Decadal-scale changes of dinoflagellates and diatoms in the anomalous baltic sea spring bloom

The algal spring bloom in the Baltic Sea represents an anomaly from the winter-spring bloom patterns worldwide in terms of frequent and recurring dominance of dinoflagellates over diatoms. Analysis of approximately 3500 spring bloom samples from the Baltic Sea monitoring programs revealed (i) that within the major basins the proportion of dinoflagellates varied from 0.1 (Kattegat) to >0.8 (central Baltic Proper), and (ii) substantial shifts (e.g. from 0.2 to 0.6 in the Gulf of Finland) in the dinoflagellate proportion over four decades. During a recent decade (1995-2004) the proportion of dinoflagellates increased relative to diatoms mostly in the northernmost basins (Gulf of Bothnia, from 0.1 to 0.4) and in the Gulf of Finland, (0.4 to 0.6) which are typically ice-covered areas. We hypothesize that in coastal areas a specific sequence of seasonal events, involving wintertime mixing and resuspension of benthic cysts, followed by proliferation in stratified thin layers under melting ice, favors successful seeding and accumulation of dense dinoflagellate populations over diatoms. This head-start of dinoflagellates by the onset of the spring bloom is decisive for successful competition with the faster growing diatoms. Massive cyst formation and spreading of cyst beds fuel the expanding and ever larger dinoflagellate blooms in the relatively shallow coastal waters. Shifts in the dominant spring bloom algal groups can have significant effects on major elemental fluxes and functioning of the Baltic Sea ecosystem, but also in the vast shelves and estuaries at high latitudes, where ice-associated cold-water dinoflagellates successfully compete with diatoms.     

An unusual bloom of the dinoflagellate Akashiwo sanguinea off the central Oregon, USA, coast in autumn 2009

An algal bloom caused by the dinoflagellate Akashiwo sanguinea was observed in October–November 2009 along the central Oregon coast (44.6°N), off Newport, Oregon, U.S.A. In this paper, the conditions are described which led to the development and demise of this bloom. The bloom was observed for 1 month from 5-October until 4-November with the peak of abundance on 19-October (347,615 cells L⁻¹). The A. sanguinea bloom followed September blooms of the diatoms Pseudo-nitzschia spp, Chaetoceros debilis, and the dinoflagellate Prorocentrum gracile. The bloom occurred when nitrate and silicate concentrations were <2 μM and <8 μM, respectively, and when the water column was stratified. This A. sanguinea dinoflagellate bloom event was closely related to the anomalous upwelling conditions in 2009: upwelling ceased early, at the end of August, whereas a normal upwelling continues into early October. This relaxation extended to near the end of September as a prolonged downwelling event, but then active upwelling reappeared in October and November. The explanation for the occurrence of the A. sanguinea bloom in October may be related to a combination of a prior diatom bloom, a stratified water column with low nutrient concentration in September, and an active upwelling event in October. As for the ultimate source of the cells, the hypothesis is that the seed stock for the A sanguinea bloom off Oregon was southward transport of cells from the Washington coast where a massive bloom of A. sanguinea was first observed in September 2009.     

Nutrient and phytoplankton dynamics in the Queen Charlotte Islands (Canada) during the summer upwelling seasons of 2001-2002

The Queen Charlotte Islands, Canada, lie at the northern extremeof the coastal upwelling system of the eastern North PacificOcean (51–54°N). In this study, the first observationsof spatial and inter-annual patterns in nutrient inventories,chlorophyll (Chl) a and phytoplankton assemblages are reportedand related to oceanographic conditions in near-shore watersof the island archipelago. Stronger and more persistent upwellingin 2002 coincided with higher nutrient and Chl a standing stockscompared to 2001 and a higher proportion of diatoms. Dinoflagellateswere more prevalent in 2001, including several potentially harmfulspecies. At sub-seasonal scales, Chl a concentrations were greatestduring downwelling conditions and smallest during upwellingconditions. On the west coast, weak water column stratification,high relative proportions of diatoms and large nutrient inventoriesaccompanied upwelling-favourable conditions, whereas on theeast coast, there was no direct relationship between the BakunUpwelling Index and water column stability. According to redundancyanalysis, variability in species composition was best explainedby sea surface temperature, the depth of the euphotic zone andnutrient inventories. The east coast supported blooms of coccolithophoreswithin protected bays, confirming previous satellite observationsthat showed bright patches in these areas. The data illustratethat moderate upwelling can have an important influence on near-shorealgal standing stocks and species composition at the northernextreme of the upwelling system off the west coast of NorthAmerica, and that topographical complexity may be importantfor the development of phytoplankton blooms.     

