Distribution and diversity of open-ocean,near-bottom macroplankton in the western Mediterranean: Analysis at different spatio-temporal scales

We analyzed the distribution, diversity, and composition of western Mediterranean macroplankton (excluding gelatinous taxa) in the water column over depths of ca. 550–850 m, with special attention to near-bottom (0–1.5 and ca. 5–77 m above the bottom, mab) levels, and including data from three areas (off the coasts of Catalonia, and to the NW, and SE of Mallorca, Balearic Islands) in the period 1991–2008. Spatio-temporal changes in macroplankton abundance were evaluated as follows: (i) by seasonal sampling in 2007 off the Catalonian coast, (ii) by comparing Catalonian and Balearic Island slopes, and (iii) by comparing a fixed station on the Catalonian slope (at 550–800 m depth) at decadal (1991/1992–2007/2008) time scales. Diversity (in terms of species richness, S) was greater (i) at ca. 5–77 mab than at 0–1.5 mab, (ii) over the insular slopes of the Balearic Island (around Mallorca) than over the mainland Catalonian slopes, and (iii) in the period 1991/1992 than in 2007, likely related to higher values of the North Atlantic Oscillation (NAO) index in 1991/1992. In most analyses species composition was strongly influenced by the degree of stratification and homogenization of the water column in summer–autumn and winter–spring respectively and by location (longitude). Changes consisted mainly of higher density of macroplankton (e.g. abundance of the dominant euphausiids Nematoscelis megalops, Meganyctiphanes norvegica and Euphausia krohni and of the fish Cyclothone braueri) between June and October, parallel to an increase in the T and S close to the bottom. This coincided with changes in the flow of Levantine intermediate water (LIW) in the area. Aggregation of adult forms of the dominant species close to the bottom in summer–autumn could be favored because summer is the period of highest density of food – copepods, mainly Calanus helgolandicus – near the bottom off the Catalan slope. The formation of a thermocline in the water column and the reinforcement of the permanent thermohaline front at the shelf-slope break during summer at ca. 400 m in the Balearic Basin may also enhance this tendency toward greater aggregation of deep macroplankton under stratified water column conditions.     

The distribution of megabenthic, invertebrate epifauna in the Balearic Basin (western Mediterranean) between 400 and 2300 m: Environmental gradients influencing assemblages composition and biomass trends

The distribution of megabenthic epifauna (invertebrates) in the Balearic Basin (western Mediterranean) has been analyzed at depths between 427 and 2265 m after compiling samplings performed in 1985–1992 and 2007–2008 with an OTSB-14 bottom trawl. 84 epibenthic taxa of invertebrates (excluded decapod crustaceans) were collected. Epibenthic assemblages were organized in five groups (n-MDS analyses) as a function of increasing depth: upper slope assemblage, U, hauls between 427 and 660 m; middle slope assemblages M1 and M2, hauls between 663–876 m and 864–1412 m, respectively; lower slope assemblages L1 and L2, hauls between 1488–1789 m and 1798–2265 m, respectively). We found significant differences in assemblage composition between all depth-adjacent pairs of groups. Trends in the distribution of biomass vs. depth and within assemblages varied when hauls taken over insular were compared to those over mainland slopes. Over insular slopes we found (n-MDS) only four distinct depth assemblages, with significant differences between all depth-adjacent group pairs, except between L1 and L2. Over the mainland slope, two peaks of biomass situated at U (427–660 m) and at L1 (1488–1789 m) were clearly identified, attributable to the echinoid Brissopsis lyrifera and holothurian Molpadia musculus at U and to the synallactid holothurian Mesothuria intestinalis at L1. The distribution of biomass vs. depth on insular slopes did not follow this pattern, showing no significant biomass peak below 1000 m and a total biomass an order of magnitude lower than adjacent to the mainland. After compiling available environmental data over the mainland slope off Barcelona, we found coincidence between the peak biomass of Mesothuria intestinalis and: i) a significant increase of labile OM (%OrgC, C/N, hydrolizable aminoacids–EHAA, and the EHAA/THAA-total hydrolizable aminoacids-ratio) over 1600 m; and ii) an increase of turbidity and T at 1500–1600 m in February 2008. We suggest that such OM inputs must likely be associated to the formation of nepheloid layers close to submarine canyons, probably associated with oceanographic processes in deep water masses in the area. This would explain why aggregations of M. intestinalis were linked to the mainland part of the Balearic basin, with highest densities located south of canyons. If hotspots of biomass as cited here for M. intestinalis are regulated by factors such as river inputs, both natural climatic changes (e.g. changes in rainfall regimes) and human impact (e.g. river damming) may affect deep-Mediterranean communities below 1000 m.     

