Composition and interannual variability of phytoplankton in a coastal upwelling region (Lisbon Bay,Portugal)

From July 2001 to May 2005, at a fixed station located in Lisbon Bay (Cascais: 38° 41′ N, 09° 24′ W), surface seawater samples were collected on a weekly basis. We aimed to describe at different temporal scales, short-term to interannual, the phytoplankton community in relation to hydrographic conditions.Maxima of the main phytoplankton groups varied according to the seasonality of upwelling/downwelling cycles and nutrient availability and were associated with particular hydrological mesoscale structures highlighted by satellite images. Short succession cycles were identified dependent on coastal upwelling events. Intermittent and weak pulses allowed the coexistence of species from different succession stages and groups, although having consecutive maxima. The interannual differences observed in the phytoplankton community, in Lisbon Bay, varied according to both the duration and strength of the upwelling events and to precipitation and Tagus river flow regimes.Diatoms developed and were dominant, during spring–summer under prevailing upwelling conditions and silicon availability. Short upwelling pulses appeared to be unfavourable for diatoms maintenance. When upwelling weakened and SST increased due to onshore advection of warmer waters, coccolithophores dominated. This assemblage was the second most abundant during the study, in particular during the short transition period from upwelling (summer) to downwelling seasons (autumn) distributing in the largest range of hydrographical conditions between diatoms (maximum turbulence) during early spring and dinoflagellates (maximum stratification) during summer to further dominate during autumn and winter. Nitrites and nitrates seemed to favour greater developments of this group. Dinoflagellates peaked mainly during summer and were the less abundant through the four years due to the decrease of lasting convergence periods. Like coccolithophores, a preference for warmer waters emerged but this group seemed to have a narrow tolerance to turbulence and temperature changes.     

The effect of water transport on nitrogen flow through a kelp-bed community

A simulation model is used to investigate possible ecological effects of up- and downwelling on nitrogen flows through a kelp-bed food web off the Cape Peninsula, South Africa. The model depicts the flow of nitrogen, which is often limiting in marine ecosystems, from kelps, other macrophytes and phytoplankton, through filter-feeders to carnivores, with a feedback loop via faeces and bacteria to detritus and filter-feeders. When modelled as a closed system, bacteria associated with detritus and animal faeces form a large component of the particulate nitrogen available to filter-feeders, and the faeces feedback loop dominates nitrogen flow. When measured rates of water transport are incorporated into the model, bacteria have little opportunity to accumulate before being transported out of the system. Animal faeces and kelp detritus are the dominant filter-feeder food components under upwelling conditions, whereas phytoplankton is the major contributor to particulate organic nitrogen under downwelling conditions. When realistic pulses of upwelling/downwelling derived from wind indices are used as model input, filter-feeders are shown to decline during the summer upwelling season when much potential food is advected out of the system, and they increase during the winter when downwelling conditions are more prevalent, bringing in nitrogen-rich phytoplankton from the blooms developing offshore.     

夏季南海北部浮游植物对上升流与羽状流的响应

文章建立了基于真实场驱动的三维物理—生态耦合模型, 利用模型定量分析了夏季南海北部上升流和羽状流过程对浮游植物生物量空间分布的影响程度及作用机制。首先, 利用2006—2008年卫星遥感数据及2006与2008年夏季观测数据对模型进行了验证, 结果表明, 模型能较好地再现夏季南海北部上升流和羽状流过程, 较好地反映出浮游植物的空间分布特征。模拟分析结果显示, 夏季南海北部浮游植物主要分布在50m等深线以内。琼州海峡东部海域和汕头海域浮游植物垂向分布较为均匀, 上升流的贡献均达到90%以上, 表层水平平流输送是浮游植物主要的汇, 生物过程是浮游植物的源。珠江口和汕尾海域浮游植物存在表层和次表层两个高值区, 羽状流贡献35%~40%, 主要促进表层浮游植物生长, 而上升流贡献60%~65%, 主要促进中底层浮游植物的生长。粤西海域上升流对浮游植物的贡献占92%, 主要促进中底层浮游植物生长, 而表层浮游植物浓度极低。整体上, 夏季南海北部上升流和羽状流主要是通过输送营养盐的方式影响浮游植物的生长。上升流对营养盐的输送作用是向岸方向的爬升输送和平行于等深线的沿岸流输送共同作用的结果。跃层的存在改变了营养盐的垂向输送过程, 是导致上升流和羽状流过程对不同水层浮游植物贡献差异的关键因素之一。整体而言, 夏季南海北部浮游植物空间分布差异是以上升流、羽状流主导, 环流—营养盐—生物过程共同作用的结果。     

