Biodiversity and community composition of sediment macrofauna associated with deep-sea Lophelia pertusa habitats in the Gulf of Mexico

Scleractinian corals create three-dimensional reefs that provide sheltered refuges, facilitate sediment accumulation, and enhance colonization of encrusting fauna. While heterogeneous coral habitats can harbor high levels of biodiversity, their effect on the community composition within nearby sediments remains unclear, particularly in the deep sea. Sediment macrofauna from deep-sea coral habitats (Lophelia pertusa) and non-coral, background sediments were examined at three sites in the northern Gulf of Mexico (VK826, VK906, MC751, 350–500 m depth) to determine whether macrofaunal abundance, diversity, and community composition near corals differed from background soft-sediments. Macrofaunal densities ranged from 26 to 125 individuals 32 cm⁻² and were significantly greater near coral versus background sediments only at VK826. Of the 86 benthic invertebrate taxa identified, 16 were exclusive to near-coral habitats, while 14 were found only in background sediments. Diversity (Fisher’s α) and evenness were significantly higher within near-coral sediments only at MC751 while taxon richness was similar among all habitats. Community composition was significantly different both between near-coral and background sediments and among the three primary sites. Polychaetes numerically dominated all samples, accounting for up to 70% of the total individuals near coral, whereas peracarid crustaceans were proportionally more abundant in background sediments (18%) than in those near coral (10%). The reef effect differed among sites, with community patterns potentially influenced by the size of reef habitat. Taxon turnover occurred with distance from the reef, suggesting that reef extent may represent an important factor in structuring sediment communities near L. pertusa. Polychaete communities in both habitats differed from other Gulf of Mexico (GOM) soft sediments based on data from previous studies, and we hypothesize that local environmental conditions found near L. pertusa may influence the macrofaunal community structure beyond the edges of the reef. This study represents the first assessment of L. pertusa-associated sediment communities in the GOM and provides baseline data that can help define the role of transition zones, from deep reefs to soft sediments, in shaping macrofaunal community structure and maintaining biodiversity; this information can help guide future conservation and management activities.     

Seasonality of particle-associated trace element fluxes in the deep northeast Atlantic Ocean

In the framework of the German contribution to the Joint Global Ocean Flux Study (JGOFS), deep-water fluxes of particle-associated trace elements were measured in the northeast Atlantic Ocean. The sinking particles were collected almost continuously from 1992 to 1996 at three time-series stations, L1 (33°N/22°W), L2 (47°N/20°W), and L3 (54°N/21°W), using sediment traps. The focus of the present study is the temporal variability of the particle-associated elemental fluxes of Al, Ca, Cd, Co, Cu, Fe, Mn, Ni, P, Pb, Ti, V, and Zn at a depth of 2000 m.A clear seasonality of the fluxes that persisted for several years was documented for the southernmost station (L1) at stable oligotrophic conditions in the area of the North Atlantic Subtropical Gyre East (NASTE). At L2 and L3, an episodic nature of the elemental fluxes was determined. Mesoscale eddies are known to frequently cause temporal and spatial variability in the flux of biogenic components in that area. These events modified the simple seasonal pattern controlled by the annual cycle at L2, in the North Atlantic Drift Region (NADR), and at L3, which was influenced by the Atlantic Arctic province (ARCT). All stations were characterized by an additional episodic lithogenic atmospheric supply reaching the deep sea.The integrated annual fluxes during the multi-year study revealed similar flux magnitudes for lithogenic elements (Al, Co, Fe, Ti, and V) at L2 and L3 and roughly twofold fluxes at L1. Biogenic elements (Cd, P, and Zn) showed the opposite trend, i.e., two to fourfold higher values at L2 and L3 than at L1. For Mn, Ni, and Cu, the spatial differences were smaller, perhaps because of the intermediate behavior, between lithogenic and biogenic, of these elements. Similarly, among the three study sites, there were no noticeable differences in the total annual flux of Pb.The respective lithogenic fractions of the deep-sea fluxes of Cd, Co, Cu, Mn, Ni, V, and Zn were subtracted based on the amount of Al, with the average composition of the continental crust as reference. This procedure allowed estimation of the labile trace element fraction (TE_exc) of the particles, i.e., TE taken up or scavenged during particle production and sedimentation. The ratios of TE_exc/P clearly demonstrated an enrichment of TE over labile P from biogenic surface material to the deep sea for Zn (factor 4–6), Mn (12–27), Ni (3–5), and Cu (9–25); an intermediate status for Co (0.5–2.2); and depletion for Cd vs. P (0.2–0.4). Surprisingly, the recycling behavior of excess Co was found to be similar to that of P. Hence, Co_exc behaved like a biogenic element; this is in contrast to total Co, which is dominated by the refractory lithogenic fraction.Moreover, it is argued that these excess elemental fluxes caused a loss of the dissolved elements in upper waters, since their transport reaches the deep-sea waters at 2000 m, a depth far below of deep-winter mixing and upwelling. The annual amount of excess TE exported from surface waters was estimated to be 1.3×10⁹ mol Zn y^?1, 4.4×10⁹ mol Mn y^?1, 4.9×10⁸ mol Ni y^?1, 2.2×10⁷ mol Cd y^?1, 7.4×10⁸ mol Cu y^?1, and 2.7×10⁷ mol Co y^?1 for the whole North Atlantic Ocean. Important primary sources that could replenish these losses are the aeolian and fluvial supply processes.     

