Dynamics of suprabenthos-zooplankton communities around the Balearic Islands (western Mediterranean): Influence of environmental variables and effects on the biological cycle of Aristeus antennatus

Dynamics of suprabenthos and zooplankton were analyzed in two areas located in the NW (off Sóller harbour) and S (off Cabrera Archipelago) of Mallorca (Balearic Islands, western Mediterranean) at depths ranging between 135–780 m. Four stations situated respectively at 150 m (shelf-slope break), and at bathyal depths of 350, 650 and 750 m were sampled at bi-monthly intervals during six cruises performed between August 2003 and June 2004. Suprabenthos showed maximum biomass in both areas from late spring to summer (April to August), while minimum biomass was found in autumn (September–November). Though variable, temporal dynamics of zooplankton showed peaks of biomass in late winter and summer (February and June), while minimals occurred in autumn (August–September) and, at bathyal depths, in April. Suprabenthos (abundance; MDS analyses) showed a sample aggregation as a function of depth (3 groups corresponding to the shelf-slope break, upper slope — over 350 m; and the middle, deeper part of the slope — over 650–750 m), without any separation of hauls by season. By contrast, zooplankton samples were separated by season and not by depth. There was evidence of three seasonal groups corresponding to summer (June 2004–August 2003), autumn–winter (September and November 2003, February 2004), and spring (April 2004), being especially well established off Sóller. In general, suprabenthos was significantly correlated with the sediment variables (e.g. total organic matter content (% OM), potential REDOX), whereas zooplankton was almost exclusively dependent on Chl a at the surface, which suggests two different food sources for suprabenthos and zooplankton. The increase of suprabenthos abundance in April–June was paralleled by a sharp increase (ca. 2.8 times) in the %OM on sediment during the same period, coupled ca. 1–2 months of delay with the peak of surface Chl a recorded in February–March (from satellite imagery data). Suprabenthos biomass was also correlated with salinity close to the bottom, suggesting a link between suprabenthos abundance and changes in the oceanographic condition of water masses close to the bottom. It is suggested that a higher suprabenthos biomass recorded off Sóller in comparison to that off Cabrera in June could, in turn, be related to a seasonal inflow of Levantine Intermediate Water (LIW) in April–June in this area at mid bathyal depths (350–650 m). This trend would be based on: 1) it was evident only at mid-slope depths between 350–750 m, coinciding with the LIW distribution, and 2) it was not recorded among zooplankton (collected throughout the water column). The possible effect of the fluctuations of suprabenthos and zooplankton on higher trophic levels has been explored studying the diet and food consumption rates of the red shrimp Aristeus antennatus, as indicator species by its dominance in bathyal communities. A. antennatus increased its food consumption from February to April–June 2004 off Sóller, which in the case of large (CL > 40 mm) specimens was found in both areas. In addition, there was a shift of diet from winter to spring–early summer. In this last period, A. antennatus preyed upon euphausiids and mesopelagic decapods and fish, while benthos (e.g. polychaetes and bivalves) decreased in the diet. This indicates an increase in the food consumption and probably in the caloric content of the diet in pre-spawning females in April–June 2004, which is synchronized with the period when gonad development begins in A. antennatus females (May–June). Anyway, macrozooplankton, and not suprabenthos, is crucial as a high energetic food source in the coupling between food intake and reproduction in the red shrimp.     

Spatio-temporal variations in deep-sea demersal communities off the Balearic Islands (western Mediterranean)

The spatial and temporal variations of deep-sea megafaunal assemblages from the western Mediterranean are analysed in the present paper. The assemblages from two locations of the Balearic Islands situated 120 km apart were compared using data collected seasonally on a bathymetric stratum covering the 150–750 m depth range during six bottom-trawl surveys. The assemblage structure, in terms of species composition, species dominance and population sizes, was differentially affected by the spatio-temporal variables analysed (depth, location and fishing period). Although depth was the main factor determining the assemblage composition, the differences obtained between the two locations were also relevant. On the upper slope these between-location differences in the dynamics of megafaunal assemblages were found to be related to the effect of fishing exploitation. Population size-based metrics and biomass spectra were good predictors of meso-scale fishing effects, and were mainly reflected by elasmobranchs and demersal teleosts. Nevertheless, the effects of fishing depended on the species considered. Two dominant large-sized fish species found on the upper slope in both localities, Galeus melastomus and Phycis blennoides, had higher biomass values associated with lower fishing effort. Although the mean body weight (MBW) of both species and also the mean maximum body weight (MMBW) of G. melastomus agreed with this pattern, the P. blennoides MMBW did not. This last case could be indicative of natural size-trends such as the bigger–deeper phenomenon which refers to the displacement of large individuals towards the deeper limit of their bathymetric distribution, beyond the maximum depth sampled in this study for this species. By contrast, the target species of the upper slope fishery, the red shrimp Aristeus antennatus, was not negatively affected by the direct impact of fishing activity and other environmental factors, such as the presence of specific water masses could also be important.     

