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ABSTRACT
This work had as objective to study the distribution and abundance of the Carangidae larvae and to analyze the
influence of the hydrological (temperature and salinity) and biological factors (phytoplanktonic biomass and
zooplanktonic biomass), on the space and temporal distribution of the larvae. Ichthyoplankton was collected during
four expeditions from the Northeast Exclusive Economic Zone. Six species (Trachurus lathami, Decapterus
punctatus, Chloroscombrus chrysurus, Selene setapinnis, Selene vomer and Elagatis bipinnulata) and Caranx-
Carangoides complex were identified. D. punctatus was the species most abundant (52% of the total), with higher
abundance during the Period 3, while the Period 2 was the period of low abundance. C. chrysurus was the second
species in abundance representing 30% of the total of carangid. This species had higher abundance during the
Period 2 and the Period 1. However, in Period 3 abundance were lesser. The third species in abundance was T.
lathami that corresponded 8% of the total of carangid larvae. S. setapinnis, S. vomer and E. bipinnulata were the
species less abundant, representing together 2% of the total identified larvae. The larvae of Caranx- Carangoides
complex represented 9% of the carangid total.
Key words: Ichthyoplankton, Carangidae, oceanographic factors, Exclusive Economic Zone, Northeast Brazilian
coast
*
Author for correspondence
little is know about the larval abundance and Parnaíba River, in the Piauí, and Todos os Santos
distribution patterns of the carangid in Brazil with Bay, in the Bahia, included the Archipelagos of
the exception of the distribution and abundance of Fernando de Noronha and Saint Peter and Saint
Chloroscombrus chrysurus larvae along the Paul. This is tropical oceanic region, constituted of
Southern coast investigated by Weiss et al. (1976). a oligothrophic system, and that it possesses a
Katsuragawa and Matsuura (1992) described nine complex alimentary web, however with low
taxa in the Southeastern Brazilian. Recent studies biological productivity (Ekau and Knoppers,
about the ichthyofauna of the Northeast Exclusive 1999). The South Equatorial Current (SEC)
Economic Zone also present information about the reaches the NE Brazilian shelf between 11 and 15º
occurrence of Carangidae (Lessa et al., 2000; S. A total of 12 Sv or more from SEC flows
Feitosa et al., 2003). northward to continue as the North Brazil Current
The distribution patterns of the fish larvae in any (NBC). Only 4 Sv from the southward flowing
region of the ocean are related to the reproductive Brazil Current (BC) (Peterson and Stramma,
activity of the adult population and to the 1991).
topographic and hydrographic features that affect The SEC comprises a broad westward flow with a
the dispersal of the larvae (Nonaka et al. 2000). In mean velocity of 10 to 15 cm.s-1 and along the
addition, information can be obtained on the equator there is a much swifter current with a
reproductive strategies adopted by these fishes in mean velocity of 30 cm.s-1 (Tchernia, 1980). The
response to the physical and biological processes North Brazil Current follows at a mean velocity of
of the region (Vorwerk et al., 2003). 75 cm.s-1 with superficial temperature between 28
Alterations in the physical conditions such as and 30 ºC, and superficial salinity between 35.0
habitat structure would be expected to result in and 37.0 psu (Medeiros et al., 1999). The Brazil
changes in the fish community structure owing to Current flows at a mean velocity of 10 to 15 cm.s-1
the effect that this change would have on the with mean superficial temperature of 26 ºC and
abundance and composition of the species salinity above of 35 psu (Tchernia, 1980).
associated with it. The interactions between the
various physical, chemical and biological
properties therefore explain why the ecology of MATERIALS AND METHODS
marine display changes in time and space
(Whitfield and Paterson, 2003). The knowledge of A total of 562 samples were collected during four
these interactions is important for a better expeditions realized between 1995 and 2000 (Fig.
understanding of the interrelationships among fish 1). The expeditions were made during August to
species during their early life stages, as well as an October of 1995 (Period 1), January to April of
understanding of adult spawning patterns, 1997 (Period 2), April to July of 1998 (Period 3)
recruitment processes and the overall dynamics of and September to December of 2000 (Period 4).
