Live (stained) benthic foraminifera from the Rhône prodelta (Gulf of Lion,NW Mediterranean): Environmental controls on a river-dominated shelf

In this paper, we investigate the ecology of live (rose Bengal stained) benthic foraminifera collected at 20 stations ranging from 15 to 100 m depth in the Rhône prodelta (Gulf of Lions, NW Mediterranean). These sites were sampled in September 2006, five months after the Rhône River annual flood. Statistical analyses based on foraminiferal communities (> 150 μm) divide our study area into six main biofacies directly related to environmental conditions. Miliolid species are abundant in the relict prodeltaic lobe which is characterised by sand with low organic matter content. Close to the river mouth, the limited oxygen penetration in the sediment combined with important hydro-sedimentary processes constitute stressful conditions for foraminiferal faunas dominated by opportunistic species (e.g. Leptohalysis scottii). With increasing distance from the river mouth, foraminiferal faunas (e.g. Nonionella turgida, Eggerella scabra) adapted to thrive in sediments enriched in Rhône-derived organic matter under more stable hydro-sedimentary conditions appear. In the distal part of the Rhône River influence, benthic species (e.g. Valvulineria bradyana, Textularia agglutinans) living in fine sediment enriched in both continental and marine organic compounds emerge. At the deepest stations located in the south-eastern part of our study area, benthic foraminiferal faunas (e.g. Bulimina aculeata, Melonis barleeanus, Bigenerina nodosaria) are highly diverse, underlining stable environmental conditions characterised by marine-derived organic matter supplies and relatively deep oxygen penetration depth in the sediment. We also compare foraminiferal faunas sampled in September 2006 with communities sampled in June 2005, one month after the Rhône River annual flood (Mojtahid et al., 2009). This comparison suggests that opportunistic species (e.g. B. aculeata, Cassidulina carinata, V. bradyana) have responded to organic matter inputs related to marine primary production in June 2005.     

Depth zonation and bathymetric trends of deep-sea megafaunal scavengers of the Hawaiian Islands

The deep sea has been shown to exhibit strong depth zonation in species composition and abundance. Examination of these patterns can offer ecological insight into how organisms adapt and respond to changing environmental parameters that co-occur with depth. Here we provide the first tropical study on bathymetric zonation and other depth-related trends (size, abundance, and species richness) spanning shelf to abyssal depths of scavenging megafauna. Baited time-lapse free-vehicle cameras were used to examine the deep-sea benthic and demersal scavenging communities of the Hawaiian Islands, an area for which the biology and ecology have remained poorly studied below 2000 m. Twenty-two deployments ranging in depth from 250 to 4783 m yielded 37 taxa attracted to bait, including the first known occurrence of the family Zoarcidae in the Hawaiian Islands. Cluster analysis of Bray–Curtis similarity of species peak abundance (n_max) revealed four main faunal zones (250–500, 1000, 1500–3000, and ?4000 m) with significant separation (ANOSIM, global R=0.907, p=0.001) between designated depth groups. A major faunal break was identified at the 500–1000 m transition where species turnover was greatest, coinciding with the location of the local oxygen minimum zone. Dominance in species assemblage shifted from decapod crustaceans to teleosts moving from shallow to deeper faunal zones. Significant size differences in total length with depth were found for two of the four fish species examined. A logarithmic decline was observed in scavenger relative abundance with depth. Evidence of interaction between scavenging species was also noted between Synaphobranchus affinis and Neolithodes sp. (competition) and Histiobranchus sp. and aristeid shrimp (predation), suggesting that interactions between scavengers could influence indices of abundance generated from baited camera data.     

Living benthic foraminifera of the Hess Rise and Suiko Seamount,central North Pacific

Rose-Bengal-stained benthic foraminifera in six pilot-core samples and one multicore sample collected from the Hess Rise and Suiko Seamount in August 1994 were studied in order to understand foraminiferal distributions between two areas divided by an oceanic front in the central North Pacific. Samples from the Hess Rise were collected in depths of 2167–3354 m under the warm, saline Kuroshio Extension, while samples from Suiko Seamount came from depths of 1811–1955 m under the cold, less-saline subarctic current. Sediment-trap results for the year prior to our sediment sampling show that organic matter fluxes were about 2.5 times greater at Suiko Seamount than at the Hess Rise. However, the hydrographic structure between 1800 and 3400 m, based on CTD observations, is almost the same at both sites. Temperature decreases from 2.2 to 1.7°C over the depth range of 1800–3400 m, salinity increases from 34.5 to 34.7, and the dissolved oxygen content gradually increases from 1.5 to 3.0 ml l⁻¹. The faunal populations at the Hess Rise are quite different from those at Suiko Seamount. The abundant species at the Hess Rise are Epistominella exigua, Brizalina pacifica, Fursenkoina cedrosensis and Alabaminella weddellensis. These species characteristically inhabit phytodetrital aggregates deposited on an oligotrophic seafloor. The populations at Suiko Seamount are dominated by Triloculina frigida, Lagenammina cf. arenulata, Reophax subfusiformis, and Reophax scorpiurus. The reason for differences between these populations is unclear. However, the typical phytodetritus-dwelling species E. exigua is dominant at the Hess Rise, which is located in a subtropical area that has a pulsed supply of settling organic matter in the spring. On the other hand, E. exigua is rare at Suiko Seamount, a subarctic site where there are more stable and greater fluxes of organic matter in summer and autumn. Occurrences of this species may be related to the seasonally short supply of organic matter that reaches the seafloor in the oceanic North Pacific.     