Habitat characteristics as determinant of the structure and spatial distribution of epibenthic and demersal communities of Le Danois Bank (Cantabrian Sea, N. Spain)

The purpose of this paper is to detect and describe general trends in the spatial distribution of epibenthic and demersal communities of Le Danois Bank (El Cachucho) in relation to the environmental variables that characterize their habitat by means of multivariate ordination. Data were derived from two multidisciplinary surveys carried out in October 2003 and April 2004 included in the ECOMARG project. The surveys were focused on the study of the physical scenario, including both geological (seabed characterization from bathymetry and backscatter data) and hydrographic features, and the different compartments of the benthic fauna (endobenthic, epibenthic, suprabenthic and demersal communities). For the present study, epibenthos and demersal species were sampled using two different gears, a 3.5 m beam trawl and a Porcupine 39/52 type baca otter trawl respectively. The total species richness combining both samplers ascended to 221, including 71 species of fishes, 65 crustaceans, 35 molluscs, 29 echinoderms, 10 cnidarians and 5 sponges.Multivariate methods were used for the study of the characteristics of communities and habitats. Hierarchical cluster analysis was applied to calculate and visualize similarities between samples in terms of species composition. To assess the amount of variation of faunal densities related to a set of eight environmental variables, a redundancy analysis (RDA) was used. The set of environmental variables used were: depth, near-bottom temperature and salinity, sedimentary typology (dry weight percentages of coarse sands, medium and fine sands, silt and organic matter) and seafloor reflectivity. Using the spatial distribution of the ranges of depth and seafloor reflectivity that characterize the habitats of the faunal assemblages we defined the spatial distribution of the different communities.The multivariate analysis of 18 beam-trawl samples and 15 otter trawl samples showed the existence of 4 main assemblages associated with the more characteristic habitats of the area. The Callogorgia–Chimaera community live mainly in the areas at the top of the Bank (425–550 m depth), where the sediment coverage is lower and there exists a high presence of rocky outcrops. The Gryphus–Galeus community were found in the areas at the top of the Bank covered by medium and fine sand sediments with low organic matter content. The Phormosoma–Trachyrincus community live on the sedimentary terraces that characterize the Bank break (550–800 m) and the Pheronema–Deania community occupy all the deeper (800–1050 m) sedimentary grounds of the inner basin.     

The annual cycle of nanoflagellates in the Central Cantabrian Sea (Bay of Biscay)

The annual cycle of nanoflagellates (NF) including autotrophic (ANF), heterotrophic (HNF) and mixotropic (MNF) flagellates carried out in a temperate sea (Central Cantabrian Sea, southern Bay of Biscay) is presented. Three stations with characteristics ranging from coastal to oceanic conditions were analysed in order to compare NF response to this gradient. Samples were monthly collected at each station at three different depths between February 2002 and December 2002. CTD profiles were also taken at each station. NF were grouped according to their trophic status into ANF, HNF and MNF. Abundance and biomass were determined for each group. The annual cycle showed a general pattern consisting in a maximum in July with secondary maxima in March and October and minimum values in May. ANF were the most important fraction, making a major contribution (nearly 75%) to total NF biomass in all stations. HNF represented over 20% along the cycle, except for a peak in spring found in every station. MNF reached less than 5%, showing low seasonability. Small flagellates (2–5 µm) dominated throughout the cycle. Microplankton community was also analysed in terms of abundance and biomass. A significant positive correlation (r² = 0.49) was obtained between 2–5 µm NF and 10–20 µm HNF–MNF biomasses, suggesting a possible trophic relationship between these groups which should be cautiously taken. No significant relationships were found between microplankton and NF or between nutrients and ANF, indicating that the regulation of NF numbers is complex and probably implicates other groups. In addition to this, the unexpected 2002 Chl a concentration pattern and the misplacing of upwelling events render necessary to perform additional studies to fully understand the precise behaviour of NF in the Cantabrian Sea. To the best of our knowledge, this is the first study of a NF cycle in a temperate sea that considers all functional groups.     

