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Pigment patterns and phytoplankton assemblages in different water masses of the Ri'o de la Plata maritime front. High chl a concentrations were recorded in the outer estuary region (up to 15. Mg m a3) and in the shelf-break front associated with Subantarctic waters.
Pigment patterns and phytoplankton assemblages in different water masses of the Ri'o de la Plata maritime front. High chl a concentrations were recorded in the outer estuary region (up to 15. Mg m a3) and in the shelf-break front associated with Subantarctic waters.
Pigment patterns and phytoplankton assemblages in different water masses of the Ri'o de la Plata maritime front. High chl a concentrations were recorded in the outer estuary region (up to 15. Mg m a3) and in the shelf-break front associated with Subantarctic waters.
Algal pigment patterns and phytoplankton assemblages in different water
masses of the Ro de la Plata maritime front
Jose I. Carreto
, Nora Montoya, Rut Akselman, Mario O. Carignan,
Ricardo I. Silva, Daniel A. Cucchi Colleoni Instituto Nacional de Investigacion y Desarrollo Pesquero, Paseo V, Ocampo No. 1, B7602HSA-Mar del Plata, Argentina a r t i c l e i n f o Article history: Received 1 November 2006 Received in revised form 24 February 2007 Accepted 28 February 2007 Available online 19 March 2008 Keywords: Phytoplankton Pigments CHEMTAX Photo-adaptation South America Argentina a b s t r a c t The composition of phytoplankton assemblages were studied in three sections across the continental shelf between the Ro de la Plata and the oceanic waters of the Subtropical Convergence, during late spring. Algal communities were examined using microscopy and HPLC-derived pigment concentrations. The CHEMTAX program was used to estimate the chlorophyll a (chl a) biomass of different algal classes. Trends in pigment ratios due to phytoplankton photo-adaptation and photo-acclimation were also examined. In order to accommodate the natural diversity of phytoplankton assemblages the original data have been split to represent ve ecosystems. In addition, the pigment data for the Brazil Current ecosystem has been split by sample depth. High chl a concentrations were recorded in the outer estuary region (up to 15.5 mg m 3 ) and in the shelf-break front associated with Subantarctic waters (24mg l 1 ). In contrast, chl a concentrations were relatively low over the continental shelf and in the oceanic region dominated by the Brazil Current, where the lowest values (0.10.2 mg l 1 ) were found. Both pigment patterns and microscopy-derived information showed ve different phytoplankton assemblages spatially segregated by the prevailing environmental conditions. In the inner estuary assemblage green algae (5456% of total chl a) were always the dominant group and most of the chl a, arises from chlorophyceans (4049%). In a decreasing order, diatoms cyanobacteria and cryptophytes were also relevant. In the outer estuary assemblage diatoms and dinoagellates were the dominant groups but cryptophytes and euglenophytes were present as sub-dominant groups. In the coastal and shelf region, the algal assemblage showed an almost total dominance (59.387.6%) of diatoms. The usual diatom-pigment pattern (chl c 1 , chl c 2 ) group (diatom I), was the more abundant and widely distributed, but in some stations, diatoms containing chl c 2 and chl c 3 (diatom II) were present as dominant group. A more complex phytoplankton community dominated by coccoid and small agellates (25mm) predominantly comprised by chlorophyceans (up to 50%) and haptophytes (up to 62%) was found near the shelf-break front. This is the rst time that high chl b concentrations associated to a bloom of a picoplanktonic (o3mm) coccal chlorophycean was reported for this area. The Brazil Current assemblage showed the dominance (55.471.9%) of the picoplanktonic cyanobacteria Synechococcus spp. (32.345%) and Prochlorococcus spp. (41.410.4%). Haptophytes were also present as sub-dominant group being particular abundant at the deep chl a maxima. A sharp transition in photo-collectors/(chl a+Dv chl a) and photo-protectors/(chl a+ Dv chl a) ratios at depth near the base of the euphotic zone was observed in the water column of this ecosystem. These results are discussed in relation to the complex environmental features of the region. & 2008 Elsevier Ltd. All rights reserved. 1. Introduction The physical oceanography and the sediment transport processes of the Ro de la Plata estuary and plume are well understood and documented (Framinan and Brown, 1996; Guerrero et al., 1997; Framinan et al., 1999; Campos et al., 1999; Piola et al., 2000, 2005; Mianzan et al., 2001; Simionato et al., 2004). Recently, Acha et al. (2008) summarize present knowledge, concluding that the estuary is a highly variable environment, strongly stratied most of the time but that can be mixed during a few hours by strong wind events that occur in an unpredictable manner, generating stratication/partially mixed pulses all along the year (Acha et al., 2008). The shallow, high turbidity tidal river is separated from the mixohaline area by a turbidity front, closely ARTICLE IN PRESS Contents lists available at ScienceDirect journal homepage: www.elsevier.com/locate/csr Continental Shelf Research 0278-4343/$ - see front matter & 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.csr.2007.02.012
E-mail address: jcarreto@inidep.edu.ar (J.I. Carreto). Continental Shelf Research 28 (2008) 15891606 related to the salinity front (Framinan and Brown, 1996). Within the nutrient rich turbid river, phytoplankton growth is lightlimited (Carreto et al., 2003; Acha et al., 2004, 2008; Huret et al., 2005). In the outer estuary zone the phytoplankton production appears to be regulated by light penetration and the assimilable nitrogen ow, while silicate and phosphate are in excess (Carreto et al., 1986; Nagy et al., 1997). In this area, the concentrations of inorganic nutrients decrease rapidly with the decrease of turbidity along the salinity gradient (Carreto et al., 2007; Huret et al., 2005). Simionato et al. (2004), showed that in the absence of winds the normal path of the estuary ow should be a buoyant plume to the NNE direction along the Uruguayan coastline. When the mean wind blows from directions between SSE and NNW this pattern is intensied. During winter, the inuence of the plume has even been noticed along the Brazilian coast as far as 281S (Campos et al., 1999), while in summer it is constrained to south of 321S (Piola et al., 2000). However, during summer its inuence may be detected sporadically in a southward direction along the coastal region of Argentina (Carreto et al., 2007, and references therein). Even though a seasonal (summer and winter) circulation pattern has been reported (Guerrero et al., 1997) recent model studies suggest that variability in the estuarine circulation is highly sensitive to the atmospheric forcing and that the dynamic condition of summer and winter are likely to occur during any season (Simionato et al., 2004, 2006). Over the continental shelf, a sharp contrast in water mass character- istics exists below the low salinity plume. The relatively cold nutrient-rich Subantarctic Shelf water dominates south of 331S, while warm, salty Subtropical Shelf water extend north of that latitude. These water masses are separated by relatively narrow frontal zone in subsurface referred to as Subtropical Shelf Front (Piola et al., 2000). Occasionally, the inuence of the Ro de la Plata waters may extend to the shelf-break front where these estuarine waters contact the nutrient-rich waters of Malvinas ow (Negri et al., 1992), and the warm, nutrient-poor subtropical Brazil Current water or it may be found between both currents (Lusquinos and Valde s, 1971; Provost et al., 1995). These waters can be carried offshore by the return ux of the Malvinas Current (Provost et al., 1995) in an area referred to as the Subtropical Convergence. However, relatively few phytoplankton studies have carried out in this region (Carreto et al., 2007 and references therein) and the recognition of the importance of delicate agellates and minute coccoid forms has been recognized only recently, after more appropriate techniques for studying the picoplankton were applied (Silva and Negri, 2000; Carreto et al., 2003). Using these approach, diatoms, cryptophytes, prasinophytes, dinoagellates, haptophytes