Turbulence, watermass stratification and harmful algal blooms: an alternative view and frontal zones as “pelagic seed banks”

Watermass stratification has been considered the essential physical condition that dinoflagellates require to bloom because of their relative inability, unlike diatoms, to tolerate the elevated shear-stress associated with water-column mixing, turbulence and high velocity, coastal currents. The swimming speeds of 71 flagellate taxa, with a focus on dinoflagellates, are compared to the turbulence fields and vertical velocities that develop during representative wind conditions, upwelling and at frontal zones. The results suggest that the classical stratification–dinoflagellate bloom paradigm needs revision. Tolerance of turbulence, growth within well-mixed watermasses and survival and dispersal while entrained within current systems are well developed capacities among dinoflagellates. Their secretion of mucous, often copious during blooms, is suggested to be an environmental engineering strategy to dampen turbulence. Biophysical tolerance of turbulence by dinoflagellates is often accompanied by high swimming speeds. Motility speeds of many species exceed in situ vertical current velocities; this also allows diel migrational patterns and other motility-based behavior to persist. Species belonging to “mixing-drift” life-form assemblages can increase their swimming speeds through chain formation, which helps to compensate for the increased turbulence and vertical water-column velocities of their habitats. The ability of dinoflagellate species to tolerate the vertical velocities of offshore, frontal zones, where abundant populations often develop, suggests that fronts may serve as “pelagic seed banks”, occurring as pelagic analogues of nearshore seed beds, from which seed stock is dispersed. The different ecologies associated with the hypothesized, “pelagic seed banks” of vegetative cells and the “seed beds” of resting stage cells deposited onto sediments are discussed. There is a contradiction in the stratification–HAB paradigm: the quiescent conditions of a stratified watermass, with its characteristic nutrient-poor conditions are expected to promote stasis of the population, rather than growth and blooms. The analyses suggest that dinoflagellate blooms do not preponderate in stratified watermasses because the bloom species are biophysically intolerant of the higher velocities and turbulence of more mixed watermasses. The watermass stratification that often accompanies flagellate blooms is probably a secondary, parallel event and less essential than some other factor(s) in triggering the observed bloom.     

Annual microplankton cycles in Villefranche Bay, Ligurian Sea, NW Mediterranean

Abundance and composition of microplankton were studied overa period of 2 years at two depths in Villefranche Bay (LigurianSea, NW Mediterranean Sea). Diatoms dominated the microplanktonin late spring and autumn, whereas dinoflagellates composedthe major part of the microplankton in summer. The silicoflagellateDictyocha fibula and the diatom Thalassionema frauenfeldii dominatedin winter. Ciliates showed low variability throughout the yearwith the lowest abundance in February and an increase whichcoincided with the diatom maxima during autumn in both years.In 1998, the spring bloom (in May) was mainly composed of dinoflagellatesnear the surface and of diatoms in deeper layers. Subsurfacediatom maxima were observed in August–September and November.In 1999, diatoms peaked in May both at the surface and at thedepth of 50 m. They showed a strong maximum in October. Dinoflagellatesand tintinnids showed maxima in early November. Comparisonswith previous studies reveal that (i) changes in species compositionhave not been significant, (ii) the silicoflagellate’sabundance is lower during the present study, (iii) the sequentialspring bloom is composed of a pico-nanoplankton bloom in Marchand microphytoplankton in May, whereas in other western Mediterraneanareas the spring microphytoplankton bloom is reported in Februaryand March, (iv) high water transport through the Corsica channelcoinciding with low or negative winter values of North AtlanticOscillation (NAO) index are associated with the anomalous strongdevelopment of the spring diatom blooms in the Bay of Villefranche,whereas the usual trend is the lack of or weak development ofthe spring diatom bloom. This feature may determine the natureand the fate of primary production and the interannual variabilityin the relative importance of the microbial food web versusthe microbial loop.     