fish populations. The ichthyoplankton samples were collected by
The present work analyzed the occurrence of the Bongo nets with 50 cm of mouth diameter, 500
Carangidae larvae, spatial and temporal and 300 µm meshes, but only larvae collected with
distribution patterns of the most abundant species 500 µm net were considered in this study. The nets
and evaluated the influence of oceanographic were equipped with two independent flowmeters
factors (temperature, salinity, primary biomass and to estimate the water volume filtered.
secondary biomass) on them in the Exclusive The sampling method used followed Smith and
Economic Zone of Brazil Northeast. Richardson (1977). The sampling was done
through oblique hauls from the maximum depth of
Study area 200m to the surface. The duration of the tow was
The Exclusive Economic Zone (EEZ) is defined as 10 minutes. The samples obtained were preserved
the area between the external limit of the in 4% buffered formalin – seawater. In the
Territorial Sea, of 12 miles of width and 200 laboratory, all the Carangidae larvae were
nautical miles of the coast (CNIO, 1998). This removed from each sample and stored in 70%
study is part of the REVIZEE Program alcohol.
(Assessment of the Sustainable Yield of the Living The carangidae larvae were identified to the lowest
Resources in the Exclusive Economic Zone). The possible taxonomic level according to the
study area was limited by the estuary of the morphological characters of each group
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
Distribution and Abundance of Carangidae (Teleostei, Perciformes) Associated 1269
(Katsuragawa, 1990). Standard densities of the 100 m3 of filtered water for each collection.
individual taxa were expressed as the number per
0º S
2º S
Parnaíba River
Fortaleza
4º S
Natal
6º S
Latitud
Recife
8º S
NORTHEAST REGION
BRAZIL
Maceió
10º S
Aracajú
12º S
Salvador
14º S
42º W 40º W 38º W 36º W 34º W 32º W 30º W 28º W
Longitud
Temperature and salinity were record by ctd. and respective sample. The canonical
The water samples for the determination of the correspondence analysis was used to study the
phytoplanktonic biomass (chlorophyll a) in 1% of relationships between the abundance of the family
light were obtained using the fluorescence method. Carangidae and oceanographic variables
The determination of the secondary biomass (dry (temperature, salinity, primary biomass and
weight) was carried through according to secondary biomass).
methodology of Omori and Ikeda (1984).
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
1270 Souza, C. S. and Mafalda Júnior, P.
30
Temperature (ºC)
28
26
24
P erio d 1 P erio d 2 P erio d 3 Perio d 4
Figure 2 - Temporal variation of temperature for investigated periods (minimum and maximum
values: vertical line; mean: horizontal bar; confidence limit: rectangle).
There were significant differences in the Salinity ranger from 35.5 to 37.3 (mean = 36.1)
temperature between the four analyzed periods during the period 1, between 34.8 and 37.2 (mean
(Kruskal-Wallis, p < 0.0001). The test of multiple = 36.2) during the period 2, between 34.5 and 37.4
comparisons of Dunn showed that the periods 1 (mean = 36.4) during the period 3 and between
and 4 differed from the periods 2 and 3. 35.2 and 37.2 (mean = 36.3), during period 4 (Fig.
3).
38
37
Salinity (psu)
36
35
34
P erio d 1 P eriod 2 Perio d 3 P eriod 4
Figure 3 - Temporal variation of salinity for investigated periods (minimum and maximum values:
vertical line; mean: horizontal bar; confidence limit: rectangle).
The salinity was significantly different among the in all the periods, indicated the presence of the
periods (Kruskal-Wallis, p < 0.0001). The test of Superficial Equatorial Water, which had a salinity
the multiple comparisons of Dunn showed that the > 35 and temperature > 26º C, and Coastal Water,
periods 1 and 2 differed from the periods 3 and 4. with salinity around 35 (Fig. 4).
The values of salinity and temperature registered
PERIOD 1 PERIOD 2
30 30
CW SEW CW SEW
Temperature (ºC)
Temperature (ºC)
29 29
28 28
27 27
26 26
25 25
34 35 36 37 38 34 35 36 37 38
Salinity (PSU) Salinity (PSU)
PERIOD 3 PERIOD 4
30 30
CW SEW
Temperature (ºC)
CW SEW
Temperature (ºC)
29 29
28 28
27 27
26 26
25 25
34 35 36 37 38 34 35 36 37 38
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
Distribution and Abundance of Carangidae (Teleostei, Perciformes) Associated 1271
Chlorophyll a (µg/l)
5
0
P erio d 1 P erio d 2 P erio d 3 P erio d 4
Figure 5 - Temporal variation of chlorophyll a for investigated periods (minimum and maximum
values: vertical line; mean: horizontal bar; confidence limit: rectangle).