and cyanobacteria were found forming different assemblages spatially segregated by the prevailing hydrographic conditions (Carreto et al., 2003). Pigment chemo-taxonomic approach based on chromato- graphic pigment analysis has been useful for distinguishing the main algal classes, especially of the small-sized phytoplankton. The signature role of the pigments has been summarized in various papers, although the interpretation of HPLC eld data is not a straightforward task (Jeffrey and Wright, 1997). The CHEMTAX program has proven to be a solid method to calculate the abundance of phytoplankton groups, if correct information on the phytoplankton groups present in the sample is loaded in the program. Screening the samples in the microscope prior to data analysis, minimize the possibility of overlooking phytoplankton groups with no or overlapping diagnostic pigments (Wright et al., 1996; Wright and van den Enden, 2000; Mackey et al., 1998; Rodrguez et al., 2002; Carreto et al., 2003; Schlu ter and Mhlenberg, 2003; Wright and Jeffrey, in press). However, choosing starting pigments/chlorophyll a (chl a) ratios for the CHEMTAX remains the biggest problem due to insufcient knowledge of these ratios in the eld (Schlu ter et al., 2000; Wright and Jeffrey, in press). Recently, concurrent analyses of microscope enumeration of phytoplankton species, HPLC-derived pigment concentrations and the CHEMTAX software were used for the rst time in this region to evaluate the surface distribution and abundance of the different phytoplankton groups (Carreto et al., 2003). Results showed that the inclusion in the pigment matrix of the most abundant members of the chlorophyll c (chl c) pigment family (chl c 1 , chl c 2 , chl c 3 and chl c 2 monogalactosyldiacylglyceride esters) improved CHEMTAX interpretation of eld data (Carreto et al., 2003; Rodrguez et al., 2002). Using this novel approach ve different surface phytoplankton assemblages, spatially segregated by the prevailing environmental conditions, were distinguished across the studied section. All of them showed a complex community structure, formed by a background of small-sized cells such as cyanobacteria, cryptophytes, haptophytes and prasinophyceans on which, in the more eutrophic waters, diatom, cryptophytes or some haptophyte blooms were overlapped (Carreto et al., 2003). In the surface phytoplankton community structure of the nutrient-poor Subtropical Brazil Current Waters the picoplanktonic cyanobacteria Synechococcus, appeared to be the most important group (Carreto et al., 2003) although the variability in the vertical distribution has not yet been reported. Recently, Barlow et al. (2002) showed that divinyl chl a (DV chl a), an exclusive biomarker of Prochloroccocus, was prominent at depth in the tropical and subtropical region of the Atlantic Ocean from 401N to 351S. The highest DV chl a concentrations were observed in the equatorial region at the chlorophyll maxima located at 7080m (Barlow et al., 2002). In this study, the structure of phytoplankton communities was studied during November 2001 in tree sections across the continental shelf between the Ro de la Plata and the oceanic waters of the Subtropical Convergence (Fig. 1), using microscopy, HPLC-pigment patterns and the chemical taxonomy software CHEMTAX. In addition, we describe the relationship between phytoplankton communities and the environmental features of the region. 2. Material and methods 2.1. Sample collection Sampling procedures were performed from 19 November to 1 December 2001, on board the R.V. Dr. E.L. Holmberg (Fig. 1). Vertical proles for temperature and salinity were obtained with a CTDSBE911. CTD data were calibrated and reduced to 1-m bins. Light penetration was measured with a PUV 500 (Biospherical Instruments Inc.) underwater radiometer. Surface water samples (0m depth) were collected with a clean plastic bucket, and discrete sampling of the water column was carried out using Niskin bottles attached to the CTD rosette. 2.2. Remote-sensing data of SST and ocean color Sea surface temperature (SST) eld was derived from data collected by the Advanced Very High Resolution Radiometer (AVHRR) installed on the NOAA 11 polar satellite and processed by the Argentine National Space Research Commission (CONAE) using a multi-channel algorithm. Chlorophyll images were pro- vided by the SeaWiFS Project, NASA/Goddard Space Flight Center and ORBIMAGE (http://seawifs.gsfs.nasa.gov/) processed to level 3. ARTICLE IN PRESS J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1590 2.3. Microscopic analysis Surface water sub-samples (0m depth) for phytoplankton identication and cell counts were obtained from selected stations and xed with neutral Lugols iodine. Sample volumes of 50 and 100ml were settled. Phytoplankton species were identied and enumerated with an Olympus IX-70 inverted microscope using bright-eld optics and 200, 400, 600 and 1000 (oil) magnications. Algal cells were identied to species level when possible, except for the small and fragile ones for which only the genus or algae group was recorded. In occasions, water samples consisting about 50100ml were ltered through polycarbonate membrane (0.2mm pore size, 47mm diameter) and left in plastic Petri slides. A portion was cut from each lter and glued to aluminum stubs, which were coated with gold/ palladium and examined by scanning electron microscopy (Jeol 6460-LV). 2.4. Pigment analysis Seawater samples of 0.53l were ltered through Whatman GF/F glass ber lters. Filters were deep-frozen immediately and stored in liquid nitrogen to prevent pigment modication. Pigments were extracted with 100% methanol and sonicated (Vibra Cell, Sonic and Materials) 01C. The extracts were ltered through GF/F lters to remove cell debris. Water was added to the extract immediately before injection to obtain an 80% methanol dilution (Jeffrey and Wright, 1997). Sample solution aliquots were automatically injected into an HPLC system Shimadzu LC 10 AC. For pigment elution we use the method of Zapata et al. (2000). A C8 column (symmetry 1504.6mm 2 , 3.5mm particle size, 100 A pore size) was used protected with a C8 (Symmetry) guard column. The mixing chamber and column were thermostated at 251C with a CTO-10 AC (Shimadzu) column oven. Mobile phases were: (A) methanol:acetonitrile:aqueous pyridine solution (0.25M, pH adjusted to 5.0 with acetic acid) (50:25:25, v/v/v), and (B) methanol:acetonitrile:acetone (20:60:20, v/v/v). A linear gradient from 0% to 40% B was pumped for 22min, followed by an increase to 95% at minute 28 and isocratic hold at 95% B for further 10min. Peak detection was carried out using a diode array detector (SPD-M10Avp). Chlorophylls were also detected by uorescence: Excitation at 440 and emission at 650nm (spec- tral-uorometer FR-10Axl). Pigments were identied by their retention time and absorption spectra obtained from the on-line diode array detector (350750nm). High-purity standards were provided by VKI (The International Agency for 14C Determination, Denmark) or isolated from cultures of Emiliania huxleyi (clone CCMP370) and Alexandrium tamarense (clone MDQ1096). The HPLC system was calibrated using genuine standards from VKI or the extinction coefcients reported by Jeffrey and Wright (1997) for prepared standards. Novel pigments were quantied consider- ing the extinction coefcients for the most similar documented chromophore (e.g. DV chl a was measured as chl a, and chl c 3 and chl c 2 -MGDG esters were measured as chl c 2 equivalents). 