Mesozooplankton species distribution in the NW and N Iberian shelf during spring 2004: Relationship with frontal structures

We have analysed the mesozooplankton community structure in the southern Bay of Biscay shelf and its relationship with the hydrographic conditions during spring 2004. According to thermohaline characteristics, we observed two frontal zones of distinct origin along the shelf (around 7° and 3°W), that allowed us to differentiate three different hydrographic domains. The westernmost part of the shelf (WC), defined by the presence of relatively warm and salty water related to the presence of the Iberian Poleward Current (IPC), the easternmost region (EC), characterized by colder and fresher water and subject to the influence of freshwater inputs from the Adour river in the French coast, and a region in the Central Cantabrian Sea (CC), where thermohaline characteristics were intermediate between these two extremes. The mixing layer depth (MLD) regime in these areas was also different: the WC region was characterized by a mixed water column, whereas in the EC region the river discharges produces stratification of the upper meters of the water column (< 10 m); in the CC region, we found a distinct vertical mixing regime that separated coastal (stratification) from shelf (mixed water column) stations, giving rise to a notorious across-shelf front. We found a good match between the aforesaid hydrographic regions and the distribution of mesozooplankton species composition and community assemblages: the Mantel correlation between physical variables and mesozooplankton distribution was highly significant (n = 63, r = 0.70, α < 0.001). In the WC region, the community was dominated by Paracalanus parvus, Oithona helgolandica, Acartia clausi and Clausocalanus pergens, while in the EC region the most dominant species were Noctiluca scintillans, Oncaea media and Temora longicornis. The CC region showed similar composition of copepods than the WC region, but larvaceans (Oikopleura spp. and Fritillaria spp.) were more abundant in the CC region than in the WC region. Within each zone, the relative abundances of the dominant species differed between coastal and shelf locations.     

A numerical investigation of phytoplankton and Pseudocalanus elongatus dynamics in the spring bloom time in the Gda sk Gulf

A nutrient–phytoplankton–zooplankton–detritus (1D-NPZD) ‘phytoplankton {Phyt} and Pseudocalanus elongatus {Zoop} dynamics in the spring bloom time in the Gda sk Gulf. The 1D-NPZD model consists of three coupled, partial second-order differential equations of the diffusion type for phytoplankton {Phyt}, zooplankton {Zoop}, nutrients {Nutr} and one ordinary first-order differential equation for benthic detritus pool {Detr}, together with initial and boundary conditions. In this model, the {Zoop} is presented by only one species of copepod (P. elongatus) and {Zoop} is composed of six cohorts of copepods with weights (W_i) and numbers (Z_i); where . The calculations were made for 90 days (March, April, May) for two stations at Gda sk Gulf with a vertical space step of 0.5m and a time step of 900 s. The flow field and water temperature used as the inputs in the biological model 1D-NPZD were reproduced by the prognostic numerical simulation technique using hydrographic climatological data. The results of the numerical investigations described here were compared with the mean observed values of surface chlorophyll-a and depth integrated P. elongatus biomass for 10 years, 1980–1990. The slight differences between the calculated and mean observed values of surface chlorophyll-a and zooplankton biomass are ca. 10–60 mg C m⁻³ and ca. 5–23 mg C m⁻², respectively, depending on the location of the hydrographic station. The 1D-NPZD model with a high-resolution zooplankton module for P. elongatus can be used to describe the temporal patterns for phytoplankton biomass and P. elongatus in the centre of the Gda sk Gulf.     

Distribution of phytoplankton in the southern Black Sea in summer 1996, spring and autumn 1998

The species composition, abundance, and biomass of micro- (>15 μm) and nano- (<15 μm) phytoplankton were studied along the southern Black Sea during June–July 1996 and March–April and September 1998. A total of 150 species were identified, 50% of them being dinoflagellates. The average total phytoplankton abundance changed from 77×10³ cells l⁻¹ in spring to 110×10³ cells l⁻¹ in autumn and biomass from 250 μg l⁻¹ in summer to 1370 μg l⁻¹ in spring. Based on the extensive sampling grid from June–July 1996, phytoplankton seemed to have a rather homogeneous biomass distribution in the southern Black Sea. In all periods, the coccolithophorid Emiliania huxleyi was the most abundant species, its contribution to the total abundance ranging from 73% in autumn to 43% in spring. However, in terms of biomass, diatoms made up the bulk of phytoplankton in spring (97%, majority being Proboscia alata) and autumn (73%, majority being Pseudosolenia calcar-avis), and dinoflagellates in summer (74%, Gymnodinium sp.). There was a remarkable similarity in the dominant species between the western and eastern regions of the southern Black Sea, indicating transport of phytoplankton within the basin.     