Spatial and temporal seasonal trends in coastal upwelling off Northwest Africa, 1981–2012

Seasonal coastal upwelling was analyzed along the NW African coastline (11–35°N) from 1981 to 2012. Upwelling magnitudes are calculated by wind speed indices, sea-surface temperature indices and inferred from meteorological station, sea-surface height and vertical water column transport data. A permanent annual upwelling regime is documented across 21–35°N and a seasonal regime across 12–19°N, in accordance with the climatology of previous studies. Upwelling regions were split into three zones: (1) the Mauritania–Senegalese upwelling zone (12–19°N), (2) the strong permanent annual upwelling zone (21–26°N) and (3) the weak permanent upwelling zone (26–35°N). We find compelling evidence in our various indices for the Bakun upwelling intensification hypothesis due to a significant coastal summer wind speed increase, resulting in an increase in upwelling-favorable wind speeds north of 20°N and an increase in downwelling-favorable winds south of 20°N. The North Atlantic Oscillation plays a leading role in modifying interannual variability during the other seasons (autumn–spring), with its influence dominating in winter. The East Atlantic pattern shows a strong correlation with upwelling during spring, while El Niño Southern Oscillation and Atlantic Multi-decadal Oscillation teleconnections were not found. A disagreement between observationally-based wind speed products and reanalysis-derived data is explored. A modification to the Bakun upwelling intensification hypothesis for NW Africa is presented, which accounts for the latitudinal divide in summer wind regimes.     

Hydrological conditions and circulation off the west coast of the North Island,New Zealand

Temperature and salinity surveys were carried out in the Tasman Sea in winter (August 1973) and summer (February‐March 1974). In both surveys the presence of the Westland Current was indicated by the distribution of surface water properties; in summer it was associated with a subsurface salinity maximum. The current extended further northwards in summer than in winter. In summer, an east‐going geostrophic flow at about latitude 35°S separated on approaching New Zealand; part of the flow passed north around the North Island and part moved slowly eastwards in the deeper off‐shore water to at least latitude 38°S. The West Auckland Current was apparent in the winter, but not in the summer. To the west of Cape Reinga, relatively low values of surface temperature and salinity are probably associated with upwelling between Cape Reinga and the Three Kings Islands. Upwelling was observed along the coast between Kaipara and Manukau Harbour.     

The 4-D structure of upwelling and Pearl River plume in the northern South China Sea during summer 2008 revealed by a data assimilation model

We analyze four-dimensional structures of upwelling and Pearl River plume in the northern South China Sea (NSCS) during the summer of 2008 based on data assimilation. An Ensemble Kalman Smoother scheme is employed in the Princeton Ocean Model. It is found that the Pearl River plume axis extended eastward along with the surface current and swerved offshore twice near (116°E, 22.6°N) and (117.5°E, 22.8°N) before reaching the Taiwan Strait. The vertical transect of salinity along the plume axis indicates that the Pearl River freshwater could affect salinity distribution down to a depth of 10–20 m. Anomalously warm water is found in the upper layer, which could be attributed to the intensified stratification and suppressed vertical mixing caused by the freshwater of the plume capping the upwelling west of 116°E. The varying winds from upwelling favorable to downwelling favorable could induce a low-salinity water lens at the center of the model domain. Upwelling in the NSCS initially occurred at 114.5°E, to the east of the Pearl River Estuary, intensified eastward, and reached its maximum near Shantou (116.7°E, 23.2°N). Since current-induced upwelling appeared mainly in Shantou due to the widened shelf, it is found that even if the wind-induced upwelling was shut down in Shanwei by downwelling favorable wind on July 4, the upwelling still existed in Shantou. Moreover, because the direction of large-scale current was in favor of upwelling in the NSCS that cannot be reversed by varying local winds over a short time period, the upwelling shutdown time is longer for both wind-induced and current-induced upwelling in Shantou than for mainly wind-induced upwelling in Shanwei. The steeper slope in Shanwei also shortens the upwelling shutdown time there.     