2.5. Data processing The contribution of different phytoplankton groups to the total chl a, at each sampled station were calculated using CHEMTAX, a matrix factorization program running under MATLAB (Mackey et al., 1996). All diagnostic pigments detected, as well as those ambiguous markers indicative of only a fewgroups, were included in the program matrix. Chl c 1 , chl c 2 , chl c 3 and chl c 2 -MGDG- esters were also included, as the HPLC method used is very useful to separate these pigments (Zapata et al., 2000). The original data has been split into ve geographic groups. The distinction of ve groups accommodated the natural diversity of phytoplankton assemblages between the different ecosystems previously distin- guished for this area (Carreto et al., 2003). These categories were delineated on the basis of: (1) identication of pigment assem- blages and their relation to the hydrography and (2) the microscopic analysis of representative samples fromeach pigment assemblage. In addition, using the photo-collector pigments (PCPs)/total chl a and the photo-protector pigments (PPPs)/total ARTICLE IN PRESS Fig. 1. AVHRR sea surface temperature (1C) mapped image from 27 November 2001, showing location of occupied stations and distribution of surface salinity (PSU). J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1591 chl a ratios as indicator of phytoplankton photo-adaptation, the pigment data for the more oceanic ecosystem has been split by sample depth. For each group, a matrix using initial pigment: chl a ratios derived from Carreto et al. (2003 and references therein), supplemented by recent surveys (Van Lenning et al., 2003; Latasa et al., 2004; Zapata et al., 2004) was constructed, since data on pigment ratios and their changes with light climate for the dominant phytoplankton species of the studied area were not available. It should be emphasized that in some cases a pigment group may be composed of several phytoplankton taxonomic classes. 3. Results and discussion 3.1. Environmental conditions The satellite-derived SST data on 29 November (Fig. 1) showed that the Brazil Current separated from the shelf-break at 361S where it encountered a narrow lament of cooler waters owing along the upper shelf-break between 391S and the latitude of Brazil Current separation. This transition zone of intermediate temperatures seems to be lled with patches and laments of Subantarctic and Subtropical waters. On the shelf the occurrence of well-dened thermal fronts was also evident: (1) a thermal front between the Ro de la Plata outow and coastal waters, (2) a thermal front separating Coastal from Subantarctic waters and (3) a thermal front between the shelf waters and the Subtropical Brazil Current waters. In coincidence with SST data, surface salinity distribution (Fig. 1) indicated that the Ro de la Plata outow forms a low-salinity tongue which extends northward along the Uruguayan coast, with salinity between 28 and 30psu (Sts. 26, 25). In the head of the estuary (Sts. 20, 19, 18), and in the more shallow outer estuarine stations (Sts. 17, 16) (Fig. 1), the vertical distribution of temperature (Fig. 2a) and salinity (Fig. 2b) showed no stratication. Under calm winds freshwater overlying sea water denes a salt wedge with a typical scale of 100km (Guerrero et al., 1997). However, during the cruise the prevailing strong onshore winds (mainly SE and ESE) eroded the quasi- permanent halocline by wind-induced vertical mixing. The inuence of the Ro de la Plata extends offshore up to the surface salinity front (Mianzan et al., 2001) capping the coastal waters (3033.5psu) produced by the mixing of the estuarine and the Subantarctic Shelf waters (Carreto et al., 2003). In the central section, the vertical light distribution showed (Fig. 2c) that in the head of the estuary the incident photo- synthetically active radiation (PAR) was reduced to less than 10% in the rst centimetres depth and that the ratio of mixing depth to euphotic depth (Z m :Z eu ) was about 3. In contrast, the highest transmission values were observed at the oceanic region of the oligotrophic Brazil Current waters, where the euphotic zone reached up to 80m depth. In this region the Z m :Z eu ratios were about 0.2. Intermediate values were observed in the coastal and shelf regions. 3.2. Chlorophyll a distribution The surface chl a distribution based on the SeaWiFS imagery (Fig. 3A) and the obtained from sampled in situ data (Fig. 3B) showed the same distributional pattern. In both cases data reveals a band along the edge of the shelf extended northward up to the latitude of the Brazil Current separation, where high surface chlorophyll concentrations (34mg l 1 ) were found (Fig. 3A). This chlorophyll band has been regularly observed from satellites on the shelf-break off Argentina during late spring and summer (Podesta , 1997; Brandini et al., 2000; Carreto et al., 2003; Romero et al., 2006). The chl a concentrations in the water column were also high, especially at St. 10 where a chl a maximum (5.81mg l 1 ) near the base of the euphotic zone was found (Fig. 4b). In contrast, at the edge of the shelf area dominated by the warm, nutrient-poor waters of the Brazil Current the lowest surface chl a concentration (0.10.2mg l 1 ) were found (Fig. 4a). Another characteristic of this area is the presence of a deep chl a maximum situated near the base of the euphotic layer at St. 24. Another noteworthy area was the Ro de la Plata estuary, where an extremely high Sea WiFS chlorophyll signal was found (Fig. 3A). However, in situ surface chl a measured was very much lower (o1.4mg l 1 ) than that obtained by the Sea WiFS (20.0mg l 1 ) and did not vary signicantly with depth (Fig. 4b). As has been previously reported, the application of the classical algorithms (OReilly et al., 2000) over-estimate the chl a concentration in the Ro de la Plata estuary, mainly at stations near the turbidity front (Armstrong et al., 2004; Carreto et al., 2003; Huret et al., 2005). In the outer estuary area in situ surface chl a was much higher (up to 15.5mg l 1 ) (Fig. 4b). However, due to constrains imposed by the light limitation, the total chl a, integrated over the euphotic layer of the outer estuary area were much lower (up to 46.3mg m 2 , St. 17) to that recorded at the shelf-break front (up to 81.2mg m 2 , St. 10). The surface chl a concentrations over the entire shelf were lower than that observed in the mentioned frontal areas. However, in some shelf stations high chl a concentrations near the base of the euphotic zone was found (Fig. 4). ARTICLE IN PRESS Fig. 2. Temperature, salinity and solar radiation along the central section. (a) Temperature distribution (1C). (b) Salinity distribution (PSU). (c) Photosynthetic active radiation (PAR) penetration distribution as percentage of the incident radiation. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1592 3.3. Phytoplankton communities, photo-adaptation and photo- acclimation Based on previous results (Carreto et al., 2003, 2007 and references therein), the obtained pigment patterns (Table 1 and Fig. 5) and the species composition of the surface phytoplankton communities (Table 2) ve phytoplankton communities (see Fig. 3B) closely related with the hydrography of the area were distinguished: (a) inner estuary, (b) outer estuary, (c) coastal, (d) Subantarctic Shelf-break front, and (e) Brazil Current. The chromatograms obtained (Fig. 5) from six selected stations showed the characteristic pigment patterns that prevailed in the considered phytoplankton assemblages. In addition to water masses and nutrient conditions, factors controlling the integrated irradiance exposure (transparency of the water and vertical mixing) should be considered to breaking the data set up (Mackey et al., 1998; Wright and van den Enden, 2000). Trends in pigment ratios due to phytoplankton photo-adaptation and phytoplankton photo-acclimation (used here to describe reversible light-induced alterations in the physiological character- istics of a population) were examined using the water column variability in the PCP and in the PPPs/chl a ratios (Fig. 6). As in most eld studies in estuarine and coastal zones (Fig. 6a), the PCP/chl a ratio observed in the water column showed small variability with irradiance, probably because the phytoplankton in the mixed layer never get the chance to acclimate their pigmentation to a constant irradiance over a generation time and because they experienced a similar average light exposure. However, the synthesis of the photo-protective carotenoids diadinoxanthin (Diadino) and diatox- anthin (Diato) is a more rapid process (hours) and the variability in the PPP/chl a ratio with the measured sample depth or irradiance appears to be evident (Fig. 6a). On the other hand, in stratied waters photo-acclimation of phytoplankton populations is usually reected by a change in pigment ratios. Dramatic changes in the PCP/Total chl a (chl a+Dv chl a) ratio was observed in the water column where subtropical waters dominated (Sts. 8 and 22), ranging from 0.4 in the surface layer to 1.1 near the base of the euphotic layer (Fig. 6b). Inversely, the PPP/Tchl a ratio decreased from 0.6 in the surface to 0.1 at the base of the euphotic layer (Fig. 6b). The sharp transition of these ratios at deep near the base of the euphotic zone may be due in part to photo-acclimation processes, but mainly to changing differently photo-adapted populations (Goericke and Repeta, 1993; Moore et al., 1995). These results indicate that the pigment data of this oceanic ecosystem should be split by sample depth. Since the small number of samples adversely affected the accuracy of the CHEMTAX calcula- tion only two pigment matrix (surface to 5% PAR and lower than 5% PAR) were considered in our calculation. ARTICLE IN PRESS Fig. 3. Surface chlorophyll a distribution (mg l 1 ). (A) SeaWiFS level 3 standard mapped image for chlorophyll, 1724 November composite. (B) Data derived from HPLC measurements of pigment extracts from surface samples. The ve identied phytoplankton communities are indicated as follows: (a) inner estuary; (b) outer estuary; (c) coastal; (d) Subantarctic Shelf-break front and (e) Brazil Current. Fig. 4. Vertical chlorophyll a concentration (mg l 1 ) from HPLC measurements of pigment extracts along the sections. (a) Northern section. (b) Central section. (c) Southern section. