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The influence of Ekman transport on

zooplankton biomass variability off
southern Brazil
C. RESGALLA JR1, C. DE LA ROCHA2 AND M. MONTÚ2
1 UNIVERSIDADE DO VALE DO ITAJAÍ – CTTMAR/UNIVALI – R. URUGUAI , CAIXA POSTAL . ITAJAÍ, SC. CEP .‒, BRAZIL, 2 FUNDAÇÃO
UNIVERSIDADE DE RIO GRANDE – DEPTO. DE OCEANOGRAFIA (FURG/DOC), CAIXA POSTAL , RIO GRANDE, RS. CEP ., BRAZIL

Cross- and long-shelf Ekman transport components were correlated with the variability of zoo-
plankton biomass from 1977 to 1990 off the southern Brazilian coast (27°S to 35°S and 46°W
to 54°W). During warm months the dominant cross-shelf Ekman transport was oriented offshore,
where an increase in zooplankton biomass occurred with intervals of 2–3 months. The presence of
the South Atlantic Central Water (SACW; the water mass that typically upwells at the continental
shelf) was confirmed by seasonal temperature–salinity diagrams. The high zooplankton biomass
values observed in this study were frequently caused by salps, which play an important ecological role
in the pelagic community in upwelling zones that has not been investigated yet.

I N T RO D U C T I O N during 12 years of continuous assessment of zooplankton

The energy that produces meso- and macro-scale circula-
tions is partially induced by wind stress, both in oceanic
and coastal environments. In coastal regions, wind action
can promote an offshore displacement of water masses by
Ekman transport. When this happens, upwelling of deep
waters to the euphotic zone may occur, resulting in modifi-
cations in the structure of the pelagic community. Such a
process is important for pelagic production, changing the
primary (phytoplankton) and secondary (zooplankton and
fish) trophic levels in the community.
One of the basic tenets of the hypothesis that explains
the oceanographic processes is the decomposition of
Ekman transport of water masses into long-shelf and
cross-shelf components (Myers and Drinkwater, 1989).
Pelagic production is mainly influenced by only one of
these two components, the cross-shelf component
vector. This component causes divergence of water
masses from the coast, resulting in upwelling of nutrients
into the pelagic zone. The long-shelf component of
water movement has little influence over the planktonic
community due to the advective process over the conti-
nental shelf. However, the latter can be related to patch
dispersion in high production areas (Mann and Lazier,
1991).
This study was initiated by the opportunity to manage
the data collected during 24 oceanographic cruises made
biomass along the southern Brazilian coast. The main
purpose of this study was to explore a possible relationship
between Ekman transport and zooplankton biomass,
observing if the data supported the Myers and Drink-
water hypothesis (Myers and Drinkwater, 1989).
METHOD
Wind vector and zooplankton biomass values were
obtained for the Santa Catarina and Rio Grande do Sul
coasts (27.0ºS to 35.0ºS and 46.0ºW to 54.0ºW) (Figure 1)
and averaged within a 1º latitude and longitude rectangu-
lar grid. The size of this grid was chosen with reference to
the availability of the wind data obtained by Diretoria de
Hidrografia e Navegação (DHN, Brazil).
A total of 18 301 data points of wind velocity and direc-
tion were analysed. Such data were obtained from the US
National Climatic Center (USA) and the DHN from 1976
to 1990. Data points with suspect values and those with
uncertain locations were removed from the set.
The Ekman transport was calculated by the ratio
between wind stress and the Coriolis effect (Apel, 1990),
using:
E = Tf (1)
where: E = Ekman Transport (ton m–1 s–1), T = wind stress

© Oxford University Press 2001

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Fig. 1. Study area. Geographical location in Latin America and the 1° latitudinal/longitudinal grid for wind (Ekman transport) and zooplankton
biomass data averaging.