Experiments on the survival of six brackish macro-invertebrates from the Baltic Sea after dredged spoil coverage and its implications for the field

Physical disturbance by disposal of dredged materials in estuarine and coastal waters may result in burial of benthic fauna. Survival rates depend on a variety of factors including the type and amount of disposed materials and the lifestyle of the organisms. Laboratory burial experiments using six common macrobenthic invertebrates from a brackish habitat of the western Baltic Sea were performed to test the organisms' escape reaction to dredged material disposal. Experimental lab-results were then extrapolated to a field situation with corresponding bottom topography and covering layer thicknesses at experimental field disposal study sites. Resulted survival rates were then verified by comparison with results of an earlier field study at the same disposal sites.Our experimental design in the lab included the disposal of two types of dredged material (i.e. ‘till’ and ‘sand/till mixture’) and two covering layer depths (i.e. 10–20 cm and 14–40 cm). All three bivalves Arctica islandica (Linnaeus), Macoma balthica (Linnaeus), Mya arenaria (Linnaeus) and the polychaete Nephtys hombergii (Savigny) successfully burrowed to the surface of a 32–41 cm deposited sediment layer of till or sand/till mixture and restored contact with the overlying water. These high escape potentials could partly be explained by the heterogeneous texture of the till and sand/till mixture with ‘voids’. The polychaete Bylgides (Harmothoe) sarsi (Malmgren) successfully burrowed through a 16 cm covering layer whereas the polychaete Lagis koreni (Malmgren) showed almost no escaping reaction. No general differences in escape behaviour after burial were detected between our test species from the brackish habitat and those reported in the literature for the same species in marine environments. However, a size-dependence in mobility of motile polychaetes and M. arenaria was apparent within our study. In comparison to a thick coverage, thin covering layers (i.e. 15–16 cm and 20 cm) increased the chance of the organisms (N. hombergii and M. arenaria) to reach the sediment surface after burial. This was not observed for the other test species. While crawling upward to the new sediment surfaces burrowing velocities of up to 8 cm d^− 1 were observed for the bivalves and up to 20 cm d^− 1 for N. hombergii. Between 17 and 79% of the test organisms showed burrowing activity after experimental burial. The survival rate (defined as the ability to regained contact with the sediment surface) ranged from 0 to 33%, depending on species and on burial depth. The organisms reached the sediment surface by burrowing (polychaetes and bivalves) and/or by extending their siphons to the new sediment surface (bivalves). The extrapolation of laboratory survival rates to the two disposal sites was obtained based on the in situ thicknesses of the dredged spoil layers measured by multi-beam echo sounder. This resulted in total average survival rate estimates for the test species of 45 and 43% for the two disposal sites. The results obtained during the laboratory tests and the following extrapolation to the field were verified by the range of results from a previous field study, using grab sampling shortly before and after a disposal event in June 2001. The effect of dredged material disposal on the tested Baltic Sea benthic macrofauna was assessed by extrapolating the verified laboratory results to the field.     

Variability of mesozooplankton spatial distribution in the North Aegean Sea, as influenced by the Black Sea waters outflow

The North Aegean Sea constitutes an important region of the Mediterranean Sea since in its eastern part the mesotrophic, low salinity and relatively cold water from the Black Sea (outflowing from the Dardanelles strait) meets the oligotrophic, warm and very saline water of Levantine origin, thus forming a thermohaline front. Mesozooplankton samples were collected at discrete layers according to the hydrology of the upper 100 m, during May 1997 and September 1998. In May highest biomass and abundance values (up to 66.82 mg m^− 3 and 14,157 ind m^− 3) were detected in the 10–20 m layer (within the halocline) of the stations positioned close to the Dardanelles strait. The front moved slightly southwards in September, characterized by high biomass and abundance values within the halocline layer. The areas moderately or non influenced by Black Sea water revealed lower standing stock values than the frontal area in both cruises and maxima were detected in the uppermost low salinity layer. Samples collected at the stations and/or layers more influenced by Black Sea water were distinguished from those collected at layers and/or stations more affected by Levantine waters in both periods. In May the former samples were characterized by the copepods Acartia clausi, Centropages typicus, Paracalanus parvus. The abundance of the above species decreased gradually with increasing salinity, in the horizontal and/or in the vertical dimension, with a parallel increase of the copepods Oithona plumifera, Oithona copepodites, Oncaea media, Ctenocalanus vanus, Farranula rostrata. During September the frontal area as well as that covered by the modified Black Sea water, were highly dominated by the cladoceran Penilia avirostris and doliolids. For both seasons, MDS plots, issued from the combination of mesozooplankton and water-type data, revealed the gradual differentiation of zooplankton composition from the frontal area towards the area covered by Levantine water, following the spreading and mixing of the Black sea water. The observed temporal and spatial variability in the distribution pattern of mesozooplankton standing stock and species composition seems to depend considerably on the variability of circulation and frontal flows.     