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1593 Changes in pigment ratios (PCP/Total chl a, and PPP/Tchl a) with depth showed a more complex scenario at the shelf-break front. Although in most of the studied stations, the PCP/total chl a ratio increased moderately with depth, variability between stations are evident, changing from about 1.070.2 in the surface layer to 1.270.2 at the base of the euphotic zone (Fig. 6c). In contrast, the high PCP/total chl a ratio observed (1.570.1) at the surface of station 10 do not changed with sample depth or water column irradiance. Nevertheless, as can be expected by the rapid synthesis of photoprotective carotenoids, the PPP/total chl a ratio decreased signicantly with decreasing irradiance in all stations, ranging from about 0.470.15 at the surface layer to 0.270.15 near the base of the euphotic zone (Fig. 6c). The studied stations are situated at the northern Subantarctic Shelf-break front, near the high SST gradient region associated to the Brazil/Malvinas Conuence where the Malvinas Current veers offshore (Podesta , 1997; Saraceno et al., 2005). Recent numerical simulations suggest that bottom friction associated to the presence of a strong slope current, such as the Malvinas Current, creates along-shelf pressure gradients and lead to robust upwelling at the shelf break (Romero et al., 2006). The heterogeneity of the frontal area and the instability related with the alternation of upwelling and vertical stratication periods appear to be relevant processes explaining the strong contrast observed in photo-acclimation stage of these phytoplankton populations. Another contributing factor is the high complexity in the pigment patterns and phytoplankton composition observed in this region. Therefore, the pigment data for this ecosystem has not been split by sample depth and only one pigment matrix was considered in our CHEMTAX calculation. 3.4. Pigment pattern and abundance of algal groups 3.4.1. Inner estuary community Pigments associated with green algae (Chlorophyceae and Prasinophyceae) were relevant in the inner estuary. Although the amount of chl b was not high in this ecosystem, luteinthe major carotenoid of chlorophyceans, was found in high proportion in this assemblage (Figs. 5a and 7). Prasinoxanthin (Pras), an unequivocal marker of some prasinophyceans (Mamiellales, Peudoscoureldiales and Prasinococcales, Latasa et al., 2004), was only detected in trace amounts. Consequently, more of the chl b arises from chlorophyceans or prasinophyceans lacking Pras. The microscopic examination of these samples showed that a not identied palmeloid chlorophycean from freshwater origin was the most abundant green algae, indicating that most of the chl b detected arises from chlorophyceans. Fucoxanthin (Fuco) and zeaxanthin (Zea) were also found in high proportion in the inner estuary (Fig. 5a). Microscopic analysis indicated that several diatoms species present in low abundance (Table 2) were co-dominant in this phytoplankton assemblage, explaining the observed Fuco concentrations. Accordingly, it is interesting to note that in our chromatograms, the chl c pigments characteristics of diatoms (chl c 1 and chl c 2 ) and other Chromo- phyte algae are present at trace levels (Fig. 5a). We estimated that these polar compounds were strongly adsorbed in the active surface of the particulate material retained by the lters in a form not accessible to methanol. Zeaxanthin, a pigment present in rather high proportion in cyanobacteria, was found in all samples, whereas canthaxanthin was measured only in one station (St. 20) where the colonial cyanobacteria Microcystis aeruginosa was detected (Table 2). In terms of chl a calculated by CHEMTAX, green algae (5658% of total chl a) were always the dominant group. Most of the chl a arises from chlorophyceans (4355%) and the contribution of prasinophyceans was small (313%). As expected from pigment results, diatoms were the subdominant group (1442%) being relatively abundant in the deeper samples of the border of the turbidity front. In a decreasing order, cyanobacteria (919%) and cryptophytes (811%) were also relevant. The contribution of other groups such as dinoagellates (08%) and haptophytes (03%) was small (Figs. 9 and 10). Similar results were reported by Go mez et al. (2004) in their study of phytoplankton samples taken in the turbidity front during the same spring cruise. 3.4.2. Outer estuary community Fucoxanthin and peridinin (Perid) were the most abundant carotenoids in the overall area of the outer estuary (Fig. 7), being particularly high (3.11 and 4.56mg l 1 , respectively) at the surface maximum chl a concentration (15.5mg l 1 , St. 17) observed in this area (Fig. 5b). Chl c 1 , chl c 2 and Mg 3,8-divinyl pheoporphyrin (MgDVP) were the major chl c pigments whereas chl c 3 was present only in minor amounts. In particular, chl c 2 and chl c 1 attained their highest concentrations of the study area (Figs. 7 and 8). Microscopic analysis (Table 2) conrmed that several diatom species (Thalassiosira spp., Thalassionema nitzschioides, Rhizosolenia setigera and Pseudo-nitzschia spp.) were very abun- dant in this phytoplankton assemblage explaining the observed Fuco dominance. In contrast, the unequivocal marker for dino- agellates, Perid, attained their highest concentration at the surface (Fig. 8) and was related with a bloom of the dinoagellate Prorocentrum minimum (up to 397,600cells l 1 ). ARTICLE IN PRESS Table 1 Peak identication and abbreviation pigment name used in the text Peak no. Pigment Abbreviation 1 Peridininol Peridol 2 Chlorophyll c 3 (PCP) Chl c 3 , 3 Monovinyl chlorophyll c 3 MV chl c 3 4 Chlorophyllide a Chlide 5 Mg 3,8-divinyl pheoporphyrin Mg DVP 6 Chlorophyll c 2 (PCP) Chl c 2 7 Chlorophyll c 1 (PCP) Chl c 1 8 Methyl-chlorophyllide a Me chlide 9 Peridinin (PCP) Perid 10 19 0 -Butanoyloxyfucoxanthin (PCP) But-fuco 11 Fucoxanthin (PCP) Fuco 12 9 0 -cis-neoxanthin Neo 13 Prasinoxanthin (PCP) Pras 14 4-Keto-19 0 -hexanoyloxyfucoxanthin 4 k-Hexa-fuco 15 Violaxanthin (PPP) Viola 16 19 0 -Hexanoyloxyfucoxanthin (PCP) Hexa-fuco 17 Diadinoxanthin (PPP) Diadino 18 Antheraxanthin Anth 19 Dinoxanthin Dino 20 Alloxanthin (PCP) Allo 21 Diatoxanthin (PPP) Diato 22 Zeaxanthin (PPP) Zea 23 Lutein (PPP) Lut 24 Gyroxanthin diester Gyr 25 Chlorophyll b (PCP) Chl b 26 Chlorophyll c 2 monogalactosyldiacylglyceride [18:4/ 14:0] ester from E. huxleyi Chl c 2 MGDG [18:4/14:0] 27 Divinyl chlorophyll a DV chl a 28 Chlorophyll a allomer Chl a allom 29 Chlorophyll a Chl a 30 Chlorophyll a epimer Chl a epi 31 Chlorophyll c 2 monogalactosyldiacylglyceride [14:0/ 14:0] ester from C. polylepis Chl c 2 MGDG [14:0/14:0] 32 Phaeophytin a Phaeo 33 b,e-Carotene (a-carotene) a-Car 34 b,b-Carotene (b-carotene) b -Car PPP: photo-protector pigments, PCP: photo-collector pigments. