Dynamics at an elongated,intermediate depth seamount in the North Atlantic (Sedlo Seamount, 40°20′N, 26°40′W)

Observations from a five-mooring array deployed in the vicinity of Sedlo Seamount over a 4-month period, together with supporting hydrographic and underway ADCP measurements, are described. Sedlo Seamount is an elongated, intermediate depth seamount with three separate peaks, rising from 2200 m water depth to summit peaks between 950 and 780 m depth, located at 40°20′N, 26°40W. Currents measured in depth range 750 and 820 m – the layer close to the summit depth of the shallowest southeast peak – showed a mean anti-cyclonic flow around the seamount, with residual current velocities of 2–5 cm s⁻¹. Significant mesoscale variability was present at this level, and this is attributed to the weak and variable background impinging flow. Stronger, more persistent currents were found at the summit mooring as a result of tidal rectification and some weak amplification. Below 1300 m, currents were extremely weak, even close to the seabed. Time series of relative vorticity for the depth layer 750–820 m showed persistent anti-cyclonic vorticity except for two periods of cyclonic vorticity. A mean relative vorticity of −0.06f (f=the local Coriolis frequency) was calculated from a triangle of current meters located at the flanks of the seamount. Modelling results confirmed that anti-cyclonic flow above the seamount was likely due to Taylor Cone generation driven by a combination of steady impinging and tidally rectified flow. The closed circulation pattern over the seamount was found to extend to ∼150 m above the summit level, consistent with simple idealised theory and the supporting hydrographic observations. At shallower depths (<500 m) model simulations predicted a predominantly cyclonic recirculation most likely controlled by topographic steering along the zonal axis of the seamount. There was some indication of flow reversal at these depths from Acoustic Doppler Current Profiler (ADCP) measurements carried out at one hydrographic survey. The model results were in good agreement with observations at the seamount summit, but were unable to reproduce the mesoscale variability patterns recorded in shallower layers. Kinetic energy patterns derived from the model revealed high variability in the oceanic far field downstream of the seamount summit probably as a result of complex flow interaction along the chain of seamount peaks. Possible impacts of the flow dynamics on the biological functioning at Sedlo Seamount and its surroundings are discussed.     

Factors influencing particulate lipid production in the East Atlantic Ocean

Extensive analyses of particulate lipids and lipid classes were conducted to gain insight into lipid production and related factors along the biogeochemical provinces of the Eastern Atlantic Ocean. Data are supported by particulate organic carbon (POC), chlorophyll a (Chl a), phaeopigments, Chl a concentrations and carbon content of eukaryotic micro-, nano- and picophytoplankton, including cell abundances for the latter two and for cyanobacteria and prokaryotic heterotrophs. We focused on the productive ocean surface (2 m depth and deep Chl a maximum (DCM)). Samples from the deep ocean provided information about the relative reactivity and preservation potential of particular lipid classes. Surface and DCM particulate lipid concentrations (3.5–29.4 μg L⁻¹) were higher than in samples from deep waters (3.2–9.3 μg L⁻¹) where an increased contribution to the POC pool was observed. The highest lipid concentrations were measured in high latitude temperate waters and in the North Atlantic Tropical Gyral Province (13–25°N). Factors responsible for the enhanced lipid synthesis in the eastern Atlantic appeared to be phytoplankton size (micro, nano, pico) and the low nutrient status with microphytoplankton having the most expressed influence in the surface and eukaryotic nano- and picophytoplankton in the DCM layer. Higher lipid to Chl a ratios suggest enhanced lipid biosynthesis in the nutrient poorer regions. The various lipid classes pointed to possible mechanisms of phytoplankton adaptation to the nutritional conditions. Thus, it is likely that adaptation comprises the replacement of membrane phospholipids by non-phosphorus containing glycolipids under low phosphorus conditions. The qualitative and quantitative lipid compositions revealed that phospholipids were the most degradable lipids, and their occurrence decreased with increasing depth. In contrast, wax esters, possibly originating from zooplankton, survived downward transport probably due to the fast sinking rate of particles (fecal pellets). The important contribution of glycolipids in deep waters reflected their relatively stable nature and degradation resistance. A lipid-based proxy for the lipid degradative state (Lipolysis Index) suggests that many lipid classes were quite resistant to degradation even in the deep ocean.     

