Nutrient and phytoplankton dynamics off the west coast of Vancouver Island during the 1997/98 ENSO event

Six research cruises were conducted off the west coast of Vancouver Island between April and October of 1997 and 1998 as part of the Canadian GLOBEC project to compare nutrient and phytoplankton dynamics between ENSO (1997) and non-ENSO (1998) years. Limited sampling also was conducted during three cruises in 1999. During the 1997 ENSO period, there was a shallow thermocline (∼10 m) that resulted in a shallower mixed layer, lower salinity and density, and stronger summer stratification. In general on the shelf, the 1997 growing season was characterized by higher nitrate (7.5 μM) and silicic acid (17 μM) concentrations, lower total chlorophyll (∼76 mg m⁻²), lower phytoplankton carbon biomass (0.2 mg C L⁻¹), and lower diatom abundance and biomass than in 1998. Phytoplankton assemblages were dominated by nanoplankton in 1997 and by diatoms in 1998. These results suggest that the 1997 ENSO was responsible for a reduction in the growth and biomass of larger phytoplankton cells. In mid-1998, the hydrographic characteristics off the west coast of Vancouver Island changed suddenly. The 1997 poleward transport of warm water reversed to an equatorward transport of coastal water in July 1998, which was accompanied by normal summer upwelling. During 1998, a large diatom bloom (mainly dominated by Chaetoceros debilis, Leptocylindrus danicus and to a lesser extent by Skeletomema and Pseudo-nitzschia sp.) was observed in July over the continental shelf. This large bloom resulted in chlorophyll concentrations of up to 400 mg m⁻², primary productivity of up to 11 g C m⁻² d⁻¹, and near undetectable dissolved nitrogen concentrations at some of the shelf stations in 1998. In contrast, during 1997, the sub-tropical waters that were advected over the slope, resulted in low chlorophyll a and primary productivity (generally <1 g C m⁻² d⁻¹). Therefore, there was a sharp contrast between the very high primary productivity on the shelf in July 1998, due to normal nutrient replenishment from summer upwelling and outflow from the Strait of Juan de Fuca, and the lower primary productivity during the 1997 ENSO year. During 1998, non-ENSO conditions resulted in phytoplankton biomass that was twice as high on the shelf as that measured in regions beyond the continental shelf of the west coast of Vancouver Island.     

Community structure across a large-scale ocean productivity gradient: Marine bird assemblages of the Southern Indian Ocean

Our objective was to understand how marine birds respond to oceanographic variability across the Southern Indian Ocean using data collected during an 16-day cruise (4–21 January 2003). We quantified concurrent water mass distributions, ocean productivity patterns, and seabird distributions across a heterogeneous pelagic ecosystem from subtropical to sub-Antarctic waters. We surveyed 5155 km and sighted 15,606 birds from 51 species, and used these data to investigate how seabirds respond to spatial variability in the structure and productivity of the ocean. We addressed two spatial scales: the structure of seabird communities across macro-mega scale (1000 s km) biogeographic domains, and their coarse-scale (10 s km) aggregation at hydrographic and bathymetric gradients. Both seabird density and species composition changed with latitudinal and onshore–offshore gradients in depth, water temperature, and chlorophyll-a concentration. The average seabird density increased across the subtropical convergence (STC) from 2.4 birds km⁻² in subtropical waters to 23.8 birds km⁻² in sub-Antarctic waters. The composition of the avifauna also differed across biogeographic domains. Prions (Pachyptila spp.) accounted for 57% of all sub-Antarctic birds, wedge-tailed shearwaters (Puffinus pacificus) accounted for 46% of all subtropical birds, and Indian Ocean yellow-nosed albatross (Thallasarche carteri) accounted for 32% of all birds in the STC. While surface feeders were the most abundant foraging guild across the study area, divers were disproportionately more numerous in the sub-Antarctic domain, and plungers were disproportionately more abundant in subtropical waters. Seabird densities were also higher within shallow shelf-slope regions, especially in sub-Antarctic waters, where large numbers of breeding seabirds concentrated. However, we did not find elevated seabird densities along the STC, suggesting that this broad frontal region is not a site of enhanced aggregation.     

