The structure of zooplankton communities,in the 2 to 2000 μm size range,in the Arabian Sea during and after the SW monsoon, 1994

Zooplankton communities, studied in the surface mixed layer on a 1600 m transect across the Arabian Sea, were found to differ in their temporal and spatial response to seasonal forcing. The transect studied, spanned seasonally eutrophic upwelling, mesotrophic downwelling and aseasonal oligotrophic waters. The nano- and microzooplankton communities constituted a relatively constant compartment in the tropical monsoon ecosystem, whilst the mesozooplankton showed a clear response to both upwelling and season. The heterotrophic nanoflagellates were concentrated in the surface mixed layer, except in the eutrophic upwelling waters of the SW monsoon. They reached maximum cell concentrations of 855 ml^-1 during the SW monsoon and a maximum biomass of 8.4 mg C m^-3 during the intermonsoon. Nanozooplankton standing stocks, in the surface mixed layer, ranged between 7 and 333 mg C m^-2, with highest stocks found during the intermonsoon. The microzooplankton community was dominated by Protozoa, particularly aloricate ciliates and heterotrophic dinoflagellates, which accounted for up to 99% in terms of numbers and up to 71% of the biomass. Sarcodines and metazoan nauplii were recorded in lower numbers (<400 l^-1). The microzooplankton were also concentrated in the surface mixed layer during both periods, except in the eutrophic coastal waters during the SW monsoon, when relatively high biomass values were found below the mixed layer depth. Their standing stocks, in the surface mixed layer, ranged between 50 and 182 mg C m^-2, with the highest concentration found in the mesotrophic offshore waters during the late monsoon period. Total mesozooplankton standing stocks, in the surface 100 m, decreased with distance from the coastal to offshore waters and between seasons, decreasing from 1248 to 238 mg C m^-2 during the late SW monsoon and 656–89 mg C m^-2 during the following intermonsoon. The largest size class, of 1000–2000 μm sized organisms, dominated throughout except at the oligotrophic station during the intermonsoon period, when the smallest class, of 200–500 μm, were more important. The shift in size structure from large to small zooplankton occurred in response to a shift in dominance from large to small phytoplankton cells both spatially, along a eutrophic–oligotrophic gradient, and seasonally. These responses are a result of the physical forcing associated with the monsoon seasons in the Arabian Sea.     

Phytoplankton community structure in the Arabian Sea during and after the SW monsoon, 1994

Diatoms, dinoflagellates, coccolithophores, nanoflagellates, picophytoplankton and procaryote algae (Synechococcus spp. and prochlorophytes) were quantified by microscopy and flow cytometry, and their biomass determined, at 12 stations along a 1600 km transect across the Arabian Sea at the end of the SW monsoon in September, and during the inter-monsoon period of November/December 1994. The transect spanned contrasting oceanic conditions that varied from seasonally eutrophic, upwelling waters through mesotrophic, downwelling waters to permanently oligotrophic, stratified waters. The overall diversity of diatoms, dinoflagellates and coccolithophores along the transect was not significantly different between the SW monsoon and inter-monsoon. However, diatoms showed greatest diversity during the SW monsoon and coccolithophores were most diverse during the inter-monsoon. Integrated phytoplankton standing stocks during the SW monsoon ranged from 3 to 9 g C m^-2 in the upwelling eutrophic waters, from 3 to 5 g C m^-2 in downwelling waters, and from 1 to 2 g C m^-2 in oligotrophic waters. Similar phytoplankton standing stocks were found in oligotrophic waters during the inter-monsoon, but were ca. 40% lower compared to the SW monsoon in the more physically dynamic waters. Phytoplankton abundance and biomass was dominated by procaryote taxa. Synechococcus spp. were abundant (often >10⁸ cells l^-1) during both the SW monsoon and inter-monsoon, where the nitrate concentration was ⩾0.1 μ mol l^-1, and often dominated the phytoplankton standing stocks. Prochlorophytes were restricted to oligotrophic stratified waters during the SW monsoon period but were found at all stations along the transect during the inter-monsoon, dominating the phytoplankton standing stocks (>40%) in the oligotrophic region during this period. Of the nano- and micro-phytoplankton, only diatoms contributed significantly to phytoplankton standing stocks, and then only in near-shore upwelling waters during the SW monsoon. There were significant changes in the temporal composition of the phytoplankton community. In nearshore waters a mixed community of diatoms and Synechococcus spp. dominated during the SW monsoon. This gave way to a community dominated by Synechococcus spp. in the intermonsoon. In the downwelling zone, a Synechococcus spp. dominated community was replaced by a mixed procaryote community of Synechococcus spp. and prochlorophytes. In the oligotrophic stratified waters, the mix of procaryote algae was replaced by one dominated by prochlorophytes alone.     