A multivariate analysis of larval fish and paralarval cephalopod assemblages at Great Meteor Seamount

Assemblage structures and distribution patterns of larval fishes and paralarval cephalopods were examined in September 1998 at Great Meteor Seamount, an isolated seamount located in the subtropical eastern North Atlantic. Early life stages of fish (n=18555) and cephalopods (n=1200) were collected at 23 stations with a multiple opening–closing net, in seven discrete depth strata from 290 m depth (close to the seamount plateau) to the surface. Oceanic species dominated in both taxonomic groups. A peak in diversity was observed at an intermediate depth, in the 100–150 m water layer. Direct and indirect gradient analyses showed distinct species assemblages in the upper and lower part of the water column, separated by approximately 150 m. The division was statistically significant, although a considerable overlap between species was also observed. Above the summit, vertical gaps were found in the distributions of the deeper assemblages, likely caused by increased predation pressure by benthopelagic fish. Horizontal distribution patterns of fish and cephalopods were similar and corresponded to the structure of closed circulation cells detected above the flanks and the flat plateau area. Fish assemblages were significantly different between the inner and outer seamount regime, which was approximately separated by the 1500 m depth contour. Differences in the taxonomic composition of cephalopods were less pronounced; for only one cephalopod species could a direct association with the seamount be assumed. The study indicates a significant retention potential at the seamount that facilitates local recruitment of resident stocks and generates self-sustainable populations isolated from the continental shelf and oceanic islands.     

Basin-scale variability of phytoplankton biomass,production and growth in the Atlantic Ocean

The latitudinal distributions of phytoplankton biomass, composition and production in the Atlantic Ocean were determined along a 10,000-km transect from 50°N to 50°S in October 1995, May 1996 and October 1996. Highest levels of euphotic layer-integrated chlorophyll a (Chl a) concentration (75–125 mg Chl m⁻²) were found in North Atlantic temperate waters and in the upwelling region off NW Africa, whereas typical Chl a concentrations in oligotrophic waters ranged from 20 to 40 mg Chl m⁻². The estimated concentration of surface phytoplankton carbon (C) biomass was 5–15 mg C m⁻² in the oligotrophic regions and increased over 40 mg C m⁻² in richer areas. The deep chlorophyll maximum did not seem to constitute a biomass or productivity maximum, but resulted mainly from an increase in the Chl a to C ratio and represented a relatively small contribution to total integrated productivity. Primary production rates varied from 50 mg C m⁻² d⁻¹ at the central gyres to 500–1000 mg C m⁻² d⁻¹ in upwelling and higher latitude regions, where faster growth rates (μ) of phytoplankton (>0.5 d⁻¹) were also measured. In oligotrophic waters, microalgal growth was consistently slow [surface μ averaged 0.21±0.02 d⁻¹ (mean±SE)], representing <20% of maximum expected growth. These results argue against the view that the subtropical gyres are characterized by high phytoplankton turnover rates. The latitudinal variations in μ were inversely correlated to the changes in the depth of the nitracline and positively correlated to those of the integrated nitrate concentration, supporting the case for the role of nutrients in controlling the large-scale distribution of phytoplankton growth rates. We observed a large degree of temporal variability in the phytoplankton dynamics in the oligotrophic regions: productivity and growth rates varied in excess of 8-fold, whereas microalgal biomass remained relatively constant. The observed spatial and temporal variability in the biomass specific rate of photosynthesis is at least three times larger than currently assumed in most satellite-based models of global productivity.     

Polychaete community structure in the South Eastern Arabian Sea continental margin (200–1000 m)