Tracing seafloor methane emissions with benthic foraminifera: Results from the Ana submarine landslide (Eivissa Channel,Western Mediterranean Sea)

The hypothesis that benthic foraminifera are useful proxies of local methane emissions from the seafloor has been verified on sediment core KS16 from the headwall of the Ana submarine landslide in the Eivissa Channel, Western Mediterranean Sea. The core MS312 from a nearby location with no known methane emissions is utilised as control. The core was analysed for biostratigraphy, benthic foraminiferal assemblages, Hyalinea balthica and Uvigerina peregrina carbon and oxygen stable isotope composition, and sedimentary structures. The upper part of the core records post-landslide deglacial and Holocene normal marine hemipelagic sediments with highly abundant benthic foraminifera species that are typical of outer neritic to upper bathyal environment. In this interval, the δ¹³C composition of benthic foraminifera indicates normal marine environment analogous to those found in the control core. Below the sedimentary hiatus caused by the emplacement of the slide, the foraminiferal assemblages are characterised by lower density and higher Shannon Index. Markedly negative δ¹³C shifts in benthic foraminifera are attributed to the release of methane through the seabed. The mean values of the ¹³C anomaly in U. peregrina are ? 0.951 ± 0.208 in the pre-landslide sediments, and ? 0.269 ± 0.152 in post-slide reworked sediments deposited immediately above the hiatus. The δ¹³C anomaly in Hyalinea balthica is ? 2.497 ± 0.080 and ? 2.153 ± 0.087, respectively. To discard the diagenetic effects on the δ¹³C anomaly, which could have been induced by Ca–Mg replacement and authigenic carbonate overgrowth on foraminifera tests, a benthic foraminifera subsample has been treated following an oxidative and reductive cleaning protocol. The cleaning has resulted, only in some cases, in a slight reduction of the anomaly by 0.95% for δ¹³C and < 0.80% for δ¹⁸O. Therefore, the first conclusion is that the diagenetic alteration is minor and it does not alter significantly the overall carbon isotopic anomaly in the core. Consequently, the pre-landslide sediments have been subject to pervasive methane emissions during a time interval of several thousand years. Methane emissions continued during and immediately after the occurrence of Ana Slide at about 61.5 ka. Subsequently, methane emissions decreased and definitely ceased during the last deglaciation and the Holocene.     

Benthic foraminiferal successions across Late Quaternary Mediterranean sapropels

Benthic foraminiferal records across Late Quaternary Mediterranean sapropels S6, S5 and S1 are reviewed and re-considered in the light of recent advances in foraminiferal ecology. It is suggested that the main factor controlling the foraminiferal successions that occur immediately before and after sapropel deposition is the amount of time involved in the onset of anoxic conditions and in the re-oxygenation of the benthic environment. Faunas dominated by deep infaunal taxa such as Globobulimina and Chilostomella reflect a very gradual decrease or increase of bottom water oxygenation. Post-sapropel faunas dominated by small biconvex, trochospiral taxa, which are inferred to have a much more opportunistic life strategy, are typical of a very rapid re-oxygenation of the benthic environment after sapropel deposition. The time involved in re-oxygenation depends upon the mechanism causing sapropel formation. The faunal succession found above sapropel S5 suggests a basin with a strong density gradient in the upper part of the water column during sapropel deposition, followed by a rapid turnover of the following column and quick re-oxygenation of the benthic environment. In contrast, the faunal succession above sapropel S6 suggests a water column with a more gradual density gradient deeper in the basin, a progressive deepening of the halocline, and a very slow increase of bottom water oxygenation.     