Feeding dynamics of the copepod Diacyclops thomasi before, during and following filamentous cyanobacteria blooms in a large, shallow temperate lake

Cyanobacteria blooms are an increasing problem in temperate freshwater lakes, leading to reduced water quality and in some cases harmful effects from toxic cyanobacteria species. To better understand the role of zooplankton in modulating cyanobacteria blooms, from 2008 to 2010 we measured water quality and plankton abundance, and measured feeding rates and prey selectivity of the copepod Diacyclops thomasi before, during and following summertime cyanobacteria blooms in a shallow, eutrophic lake (Vancouver Lake, Washington, USA). We used a combined field and experimental approach to specifically test the hypothesis that copepod grazing was a significant factor in establishing the timing of cyanobacteria bloom initiation and eventual decline in Vancouver Lake. There was a consistent annual succession of zooplankton taxa, with cyclopoid copepods (D. thomasi) dominant in spring, followed by small cladocerans (Eubosmina sp.). Before each cyanobacteria bloom, large cladocerans (Daphnia retrocurva, Daphnia laevis) peaked in abundance but quickly disappeared, followed by brief increases in rotifers. During the cyanobacteria blooms, D. thomasi was again dominant, with small cladocerans abundant in autumn. Before the cyanobacteria blooms, D. thomasi substantially consumed ciliates and dinoflagellates (up to 100% of prey biomass per day), which likely allowed diatoms to flourish. A shift in copepod grazing toward diatoms before the blooms may have then helped to facilitate the rapid increase in cyanobacteria. Copepod grazing impact was the highest during the cyanobacteria blooms both years, but focused on non-cyanobacteria prey; copepod grazing was minimal as the cyanobacteria blooms waned. We conclude that cyclopoid copepods may have an indirect role (via trophic cascades) in modulating cyanobacteria bloom initiation, but do not directly contribute to cyanobacteria bloom decline.     

Harmful phytoplankton diversity and dynamics in an upwelling region (Sagres,SW Portugal) revealed by ribosomal RNA microarray combined with microscopy

The study region in Sagres, SW Portugal, is subject to natural eutrophication of coastal waters by wind-driven upwelling, which stimulates high primary productivity facilitating the recent economic expansion of bivalve aquaculture in the region. However, this economic activity is threatened by harmful algal blooms (HAB) caused by the diatoms Pseudo-nitzschia spp., Dinophysis spp. and other HAB dinoflagellates, all of which can produce toxins, that can induce Amnesic Shellfish Poisoning (ASP), Diarrhetic Shellfish Poisoning (DSP) and Paralytic Shellfish Poisoning (PSP). This study couples traditional microscopy with 18S/28S rRNA microarray to improve the detection of HAB species and investigates the relation between HAB and the specific oceanographic conditions in the region. Good agreement was obtained between microscopy and microarray data for diatoms of genus Pseudo-nitzschia and dinoflagellates Dinophysis spp., Gymnodinium catenatum and raphidophyte Heterosigma akashiwo, with less effective results for Prorocentrum. Microarray provided detection of flagellates Prymnesium spp., Pseudochattonella spp., Chloromorum toxicum and the important HAB dinoflagellates of the genera Alexandrium and Azadinium, with the latter being one of the first records from the study region. Seasonality and upwelling induced by northerly winds were found to be the driving forces of HAB development, with Pseudo-nitzschia spp. causing the risk of ASP during spring and summer upwelling season, and dinoflagellates causing the risk of DSP and PSP during upwelling relaxation, mainly in summer and autumn. The findings were in agreement with the results from toxicity monitoring of shellfish by the Portuguese Institute for Sea and Atmosphere and confirm the suitability of the RNA microarray method for HABs detection and aquaculture management applications.     