Mesoscale distribution and advection of overwintering Calanus finmarchicus off the shelf of northern Norway

Data collected on a cruise in January 2008, using a laser optical plankton counter, conductivity–temperature–depth sensors, and a lowered acoustic Doppler current profiler, was used to study the mesoscale distribution and advection of overwintering Calanus finmarchicus in its deep water winter habitat off the shelf of northern Norway. The overwintering animals were generally located immediately below the Atlantic Water (AW) in Arctic Intermediate Water (AIW), in the 600–1200 m depth range. The depth of the interface between AW and AIW varied considerably in the area and this was clearly reflected in the C. finmarchicus distribution. Maximum abundance varied from about 80 ind m^?3 to more than 200 ind m^?3 at the different stations. Current measurements showed the richness of mesoscazle eddies, with speeds exceeding 70 cm s^?1 at the surface and rapidly decreasing with depth. In the main overwintering layer the eddy features were also clearly seen, but with speeds generally below 20 cm s^?1. C. finmarchicus were found in the whole survey area, but they were not homogeneously distributed. Advection of the copepods resulted in relatively high local rates of change in overwintering C. finmarchicus abundance with mean value of 8% per day in the area. It is clear that mesoscale physical processes greatly contribute to the variability in the abundance of overwintering C. finmarchicus off the shelf of northern Norway. The collected data are also a valuable addition to the generally sparse datasets on the C. finmarchicus winter distribution and the role of the Lofoten basin in the large scale system is also discussed.     

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.     

Total organic and inorganic carbon exchange through the Strait of Gibraltar in September 1997