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.     

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.     

Zooplankton herbivory in the Arabian Sea during and after the SW monsoon, 1994

Microzooplankton herbivory in the Arabian Sea was measured using dilution experiments towards the end of the SW monsoon in September and during the intermonsoon to NE monsoon period in November–December 1994. Microzooplankton grazing resulted in a turnover of phytoplankton stocks that ranged from 11 to 49% per day. This was equivalent to grazing fluxes of between 1 and 17 mg C m^-3 d^-1. Depth-integrated microzooplankton herbivory ranged between 161 and 415 mg C m^-2 d^-1 during the SW monsoon cruise, and between 110 and 407 mg C m^-2 d^-1 during the intermonsoon period. Microzooplankton grazed between 4 and 60% of daily primary production, with higher percentages found during the intermonsoon season. Phytoplankton growth rates during the SW monsoon ranged from 0.3 to 1.8 d^-1, with lower values in upwelling waters and higher values in downwelling and oligotrophic areas. During the intermonsoon period, phytoplankton growth was more uniform across the basin and averaged 0.68±0.15 d^-1. Microzooplankton abundance in experimental samples varied between 2800 and 16 162 cells l^-1, equivalent to a biomass of between 1.1 and 7.2 mg C m^-3. The mean cell carbon content of microzooplankton was similar in both periods and ranged from 0.33 to 0.55 ng C cell^-1. Microzooplankton were smallest in downwelling waters and largest in oligotrophic waters. Average clearance rates in those taxa that took up fluorescently-labelled algae ranged from 0.2 to 14 μl ind^-1 hr^-1. Average mesozooplankton grazing rates, derived from biomass data, varied from 19 to 92 mg C m^-2 d^-1; these rates accounted for removal of between 4 and 12% of the daily primary production. Mesozooplankton herbivory was most pronounced in upwelling and downwelling waters and reduced in stratified oligotrophic waters during the SW monsoon period. Microzooplankton herbivory was greater than the average mesozooplankton herbivory at all stations, during both the SW monsoon and intermonsoon periods.     

Physical-biological coupling in the Algerian Basin (SW Mediterranean): Influence of mesoscale instabilities on the biomass and production of phytoplankton and bacterioplankton

The biomass and production of phytoplankton and bacterioplankton was investigated in relation to the mesoscale structures found in the Algerian Current during the ALGERS'96 cruise (October 1996). Biological determinations were carried out in three transects between 0° and 2°E aimed at crossing a so-called event, formed by a coastal anticyclonic eddy associated with an offshore cyclonic eddy to the west. The concentration of chlorophyll a (Chl) was maximum (>1.2 mg m⁻³) within the cyclonic eddy and at the frontal zones between the Modified Atlantic Water (MAW) of the Algerian Current and the Mediterranean waters further north. Chl (total and >2 μm) was significantly correlated with proxies of nutrient flux into the upper layers. Autotrophic picoplankton and heterotrophic bacterial abundance and production presented clear differences between MAW and Mediterranean water, with higher values at those stations under the influence of the Algerian Current. In general, greater differences were observed in production than in biomass variables. The photosynthetic parameters (derived from P–E relationships) and integrated primary production (range 189–645 mg m⁻² d⁻¹) responded greatly to the different hydrological conditions. The mesoscale phenomena inducing fertilization caused a 2 to 3-fold increase in primary production rates. The relatively high values found within the cyclonic eddy suggest that, although short-lived in comparison with anticyclonic eddies, these eddies may produce episodic increases of biological production not accounted for in previous surveys in the region.     