The influence of weather conditions (temperature and wind) on cyanobacterial bloom development in the Gulf of Finland (Baltic Sea)

Cyanobacterial blooms are common in the Baltic Sea. They are dominated by Aphanizomenon flos-aquae and Nodularia spumigena and take place in July–August. Investigations of bloom development using different approaches have been carried out in the Gulf of Finland during recent years. The ship-of-opportunity technique allows to observe the upper layer dynamics from meso- to basin-wide scale with high temporal and spatial frequency at low cost. Unattended measurements on board a commercial ferry along a transect between Tallinn and Helsinki have been conducted for 3 years (1997–1999). The influence of weather conditions—temperature and wind—on the cyanobacterial bloom development was investigated. The formation of cyanobacterial blooms was favoured by warm and calm weather, while in cold and windy conditions other species formed mass occurrences. Water temperature has been found to be the main factor controlling the initiation of the bloom, in general, while vertical stratification appeared to be the critical factor determining the intensity of the bloom at species level. The spatial distribution of the cyanobacterial bloom was determined rather by the wind-forced advection than by the possible vertical transport of nutrients in the areas of the observed upwelling events.     

Predictive Ecological Modeling of Harmful Algal Blooms

We have thus far constructed a series of coupled ecological/physical models to predict the origin and fate of harmful algal blooms of the toxic dinoflagellate, Gymnodinium breve, on the West Florida shelf. We find that (1) the maximal population growth rate of G. breve must be ～0.80 day^?1 during initiation of a red tide, but as little as 0.08 day^?1 during its decay, (2) diatoms dominate when estuarine and shelf-break supplies of nitrate are made available to a model community of small and large diatoms, coccoid cyanophytes and Trichodesmium, non-toxic and red-tide dinoflagellates, microflagellates, and coccolithophores, (3) a numerical recipe for large red tides of G. breve requires DON supplies, mediated by iron-starved, nitrogen-fixers, while small blooms may persist on sediment sources of DON, (4) selective grazing must be exerted on the non-toxic dinoflagellates, and (5) vertical migration of G. breve in relation to seasonal changes of summer downwelling and fall/winter upwelling flow fields determines the duration and intensity of red tide landfalls along the barrier islands and beaches of the west coast of Florida, once other losses are specified. Given poorly known initial and boundary conditions and the expense of shipboard monitoring programs, however, bio-optical moorings or remote sensors are the most likely sources of model validation and improved HAB forecasts. Accordingly, the bio-optical implications of our ecological models must be included in future simulation analyses of HABs on both the West Florida shelf and within other coastal regions. Of particular importance for initiation of the coupled biophysical models is an improved understanding of the relationship of remotely sensed surface signals to shade-adapted dinoflagellates, aggregating below the first optical depth of the water column.     

Bacterioplankton composition of the coastal upwelling system of 'Ría de Vigo', NW Spain

Catalysed reported deposition-FISH and clone libraries indicated that Roseobacter , followed by Bacteroidetes , and some gammaproteobacterial groups such as SAR86, dominated the composition of bacterioplankton in Ría de Vigo, NW Spain, in detriment to SAR11 (almost absent in this upwelling ecosystem). Since we sampled four times during the year, we observed pronounced changes in the structure of each bacterioplankton component, particularly for the Roseobacter lineage. We suggest that such variations in the coastal upwelling ecosystem of Ría de Vigo were associated with the characteristic phytoplankton communities of the four different hydrographical situations: winter mixing, spring bloom, summer stratification, and autumn upwelling. We retrieved new sequences among the major marine bacterial lineages, particularly among Roseobacter , SAR11, and especially SAR86. The spring community was dominated by two Roseobacter clades that had previously been related to phytoplankton blooms. In the other seasons, communities with higher diversity than the spring one were detected.     