Pigments, size and distribution of Synechococcus spp. in the Black Sea

Pigments, size and distribution of Phycoerythrin-containing unicellular cyanobacteria Synechococcus spp. within the euphotic zone were studied for the first time in April–May 1994 in the western and southwestern Black Sea by epifluorescence microscopy and flow-cytometry. Synechococcus was present in varying quantities at every station and depth studied. Surface spatial distribution of Synechococcus revealed that cells were much more abundant in offshore waters than near coastal regions under the direct influence of the Danube river. Minimum and maximum cell concentrations ranged between 9×10² and 1.45×10⁵ cells/ml at the surface, between 2×10³ and 1.23×10⁵ cells/ml at the chlorophyll sub-maximum layer, and between 1.3×10² and 3.5×10² at the nitrite maximum layer. Cells at the chlorophyll sub-maximum layer (based on in-situ fluorometer readings) fluoresce brighter and longer than the ones at the surface and lower depths. Spectral properties of chromophore pigment types of total 64 clonal isolates from different depths down to the lower layer of the euphotic zone (60 m) in the southern Black Sea coast revealed that all have type 2 phycoerythrobilin in common, lacking in phycourobilin. In vivo fluorescence emission maxima for the phycoerythrobilin were about the same (578 nm) for all isolates. All isolates examined showed in vivo absorption maxima at between 435 and 442 nm and at about 681 nm due to chlorophyll-a. Based on the flow cytometer mean forward light scatter data for size distribution, it could be concluded that cells at the surface mixed layer (0–10 m) were larger in cell size than the cells at lower depths (20–60 m).     

Summer distribution of seabirds in the North-East Water polynya, Greenland

The distribution at sea of seabirds was studied in the North-East Water (NEW) polynya, Greenland, during transect counts in the summers of 1991, 1992 and 1993 on board the ice-breaking RVs Polarstern and Polar Sea. Data collected within the polynya ‘box’ (78–82°N; 5–18°W) concern observations of 8000 birds counted during 1350 half-hour counts. Distribution is presented as density (N/km²) and calculated daily food intake. Five bird species were selected for discussion, representing more than 95% of the total numbers encountered: Fulmar (Fulmarus glacialis), Ivory Gull (Pagophila eburnea), Kittiwake (Rissa tridactyla), Glaucous Gull (Larus hyperboreus) and Ross's Gull (Rhodostethia rosea). For these species, densities are comparable in the NE Greenland polynya and in other European Arctic seas. The main difference is the absence in NEW of the species playing the main role in Arctic seas: Brünnich's Guillemot (Uria lomvia) and Little Auk (Alle alle). In the absence of fish-eating birds and of birds consuming zooplankton in the water column, the NEW polynya ecosystem is thus dominated by surface feeders and, closer to the coast, by benthic feeders like eiders, Somateria mollissima and S. spectabilis, and walrus, Odobenus rosmarus. The density and daily food intake for all seabirds are one order of magnitude lower in the polynya than in the Arctic seas. The distribution and abundance of seabirds in the NEW polynya seems to reflect a very low density of pelagic fish and Zooplankton in the water column, while Zooplankton must be present at ‘normal’ concentrations in the upper layer.     

Westward moving waves or eddies (Storms) on the Subtropical/Azores Front near 32.5°N? Interpretation of the Eulerian currents and temperature records at moorings 155 (35.5°W) and 156 (34.4°W)