Macrofaunal polychaete communities (>500 µm) in the South Eastern Arabian Sea (SEAS) continental margin (200–1000 m) are described, based on three systematic surveys carried out in 9 transects (at ~200 m, 500 m and 1000 m) between 7°00′and 14°30′N latitudes. A total of 7938 polychaetes belonging to 195 species were obtained in 136 grab samples collected at 27 sites. Three distinct assemblages were identified in the northern part of the SEAS margin (10–14°30′N), occupying the three sampled depth strata (shelf edge, upper and mid-slope) and two assemblages (shelf edge and slope) in the south (7–10°N). Highest density of polychaetes and dominance of a few species were observed in the shelf edge, where the Arabian Sea oxygen minimum zone (OMZ) impinged on the seafloor, particularly in the northern transects. The resident fauna in this region (Cossura coasta, Paraonis gracilis, Prionospio spp. and Tharyx spp.) were characteristically of smaller size, and well suited to thrive in the sandy sediments in OMZ settings. Densities were lowest along the most northerly transect (T9), where dissolved oxygen (DO) concentrations were extremely low (<0.15 ml l⁻¹, i.e.<6.7 μmol l⁻¹). Beyond the realm of influence of the OMZ (i.e. mid-slope, ~1000 m), the faunal density decreased while species diversity increased. The relative proportion of silt increased with depth, and the dominance of the aforementioned species decreased, giving way to forms such as Paraprionospio pinnata, Notomastus sp., Eunoe sp. and lumbrinerids. Relatively high species richness and diversity were observed in the sandy sediments of the southern sector (7–9°N), where influence of the OMZ was less intense. The area was also characterized by certain species (e.g. Aionidella cirrobranchiata, Isolda pulchella) that were nearly absent in the northern region. The gradients in DO concentration across the core and lower boundary of the OMZ, along with bathymetric and latitudinal variation in sediment texture, were responsible for differences in polychaete size and community structure on the SEAS margin. Spatial and temporal variations were observed in organic matter (OM) content of the sediment, but these were not reflected in the density, diversity or distribution pattern of the polychaetes.     

Bioluminescence in the deep sea: Free-fall lander observations in the Atlantic Ocean off Cape Verde

A novel autonomous free-fall lander vehicle, with a capability down to 6000 m, was deployed off Cape Verde for studies on bioluminescence in the deep sea. The system was equipped with a high-sensitivity Intensified Silicon Intensified Target (ISIT) video camera, a programmable control-recording unit and an acoustic current meter with depth and temperature sensors. The ISIT lander was used in three modes: (1) free falling at 34 m min⁻¹, with the camera looking downwards at a mesh screen, recording impacts of luminescent organisms to obtain a vertical profile down to the abyssal sea floor, sampling at >100 l s⁻¹; (2) rotating, with the lander on the sea floor and the camera orienting to the bottom current using a servo-controlled turntable, impacts of luminescent organisms carried by the bottom current onto a mesh screen mounted 0.5 m in front of the camera were recorded to estimate abundance in the benthic boundary layer; (3) baited, with the camera focused on a bait placed on the sea floor.Profiles recorded abundance of luminescent organisms as 26.7 m⁻³ at 500–999 m depth, decreasing to 1.6 m⁻³ at 2000–2499 m and 0.5 m⁻³ between 2500 m and the sea floor at 4046 m, with no further detectable significant change with depth. Rotator measurements at a 0.5 m height above the sea floor gave a mean abundance of 0.47 m⁻³ in the benthic boundary layer at 4046 m and of 2.04 m⁻³ at 3200 m. Thirty five minutes after the bait was placed on the sea floor at 3200 m, bioluminescent fauna apparently arrived at the bait and produced luminescent displays at a rate of 2 min⁻¹. Moving, flashing light sources were observed and luminescent material was released into the bottom current.     

Occurrence and turnover of DMSP and DMS in deep waters of the Ross Sea,Antarctica