Dispersal,survival and delayed growth of benthic foraminiferal propagules

New data support our previously published propagule dispersal hypothesis and show that propagules of some benthic foraminiferal species can survive for two years before growth commences. Following exposure to simulated shallow-water conditions, shallow-water species of benthic foraminifera appeared and grew in large numbers (commonly >100 ind/12 ml sediment) in the <32 µm-size sediment fraction collected from 320 m water depth in the Skagerrak basin (North Sea). None of the shallow-water species that grew abundantly (Planorbulina mediterranensis, Morulaeplecta bulbosa, Bolivina pseudoplicata, Cuneata arctica, Eggerelloides scaber, Gavelinopsis praegeri) seem to grow or reproduce at or in the vicinity of the sampling site. Consequently, they must have been transported there as <32 µm-sized individuals. Their sudden appearance when exposed to shallow-water conditions suggests that they had been transported to the sampling site as propagules and that they could survive in the sediments until conditions became suitable for growth and, for some, reproduction. The lack of agglutination on the proloculi of the agglutinated taxa that appeared in the growth-chambers may enhance their passive transport via currents and, thereby, dispersal. Of all the indigenous foraminiferal species that occur at the sampling site, only Textularia earlandi and Bolivinellina pseudopunctata continued to grow and reproduce when transferred from bathyal (320 m) to simulated shallow-water (0 m) conditions. The former is considered a highly opportunistic species. According to the literature, most of the morphospecies which grew in the experiments are cosmopolitan. Our results indicate substantial inter-specific differences in dispersal potential and support previous suggestions that among free-living species, some serial forms have the potential for long-distance dispersal. Still, oceanographic, physical and ecological boundaries and barriers constrain the distribution of most species. In addition to benthic foraminifera, Gromia spp. (rhizarian protists related to the foraminifera) grew in >60% of the experimental growth-chambers.     

Regional variations in the fluxes of foraminifera carbonate,coccolithophorid carbonate and biogenic opal in the northern Indian Ocean

Mass fluxes of diatom opal, planktonic foraminifera carbonate and coccolithophorid carbonate were measured with time-series sediment traps at six sites in the Arabian Sea, Bay of Bengal and Equatorial Indian Ocean (EIOT). The above fluxes were related to regional variations in salinity, temperature and nutrient distribution. Annual fluxes of diatom opal range between 3 and 28 g m⁻² yr⁻¹, while planktonic foraminifera carbonate fluxes range between 6 and 23 g m⁻² yr⁻¹ and coccolithophorid carbonate fluxes range between 4 and 24 g m⁻² yr⁻¹. Annual planktonic foraminifera carbonate to coccolithophorid carbonate ratios range between 0.8 and 2.2 and coccolithophorid carbonate to diatom opal ratios range between 0.5 and 3.3.In the western Arabian Sea, coccolithophorids are the major contributors to biogenic flux during periods of low nutrient concentrations. Coccolithophorid carbonate fluxes decrease and planktonic foraminiferal carbonate and diatom opal fluxes increase when nutrient-rich upwelled waters are advected over the trap site. In the oligotropic eastern Arabian Sea, coccolithophorid carbonate fluxes are high throughout the year. Planktonic foraminiferal carbonate fluxes are the major contributors to biogenic flux in the EIOT. In the northern and central Bay of Bengal, when surface salinity values drop sharply during the SW monsoon, there is a drastic reduction in planktonic foraminiferal carbonate fluxes, but coccolithophorid carbonate and diatom opal fluxes remain steady or continue to increase. Distinctly higher annual molar Si_bio/C_inorg (>1) and C_org/C_inorg (>1.5) ratios are observed in the northern and central Bay of Bengal mainly due to lower foraminiferal carbonate production as a result of sharp salinity variations. We can thus infer that the enhanced freshwater supply from rivers should increase oceanic CO₂ uptake. Its silicate supply favours the production of diatoms while the salinity drop produces conditions unfavourable for most planktonic foraminifera species.     

Deep-sea macrofaunal assemblages within the Benthic Boundary Layer of the Cap-Ferret Canyon (Bay of Biscay,NE Atlantic)

The Benthic Boundary Layer (BBL) assemblages from the Cap-Ferret Canyon (Bay of Biscay) were quantitatively sampled at two sites located within its main channel near mooring deployments (Mooring Sites MS 1: ca. 2400 m; MS 2: ca. 3000 m) with a suprabenthic sled equipped with four nets fishing at different heights above the bottom. The macrofaunal abundance above the sea-floor was mainly represented by Isopoda (42.2%), Amphipoda (19.0%), Euphausiacea (17.3%), Cumacea (13.5%), Mysidacea (2.8%) and Tanaidacea (2.6%). At both sampling sites, the highest total densities were generally recorded in the immediate vicinity of the sea floor (10–40 cm water layer), and a drastic decrease occurred higher in the BBL community. The BBL assemblages from the two sampling sites were similar in their faunal composition (major taxa), and their mean density estimates were not statistically different (MS 1 : 525.3 ind. 100 m⁻²; MS 2 : 283.3 ind. m⁻²) although the recorded values during each cruise were always lower at the deeper site. The BBL macrofauna abundance showed obvious temporal fluctuations at both sites, probably linked with a seasonal organic input from the euphotic zone (vertical flux) via phytodetritus deposition on the sea bottom.     

