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.     

Stock and losses of trace metals from salt marsh plants

Pools of Zn, Cu, Cd and Co in the leaf, stem and root tissues of Sarcocornia fruticosa, Sarcocornia perennis, Halimione portulacoides and Spartina maritima were analysed for a Tagus estuary (Portugal) salt marsh. Pools of Cu and Cd in the salt marsh were higher in spring/summer, indicating a net uptake of these metals during the growing season. Standing stocks of Zn, Cu, Cd and Co in the leaf and stem biomass of S. fruticosa, S. perennis and H. portulacoides showed a strong seasonal variation, with higher values recorded in autumn. The metal-containing leaves and stems that shed in the autumn become metal-containing detritus. The amount of this material washed out from the total marsh area (200 ha) was estimated as 68 kg of Zn, 8.2 kg of Cu, 13 kg of Co and 0.35 kg of Cd. The high tidal amplitude, a branched system of channels and semi-diurnal tidal cycle greatly favour the export of the organic detritus to adjoining marsh areas.     

Comparative study on microphytobenthic pigments of muddy and sandy intertidal sediments of the Tagus estuary

The abundance and distribution of microphytobenthic pigments determined by HPLC (chlorophylls and carotenoids) were compared between muddy and sandy sediments of the Tagus estuary (Portugal). In the two types of sediment, with similar periods of illuminated emersion, chlorophyll a concentrations on a per area basis (mg m⁻²) were comparable (down to 2 mm). Pigment analysis also revealed similar microphytobenthic communities in terms of algal classes. Diatoms were the dominant microalgae, but cyanophytes, euglenophytes and phanerogam debris were also present. For both muddy and sandy sediments, microphytobenthic biomass showed a high level of variability both within and between two consecutive years. Microphytobenthos was highly stratified in the mud, with most of the chlorophyll a occurring in the top 500 μm. In the sand, relatively constant concentrations were found throughout the sediment profile down to 3 mm. This is probably related to deeper light penetration in sandy sediment and/or increased physical mixing caused by invertebrate activity or overlying currents, leading to the burial of an important fraction of the microphytobenthic cells. Differences observed in the intensity of sediment coloration of muddy and sandy sediments might have resulted from the different vertical distribution of benthic biomass.     

Critical thermal maxima of common rocky intertidal fish and shrimps — A preliminary assessment

Rocky shore ecosystems are considered sentinels of climate warming because they are in close contact with the atmosphere and their shallow waters present low thermal inertia. Concerns on the vulnerability of rocky shore species subject to climate warming make the investigation of their thermal tolerance an urgent topic. The aim of this study was to determine the upper thermal limits of species that are common in tidal pools of rocky shore ecosystems of the Northeast Atlantic. The method used was the Critical Thermal Maximum (CTMax), which allowed the ranking of species in terms of their upper thermal limits as follows: Coryphoblennius galerita (32.0 °C), Palaemon serratus (33.0 °C), Gobius paganellus (33.1 °C), Palaemon elegans (33.4 °C), Lipophrys pholis (33.9 °C) and Paralipophrys trigloides (35.0 °C). Intraspecific variability was always lower than 2%.     

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.     

A re-appraisal of the total biomass and annual production of Antarctic krill

Despite much research on Euphausia superba, estimates of their total biomass and production are still very uncertain. Recently, circumpolar krill databases, combined with growth models and revisions in acoustics have made it possible to refine previous estimates. Net-based databases of density and length frequency (KRILLBASE) yield a summer distributional range of ～19×10⁶ km² and a mean total abundance of 8×10¹⁴ post-larvae with biomass of 379 million tonnes (Mt). These values are based on a standardised net sampling method but they average over the period 1926–2004, during which krill abundance has fluctuated. To estimate krill biomass at the end of last century we combined the KRILLBASE map of relative krill density around Antarctica with an acoustics-derived biomass estimate of 37.3 Mt derived for the Scotia Sea area in 2000 by the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR). Thus the CCAMLR 2000 survey area contains 28% of the total stock, with total biomass of ～133 Mt in January–February 2000. Gross postlarval production is estimated conservatively at 342–536 Mt yr^?1, based on three independent methods. These are high values, within the upper range of recent estimates, but consistent with the concept of high energy throughput for a species of this size. The similarity between the three production estimates reflects a broad agreement between the three growth models used, plus the fact that, for a given population size, production is relatively insensitive to the size distribution of krill at the start of the growth season. These production values lie within the envelope of what can be supported from the Southern Ocean primary production system and what is required to support an estimated predator consumption of 128–470 Mt yr^?1. Given the range of recent acoustics estimates, plus the need for precautionary management of the developing krill fishery, our net-based data provide an alternative estimate of total krill biomass.     

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.     