Changes in the diet and feeding of the hake Merluccius merluccius at the shelf-break of the Balearic Islands: Influence of the mesopelagic-boundary community

Short spatio–temporal variations in the feeding intensity and the diet of the European hake, Merluccius merluccius, together with the abundance of their potential prey were studied between August 2003 and June 2004 at two locations, northwest (Sóller) and south (Cabrera), off the island of Mallorca (Balearic Islands, Western Mediterranean) at depths between 150 and 750 m. The two areas present different oceanographic conditions. Hake was mainly distributed along the shelf-slope break and the upper slope (between 166 and 350 m) where recruits (TL<18 cm) were dominant. The hake's diet varied as a function of size. Recruits fed mainly on micronektonic prey, and the diet was influenced primarily by seasonality, with two dietary patterns (identified by MDS analyses) corresponding to August–September 2003 (summer) and to November 2003/February–April 2004 (autumn–winter). The summer pattern was consistent with a thermally stratified water column, while November and April were consistent with homogenized temperature and salinity throughout all the water column. The main prey of recruits were the euphausiid Meganyctiphanes norvegica and the midwater fish Maurolicus muelleri in autumn–winter and myctophids (mainly Ceratoscopelus maderensis) in summer. In contrast to recruits, the geographic factor (NW vs. S) was the main factor influencing the diets of post-recruits (TL between 18 and 21.9 cm) and adults (TL?22 cm). Hake recruits (and to a lesser extent post-recruits) and their preferred prey occupied different depth ranges during daylight periods. Meganyctiphanes norvegica and Ceratoscopelus maderensis were, for instance, distributed as much as 500 m deeper than hake that had eaten them. All these trends were especially obvious at NW, an area with a more abrupt slope and with a greater influence by northern winter intermediate water (WIW) inflow in early spring than the S area. These factors probably enhanced micronekton aggregation in April, when feeding intensity (stomach fullness) increased among recruits and post-recruits only at NW. All these factors may have a crucial role in the diet, distribution and probably recruitment success of small hake. Biological factors were also important in trophic shifts in the diet and feeding of hake. Multi-linear regression models pointed to a trend of higher fullness with higher hepato-somatic index (HSI). Therefore greater food consumption by hake may enhance its metabolic condition. Within the framework of shelf-break and slope ecology, we show how the ‘boundary’ mesopelagic community inhabiting the middle slope sustains the trophic requirements of hake, a species distributed at shallower depths along the shelf-slope break. Mesopelagic euphausiids and myctophids are often found in the diets of shelf-break fish. Because the boundary mesopelagic community is distributed worldwide, the high levels of fish biomass often found at shelf-slope breaks could be sustained trophically by deeper, offshore mesopelagic communities, an inverse energy transfer from deep to shallow-water marine ecosystems.     

Life history of the bathyal octopus Pteroctopus tetracirrhus (Mollusca,Cephalopoda) in the Mediterranean Sea

The life cycle of the deep-sea octopus Pteroctopus tetracirrhus was studied from monthly samples obtained throughout the year in different areas of the western Mediterranean (mainly around the Balearic Islands and along the coast of the Iberian Peninsula). A total of 373 individuals (205 females, 168 males) were analyzed; females ranged from 4.5 to 14.0 cm mantle length (ML) and males from 4.5 to 11.5 cm ML. There were few small-sized octopuses (<7 cm ML) in the samples, which might indicate that these individuals inhabit rocky grounds that are not accessible to trawlers or waters deeper than the maximum depth sampled (800 m). The species occurred more frequently around the Balearic Islands than along the Iberian Peninsula as they appeared in 20% and 7%, respectively, of the hauls in these areas. The octopus inhabits the lower continental shelf and upper slope in both areas, primarily between 200 and 500 m depth. Modal lengths were followed from autumn, when recruits were caught by trawlers, to summer, when reproduction took place. Females grew from 8 to 10 cm ML from winter to spring, but this modal size did not increase further in summer; males grew from 7 to 9 cm ML from winter to spring. The total disappearance of large individuals after summer suggests a life cycle lasting a single year. The evolution of the monthly mean sizes showed that the growth was best described by log-linear functions in both sexes. The length at first maturity was clearly higher in females (12 cm ML) than in males (8 cm ML). A total of 30 different prey items, belonging to four major taxonomic groups (crustaceans, osteichthyes, cephalopods and gastropods), were identified in the stomach contents. The diet of the octopus was based on crustaceans and teleosts, which accounted for 75% and 23% of the prey items, respectively. Cephalopods and gastropods were accessory prey as they only represented 1.6% and 0.7%, respectively, of the total. The octopus showed a marked preference for the benthic fish Symphurus nigrescens and the endobenthic crustacean Alpheus glaber. The bathymetric distribution of P. tetracirrhus coincides with those of these two main prey, which suggests that the distribution of the octopus might be strongly linked to its trophic resources.     