The ecology and distribution of benthic foraminifera at the Håkon Mosby mud volcano (SW Barents Sea slope)

To investigate a possible influence of submarine methane seepage on benthic foraminiferal communities, Rose Bengal stained (“live”) and empty tests of benthic foraminifera were studied from the sediment surface down to 15 cm sub-bottom depth of 12 sites at the Håkon Mosby mud volcano (HMMV). In addition, one reference site well away from the seep sites, but from similar water depths and the same general hydrographic setting was occupied for comparison. The HMMV is located at 1265 m water depth on the SW Barents Sea continental slope. Distinct living foraminiferal associations at the HMMV are linked to specific sedimentary, microbial, and macrofaunal habitats. In the center of the crater, and in crater areas completely covered by bacterial mats, Cassidulina reniforme is the only, albeit rare, living species. Below the top few millimeters, sediments are anoxic and devoid of living specimens. At the rim of the mud volcano, at sites densely populated by pogonophoran tube worms, three benthic foraminiferal associations are found; (i) a Fontbotia wuellerstorfi–Lobatula lobatula association living attached to the upper parts of pogonophoran tubes, which protrude into oxic water, (ii) a diverse Cassidulina neoteretis association populating dysoxic sediments of the surface centimeter, and (iii) a species-poor Bolivina pseudopunctata association colonizing the subsurface sediments down to four centimeters. Generally, we did not find endemic or seep indicative species or associations at the HMMV. However, the HMMV live faunas dominated by B. pseudopunctata are not found at the reference site nor are they described from comparable water depths and environments without gas seepages from the Norwegian-Greenland Seas.In the center and outer rim of the mud volcano, a C. neoteretis–Reophax guttifer dead association, similar to the one at the reference site, characterizes an assemblage of strongly corroded and partly displaced tests. At bacterial mat sites, a C. reniforme dead association corresponds to the living one. Thus both the living and the dead associations are indicative of a specific bacterial mat environment at the HMMV.     

Blake Ridge methane seeps: characterization of a soft-sediment,chemosynthetically based ecosystem

Observations from the first submersible reconnaissance of the Blake Ridge Diapir provide the geological and ecological contexts for chemosynthetic communities established in close association with methane seeps. The seeps mark the loci of focused venting of methane from the gas hydrate reservoir, and, in one location (Hole 996D of the Ocean Drilling Program), methane emitted at the seafloor was observed forming gas hydrate on the underside of a carbonate overhang. Megafaunal elements of a chemosynthetically based community mapped onto dive tracks provide a preliminary overview of faunal distributions and habitat heterogeneity. Dense mussel beds were prominent and covered 20×20 m areas. The nearly non-overlapping distributions of mussels and clams indicate that there may be local (meter-scale) variations in fluid flux and chemistry within the seep site. Preliminary evidence suggests that the mussels are host to two symbiont types (sulfide-oxidizing thiotrophs and methanotrophs), while the clams derive their nutrition only from thiotrophic bacteria. Invertebrate biomass is dominated by mussels (Bathymodiolus heckerae) that reach lengths of up to 364 mm and, to a lesser extent, by small (22 mm length) vesicomyid clams (Vesicomya cf. venusta). Taking into account biomass distributions among taxa, symbiont characteristics of the bivalves, and stable-isotope analyses, the relative importance of methanotrophic vs thiotrophic bacteria in the overall nutrition of the invertebrate assemblage is on the order of 60% vs 40% (3:2).     

An octo-generator for energy harvesting based on the piezoelectric effect

A renewable energy harvester using the piezoelectric effect is developed for the ocean tidal and wind flow. The harvester is made of connected driving blades to an octo-generator, which has a rotator with n blades and a stator attached by eight mass-spring-piston-cylinder-piezoelectricity devices. The resonance and force magnification are utilized to increase the power output of the harvester. A corresponding mathematical model is developed to calculate the root mean square of the generated electric power. The simulation results indicate that the generated power is largely enhanced when the near-resonant condition is established. The power increases with increases in the magnetic flux density, the large-to-small diameter ratio of the cylinder, the size of magnetic bar face, and decreases in the gap between two magnetic faces and the size of the piezoelectric bar face. A generated power of 5 kW is realized by the harvester working under an ocean tidal speed, V = 1.75 m/s, and its geometric and material properties of driving length L = 7.5 m, spring constant k_v = 65000 N/m, gap between the two magnets s = 0.0015 m, large to small diameter ratio of the cylinder z = 6, and magnetic flux density Br = 1.45 T.     

