Algal Pigment Patterns and Phytoplankton Assemblages in Different Water

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
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Continental Shelf Research
0278-4343/$ - see front matter & 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.csr.2007.02.012
Corresponding author. Tel.: +542234862586; fax: +542234861830.

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).
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.
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.
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
).
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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.
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.
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
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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.
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.
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.
(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.
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.
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),
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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.
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
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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.
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.
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
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.
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Algal pigment patterns and phytoplankton assemblages in different water

masses of the Ro de la Plata maritime front

, Nora Montoya, Rut Akselman, Mario O. Carignan,

Corresponding author. Tel.: +542234862586; fax: +542234861830.

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