The total organic carbon (TOC) and total inorganic carbon (C_T) exchange between the Atlantic Ocean and the Mediterranean Sea was studied in the Strait of Gibraltar in September 1997. Samples were taken at eight stations from western and eastern entrances of the Strait and at the middle of the Strait (Tarifa Narrows). TOC was analyzed by a high-temperature catalytic oxidation method, and C_T was calculated from alkalinity–pH_T pairs and appropriate thermodynamic relationships. The results are used in a two-layer model of water mass exchange through the Strait, which includes the Atlantic inflow, the Mediterranean outflow and the interface layer in between. Our observations show a decrease of TOC and an increase of C_T concentrations from the surface to the bottom: 71–132 μM C and 2068–2150 μmol kg⁻¹ in the Surface Atlantic Water, 74–95 μM C and 2119–2148 μmol kg⁻¹ in the North Atlantic Central Water, 63–116 μM C and 2123–2312 μmol kg⁻¹ in the interface layer, and 61–78 μM C and 2307–2325 μmol kg⁻¹ in the Mediterranean waters. However, within the Mediterranean outflow, we found that the concentrations of carbon were higher at the western side of the Strait (75–78 μM C, 2068–2318 μmol kg⁻¹) than at the eastern side (61–69 μM C, 2082–2324 μmol kg⁻¹). This difference is due to the mixing between the Atlantic inflow and the Mediterranean outflow on the west of the Strait, which results in a flux of organic carbon from the inflow to the outflow and an opposite flux of inorganic carbon. We estimate that the TOC input from the Atlantic Ocean to the Mediterranean Sea through the Strait of Gibraltar varies from (0.97±0.8)10⁴ to (1.81±0.90)10⁴ mol C s⁻¹ (0.3×10¹² to 0.56×10¹² mol C yr⁻¹), while outflow of inorganic carbon ranges from (12.5±0.4)10⁴ to (15.6±0.4)10⁴ mol C s⁻¹ (3.99–4.90×10¹² mol C yr⁻¹). The high variability of carbon exchange within the Strait is due to the variability of vertical mixing between inflow and outflow along the Strait. The prevalence of organic carbon inflow and inorganic carbon outflow shows the Mediterranean Sea to be a basin of active remineralization of organic material.     

Wave climate variability in the North-East Atlantic Ocean over the last six decades

Ocean surface gravity waves play a major role in many engineering and environmental problems, both in the open ocean and in coastal zones. Therefore, it is essential to improve our knowledge on spatial and temporal variability of wave climate. This study aims at investigating this variability in the North-East Atlantic Ocean (25°W–0°W and 30°N–60° N), using a 57-year hindcast (1953–2009) obtained with a spectral wave model forced with reanalysis wind fields. The hindcast analysis reveals firstly strong seasonal fluctuations of wave climate, with winters characterized by large and long-period waves of mean direction spreading from south-west to north-west, and summers characterized by smaller and shorter-period waves originating from norther directions. From northern (55°N) to southern (35°N) latitudes, the significant wave height (Hs) decreases by roughly 40%, the mean wave direction (Mwd) rotates clockwise by about 25% while the peak period (Tp) only grows by 5%. These three parameters also exhibit a strong inter-annual variability, particularly when winter-means (from 1st of December to 1st of April) are considered. Linear trend analysis over the studied period shows spatially variable long-term trends, with a significant increase of Hs (up to 0.02 m yr^?1) and a counterclockwise shift of Mwd (up to ?0.1° yr^?1) at northern latitude, contrasting with a fairly constant trend for Hs and a clockwise shift of Mwd (up to +0.15° yr^?1) at southern latitudes. Long-term trends of Tp are less significant, with still a slight increase in the north-eastern part of the study area (up to +0.01 s yr^?1). Eventually, a comparison between the inter-annual variability of the winter-means of the three selected wave parameters and the North Atlantic Oscillation (NAO) reveals: (1) a strong positive correlation between Hs and the NAO index at northern latitudes (correlation coefficient up to R = 0.91) and a significant negative correlation at southern latitudes (up to R = ?0.6); (2) no significant correlation for Mwd north of 40°N and a clear positive correlation southward of 40°N (up to R = 0.8) and (3) a northward increasing correlation for Tp (up to R = 0.8). Long-term trends for Hs, Mwd and Tp are finally explained by a significant increase in the NAO index over the studied period.     

Growth rates,densities, and distribution of Lophelia pertusa on artificial structures in the Gulf of Mexico