大气氮沉降对南黄海初级生产过程影响的数值研究

本研究利用三维物理-生物耦合模型模拟了大气氮沉降对南黄海主要初级生产过程的影响,并通过数值实验区分了不同季节大气氮沉降的贡献。模拟结果显示,大气氮沉降明显增大了南黄海表层溶解无机氮的浓度,近岸海域增加量较大,可以达到3.0 mmol/m^3,且由近岸海域到黄海中部海域有明显的递减趋势,这主要是由于近岸海域无机氮来源众多,导致浓度较高,大气沉降的氮不会被浮游植物生长吸收,出现氮累积。大气氮沉降明显促进了黄海中部春季表层水华和夏季次表层叶绿素最大值两个重要初级生产过程,春季表层叶绿素增加量最大,可达0.20 mg/m^3,夏季次表层叶绿素浓度增加最显著,可达0.10 mg/m^3,分别约为峰值浓度的10%和6%。不同季节大气氮沉降对初级生产过程的贡献不同,冬季氮沉降可以存留下来影响春季水华过程,但作用小于春季氮沉降;夏季,由于水体层化较强,本季的氮沉降对次表层叶绿素最大值的促进作用并不明显,反而冬季氮沉降的影响大于春季和夏季的氮沉降。同时,大气氮沉降也促进了氮循环的各个过程,包括浮游植物生长吸收、呼吸释放和矿化过程。     

Numerical investigation of the effects of upwelling on harmful algal blooms off the west Florida coast

The initiation of the toxic harmful algal bloom (HAB), Karenia brevis, along the west Florida coast has been associated with upwelling events. Upwelling processes may be responsible for the transport of nutrients or algae from deep offshore locations across the Florida shelf to the coast. The influence of coastal wind-driven upwelling on the onset and occurrences of K. brevis in this region was numerically investigated using Rutgers University's Regional Ocean Modeling System. Computations were carried out in an idealized model domain, a two-dimensional slice in the cross-shore and vertical directions. The surface forcing data used was from several offshore meteorological buoys. The motion of the algae was simulated using Lagrangian particles and a passive tracer. The numerical simulations of three K. brevis events in 2000–2002 showed that the particles respond (with upwelling/downwelling) to the along-shore wind stresses as expected and some upwelling was present during the events. Comparison of the passive tracer fields with measured fluorescence data exposed the model's sensitivity to the particular surface forcing data employed and the relatively more significant role played by surface forcing over initial conditions. The present model set-up constitutes a useful predictive tool for conditions conducive to the onset of HABs. It is planned to be used in a real-time mode to aid the NOAA HAB monitoring and forecasting system.     

印尼爪哇上升流海域次表层叶绿素最大值层的浮游植物色素分布特征

Upwelling occurs on the coast of Java between June and October, forced by local alongshore winds associated with the southeasterly monsoon. This causes variations in phytoplankton community composition in the upwelling zone compared with the surrounding offshore area. Based on pigments analysis with subsequent calculations of group contributions to total chlorophyll a(Chl a) using CHEMTAX, we studied the distribution and composition of phytoplankton assemblages in the subsurface chlorophyll maximum along the south coast of Java and the influence of upwelling. Nineteen phytoplankton pigments were identified using high-performance liquid chromatography, and CHEMTAX analysis associated these to ten major phytoplankton groups. The phytoplankton community in the coastal area influenced by upwelling was characterized by high Chl a and fucoxanthin concentrations, indicating the dominance of diatoms. In contrast, in the offshore area, the Chl a and fucoxanthin concentrations declined to very low levels and the community was dominated by haptophytes represented by 19′-Hexanoyloxyfucoxanthin. Accordingly, microphytoplankton was found to be the major size class in the coastal area influenced by upwelling, while nanophytoplankton was most abundant in the offshore area. Low concentrations of other accessory pigments indicated less contribution from dinoflagellates,prasinophytes, chlorophytes and cryptophytes. Photo-pigment indices revealed that photosynthetic carotenoids(PSCs) were the largest component of the pigment pool, exceeding the proportion of Chl a, with the average PSCTP up to 0.62. These distribution trends can mainly be explained by phytoplankton adaption strategies to upwelling and subsurface conditions by changing species composition and adjusting the pigment pool.     