High concentrations of the phytoplankton metabolite dimethylsulfoniopropionate (DMSP) and its degradation product dimethylsulfide (DMS) are associated with blooms of Phaeocystis antarctica in the Ross Sea, Antarctica. Episodic and rapid vertical export of Phaeocystis biomass to deep water has been reported for the Ross Sea, therefore we examined the distribution and microbial consumption rates of DMSP and DMS throughout the sub-euphotic water column. Total DMSP (dissolved+particulate; DMSPt) was present at 0.5–22 nM at depths between 70 and 690 m during both the early bloom (November) and the late bloom (January). Sub-euphotic peaks of DMSP were sometimes associated with mid-water temperature maxima, and elevated DMSP below 70 m was found mainly in water masses characterized as Modified Circumpolar Deep Water or Antarctic Shelf Water. Overall, 50–94% of the integrated water-column DMSPt was found below the euphotic zone. At one station during the early bloom, local maxima of DMSPt (14 nM) and DMS (20 nM) were observed between 113 and 240 m and these maxima corresponded with high chlorophyll a concentrations, P. antarctica cell numbers, and Fv/Fm (the quantum yield of photosystem II). During the late bloom, a sub-euphotic maximum of DMSPt (15.8 nM) at 250 m cooccurred with peaks of chlorophyll a concentration, DMSP lyase activity, bacterial production and dissolved DMSP consumption rates. DMSP turnover contributed ～12% of the bacterial carbon demand between 200 and 400 m. DMS concentrations peaked at 286 m but the maximum concentration (0.42 nM) was far lower than observed during the early bloom, probably because of relatively rapid biological consumption of DMS (1–3 turnovers per day) which, in turn, contributed to elevated dissolved dimethylsulfoxide (DMSO) concentrations. Relatively stable DMSPt distributions at some sites suggest that rapid sinking of Phaeocystis biomass is probably not the major mechanism responsible for mesopelagic DMSP accumulations. Rather, subduction of near-surface water masses, lateral advective transport or trapping of slowly sinking P. antarctica biomass in intermediate water masses are more likely mechanisms. We found that a culture of P. antarctica maintained cellular integrity during 34 days of darkness, therefore the presence of intact cells (and DMSP) at depth can be explained even under a slow sinking/advection scenario. Whatever the mechanism, the large pools of DMSP and DMS below the euphotic zone suggest that export exerts a control on potential DMS emission from the surface waters of the Ross Sea.     

Microbial respiration in the Levantine Sea: evolution of the oxidative processes in relation to the main Mediterranean water masses

First data on microbial respiration in the Levantine Sea are reported with the aim of assessing the distribution of oxidative processes in association with the main Mediterranean water masses and the changing physical structure determined by the Eastern Mediterranean Transient. Respiratory rates, in terms of metabolic carbon dioxide production, were estimated from measured electron transport system activities in the polygonal area of the Levantine Sea (32.5–36.5 N Latitude, 26.0–30.25 E Longitude) and at Station Geo’95, in the Ionian Sea (35°34.88 N; 17°14.99 E). At the Levantine Sea, the mean carbon dioxide production rate decreased from the upper to the deeper layers and varied from 22.0±12.4 μg C h⁻¹ m⁻³ in the euphotic layer to 1.30±0.5 μg C h⁻¹ m⁻³ in the depth range between 1600 and 3000 m. Significant differences were found among upper, intermediate and bottom layers. The euphotic zone supported a daily carbon dioxide production of 96.6 mg C d⁻¹ m⁻² while the aphotic zone (between 200 and 3000 m) sustained a 177.1 mg C d⁻¹ m⁻² carbon dioxide production. In Station Geo’95, the carbon dioxide production rates amounted to 170.4 and 102.2 mg C d⁻¹ m⁻² in the euphotic and aphotic zones, respectively. The rates determined in the identified water masses showed a tight coupling of respiratory processes and Mediterranean circulation patterns. The increasing respiratory rates in the deep layers of the Levantine Sea are explained by the introduction of younger waters recently formed in the Aegean Sea.     

Monsoonal influence on the distribution of phytoplankton pigments in the Arabian Sea

Variations in the distribution of chemotaxonomic pigments were monitored in the Arabian Sea and the Gulf of Oman at the end of the SW monsoon in September 1994 and during the inter-monsoon period in November/December 1994 to determine the seasonal changes in phytoplankton composition. The Gulf of Oman was characterized by sub-surface chlorophyll maxima at 20-40 m during both seasons, and low levels of divinyl chlorophyll a indicated that prochlorophytes did not contribute significantly to the total chlorophyll a. Prymnesiophytes (19′-hexanoyloxyfucoxanthin), diatoms (fucoxanthin) and chlorophyll b containing organisms accounted for most of the phytoplankton biomass in September, while prymnesiophytes dominated in November/December. In the Arabian Sea in September, high total chlorophyll a concentrations up to 1742 ng l^-1 were measured in the coastal upwelling region and a progressive decline was monitored along the 1670 km offshore transect to oligotrophic waters at 8°N. Divinyl chlorophyll a was not detected along this transect except at the two most southerly stations where prochlorophytes were estimated to contribute 25–30% to the total chlorophyll a. Inshore, the dominance of fucoxanthin and/or hexanoyloxyfucoxanthin indicated that diatoms and prymnesiophytes generally dominated the patchy phytoplankton community, with zeaxanthin-containing Synechococcus also being important, especially in surface waters. At the southern oligotrophic localities, Synechococcus and prochlorophytes dominated the upper 40 m and prymnesiophytes were the most prominent at the deep chlorophyll maximum. During the inter-monsoon season, total chlorophyll a concentrations were generally half those measured in September and highest levels were found on the shelf (1170 ng l^-1). Divinyl chlorophyll a was detected at all stations along the Arabian Sea transect, and we estimated that prochlorophytes contributed between 3 and 28% to the total chlorophyll a, while at the two oligotrophic stations this proportion increased to 51–52%. While procaryotes were more important in November/December than September, eucaryotes still accounted for >50% of the total chlorophyll a. Pigment/total chlorophyll a ratios indicated that 19′-hexanoyloxyfucoxanthin-containing prymnesiophytes were the dominant group, although procaryotes accounted for 65% at the two southerly oligotrophic stations.     