Using industry inspection video and ROV imaging, we examined Lophelia pertusa (Linnaeus 1758) on 10 artificial structures of known ages (9 to 100 years) in the northern Gulf of Mexico (GoM). Five different types deep-water energy installations with depths ranging from 320 to 995 m, and three shipwrecks with depths ranging from approximately 530 to 615 m, were examined. Density, depth ranges, and growth rates of L. pertusa colonies were calculated from video and image analysis. L. pertusa colonies were present on all structures examined. Minimum calculated growth rates for the largest colonies ranged from 0.32 to 3.23 cm/yr on the different structures. The shallowest depth at which L. pertusa was observed was 201 m and the deepest was 801 m, considerably expanding the known depth range of this species in the northern GoM. Colony density varied with structure type, age, and depth, with the highest density between 503 and 518 m on the single structure that spanned the entire depth range of occurrence of L. pertusa observed in this study. L. pertusa colonies growing on thinner and deeper installations appear to have higher colonization rates, i.e. to develop higher densities over a shorter time period, compared to those on shallower and more massive types of installations. However, on average, colonies have slower growth rates on these installations than colonies on more massive, shallower installations (compliant and solid installations). In general, the calculated minimum growth rates were higher on the installations than on the shipwrecks, which were substantially older. A continuum of colony sizes was documented on all installations, suggesting multiple settlement events. L. pertusa thickets were observed on the oldest anthropogenic structures, with most of the components of these structures covered by colonies of L. pertusa. Brown, orange, and mottled color-variants were documented for the first time in the GoM. All installations examined for this study were colonized by L. pertusa and it is likely that most artificial surfaces in appropriate depths in the GoM will be as well.     

Intra-annual variability of extremely rapid sedimentation onto Gardar Drift in the northern North Atlantic

North Atlantic sediment drifts are valuable archives for paleoceanographic reconstructions spanning various timescales. However, the short-term dynamics of such systems are poorly known, and this impinges on our ability to quantitatively reconstruct past change. Here we describe a high-resolution 319-day time-series of hydrodynamics and near-bottom (4 m) particulate matter flux variability at a 2600 m deep site with an extremely high sediment accumulation rate on the southern Gardar Drift in the North Atlantic. We compare our findings with the actual deposits at the site. The total annual particle flux amounted to ～360 g m^?2 yr^?1, varied from ～0.15 to >5.0 g m^?2 day^?1 and displayed strong seasonal compositional changes, with the highest proportion of fresh biogenic matter arriving after the spring bloom in June and July. Flux variability also depended on the changing input of lithogenic matter that had been (re)suspended for a longer time (decades). Active focussing of material from both sources is required to account for the composition and the magnitude of the total flux, which exceed observations elsewhere by an order of magnitude. The enhanced focussing or increased delivery appeared to be positively related to current velocity. The intercepted annual particle flux accounted for only 60% of the sediment accumulation rate of 600±20 g m^?2 yr^?1 (0.20±0.07 cm yr^?1), indicating higher intra- and inter-annual variability of both the biogenic and lithogenic fluxes and/or advection of additional sediment closer to the seafloor (i.e. <4 m). This temporal variability in the composition and amount of material deposited highlights intra-annual changes in the flux of lithogenic material, but also underscores the importance of (reworked) sediment focussing and seasonality of the biogenic flux. All should be taken into account in the interpretation of the paleorecord from such depositional settings.     

Variability of chlorophyll and primary production in the Eastern North Atlantic Subtropical Gyre: potential factors affecting phytoplankton activity

Size-fractionated chlorophyll-a and carbon incorporation rates were determined on a series of 13 cruises carried out from 1992 to 2001with the aim of investigating the patterns and causes of variability in phytoplankton chlorophyll and production in the Eastern North Atlantic Subtropical Gyral Province (NASE). Averaged (±SE) integrated chlorophyll-a concentration and primary production rate were 17±1 mg m⁻² and 253±22 mg C m⁻² d⁻¹. Small-sized cells (<2 μm) formed the bulk of phytoplankton biomass (71%) and accounted for 54% of total primary production. A clear latitudinal gradient in these variables was not detected. By contrast, large seasonal variability was detected in terms of primary production, although integrated phytoplankton biomass, as estimated from chlorophyll-a concentration, remained rather constant and did not display significant changes with time. Variability in primary production (PP) was related mainly to variability in surface temperature and surface chlorophyll-a concentration. The control exerted by surface temperature was related to nutrient availability. By contrary, euphotic-zone depth, depth of maximum concentration of chlorophyll-a and integrated chlorophyll-a did not contribute significantly to the high variability in primary production observed in this oligotrophic region.     

