kinematics with increasing turbulence. These species-specific trends and threshold quantities provide a data set for future comparative analyses of copepod responses to turbulence of varying duration as well as intensity.     

Effects of closed trawl fishery areas on some elasmobranch species in the Cantabrian Sea

This study examines the effects of certain closed areas to the trawl fishery on some Rajidae species and small-spotted catshark in the southern area of the Bay of Biscay. The results are based on abundance index (number/haul) and catch length composition obtained from a series of bottom trawl surveys carried out in the area (1993–2005). Abundance trends are similar considering or not closed areas, but with much lower values excluding closed areas. The length distributions revealed that in some of the species like Scyliorhinus canicula the larger specimens were the most affected by this decrease in number. Finally, the effects and consequences of one of these marine protected areas (MPAs), an artificial reef were examined in more detail for all the fishes and invertebrates. The analysis of the yield series showed an increase in total biomass after the placement of the reefs not only for S. canicula and rays but also teleosts fishes like Pagellus acarne, Trachurus trachurus, Mullus surmuletus and Trachinus draco and particularly invertebrates (sea urchins). On the contrary, the main target species of the trawl fishery (hake, monkfish and megrim among others) did not increase.     

The role of copepods in the Baikal ecosystem

This describes a study of the pelagic zone of Lake Baikal with particular reference to copepods. In addition the cycling of matter and energy in lacustrine basins is described.     

Plankton distribution and the impact of copepod grazing on primary production in Fram Strait, Greenland Sea

Two hydrobiological transects across the East Greenland Shelf and the open waters of Fram Strait in summer were chosen to illustrate the distribution and production of phyto- and zooplankton in relation to water masses and ice cover. The parameters used were temperature and salinity, inorganic nutrients, chlorophyll a, primary production, phytoplankton species composition, abundance of the dominant herbivorous copepods Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa and egg production of C. finmarchicus and C. glacialis. Grazing impact of copepodites and adults of these four species was modelled for each station by using egg production rates as an index of growth. Seasonal development of plankton communities was closely associated with the extent of the ice cover, hydrographic conditions and the water masses typical of the different hydrographic domains. Four regions were identified from their biological activities and physical environment: The Northeast Water polynya on the East Greenland Shelf, with a springbloom of diatoms and active reproduction of herbivorous copepods. The pack ice region, dominated by small flagellates and negligible grazing activities. The marginal ice zone, with high variability and strong gradients of autotroph production related to eddies and ice tongues, an active microbial loop and low egg production. The open water, with high station-to-station variability of most of the parameters, probably related to hydrographic mesoscale activities. Here, Phaeocystis pouchetii was a prominent species in the phytoplankton communities. Its presence may at least partly be responsible for the generally low egg production in the open waters. Grazing impact on primary production was always small, due to low zooplankton biomass in the polynya and due to low ingestion in the remaining regions.     

Community structure of mesopelagic copepods (>500 μm) in the Ligurian Sea (Western Mediterranean)

The distribution of mesopelagic copepods between 250 and 1000 m depth was studied along a transect off the coast of Villefranche-sur-Mer in June 1991 by means of a BIONESS multiple-net sampler. Among the most abundant species at all stations were Pleuromamma gracilis, Paraeuchaeta acuta and overwintering Calanus helgolandicus CV copepodites, each species inhabiting different depth layers with maximum abundances in the 250–350 m, 450–550 m and the 700–850 m layer, respectively. Cluster analysis of the vertical distribution of all species caught revealed three distinct strata (‘TOP', ‘INTERMEDIATE', ‘BOTTOM') consisting of characteristic species that occurred in the same stratum at all stations. Differences in total abundances of the species assemblage occupying each layer are discussed with regard to predation pressure by mesopelagic macroplanktonic crustaceans and fish. We show that each stratum is characterized by distinct trophic interactions and life strategies, which play an important role in the structuring of mesopelagic zooplankton communities.     

New species of the genus Xanthocalanus (Copepoda, Calanoida, Phaennidae) from the Laptev Sea

Sampling of the near-bottom calanoid copepods was performed on board of the research vessel Polarstern in 1993 in the Laptev Sea (Siberian Arctic). Three new species of Xanthocalanus were identified as such, and are described and figured here: X. spinodenticulatus sp. nov., X. laptevorum sp. nov. and X. polarsternae sp. nov.     

Harpacticoid and cyclopoid fauna of groundwater and springs in southern Finland

The groundwater fauna, of which scarcely anything has previously been studied in formerly glaciated areas, was richest close to the water table and diminished markedly at greater depths. However, few individuals were generally recorded per 1 m³ of water. Bryocamptus minutus was the most abundant species of copepod at an esker site, the other species recorded beingAttheyella crassa, Bryocamptus pygmaeus, Moraria brevipes and Parastenocaris phyllura, and some species from the genus Diacyclops. At a shore bank infiltration site M. brevipes was the most numerous species. However, one esker site examined did not have any animals. Springs, which were grouped into those in a natural state and those under anthropogenic influence, had some species in common with esker groundwater, but also Paracyclops fimbriatus, Acanthocyclops robustus, Acanthocyclops vernalis, Diacyclops bicuspidatus, Megacyclops viridis, and the harpacticoid species Bryocamptus cuspidatus, Bryocamptus echinatus and Canthocamptus staphylinus. Of the spring copepods, P. fimbriatus appeared to withstand the influence of road de-icing with NaCl, but numbers of Moraria brevipes were reduced under this influence. Bryocamptus echinatus was more numerous in springs having a high oxygen content and high pH. The species found in esker groundwaters and springs were in part the same evidently cold-stenotherm species which inhabit the profundal zone of oligotrophic lakes.     

Distribution, reproduction and development of Calanus species in the Northeast water in relation to environmental conditions

The vertical and horizontal distribution of mesozooplankton biomass and its composition, together with the reproduction and development of the three dominant herbivorous copepods Calanus finmarchicus, C. glacialis and C. hyperboreus, were studied in the Northeast Water between the end of May and end of July, 1993. Biomass varied between 0.15 and 10.6 g m⁻² and was lower on the shallow banks. Highest biomass was found on the shelf slope and in the entrances to Belgica and Westwind Troughs. There was no seasonal trend during the study period. Among the zooplankton taxa, copepods were dominant, with 84% biomass of all other taxa, followed by chaetognaths with 14%. The large Calanus species made up 91% of copepod biomass. The most abundant species, C. glacialis, inhabited areas of low current speeds on Belgica and Ob Bank, C. hyperboreus dominated shelf slope and trough stations, while C. finmarchicus was most abundant in the Return Atlantic Current along the shelf slope and also eastern Belgica Trough. C. glacialis was the only Calanus species spawning during this period, but young copepodites of the other species were also abundant. Egg production of C. glacialis was at a maximum by our arrival and continued to at least mid August. According to the results from starvation experiments, its egg production was fuelled by food uptake, but was decoupled from phytoplankton chlorophyll until July, indicating ice-algae and microzooplankton as an alternative food source. Only when the polynya approached its maximum extent was a close relationship to phytoplankton established. Due to both spatial and temporal heterogeneity of the occurrence of young copepodite stages it was impossible to follow the growth of cohorts of developmental stages. Herbivorous carbon requirements estimated from egg production rates were mostly less than one third of the phytoplankton stock. From egg production and the distribution of young stages, the outer Westwing Trough seems to be the centre of biological activities. This may be related to the formation of young ice in winter in this area, which was found to carry a large mass of under-ice algae. High biomass but low production in Belgica Trough indicate this as an advective and expatriated community of C. finmarchicus and C. hyperboreus, where grazing is negligible, at least during early summer.