Table I: List of 24 oceanographic cruises from 1977 to 1990 for which zooplankton biomass was
estimated in the study area

Period No.
Project Area Cruises (day/month) Year samples Reference

CONVERSUT 29°S to 34°S I 25/08–14/11 1977 181 Hubold, 1980a

II 08/04–27/06 1978 169 Hubold, 1980b
III 23/01–17/03 1981 108 –
AREPE 28°S to 34°S I 30/04–22/04 1980 100 –
II 29/07–18/08 1980 86 –
III 25/10–14/11 1980 90 –
IV 04/11–25/11 1981 59 –
V 14/10–05/11 1982 98 –
BONITO 28°S to 35°S I 24/03–31/03 1983 23 –
II 21/05–31/05 1983 15 –
III 11/10–20/10 1983 14 –
IV 11/12–11/12 1983 4 –
V 07/01–12/01 1984 5 –
VI 17/05–22/05 1984 9 –
VII 12/04–19/04 1985 6 –
VIII 15/08–15/08 1985 1 –
IX 11/12–21/12 1985 10 –
X 15/01–18/01 1986 9 –
XI 08/04–14/04 1986 5 –
XII 03/10–03/10 1986 5 –
XIII 21/01–22/01 1987 3 –
ECOPEL 32°S to 34°40’S I 10/10–28/10 1987 45 –
II 07/09–15/09 1988 50 –
III 06/02–21/02 1990 51 –
Total 1146

Many of these data have not been published and were obtained from internal reports (–). The projects full names are: CONVERSUT, Study of the Sub-
tropical Convergence Area; AREPE, Pelagic Stock Evaluation; BONITO, Study of tuna in the southern Brazilian coast; and ECOPEL, Study of the Pelagic
Ecosystems in Southern Brazil.

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C. RESGALLA JR ET AL. EKMAN TRANSPORT AND ZOOPLANKTON BIOMASS

(dyne cm–2), and f = Coriolis parameter as a function of

latitude.
The wind stress was calculated by the following equa-
tion:
2
T =  air C d V (2)

where: T = wind stress (dyne cm–2), air = air density

(0.00122 g cm–3), Cd = drag coefficient between water and
air (0.0013), and V = wind velocity vector (cm s–1). The
constants used in this work followed those of Lima and
Castello (Lima and Castello, 1995).
The Ekman transport long- and cross-shelf com-
ponents were obtained by vector decomposition on Car-
tesian co-ordinates, with the x-axis oriented along the
coastline. The results were positive for the long-shelf com-
ponent when the movement of the water mass was north-
ward and negative when water movement was southward,
and for cross-shelf components the results were positive
when movement was in an offshore direction and negative
when it was towards the coast.
The zooplankton biomass was assessed from 1146 data
points. Data were obtained from 24 oceanographic
cruises, made between 1977 and 1990 (Table I). The zoo-
plankton biomass was expressed in ml m–3, calculated by
the volume displacement method (Beers, 1976). Zoo-
plankton sampling varied in each cruise, with vertical
hauls using a 220 µm mesh (2.27% of the data set) and
oblique hauls using a 330 µm Bongo net (97.73%). No cor-
rections on the biomass values were made for the mesh
size utilized.
The relationship between Ekman transport com-
ponents and zooplankton biomass was assessed using
cross-correlation analysis, aiming to identify time lags
between events. In order to validate the results both para-
meters (Ekman transport components and biomass aver-
ages) were grouped in 2 and 3 month periods over the total
area.

R E S U LT S
The frequency distribution of the Ekman transport
analysed over the total study area showed an offshore
movement predominance, observed by the modal ten-
dency of the cross-shelf components (∑ = 68.2%, Figure
2). These results are a reflection of the northeast wind,
which is the most frequent on the studied area, except in
Fig. 2. Frequency distribution of the cross- and long-shelf Ekman trans-
winter. During winter the southern wind is dominant, pro- port intensity over the study area between 1976 and 1990. ∑ is the
moting an Ekman transport towards the shelf, but with probability sum of the Ekman transport distribution different from zero.
less intensity. No predominant water movement was
observed on the long-shelf component. Large northward
transport values were observed as a result of the strong
cold fronts which arrive in the area during winter.

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Fig. 3. Seasonal zooplankton biomass distributions averaged over 3 months in the 1° longitudinal–latitudinal grid. Data are in ml m–3 and the
number in parentheses is the total average by season.