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.     

Increase in anthropogenic CO2 in the Atlantic Ocean in the last two decades

Data from the first systematic survey of inorganic carbon parameters on a global scale, the GEOSECS program, are compared with those collected during WOCE/JGOFS to study the changes in carbon and other geochemical properties, and anthropogenic CO₂ increase in the Atlantic Ocean from the 1970s to the early 1990s. This first data-based estimate of CO₂ increase over this period was accomplished by adjusting the GEOSECS data set to be consistent with recent high-quality carbon data. Multiple Linear Regression (MLR) and extended Multiple Linear Regression (eMLR) analyses to these carbon data are applied by regressing DIC with potential temperature, salinity, AOU, silica, and PO₄ in three latitudinal regions for the western and eastern basins in the Atlantic Ocean. The results from MLR (and eMLR provided in parentheses) indicate that the mean anthropogenic CO₂ uptake rate in the western basin is 0.70 (0.53) mol m^?2 yr^?1 for the region north of 15°N; 0.53 (0.36) mol m^?2 yr^?1 for the equatorial region between 15°N and 15°S; and 0.83 (0.35) mol m^?2 yr^?1 in the South Atlantic south of 15°S. For the eastern basin an estimate of 0.57 (0.45) mol m^?2 yr^?1 is obtained for the equatorial region, and 0.28 (0.34) mol m^?2 yr^?1 for the South Atlantic south of 15°S. The results of using eMLR are systematically lower than those from MLR method in the western basin. The anthropogenic CO₂ increase is also estimated in the upper thermocline from salinity normalized DIC after correction for AOU along the isopycnal surfaces. For these depths the results are consistent with the CO₂ uptake rates derived from both MLR and eMLR methods.     

Abundance and size distribution of transparent exopolymer particles (TEP) in a coccolithophorid bloom in the northern Bay of Biscay

The distribution of transparent exopolymer particles (TEP) was investigated during a coccolithophorid bloom in the northern Bay of Biscay (North Atlantic Ocean) in early June 2006. MODIS chlorophyll-a (Chl-a) and reflectance images before and during the cruise were used to localize areas of important biological activity and high reflectance (HR). TEP profiles along the continental margin, determined using microscopic (TEP_micro) and colorimetric (TEP_color) methods, showed abundant (6.1×10⁶–4.4×10⁷ L^?1) and relatively small (0.5–20 μm) particles, leading to a low total volume fraction (0.05–2.2 ppm) of TEP_micro and similar vertical profiles of TEP_color. Estimates of carbon content in TEP (TEP-C) derived from the microscopic approach yielded surface concentration of 1.50 μmol C L^?1. The contribution of TEP-C to particulate organic carbon (POC) was estimated to be 12% (molar C ratio) during this survey. Our results suggest that TEP formation is a probable first step to rapid and efficient export of C during declining coccolithophorid blooms.     

The Manila clam population in Arcachon Bay (SW France): Can it be kept sustainable?

The venerid clam Ruditapes philippinarum is the most prominent suspension-feeding bivalve inhabiting muddy intertidal seagrass beds in Arcachon Bay (SW France). It is exploited by fishermen, and Arcachon Bay ranks number one in France in terms of production and total biomass of this species. Previous studies revealed a decrease in the standing stock of R. philippinarum since 2003 and unbalanced length–frequency distributions with a lack of juveniles and of adults > 40 mm. Consequently, the population dynamics of this bivalve were studied at four intertidal sites and one oceanic site in Arcachon Bay. As clam size structure did not allow classical dynamics computations, field monitoring was coupled with field experiments (tagging–recapture) over two years. Monitoring of condition index and gonadal maturation stages highlighted a high variability in spawning number and intensity between sites. Recruitment events in the exploited area varied spatially but with uniformly low values. Von Bertalanffy Growth Function (VBGF) parameters (K, L_∞) were determined using Appeldoorn and ELEFAN methods. In the exploited sites in the inner lagoon, K was relatively high (mean = 0.72 yr^? 1) but L_∞ was low (mean = 41.1 mm) resulting in a moderate growth performance index (Φ′ = 2.99). Growth parameters were not correlated with immersion time and L_∞ was different between sites. Comparison of mortality coefficients (Z) between cage experiments and field monitoring suggested that fishing accounts for 65–75% of total adult mortality. Low recruitment, a low growth rate and a normal mortality rate led to low somatic production (4.1 and 8.7 g Shell-Free Dry Weight (SFDW) m^? 2 yr^? 1) and an annual P/B ratio from 0.44 to 0.92 yr^? 1. Under current conditions, the possibility of a sustainable population in Arcachon Bay will strongly depend on recruitment success and fishing management.     