Spatial variability in the trophic ecology and biology of the deep-sea shrimp Aristaeomorpha foliacea in the Mediterranean Sea

The trophic ecology, energy and reproductive states of the deep-water shrimp Aristaeomorpha foliacea, widely distributed along the slopes of the Mediterranean Sea Basins, were analysed in eight areas spread along ca. 3000 km in order to identify patterns in the habitat conditions supporting the species. From W to E the areas were situated between the north side of Eivissa (39°12′N, 1°20′E, in the Balearic Basin) and off Mersin, Turkey (36°15′N, 34°19′E, in the Levantine Sea). Trends identified mainly as a function of longitude from west to east were: (i) higher δ¹⁵N, parallel to δ¹⁵N shifts in the top 200 m of the water column for particulate organic N (Pantoja et al., 2002). The δ¹⁵N trend indicates that the deep trophic web, i.e. A. foliacea at 400–600 m, reflects the δ¹⁵N signal of the photic zone; (ii) a similar significant trend of δ¹³C, related with exploitation of pelagic versus benthic resources by A. foliacea in each area (i.e. by local variability of terrigenous inputs via submarine canyons). More depleted δ¹³C was found at mid-longitudes (Tyrrhenian Sea and Sicily Channel) linked to higher consumption of macroplankton prey (Pasiphaea spp., euphausiids and mesopelagic fishes). The feeding intensity (gut fullness, F) and prey diversity (J) of A. foliacea were related, according to generalized linear models, with the temperature and salinity of intermediate waters, variables in turn associated with latitude and longitude. Both F and J were higher in areas with greater shrimp density. The optimal ecological habitat of A. foliacea appears to be located in the Tyrrhenian Sea and the Sicily Channel, where we found the highest F, the greatest trophic diversity and A. foliacea in the best biological condition (i.e. with higher hepato-somatic index, HSI). These are also the areas with the highest densities of A. foliacea. In contrast, in the western Mediterranean Sea (Balearic Basin and the southern Balearic Islands), where A. foliacea has low densities, the shrimp showed generally lower values of trophic indicators and biological condition.     

Geographical variation in metazoan parasites of the deep-sea fish Bathypterois mediterraneus Bauchot, 1962 (Osteichthyes: Ipnopidae) from the Western Mediterranean

This study examines the parasite fauna of Bathypterois mediterraneus, the most common fish below 1500 m in Western Mediterranean waters. Samples were obtained during July 2010 from the continental slope of two different areas (off Catalonia and Balearic Islands) in three different bathymetric strata at depths between 1000 and 2200 m. The parasite fauna of B. mediterraneus included a narrow range of species: Steringophorus cf. dorsolineatum, Scolex pleuronectis, Hysterothylacium aduncum, Anisakis sp. larva 3 type II and Sarcotretes sp. Steringophorus cf. dorsolineatum and H. aduncum were the most predominant parasites. H. aduncum showed significant differences in abundance between depths of 2000–2200 m with 1000–1400 m and 1400–2000 m, irrespective of locality, whereas S. cf. dorsolineatum showed significant differences between the two localities at all depths except for 2000–2200 m. We suggest the possible usefulness of these two parasites as geographical indicators for discriminating discrete stocks of B. mediterraneus in Western Mediterranean waters.     

