Journal
Journalof
ofCoastal
CoastalResearch
Research
SI 64
pg -- pg
354
358
ICS2011
ICS2011 (Proceedings)
Poland
ISSN 0749-0208
Effects of a high energy coastal environment on the structure and
dynamics of phytoplankton communities (Brazilian Amazon littoral)
V. B. da Costa†, E. B. de Sousa‡, S. C. C. Pinheiro†, L. C. C. Pereira‡ and R. M. da Costa‡*
†Instituto Evandro Chagas/SVS/MS,
67030-070, Ananindeua, Pará, Brazil
e-mail: [email protected]
‡Instituto de Estudos Costeiros,
Universidade Federal do Pará,
Bragança 68600-000, Brazil.
*email: [email protected]
ABSTRACT
Costa, V.B. da, Sousa, E.B de, Pinheiro, S.C.C., Pereira, L.C.C. and Costa, R.M. da, 2011. Effects of a high
energy coastal environment on the structure and dynamics of phytoplankton communities (Brazilian Amazon
littoral. Journal of Coastal Research, SI 64 (Proceedings of the 11th International Coastal Symposium), .
Szczecin, Poland, ISSN 0749-0208
The present study investigated the spatial-temporal dynamics of phytoplankton communities at three stations at
Ajuruteua Beach in Bragança (Pará, Brazil) between August, 2004, and July, 2005. Water temperature, salinity,
pH and dissolved oxygen concentrations were measured in situ using a multiparameter probe, while water
transparency was estimated using a Secchi disk. Samples for quantitative analyses were obtained from the
subsurface water and fixed with 4% formaldehyde. Phytoplankton density was estimated using Utermöhl’s
sedimentation method. Chlorophyll a concentrations, the frequency and relative abundance of different taxa, and
diversity and evenness were also determined. The data were analyzed using a one-way ANOVA, followed by the
Fisher’s test. Water transparency ranged from 10 to 38 cm, while water temperature varied between 26.9 and
31.3ºC, and salinity from 15.3 to 36.2 psu. Dissolved oxygen ranged from 4.8 to 8.4 mg/l, while pH values
ranged from 7.20 to 8.26. A total of 82 taxa were recorded, most of which (93.7%) belonged to the
Bacillariophyta. Total phytoplankton density ranged from 719.13 x 103 cell/l to 2,675.81 x 103 cell/l with higher
values being recorded in the rainy season. Phytoflagellates comprised approximately 84% of the phytoplankton
collected, followed by diatoms. Phytoplankton biomass varied from 1.16 mg/m³ to 17.63 mg/m³ with higher
values being recorded in the rainy season. The results indicated that precipitation is the main factor influencing
the composition, density, biomass and diversity of the phytoplankton community at the study site, although local
hydrodynamics favored the resuspension of sediments and tychoplankton species such as Dimmeregrama minor.
ADITIONAL INDEX WORDS: Plankton, Spatiotemporal variability, Amazon Beaches.
INTRODUCTION
The coast zone of the Amazonian states of Amapá, Pará, and
Maranhão encompasses more than a third of the 7,400 km-long
coastline of Brazil (Isaac and Barthem, 1995). This region is
characterized by complex hydrodynamic processes resulting from
the action of winds and currents (Nittrouer and DeMaster, 1996)
and the discharge of freshwater, solutes and suspended particulate
material from the Amazon and Pará rivers (Smith and Demaster,
1996) and its many minor estuaries.
As in most aquatic ecosystems, the phytoplankton community
forms the base of the food web (Chiu et al., 1994; Lalli and
Parsons, 1993) and is responsible for around 90% of the annual
organic production and fixing most of the available inorganic
carbon (Raymont, 1980; Lalli and Parsons, 1993). These
communities are highly dynamic, responding rapidly to
fluctuations in the physical and chemical properties of the water
through high rates of reproduction and mortality (Valiela, 1995).
The composition, productivity, and biomass of a community are
thus related directly to the climatic and hydrological
characteristics of the environment it inhabits, as well as regional
and seasonal variations in these qualities (Brandini et al., 1997).
Despite the potential economic importance of the phytoplankton
community, and its unique hydrodynamic characteristics, few data
are available for the vast northern coast of Brazil, and most studies
have focused on the oceanic areas influenced by the discharge of
the Amazon River and other estuary systems (Smith and DeMaster
1996; Paiva et al., 2006; Sousa et al., 2008, 2009). The present
study aimed to evaluate the spatial-temporal variation in the
structure (composition, frequency, abundance, biomass, density,
diversity and evenness) of the phytoplankton community of a
macrotidal beach environment, and in particular the effects of
climatic and hydrological variables on the dynamics of its
populations over an annual cycle.