Regarding the seasonal variation of the zooplankton was characterized by higher biomass values during spring.
biomass, higher values were observed in summer, mainly However, this result was influenced by data from a single
in the near-shore zone (Figure 3). This spatial tendency cruise and could be an over-estimation for this region.
was similar for all seasons, but the southern open-sea zone The high correlation coefficient (>0.70) obtained by

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C. RESGALLA JR ET AL. EKMAN TRANSPORT AND ZOOPLANKTON BIOMASS

Table II: Cross-correlation coefficients of Ekman transport components (input) and zooplankton
biomass (output), averaged over the total area and grouped over 2 and 3 months

2 months 3 months

Lag Long Cross Lag Long Cross

0 –0.425 0.122 0 –0.553 0.921

(0.401) (0.818) (0.447) (0.079)
1 –0.737 0.830 1 –0.240 –0.153
(0.095) (0.041) (0.760) (0.847)
2 0.528 –0.092 2 0.801 –0.285
(0.282) (0.863) (0.199) (0.715)
3 0.056 –0.049 3 –0.007 0.483
(0.916) (0.926) (0.993) (0.517)
4 0.353 –0.501
(0.493) (0.312)
5 0.225 –0.409
(0.668) (0.421)

Significance levels are in parentheses (calculated by analysis of variance). Values in bold indicate high correlation coefficient.

cross-correlation analysis suggests a strong relationship DISCUSSION

between the Ekman transport cross-shelf component and
zooplankton biomass (Table II). The temporal treatments
pointed out a time lag of 2–3 months between the events
and their effects, with a positive correlation in both cases.
The Ekman transport long-shelf component did not show
coherent results. Despite the smaller correlation co-
efficients obtained (and significance level), the 2 and 3
month temporal treatments did not agree with each other,
with an observed time lag of 2 months (lag 1) and 6
months (lag 2) for each treatment, respectively. Addition-
ally, the highest correlation coefficients showed different
signs, negative for 2 month and positive for 3 month data
groups.
Zooplankton biomass increased with the intensity of
cross-shelf transport. Large values in January/February
were related to high biomass values in March/April (lag
1), and large transport in summer had large biomass in
the same period (lag 0) (Figure 4). This supports the
hypothesis that zooplankton biomass increases with an
increment of the offshore Ekman transport component.
The long-shelf transport component displayed a positive
relationship with zooplankton biomass when the trans-
port was southward, i.e. strong longshore transport in
January/February resulting in large zooplankton biomass
in March/April (lag 1) (Figure 5). During the winter,
small southward transport was related to high biomass in
summer (lag 2). Long-shelf transport and biomass were
weakly related.
The results of this study support the idea of the dynamic
processes presented by Myers and Drinkwater and
Wickett on production in pelagic communities (Wickett,
1967; Myers and Drinkater, 1989). The predominance of
the cross-shelf Ekman transport component all year long
suggested a strong tendency towards a positive upwelling
index in this area (Bailey, 1981; Livingstone and Royer,
1980). The same fact was observed both in this area (Lima
and Castello, 1995) and in the São Paulo Bight (Bakun
and Parrish, 1990). These studies found that reproductive
strategies of coastal pelagic fish are related to seasonal
variability of water mass movement over the continental
shelf.
The cross-shelf water mass component showed a more
positive mode in its frequency distribution than the long-
shelf component (Figure 2). These results above suggest a
predominance of water mass movement offshore, result-
ing in cold nutrient-rich water upwelling in the euphotic
zone between spring and autumn. This creates conditions
for the development of primary (phytoplankton) and
secondary production (zooplankton). The phytoplankton
responds quickly to the nutrient input (on a daily time
scale), but the response of the herbivorous community
takes more time to develop (with a weekly or monthly time
scale) (Parsons et al., 1984).
The multiple cross-correlation analysis tends to
produce random high correlation indices (Chelton, 1982).

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Fig. 4. Relationship between zooplankton biomass and cross-shelf Ekman transport averaged over the whole study area at two temporal scales (2
and 3 months). The months and seasons near the points refer to the Ekman transport period and the time lag biomass, where: J-F, January and
February; M-A, March and April; M-J, May and June; J-A, July and August; S-O, September and October; N-D, November and December; SM,
summer; A, autumn; W, winter and SP, spring.

In the present case the validation step involved changing
the values of the temporal treatment to obtain similar
results. This was done by averaging the data over groups
of 2 and 3 months. In both cases the results confirmed the
trend of a positive relationship between cross-shelf trans-
port and biomass.
The northern region of the study area (27°S to 31°S) is
characterized by an upwelling process over the continen-
tal shelf (Castello and Moller, 1977; Hubold, 1980a,
1980b; Brandini, 1986; Matsuura, 1986; Odebrecht and
Djurfeldt, 1996), but these studies were limited to data col-
lection and oceanographic aspects. The primary and
secondary production in the southern coastal region (31°S
to 35°S) are supplied with nutrients by the seasonally vari-