Latitudinal distribution of microbial plankton abundance,production, and respiration in the Equatorial Atlantic in autumn 2000

Phytoplankton and bacterial abundance, size-fractionated phytoplankton chlorophyll-a (Chl-a) and production together with bacterial production, microbial oxygen production and respiration rates were measured along a transect that crossed the Equatorial Atlantic Ocean (10°N–10°S) in September 2000, as part of the Atlantic Meridional Transect 11 (AMT 11) cruise. From 2°N to 5°S, the equatorial divergence resulted in a shallowing of the pycnocline and the presence of relatively high nitrate (>1 μM) concentrations in surface waters. In contrast, a typical tropical structure (TTS) was found near the ends of the transect. Photic zone integrated ¹⁴C primary production ranged from ∼200 mg C m⁻² d⁻¹ in the TTS region to ∼1300 mg C m⁻² d⁻¹ in the equatorial divergence area. In spite of the relatively high primary production rates measured in the equatorial upwelling region, only a moderate rise in phytoplankton biomass was observed as compared to nearby nutrient-depleted areas (22 vs. 18 mg Chl-a m⁻², respectively). Picophytoplankton were the main contributors (>60%) to both Chl-a biomass and primary production throughout the region. The equatorial upwelling did not alter the phytoplankton size structure typically found in the tropical open ocean, which suggests a strong top-down control of primary producers by zooplankton. However, the impact of nutrient supply on net microbial community metabolism, integrated over the euphotic layer, was evidenced by an average net microbial community production within the equatorial divergence (1130 mg C m⁻² d⁻¹) three-fold larger than net production measured in the TTS region (370 mg C m⁻² d⁻¹). The entire region under study showed net autotrophic community metabolism, since respiration accounted on average for 51% of gross primary production integrated over the euphotic layer.     

Modelling phytoplankton deposition to Chesapeake Bay sediments during winter–spring: interannual variability in relation to river flow

The often-rapid deposition of phytoplankton to sediments at the end of the spring phytoplankton bloom is an important component of benthic–pelagic coupling in temperate and high latitude estuaries and other aquatic systems. However, quantifying the flux is difficult, particularly in spatially heterogeneous environments. Surficial sediment chlorophyll-a, which can be measured quickly at many locations, has been used effectively by previous studies as an indicator of phytoplankton deposition to estuarine sediments. In this study, surficial sediment chlorophyll-a was quantified in late spring at 20–50 locations throughout Chesapeake Bay for 8 years (1993–2000). A model was developed to estimate chlorophyll-a deposition to sediments using these measurements, while accounting for chlorophyll-a degradation during the time between deposition and sampling. Carbon flux was derived from these estimates via C:chl-a = 75.Bay-wide, the accumulation of chlorophyll-a on sediments by late spring averaged 171 mg m⁻², from which the chlorophyll-a and carbon sinking fluxes, respectively, were estimated to be 353 mg m⁻² and 26.5 gC m⁻². These deposition estimates were ∼50% of estimates based on a sediment trap study in the mid-Bay. During 1993–2000, the highest average chlorophyll-a flux was in the mid-Bay (248 mg m⁻²), while the lowest was in the lower Bay (191 mg m⁻²). Winter–spring average river flow was positively correlated with phytoplankton biomass in the lower Bay water column, while phytoplankton biomass in that same region of the Bay was correlated with increased chlorophyll-a deposition to sediments. Responses in other regions of the Bay were less clear and suggested that the concept that nutrient enrichment in high flow years leads to greater phytoplankton deposition to sediments may be an oversimplification. A comparison of the carbon flux associated with the deposition of the spring bloom with annual benthic carbon budgets indicated that the spring bloom did not contribute a disproportionately large fraction of annual carbon inputs to Chesapeake Bay sediments. Regional patterns in chlorophyll-a deposition did not correspond with the strong regional patterns that have been found for plankton net community metabolism during spring.     