Distribution of deep-water corals along the North American continental margins: Relationships with environmental factors

Despite the increasing attention to assemblages of deep-water corals in the past decade, much of this research has been focused on documenting and enumerating associated fauna. However, an understanding of the distribution of most species of coral and the ecological processes associated with these assemblages is still lacking. In this study, we qualitatively and quantitatively described the habitats of two families of deep-water corals in relation to six oceanographic factors (depth, slope, temperature, current, chlorophyll a concentration and substrate) on the Pacific and Atlantic Continental Margins of North America (PCM and ACM study areas, respectively). This study focused primarily on the distributions of Primnoidae and Paragorgiidae because of the large number of documented occurrences. For each environmental factor, deep-water coral locations were compared to the surrounding environment using χ² tests. On both continental margins, coral locations were found to be not randomly distributed within the study areas, but were within specific ranges for most environmental factors. In the PCM study area, Paragorgiidae and Primnoidae locations were found in areas with slopes ranging from 0° to 10.0°, temperature from −2.0 to 11.0 °C and currents from 0 to 143 cm s⁻¹. In the ACM study area, Paragorgiidae and Primnoidae locations were found in areas with slopes ranging from 0° to 1.4°, temperature ranging from 0 to 11.0 °C and currents ranging from 0 to 207 cm s⁻¹. Although the patterns in habitat characteristics were similar, differences existed between families with respect to particular environmental factors. In both study areas, most environmental parameters in locations where corals occurred were significantly different from the average values of these parameters as determined with χ² tests ($p<0.05$) except for substrate in Paragorgiidae locations and depth in Primnoidae locations on the PCM. This is the first study to show coral distributional patterns at the continental shelf/slope scale.     

Seasonal POC fluxes at BATS estimated from 210Po deficits

In this study at the Bermuda Atlantic Time-series Study (BATS) site we demonstrate that the polonium–lead disequilibrium system may perform better as a tracer of organic carbon export under low-flux conditions (in this case, <2.5 mmol C m^?2 d^?1) than under bloom conditions in an oligotrophic setting. With very few exceptions, the POC flux predictions calculated from the water-column ²¹⁰Po deficit were within a factor of 2 of the POC flux caught in surface-tethered sediment traps. However, we found higher correlation between size-fractionated particulate ²¹⁰Po activity and POC concentration in November 2006 (r=0.93) than in January (r=0.79) and during the spring bloom in March 2007 (r=0.80). We suggest that this is due to the ability of polonium to distinguish between bulk mass flux and organic carbon export under oligotrophic and lithogenic-driven flux regimes. Further, we found that the POC/Po ratio on particles was largely independent of size class between 10 and 100 μm (P=0.13) during each season, supporting the notion that export in this oligotrophic system is driven by sinking aggregates of smaller cells and not by large, individual cells.     