The role of mixotrophic protists in the population dynamics of the microbial food web in a small artificial pond

• 1 The seasonal variations of protist and rotifer populations were monitored over 1 year in a small artificial pond. Grazing rates on fluorescently labelled bacteria were also determined.
• 2 The data showed population dynamics similar to other small freshwater bodies; diatoms were numerous during the spring, chlorophytes dominated during the summer months, and mixotrophs, in particular Gymnodinium, dominated during the autumn and winter.
• 3 The mixotrophic dinoflagellates were responsible for a high chlorophyll concentration during the autumn and winter. Mixotrophs were important consumers of bacteria, particularly during the autumn when population densities of pure heterotrophs were low. Mixotrophs were an important component of the microbial food web in this pond.

     

Parasitism as a biological control agent of dinoflagellate blooms in the California Current System

Amoebophrya is a marine parasite recently found to infect and kill bloom-forming dinoflagellates in the California Current System (CCS). However, it is unknown whether parasitism by Amoebophrya can control dinoflagellate blooms in major eastern boundary upwelling systems, such as the CCS. We quantified the abundance of a common bloom-forming species Akashiwo sanguinea and prevalence of its parasite (i.e., % infected cells) in surface water samples collected weekly from August 2005 to December 2008 at the Santa Cruz Wharf (SCW), Monterey Bay, CA. Additionally, we measured physical and chemical properties at the SCW and examined regional patterns of wind forcing and sea surface temperature. Relative abundance of the net phytoplankton species was also analyzed to discern whether or not parasitism influences net phytoplankton community composition. Epidemic infection outbreaks (>20% parasite prevalence in the host species) may have contributed to the end or prevented the occurrence of A. sanguinea blooms, whereas low parasite prevalence was associated with short-term (≤2 weeks) A. sanguinea blooms. The complete absence of parasitism in 2007 was associated with an extreme A. sanguinea bloom. Anomalously strong upwelling conditions were detected in 2007, suggesting that A. sanguinea was able to outgrow Amoebophrya and ‘escape’ parasitism. We conclude that parasitism can strongly influence dinoflagellate bloom dynamics in upwelling systems. Moreover, Amoebophrya may indirectly influence net phytoplankton species composition, as species that dominated the net phytoplankton and developed algal blooms never appeared to be infected.     

大亚湾澳头水域浮游植物群落结构及周年数量动态

对1997年至1998年广东省大亚湾澳头水域的浮游植物群落进行调查和分析。结果发现浮游植物65属198种;硅藻在种类组成和数量上都比甲藻占有优势,存在春季和秋季高峰,主要优势类群依次是角毛藻、骨条藻、拟菱形藻等;甲藻只存在春季高峰,代表种类有裸甲藻、原甲藻等。主要优势种类的生长与调查水域的盐度没有明显关系,但全年水温的季节性变化对优势种类的消长影响显著。Simpson多样性指数、Shannon-Weaver多样性指数、均匀度的年平均值分别是0.611、2.107、0.557,多样性指数没有明显的季节变化规律和水平分布规律。       

Regulation of blue-green algal buoyancy and bloom formation by light,inorganic nitrogen,C02, and trophic level interactions

In highly eutrophic ponds, buoyancy of the gas-vacuolate blue-green alga Anabaenopsis Elenkinii (Miller) was regulated by complex interactions between chemical and physical parameters, as well as by biological interactions between various trophic levels. Algal buoyancy and surface bloom formation were enhanced markedly by decreased light intensity, and to a lesser extent by decreased CO₂ availability and increased availability of inorganic nitrogen. In the absence of dense populations of large-bodied Cladocera, early season blooms of diatoms and green algae reduced light availability in the ponds thus creating conditions favorable for increased buoyancy and bloom formation by A. Elenkinii. The appearance of blue-green algal blooms could be prevented by a reduced density of planktivorous fish, which allowed development of dense cladoceran populations. The cladocerans limited the growth of precursory blooms of diatoms and green algae, and given the resulting clear-water conditions, buoyancy of A. Elenkinii was reduced, and blue-green algal blooms never appeared.     