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1594 Other dinoagellate species were also present but in lower concentrations (Table 2). In coincidence with the distribution of dinoagellates, the relative concentration of the unequivocal Cryptophyceae marker alloxanthin (Allo) was high in the surface samples of the outer estuarine area, but sharply dropped with depth (Figs. 7 and 8). Microscopy (Table 2) showed that free-living cryptophytes (Hemiselmis sp. and Plagioselmis sp.) were abundant in outer estuary waters, explaining the observed distribution of Allo. It is interesting to note that some cryptomonads may be found in endosymbiotic association with other organisms (Hackett et al., 2004), as for example some toxic dinoagellate species of the genus Dinophysis (D. caudata and D. acuminata) that also were present at relatively high abundance in our samples (Table 2). Although red-tide concentrations of the autotrophic ciliate Myrionecta rubra has been reported (Montoya et al., 2006 and references therein), in our samples was present only in low numbers. Another pigment pattern can be dened in association with the detected bloom (St. 14) of the silicoagellate Dictyocha bula (up to 14,700cells l 1 ; Table 2). The pigment data showed that in addition to Fuco and chl c 3 , 19 0 -butanoyloxyfucoxanthin (But- fuco) was the dominant acyl-fucoxanthin (Fig. 7) with minor contributions of 19 0 -hexanoyloxyfucoxanthin (Hexa-fuco). This ARTICLE IN PRESS Fig. 5. HPLC chromatogram of pigment extracts from surface samples (0m depth) of six selected representative stations. (a) Inner estuary (station 18). (b) Outer estuary (station 17). (c) Coastal (station 13). (d) Subantarctic Shelf-break front (station 10). (e) Subantarctic Shelf-break front (station 5). (f) Brazil Current waters (station 21). Detection was by absorbance at 436nm; peak identication as in Table 1. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1595 pigment prole is similar to that reported for the silicoagellate D. speculum (Daugbjerg and Henriksen, 2001), but also with that described for some pelagophytes (Jeffrey and Wright, 1997). Pigments associated with green microalgae (Chlorophyceae, Prasinophyceae and Euglenophyceae) were not relevant in the outer estuary phytoplankton community. In this assemblage, the amount of chl b with respect to chl a was low and lutein (Lut), the major carotenoid of chlorophyceans, was also found in low proportion (Fig. 5b). Prasinoxanthin, an unequivocal marker of some prasinophyceans (Latasa et al., 2004), was only detected in trace amounts. However, the high amount of Diadino (3.4mg l 1 ), the major carotenoid of marine euglenoids, and the Diadino/Fuco ratio (Fig. 5b) are indicative that part of the chl b arises from euglenophyceans. The microscopic examination showed a patchy distribution of a non-identied microplanktonic euglenophyte, in concentration of up to 30,271cells l 1 at the chl a maximum (St. 17). The nanoplanktonic euglenophyte Eutreptiella sp. was also detected in this estuarine area (up to 10.710 4 cells l 1 ) but the diagnostic pigment siphonein, which is present in some eugleno- phyte species, was not detected. On the other hand, Pyramimonas sp., a prasinophycean lacking Pras, was also detected as numeri- cally abundant (13.610 4 cells l 1 ). These results suggest that chl b could be shared between euglenophyceans and prasinophyceans lacking prasinoxanthin. Zeaxanthin, a pigment present in rather high proportion in cyanobacteria, was found in small amounts in all samples. However, in the Uruguayan coast associated with the river plume, Zea was the most abundant carotenoid. Minor contributions by haptophytes were also detected in stations 14, 15 and 16. The presence of a non-identied species of Prymnesiaceae (Imantonia/ Chrysochromulina) (Table 2) was in coincidence with the presence of both chl c 2 -MGDG (18:4/14:0) and chl c 2 -MGDG (14:0/14:0) in trace amounts. These haptophyte markers appear to be associated with the presence of chl c 3 , Fuco, Hexa-fuco and 4-keto-19 0 - hexanoyloxyfucoxanthin (4 k-Hexa-fuco), as in the Pigment-type 7 described by Zapata et al. (2004). As expected from pigment results, the relative contribution of the various groups to chl a calculated by CHEMTAX (Figs. 9 and 10), showed that with exception of some surface samples (St. 26), diatoms and dinoagellates were the dominant groups in the phytoplankton community of the outer estuary. The highest specic diatom-chl a concentration was attained at station 17 (7.24mg l 1 , 47% of total chl a). Also the highest absolute (4.81mg l 1 ) and relative contribution (31% of total chl a) of dinoagellates to the total chl a calculated by CHEMTAX was attained at the surface of St. 17, and was related with a bloom of the dinoagellate Prorocentrum minimum (up to 397,600cells l 1 ). However, in some surface samples cyanobacteria was the most ARTICLE IN PRESS Table 2 Main identied taxa of surface microplankton, nanoplankton and picoplankton rank abundance values in the ve geographic areas considered Shelf-break front Brazil Coastal Outer estuary Inner estuary Microplankton Bacillariophyceae Pseudo-nitzschia spp. 0960 a 1604800 0200 Rhizosolenia setigera Brightwell 240680 48012,800 203460 Thalassionema nitzschioides (Grunow) Grunow ex Hustedt 1202520 1202520 0200 Thalassiosira spp. 0200 040 8401000 4004720 0400 Total diatoms 0200 401840 216015,120 392011,200 01600 Dinophyceae Ceratium furca (Ehrenberg) Clapare` de et Lachmann 0120 3602120 Ceratium tripos (O.F. Mu ller) Nitzsch 040 40400 02160 Dinophysis acuminata Clapare` de et Lachmann 601760 4802500 Dinophysis caudata Saville-Kent 040 802340 Prorocentrum minimum (Pavillard) Schiller 0440 40160 080 1120397,600 01800 Prorocentrum scutellum Schro der 020 0180 280540 805400 Total photosynthetic dinoagellates 0980 280360 13605480 6400398,490 01800 Dictyochophyceae Dictyocha bula 576014730 Dictyocha speculum 402360 Euglenophyceae 4030,270 Nano and picoplankton Cryptophyceae Hemiselmis or Plagioselmis 00.687 021.8 Prymnesiophyceae Imantonia or Chrysocromulina 027.7 01.37 013.8 Gephyrocapsa or Emiliania 013.0 00.687 Other Coccolithophore species 09.09 076.3 Prasinophyceae Pyramimonas spp. 013.6 Chlorophyta Coccal cells 05910 Euglenophyceae Eutreptiella spp. 010.7 Bacillariophyta Centric cells (o5mm) 066.5 Chrysophyta Ollicola spp. 01.38 Microplankton abundance in cells per liter; nano- and picoplankton abundance in cells per liter 10 4 . a 219,260cells l 1 at 20m depth. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1596 important group (St. 26: 39.0% of total chl a). In decreasing order, green algae (euglenophyceans+prasinophyceans lacking Pras, 30.71%), cryptophytes (19.23.4%) and silicoagellates (14.23.8) were also relevant. The contribution of other groups such as haptophytes (100%) and prasinophyceans containing Pras (10.20%) was small. 3.4.3. Coastal community In contrast to the estuarine waters, the pigment markers for green algae (chl b, Lut), dinoagellates (Perid) and cryptophytes (Allo) were in relatively low concentrations in the coastal ecosystem (Fig. 7). Fuco was largely the most abundant carotenoid in the whole area, being particularly high at the chl a maximum observed at station 12 (3.81mg l 1 ) near the base of the euphotic zone (Fig. 8). In a decreasing order, chl c 2 , chl c 3 and chl c 1 were the major chl c pigments of this assemblage. In the surface, chl c 3 was detected in relatively low amounts but a sharp increase in the chl c 3 /chl a ratio was observed at the chl a subsurface maximum, where the highest chl c 3 concentration (St. 12, 0.35mg l 1 ) was found. However, Hexa-fuco was present at very low concentra- tions (Fig. 8), and some other haptophyte markers such as chl c 2 -MGDG (18:4/14:0) and chl c 2 -MGDG (14:0/14:0) were only detected at trace levels. Moreover, in this station the vertical distribution of phytoplankton showed that diatoms prevailed at all depths of the water column whereas haptophytes were absent. However, associated with the subsurface chl c 3 maximum, a sharp change in the diatom species composition was observed. In coincidence with the chl c 3 maximum, a mono-specic bloom of the diatom Pseudo-nitzschia sp. was found (Fig. 11), explaining the observed pigment pattern (Zapata et al., 2000. These results allowed us to distinguish between two pigment-types diatom populations (Diatom I and Diatom II) within the coastal phytoplankton community. The algal group composition obtained by CHEMTAX for the coastal area showed an almost total dominance (59.393.5% of total chl a) of diatoms. With the exception of some samples (St. 12: 20 and 30m, and St. 8: 30m) the usual diatom-pigment pattern group (chl c 1 , chl c 2 and Fuco, Diatom I), was the more abundant and widely distributed in the coastal area. However, the CHEMTAX calculation showed total dominance (87.3%) of the pigment pattern group Diatom II (chl c 2 and chl c 3 ) within the Pseudo-nitzschia spp. bloom. In a decreasing order, dinoagellates (160%), haptophytes (121.6%), cryptophytes (160%) and cya- nobacteria (110.5%) were also relevant. The contribution of green algae (prasinophyceans and chlorophyceans) was small. 