WEST: A northern California study of the role of wind-driven transport in the productivity of coastal plankton communities

The “Wind Events and Shelf Transport” (WEST) program was an interdisciplinary study of coastal upwelling off northern California in 2000–03. WEST was comprised of modeling and field observations. The primary goal of WEST was to better describe and understand the competing influences of wind forcing on planktonic productivity in coastal waters. While increased upwelling-favorable winds lead to increased nutrient supply, they also result in reduced light exposure due to deeper surface mixed layers and increased advective loss of plankton from coastal waters. The key to understanding high levels of productivity, amidst these competing responses to wind forcing, is the temporal and spatial structure of upwelling. Temporal fluctuations and spatial patterns allow strong upwelling that favors nutrient delivery to be juxtaposed with less energetic conditions that favor stratification and plankton blooms. Observations of winds, ocean circulation, nutrients, phytoplankton and zooplankton off Bodega Bay and Point Reyes (38°N) were combined with model studies of winds, circulation and productivity. This overview of the WEST program provides an introduction to the WEST special issue of Deep-Sea Research, including the motivation for WEST, a summary of study components, an integrative synthesis of major research results to-date, and background on conditions during field studies in May–June 2001 (the upwelling period on which this special issue is focused).     

南黄海浮游植物季节性变化的数值模拟与影响因子分析

用三维物理-生物耦合模式研究南黄海浮游植物(以叶绿素a为指标)的季节变化.对于物理模式采用Princeton ocean model(POM),对于生物模式考虑溶解无机营养盐(氮、磷、硅)、浮游植物、食草性浮游动物和碎屑.给定已知的初始场和外加边界强迫,模拟了观测到叶绿素a的主要时、空分布特征,如浮游植物的春、秋季水华和夏季次表层叶绿素a极大值现象等.研究表明,浮游植物春季水华最先发生于黄海中央海域,主要原因是该海域透明度较高,流速较小.春季水华开始于垂直对流减弱和层化开始形成之前(约3月底至4月上旬),显著地依赖水层的稳定性.水体层化以后(约5～9月)叶绿素a浓度高值区分布在南黄海的南部和锋区.夏季的南黄海中央海域,由于上混合层营养盐几乎耗尽,限制了浮游植物的生长,在紧贴温跃层下部的真光层,具有丰富的营养盐和合适的光照,次表层叶绿素a极大值得以形成.秋季(约9～11月份,略迟于海表面开始降温的时间,随地点不同而异)随垂直混合的增强,有利于营养盐向上输运,浮游植物出现一次较小的峰值.     

Seasonal cooling and blooming in tropical oceans

The relative importance of tropical pelagic algal blooms in not yet fully appreciated and the way they are induced not well understood. The tropical Atlantic supports pelagic blooms together equivalent to the North Atlantic spring bloom. These blooms are driven by thermocline tilting, curl of wind stress and eddy upwelling as the ocean responds to intensified basin-scale winds in boreal summer. The dimensions of the Pacific Ocean are such that seasonal thermocline tilting does not occur, and nutrient conditions are such that tilting might not induce bloom, in any case. Divergence at the equator is a separate process that strengthens the Atlantic bloom, is more prominent in the eastern Pacific, and in the Indian Ocean induces a bloom only in the western part of the ocean. Where western jet currents are retroflected from the coast off Somalia and Brazil, eddy upwelling induces prominent blooms. In the eastward flow of the northern equatorial countercurrents, positive wind curl stress induces Ekman pumping and the induction of algal blooms aligned with the currents. Some apparent algal bloom, such as that seen frequently in CZCS images westwards from Senegal, must be due to interference from airborne dust.     