Late Holocene paleoceanography and climate variability over the Mackenzie Slope (Beaufort Sea,Canadian Arctic)

Late Holocene paleoceanography and climate variability of the Southeastern Beaufort Sea (Canadian Arctic) have been investigated on the basis of sedimentary cores collected over the Mackenzie Slope. Piston, trigger and box cores were sampled at station 803 in 2004 aboard the CCGS Amundsen at 218 m water depth. The chronology of the piston core is constrained by 4 AMS-¹⁴C dates, as the sedimentation rate in the box core is assessed from ²¹⁰Pb data. We obtain a continuous composite sequence covering the last 4600 years, with a sedimentation rate of ~ 140 cm.kyr^? 1. Transfer functions (modern analogue technique) based on dinoflagellate cyst (dinocyst) assemblages were used to reconstruct the evolution of sea-surface conditions over the time period covered by the cores.Palynological data reveal that dinocyst assemblages are dominated by Operculodinium centrocarpum sensu lato (mean of 43.3%) throughout the core, with the accompanying taxa Brigantedinium spp. (19.6%), Islandinium minutum (15.6%) and cysts of Pentapharsodinium dalei (13.7%). Four zones have been established on the basis of dinocyst relative abundances. Dinocyst assemblage zone 1 (D1), from 4600 to 2700 cal years BP, is dominated by O. centrocarpum (mean of 49.0%). In zone D2 (2700–1500 cal years BP), the relative abundances of O. centrocarpum decrease (34.4%) in favour of the opportunistic, heterotrophic taxa Brigantedinium spp. (28.8%) and cysts of Polykrikos sp. var. arctic/quadratus (2.8%). Dinocyst zone D3 (1500–30 cal years BP or 450–1920 AD) is characterised by the high relative abundance of the peridinioid taxa I. minutum (19.9%). The last zone (D4), spanning from 1920 to 2004 AD, is again dominated by O. centrocarpum (44.5%), and shows low relative abundances of Brigantedinium spp. and cf. Echinidinium karaense.Quantitative reconstructions of past sea-surface parameters (August sea-surface temperature: SST, August sea-surface salinity: SSS, and duration of sea-ice cover) indicate relatively stable conditions over the last 4.6 kyr, with episodic cooling events (SST of ~ 1.5 °C below the modern value of 6 °C) that took place between 700 and 1820 AD. We associate the last and the longest of these cooling events (1560–1820 AD) with the Little Ice Age. Reconstructed SSS shows decadal oscillations since 1920 AD that we tentatively associate with the accumulation of freshwater by the Beaufort Gyre and the subsequent Great Salinity Anomalies. Our data suggest that similar salinity anomalies could have occurred ca. 1860 and 1790 AD.Stable isotopic data show a slight increase in δ¹³C values (from ~?27.1‰ at the base to ~?25.8‰ at the top) over the last 4.6 kyr that we associate with the gradual increase in atmospheric CO₂ concentration as recorded by Antarctic ice cores. Variations in the δ¹⁵N profile suggest variations in Pacific water influence from 4600 to ~ 1300 cal years BP, associated with centennial scale shifts of the Arctic Oscillation phases.     

How much data is enough? The importance of morphological sampling interval and duration for calibration of empirical shoreline models

The ability to robustly predict future shoreline position under the influence of changing waves and sea-level rise is a key challenge to scientists and engineers alike. While extrapolating a linear trend out in time is a common baseline approach, the recent development of a number of empirical shoreline models allows the prediction of storm and annual-scale variability as well. The largest constraint in applying these models is the availability of high quality, adequate duration data sets in order to calibrate model free parameters. This contribution outlines several such models and discusses the monitoring programs required to calibrate and hindcast shoreline change from 1 to 10 years at two distinct beach types: a storm-dominated site and the second exhibiting a large seasonal variability. The seasonally-dominated site required longer data sets but was less sensitive to sampling interval, while the storm-dominated site converged on shorter, more frequently sampled data sets. In general, calibration based on a single year of observed shorelines resulted in a large range of model skill and was not considered robust. Monitoring programs of at least two years, with shorelines sampled at dt ≤ 30 days were sufficient to determine initial estimates of calibration coefficients and hindcast short-term (1–5 years) shoreline variability. In the presence of unresolved model processes and noise, hindcasting longer (5 + years) data sets required longer (5 + years) calibration data sets, particularly when sampling intervals exceeded 60 days.     