Deep-sea suprabenthos assemblages (Crustacea) off the Balearic Islands (western Mediterranean): Mesoscale variability in diversity and production

The composition of suprabenthic crustacean assemblages, their diversity, production (P) and production/biomass (P/B) ratios, were analyzed at species level along two transects situated to the north (N) and south (S) of Mallorca (Balearic Islands, western Mediterranean) at depths between 134 m and 760 m, based on a ca. bi-monthly sampling performed between August 2003 and June 2004. Differences with depth and season in assemblage composition and diversity were analyzed as a function of the contrasting environmental features (e.g. water mass dynamics) of the two areas. We identified 187 species (18 decapods, 5 euphausiids, 16 mysids, 76 gammaridean amphipods, 13 hyperiids, 1 caprellid, 21 isopods and 37 cumaceans). Substantial mesoscale variability in the deep-sea suprabenthic assemblages coupled with diversity trends between the N and S transects were found. Seasonality was the most important gradient influencing the dynamics of suprabenthos over the upper (350 m) and middle (650–750 m) slope in the N area. Conversely, the S area appeared to be more stable temporally with depth as the main gradient inducing assemblage differences. Different depth-related patterns were observed both for diversity and P/B. To the north diversity was very low at the shelf-break, increasing on the upper-slope (H′ > 3.00) and then decreasing again on the middle-slope. To the south diversity increased smoothly downward, reaching the highest values on the middle-slope. Regarding productivity, P/B was highest at intermediate depths to the north (over ca. 450–500 m), while to the south highest P/Bs were found deeper (over ca. 600–650 m). The higher P/B at intermediate depths found along N are likely due to higher % of organic matter (OM) in sediments, a product of oceanographic frontal systems. In particular, P/B was higher along N among omnivores and detritus feeders (e.g. Andaniexis mimonectes, Lepechinella manco and combined cumaceans), coupled to enriched OM in sediments, while along S mesoplanktonic carnivores (Rhachotropis spp.) had higher P/Bs. We conclude that on the north slope the influence of frontal systems and more active flow dynamics of different water masses (WIW and LIW) increases natural disturbance in the area, increasing productivity and diversity of suprabenthic peracarids in the Benthic Boundary Layer. Also, species showed a displacement of their average distributions (their Centres of Gravity, CoG) to shallower depths along N, which is another indicator of more favorable habitat conditions for suprabenthos in the 400–500 m range at N.     

Bioaccumulation and trophic transfer of mercury in striped bass (Morone saxatilis) and tautog (Tautoga onitis) from the Narragansett Bay (Rhode Island,USA)

We examined the bioaccumulation and trophic transfer of mercury in two marine finfish species, striped bass (Morone saxatilis) and tautog (Tautoga onitis), collected from the Narragansett Bay (Rhode Island, USA). For each of these target fish, white muscle tissue was analyzed for total mercury (Hg) and results were evaluated relative to fish age, body size, and Hg content of preferred prey. Dietary and stable isotope analysis was also used to elucidate the effect of trophic processes on Hg concentrations in fish. The Hg content of muscle tissue was positively correlated with fish age and length for both species, although striped bass accumulated Hg faster than tautog. Accelerated Hg bioaccumulation in striped bass is consistent with its high trophic level (trophic level = 4.07) and Hg-enriched prey (forage fish and macrocrustaceans; mean Hg content = 0.03 mg Hg kg wet wt^?1). In contrast, tautog maintain a lower trophic status (trophic level = 3.51) and consume prey with lower Hg levels (mussels and crabs; mean Hg content = 0.02 mg Hg kg wet wt^?1). Despite differences in Hg bioaccumulation between target fish, the mean Hg concentration of tautog exceeded levels in striped bass (0.24 and 0.16 mg Hg kg wet wt^?1, respectively) due to a disparity in age-at-catch between sampled groups (mean age of tautog and bass = 11.3 and 4.3 yr, respectively). Taking into account legal minimum catch lengths further revealed that 75.0% of legal-size striped bass (>70.2 cm TL; n = 4) and 44.8% of tautog (>40.6 cm TL; n = 29) had Hg levels beyond the US EPA regulatory threshold of 0.3 mg Hg kg wet wt^?1. Moreover, Hg-length relationships suggest that each target fish meets this threshold near their minimum legal catch length. Our findings reiterate the value of species ecology to improve predictions of fish Hg and permit better management of human contamination by this important dietary source.     