Spatial and temporal patterns in the distribution and abundance of macrozooplankton on the southern North West Shelf, Western Australia

We describe the spatial and temporal dynamics of macrozooplankton on the southern North West Shelf (NWS) of Australia over two consecutive summers (1997/1998 and 1998/1999). Sampling was conducted using submersible light traps, deployed at the surface and at depth, along one cross-shelf transect in 1997/1998, and along two cross-shelf transects and one long-shore transect in 1998/1999. Our results revealed large inter-annual changes in macrozooplankton assemblages during the two summers. An upwelling regime associated with El Niño conditions prevailed during the first summer, resulting in high chlorophyll concentrations and an abundance of both meso- and macrozooplankton. During this time, there were distinct inshore and offshore macrozooplankton assemblages that reflected an abrupt transition from upwelling-enriched shelf waters to oligotrophic oceanic waters. In contrast, the second summer was characterised by a downwelling regime associated with La Niña conditions that resulted in low chlorophyll concentrations and decreased abundances of meso- and macrozooplankton. The lack of upwelling activity weakened horizontal gradients in macrozooplankton assemblages so that differences between surface and deep assemblages were more pronounced. Catches in light traps were dominated by hyperiid amphipods during the first summer and by the neritic euphausiid Pseudeuphausia latifrons during the second. There was little within-season change in macrozooplankton assemblages during both summers, and only weak relationships were found between the environmental parameters and the distribution of individual macrozooplankton taxa.     

Eddy-induced cross-shelf phytoplankton transport along the downwelling coast off Western Australia

Along the downwelling coast off Western Australia, late-autumn/early-winter chlorophyll a blooms are observed on the continental shelf south of Shark Bay (26°S), in contrast with summer blooms in the north. The late-autumn/early-winter blooms are in phase with seasonal strengthening of the Leeuwin Current and its eddy field. Anticyclonic eddies entrain the high phytoplankton biomass waters from the shelf and transport offshore into the oligotrophic, subtropical marine environment, as revealed by coalescing the finite-size Lyapunov exponent (FSLE) of the surface geostrophic flow field and the satellite chlorophyll a images.     

Hydrographic and atmospheric analysis of an autumnal upwelling event in the Ria of Vigo (NW Iberian Peninsula)

An autumnal upwelling event was observed in the Ria of Vigo (NW Iberian Peninsula) on 15th November 2001. This event was analyzed by means of thermohaline variables measured at CTD stations located in the study area, satellite sea surface temperature and wind data provided by QuikSCAT. Salinity and temperature distributions revealed that the upwelled water mass was Eastern North Atlantic Central Water (ENACW), typically observed during summer upwelling events. However, previous to the upwelling event, the characteristic autumnal body of water was recorded on 31st October 2001. Subsequent sea surface temperature and upwelling index corroborated the presence of an autumnal upwelling event of approximately 15 days duration. The probability of upwelling-favorable winds (from 1999 to 2004) was lower during autumn–winter than during spring–summer, although they may occur at any time of the year. Probabilities of 45% were calculated for February and November, with the highest probability (65%) corresponding to July.     

Spatial and temporal aspects of phytoplankton blooms in complex ecosystems off the korean coast from satellite ocean color observations

Complex physical, chemical and biological interactions off the Korean coast created several striking patterns in the phytoplankton blooms, which became conspicuous during the measurements of ocean color from space. This study concentrated on analyzing the spatial and temporal aspects of phytoplankton chlorophyll variability in these areas using an integrated dataset from a Sea-viewing Wide Field-of-view Sensor (SeaWiFS), Advanced Very High Resolution (AVHRR) sensor, and Conductivity Temperature Depth (CTD) sensor. The results showed that chlorophyll concentrations were elevated in coastal and open ocean regions, with strong summer and fall blooms, which appeared to spread out in most of the enclosed bays and neighboring waters due to certain oceanographic processes. The chlorophyll concentration was observed to range between 3 and 54 mg m^-3 inside Jin-hae Bay and adjacent coastal bays and 0.5 and 8 mg m^-3 in the southeast sea offshore waters, this gradual decrease towards oceanic waters suggested physical transports of phytoplankton blooms from the shallow shelves to slope waters through the influence of the Tsushima Warm Current (TWC) along the Tsushima Strait. Horizontal distribution of potential temperature (θ) and salinity (S) of water off the southeastern coast exhibited cold and low saline surface water (θ<19°C; S<32.4) and warm and high saline subsurface water (θ>12°C; S>34.4) at 75dBar, corroborating TWC intrusion along the Tsushima Strait. An eastward branch of this current was called the East Korean Warm Current (EKWC), tracked with the help of CTD data and satellite-derived sea surface temperature, which often influenced the dynamics of mesoscale anticyclonic eddy fields off the Korean east coast during the summer season. The process of such mesoscale anticyclonic eddy features might have produced interior upwelling that could have shoaled and steepened the nutricline, enhancing phytoplankton population by advection or diffusion of nutrients in the vicinity of Ulleungdo in the East Sea.     