Three Argos buoy-years of Lagrangian data in westward-moving cyclonic eddies, or Storms, near 32.5°N, together with hydrographic measurements, have shown that Storms move westward at nearly 3 km day⁻¹. Water in eddies can be trapped and moved westward by advection within the eddy or by phase propagation of the eddy pattern, so we cannot say that the flow field (or Eulerian mean) is 3 km day⁻¹ westward. Two moorings (155 and 156) deployed in the Storm Corridor have provided a further 8 instrument-years of Eulerian data. The temperature and current records confirmed that two Storms a year passed each mooring over the 2-year measurement period. As expected, there is a lag of 1.3 month at mooring 155 (which is 102 km to the west of mooring 156) with respect to conditions at mooring 156. The progressive vector diagrams (PVDs) derived from the current meter records exhibit fairly regular X (east or zonal) and Y (north or meridional) displacement scales that repeat with semi-annual periodicity (SAP). The dominant current signal is the north component of the SAP, which reaches an amplitude of 18 cm s⁻¹ for the upper layer or near surface current record (242-m depth). The geostrophic north component values derived from altimetry were in good agreement with the upper layer current meter measurements. The large north component amplitude was not interpreted as evidence for Rossby Waves but rather due to the passage of nine eddies (eight complete) of alternate sign (cyclonic, anticyclonic) passing the mooring rigs during the 2-year deployment period. The Y scale shows that the near surface characteristic or mean maximum azimuthal speed is about 35 cm s⁻¹ for cyclonic eddies or Storms, and that this value is reduced to 4 cm s⁻¹ at 1400-m depth. The residual or mean Eulerian currents range from 8 cm s⁻¹ for the upper layer currents to 1 cm s⁻¹ for the deeper currents at 1400-m depth and are predominantly westward. Simple theoretical considerations and idealised numerical simulations show that the mean westward Eulerian current depends markedly on whether the eddy centres pass to the north or south of the rigs. This raises the question as to what do we mean by Eulerian residual currents, even for relatively long records (2 years). It is shown that the strong near surface westward current (6 km day⁻¹) measured at mooring 155 is largely due to a westward-moving eddy field with variable centre offsets. The magnitude of the near surface east–west component of flow was estimated as eastward at 2 cm s⁻¹. The north–south component of mean flow was southward at 2 cm s⁻¹. The deeper records gave a weak westward flow of 1 cm s⁻¹ but did not show a significant southward component for the mean Eulerian flow field. 7.4 float-years of Lagrangian ALACE data in the Subtropical Front region near 740 dbar gave mean east–west flows that were <0.5 cm s⁻¹. Overall, it is shown that the eddy structures propagate westward mainly by phase propagation (i.e. a westward-moving pattern with no westward advection for the current meter to measure), though plane Rossby Wave dynamics appeared inappropriate. Theoretical and modeling considerations show that a speed of 3-km day⁻¹ westward is too large a value for the self-advection of eddies due to the beta effect.     

Influence of the Mackenzie River plume on the sinking export of particulate material on the shelf

We examined the influence of the Mackenzie River plume on sinking fluxes of particulate organic and inorganic material on the Mackenzie Shelf, Canadian Arctic. Short-term particle interceptor traps were deployed under the halocline at 3 stations across the shelf during fall 2002 and at 3 stations along the shelf edge during summer 2004. During the two sampling periods, the horizontal patterns in sinking fluxes of particulate organic carbon (POC) and chlorophyll a (chl a) paralleled those in chl a biomass within the plume. Highest sinking fluxes of particulate organic material occurred at stations strongly influenced by the river plume (maximum POC sinking fluxes at 25 m of 98 mg C m^− 2 d^− 1 and 197 mg C m^− 2 d^− 1 in 2002 and 2004, respectively). The biogeochemical composition of the sinking material varied seasonally with phytoplankton and fecal pellets contributing considerably to the sinking flux in summer, while amorphous detritus dominated in the fall. Also, the sinking phytoplankton assemblage showed a seasonal succession from a dominance of diatoms in summer to flagellates and dinoflagellates in the fall. The presence of the freshwater diatom Eunotia sp. in the sinking assemblage directly underneath the river plume indicates the contribution of a phytoplankton community carried by the plume to the sinking export of organic material. Yet, increasing chl a and BioSi sinking fluxes with depth indicated an export of phytoplankton from the water column below the river plume during summer and fall. Grazing activity, mostly by copepods, and to a lesser extent by appendicularians, appeared to occur in a well-defined stratum underneath the river plume, particularly during summer. These results show that the Mackenzie River influences the magnitude and composition of the sinking material on the shelf in summer and fall, but does not constitute the only source of material sinking to depth at stations influenced by the river plume.     