Analysis of a PAH-degrading bacterial population in subsurface sediments on the Mid-Atlantic Ridge

Little is known about the types and concentrations of polycyclic aromatic hydrocarbons (PAHs) existing in the deep-sea subsurface environment, which is believed to be cold, oligothrophic and of high static pressure. PAHs in the upper layers of the water column are unavoidably subjected to degradation while they are deposited to the sea floor and become embedded in the deep-sea sediment. In this report, a high concentration of PAHs was discovered in the sediment 2.7 m beneath the bottom surface at a water depth of 3962 m on the Mid-Atlantic Ridge (MAR). The total concentration of PAHs was 445 ng (g dry wt sediment)^-1. Among the seven detected PAHs, the concentrations of phenanthrene (222 ng g^-1) and fluorene (79 ng g^-1) were relatively high. In addition, PAH-degrading bacteria were found within the sediments. As in a previously detected site on the MAR, in the PAH-enriched region of this site, a bacterium of the genus Cycloclasticus was found to be the predominant isolate detected by PCR-DGGE analysis. In addition, bacteria of the Halomonas, Marinobacter, Alcanivorax, Thalassospira and Maricaulis genera, were also included in the PAH-degrading community. In summary, a high concentration of PAHs was detected in the subsurface of the deep-sea sediment, and once again, the Cycloclasticus bacterium was confirmed to be a ubiquitous marine PAH degrader even in the subsurface marine environment. Considering the abundance of PAHs therein, biodegradation is thus thought to be inactive, probably because of the low temperature, limited oxygen and/or limited nutrients.     

Meiofaunal distributions on the Peru margin:: relationship to oxygen and organic matter availability

A quantitative study of metazoan meiofauna was carried out on bathyal sediments (305, 562, 830 and 1210 m) along a transect within and beneath the oxygen minimum zone (OMZ) in the southeastern Pacific off Callao, Peru (12°S). Meiobenthos densities ranged from 1517 (upper slope, middle of OMZ) to 440–548 ind. 10 cm⁻² (lower slope stations, beneath the OMZ). Nematodes were the numerically dominant meiofaunal taxon at every station, followed by copepods and nauplii. Increasing bottom-water oxygen concentration and decreasing organic matter availability downslope were correlated with observed changes in meiofaunal abundance. The 300-m site, located in the middle of the OMZ, differed significantly in meiofaunal abundance, dominance, and in vertical distribution pattern from the deeper sites. At 305 m, nematodes amounted to over 99% of total meiofauna; about 70% of nematodes were found in the 2–5 cm interval. At the deeper sites, about 50% were restricted to the top 1 cm. The importance of copepods and nauplii increased consistently with depth, reaching ∼12% of the total meiofauna at the deepest site. The observation of high nematode abundances at oxygen concentrations <0.02 ml l⁻¹ supports the hypothesis that densities are enhanced by an indirect positive effect of low oxygen involving (a) reduction of predators and competitors and (b) preservation of organic matter leading to high food availability and quality. Food input and quality, represented here by chloroplastic pigment equivalents (CPE) and sedimentary labile organic compounds (protein, carbohydrates and lipids), were strongly, positively correlated with nematode abundance. By way of contrast, oxygen exhibited a strong negative correlation, overriding food availability, with abundance of other meiofauna such as copepods and nauplii. These taxa were absent at the 300-m site. The high correlation of labile organic matter (C-LOM, sum of carbon contents in lipids, proteins and carbohydrates) with CPE (Pearson's r=0.99, p<0.01) suggests that most of the sedimentary organic material sampled was of phytodetrital origin. The fraction of sediment organic carbon potentially available to benthic heterotrophs, measured as C-LOM/Total organic carbon, was on average 17% at all stations. Thus, a residual, refractory fraction, constitutes the major portion of organic matter at the studied bathyal sites.     