Zooplankton grazing in the Atlantic Ocean: A latitudinal study

Mesozooplankton and 63–200 μm net-collected microzooplankton grazing on phytoplankton and protozoans was evaluated by 24-h incubations on a latitudinal transect in the Atlantic Ocean, from 35°N to 38°S (AMT-15; September–October 2004). The sampling area comprised contrasting ecosystems, including upwelling zones and oligotrophic subtropical gyres. Grazing impacts of mesozooplankton and 63–200 μm microzooplankton on total chlorophyll a (Chl a), >5 μm Chl a, ciliates, and dinoflagellates were low for both zooplankton size fractions, always removing<1.5% of the standing stocks of these groups. Grazing had a slightly greater impact upon primary production (up to 10% of primary production consumed daily), although on most occasions grazing removed<1% of primary production per day. To account for the reduction of micrograzers by predators in the experimental bottles and the consequent reduction of grazing pressure, the data were corrected with knowledge on the decrease of microzooplankton during incubations and global estimates of microzooplankton grazing. The corrected grazing rates for mesozooplankton ranged from 4% to 28% of the primary production consumed daily, and from 1% to 2% of the standing stock of Chl a removed every day. The 63–200 μm microzooplankton corrected grazing impact was always<5% of the primary production and standing stock consumed per day. The corrected grazing activity of 63–200 μm microzooplankton and mesozooplankton rendered daily rations ranging from 3% to 38% of the body carbon consumed daily, not sufficient for basal metabolism in most of the areas studied. Finally, the data on mesozooplankton grazing on primary production confirm the recent hypothesis of a decline of the relative importance of mesozooplankton grazing on primary producers with increasing primary production [Calbet, A., 2001. Mesozooplankton grazing effect on primary production: a global comparative analysis in marine ecosystems. Limnology and Oceanography 46, 1824–1830].     

Zooplankton metabolism and carbon demand at two seamounts in the NE Atlantic

Zooplankton metabolic rates, determined from electron transfer system (ETS) activity, were studied at two seamounts (Seine: 34°N, 14°W, summit depth ∼170 m; Sedlo: 40°N, 27°W, summit depth ∼750 m) in the northeast (NE) Atlantic during three cruises in November 2003, April 2004 and July 2004. ETS activity and respiratory carbon demand were measured for samples taken at seamount and open-ocean locations in order to probe the hypothesis of locally enhanced seamount productivity. ETS activity and biomass revealed no consistent diel patterns of feeding activity and vertical migration at Seine and Sedlo Seamounts. Spatial differences of biomass-specific ETS activity were observed at both seamounts and coincided with differences in food abundance and quality. At Seine Seamount in April 2004, biomass-specific ETS activity was on average higher at the seamount locations compared to the open ocean, though the enhancement was of a lower magnitude than spatial and temporal variability and had no apparent influence on zooplankton respiratory carbon demand or biomass. A persistent pattern of reduced zooplankton biomass above the summit location at Seine Seamount in April 2004 and July 2004 resulted in a local reduction of respiratory carbon demand. At Sedlo Seamount in November 2003, large spatial differences in biomass-specific ETS activity observed at the seamount locations resulted in a large range of respiratory carbon demand at the seamount, but were not reflected in zooplankton biomass. The depth-integrated (0–150 m) median respiratory carbon demand of the zooplankton community estimated from day and night hauls was 2.1 mg C m⁻² d⁻¹ at Seine Seamount (range: 0.3–6.3) and 2.9 mg C m⁻² d⁻¹ at Sedlo Seamount (range: 1.6–12.0). The sporadic nature and low magnitude of locally higher zooplankton respiration rates at the seamounts, which did not result in locally higher zooplankton standing stock biomass, lead us to reject the hypothesis that locally enhanced seamount productivity provides an autochthonous food supply to the resident faunas at Seine and Sedlo Seamounts. Instead, we conclude that the faunas at both seamounts are more likely supported by advection of food from the surrounding ocean.     