Actinian dominated intertidal mudflats: A new case of an extraordinary rare phenomenon from Southern Chile

Generally, estuarine intertidal mudflats constitute important nurseries for fish and foraging grounds for coastal birds by providing a plenitude of mollusks, worms, and crustaceans as prey, which in turn mostly feed on suspended and benthic microalgae, bacteria, and detritus. Despite the high productivity of such habitats, pronounced variability in both salinity and temperature results typically in low diversity. The only sea anemone reported from estuarine mud is the edwardsiid Nematostella vectensis Stephenson, 1935. It occurs widely in the northern hemisphere, and occasionally in extremely high density. Here we document another sea anemone from estuarine mud and muddy sand found in Southern Chile which has similar ecological attributes. Taxonomic confusion has impeded the reporting on this small but prominent member in a macrozoobenthic assemblage, the brooding Anthopleura hermaphroditica (Carlgren, 1899; Anthozoa: Actiniidae). It differs from N. vectensis by the presence of symbiotic algae. Average density under poly- to euhaline conditions in mud and muddy sand at around mid tide level was about 3 actinians per cm². An average abundance of 11,000 m^? 2, a biovolume of 487 cm³ m^? 2, and a biomass of 35.5 g dry organic weight m^? 2 were found in mud and muddy sand in two surveys 20 years apart. The mean fishing area of fully expanded individuals covers 42 ± 25 mm², corresponding to a circular area with a diameter of 7.3 ± 5.7 mm. Preliminary experiments indicate that associated benthos may be relegated to life below surface by the net of tentacles above the sediment. As no predators on A. hermaphroditica could be found on the mudflat, the success of this mixotrophic sea anemone may entail a trophic dead end.     

Continuous shipboard sampling of gas tension,oxygen and nitrogen

A novel shipboard gas tension device (GTD) that measures total dissolved air pressure in ocean surface waters is described and demonstrated. In addition, an improved method to estimate dissolved N₂ levels from simultaneous measurements of gas tension, dissolved O₂, water temperature, and salinity is described. Other than a flow-through plenum, the shipboard GTD is similar to the previously described moored-mode GTD (McNeil et al., 1995, Deep-Sea Research I 42, 819–826). The plenum has an integrated water-side screen to protect the membrane, and prevent the membrane from flexing in super-saturated near surface waters. The sampling scheme uses a well mixed and thermally insulated 15 L container that is flushed by the ship's seawater intake at a rate of 3–15 L min⁻¹. Dissolved gas sensors are placed inside this container and flushed with a small recirculation pump. Laboratory data that characterize the response of the modified GTD are presented. The modified GTD has a constant, isothermal, characteristic (e-folding) response time of typically 11±2 min at 20 °C. The response time decreases with increasing temperature and varies by ±35% over a temperature range of 5–35 °C. Results of field measurements, collected on the R.V. Brown between New York and Puerto Rico during September 2002, are presented, and provide the first look at co-variability in surface ocean N₂, O₂, and CO₂ levels over horizontal length scales of several kilometers. Dissolved N₂ concentrations decreased by approximately 16% as the ship sailed from the colder northern continental shelf waters, across the Gulf Stream, and into the warmer northwestern Atlantic Ocean. Historical database measurements, buoy time series, and satellite imagery, are used to aid interpretation of the dissolved gas levels.     

Air–sea CO2 fluxes in the north-eastern shelf of the Gulf of Cádiz (southwest Iberian Peninsula)

An intra-annual investigation of the fugacity of CO₂ (fCO₂) has been conducted in surface waters of the north-eastern shelf of the Gulf of Cádiz (SW Iberian Peninsula) in four cruises made in 2006 and 2007. Intra-annual variability of fCO₂ was assessed and is discussed in terms of mixing, temperature and biology. In the study area of the shelf, thermodynamic control over fCO₂ predominates from early May to late November, and this is opposite and similar in magnitude to the net biological effect. However, biological control over fCO₂ predominates during winter. The results suggest that surface waters in the coastal area are under-saturated with respect to atmospheric CO₂ during most of the year; therefore they represent a sink for atmospheric CO₂ between November and May (? 1.0 mmol m^? 2 day^? 1), but a weak source in June (1.3 mmol m^? 2 day^? 1). In contrast, the coastal ecosystems studied (the lower estuary of Guadalquivir Estuary and Bay of Cádiz) acted as a weak sink for atmospheric CO₂ during February (? 1.3 mmol m^? 2 day^? 1) and as a source between May and November (2.6 mmol m^? 2 day^? 1). The resulting mean annual CO₂ flux in the north-eastern shelf of the Gulf of Cádiz was ? 0.07 mol m^? 2 year^? 1 (? 0.2 mmol m^? 2 day^? 1), indicating that the area acts as a net sink on an annual basis.     