Thermohaline variability and geostrophic circulation in the southern portion of the Gulf of California

Data from seven oceanographic cruises in the southern Gulf of California from 1997 to 2002 are used to describe the thermohaline variability and the geostrophic circulation. Baroclinic patterns exhibited spatial and temporal variability. A deepening of isotherms at the center of the section was evident in February 1999, suggesting anticyclonic flow. In May 1998 and November 1997, cyclonic flow was suggested by shoaling of isotherms at the center of the section. Other cruises showed alternating cores of flow into and out of the Gulf (August 1998, September 1997 and October 2002). Neither a seasonal nor a spatial pattern in geostrophic flows was apparent, suggesting that the exchange of waters between the cyclonic flow of Pescadero basin and the interior of the Gulf is complex. Relatively high salinities were recorded during most of the cruises indicating that Gulf of California Water (GCW) was present most of the year. Higher salinities were observed during winter and spring, although during summer, relatively high and low salinities were both observed as surface and subsurface cores. Temperature and salinity characteristics of California Current waters were observed only in August 1995 when they reached as far north as Cerralvo Island at ∼50 dbar. During El Niño conditions in November 1997, a mixed layer (∼70 dbar) and deepening of the thermocline (∼50 dbar) characterized anomalous conditions; during this cruise an asymmetric salinity pattern was observed with low salinities characteristic of Tropical Surface waters at the center and east of the section, while maximum salinities (34.9<S<35.0) and Gulf waters were located in an 80 km wide core next to the Baja California Sur shelf as far north as San Jose Island.     

Drift algae reduce foraging efficiency of juvenile flatfish

Although flatfish species utilise a wide range of habitats as adults, several species are confined to a very limited habitat as juveniles. Recruitment levels are dependent on the quality and quantity of these nursery areas and changes therein. In the Baltic Sea, these shallow environments are often subject to influxes of drifting macroalgae, which add structure to otherwise bare sandy substrate. Structure, such as vegetation, alters predator–prey interactions of a wide range of fauna and in an array of marine, freshwater, and terrestrial systems. The aim of our study was to assess the inhibition potential of drifting macroalgae on the foraging efficiency of juvenile flatfish (young of the year Scophthalmus maximus L., young of the year- and group 1 + Platichthys flesus L.) through a series of microcosm experiments. Our results show that foraging success is restricted by drift algae as predation efficiency of all predator species and size classes was negatively affected by the presence of macroalgae. Overall, there was a reduction in predation success by 80 ± 12% due to structural effects and/or the induced changes in water chemistry associated with the algae. Flatfish depend on shallow sandy areas as feeding and nursery grounds during their juvenile stage and frequently occurring macroalgal assemblages drastically change the features of the otherwise bare substrate, setting the stage for small-scale, localised processes potentially affecting population dynamics.     

Inter-cohort differences in spatial and temporal settlement patterns of young-of-the-year windowpane (Scophthalmus aquosus) in southern New Jersey