Comparison of benthic foraminifera inside and outside a sulphur-oxidizing bacterial mat from the present oxygen-minimum zone off Pakistan (NE Arabian Sea)

Assemblages of live (Rose-Bengal-stained) and dead benthic foraminifera and stable oxygen and carbon isotopic composition of live benthic foraminifera were studied in and outside a bacterial mat composed of the large sulphur-oxidizing bacteria Thioploca and Beggiatoa from the oxygen-minimum zone off Pakistan (NE Arabian Sea). Two cores from the same Multicorer retrieved a bacterial mat and ambient sediment. The dominant species (Globobulimina affinis, G. turgida, Bolivina pacifica, B. pseudopunctata, Uvigerina peregrina and Buliminella tenuata) in both cores are characteristic for dysoxic oxygen minimum zone conditions. The most significant difference between the two cores is the reduced number of stained benthic foraminifera (SBF) in the top 0.5 cm of the bacterial mat. Faunal densities of stained species are more than four times higher in the sediment surface sample (0–0.5 cm) outside the bacterial mat, at a distance of only 1.5 m. All stained species, however, observed outside the Beggiatoa/Thioploca mat were also observed in the core with the mat. Two species, Virgulinella fragilis and Bolivina dilatata, occur exclusively in the core with the bacterial mat. The diversity within the bacterial mat core is thus slightly higher than outside. Furthermore, the abundances of the species Buliminella morgani, B. tenuata and Alliatina primitiva are substantially higher in the bacterial mat than outside. Globobuliminids, on the other hand, seem to prefer the conditions outside the bacterial mat and are five times more frequent in the core taken outside the bacterial mat. Benthic foraminifers inhabit a broader microhabitat range outside the bacterial mat (∼5 cm) than within (3.5 cm). A marked decrease in SBF abundances was observed at the level of a black sulphur-rich layer which is interpreted to mark the shallow redox front below the bacterial mat. Stable carbon isotope analyses on live benthic foraminifera do not support a relation of the investigated Beggiatoa/Thioploca mat to a constant or seasonal seepage of methane at the continental slope off Pakistan. Surprisingly, however, stable oxygen isotope values of many species and especially of U. peregrina decrease with depth, which calls into question the suitability of U. peregrina as a recorder of bottom-water δ¹⁸O.     

Population dynamics and secondary production of the cockle Cerastoderma edule: A comparison between Merja Zerga (Moroccan Atlantic Coast) and Arcachon Bay (French Atlantic Coast)

Cockle (Cerastoderma edule) population dynamics were studied at the southern limit of the distribution of this marine bivalve in Merja Zerga, Morocco. Parameters such as growth, mortality, and production were compared with those of a population at Arcachon Bay (France) a site in the center of the cockle's range. At each sampling period between two and three cohorts were simultaneously observed at each site and the average total abundance was usually higher at Merja Zerga. Recruitment occurred at both sites in spring when temperature rose above 19 °C, independently of the month. In Merja Zerga, winter recruitment was also observed at one occasion, following high sediment disturbance. The first year (2005–06) at Merja Zerga, the mortality rate was close to nil for juveniles and was Z = 1.5 yr^? 1 for adults, providing a high production (64 g dry weight m^? 2 yr^? 1). At Arcachon during the same period, the juvenile mortality rate was Z = 10.9 yr^? 1, the adult mortality rate was 3.4 yr^? 1 and production was 26 gDW m^?2 yr^? 1. The second year (2006–07), mortality after recruitment was much higher (Z = 8.6 yr^? 1, for juveniles) and similar to what was observed at Arcachon (Z = 8.4 yr^? 1). Mortality rate of adults was higher at Merja Zerga (Z = 3.0 yr^? 1) than at Arcachon (Z = 1.5 yr^? 1). Production was lower at Arcachon than at Merja Zerga although growth performances were higher at Arcachon. The higher growth performance at Arcachon (Φ′ = 3.3) was mainly due to high asymptotic length (L_∞ = 38 mm) and was related to low intraspecific competition compared to Merja Zerga where cockle abundance was higher (Φ′ = 3.1, L_∞ = 31 mm). P/B was low in both sites and slightly higher at Arcachon (1.1–1.5 against 1.0–1.1 yr^? 1). At Arcachon, recruitment was correlated with temperature, a peak occurring when temperature rose above 19 °C (June–July). At Merja Zerga, recruitment was already 2–3 months earlier but was not significantly correlated to temperature.This study showed that population dynamics of cockles at the southern limit of this distribution fell in the range of what was observed elsewhere in the North-Eastern Atlantic coast. Most factors that were involved in population regulation (intraspecific competition, predation and sediment dynamics) were not strictly dependent on latitude. The direct role of temperature (latitude dependent factor) was not obvious. Variation in temperature could explain the recruitment delay between Arcachon and Merja Zerga and the low maximum shell length at Merja Zerga.     