Light absorption by phytoplankton,non-algal particles and dissolved organic matter at the Patagonia shelf-break in spring and summer

Satellite image studies and recent in situ sampling have identified conspicuous phytoplankton blooms during spring and summer along the Patagonia shelf-break front. The magnitudes and spectral characteristics of light absorption by total particulate matter (phytoplankton and detritus) and colored dissolved organic matter (CDOM) have been determined by spectrophotometry in that region for spring 2006 and late summer 2007 seasons. In spring, phytoplankton absorption was the dominant optical component of light absorption (60–85%), and CDOM showed variable and important contributions in summer (10–90%). However, there was a lack of correlation between phytoplankton biomass (chlorophyll-a concentration or [chl a]) and the non-algal compartment in both periods. A statistically significant difference was found between the two periods with respect to the CDOM spectral shape parameter (S_cdom), with means of 0.015 (spring) and 0.012 nm^?1 (summer). Nonetheless, the mean S_cdm values, which describe the slope of detritus plus CDOM spectra, did not differ between the periods (average of 0.013 nm^?1). Phytoplankton absorption values in this work showed deviations from mean parameterizations in previous studies, with respect to [chl a], as well as between the two study periods. In spring, despite the microplankton dominance, high specific absorption values and large dispersion were found (a*_ph(440)=0.04±0.03 m² mg [chl a]^?1), which could be attributed to an important influence of photo-protector accessory pigments. In summer, deviations from general trends, with values of a*_ph(440) even higher (0.09±0.02 m² mg [chl a]^?1), were due to the dominance of small cell sizes and also to accessory pigments. These results highlight the difficulty in deriving robust relationships between chlorophyll concentration and phytoplankton absorption coefficients regardless of the season period. The validity of a size parameter (S_f) derived from the absorption spectra has been demonstrated and was shown to describe the size structure of phytoplankton populations, independently of pigment concentration, with mean values of 0.41 in spring and 0.72 in summer. Our results emphasize the need for specific parameterization for the study region and seasonal sampling approach in order to model the inherent optical properties from water reflectance signatures.     

Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Ectoenzymatic activity in surface waters: A transect from the Mediterranean Sea across the Indian Ocean to Australia

The activities of two hydrolytic enzymes (leucine aminopeptidase and β glucosidase), belonging to the particle-bound enzymatic fraction, were measured in open-sea surface waters. Samples were collected along a transect crossing the Indian Ocean during the early NW monsoon period (November and December 2001). The latitudinal pattern of the ectoenzymatic activities highlighted a generally increasing trend of glycolysis approaching the equator, with significantly higher β glucosidase activity (0.79–3.00 nmol l⁻¹ h⁻¹) within the latitudinal range from 12°N to 16°S. In this area, the surface waters coming from the Indonesian Throughflow and the Bay of Bengal carry a considerable quantity of carbohydrates (38.9–41.9 μg l⁻¹), which stimulated glycolytic activity and its cell-specific rates scaled to bacterial abundance. On the other hand, in the Central Indian Ocean, the proteolytic activity was considerable (0.91–2.03 nmol l⁻¹ h⁻¹), although the particulate proteins did not show significant increases and the dissolved proteinlike signal was one of the lowest of the entire transect (0.7 mg l⁻¹ on average compared to the 1.4–1.6 mg l⁻¹ of the adjacent areas). Therefore, in this area, the two ectoenzymes studied did not respond to the same stimulatory effect (namely the specific substrate concentrations). The time needed for the hydrolysis of macromolecules within the particulate and dissolved organic substrate fractions, although these measures are affected by a number of assumptions starting with the potential nature of the ectoenzymatic determinations, confirms these observations. The Central Indian Ocean displayed the lowest values, from 8 to 26 days for particulate and dissolved organic carbon, respectively. As observed in the equatorial areas of the Atlantic Ocean, the relevant degradation activity of the central area of the Indian Ocean Basin suggests a notable heterotrophy based on a faster turnover of organic substrates.     

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

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

Nitrogen assimilation and the f-ratio in the northwestern Indian Ocean during an intermonsoon period

Rates of nitrogen assimilation by phytoplankton were measured at 13 stations along a transect in the northwestern Indian Ocean, from the Gulf of Oman, southwards to approximately 8°N, during November and December 1994. Nitrate (NO₃), ammonium (NH₄) and urea assimilation were measured using simulated in situ ¹⁵N incubation techniques. These measurements were supported by simultaneous rate measurements of primary production using ¹⁴C incubation techniques and detailed vertical distributions of temperature and chlorophyll concentrations. Euphotic zone integrated nitrogen assimilation rates varied between 1.1 and 23.6 mmol N m^-2 day^-1, with generally higher rates occurring at the northern and southern ends of the transect. At the majority of stations ammonium was the preferred nitrogen substrate assimilated; the average integrated assimilation rate of ammonium being 3.7 mmol N m^-2 day^-1 compared to 1.6 and 1.8 mmol N m^-2 day^-1 for urea and nitrate respectively. This general preference is reflected in the low f-ratios, which were ⩽0.52 for all stations and in the relative preference indices (RPI) values which were consistently >1 for ammonium and <1 for nitrate. A further examination of the data has lead to an apparent partitioning of the northwestern Indian Ocean into 2 regions; a region north of 17°30′N and a region south of this, to about 8°N. This division is based on: (i) the relationship between the f-ratio and ambient nitrate levels; (ii) nitrogen assimilation and primary production and (iii) the biomass distribution. It is suggested that this partitioning should be investigated further with the development of biogeochemical provinces in mind and the estimation of f-ratios on much larger, horizontal scales.     