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.     

Estimates of primary production by remote sensing in the Arctic Ocean: Assessment of accuracy with passive and active sensors

The Arctic Ocean, including its regional shelf seas, is assumed to play an important role in the global carbon cycle. However, the true magnitude of annual production is unknown, as in situ data are sparse in time and space. Remote sensing technology has the potential to provide large scale estimates of phytoplankton biomass at much higher frequency and spatial coverage than shipboard observations in this remote region. Subsurface peaks in both biomass and primary production (PP), which are the characteristics of the Arctic, are shown to limit the reliability of ocean color based integrated PP (IPP) models in the Chukchi Sea. Here we report that the retrievals of IPP from remotely sensed ocean color data were accurate only when limited to 1.2 optical depths, which severely constrains the utility of ocean color remote sensing for the assessment of Arctic Ocean dynamics.Active sensors such as LIDAR, can, in combination with passive ocean color, dramatically improve our ability to estimate IPP for the Arctic. IPP retrievals were improved to within a factor of 2–3 of the measured values, when the vertical distribution of Chl a was determined to a resolution of 1 m using modeled LIDAR retrievals of the beam attenuation coefficient. This was far better than models using only passive ocean color. The instrument specifications of the current NASA spaceborne LIDAR (CALIOP) allow for the retrieval of K_d at a depth resolution of 23 m. Even with this constraint, however, the accuracy of the modeled IPP was improved over passive ocean color retrievals to approximately a factor of 3. The Arctic is a perfect location to merge ocean color and LIDAR measurements as the polar orbit of CALIOP provides complete grid coverage of the area every 8 days, crossing the horizontal gradients in Chl a already known to exist from passive ocean color observations.     

DOC dynamics in the meso and bathypelagic layers of the Mediterranean Sea

Seven years (2001–2008) of dissolved organic carbon (DOC) vertical profiles were examined in order to assess the main processes determining DOC concentration and distribution in the meso- and bathypelagic layers of the Mediterranean Sea. As expected, DOC showed high and highly variable concentrations in the surface layer of 57–68 μM (average values between 0 and 100 m), with a decrease to 44–53 μM between 200 and 500 m. Deep DOC distribution was strongly affected by deep-water formation, with a significant increase to values of 76 μM in recently ventilated deep waters, and low concentrations, comparable to those observed in the open oceanic waters (34–45 μM), where the oldest, deep waters occurred. In winter 2004/2005 a deep-water formation event was observed and the consequent DOC export at depth was estimated to range between 0.76–3.02 Tg C month^–1. In the intermediate layer, the main path of the Levantine Intermediate Water (LIW) was followed in order to estimate the DOC consumption rate in its core. Multiple regression between DOC, apparent oxygen utilization (AOU), and salinity indicated that 38% of the oxygen consumption was related to DOC mineralization when the effect of mixing was removed. In deep waters of the southern Adriatic Sea a DOC decrease of 6 μM, together with an AOU increase of 9 μM, was observed between the end of January 2008 and the end of June 2008 (5 months). These data indicate a rate of microbial utilization of DOC of about 1.2 μM C month⁻¹, with 92% of the oxygen consumption due to DOC mineralization. These values are surprisingly high for the deep sea and represent a peculiarity of the Mediterranean Sea.     

Consumption and production on scales of a few days of inorganic carbon,nitrate and oxygen by the planktonic community: results of continuous measurements at the Dyfamed Station in the northwestern Mediterranean Sea (May 1995)