The timing and location of settlement of two cohorts (spring and fall) of windowpane (Scophthalmus aquosus) were identified based on collections from 64 sampling locations along a corridor from the lower estuary, through the inlet, and on to the adjacent inner continental shelf in southern New Jersey. Spatio-temporal patterns of settlement during 1989–1998 were determined based on capture location and timing, and eye migration stage. Spring-spawned windowpanes were collected in estuarine, inlet and ocean habitats as larvae, during settlement, and after settlement. Densities of spring-spawned larvae (∼2–10 mm standard length (SL)) peaked in May in all habitats (estuary, inlet, and ocean). Initial settlement of spring-spawned windowpane occurred during May in the inlet and ocean when fish had grown to ∼7–8 mm SL (mid-point of eye migration), but fish did not appear in demersal estuarine collections until June when they were larger and more developmentally advanced (∼24–32 mm SL; post-eye migration). A transitional settlement period, comprised of a progressive habitat shift from pelagic to demersal habitats, is proposed for the spring cohort to explain the observed patterns. Fall-spawned fish of all developmental stages and sizes were virtually absent from estuarine collections. Fall-spawned larval (∼2–10 mm SL) densities peaked in October in inlet and ocean habitats and fish began settling there during the same month at sizes similar to the spring cohort (∼7–8 mm SL). This research confirms that there are important cohort-specific and life-stage dependent differences in young-of-the-year (YOY) windowpane habitat use in southern New Jersey and perhaps in other east coast US estuaries. These differences may affect the overall contribution that each cohort makes to a given year class and thus, may have an important role in determining the recruitment dynamics of this species.     

Effects of solar radiation on dimethylsulfide cycling in the western Atlantic Ocean

The influence of solar radiation on springtime rates of photochemical and biological consumption of dimethylsulfide (DMS) in surface waters from the western Atlantic Ocean was examined by exposing 0.2 μm filtered and unfiltered surface seawater to natural sunlight at five depths in the upper 30 m. Parallel deck incubations of 0.2 μm filtered seawater under various long-pass optical filters were also carried out to aid in assessing the wavelength dependence of DMS photolysis. DMS photolysis rate constants for mid-day exposure (∼10:30–17:30 local time) to surface irradiance ranged from 0.026 to 0.086 h⁻¹ and were highest in coastal and shelf waters. Photolysis rate constants decreased with increasing irradiation depth, in accordance with the attenuation of ultraviolet radiation (UVR, 280–400 nm). Total DMS consumption rates (photochemical+biological) in unfiltered surface samples also decreased with increasing incubation depth and were larger than photolysis rates at nearly all depths and all stations. The decrease in photolysis rate constants with exposure depth was mirrored by biological DMS consumption rate constants that were severely inhibited at surface irradiances, and approached or exceeded dark rate constants at deeper exposure depths. Photolysis rates were 2–19 times greater than estimated biological consumption rates in the surface light exposed samples, while biological consumption rates were significantly larger than photolysis rates at incubation depths below the 1% light level for UV–B radiation (280–320 nm). Total DMS loss rates increased up to nine-fold with UVR exposure, but changes in DMS concentrations were not strongly correlated to light dose, presumably due to parallel, light-mediated DMS production processes. The primary loss process for DMS depended mainly on the depth interval considered and the attenuation of UVR; in general, photochemical removal dominated shallow layers characterized by high UV–B intensities, whereas biological removal dominated in deeper layers where UV–B was absent, but UV–A (320–400 nm) and visible (400–700 nm) light fluxes were still relatively high. These results demonstrate that UVR exposure significantly influences the spatial and temporal pattern of DMS production and loss processes, and ultimately the DMS flux to the atmosphere.     

Fish settlement in the ocean vs. estuary: Comparison of pelagic larval and settled juvenile composition and abundance from southern New Jersey,U.S.A.

To describe the larval and juvenile fish fauna and to evaluate the relative contribution of the ocean and the estuary as settlement areas for benthic species, we compared the composition and abundance of larval fish supply to that of recently settled juvenile fishes in both ocean and an adjacent estuary habitats in southern New Jersey. The study was conducted from May to November 1992 in the Great Bay–Little Egg Harbor estuary (<1–8 m sampling depth) and on the adjacent inner continental shelf in the vicinity of Beach Haven Ridge (8–16 m). During the study more larvae nearing settlement (postflexion) were captured in the estuary than in the ocean. Settlement occurred earlier in the estuary than in the ocean perhaps under the influence of earlier, seasonal warming of estuarine waters. There appeared to be two spatial patterns of settlement in the study area based on the dominant species (n = 17) represented by a sufficient number of individuals (n ≥ 25 individuals). There were species that primarily settle in the estuary, as represented by both estuarine residents (n = 3) and transients (n = 4), and those that settle in both the estuary and the ocean (n = 10). However, there were no species whose larvae were present in the estuary yet settle in the ocean. The fact that many of the species settle in both the estuary and the ocean indicates an overlap between these habitats because, at least for some species, these habitats may function in the same way. Further resolution of fish settlement patterns, and its influence on recruitment will need to rely on synoptic comparisons between estuaries and the ocean over multiple years.     