Seasonal carbon distribution of copepods in the eastern Weddell Sea, Antarctica

Copepods were sampled by a multiple opening-closing net in the eastern Weddell Sea during various seasons (late winter/early spring, summer, autumn). Total copepod biomass integrated over the upper 1000 m varied seasonally between 1.7 mg C m⁻³ in late winter/early spring and 3.7 mg C m⁻³ in autumn. After the dark season the copepods were rather evenly distributed vertically and highest biomass levels were found in the mid-water layers between about 200 m and 500 m. By contrast, especially in summer but also in autumn copepod biomass concentrated in the uppermost water layer. A total of 64 calanoid species were identified in the upper 1000 m with maximum species numbers in the deepest layer. The large calanoids Calanus propinquus, Calanoides acutus, Metridia gerlachei, Euchaeta antarctica and the small calanoid Microcalanus pygmaeus prevailed and accounted for 60–70% of total copepod biomass, while the small poecilostomatoid Oncaea and the cyclopoid Oithona species comprised about 20%. Hence, the distribution pattern of the entire copepod biomass is strongly influenced by the life cycles of a few dominant species.     

Picophytoplankton and nanophytoplankton abundance and distribution in the southeastern Beaufort Sea (Mackenzie Shelf and Amundsen Gulf) during Fall 2002

The distribution of picophytoplankton (0.2–2 µm) and nanophytoplankton (2–20 µm) in the Beaufort Sea–Mackenzie Shelf and Amundsen Gulf regions during autumn, 2002 is examined relative to their ambient water mass properties (salinity, temperature and nutrients: nitrate + nitrite, phosphate, and silicate) and to the ratio of variable to maximum fluorescence, Fv/Fm. Total phytoplankton and cell abundances (< 20 µm) were mainly correlated with salinity. Significant differences in picophytoplankton cell numbers were found among waters near the mouth of the Mackenzie River, ice melt waters and the underlying halocline water masses of Pacific origin. Picophytoplankton was the most abundant phytoplankton fraction during the autumnal season, probably reflecting low nitrate concentrations (surface waters average ~ 0.65 µM). The ratio Fv/Fm averaged 0.44, indicating that cells were still physiologically active, even though their concentrations were low (max Chl a = 0.9 mg m^− 3). No significant differences in Fv/Fm were evident in the different water masses, indicating that rate limiting conditions for photosynthesis and growth were uniform across the whole system, which was in a pre-winter stage, and was probably already experiencing light limitation as a result of shortening day lengths.     

Seasonal variability of the Black Sea chlorophyll-a concentration

The optimal spectral decomposition (OSD) method is used to reconstruct seasonal variability of the Black Sea horizontally averaged chlorophyll-a concentration from data collected during the NATO SfP-971818 Black Sea Project in 1980–1995. During the reconstruction, quality control is conducted to reduce errors caused by measurement accuracy, sampling strategy, and irregular data distribution in space and time. A bi-modal structure with winter/spring (February–March) and fall (September–October) blooms is uniquely detected and accurately documented. The chlorophyll-a enriched zone rises to 15 m depth in winter and June, and deepens to 40 m in April and 35 m in August. The June rise of the chlorophyll-a enriched zone is accompanying by near-continuous reduction of upper layer maximum chlorophyll-a concentration.     

Hydrographic structure and zooplankton abundance and diversity off Paita, northern Peru (1994 to 2004) — ENSO effects, trends and changes

The objective of the present study was to verify possible spatial, seasonal, and inter-annual changes in the zooplankton off Paita (northern Peru), an upwelling area located closely to the limits of cold Humboldt Current and warm Equatorial Surface Waters. Zooplankton was sampled at subsurface on 53 occasions from August 1994 to December 2004 at four stations located 2 to 30 km offshore with a WP-2 net (300 µm). Extremely high surface water temperatures combined with low salinities were observed during the 1997/98 El Niño up to 29.0 °C) and in April 2002 (up to 25.0 °C). Temperatures more than 2 °C above monthly average were also observed in October 1994, in April 2000, and in November 2004. Significant trends were observed for oxygen concentration (increase) and several horizontal and vertical gradients. Among the copepods (72% of all individuals), the most abundant species were Paracalanus parvus (28%), Acartia tonsa (26%), and Calanus sp. (10%). The strong 1997–98 El Niño (EN) event led to drastic changes in species composition that were reversed during the 1998–99 La Niña (LN) event. Community parameters such as total abundance, diversity, equitability and species richness displayed marked variations associated with the 1997–98 EN and long-term trends. Long-term trends were significant for several vertical and horizontal temperature and oxygen gradients, indicating an increase in upwelling intensity at the shelf during the study period. 10-year-trends were also significant for total zooplankton abundance (increase) and community evenness (J, decline). Our data confirmed the importance of the weak EN in 2002/03 for the study region. Within the trend of increasing zooplankton abundance, a sharp step or shift was observed from 1999 to 2000. When using sequential t-tests to detect shifts in (x + 1) transformed abundance data, a significant rupture was found between the last sampling in 1999 and the first sampling in 2000. Also, a substantial decrease in diel variability occurred after 1999, probably due to changes in vertical migration patterns. The considerable increase in zooplankton abundance over the study period, the ENSO effect, and the 1999–2000 transition are discussed with regard to synchronicity with other zooplankton time series. The present study contributes with the first evidence from an important area located in the Humboldt Current for synchronous trends and changes that were previously observed elsewhere in the Pacific. Our results demonstrated the importance of long-term zooplankton monitoring studies in upwelling areas, and confirms the idea of dramatic changes in pelagic ecosystem structure occurring in the East Pacific.     