Selective grazing of Temora longicornis in different stages of a Phaeocystis globosa bloom—a mesocosm study

Selective grazing of a calanoid copepod Temora longicornis was measured during different stages of a Phaeocystis globosa bloom, in order to reveal (1) if T. longicornis feeds on single cells and/or colonies of P. globosa in the presence of alternative food sources, (2) if copepod food selection changes during the initiation, maintenance, collapse and decay of a P. globosa bloom and (3) if P. globosa dominated food assemblage provides a good diet for copepod egg production. Our results show low but constant feeding on small colonies of P. globosa, irrespective of the type or concentration of alternative food sources. In contrast, feeding on single cells was never significant, and the total contribution of P. globosa to carbon ingestion of T. longicornis was minor. T. longicornis fed most actively on the decaying colonies, whereas during the peak of the bloom copepods selected against P. globosa. Mostly, T. longicornis fed unselectively on different food particles: before the bloom, the major part of the diet consisted of diatoms, whereas during and after the bloom copepod diet was dominated by dinoflagellates and ciliates. Egg production was highest during the decay of the bloom, coinciding with highest proportional ingestion of heterotrophic organisms, but was not seriously reduced even during the peak of the bloom. We conclude that P. globosa blooms should not threaten survival of copepod populations, but the population recruitment may depend on the type (and concentration) of the dominant heterotrophs present during the blooms. Due to relatively unselective grazing, the impact of T. longicornis to the initiation of a Phaeocystis bloom is considered small, although grazing on decaying colonies may contribute to the faster termination of a bloom.     

A spring poleward current and its influence on microplankton assemblages and harmful dinoflagellates on the western Iberian coast

Along-shore currents can propagate harmful algal blooms (HABs) over long distances in many coastal areas of the ocean. Harmful dinoflagellate blooms on the west coast of Iberia frequently occur when the Iberian poleward current (IPC) establishes on the continental slope. This has led to the suggestion that HABs could be transported northward by the IPC. To examine this possibility, the microplankton composition along the west coast of Iberia was studied in May 1993 coinciding with the presence of the IPC. The microplankton of the IPC was almost exclusively composed of small flagellates, with the notable absence of the harmful species usually associated with coastal waters. The primary influence of the IPC was to confine coastal microplankton populations to the shelf, where a downwelling convergence prevented their export from the coastal environment. Microplankton assemblages on the shelf revealed a north–south gradient related to different stages of succession. Earlier stages of succession in which diatoms were prominent were found on the northern shelf, whereas dinoflagellates were more abundant in the south. The toxic species Gymnodinium catenatum, which was only present in the southern shelf, did not show a northward transport associated with the IPC. It is suggested that the northward spreading of HABs along the west coast of Iberia must be related to the interaction between the IPC, which accumulates coastal populations on the shelf, and the latitudinal progress of microplankton succession that determines species composition. Thus, during the course of the season, HABs are likely to be observed in the south prior to their development in the north.     

Community-Level and Species-Specific Associations between Phytoplankton and Particle-Associated Vibrio Species in Delaware's Inland Bays

Vibrio species are an abundant and diverse group of bacteria that form associations with phytoplankton. Correlations between Vibrio and phytoplankton abundance have been noted, suggesting that growth is enhanced during algal blooms or that association with phytoplankton provides a refuge from predation. Here, we investigated relationships between particle-associated Vibrio spp. and phytoplankton in Delaware''s inland bays (DIB). The relative abundances of particle-associated Vibrio spp. and algal classes that form blooms in DIB (dinoflagellates, diatoms, and raphidophytes) were determined using quantitative PCR. The results demonstrated a significant correlation between particle-associated Vibrio abundance and phytoplankton, with higher correlations to diatoms and raphidophytes than to dinoflagellates. Species-specific associations were examined during a mixed bloom of Heterosigma akashiwo and Fibrocapsa japonica (Raphidophyceae) and indicated a significant positive correlation for particle-associated Vibrio abundance with H. akashiwo but a negative correlation with F. japonica. Changes in Vibrio assemblages during the bloom were evaluated using automated ribosomal intergenic spacer analysis (ARISA), which revealed significant differences between each size fraction but no significant change in Vibrio assemblages over the course of the bloom. Microzooplankton grazing experiments showed that losses of particle-associated Vibrio spp. may be offset by increased growth in the Vibrio population. Moreover, analysis of Vibrio assemblages by ARISA also indicated an increase in the relative abundance for specific members of the Vibrio community despite higher grazing pressure on the particle-associated population as a whole. The results of this investigation demonstrate links between phytoplankton and Vibrio that may lead to predictions of potential health risks and inform future management practices in this region.     