Population structure and species dynamics of Spisula solida,Diogenes pugilator and Branchiostoma lanceolatum along a temporal–spatial gradient in the south coast of Portugal

The population structure, dynamics and distribution of Spisula solida, Diogenes pugilator and Branchiostoma lanceolatum, common species in the south coast of Portugal, were studied in a spatial–temporal manner in order to understand the influence of cross-shore sediment transport and anthropogenic activities. Spisula solida is harvested commercially, whereas D. pugilator and B. lanceolatum are non-target species, with little information available on the population dynamics of these species. The study was performed in 2001–2002, and along a gradient of 100–5000 m from the coastline, corresponding to a depth gradient of between 1 and 32 m deep. Spisula solida was distributed preferentially at 3–12 m deep, and its distribution appeared to be influenced seasonally by the cross-shore sediment dynamics. Results suggested benthic recruitment in June. Some recruitments had no expression (year 2001) since adults were not present, which seems to be a direct impact of clams' fisheries. Diogenes pugilator showed preferential distribution at shallow depths, from 1.3 to 8 m. Results suggested recruitments every 4 months, in June, February and October. Branchiostoma lanceolatum showed the widest distribution, from 7 to 26 m deep. Recruitment seemed to start in June until October, when it attained an abundance peak of juveniles. For both non-target species no clear effects of the cross-shore dynamics or the clams' fisheries impact were visible.     

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

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

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

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

Seasonal dynamics of primary and new production in the northern South China Sea: The significance of river discharge and nutrient advection

Biochemical and productivity measurements and nutrient enrichment experiments were conducted on three cruises in summer and two cruises in winter on the shelf and the basin of the northern South China Sea (SCS) between 2001 and 2004. Phytoplankton production, in terms of depth-integrated new production (INP) or depth-integrated primary production (IPP), was higher in winter than in summer and on the shelf than in the basin. In winter, with deepening of the mixed layer, nitrate from the shallow nitracline that characterized the SCS waters was made available in the surface and supported the highest production of the year. Averaged INP measured in winter (0.25 g C m⁻² d⁻¹) was about twice the summer average (0.12 g C m⁻² d⁻¹) and was 0.19 g C m⁻² d⁻¹ on the shelf compared with 0.15 g C m⁻² d⁻¹ in the basin. In winter, average INP on the shelf was higher than the basin (0.34 versus 0.21 g C m⁻² d⁻¹); whereas in summer, averaged INP on the shelf (0.13 g C m⁻² d⁻¹) and the basin (0.11 g C m⁻² d⁻¹) were similar. While averaged IPP measured in the basin was higher in winter than in summer (0.53 versus 0.35 g C m⁻² d⁻¹), IPP on the shelf showed little temporal variation (0.82 in winter versus 0.84 g C m⁻² d⁻¹ in summer). Considerable spatial and inter-annual variation in production was measured in the shelf waters during summer, which could be linked to discharge volume and plume flow direction of the Zhujiang River. While the shelf waters in summer were mostly nitrogen starved or nitrogen and phosphorus co-limited, excessive river runoff may cause the nutritive state to shift to phosphorus deficiency. Waters with low surface salinities and high fluorescence from riverine mixing could be found extending from the Zhujiang mouth to as far as offshore southern Taiwan after a typhoon passed the northern SCS and brought heavy rainfall. Overall, both nutrient advection in winter and river discharge from the China coast in summer made new nitrogen available and shaped the dynamics of phytoplankton production in these oligotrophic waters.     