Phytoplankton of the western Arctic in the spring and summer of 2002: Structure and seasonal changes

We analyzed the taxonomic structure and spatial variability of phytoplankton abundance and biomass in the Chukchi and Beaufort Seas during spring and summer seasons of the SBI program. Phytoplankton samples were collected during two surveys from May 10 to June 13 and from July 19 to August 21 of 2002. In May and June, ice cover exceeded 80% over most of the study area and there was no vertical stratification, indicating that the successional state of the phytoplankton corresponded to the end of the winter biological season. The phytoplankton abundance ranged from a few tens to a few thousands of cells per liter, while biomass varied from 0.1 to 3.0 mg C m⁻³. Small areas of high phytoplankton abundance (0.13–1.3×10⁶ cells L⁻¹) and biomass (22–536 mg C m⁻³), dominated by early spring diatoms Pauliella taeniata and Fragilariopsis oceanica in the surface waters, which indicated the beginning of the spring bloom, were observed only in the southeastern part of the Chukchi shelf and off Point Barrow. In July and August summer period, more than a half of the study area had <50% ice cover and the water column was stratified by temperature and salinity. Over the Chukchi shelf and continental slope of the Beaufort Sea, the phytoplankton abundance and biomass were an order of magnitude higher in July–August than in May–June. The taxonomic diversity of algae also increased due to the appearance of late-spring and summer diatoms, dinoflagellates, and coccolithophorids (Emiliania huxleyi). Interestingly, the seasonal differences between phytoplankton abundance and taxonomic composition in the spring and summer periods varied the least over the Chukchi Sea slope and in the deep-water area of the Arctic Ocean. High algae concentrations in summer were located in the lower layers of the euphotic zone, suggesting that the spring bloom on both the Chukchi shelf and in the western part of the Beaufort Sea occurred in late June/early July. In the spring and summer, the microalgal community was characterized by a high abundance of 4–10 μm flagellates, which exceeded the abundance of all other taxonomic groups. In both seasons studied, phytoplankton reached its maximum abundance within restricted areas in the southern part of the Chukchi Sea southwest of Point Hope, in the northern part of the Chukchi shelf between the 50- and 100-m isobaths, on the shelf northwest of Point Barrow, and over the continental slope in the Beaufort Sea. The pronounced spatial difference in the seasonal state was a characteristic feature of the phytoplankton community in the western Arctic.     

Full-depth profiles of prokaryotes,heterotrophic nanoflagellates,and ciliates along a transect from the equatorial to the subarctic central Pacific Ocean

Studies in epipelagic waters report higher heterotrophic microbial biomass in the productive high latitudes than in the oligotrophic low latitudes; however, biogeographical data are scarce in the deep ocean. To examine the hypothesis that the observed latitudinal differences in heterotrophic microbial biomass in the epipelagic zone also occur at depth, abundance and biomass of heterotrophic prokaryotes, nanoflagellates (HNF), and ciliates were determined at depths of 5–5000 m in the central Pacific between August and September of 2005. Heterotrophic microbial biomass increased from the tropical to the subarctic region over the full water column, with latitudinal differences in prokaryotic biomass increasing from 2.3-fold in the epipelagic zone to 4.4-fold in the bathypelagic zone. However, the latitudinal difference in HNF and ciliate biomass decreased with depth. In the mesopelagic zone, the vertical attenuation rate of prokaryotic abundance, which was calculated as the linear regression slope of log-log plot of abundance versus depth, ranged from –0.55 to –1.26 and was more pronounced (steeper slope) in the lower latitudes. In contrast, the vertical attenuation rate of HNF in the mesopelagic zone (–1.06 to –1.27) did not differ with latitude. In the subarctic, the attenuation rate of HNF was 1.7 times steeper than for prokaryotes. These results suggest the accumulation of prokaryotes in the deep subarctic Pacific, possibly due to low grazing pressure. Although the vertical attenuation rate of ciliates was steepest in the bathypelagic zone, HNF abundance did not further decrease at depths below 1000 m, except for at 2000 m where HNF was lowest across the study area. Ciliate abundance ranged 0.3–0.8 cells l^–1 at 4000 m, and were below the detection limit (<0.1 cells l ^–1) at 5000 m. To our knowledge, this study presents the first data for ciliates below 2000 m.     