15
Dry Weight (g/100 m3)
10
0
P erio d 1 P erio d 2 P erio d 3 P erio d 4
Figure 6 - Temporal variation of dry weight for investigated periods (minimum and maximum
values: vertical line; mean: horizontal bar; confidence limit: rectangle).
Carangidae larvae occurrence and composition abundance, representing 29.7%. The higher
A total of 313 larvae representing six species abundance occurred during period 2. T. lathami
(Trachurus lathami, Decapterus punctatus, represented 7.7% of total Carangidae larvae and
Chloroscombrus chrysurus, Selene setapinnis, was more abundance during period 4. Selene
Selene vomer and Elagatis bipinnulata) and two setapinnis, Selene vomer and Elagatis bipinnulata
genera (Caranx and Carangoides) were identified were less abundant, contributing 2.2% of total
in the EEZ Northeast. D. punctatus, C.oscombrus larvae. The members of Caranx and Carangoides
chrysurus and Trachurus lathami were the most complex comprised less than 9% of the total (Fig.
dominant species comprising 89.1% of the larvae 7).
composition.
D. punctatus was the most abundance species Spatial and temporal distribution of species
(51.8%). The higher abundance were observed The carangidae larvae were collected more
during periods 1 and 3, while period 2 was lower frequently within the neritic zone of the Northeast
abundance. C. chrysurus was the second species in Brazil.
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
1272 Souza, C. S. and Mafalda Júnior, P.
100
80 Period 1
60 Period 2
%
40
Period 3
20
Period 4
0
Decapterus
Trachurus
setapinnis
Carangoids
bipinnulata
Chloroscombrus
Selene vomer
punctatus
lathami
Complex
Elagatis
Selene
Caranx
chrysurus
Figure 7 - Relative abundance of species for investigated periods.
D. punctatus presented large distribution occurring the Parnaíba river mouth, during the period 2. This
in the neritic and oceanic region. D. punctatus species did not occur during period 1 (Fig. 8). T.
larvae were collected in all the periods with the lathami occurred mainly in the neritic stations,
maximum density during period 3 in the area between Natal and Aracaju, during the period 3
between Parnaíba and Salvador (Fig. 8).C. and in the Fernando de Noronha Archipelago
chrysurus larvae were found predominantly in the during the period 4 (Fig. 8).
neritic stations. The highest density occurred near
F. Noronha F. Noronha
Parnaíba Archipelago Parnaíba Archipelago
4ºS Fortaleza 4ºS Fortaleza
8ºS
PERIOD 1 Recife
8ºS PERIOD 2 Recife
08/02 to 10/26 01/21 to 04/13
Trachurus lathami
1995 Maceió
Chloroscombrus chrysurus 1997 Maceió
10ºS Decapterus punctatus 10ºS
3
Aracajú Larvae/100m Aracajú
12ºS 0,1 - 5 12ºS
Salvador 5 - 10 Salvador
10 - 35
14ºS 14ºS
42ºW 40ºw 38ºW 36ºW 34ºW 32ºW 30ºW 28ºW 42ºW 40ºW 38ºW 36ºW 34ºW 32ºW 30ºW 28ºW
Aracajú Aracajú
12ºS 12ºS
Salvador Salvador
14ºS 14ºS
42ºW 40ºW 38ºW 36ºW 34ºW 32ºW 30ºW 28ºW 42ºW 40ºW 38ºW 36ºW 34ºW 32ºW 30ºW 28ºW
Correlations between Carangidae larvae and between 0.2 and 1.4 µg.L-1 and between 0.5 and
oceanographic factors 2.3 g.100m-3, respectively.
The higher Decapterus punctatus densities were The highest Trachurus lathami densities occurred
found in a wide temperature range between 26.1º in the temperature between 25.4 to 26.8º C.
C and 28.9º C. The salinity of all positive stations The salinity variation was between 34.99 and
varied from 36.7 to 37.3, the primary biomass 36.36. T. lathami larvae were found in the
varied from 0.69 to 2.70 µg.L-1 and secondary secondary and primary biomass range between
biomass 0.28 to 0.90 g.100m-3. 0.95 and 1.4 µg.L-1 and between 0.31 and 1.2
Temperature variation in positive stations of g.100m-3, respectively.