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.     

Nitrogen isotopic composition of sinking particles from the southern Bay of Bengal: Evidence for variable nitrogen sources

An 8-year record of N fluxes and δ¹⁵N of sinking particles from the deep southern Bay of Bengal, northern Indian Ocean, is presented. Fluxes and δ¹⁵N vary between ∼0.1 and 3 mg m⁻² day⁻¹ and ∼2‰ and 8‰, respectively. The seasonal variation is determined mainly by oceanographic processes coupled to the Indian monsoon system. The annual pattern of δ¹⁵N is characterized by minima during spring intermonsoon (∼March–May), when nutrient inputs to the euphotic zone should be low because of stratification, and lighter nitrate/particulate matter is expected to be advected from the central Bay. Highest δ¹⁵N are associated with peak fluxes during southwest monsoon (∼June–September), when the southern Bay comes under the influence of the SW monsoon current, which appears to advect particulate matter with distinctly higher δ¹⁵N. The impact of this process, however, varies interannually under the influence of factors such as ENSO and the Indian Dipole Mode. Weakened advection leads to relatively low N fluxes and reduced δ¹⁵N. The data highlight the necessity of multi-annual studies to comprehend the natural variability of a system.     

Diatom biomass and productivity in oceanic and plume-influenced waters of the western tropical Atlantic ocean

Whereas diatoms (class Bacillariophyceae) often dominate phytoplankton taxa in the Amazon estuary and shelf, their contribution to phytoplankton dynamics and impacts on regional biogeochemistry are poorly understood further offshore in the western tropical Atlantic Ocean (WTAO). Thus, relative contribution of diatoms to phytoplankton biomass and primary production rates and associated environmental conditions were quantified during three month-long cruises in January–February 2001, July–August 2001, and April–May 2003. The upper water column was sampled at 6 light depths (100%, 50%, 25%, 10%, 1% and 0.1% of surface irradiance) at 64 stations between 3° and 14°N latitude and 41° and 58°W longitude. Each station was categorized as ‘oceanic’ or ‘plumewater’, based on principal component analysis of eight physical, chemical and biological variables. All stations were within the North Brazil Current, and plumewater stations were characterized by shallower mixed layers with lower surface salinities and higher dissolved silicon (dSi) concentrations than oceanic stations. The major finding was a much greater role of diatoms in phytoplankton biomass and productivity at plumewater stations relative to oceanic stations. Mean depth-integrated bSi concentrations at the plumewater and oceanic stations were 14.2 and 3.7 mmol m⁻², respectively. Mean depth-integrated SiP rates at the plumewater and oceanic stations were 0.17 and 0.02 mmol m⁻² h⁻¹, respectively. Based on ratios of SiP and PP rates, and typical Si:C ratios, diatoms contributed on average 29% of primary productivity at plumewater stations and only 3% of primary productivity at oceanic stations. In contrast, phytoplankton biomass (as chlorophyll a concentrations) and primary production (PP) rates (as ¹⁴C uptake rates) integrated over the euphotic zone were not significantly different at plumewater and oceanic stations. Chlorophyll a concentrations ranged from 8.5 to 42.4 mg m⁻² and 4.0 to 38.0 mg m⁻² and PP rates ranged from 2.2 to 11.2 mmol m⁻² h⁻² and 1.8 to 10.8 mmol m⁻² h⁻² at plumewater and oceanic stations, respectively. A conservative estimate of annual integrated SiP in offshore waters of Amazon plume between April and August is 0.59 Tmol Si, based on mean SiP rates in plumewaters and satellite-derived estimates of the area of the Amazon plume. In conclusion, river plumewaters dramatically alter the silicon dynamics of the WTAO, forming extensive diatom-dominated phytoplankton blooms that may contribute significantly to the global Si budget as well as contributing to energy and matter flow off of the continental shelf.     