Distribution of bacterial abundance and cell-specific nucleic acid content in the Northeast Pacific Ocean

We tested the idea that bacterial cells with high nucleic acid content (HNA cells) are the active component of marine bacterioplankton assemblages, while bacteria with low nucleic acid content (LNA cells) are inactive, with a large data set (>1700 discrete samples) based on flow cytometric analysis of bacterioplankton in the Northeast Pacific Ocean off the coast of Oregon and northern California, USA. Samples were collected in the upper 150 m of the water column from the coast to 250 km offshore during 14 cruises from March 2001 to September 2003. During this period, a wide range of trophic states was encountered, from dense diatom blooms (chlorophyll-a concentrations up to 43 μg l⁻¹) at shelf stations during upwelling season (March–September) to lower chlorophyll-a concentrations (0.1–5 μg l⁻¹) during winter (November–February) and at basin stations (>1700 m depth). We found only weakly positive relations of log total bacterial abundance to log chlorophyll-a concentration (as a proxy for availability of organic substrate), and of HNA bacteria as a fraction of total bacteria to log chlorophyll-a. Abundance of HNA and LNA bacteria co-varied positively in all regions, although HNA bacteria were more responsive to high phytoplankton biomass in shelf waters than in slope and basin waters. Since LNA cell abundance in general showed responses similar to those of HNA cell abundance to changes in phytoplankton biomass, our data do not support the hypothesis that HNA cells are the sole active component of marine bacterioplankton.     

Stable abundance,but changing size structure in grenadier fishes (Macrouridae) over a decade (1998–2008) in which deepwater fisheries became regulated

A ten-year time series (1998–2008) from a trawl survey of the continental slope of the NE Atlantic was analyzed to assess temporal variation in the abundance and length frequency of seven species of deepwater grenadier fish. This period coincided (in 2003) with the regulation of deepwater fisheries in this area. None of the species declined in numbers or biomass over the period, and 2 species significantly increased. This suggests that the declines in abundance of these deepwater species following the onset of fishing in the 1970s may now have stabilized, albeit at much lower levels than the virgin biomass. Although two metrics of body size (mean length and maximum length) did not show any evidence for consistent decrease over time, there were significant changes in the overall length–frequency distributions. The species found in shallower depths (500 m) had a greater number of larger individuals in 2008 whereas those found deeper (1500 m) tended to have a greater number of smaller individuals. This suggests the presence of a lagged indirect effect of fishing on species that live beyond the actual depths that fishing takes place.     

Biomass size-spectra of macrobenthic communities in the oxygen minimum zone off Chile

Estimates of macrofaunal secondary production and normalized biomass size-spectra (NBSS) were constructed for macrobenthic communities associated with the oxygen minimum zone (OMZ) in four areas of the continental margin off Chile. The presence of low oxygen conditions in the Humboldt Current System (HCS) off Chile was shown to have important effects on the size structure and secondary production of the benthic communities living in this ecosystem. The distribution of normalized biomass by size was linear (log₂–log₂ scale) at all stations. The slope of the NBSS ranged from −0.481 to −0.908. There were significant differences between the slopes of the NBS-spectra from the stations located in the OMZ (slope = −0.837) and those located outside the OMZ (slope = −0.463) (p < 0.05). The results of this study suggest that low oxygen conditions (<0.5 ml L⁻¹) appear to influence biomass size-spectra, because small organisms are better able to satisfy their metabolic demands. The annual secondary production was higher off central Chile (6.8 g C m⁻² y⁻¹) than off northern Chile (2.02 g C m⁻² y⁻¹) and off southern Chile (0.83 g C m⁻² y⁻¹). A comparison with other studies suggests that secondary production in terms of carbon equivalents was higher than in other upwelling regions.     

Nematode diversity and its relation to the quantity and quality of sedimentary organic matter in the deep Nazaré Canyon,Western Iberian Margin

Samples collected in the deep Nazaré Canyon and at the adjacent slope, during the HERMES RRS Discovery D297 cruise (2005), were analysed for metazoan meiofauna, nematode structure and diversity and its relation to quality and quantity of sedimentary organic material. The amount and quality of organic matter available for direct consumption was much higher in the canyon compared to the slope and positively correlated with high nematode abundances (795–1171 ind. 10 cm^?2) and biomass (93.2–343.5 μg dry weight 10 cm^?2), thus leading to higher standing stocks. Canyon nematode assemblages also showed particular adaptations (e.g. higher trophic complexity, variability of nematode morphology, and presence of opportunistic genera) to canyon conditions, particularly in the deeper sediment layers. The Nazaré Canyon's nematode diversity was slightly lower than that of the adjacent slope and its assemblages were characterised by a higher dominance of certain genera. Still, the canyon contributes considerably to total Western Iberian Margin diversity due to different assemblages present compared to the slope. Furthermore, the harsh conditions in terms of hydrodynamic disturbance and the high organic matter flux are likely to have a negative impact on the establishment of species rich meiobenthic communities, especially in the canyon axis.     