Does Encope emarginata (Echinodermata: Echinoidea) affect spatial variation patterns of estuarine subtidal meiofauna and microphytobenthos?

Foraging macrofauna, such as the sand dollar Encope emarginata, can modify sediment properties and affect spatial distribution patterns of microphytobenthos and meiobenthos at different spatial scales. We adopted a spatial hierarchical approach composed of five spatial levels (km, 100 s m, 10 s m, 1 s m and cm) to describe variation patterns of microphytobenthos, meiobenthos and sediment variables in shallow subtidal regions in the subtropical Paranaguá Bay (Southern Brazil) with live E. emarginata (LE), dead E. emarginata (only skeletons — (DE), and no E. emarginata (WE). The overall structure of microphytobenthos and meiofauna was always less variable at WE and much of variation at the scale of 100 s m was related to variability within LE and DE, due to foraging activities or to the presence of shell hashes. Likewise, increased variability in chlorophyll-a and phaeopigment contents was observed among locations within LE, although textural parameters of sediment varied mainly at smaller scales. Variations within LE were related to changes on the amount and quality of food as a function of sediment heterogeneity induced by the foraging behavior of sand dollars. We provide strong evidence that top-down effects related to the occurrence of E. emarginata act in synergy with bottom-up structuring related to hydrodynamic processes in determining overall benthic spatial variability. Conversely, species richness is mainly influenced by environmental heterogeneity at small spatial scales (centimeters to meters), which creates a mosaic of microhabitats.     

Gouge marks on deep-sea mud volcanoes in the eastern Mediterranean: Caused by Cuvier's beaked whales?

Enigmatic seafloor gouge marks at depths of 1700–2100 m have been observed from submersible during geological survey work studying mud volcanoes in the eastern Mediterranean Sea. The marks consist of a central groove (about 10 cm deep and 1–2 m long), superimposed on a broader bowl-shaped depression (1–2 m long by about 50 cm wide) with raised rims (up to 10 cm high) to either side of the central groove. We discuss the potential biological causes of these marks, and conclude that they are probably created by Cuvier's beaked whales (Ziphius cavirostris) during foraging dives to these depths. The mud volcanoes have a comparatively rich and diverse benthic ecology associated with methane-rich fluid seeps and thus could be the base of food chains that reach top predators like the deep-diving whales. The characteristic high acoustic backscatter of the mud volcanoes would facilitate their detection by the echolocation system of these whales.     

Nutrient gradients in the western North Atlantic Ocean: Relationship to microbial community structure and comparison to patterns in the Pacific Ocean