STUDY AREA
The study area is located in the Bragança coastal plain, in the
Brazilian state of Pará, which extends from Maiaú Point to the
mouth of the Caeté River (00º46’00”-1º00’00”S, 46º36’00”46º44’00”W), with an area of approximately 1.570 km² (SouzaFilho and El-Robrini, 1996).
The climate is equatorial humid with two main seasons, a dry
season (August to December) and a rainy season, from January to
July (Moraes et al., 2005). Mean annual rainfall ranges from 2,500
to 3,000 mm, the humidity of the air oscillates between 80 and
91%, and mean annual air temperature is about 25.7°C, ranging
from 20.4°C to 32.8°C (Martorano et al., 1993).
Like the rest of the northern coast of Brazil, this coastal plain is
dominated by semi-diurnal macrotides (Souza-Filho and El-
Journal of Coastal Research, Special Issue 64, 2011
354
Structure and dynamics of equatorial phytoplankton communities
Robrini, 1996), with heights ranging from 4 to 6 m. Ajuruteua
Beach (Figure 1) is located in northeastern Pará, 36 km from the
town of Bragança, and is classified as a barrier-ridge beach, which
forms a NW-SE arc about 2.5 km long and 300-400m wide
(Souza-Filho et al., 2003). It is a macrotidal dissipative beach
composed mainly of well-selected, fine quartzose sands (Alves
and El-Robrini, 2006).
The statistical analysis of data included a one-way analysis of
variance (ANOVA) followed by Fisher’s post-hoc test with a 5%
significance level. All analyses were run in the STATISTICA 6.0
package. Additional multivariate cluster analyses (Weighted Pair
Group Method with Averaging, WPGMA) were run in PRIMER
(Plymouth Routines in Multivariate Ecological Research), version
6.0.
RESULTS
Climatic and hydrological variables
Figure 1. Location of the study area, showing the survey stations
(black circles: 1, 2 and 3).
Total monthly precipitation varied from 0.20 mm in November,
2004, to 501.60 mm in March, 2005, with a total of 1,871.4 mm
over the 12-month period between August, 2004, and July, 2005.
Mean monthly air temperature ranged from 26.0ºC in August,
2004, and September, 2004, to 27.2ºC in December, 2004. Mean
monthly relative humidity varied from 73% in October, 2004, to
90% (March, April, and May, 2005), accompanying precipitation
levels.
No significant spatial or tidal patterns were recorded in the
variables analyzed during the study period. Water transparency
ranged from 10 cm in March, 2005, to 38 cm in February, with the
highest values being recorded during the dry season (p < 0.001).
Water temperature ranged between 26.9ºC (March, 2005) and
31.3ºC (February, 2005), with an amplitude of 4.4ºC. Salinity
increased significantly (p < 0.001) between the rainy season
(minimum of 15.3 psu in March, 2005) and the dry (maximum of
36.2 psu in December, 2004), with an amplitude of 20.9 psu.
Dissolved oxygen concentrations ranged from 4.8 mg/l in March,
2005, corresponding to a saturation of 106.24%, to 8.4 mg/l in
June, with a saturation of 185.90%. Significantly higher pH values
(7.29-8.26) were recorded during the dry season (p < 0.001).
Phytoplankton
METHODS
Climate data (rainfall, air temperature and relative humidity)
were obtained from the meteorological station of the National
Meteorological Institute (INMET) located at Tracuateua (01º04’S,
46º54’W), around 50 km from Bragança.
Water samples were collected for the analysis of biotic and
abiotic variable each month between August, 2004, and July,
2005, during spring tide periods (ebb and flood tides) at three
stations distributed along Ajuruteua beach (Figure 1). Water
temperature, salinity, pH, and disssolved oxygen levels were
measured in situ trough the use of a multiparameter probe (WTW
- model 340i), while water transparency was determined using a
Secchi disk.
Samples for the quantitative study of the phytoplankton were
collected using 250 ml plastic flasks and preserved in a 4%
buffered formalin-seawater solution. Additional subsurface water
samples were collected for the measurement of chlorophyll a
concentrations. The Utermöhl (1958) sedimentation method was
used for the determination of phytoplankton density.
Phytoflagellates were identified to the group level and counted.
The criteria for the identification and classification of the
microphytoplankton were obtained from the relevant scientific
literature. Once the organisms were identified and quantified,
frequency and relative abundance were estimated, and the
Shannon-Wiener diversity index and the evenness index were
calculated. Chlorophyll a levels were also determined using a
spectrophotometrical analysis (Parsons and Strickland, 1963).
A total of 82 phytoplanktonic taxa were identified, most of
which (93.68%) belonged to the Bacillariophyta, with the
remainder from the Dinophyta (6.10%) and Cyanophyta (1.22%).
The species Campylosira cymbelliformis, Coscinodiscus sp.