Parameterization of nonlinear shallow water waves over sloping bottoms

Investigation of the bottom slope effects on the nonlinear transformation of irregular waves, which are generated based on JONSWAP spectra, is carried out in a physical wave flume with three slopes (β = 1/15, 1/30, 1/45). The slope effects on the estimation of representative wave height are examined first. To obtain a better estimation of wave height, the slope effect should be considered when slope is larger than 1/30. The nonlinear parameters (bicoherence, skewness and asymmetry) are estimated by using the wavelet-based bispectrum, and the empirical formulae regarding these nonlinear parameters as a function of the local Ursell number are derived based on the present data measured on each slope. The results indicate that the slopes have a negligible effect on the variations of the skewness. The fitted coefficients of the formulae for the other parameters on slope β = 1/15 are clearly different from the results on the slopes β = 1/30 and 1/45, indicating that slope influence on the parameterization cannot be ignored when β > 1/30. Hence, new formulae considering the slope effect are presented. Furthermore, the empirical formulae for the data in surf zone are recommended.     

Vertical changes in abundance,biomass and community structure of copepods down to 3000 m in the southern Bering Sea

Vertical changes in abundance, biomass and community structure of copepods down to 3000 m depth were studied at a single station of the Aleutian Basin of the Bering Sea (53°28′N, 177°00′W, depth 3779 m) on the 14th June 2006. Both abundance and biomass of copepods were greatest near the surface layer and decreased with increase in depth. Abundance and biomass of copepods integrated over 0–3000 m were 1,390,000 inds. m^?2 and 5056 mg C m^?2, respectively. Copepod carcasses occurred throughout the layer, and the carcass:living specimen ratio was the greatest in the oxygen minimum layer (750–100 m, the ratio was 2.3). A total of 72 calanoid copepod species belonging to 34 genera and 15 families occurred in the 0–3000 m water column (Cyclopoida, Harpacticoida and Poecilostomatoida were not identified to species level). Cluster analysis separated calanoid copepod communities into 5 groups (A–E). Each group was separated by depth, and the depth range of each group was at 0–75 m (A), 75–500 m (B), 500–750 m (C), 750–1500 m (D) and 1500–3000 m (E). Copepods were divided into four types based on the feeding pattern: suspension feeders, suspension feeders in diapause, detritivores and carnivores. In terms of abundance the most dominant group was suspension feeders (mainly Cyclopoida) in the epipelagic zone, and detritivores (mainly Poecilostomatoida) were dominant in the meso- and bathypelagic zones. In terms of biomass, suspension feeders in diapause (calanoid copepods Neocalanus spp. and Eucalanus bungii) were the major component (ca. 10–45%), especially in the 250–3000 m depth. These results are compared with the previous studies in the same region and that down to greater depths in the worldwide oceans.     

Distribution of zooplankton biomass at three seamounts in the NE Atlantic

During different seasons of the years 2003–2005 in the NE Atlantic, zooplankton were sampled with a MOCNESS (multiple opening/closing net and environmental sensing system, mesh size 333 μm) above the slopes and summits of Seine, Sedlo and Ampère seamounts and at remote reference sites outside the influence of the seamounts (far field). Wet weights of different zooplankton size classes (<0.5, 0.5–2, >2 cm) were measured. Night and day hauls were analysed in order to detect diel vertical migrations of the zooplankton, as well as a possible trapping effect due to the shallow topography.Biomass concentrations, independent of daytime, season and summit height, were reduced above the summits at all three seamounts compared to the slope and far-field sites. No trapping effect or retention of biomass was apparent above the seamounts. The vertical distribution patterns of the size class <0.5 cm did not differ between night and day hauls at most sites, but indications of diel vertical migrations were found in the larger size fractions. With the exception of gelatinous organisms, zooplankton >0.5 cm were nearly absent above the summits of Seine and Ampère seamounts, but considerable numbers were found above the slopes and at the far-field sites. Possible explanations for the observed distribution patterns of zooplankton biomass and size classes are discussed, including retention and lateral advection due to the hydrography at the seamounts, as well as predation by resident seamount fish.     