Seasonal and spatial variation of phytoplankton assemblages, biomass and cell size from spring to summer across the north-eastern New Zealand continental shelf

The composition, biomass and cell size of phytoplankton taxonomicgroups were determined in the Hauraki Gulf and adjacent shelfof north-eastern New Zealand. In early spring, on the innershelf, over-winter mixing and upwelling supported a bloom dominatedby large, chain-forming diatoms in a moderately turbulent watercolumn. The bloom declined in late spring because of nutrientlimitation, and the assemblage evolved initially toward smalldiatoms, and eventually to co-occurrence of dinoflagellates,small nanoflagellates and picophytoplankton in early and latesummer. Mid- to outer-shelf biomass was much lower than inshore,and was dominated by small or motile taxa which were probablylimited by grazing and light. In early summer, strong upwellingdisplaced inner shelf phytoplankton to beyond the shelf edge,whilst enriching the shelf with nutrients. However, shelf phytoplanktonbiomass increased only after the relaxation of upwelling. TheHauraki Gulf was strongly stratified from early spring throughlate summer. The flora were seasonally less variable than onthe shelf, with a thecate dinoflagellate-dominated flora inearly spring, replaced post-bloom by the co-occurrence of presumablylow-nutrient-adapted autotrophic and/or heterotrophic dinoflagellates(most of which were non-thecate, and some toxic), nanoflagellatesand picophytoplankton. The succession in floristics was consistentwith a change from an autotrophic toward a heterotrophic ecosystemfrom spring to summer. Implications for secondary production,and vertical and lateral organic carbon export on the shelf,are discussed.     

Phytoplankton biomass and production in shelf waters off NW Spain: spatial and seasonal variability in relation to upwelling

Summary Chlorophyll-a and primary production on the euphotic zone of the N-NW Spanish shelf were studied at 125 stations between 1984 and 1992. Three geographic areas (Cantabrian Sea, Rías Altas and Was Baixas), three bathymetric ranges (20 to 60 m, 60 to 150 m and stations deeper than 200 m), and four oceanographic stages (spring and autumn blooms, summer upwelling, summer stratification and winter mixing) were considered. One of the major sources of variability of chlorophyll and production data was season. Bloom and summer upwelling stages have equivalent mean and maximum values. Average chlorophyll-a concentrations approximately doubled in every step of the increasing productivity sequence: winter mixing — summer stratification — high productivity (upwelling and bloom) stages. Average primary production rates increased only 60% in the described sequence. Mean (± sd) values of chlorophyll-a and primary production rates during the high productivity stages were 59.7 ± 39.5 mg Chl-a m^–2 and 86.9 ± 44.0 mg C m^–2 h^–1, respectively. Significant differences in both chlorophyll and primary production resulted between geographic areas in most stages. Only 27 stations showed the effects of the summer upwelling that affected coastal areas in the Cantabrian Sea and Rías Baixas shelf, but also shelf-break stations in the Rías Altas area. The Rías Baixas area had lower chlorophyll than both the Rías Altas and the Cantabrian Sea areas during spring and autumn blooms, but higher during summer upwelling events. On the contrary, primary production rates were higher in the Rías Baixas area during blooms in spring and autumn. Mid-shelf areas showed the highest chlorophyll concentrations during high productivity stages, probably due to the existence of frontal zones in all geographic areas considered. The estimated phytoplankton growth rates were comparable to those of other coastal upwelling systems, with average values lower than the maximum potential growth rates. Doubling rates for upwelling and stratification stages in the northern and Rías Altas shelf areas were equivalent, despite larger biomass accumulations during upwelling events. Low turnover rates of the existing biomass in the Rías Baixas shelf in upwelling stages suggests that the accumulation of phytoplankton was due mainly to the export from the highly productive rías, while the contribution of in situ production to these accumulations was relatively lower.