Persian Gulf response to a wintertime shamal wind event

The results from a～1 km resolution HYbrid Coordinate Ocean Model (HYCOM), forced by 1/2° Navy Operational Global Atmospheric Prediction System (NOGAPS) atmospheric data, were used in order to study the dynamic response of the Persian Gulf to wintertime shamal forcing. Shamal winds are strong northwesterly winds that occur in the Persian Gulf area behind southeast moving cold fronts. The period from 20 November to 5 December 2004 included a well defined shamal event that lasted 4–5 days. In addition to strong winds (16 m s^?1) the winter shamal also brought cold dry air (T_a=20 °C, q_a=10 g kg^?1) which led to a net heat loss in excess of 1000 W m^?2 by increasing the latent heat flux. This resulted in SST cooling of up to 10 °C most notably in the northern and shallower shelf regions. A sensitivity experiment with a constant specific humidity of q_a=15 g kg^?1 confirmed that about 38% of net heat loss was due to the air–sea humidity differences. The time integral of SST cooling closely followed the air–sea heat loss, indicating an approximate one-dimensional vertical heat balance. It was found that the shamal induced convective vertical mixing provided a direct mechanism for the erosion of stratification and deepening of the mixed layer by 30 m. The strong wind not only strengthened the circulation in the entire Persian Gulf but also established a northwestward flowing Iranian Coastal Current (ICC, 25–30 cm s^?1) from the Strait of Hormuz to about 52°E, where it veered offshore. The strongest negative sea level of 25–40 cm was generated in the northernmost portion of the Gulf while the wind setup against the coast of the United Arab Emirates established a positive sea level of 15–30 cm. The transport through the Strait of Hormuz at 56.2°E indicated an enhanced outflow of 0.25 Sv (Sv≡10⁶ m³ s^?1) during 24 November followed by an equivalent inflow on the next day.     

Seasonal and inter-annual biogeochemical variations in the Porcupine Abyssal Plain 2003–2005 associated with winter mixing and surface circulation

We present a 3-year multidisciplinary biogeochemical data set taken in situ at the Porcupine Abyssal Plain (PAP) time-series observatory in the Northeast Atlantic (49°N, 16.5°W; water depth ∼4850 m) for the period 2003 to 2005. The high-resolution year-round autonomous measurements include temperature, salinity, chlorophyll-a (derived from in situ chlorophyll-fluorescence) and inorganic nitrate, all at a nominal depth of 30 m on an Eulerian observatory mooring. This study compares these in situ time-series data with satellite chlorophyll-a data, regional data from a ship of opportunity, mixed-layer depth measurements from profiling Argo floats and lateral advection estimates from altimetry. This combined and substantial data set is used to analyse seasonal and inter-annual variability in hydrography and nitrate concentrations in relation to convective mixing and lateral advection. The PAP observatory site is in the inter-gyre region of the North Atlantic where convective mixing ranges from 25 m in the summer to over 400 m in winter when nutrients are supplied to the surface. Small inter-annual changes in the winter mixed layer can result in large changes in nitrate supply and productivity. However the decrease in maximum winter nitrate over the three-year period, from 10 to 7 mmol m⁻³, cannot be fully explained by convective mixing. Trajectories leading to the PAP site, computed from altimetry-derived geostrophic velocities, confirm that lateral advection cannot be ignored at this site and may be an important process along with convective mixing. Over the three years, there is an associated decrease in new production calculated from nitrate assimilation from 85.4 to 40.3±4.3 gCm⁻² a⁻¹. This confirms year-to-year variability in primary production seen in model estimates for the region. The continuous in situ dataset also shows inter-annual variation in the timing of the spring bloom due to variations in heat flux; the 2005 bloom occurred earlier than in 2004.