Studies of nitrogen and phosphorus dynamics in the oligotrophic surface waters of the western North Atlantic Ocean have been constrained because ambient concentrations are typically at or below the detection limits of standard colorometric methods, except during periods of deep vertical mixing. Here we report the application of high-sensitivity analytical methods—determinations of nitrate plus nitrite (N+N) by chemiluminescence and soluble reactive phosphorus (SRP) by the magnesium induced co-precipitation (MAGIC) protocol—to surface waters along a transect from the Sargasso Sea at 26°N through the Gulf Stream at 37°N, including sampling at the JGOFS Bermuda Atlantic Time-series Study (BATS) station. The results were compared with data from the BATS program, and the HOT station in the Pacific Ocean, permitting cross-ecosystem comparisons. Microbial populations were analyzed along the transect, and an attempt was made to interpret their distributions in the context of the measured nutrient concentrations.Surface concentrations of N+N and SRP during the March 1998 transect separated into 3 distinct regions, with the boundaries corresponding roughly to the locations of the BATS station (∼31°N) and the Gulf Stream (∼37°N). Although N+N and SRP co-varied, the [N+N] : [SRP] molar ratios increased systematically from ∼1 to 10 in the southern segment, remained relatively constant at ∼40–50 between 31°N and 37°N, then decreased again systematically to ratios <10 north of the Gulf Stream. Dissolved organic N (DON) and P (DOP) dominated (⩾90%) the total dissolved N (TDN) and P (TDP) pools except in the northern portion of the transect. The [DON] : [DOP] molar ratios were relatively invariant (∼30–60) across the entire transect.Heterotrophic prokaryotes (operationally defined as “bacteria”), Prochlorococcus, Synechococcus, ultra- and nanophytoplankton, cryptophytes, and coccolithophores were enumerated by flow cytometry. The abundance of bacteria was well correlated with the concentration of SRP, and that of the ultra- and nanophytoplankton was well correlated with the concentration of N+N. The only group whose concentration was correlated with temperature was Prochlorococcus, and its abundance was unrelated to the concentrations of nutrients measured at the surface.We combined our transect results with time-series measurements from the BATS site and data from select depth profiles, and contrasted these North Atlantic data sets with time-series of N and P nutrient measurements from a station in the North Pacific subtropical gyre near Hawaii [Hawaii Ocean Time-series (HOT) site]. Two prominent differences are readily observed from this comparison. The [N+N] : [SRP] molar ratios are much less than 16 : 1 during stratified periods in surface waters at the BATS site, as is the case at the HOT site year round. However, following deep winter mixing, this ratio is much higher than 16 : 1 at BATS. Also, SRP concentrations in the upper 100 m at BATS fall in the range 1–10 nM during stratified periods, which is at least one order of magnitude lower than at the HOT site. That two ecosystems with comparable rates of primary and export production would differ so dramatically in their nutrient dynamics is intriguing, and highlights the need for detailed cross ecosystem comparisons.     

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.     

Biogeochemical study of a coccolithophore bloom in the northern Bay of Biscay (NE Atlantic Ocean) in June 2004

The present paper synthesizes data obtained during a multidisciplinary cruise carried out in June 2004 at the continental margin of the northern Bay of Biscay. The data-set allows to describe the different stages of a coccolithophore bloom dominated by Emiliania huxleyi. The cruise was carried out after the main spring phytoplankton bloom that started in mid-April and peaked in mid-May. Consequently, low phosphate (PO₄ < 0.2 μM) and silicate (DSi < 2.0 μM) concentrations, low partial pressure of carbon dioxide (pCO₂) and high calcite saturation degree in surface waters combined with thermal stratification, probably favoured the blooming of coccolithophores. During the period of the year our cruise was carried out, internal tides induce enhanced vertical mixing at the continental shelf break leading to the injection of inorganic nutrients to surface waters that probably trigger the bloom. The bloom developed as the water-column stratified and as the water mass was advected over the continental shelf, following the general residual circulation in the area. The most developed phase of the bloom was sampled in a remote sensed high reflectance (HR) patch over the continental shelf that was characterized by low chlorophyll-a (Chl-a) concentration in surface waters (<1.0 μg L^?1), high particulate inorganic carbon (PIC) concentration (～8 μmol L^?1) and coccolithophore abundance up to 57 × 10⁶ cells L^?1. Transparent exopolymer particles (TEP) concentrations ranged between 15 and 75 μg C L^?1 and carbon content of TEP represented up to 26% of the particulate organic carbon (POC; maximum concentration of 15.5 μmol L^?1 in the upper 40 m). Integrated primary production (PP) ranged between 210 and 680 mg C m^?2 d^?1 and integrated calcification (CAL) ranged between 14 and 140 mg C m^?2 d^?1, within the range of PP and CAL values previously reported during coccolithophore blooms in open and shelf waters of the North Atlantic Ocean. Bacterial protein production (BPP) measurements in surface waters (0.3–0.7 μg C L^?1 h^?1) were much higher than those reported during early phases of coccolithophore blooms in natural conditions, but similar to those during peak and declining coocolithophorid blooms reported in mesocosms. Total alkalinity anomalies with respect to conservative mixing (ΔTA) down to ?49 μmol kg^?1 are consistent with the occurrence of biogenic precipitation of calcite, while pCO₂ remained 15–107 μatm lower than atmospheric equilibrium (372 μatm). The correlation between ΔTA and pCO₂ suggested that pCO₂ increased in part due to calcification, but this increase was insufficient to overcome the background under-saturation of CO₂. This is related to the biogeochemical history of the water masses due to net carbon fixation by the successive phytoplankton blooms in the area prior to the cruise, hence, the investigated area remained a sink for atmospheric CO₂ despite calcification.     