3.4.4. Shelf-break front community Pigments associated with green algae were relevant in this phytoplankton assemblage, particularly at the chl a maximum detected in the central stations (Sts. 6 and 10) of this frontal area. Chromatograms showed (Fig. 5d and e) high absolute and relative concentrations of chl b (chl b:chl a ratio up to 0.65) and lutein (Lut:chl a ratio up to 0.25), associated with minor amounts of neoxanthin (Neo), violaxanthin (Viola) and Zea. Prasinoxanthin, an unequivocal marker of some prasinophyceans was also detected in trace or minor amounts, being relevant at the deep euphotic zone. The Lut/chl b ratio has found to be higher in chlorophytes (0.301.77) than in prasinophytes lacking Pras (00.18) (Schlu ter et al., 2000; Henriksen et al., 2002; Latasa et al., 2004). The observed Lut/chl b ratios averaged 0.3770.2 indicating that mainly chlorophyceans contributed to the mea- sured chl b. In coincidence with pigment analysis, electronic and light microscopic examination of these samples revealed a bloom (up to 59.110 6 cells l 1 ) of a picoplanktonic (o3mm) coccal cells of a not identied chlorophycean. Therefore, most of the chl b appears to be related to picoplanktonic chlorophyceans. The high abundance of chl b-containing organisms in the shelf-break area during this study is in contrast with a previous investigation ARTICLE IN PRESS Fig. 6. Changes in photo-collector pigments/chlorophyll a (empty symbols) and photo-protector pigments/chlorophyll a ratios (full symbols) with PAR irradiance as percentage of the incident radiation, in some selected representative stations. (a) Coastal. (b) Brazil Current waters. (c) Subantarctic Shelf-break front. Drawn lines show general tendencies in photo-adaptive processes. No tting was done. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1597 ARTICLE IN PRESS Fig. 7. Surface distribution of principal accessory biomarker pigments concentration (mgl 1 ) grouped in separate panels: (a) fucoxanthin, peridinin, chlorophyll b and 19 0 -hexanoyloxyfucoxanthin and (b) chlorophyll c 1 and c 3 , prasinoxanthin, alloxanthin, lutein, 19 0 -butanoyloxyfucoxanthin, divinyl chlorophyll a and zeaxanthin. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1598 (Carreto et al., 2003) reporting insignicant concentrations of chl b during late spring 1999 in this area. At the maximum chl a detected in the central stations of the shelf-break front, the concentration of chl c 3 , Fuco, Hexa-fuco and But-fuco were high (Fig. 5d and e) indicating that haptophytes and/or pelagophyceans could be also abundant. However, some species of Parmales with the same pigment signature, were observed in a sample from the Ross Sea (Wright and van den Enden, 2000), indicating that sources other than haptophytes and pelagophytes could be related with the presence of the mentioned pigment pattern. In our samples, the additional useful haptophyte marker pigments, chl c 2 -MGDG (18:0/14:0), chl c 2 -MGDG (14:0/ 14:0), monovinyl chl c 3 and 4 k-Hexa-fuco (Zapata et al., 2004), were also detected in minor or trace amounts. Using these pigment markers, at least two haptophyte pigment types were distinguished in the shelf-break area. Chl c 2 -MGDG (18:0/14:0) was the dominant non-polar chl c 2 at the sub-surface chl a maximum detected in the central St. 10, whereas at the borders of the front (Sts. 5, 7 and 9), in correspondence with the lowest chl a values, both non-polar chl c 2 [chl c 2 -MGDG (18:0/14:0) and chl c 2 -MGDG (14:0/14:0)] were found in similar amounts. The presence of a non-identied prymnesiophycean species (possibly Chrysochromulina) in concentrations of up to 27.710 4 cells l 1 (Table 2), can explain the presence of both non-polar chl c 2 (Pigment type 7 of Zapata et al., 2004) and the relative amounts of Hexa-fuco detected at the borders of the front. In contrast, in ARTICLE IN PRESS Fig. 8. Vertical distribution of principal accessory biomarker pigments concentration (mgl 1 ) along the central section. (a) Fucoxanthin, peridinin, chlorophyll b and 19 0 - hexanoyloxyfucoxanthin grouped in separate panels. (b) Chlorophyll c 1 and c 3 , prasinoxanthin, alloxanthin, lutein, 19 0 -butanoyloxyfucoxanthin, divinyl chlorophyll a and zeaxanthin. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1599 some stations (Sts. 6 and 7) at the shelf-break front the haptophyte populations were mainly comprised by coccolitho- phorids (E. huxleyi and/or Gephyrocapsa oceanica), a family characterized by a single pigment type (Type 6) containing chl c 2 -MGDG (18:0/14:0) as unique non-polar chl c 2 (Zapata et al., 2004). Similarly to that observed previously in this area (Carreto et al., 2003), the Hexa-fuco biomass estimated by microscopic coccolithophorid cell counts (assuming a cell content of 0.1mg Hexa-fuco per 10 6 cells, Stolte et al., 2000; Galmozzi et al., 2001) was three orders of magnitude lower than that determined by HPLC. Moreover, no other haptophytes were microscopically observed in samples containing the highest acyl-fucoxanthins concentration (St. 10, 1.31mg l 1 ). The dominance of naked or scaly coccolithophorid cells (Paasche, 2001) and the presence of other ARTICLE IN PRESS Fig. 9. Surface distribution of percentage of different phytoplankton groups contributing to the total chlorophyll a concentration, estimated by interpretation of pigment HPLC data using CHEMTAX program, grouped in separate panels. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1600 haptophytes too fragile to be adequately preserved and therefore undetected by microscopy, could explain such inconsistency between E. huxleyi/G. oceanica cell biomass and Hexa-fuco concentrations. For instance, recent molecular studies in photic oceanic regions have revealed a large diversity of photosynthetic picoeukaryotic heterokonts, including haptophyceans (Moon-van der Staay et al., 2000). Our chromatograms also showed (Fig. 5d) a signicant But-fuco/Hexa-fuco ratio, indicating that Pigment type 8 of haptophytes was probably present (Zapata et al., 2004). However, the presence of pelagophyceans and/or other But-fuco- containing picoeukaryotes, such as the small and too fragile agellate Florenciella parvula (Dictyochophyceae; Eikrem et al., 2004), could not be discarded. Microscopy also showed that in one station of this area, some non-identied, very small diatom cells (5mm) were present in high concentrations (up to 66.510 4 cells l 1 at St. 10). These results indicated that in addition to haptophytes, the Fuco concentration was also related with the abundance of these small diatoms. It is interesting to note that signicant amounts of chlorophyllide a and lesser amounts of their methyl-ester, were found in our chromatograms (Fig. 5d) indicating that the diatom population probably was in a senescent stage. In addition to the major pigment markers, Pras, Allo, Zea and Perid were also detected in minor or trace amounts. As expected from pigment results, the relative contribution of the various groups to the total chl a calculated by CHEMTAX, showed that in contrast with the almost exclusive diatom dominance observed in the estuarial and coastal systems, the shelf-break community was more complex, being principally comprised by chlorophyceans (up to 50% of total chl a in St. 6) and haptophytes (up to 62% in St. 5). Three categories of haptophytes (of pigment types 6, 7 and 8; Table 3) were considered in our calculations in view of the high complexity in the pigment patterns and phytoplankton composition observed in this region. Results showed that haptophyte pigment type 7 was abundant in all the studied stations, being dominant at the surface samples (75100% of total haptophytes). However, at the chl a sub-surface maximum a noticeable increase in the relative contribution of haptophyte pigment types 6 and 8 was found in some stations (Fig. 12a and b). The CHEMTAX calculation showed that diatoms were relatively abundant in the deeper samples of the borders of the front (St. 5: 47.7% and St. 7: 44.5%). Cryptophytes, prasinophyceans and pelagophyceans were also present in surface samples of this area but in relatively low proportions. Furthermore, the CHEMTAX calculation showed a marked increase in the relative contribution of cryptophytes (up to 20.4%) and prasinophyceans (up to 36.6%) at the base of the euphotic layer. The cyanobacteria (Synechococ- cus) was only detected in minor amounts with the exception of station 7 (21.5%). 