Diel vertical migration and feeding in adult female Calanus pacificus, Metridia lucens and Pseudocalanus newmani during a spring bloom in Dabob Bay, a fjord in Washington USA

Diel vertical migration and feeding on phytoplankton by adult female Calanus pacificus, Metridia lucens and Pseudocalanus newmani were simultaneously measured near the end of a phytoplankton bloom. Almost the entire Calanus population migrated out of the deep layer (108–50 m) at night but only about 30% came to the surface (25–0 m). Feeding occurred only at night and was equally high in the surface and intermediate layers, in spite of much higher food concentrations in the surface. Like Calanus, the entire Metridia population was found in the deep layer during the day but unlike Calanus, 20–50% remained in the deep layer at night and most migratory Metridia were collected from the surface layer. Metridia feeding at night was highest in the surface layer but significant feeding also occurred in both the intermediate and deep layers. Migratory behavior of Pseudocalanus was weak, with the proportion of the population in the surface layer increasing from slightly <10% during the day to approx 30% at night. Feeding occurred in both surface and intermediate layers throughout the 24 h but was greater in both layers at night. The different migratory patterns are discussed in the context of our current understanding of the contributions of predator avoidance and feeding to diel vertical migration.     

Thorium-234 derived information on particle residence times and sediment deposition in shallow waters of the south-western Baltic Sea

Activities of the naturally occurring, short-lived and highly particle-reactive radionuclide tracer ²³⁴Th in the dissolved and particulate phase were measured at three shallow-water stations (maximum water depths: 15.6, 22.7 and 30.1 m) in Mecklenburg Bay (south-western Baltic Sea) to constrain the time scales of the dynamics and the depositional fate of particulate matter. Activities of particle-associated (> 0.4 μm) and total (particulate + dissolved) ²³⁴Th were in the range of 0.08–0.11 dpm L^− 1 and 0.11–0.20 dpm L^− 1, respectively. The activity ratio of total ²³⁴Th and its long-lived and conservative parent nuclide ²³⁸U was well below unity (range: 0.09–0.19) indicating substantial radioactive disequilibria throughout the water column, very dynamic trace-metal scavenging and particle export from the water column at all three stations. For the discussion the ²³⁴Th data of this study were combined with previously published water-column ²³⁴Th and particulate-matter data from Mecklenburg Bay (Kersten et al., 1998. Applied Geochemistry 13, 339–347). The resulting average vertical distribution of total ²³⁴Th/²³⁸U disequilibria was used to estimate the depositional ²³⁴Th flux to the sediment. There was a virtually constant net downward flux of ²³⁴Th of about 28 dpm m^− 2 d^− 1 leaving each water layer of one meter thickness. Thorium-234-derived net residence times of particulate material regarding settling from a given layer in the water column were typically on the order of days, but with maximum values of up to a couple of weeks. Based on an average ratio of particulate matter (PM) to particle-associated ²³⁴Th a net flux of about 145 mg PM m^− 2 d^− 1 was estimated to leave each water layer of one meter thickness. The estimated cumulative water-column-derived particulate-matter fluxes at the seafloor are higher by a factor of about 2 than previously published sediment-derived estimates for Mecklenburg Bay. This suggests that about half of the settling particulate material is exported from the study area and/or subject to processes such as mechanical breakdown, remineralisation and dissolution. Lateral particulate-matter redistribution and particle breakdown in the water column (as opposed to the sediment) seem to be favoured by (repeated) particle resuspension from and resettling to the seafloor before ultimate sedimentary burial. The importance of net lateral redistribution of particulate material seems to increase towards the seafloor and be particularly high within the bottommost few meters of the water column.