Planktic foraminiferal production stimulated by chlorophyll redistribution and entrainment of nutrients

During September and October 1996 planktic foraminifers and pteropods were sampled from the upper 2500 m of the water column in the BIOTRANS area (47°N, 20°W), eastern North Atlantic, as part of the JGOFS program. Hydrography, chlorophyll fluorescence, and nutrient content were recorded at high spatial and temporal resolution providing detailed information about the transition time between summer and fall. At the beginning of the cruise a shallow pycnocline was present and oligotrophic conditions prevailed. Over the course of the cruise, the mixed layer depth increased and surface water temperature decreased by 1.5°C. Both chlorophyll-a dispersed in the upper 50 m by vertical mixing and chlorophyll-a concentrations at the sea surface increased. The nitracline shoaled and nutrient enriched waters were entrained into the mixed layer. Planktic foraminifers and pteropods closely reflected the changes in the hydrography by increased growth rates and changes in species composition. Three main groups of planktic foraminiferal species were recognized: (1) a temperate and low-productivity group dominated by Neogloboquadrina incompta characterized the shallow mixed layer depths. (2) A temperate and high-productivity group dominated by Globigerina bulloides characterized the period with wind-induced dispersal of chlorophyll-a and entrainment of nutrient-enriched waters. (3) A warm water group containing Globigerinoides sacculifer, Orbulina universa, Globigerinoides ruber (white), and Globigerinella siphonifera was most common during the first days of sampling. Synchronous with the hydrographic change from summer to fall, planktic foraminiferal and pteropod growth was stimulated by redistribution of chlorophyll-a and entrainment of nutrient-enriched waters into the mixed layer. In addition, the seasonal change in the eastern North Atlantic resulted in a transition of the epipelagic faunal composition and an increased calcareous particle flux, which could be used to trace seasonality in fossil assemblages and allow for better paleoceanographic interpretation of the boreal Atlantic.     

Effects of mesoscale phytoplankton variability on the copepods Neocalanus flemingeri and N. plumchrus in the coastal Gulf of Alaska

The copepods Neocalanus flemingeri and N. plumchrus are major components of the mesozooplankton on the shelf of the Gulf of Alaska, where they feed, grow and develop during April–June, the period encompassing the spring phytoplankton bloom. Satellite imagery indicates high mesoscale variability in phytoplankton concentration during this time. Because copepod ingestion is related to food concentration, we hypothesized that phytoplankton ingestion by N. flemingeri and N. plumchrus would vary in response to mesoscale variability of phytoplankton. We proposed that copepods on the inner shelf, where the phytoplankton bloom is most pronounced, would be larger and have more lipid stores than animals collected from the outer shelf, where phytoplankton concentrations are typically low. Shipboard feeding experiments with both copepods were done in spring of 2001 and 2003 using natural water as food medium. Chlorophyll concentration ranged widely, between 0.32 and 11.44 μg l⁻¹ and ingestion rates varied accordingly, between 6.0 and 627.0 ng chl cop⁻¹ d⁻¹. At chlorophyll concentrations<0.50 μg l⁻¹, ingestion is always low, <40 ng cop⁻¹ d⁻¹. Intermediate ingestion rates were observed at chlorophyll concentrations between 0.5 and 1.5 μg l⁻¹, and maximum rates at chlorophyll concentrations>1.5 μg l⁻¹. Application of these feeding rates to the phytoplankton distribution on the shelf allowed locations and time periods of low, intermediate and high daily feeding to be calculated for 2001 and 2003. A detailed cross-shelf survey of body size and lipid store in these copepods, however, indicated they were indistinguishable regardless of collection site. Although the daily ingestion of phytoplankton by N. flemingeri and N. plumchrus varied widely because of mesoscale variability in phytoplankton, these daily differences did not result in differences in final body size or lipid storage of these copepods. These copepods efficiently dealt with small and mesoscale variations in their food environment such that mesoscale structure in phytoplankton did not affect their final body size.     

Field and laboratory studies on the effect of particle size and composition on optical backscattering measurements in hydrothermal plumes

We describe the performance of an inexpensive but highly sensitive light backscattering sensor (LBSS) suitable for use in deep-sea waters, where particle concentrations are typically <∼0.1 mg/l. Laboratory calibrations using aluminosilicate particles and latex spheres show that the concentration-normalized backscattering of the LBSS, K_bs, is greatest for particles with a diameter close to the wavelength of light emitted by the LBSS (0.88 μm), declining by more than a factor of five for particles <0.1 μm or >10 μm. Field studies indicate that in hydrothermal plumes dominated by fine-grained metal precipitates the 95% confidence interval for predicting mass concentration with an LBSS ranges from ±0.004 mg/l for 0.03 mg/l suspensions to ±0.008 mg/l at 0.12 mg/l. Comparisons among a group of 19 LBSSs found between-instrument variability to reach 70%, but normalizing the raw data using a laboratory calibration procedure reduced the maximum variability to ∼8%.