Contrasting life-histories,secondary production,and trophic structure of Peracarid assemblages of the bathyal suprabenthos from the Bay of Biscay (NE Atlantic) and the Catalan Sea (NW Mediterranean)

The life-histories and the secondary production of four dominant peracarid crustaceans (the mysids Boreomysis arctica and Parapseudomma calloplura, the amphipod Rhachotropis caeca, and the isopod Ilyarachna longicornis) in bathyal depths of the Bay of Biscay (NE Atlantic; between 383 and 420 m) and the Catalan Sea (Northwestern Mediterranean; between 389 and 1355 m) were established. Both the Atlantic and the Mediterranean populations of the major part of the target-species had two generations/year with mean cohort-production intervals (CPI) ranging from 5.5 mo for Ilyarachna longicornis to 6.3 mo for Parapseudomma calloplura. The Hynes method showed secondary production to vary in the Bay of Biscay between 0.113 mg DW m⁻² yr⁻¹ for I. longirostris and 3.069 mg DW m⁻² yr⁻¹ for P. calloplura, with P/B ratios between 4.57 (I. longirostris) and 7.93 (Boreomysis arctica). In the Catalan Sea, production varied between 0.286 mg DW m⁻² yr⁻¹ for I. longirostris and 1.096 mg DW m⁻² yr⁻¹ for P. calloplura, with P/B between 5.72 (I. longirostris) and 6.66 (P. calloplura). Application of two different empiric models to the whole peracarid assemblage gave similar levels of secondary production in both study areas (between 29.26 and 32.14 mgDWm⁻² yr⁻¹ in the Bay of Biscay; between 26.23 and 26.54 mg DW m⁻² yr⁻¹ in the Catalan Sea). From the analysis of gut contents of 22 species the dominant species in each study area were assigned to two basic trophic levels, detritus feeders and predators. Also, cumulative curves of dominance showed high diversity (low dominance) for peracarid assemblages distributed at mid-bathyal depths (524–693 m) both in the Bay of Biscay off Arcachon and in the Catalan Sea off Barcelona. We also discuss and compare, both within and between areas, how environmental features may explain the observed diversity patterns, the trophic structure, and the production results obtained for the suprabenthos assemblages.     

Bacterioplankton and phytoplankton biomass and production during summer stratification in the northwestern Mediterranean Sea

We examined bacterioplankton biomass and heterotrophic production (BHP) during summer stratification in the northwestern Mediterranean in four successive stratification seasons (June–July of 1993–1996). Values of phytoplankton biomass and primary production were determined simultaneously so that the data sets for autotrophic and heterotrophic microbial plankton could be compared. Three standard stations were set along a transect from Barcelona to the channel between Mallorca and Menorca, representing coastally influenced shelf waters, frontal waters over the slope front, and open sea waters. Conversion factors from ³H-leucine incorporation to BHP were empirically determined and varied between 0.29 and 3.25 kg C mol^-1. Bacterial biomass values were among the lowest found in any marine environment. BHP values (between 0.02 and 2.5 μg C L^-1 d^-1) were larger than those of low nutrient low chlorophyll areas such as the Sargasso Sea and lower than those from high nutrient low chlorophyll areas such as the equatorial Pacific. Growth rates of bacterioplankton were highest at the slope front (0.20 d^-1) and lowest at the open sea station (0.04 d^-1). Phytoplankton growth rates were similar at the three stations (∼0.50 d^-1). Integrated values of bacterioplankton biomass, BHP and bacterial growth rates did not show significant differences among years, but differences between the three stations were clearly significant. Phytoplankton biomass, primary production, and phytoplankton growth rates did not show significant differences either with year or with station. As a consequence the bacterioplankton to phytoplankton biomass (BB/BPHY) and production (BHP/PP) ratios varied from the coastal to the open sea stations. The BB/BPHY ratio was 0.98 at the coast and ∼0.70 at the other two stations. These ratios are similar to those found in other oligotrophic marine environments. The BHP/PP ratio was 0.83 at the coast, 0.36 at the slope and 0.09 at the open sea station. The last value is also similar to values found in other oligotrophic marine environments. Vertical distribution of these ratios was also examined.The comparison of microbial parameters at the three stations indicates a different kind of relationship between bacterioplankton and phytoplankton in oligotrophic open sea waters and in coastal, nutrient-richer waters. According to such parameters and to the values of the BB/BPHY and BHP/PP ratios, open waters in the northwestern Mediterranean (despite their relatively short distance from the shore) were intermediate between the extremely oligotrophic waters of the eastern Mediterranean or the Sargasso Sea and the more productive waters of the equatorial Pacific.