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

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

Arctic ocean shelf–basin interaction: An active continental shelf CO2 pump and its impact on the degree of calcium carbonate solubility

The Arctic Ocean has wide shelf areas with extensive biological activity including a high primary productivity and an active microbial loop within the surface sediment. This in combination with brine production during sea ice formation result in the decay products exiting from the shelf into the deep basin typically at a depth of about 150 m and over a wide salinity range centered around S ～33. We present data from the Beringia cruise in 2005 along a section in the Canada Basin from the continental margin north of Alaska towards the north and from the International Siberian Shelf Study in 2008 (ISSS-08) to illustrate the impact of these processes. The water rich in decay products, nutrients and dissolved inorganic carbon (DIC), exits the shelf not only from the Chukchi Sea, as has been shown earlier, but also from the East Siberian Sea. The excess of DIC found in the Canada Basin in a depth range of about 50–250 m amounts to 90±40 g C m^?2. If this excess is integrated over the whole Canadian Basin the excess equals 320±140×10¹² g C. The high DIC concentration layer also has low pH and consequently a low degree of calcium carbonate saturation, with minimum aragonite values of 60% saturation and calcite values just below saturation. The mean age of the waters in the top 300 m was calculated using the transit time distribution method. By applying a future exponential increase of atmospheric CO₂ the invasion of anthropogenic carbon into these waters will result in an under-saturated surface water with respect to aragonite by the year 2050, even without any freshening caused by melting sea ice or increased river discharge.     

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.     

Hydrography and circulation of the West Antarctic Peninsula Continental Shelf

The water mass structure and circulation of the continental shelf waters west of the Antarctic Peninsula are described from hydrographic observations made in March–May 1993. The observations cover an area that extends 900 km alongshore and 200 km offshore and represent the most extensive hydrographic data set currently available for this region. Waters above 100–150 m are composed of Antarctic Surface Water and its end member Winter Water. Below the permanent pycnocline is a modified version of Circumpolar Deep Water, which is a cooled and freshened version of Upper Circumpolar Deep Water. The distinctive signature of cold and salty water from the Bransfield Strait is found at some inshore locations, but there is little indication of significant exchange between Bransfield Strait and the west Antarctic Peninsula shelf. Dynamic topography at 200 m relative to 400 m indicates that the baroclinic circulation on the shelf is composed of a large, weak, cyclonic gyre, with sub-gyres at the northeastern and southwestern ends of the shelf. The total transport of the shelf gyre is 0.15 Sv, with geostrophic currents of order 0.01 m s^-1. A simple model that balances across-shelf diffusion of heat and salt from offshore Upper Circumpolar Deep Water with vertical diffusion of heat and salt across the permanent pycnocline into Winter Water is used to explain the formation of the modified Circumpolar Deep Water that is found on the shelf. Model results show that the observed thermohaline distributions across the shelf can be maintained with a coefficient of vertical diffusion of 10^-4 m² s^-1 and horizontal diffusion coefficients for heat and salt of 200 and 1200 m² s^-1, respectively. When the effects of double diffusion are included in the model, the required horizontal diffusion coefficients for heat and salt are 200 and 400 m² s^-1, respectively.