Dimeregramma minor, Ditylum brightwellii, Navicula sp.,
Odontella aurita, Pleurosigma sp., Skeletonema spp.,
Thalassionema nitzschioides, Thalassiosira subtilis, and
Thalassiosira sp. were recorded in all samples, and were also the
most abundant species overall. The only species considered both
dominant and abundant was D. minor, a marine tychoplankton
(Figure 2). The community was dominated by marine species
(79.31%), and to a lesser extent by marine littoral (tychoplankton)
species (20.69%).
Total phytoplankton density ranged from 719.1 x 10³ cell/l in
February, 2005, to 2,675.8 x 10³ cell/l in March, with significantly
higher values being recorded during the rainy season (p < 0.001).
Phytoflagellates comprised about 84% of the total phytoplankton,
with densities varying from 539.0 x 10³ cell/l (February) to
1,715.8 x 10³ cell/l in March. Diatoms were the group with the
second highest density, in particular D. minor, Skeletonema spp.
and A. glacialis, which returned densities of 295.0 x 103 cell.l-1;
33.7 x 103 cell.l-1 and 14,2 x 103 cell.l-1. Dimeregramma minor
occurred at the highest densities in the dry season (629.1 x 103
cell.l-1, September, 2004), and the lowest ones in the rainy season
(87.4 x 103 cell.l-1), when Skeletonema spp. (106.1 x 103 cell.l-1)
and A. glacialis (96.5 x 103 cell.l-1) occurred at higher densities.
Phytoplankton biomass varied from 1.16 mg/m³ in January,
2005, to 17.63 mg/m³ in March, with significantly higher
chlorophyll a concentrations (p < 0.001) recorded during the rainy
Journal of Coastal Research, Special Issue 64, 2011
355
Costa et al.
season. Species diversity ranged from 0.64 (December, 2004) to
3.13 bits/cell in August, 2004, while evenness varied between 0.17
in November and December, 2004, and 0.74 in August, 2005.
Both parameters returned significantly higher values in the rainy
season (p < 0.05).
Figure 2. Relative abundance (%) of main microphytoplankton
species in Ajuruteua beach.
The cluster analysis revealed the formation of three groups,
with 70% similarity (Figure 3). The first group was made up of
samples from the rainy season (March to July), and the second by
dry-season samples (August to November). While the third group
composed by samples from both seasons, it covered a continuous
period, from December to February.
Figure 3. Samples association in Ajuruteua beach during the study
period.
DISCUSSION
Climatic conditions have a considerable effect on hydrological
variables, and thus on the biological characteristics of both pelagic
and benthic organisms (Aidar et al., 1993). In tropical and
subtropical regions, rainfall appears to be the main factor
controlling the distribution, abundance and seasonal dynamics of
estuarine phytoplankton (Lacerda et al., 2004), by modifying the
physical and chemical properties of the water, in particular the
availability of nutrients, and the optical qualities of the water
(Bastos et al., 2005).
During the present study period, while precipitation presented
the typical seasonal pattern (Moraes et al., 2005), with 92% falling
between January and July, there was a 20.2% reduction in total
rainfall in comparison with the historical average for the period
1996-2005. In particular, precipitation was 66.6% and 45.3%
lower in January and February, respectively, in comparison with
the historical mean values (months with atypical characteristics of
dry season in the present study). This variation would have
affected both hydrological variables and the phytoplankton
community.
Water transparency and salinity varied considerably over the
study period, in a pattern similar to that observed in other
Brazilian estuaries and coastal systems (Resurreição et al., 1996;
Brandini, 1985; Branco et al., 2002; Koening et al., 2003; Losada
et al., 2003; Sousa et al., 2009), with the lowest values being
recorded during the period of highest precipitation.
The dissolved oxygen concentrations recorded during the study
indicate that Ajuruteua Beach is a supersaturated environment
(Macedo and Costa, 1978), with no clear seasonal pattern. This
may be related to the highly dynamic nature of the environment,
with strong winds and tidal currents that oxygenate the water
column continuously throughout the year, a pattern observed in
other, similarly turbulent Brazilian coastal systems (Campelo et
al., 1999; Losada et al., 2003).
Little variation was found in the pH of the water, which was
alkaline throughout the study period. This is a common pattern in
marine and estuarine ecosystems, in which the pH is controlled
primarily by the buffer effect of the seawater (Costa et al., 2008;
Sousa et al., 2009), as well as the high primary productivity which
increases the consumption of CO2, which leads to an increase in
the pH of the water (Branco et al., 2002; Bastos et al., 2005). A
similar pattern has been recorded at other sites in the Amazon
coastal zone (Costa et al., 2009; Pereira et al., 2010).