Depth distribution of algal species on the deep insular fore reef at Lee Stocking Island,Bahamas

Deep-water benthic algal composition and cover were studied with a submersible on the deep fore reef of Lee Stocking Island, Bahamas, from 45 to 150 m. Algal cover decreased from 57% to 16% over this depth range. Although there was substantial overlap in depth distributions, different species or groups of species dominated benthic cover at different depths. Lobophora and Halimeda copiosa co-dominated the fore reef from 45 to 60 m. A Corallinales/Peyssonnelia group was abundant from 60 to 120 m. The Corallinales/Peyssonnelia group shared dominance with Ostreobium between 90 and 120 m. Ostreobium was the only alga observed below 150 m and remained abundant below 200 m. Movement of sand down the fore reef is recognized as having substantial influence on algal cover.     

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.     

Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

Mesozooplankton in the Arctic Ocean in summer

The biomass, species and chemical composition of the mesozooplankton and their impact on lower food levels were estimated along a transect across the Arctic Ocean. Mesozooplankton biomass in the upper 200 m of the water column was significantly higher (19–42 mg DW m^-3) than has previously been reported for the Arctic Ocean, and it reached a maximum at ca. 87°N in the Amundsen Basin. The lowest values were recorded in the Chukchi Sea and Nansen Basin, where ice cover was lower (50–80%) than in the central Arctic Ocean. In the deeper strata (200–500 m) of the Canadian and Eurasian Basins, the biomass was always much lower (4.35–16.44 mg DW m^-3). The C/N (g/g) ratio for the mesozooplankton population was high (6.5–8.5) but within the documented range. These high values (when compared to 4.5 at lower latitudes) may be explained by the high lipid content. Mesozooplankton accounted for approximately 40% of the total particulate organic carbon in the upper 100 m of the water column. Mesozooplankton species composition was homogeneous along the transect, consisting mainly of copepods (70–90% of the total number). It was dominated by four large copepod species (Calanus hyperboreus, C. glacialis, C. finmarchicus and Metridia longa), which together accounted for more than 80% of the total biomass. According to measurements of gut pigment and gut turnover rates, the mesozooplankton on average ingested between 6 and 30% of their body carbon per day as phytoplankton. Microzooplankton may have provided an additional source of energy for the mesozooplankton community. These data emphasize the importance of mesozooplankton in the arctic food web and reinforce the idea that the Arctic Ocean should no longer be considered to be a “biological desert”.     

Vertically migrating micronekton and macrozooplankton communities around Guam and the Northern Mariana Islands

The distribution, biomass, and assemblages of vertically migrating micronekton/macrozooplankton were studied in relation to oceanographic conditions around Guam and the adjacent Northern Mariana Islands during Spring 2010, using 3-m² Isaacs-Kidd Midwater Trawl (IKMT). The study area was located within the oligotrophic waters of the westward flowing North Equatorial Current (NEC). However, southern stations of the survey were situated close to the northern boundary of the more productive North Equatorial Countercurrent (NECC), where we observed the highest biomass, abundance, species richness, and diversity of pelagic organisms. Overall, we recorded 85 species from 20 families of mostly mesopelagic species in the area, with lanternfishes (Myctophidae—40 species) and dragonfishes (Stomiidae—18 species) being the most taxonomically diverse groups. Three genera of mesopelagic shrimps, Sergestes, Janicella and Sergia, dominated the decapod crustacean component of the micronekton community numerically and by biomass, while the contribution from cephalopods was relatively minor. Assemblages of major micronekton/macrozooplankton groups, based on biomass and abundance showed principal changes with latitude. However, the classification and ordination analysis, based on taxonomically resolved taxa (fishes and decapod shrimps), indicated additional zonal variation, with areas east and west of the island chain showing different community structure. The mean total micronekton biomass for the area near the productive boundary region between the NEC and NECC was 5.8 mg/m³, with a mean biomass of 1.2 mg/m³ obtained for stations in the oligotrophic NEC area. The corresponding biomass of mesopelagic fishes was 0.88 mg/m³ and 0.24 mg/m³ for these two areas, respectively. We reviewed and compared the available information on the quantitative distribution of midwater fish biomass in the western tropical Pacific and outlined major patterns of variation in the equatorial Pacific in general.     

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.     