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C. RESGALLA JR ET AL. EKMAN TRANSPORT AND ZOOPLANKTON BIOMASS

Fig. 5. Relationship between zooplankton biomass and the long-shelf Ekman transport averaged over the whole study area at two temporal scales
(2 and 3 months). The months and seasons near the points refer to the Ekman transport period and the time lag biomass, where: J-F, January and
February; M-A, March and April; M-J, May and June; J-A, July and August; S-O, September and October; N-D, November and December; SM,
summer; A, autumn; W, winter and SP, spring.

able continental water inputs from the La Plata River, the
Patos Lagoon estuary and the coastal branch of the Mal-
vinas Current (Miranda, 1972; Castello and Moller, 1977;
Hubold, 1980a, 1980b). The high zooplankton biomass
observed in this region may be due to local primary pro-
duction or to transport from the Argentinean coastal shelf.
In the São Paulo bight, north of the study area, the
Cabo Frio and Ubatuba upwelling centres are found (Mat-
suura, 1986; Valentin, 1988; Pires-Vanin and Matsuura,
1993; Pires-Vanin et al., 1993). The high zooplankton
biomass found in this area may be transported south,
which would explain the observed negative correlation
between long-shelf transport and biomass mainly in the
northern coastal region of the study area.

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Fig. 6. Temperature–salinity diagrams grouped by north and south coastal regions of the study area at the seasonal scale (3 months) between 1976
and 1990; data collected by DHN.

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C. RESGALLA JR ET AL. EKMAN TRANSPORT AND ZOOPLANKTON BIOMASS
The water mass that upwells in the Brazilian
south–southwest coast is the South Atlantic Central Water
(SACW), characterized by temperatures <20°C and salin-
ity between 34.2 and 36.0 ppt (Ciotti et al., 1995). This
water mass frequently occurs in the region of the deep
shelf slope in southern Brazil. To verify the presence and
variability of this water mass along the north (27°S to
31°S, <200 m deep) and south (31°S to 35°S, <200 m
deep) coasts of the study area, temperature and salinity
data were obtained from DHN for the same period, and
plotted in temperature–salinity diagrams, grouped seas-
onally (Figure 6). The main difference observed between
regions was the strong presence of the Coastal Water (with
influence of continental water, salinity <30 ppt) in the
south region during most of the year, but mainly in winter.
The presence of the SACW in both regions may be
related to data from the shelf break. This water is related
to the northern limit of the west boundary of the south-
ern subtropical convergence zone (Godoi, 1982), where
the presence of subantarctic water is observed during the
winter. In the northern area, the SACW was probably
related to upwelling processes. It probably had a reduced
influence during the winter due to the offshore position of
the southern subtropical convergence zone and the higher
influence of the coastward cross-shelf Ekman transport
(Castello et al., 1997).
It should be noted that the zooplankton biomass data
used in this study do not provide information about the
specific composition of the samples. The large values
observed in the coastal zone were frequently related to
groups with gelatinous organisms. Salps were the most
common organism in the samples, and their larger size prob-
ably biased the data, causing overestimation of the biomass
values if displacement volume methods were used
(Meneghetti, 1973). The energetic value of salps is low (high
water content), but the success of salps in upwelling regions
is a consequence of the high individual and populational
growth rates (Heron, 1972a, 1972b). Such behaviour could
thus explain the events on the temporal scale presented in
this paper. Another point is that the high grazing and low
assimilation rate of salps can change the pelagic community
structure size and assist the transport of primary production
to the benthic community through faecal pellets of non-
digested phytoplankton (Esnal, 1981; Blackburn, 1979).
High density of salps has also been observed in upwelling
regions in the São Paulo bight (Pires-Vanin et al., 1993), but
the quantification of these processes along the Brazilian
coast waters has not yet been investigated.
AC K N O W L E D G E M E N T S
This study was supported by FAPERGS – Scientific Fund
of Rio Grande do Sul, Brazil. The authors acknowledge

the Scientific Fund of the Rio Grande do Sul State
(FAPERGS) for a scholarship for C. R. J., DHN (Brazilian
Navy) and Dr Sinque (Icthioplankton Laboratory of the
FURG) for supplying, respectively, wind and zooplankton
biomass data. Thanks to Carlos Augusto Schetini, André
Barreto and the two anonymous referees, who helped in
reviewing an early manuscript.
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Received on August 15, 2000; accepted on March 12, 2001