Picophytoplankton in the equatorial Pacific: vertical distributions in the warm pool and in the high nutrient low chlorophyll conditions

Abundance distribution and cellular characteristics of picophytoplankton were studied in two distinct regions of the equatorial Pacific: the western warm pool (0°, 167°E), where oligotrophic conditions prevail, and the equatorial upwelling at 150°W characterized by high-nutrient low-chlorophyll (HNLC) conditions. The study was done in September–October 1994 during abnormally warm conditions. Populations of Prochlorococcus, orange fluorescing Synechococcus and picoeukaryotes were enumerated by flow cytometry. Pigment concentrations were studied by spectrofluorometry. In the warm pool, Prochlorococcus were clearly the dominant organisms in terms of cell abundance, estimated carbon biomass and measured pigment concentration. Integrated concentrations of Prochlorococcus, Synechococcus and picoeukaryotes were 1.5×10¹³, 1.3×10¹¹ and 1.5×10¹¹ cells m⁻², respectively. Integrated estimated carbon biomass of picophytoplankton was 1 g m⁻², and the respective contributions of each group to the biomass were 69, 3 and 28%. In the HNLC waters, Prochlorococcus cells were slightly less numerous than in the warm pool, whereas the other groups were several times more abundant (from 3 to 5 times). Abundance of Prochlorococcus, Synechococcus and picoeukaryotes were 1.2×10¹³, 6.2×10¹¹ and 5.1×10¹¹ cells m⁻², respectively. The integrated biomass was 1.9 g C m⁻². Prochlorococcus was again the dominant group in terms of abundance and biomass (chlorophyll, carbon); the respective contributions of each group to the carbon biomass were 58, 7 and 35%. In the warm pool the total chlorophyll biomass was 28 mg m⁻², 57% of which was divinyl chlorophyll a. In the HNLC waters, the total chlorophyll biomass was 38 mg m⁻², 44% of which was divinyl chlorophyll a. Estimates of Prochlorococcus, Synechococcus and picoeukaryotes cell size were made in both hydrological conditions.     

Island mass effect in the Juan Fernández Archipelago (33°S), Southeastern Pacific

Spatial and temporal variability of the island mass effect (IME; defined as local increases of phytoplankton associated with the presence of islands) at the Juan Fernández Archipelago (JFA) is analyzed using chlorophyll-a (Chl-a) satellite data, altimetry, sea surface temperature, wind, geostrophic currents and net heat flux over a ten year period (2002–2012). The the JFA islands (Robinson Crusoe-Santa Clara (RC-SC) and Alejandro Selkirk (AS)) present wakes with significant Chl-a increases, mainly during spring time. These wakes can reach Chl-a values of one order of magnitude higher (~1 mg m⁻³) than the surrounding oligotrophic waters (<0.1 mg m⁻³). The wakes are similar to von Kármán vortex streets which have been used to explain the impact of IME on Chl-a increases in numerical models. The wakes are formed from a high productivity area in the lee of the island, extending to the oceanic region as high Chl-a patches associated with submesoscale eddies that are detached from the islands and connected by less-productive zones. This pattern coincides with previous models that predict the effects of island-generated flow perturbations on biological production variability. The IME is a recurrent feature of islands that has even been observed in decadal average fields. In such average fields, the Chl-a values in RC-SC and AS islands can exceed values found in a Control Zone (a zone without islands) by ~50% and 30%, respectively. Seasonal and interannual variability reveals that, as a consequence of the IME, the winter Chl-a maximum associated with the development of winter convection and mesoscale eddies that propagate from the continental zone, promote that the Chl-a maximum extends towards spring. The IME has an impact on the island on both a local as well as a more regional scale that affects an area of ~40,000 km² (1°Latitude×4°Longitude) centered on the islands. The transport of high productivity patches associated with submesoscale eddies may be responsible for IME propagation at a regional scale. Around the islands, the presence of a weak oceanic incident flow and strong and recurrent wind-wakes, suggest that the generation of Chl-a wakes result from a combined effect between both forcings.