3.4.5. Brazil Current community Zeaxanthin was the principal carotenoid in the phytoplankton community of the oligotrophic Brazil Current waters (Fig. 5f), ARTICLE IN PRESS Fig. 10. Vertical distribution of percentage of different phytoplankton groups contributing to the total chlorophyll a concentration along the central section, estimated by interpretation of pigment HPLC data using CHEMTAX program grouped in separate panels. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1601 indicating that cyanobacteria were the most abundant cells in the phytoplankton assemblage. Zeaxanthin is found in both Synecho- coccus and Prochlorococcus, whereas divinyl chl a is exclusive to Prochlorococcus. Although in our samples Synechococcus appears to be dominant, the relative proportion of Prochlorococcus was important, particularly at the surface of the more oceanic station (St. 21) where a high contribution of DV chl a (37%) to the total chl a (chl a +DV chl a) was found (Fig. 5f). Divinyl chl b (DV chl b) could not be separated from chl b with the method employed (Zapata et al., 2000), and our data refer to the sum of DV chl b and chl b (chl b 1+2 ). The co-elution of these pigments could be a drawback if green algae without specic marker was also present in the assemblage and its contribution to the total chl a should also be evaluated. In this phytoplankton assemblage, Lut was not detected and consequently the chl b of green algae arises only from prasinophyceans containing prasinoxanthin. On the other hand, DV chl b is a less specic pigment marker for Prochlor- ococcus since some clones contain both DV chl b and chl b in a similar proportion (Moore et al., 1995; Moore and Chisholm, 1999). Although prochlorophytes were dominant at depth in the tropical and subtropical regions of the Atlantic Ocean (Barlow et al., 2002), differences among ecotypes could be important in determine the relative distribution of these cyanobacteria in the water column and throughout the oceans (Moore et al., 1995). The recent divergence of the Prochlorococcus lineage into a distinct clade that is better adapted for growth at higher irradiance levels could explain the ability of this genus to dominate at several different depths in the water column (Moore and Chisholm, 1999; Ting et al., 2002). Our results showed that in the Brazil Current waters during spring, the concentration of DV chl a was high at the surface (Fig. 7), and uniformly distributed throughout the ARTICLE IN PRESS Fig. 11. Vertical distribution of chlorophyll c 3 concentration (mg l 1 ), the concen- tration of the diatom Pseudo-nitzschia sp. (cells ml 1 ) and of the others diatoms counts (cells ml 1 ) in the Station 11 of the coastal area. Table 3 Pigment: chlorophyll a ratio in the haptophyte pigment-types considered for calculations after tting by CHEMTAX program Haptophyte pigment type Fuco Hexa-fuco But-fuco Chl c 3 Chl c 2 Chl c 2 MGDG [18:4/14:0] Chl c 2 MGDG [14:0/14:0] Type 6 0.10 1.06 0.021 0.24 0.23 0.087 Type 7 0.36 0.72 0.062 0.09 0.11 0.034 0.023 Type 8 0.48 0.45 0.22 0.31 0.10 0.049 Fig. 12. Vertical distribution of chlorophyll a (mg l 1 ) in the different haptophyte pigment types (6, 7 and 8), estimated by interpretation of pigment HPLC data using CHEMTAX program, in some selected stations of the shelf-break area: (a) Station 5 and (b) Station 1. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1602 euphotic zone (Fig. 8), suggesting the existence of at less one high light adapted ecotype that is capable to growth at great irradiance levels and that posses a lowchl b 1+2 : DV chl a ratio (Fig. 13). At the surface of station 21 where no traces of other green pigment markers was found, the chl b 1+2 :DV chl a ratio was 0.22 (Fig. 5f). However, the range of chl b 1+2 :DV chl a ratios measured in the deep euphotic zone (70150m) of station 22 (Fig. 13) was similar (2.012.99) to that of members of the more deeply branching clades (Ting et al., 2002). As suggested by Goericke and Repeta (1993), the sharp transition of chl b 1+2 :chl a ratio observed at depth may be due in part to photoacclimation processes, but mainly to changes in differently photoadapted Prochlorococcus populations (Moore et al., 1995). However, in this assemblage, Pras concentrations were also important, indicating that chl b 1+2 should be shared between prasinophytes and Prochlorococcus. Neverthless, picoplanktonic prasinophytes and Prochlorococcus could not be detected in the microscope. The Pras-containing prasinophytes are represented by three phylogenetic groups: Mamiellales, Pseudoscoureldiales and Prasinococcales (Guillou et al., 2004). These groups can be classied based in the presence/absence of uriolide, micromonal and other pigments (Latasa et al., 2004). We could not detect these pigments in our samples and therefore some members of Pseudoscoureldiales were probably present (Latasa et al., 2004). The relative proportions of chl c 3 , Fuco, Hexa-fuco and But-fuco were also relatively high (Fig. 5f) indicating that haptophytes and/ or pelagophytes were abundant. Chl c 2 -MGDG (18:4/14:0) was the only non-polar chl c 2 detected at minor or trace levels. Although interpretation is complicated by the fact that some additional markers for distinguishing between the different haptophyte types (Zapata et al., 2004) were below the limits of detection, the observed proles can be associated with the Pigment-type 6. Microscopy showed that although E. huxleyi and/or G. oceanica were present in this assemblage, some unidentied coccolito- phorid species were the dominant haptophyte (up to 760cells ml 1 ; Table 2). The ratio of But-fuco to total fucoxanthins (Fig. 5f) was signicantly higher than that observed for any studied haptophyte species (Zapata et al., 2004), indicating that pelagophyceans were probably present. Minor amounts of Perid, the unequivocal marker for dino- agellates, were also detected in the Brazil Current assemblage (Fig. 5f). Microscopy (Table 2) conrmed that several dinoagellate species of the genera Prorocentrum, Ceratium and Torodinium including thermophilous species such as Ceratium lanceolatum and Oxytoxum scolopaxwere present in low abundance in this phytoplankton assemblage, in coincidence with the observed distribution of Perid. Microscopy also showed that several diatom species were found in some samples at very low numerical abundance (up to 1.5cells ml 1 ) indicating their minimal con- tribution to the observed Fuco concentration. The algal group composition obtained by CHEMTAX for the Brazil Current assemblage showed the dominance of the pico- planktonic cyanobacteria Synechococcus (23.654.5% of total chl a) and Prochlorococcus (10.438% of total chl a). Haptophytes were also present as sub-dominant group, being particularly abundant (up to 41.8%) in the deep chl a maximum situated near the base of the euphotic layer (Fig. 14). Prasinophyceans and pelagophyceans were present in surface samples but in relatively low proportions (up to 6.2% and 3.7% respectively). However, at the base of the euphotic layer the CHEMTAX calculation showed a marked increase in the relative contribution of prasinophyceans (up to 16.0%) and pelagophyceans (up to 13.3%). ARTICLE IN PRESS Fig. 13. Vertical distribution of chlorophyll a, divinyl chlorophyll a and total chlorophyll b (chlorophyll b 1 +divinyl chlorophyll b 2 ) concentrations (mg l 1 ) in station 22 at the Brazil Current area. Fig. 14. Vertical distribution of chlorophyll a contribution (mg l 1 ) of the major phytoplankton groups, estimated by interpretation of pigment HPLC data using CHEMTAX program, in station 22 at the Brazil Current area. J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1603 3.5. Phytoplankton communities and environmental conditions Our results are consistent with the view that the size structure and taxonomic composition of phytoplankton communities are principally regulated by the linkage between nutrient availability and turbulent mixing (Margalef, 1978). However, this linkage is expressed differently in the different studied ecosystems (Cullen et al., 2002). Although large diatoms occupy the high-turbulence/ high-nutrient ecosystems (Margalef, 1978), our study showed that within the tidal-mixed and