The structure and dynamic of the phytoplankton community at
Ajuruteua Beach were influenced primarily by hydrological
variables. Diatoms were the primary group responsible for the
high diversity and relative abundance recorded in both seasons.
The dominance of these organisms appears to be typical of
turbulent coastal areas influenced by strong winds and current
(Smayda, 1980), in which they may account for as much as 80%
of the composition of the phytoplankton community (Aidar et al.,
1993; Koening et al., 2003; Huang et al., 2004; Sousa et al.,
2008).
Dimeregramma minor was a dominant and very common
species throughout the study period. This species was most
abundant during the dry season, probably as a function of the
increased salinity during this period, given that D. minor is a
polyhalobe species, well-adapted to saline conditions (ValenteMoreira et al., 1994; Hassan et al., 2006). The strong local
hydrodynamic conditions were responsible for the resuspension of
sediments and associated benthic organisms, which explains their
abundance within the study area.
During the rainy season, when salinity was lowest and wind
speeds highest, A. glacialis and Skeletonema spp. were relatively
abundant, while D. minor was relatively scarce, indicating a
process of ecological succession during the annual cycle.
Santander et al. (2003) suggest that knowledge of the adaptability
of different phytoplankton species is essential for the
understanding the local dynamics of phytoplankton communities,
Journal of Coastal Research, Special Issue 64, 2011
356
Structure and dynamics of equatorial phytoplankton communities
especially in turbulent environments such as that of Ajuruteua,
because each algal population achieves a high rate of growth at a
different points in time. Given this, temporal distribution patterns
are related to the response of certain populations, rather than the
phytoplankton community as a whole, to fluctuations in
environmental factors such as precipitation (Branco et al. 2002;
Lacerda et al., 2004), salinity (Koening et al., 2003; Bastos et al.,
2005), and wind speed (Smayda, 1983; Sassi, 1991; Campelo et
al., 1999; Branco et al., 2002).
Phytoplankton biomass (chlorophyll a) was significantly higher
in the rainy season (March), and lowest in February, which in the
present study corresponded to the end of the dry season in 2005.
According to Sassi and Kutner (1982), the increase in
phytoplankton biomass in coastal environments during the rainy
season is related to the increased availability of dissolved nutrients
originating from the increased drainage of continental areas, as
observed in other Brazilian coastal areas (Campelo et al., 1999;
Branco et al., 2002; Bastos et al., 2005). During the present study
period, however, precipitation levels in January and February were
more typical of those of dry season months, which presumably
accounts for the relatively low chlorophyll a concentrations
observed in these months.
Phytoplankton densities were significantly higher during the
rainy season, in particular March and July, a pattern also observed
at a similarly dynamic beach on Canela Island in Pará (Sousa et
al., 2009). The group with the highest densities was the
phytoflagellates, which made up around 84% of the
phytoplankton. Melo et al. (2005) recorded a similar value (82%)
in a previous study at the same site. While phytoflagellates are
assumed to prefer calm oceanic environments, a similar pattern
has been recorded at other Brazilian sites (Brandini, 1985;
Koening et al., 2002; Lacerda et al., 2004; Sousa et al., 2009),
indicating that this is a common pattern in tropical and subtropical
coastal environments.
Diversity and evenness were the highest during the rainy
season, when A. glacialis, C. cymbeliformis, D. minor,
Skeletonema spp. and T. subtilis were relatively common. The
dominance of D. minor during the dry season was the main factor
responsible for the reduced diversity and evenness recorded during
this season.
The three groups identified by the cluster analysis reflected (i)
the influence of the highest precipitation (rainy season), i.e. the
lowest salinity, highest chlorophyll a concentrations and the
highest diversity and evenness; (ii) the reduced precipitation
during the dry season, when the highest salinity, the lowest
chlorophyll a concentrations, diversity and evenness, and the
dominance of D. minor was recorded, and (iii) the atypical
climatic conditions between December and February, when
abnormally low precipitation rates resulted in intermediate results,
with low chlorophyll a concentrations, diversity and evenness
more typical of dry seasons months.
FINAL CONSIDERATIONS
Overall, the results indicated that precipitation is the main factor
influencing the composition, density, biomass and diversity of the
phytoplankton community at the study site (Ajuruteua beach),
through seasonal fluctuations in the hydrological cycle. However,
other factors such as wind action, waves, and tidal currents also
appear to exert a strong influence on the hydrodynamics of the
Amazon coast, and must be taken into account in any analysis. In
particular, these factors promote the mixing of the water column,
which favors the resuspension of sediments and marine littoral
species such as D. minor, which was a crucial factor in the
structure of the phytoplankton community during some parts of
the year.
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ACKNOWLEDGEMENT
This study was supported by the Brazilian National Council for
Scientific and Technological Development (CNPq) and CAPES.
We thank Steve Ferrari for improvements to the manuscript’s
English.
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