Modeling the seasonal variability of marine ecosystem in the region of the Central-Eastern Atlantic

A 3D eco-hydrodynamical model of high resolution (0.25° × 0.25°, 27 σ-levels) is used to simulate the seasonal variability of the ocean circulation and marine ecosystem in the Central-Eastern Basin of the North Atlantic including the Canary upwelling system. According to the model results, in the winter period, the “patches” of maximal phytoplankton and zooplankton biomass are often located in upwelling zones in the open ocean on the periphery of cyclonic eddies rather than in the coastal upwelling zones. In the summer period, when the phytoplankton biomass reaches maximal (in the annual cycle) values, the maxima of the phytoplankton are located in the coastal upwelling zones. As shown, there is no simple relationship between the nitrate distributions, on the one hand, and the phytoplankton and zooplankton ones, on the other hand.     

A four-component ecosystem model of biological activity in the Arabian Sea

A coupled, physical-biological model is used to study the processes that determine the annual cycle of biological activity in the Arabian Sea. The physical model is a system with a surface mixed layer imbedded in the upper layer, and fluid is allowed to move between layers via entrainment, detrainment and mixing processes. The biological model consists of a set of advective-diffusive equations in each layer that determine the nitrogen concentrations in four compartments: nutrients, phytoplankton, zooplankton and detritus. Coupling is provided by the horizontal-velocity, layer-thickness, entrainment and detrainment fields from the physical solution. Surface forcing fields (such as wind stress and photosynthetically active radiation) are derived from monthly climatological data, and the source of nitrogen for the system is upward diffusion of nutrients from the deep ocean into the lower layer. Our main-run solution compares favorably with observed physical and biological fields; in particular, it is able to simulate all the prominent phytoplankton blooms visible in the CZCS data. Three bloom types develop in response to the physical processes of upwelling, detrainment and entrainment. Upwelling blooms are strong, long-lasting events that continue as long as the upwelling persists. They occur during the Southwest Monsoon off Somalia, Oman and India as a result of coastal alongshore winds, and at the mouth of the Gulf of Aden through Ekman pumping. Detrainment blooms are intense, short-lived events that develop when the mixed layer thins abruptly, thereby quickly increasing the depth-averaged light intensity available for phytoplankton growth. They occur during the fall in the central Arabian Sea, and during the spring throughout most of the basin. In contrast to the other bloom types, entrainment blooms are weak because entrainment steadily thickens the mixed layer, which in turn decreases the depth-averaged light intensity. There is an entrainment bloom in the central Arabian Sea during June in the solution, but it is not apparent in the CZCS data. Bloom dynamics are isolated in a suite of diagnostic calculations and test solutions. Some results from these analyses are the following. Entrainment is the primary nutrient source for the offshore bloom in the central Arabian Sea, but advection and recycling also contribute. The ultimate cause for the decay of the solution's spring (and fall) blooms is nutrient deprivation, but their rapid initial decay results from grazing and self shading. Zooplankton grazing is always an essential process, limiting phytoplankton concentrations during both bloom and oligotrophic periods. Detrital remineralization is also important: in a test solution without remineralization, nutrient levels drop markedly in every layer of the model and all blooms are severely weakened. Senescence, however, has little effect: in a test solution without senescence, its lack is almost completely compensated for by increased grazing. Finally, the model's detrainment blooms are too brief and intense in comparison to the CZCS data; this difference cannot be removed by altering biological parameters, which suggests that phytoplankton growth in the model is more sensitive to mixed-layer thickness than it is in the real ocean.     