nutrient-rich turbidity front, phyto- plankton growth is light-limited and freshwater chlorophytes, diatoms and cyanophytes were the dominant groups . These results are consistent with previous records for this area (Go mez et al., 2004). A high-nutrient/low-turbulent regimen, as can be expected along the river plume (Acha et al., 2004; Huret et al., 2005; Carreto et al., 2007), was associated with specially adapted dinoagellate species that can form red tides. However, phyto- plankton community in the outer estuary, as in most of the previous reports for this area (Negri et al., 1988; Carreto et al., 2003; Carreto et al., 2007; Go mez et al., 2004), was dominated by large diatoms. Nevertheless, bloom-forming dinoagellate species were also abundant, co-dominating in some cases with the diatom ora. Small-scale temporal variability in the turbulence/nutrient ratio, produced by changes in the wind-induced vertical mixing, can modulate the relative abundance of both life forms in the phytoplankton assemblage of this ecosystem (Acha et al.,2008). According to Guerrero et al. (1997), a total disruption of water column stratication and mixing of the salt wedge occurs after several hours of strong onshore winds (411ms 1 ). During the more turbulent period, mucous secretion, chain formation and high swimming speeds may be environmental engineering strategies that dinoagellates and other agellatesas those present in this ecosystemcan use to offset physical displace- ment through mixing, advection and turbulence (Smayda, 2002). Along the shelf region, seasonal changes in the turbulence/ nutrient relationship drive the typical phytoplankton succession characteristics of temperate ecosystems (Carreto et al., 1995, 2004; Lutz et al., 2006). After the spring bloom, the slightly coastal stratied low-nutrient waters (Carreto et al., 2007) showed low chl a concentration and the dominance of diatoms Type I (Thalassiosira sp.). Their abundance in the subsurface and deep samples of the more coastal stations was probably related to their transport in the buoyant plume from the estuary and, then, sedimentation. However, in the offshore stations, a more ad- vanced succession stage forms of diatoms (Type II) specially adapted to minimize sedimentation (Pseudo-nitzschia sp. and Rhizosolenia setigera) were the dominant ora, suggesting their local origin. Typical phytoplankton succession can also provide a basis for understand large-scale horizontal distribution of phytoplankton communities. It is well known that growth of phytoplankton in the quasi permanent low-turbulence/low nutrient ecosystems is largely supported by regeneration of nutrients and that nano- and picoplankton are the main components in these communities (Cullen et al., 2002). Along the offshore trajectory from the eutrophic estuarine waters (Carreto et al., 2007) to oligotrophic subtropical waters (Brandini et al, 2000) our results showed a notably gradient in abundance, size structure and taxonomic composition of phytoplankton communities from high micro- plankton biomass to low picoplankton biomass. However, in spite that specic growth rates of larger phytoplankton are relatively low, some forms (Ceratium spp.) can persist through special strategies, such as vertical migration (Margalef, 1978; Smayda, 2002). At the central and southern sections the deepening of the stratied layer was interrupted at the shelf-break-front by a moderate upwelling of subantarctic waters that forms a two-layer water column with a well-developed, relatively shallow thermo- cline. Fast-growing more r-selected small agellates (haptophytes and chlorophytes) were abundant in this environment, explaining the characteristic high surface chlorophyll concentration band observed from satellites along the edge of the shelf, during late spring (Podesta , 1997; Carreto et al., 2003; Huret et al., 2005; Romero et al., 2006). The occurrence of patches of water with spectral signatures identical to those of blooms of the coccolitho- phore E. huxleyi have been regularly observed on the shelf-break off Argentina during late spring and summer (Brown and Podesta , 1997). In this area the most remarkable hydrographic feature associated with previously reported E. huxleyi bloom was the existence of a well-developed shallow pycnocline separating surface water from deep nutrient-rich Subantarctic waters (Gayoso and Podesta , 1996; Carreto et al., 2003). Recently, the ubiquitous coccolithophorid species G. oceanica was also observed in high abundance (up to 310 5 cells l 1 ) at the edge of the shelf- break (Negri et al., 2003). On the other hand, this is the rst time that high chl b concentrations associated to a bloom of a picoplanktonic (o3mm) coccal chlorophycean cells was reported for this region. Notably, picoplankton cells (o2mm) were dominant in the Antarctic Circumpolar Current and Chlorella like organisms dominated the autotrophic pico- and nanoplank- ton (Peeken, 1997; Detmer and Bathmann, 1997). Preliminary studies showed that the high surface chlorophyll concentration band observed along the shelf-break front during October 2005, was produced by a typical temperate diatom spring bloom. The predominance of small agellates in the shelf-break area during this study suggests that the observed scenario corresponded to a more advanced phytoplankton succession stage (Margalef, 1978). These results are in coincident with recent studies using satellite-derived data (Signorini et al., 2006). In the warm, highly stratied nutrient-poor oceanic waters of the Brazil Current, picoplanktonic cyanobacteria (Synechococcus and Prochlorococcus) were the dominant groups, being more abundant in the deep chl a maximum situated near the base of the euphotic layer. As was observed in other subtropical waters (Bouman et al., 2006), in this ecosystem, the strong stratication isolate deep illuminated waters from the surface, producing selection of genetic and physiological traits of marine microbial communities. In particular, the high light plasticity of the genus Prochlorococcus has been explained by the discovery of genetically and physiologically distinct populations, commonly referred to as high light (HL)- and low light (LL)-adapted ecotypes (Goericke and Repeta, 1993; Moore et al., 1995; Bouman et al., 2006). Our results indicate the existence of at least one HL-adapted ecotype that is capable to growth at great irradiance levels, posses a low chl b 1+2 :DV chl a ratio and contains relatively high amounts of UV photo-protective substances, mycosporine-like amino acids (Carreto et al., 2005). Recently, it has shown that the HL strain has a photolyase gene, which serves to repair ultraviolet damage, and which is absent in the LL-adapted strain (Rocap et al., 2003). The observed variations in community structure along the studied transects highlighted the adaptability of the phytoplank- ton to changing environmental conditions. In turn, the high variability in the size structure and pigment composition of the communities strongly inuences the pattern of absorption in phytoplankton cells (Yentsch and Phinney, 1989; Barlow et al., 2002; Bricaud, 2004; Sathyendranath et al., 2004). A more frequent sampling strategy in space and time combined with analysis of satellite images should be undertaken to improve our understanding of phytoplankton dynamics in this region, and therefore to improve the synoptic census of phytoplankton by ocean-colour remote sensing. ARTICLE IN PRESS J.I. Carreto et al. / Continental Shelf Research 28 (2008) 15891606 1604 Acknowledgments We are grateful to Dr. M.D. Mackey for providing the computer program CHEMTAX. We wish also to thank Dr. Domingo Gagliardini for NOAA images processing and to SeaWiFS Project NASA Goddard Space Flight Centre and ORBIMAGE for chl a images. This study was partially supported by Antorchas Founda- tion (Grant 13900-13) and CONICET PIP 5009. This is contribution no. 1443 of INIDEP. References Acha, E.M., Mianzan, H., Guerrero, R., Carreto, J.I., Giberto, D., Montoya, N., Carignan, M.O., 2008. An overview of ecological processes in the Ro de la Plata estuary Cotinental Shelf Research, in press, doi:10.1016/j.csr.200701.031. Acha, E.M., Mianzan, H., Guerrero, R., Favero, M., Bava, J., 2004. Marine fronts at the continental shelves of austral South America. Physical and ecological Processes. Journal of Marine Systems 44, 83105. 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