Molluscan assemblages of seagrass-covered and bare intertidal flats on the Banc d'Arguin,Mauritania, in relation to characteristics of sediment and organic matter

The Banc d'Arguin, a non-estuarine area of shallows and intertidal flats off the tropical Saharan coast of Mauritania, is characterised by extensive intertidal and subtidal seagrass beds. We examined the characteristics of intertidal seagrass (Zostera noltii) meadows and bare areas in terms of the presence and abundance of molluscs (gastropods and bivalves). To explain observed differences between molluscan assemblages in seagrass and bare patches, some aspects of the feeding habitat (top-5 mm of the sediment) and of food (organic materials) of molluscs were examined. The novelty of this study is that phytopigments were measured and identified to assess source and level of decay (freshness) of organic material in the sediment and to study their importance as an explanatory variable for the distribution of molluscs. Over an area of 36 km² of intertidal flats, at 12 sites, paired comparisons were made between seagrass-covered and nearby bare patches. Within seagrass meadows, dry mass of living seagrass was large and amounted to 180 ±10 g AFDM m^− 2 (range 75–240). Containing twice the amount of silt per unit dry sediment mass, seagrass sediments were muddier than bare areas; the relative amount of organic material was also larger. The total number of species of bivalves and gastropods amounted to 27, 14 of which were found only in seagrass areas, 4 only in bare and 9 in both types of habitat. Among the three numerically most abundant species, the bivalves Anadara senilis, Dosinia hepatica and Loripes lacteus, the first was numerically most abundant in bare and the other two in seagrass-covered areas. Bare intertidal areas had greater mean total biomass of molluscs (80.5 g AFDM m^− 2) than seagrass meadows (30.0 g AFDM m^− 2). In both habitats, the bulk of the biomass was made up by A. senilis. Excluding this species, bare mudflats contained on average only 3.1 g AFDM m^− 2 and seagrass meadows 6.9 g AFDM m^− 2. As compared to previous surveys in 1980–1986, the biomass of A. senilis had increased almost 10-fold and D. hepatica, previously found in very small numbers, had become the most numerous species. However, the total biomass excluding that of A. senilis was similar. Concentrations of phytopigments were similar to those observed at temperate mudflats, indicating that the Banc d'Arguin might not be as oligotrophic as previously thought. Per unit of dry sediment mass, smaller amounts of phytopigments were found in bare than in seagrass areas. Per unit of dry organic material, bare sediments contained most (fresh) phytopigments. This suggests that in seagrass-covered meadows the organic material is more degraded than in bare sediments. Overall, the composition of phytopigments, quite surprisingly, indicated a benthic-diatom-dominated trophic system. Multivariate statistics revealed that patterns of zoobenthic assemblages were correlated with patterns of a combination of four environmental parameters: grain size of the sediment, amount of fresh phytopigments and amounts of leaves and roots of seagrass.     

Nutrient retention in the sediments and the submerged aquatic vegetation of the coastal lagoon of the Ria de Aveiro,Portugal

A decrease in the areas covered by seagrasses within the Ria de Aveiro, Portugal, has been observed over the past five decades, resulting in a corresponding increase of the areas of uncovered sediment supporting the growth of sparse macroalgae populations only. Presently, several macroalgae (Ulva spp., Gracilaria sp.) and one seagrass species (Nanozostera noltii (Hornem.) Toml. & Posl.) comprise the submerged aquatic vegetation (SAV) adapted to this shallow, high-energy environment, characterised by fast tidal currents and turbid waters and in which large areas of the bed are exposed during low tide. This study shows that there is a strong inter-relation between the SAV and the surface sediment in intertidal areas. The sediment covered by N. noltii was finer (median grain size 95 µm) and had a high percentage of organic matter (mean value 7.6%), compared with the sediment colonised by macroalgae (median grain size 239 µm; mean organic content 3.2%). The concentrations of both total nitrogen and phosphorous were significantly greater (P < 0.001) in surface sediments covered by N. noltii. Thus, sediments within N. noltii appear to act as a large reservoir of N and P by accumulating greater concentrations of fine sediment particles (silt and clay) and organic matter when compared with the coarser sediment covered with macroalgae only. Hence, the reduction in the area covered by seagrasses will likely result in a gradual loss of nutrients and fine sediment from the Ria de Aveiro channels.     

Microzooplankton composition,biomass and grazing rates along the WOCE SR3 line between Tasmania and Antarctica