Consequences of winter upwelling events on biogeochemical and phytoplankton patterns in a western Galician ria (NW Iberian peninsula)

The consequences of two upwelling events in mid- (MW) and late (LW) winter on biogeochemical and phytoplankton patterns were studied in the Pontevedra Ria and compared with the patterns measured under typical winter conditions and under a summer upwelling event. Thermohaline patterns measured during the mid-winter upwelling event (MW-up) revealed the intrusion of saltier seawater (35.9) into the ria associated with the Iberian Poleward Current (IPC). During the late-winter upwelling event (LW-up), the seawater which had welled up into the ria showed characteristics of the Eastern North Atlantic Central Water mass (ENACW). In both cases the measured water residence time (4 days during MW-up and 10 days during LW-up) was related to both meteorological and fluvial forcing. This residence time contrasts with that of summer upwelling (7 days) and with that estimated under unfavorable upwelling atmospheric conditions (2–4 weeks). During MW-up, the ria became poor in nutrients due to continental freshwater dilution, associated with the shorter residence time of the water, and the intrusion of IPC, which is a water body poor in nutrient salts: 2.9 μM of nitrate, 0.1 μM of phosphate and 1.5 μM of silicate. During this event, the ria exported 3.4 mol_DIN s⁻¹, compared with 6.9 mol_DIN s⁻¹ in non-upwelling conditions. Phytoplankton showed a uniform distribution throughout the ria, as during unfavorable upwelling conditions, and was characterized by the dominance of diatoms, mainly Nitzschia longissima and Skeletonema costatum. During LW-up, a nutrient depletion in the photic layer also occurred, but as a result of a phytoplankton spring bloom developing at this time. The ria was a nutrient trap where 4.1 mol_DIN s⁻¹ were processed by photosynthesis. This budget is three times higher than the one under non-upwelling conditions. In contrast with the MW-up, which had no effect on primary production, during LW-up the ria became more productive, although not as productive as during a summer upwelling event (9.9 mol_DIN s⁻¹). The taxonomic composition of the phytoplankton community did not change noticeably during LW-up and the summer upwelling, with the same species present and changing only in relative proportions. Diatoms were always the dominant microphytoplankton community, with Pseudonitzschia pungens, Thalassionema nitzschioides and several species of Chaetoceros as characteristic taxons.     

Global distribution of summer chlorophyll blooms in the oligotrophic gyres

Chlorophyll blooms consistently develop in the oligotrophic NE Pacific in late summer, isolated from land masses and sources of higher chlorophyll waters. These blooms are potentially driven by nitrogen fixation, or by vertically migrating phytoplankton, and a better understanding of their ubiquity could improve our estimate of the global nitrogen fixation rate. Here, global SeaWiFS chlorophyll data from 1997 to 2007 are examined to determine if similar blooms occur in other oligotrophic gyres. Our analysis revealed blooms in five other areas. Two of these are regions where blooms have been previously identified: the SW Pacific and off the southern tip of Madagascar. Previously, unnoticed summer blooms were also identified in the NE and SW Atlantic and in a band along 10°S in the Indian Ocean. There is considerable variation in the intensity and frequency of blooms in the different regions, occurring the least frequently in the Atlantic Ocean. The blooms that develop along 10°S in the Indian Ocean are unique in that they are clearly associated with a hydrographic feature, the 10°S thermocline ridge, which explains the bloom within a conventional upwelling scenario. The environment and timing of the blooms, developing in oligotrophic waters in late summer, are conducive to both nitrogen fixers and vertically migrating phytoplankton, which require a relatively stable water column. However, the specific locations of the chlorophyll blooms generally do not coincide with areas of maximum levels of nitrogen fixation or Trichodesmium. The NE Pacific chlorophyll blooms develop in a region with a very high SiO₄/NO₃ ratio, where silicate will not be a limiting nutrient for diatoms. The blooms often develop between eddies, wrapping around the periphery of anti-cyclonic features. However, none of the areas where the blooms develop have particularly high eddy kinetic energy, from either a basin-scale or a mesoscale perspective, suggesting that other factors, such as interactions with a front or dynamics associated with the critical latitude, operate in conjunction with the eddy field to produce the observed blooms.