Continuous measurements between 0 and 200 m depth were performed every 2 h over two separate periods of four days at a station in the open northwestern Mediterranean Sea (Dyfamed Station) during the Dynaproc cruise in May 1995. Estimates of the daily variations in profiles of temperature, partial pressure of CO₂, oxygen, chlorophyll a and nutrients were obtained. The distributions of the various physical and chemical properties were clearly different during the two time series, which were separated by a period of 11 days during which a wind event occurred. The mean daily utilization or production due to biological processes of dissolved inorganic carbon (DIC), nitrate+nitrite and oxygen were calculated along isopycnals using a vertical diffusion model. Between the surface and about 20 m depth, DIC was consumed and O₂ released during the two time series while the nitrate+nitrite concentrations as well as supplies were zero. After the wind event, the O₂ : C : N ratios of consumption (or production) were, on average, near the Redfield ratios, but during the first time series, the C : N utilization ratio between 20 and 35 m was two to three times that of Redfield stoichiometry and the oxygen release was low. The integrated net community production (NCP) in terms of carbon was equivalent during the two time series, whereas the chlorophyll a biomass was twice as high, on average, during the first time series but did decrease. These results imply that the production systems were different during the two periods. The first time series corresponds to a period at the end of production, due to the nutrient depletion in the euphotic layer. The formation of degradation products of the living material in dissolved organic form is probably important as indicated by the high C : N utilization ratios. The second time series corresponds to a reactivation of the primary production due to the upward shift of nutrients after the wind event.     

First in situ observations of the deep-sea carnivorous ascidian Dicopia antirrhinum Monniot C., 1972 in the Western Mediterranean Sea

Dicopia antirrhinum C. Monniot, 1972 is a rare species of deep-sea ascidian belonging to the Family Octacnemidae, reported at depths of 1000–2500 m in European Atlantic waters. Adult individuals have never been reported before in the Mediterranean Sea, where only seven juvenile specimens were found in 1975 at 500 m water depth in the Central basin (Malta). The affinities of these specimens with D. antirrhinum were noted, but lack of some typical characters of the species in juveniles prevented a definite taxonomical identification. No other member of the Octacnemidae has ever been found in the Mediterranean. In this study we describe the sampling of an adult specimen of D. antirrhinum at around 1100 m water depth on the flank of the La Fonera (Palamós) canyon, Northwestern Mediterranean, confirming their presence in the Mediterranean Sea. We also observed 5 individuals of this species on their natural habitat with a Remotely Operated Vehicle (ROV). Our results highlight the potential occurrence of Octacnemidae, the presence of which has been largely overlooked, in several deep-sea canyon areas within the Western Mediterranean basin. These observations are important because they indicate the need for increased sampling effort with new technologies, such as ROVs, in ecologically relevant habitats such as canyons, in order to obtain a more accurate picture of deep-sea biodiversity in the Mediterranean Sea.     

Picophytoplankton dynamics and production in the Arabian Sea during the 1995 Southwest Monsoon

Phytoplankton community structure is expected to shift to larger cells (e.g., diatoms) with monsoonal forcing in the Arabian Sea, but recent studies suggest that small primary producers remain active and important, even in areas strongly influenced by coastal upwelling. To better understand the role of smaller phytoplankton in such systems, we investigated growth and grazing rates of picophytoplankton populations and their contributions to phytoplankton community biomass and primary productivity during the 1995 Southwest Monsoon (August–September). Environmental conditions at six study stations varied broadly from open-ocean oligotrophic to coastal eutrophic, with mixed-layer nitrate and chlorophyll concentrations ranging from 0.01 to 11.5 μM NO₃ and 0.16 to 1.5 μg Chl a. Picophytoplankton comprised up to 92% of phytoplankton carbon at the oceanic stations, 35% in the diatom-dominated coastal zone, and 26% in a declining Phaeocystis bloom. Concurrent in situ dilution and ¹⁴C-uptake experiments gave comparable ranges of community growth rates (0.53–1.05 d⁻¹ and 0.44–1.17 d⁻¹, to the 1% light level), but uncertainties in C:Chl a confounded agreement at individual stations. Microzooplankton grazing utilized 81% of community phytoplankton growth at the oligotrophic stations and 54% at high-nutrient coastal stations. Prochlorococcus (PRO) was present at two oligotrophic stations, where its maximum growth approached 1.4 d⁻¹ (two doublings per day) and depth-integrated growth varied from 0.2 to 0.8 d⁻¹. Synechococcus (SYN) growth ranged from 0.5 to 1.1 d⁻¹ at offshore stations and 0.6 to 0.7 d⁻¹ at coastal sites. Except for the most oligotrophic stations, growth rates of picoeukaryotic algae (PEUK) exceeded PRO and SYN, reaching 1.3 d⁻¹ offshore and decreasing to 0.8 d⁻¹ at the most coastal station. Microzooplankton grazing impact averaged 90, 70, and 86% of growth for PRO, SYN, and PEUK, respectively. Picoplankton as a group accounted for 64% of estimated gross carbon production for all stations, and 50% at high-nutrient, upwelling stations. Prokaryotes (PRO and SYN) contributed disproportionately to production relative to biomass at the most oligotrophic station, while PEUK were more important at the coastal stations. Even during intense monsoonal forcing in the Arabian Sea, picoeukaryotic algae appear to account for a large portion of primary production in the coastal upwelling regions, supporting an active community of protistan grazers and a high rate of carbon cycling in these areas.     