Microzooplankton species composition and grazing rates on phytoplankton were investigated along a transect between ∼46 and 67°S, and between 140 and 145°E. Experiments were conducted in summer between November 2nd and December 14th in 2001. The structure of the microbial food web changed considerably along the transect and was associated with marked differences in the physical and chemical environment encountered in the different water masses and frontal regions. On average microzooplankton grazing experiments indicated that 91%, 102%, and 157%, (see results) of the phytoplankton production would be grazed in the <200, <20 and <2 μm size fractions, respectively, indicating microzooplankton grazing was potentially constraining phytoplankton populations (<200 μm) along most of the transect. Small ciliates in general and especially oligotrich species declined in importance from the relatively warm, Southern Subtropical Front waters (6.8 μg C/L) to the colder waters of the southern branch of the Polar Front (S-PF), (∼0.5 μg C/L) before increasing again near the Antarctic landmass. Large changes in microzooplankton dominance were observed, with heterotrophic nanoflagellates (HNF), ciliates and larger dinoflagellates having significant biomass in different water masses. HNF were the dominant grazers when chlorophyll a was low in areas such as the Inter-Polar Frontal Zone (IPFZ), while in areas of elevated biomass such as the S-PF and Southern Antarctic Circumpolar Current (SACC), a mix of copepod nauplii and large heterotrophic and mixotrophic dinoflagellates tended to dominate the grazing community. In the S-PF and SACC water masses the tight coupling observed between the microzooplankton grazers and phytoplankton populations over most of the rest of the transect was relaxed. In these regions grazing was low on the >20 μm size fraction of chlorophyll a, which dominated the biomass, while smaller diatoms and nanoplankton in the <20 μm size fraction were still heavily grazed. The lack of grazing pressure on large phytoplankton contributes to this region's potential to export carbon with larger cells known to have higher sinking rates.     

Bacterioplankton and phytoplankton biomass and production during summer stratification in the northwestern Mediterranean Sea

We examined bacterioplankton biomass and heterotrophic production (BHP) during summer stratification in the northwestern Mediterranean in four successive stratification seasons (June–July of 1993–1996). Values of phytoplankton biomass and primary production were determined simultaneously so that the data sets for autotrophic and heterotrophic microbial plankton could be compared. Three standard stations were set along a transect from Barcelona to the channel between Mallorca and Menorca, representing coastally influenced shelf waters, frontal waters over the slope front, and open sea waters. Conversion factors from ³H-leucine incorporation to BHP were empirically determined and varied between 0.29 and 3.25 kg C mol^-1. Bacterial biomass values were among the lowest found in any marine environment. BHP values (between 0.02 and 2.5 μg C L^-1 d^-1) were larger than those of low nutrient low chlorophyll areas such as the Sargasso Sea and lower than those from high nutrient low chlorophyll areas such as the equatorial Pacific. Growth rates of bacterioplankton were highest at the slope front (0.20 d^-1) and lowest at the open sea station (0.04 d^-1). Phytoplankton growth rates were similar at the three stations (∼0.50 d^-1). Integrated values of bacterioplankton biomass, BHP and bacterial growth rates did not show significant differences among years, but differences between the three stations were clearly significant. Phytoplankton biomass, primary production, and phytoplankton growth rates did not show significant differences either with year or with station. As a consequence the bacterioplankton to phytoplankton biomass (BB/BPHY) and production (BHP/PP) ratios varied from the coastal to the open sea stations. The BB/BPHY ratio was 0.98 at the coast and ∼0.70 at the other two stations. These ratios are similar to those found in other oligotrophic marine environments. The BHP/PP ratio was 0.83 at the coast, 0.36 at the slope and 0.09 at the open sea station. The last value is also similar to values found in other oligotrophic marine environments. Vertical distribution of these ratios was also examined.The comparison of microbial parameters at the three stations indicates a different kind of relationship between bacterioplankton and phytoplankton in oligotrophic open sea waters and in coastal, nutrient-richer waters. According to such parameters and to the values of the BB/BPHY and BHP/PP ratios, open waters in the northwestern Mediterranean (despite their relatively short distance from the shore) were intermediate between the extremely oligotrophic waters of the eastern Mediterranean or the Sargasso Sea and the more productive waters of the equatorial Pacific.     

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.     

Biomass size-spectra of macrobenthic communities in the oxygen minimum zone off Chile

Estimates of macrofaunal secondary production and normalized biomass size-spectra (NBSS) were constructed for macrobenthic communities associated with the oxygen minimum zone (OMZ) in four areas of the continental margin off Chile. The presence of low oxygen conditions in the Humboldt Current System (HCS) off Chile was shown to have important effects on the size structure and secondary production of the benthic communities living in this ecosystem. The distribution of normalized biomass by size was linear (log₂–log₂ scale) at all stations. The slope of the NBSS ranged from −0.481 to −0.908. There were significant differences between the slopes of the NBS-spectra from the stations located in the OMZ (slope = −0.837) and those located outside the OMZ (slope = −0.463) (p < 0.05). The results of this study suggest that low oxygen conditions (<0.5 ml L⁻¹) appear to influence biomass size-spectra, because small organisms are better able to satisfy their metabolic demands. The annual secondary production was higher off central Chile (6.8 g C m⁻² y⁻¹) than off northern Chile (2.02 g C m⁻² y⁻¹) and off southern Chile (0.83 g C m⁻² y⁻¹). A comparison with other studies suggests that secondary production in terms of carbon equivalents was higher than in other upwelling regions.     

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

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