Submarine groundwater discharge (SGD) as a main nutrient source for benthic and water-column primary production in a large intertidal environment of the Yellow Sea

The biogeochemistry and magnitude of submarine groundwater discharge (SGD) was investigated in one of the largest tidal flat ecosystems worldwide, along the Yellow Sea coast. A representative semi-enclosed embayment located in the south eastern Yellow Sea, Hampyeong Bay, was chosen for this purpose. Groundwater and seawater samples were collected in three seasons (May, July, and September) and analyzed for Ra isotopes, nutrients, and photosynthetic pigments. The biogeochemistry of SGD was strongly influenced by tidal oscillations and seasonal precipitation changes and switched from a brackish, nutrient-enriched regime in May and July to an exclusively saline regime, with lower nutrient concentrations, in September. SGD magnitudes, calculated by using a ²²⁶Ra mass balance model, were 0.14 m³ m^? 2 d^? 1 in May and 0.35 m³ m^? 2 d^? 1 in September. A nutrient mass balance was established for the two campaigns, which suggests that SGD causes the flushing of substantial amounts of pore water nutrients into this embayment; because of SGD, the embayment acts as a source of dissolved inorganic silicates (DSi) that are transported to the open ocean. Potential C fixation rates derived from this nutrient mass balance were compared with two different models for water-column phytoplankton productivity based on water-column Chl a and local irradiation levels. The Chl a-based models generally showed lower C fixation rates than the nutrient-based mass balance, indicating removal of up to 70% of the nutrients by other primary producers, such as benthic algae. During monsoon season, when benthic algal biomass is high and nutrient fluxes are substantial due to a terrestrial component, SGD — driven benthic primary production could play a significant role in this large tidal flat ecosystem.     

Characterization of the solar light field within the ocean mesopelagic zone based on radiative transfer simulations

The solar light field within the ocean from the sea surface to the bottom of the mesopelagic zone was simulated with a radiative transfer model that accounts for the presence of inelastic radiative processes associated with Raman scattering by water molecules, fluorescence of colored dissolved organic matter (CDOM), and fluorescence of chlorophyll-a contained in phytoplankton. The simulation results provide a comprehensive characterization of the ambient light field and apparent optical properties (AOPs) across the entire visible spectral range within the depth range 200–1000 m of the entire mesopelagic zone for varying chlorophyll-a concentration and seawater optical properties in the mixed surface layer of the ocean. With increasing depth in the mesopelagic zone, the solar irradiance is reduced by ~9–10 orders of magnitude and exhibits a major spectral maximum in the blue, typically centered around a light wavelength of 475 nm. In the green and red spectral regions, the light levels are significantly lower but still important owing to local generation of photons via inelastic processes, mostly Raman scattering and to a lesser extent CDOM fluorescence. The Raman scattering produces a distinct secondary maximum in irradiance spectra centered around 565 nm. Comparisons of our results with light produced by the radioactive decay of the unstable potassium isotope contained in sea salt (⁴⁰K) indicates that the solar irradiance dominates over the ⁴⁰K-produced irradiance within the majority of the mesopelagic zone for most scenarios considered in our simulations. The angular distribution of radiance indicates the dominance of downward propagation of light in the blue and approach to uniform distribution in the red throughout the mesopelagic zone. Below the approximate depth range 400–500 m, the shape of the angular distribution is nearly invariant with increasing depth in the green and red and varies weakly in the blue. The AOPs at any light wavelength also assume nearly constant values within the deeper portion of the mesopelagic zone. These results indicate that the mesopelagic light field reaches a nearly-asymptotic regime at depths exceeding ~400–500 m.