Chloroscombrus chrysurus was between 28.4 and Eigenvalues, measures of the importance for the
29.2º C. Salinity variation was between 36.5 and CCA axes that can vary between zero and one,
37.2. ranged from 0.022 for CCA 2 to 0.701 for CCA 3
Secondary and primary biomass variation was (Table 1).
The first two CCA axes explained 89.5% of the gradients were important correlates with the
cumulative percentage variance of the species- abundance of Carangidae family in the CCA. The
environmental. The Monte Carlo test was temperature water, primary biomass and secondary
significant for the first CCA axes (p = 0.012) and biomass correlated particularly well with the CCA
all CCA axes (p = 0.016). The low multiple axes 1, and the salinity correlated well with the
regression coefficients of the environmental CCA axes 2 (Fig. 9). These environmental
variables indicated that there was no collinear gradients also reflected the spatial and temporal
variables. This was important because changes in the density species.
multicollinear variables might be deleted from the C. chrysurus was positively correlated with first
analysis. CCA axes. D. punctatus was negatively correlated
The plot of CCA sample and species scores with the first and second CCA axes and T. lathami
illustrates their dispersion pattern and the plot of was positively correlated with the second CCA
the oceanographic variables vectors illustrat the axes. This implied that C. chrysurus (Chl)
directions and strengths of the environmental occurred mostly at high water temperature and
relationships within the first two dimensions of the secondary biomass during the period 2. D.
CCA ordination (Fig. 9). punctatus (Dec) appeared preferentially during the
The dispersion of CCA sample and species scores period 3 at high salinity and primary biomass,
resulted from the linear combinations of the whereas T. lathami (Tra) predominantly occurred
oceanography variables. Strong environmental at low salinity during the period 4 (Fig. 9).
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
1274 Souza, C. S. and Mafalda Júnior, P.
3
2
Tra
3 11
31 3
2 4
2 Temperature 3 4
Secondary Biomass Chl
2 3 4 4 4 3
2 4 4 4 4
2 2 24 3 4 4
2 1 4 4
2 2 2
4 4
3 3 13 3
3 3 3 3 4 3 Primary Biomass
4 3 3 33
3 3
4 3 Dec
2
Salinity
-2
-3 3
Figure 9 - Ordination diagram of the Canonical Correspondence Analysis (CCA) (1: Period 1; 2:
Period 2; 3: Period 3; 4: Period 4).
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
Distribution and Abundance of Carangidae (Teleostei, Perciformes) Associated 1275
reduce the co-occurrence of the larvae. Several (C. chrysurus, T. lathami, D. punctatus, C. crysos,
species spawn offshore, but their larvae return to O. saurus and H. amblyrhynchus) in
nursery grounds located in the shallow, inshore hydrographically dynamic areas.
waters. This reproductive style appeared to Variation in the oceanographic environmental on
maximize the probability of the survival through the annual scale may cause interannual changes in
the planktonic phase and has been well both the distributional range of the adults fishes
documented for some species of the coastal fishes and the features of their spawning environment
in the tropics, including the Carangidae (Vásquez- such as the timing, duration and location of the
Yeomans, 2000). The period of occurrence of spawning (Doyle et al., 1993). Thus, studies have
some groups was restricted to the Period 2, like the examined the larval distributions within a dynamic
cases of E. bipinnulata and S. vomer. E. framework as simple hydrographic correlations
bipinnulata spawning in the oceanic waters have not explained the significant amounts of the
(Aprieto, 1974) and adults were common offshore variability in the distribution of the assemblages or
(Nakamura, 1980). in the recruitment process (Govoni, 1993; Cowen
The cases where the occurrence was during all the et al., 1993; Dempster et al., 1999). The
year, was observed the existence of a preferential meteorological and oceanographic conditions
period; Period 4 for Caranx-Carangoides influence the availability of the food organisms
complex, Period 3 and Period 4 for T. lathami and and the eggs and larval transport by the residual
D. punctatus, and Period 2 for C. chrysurus. This current and thus affect the reproductive success,
study showed that the Carangidae fish presented resulting in the recruitment variation (Norcross
similar reproductive strategies, spawning at a and Shaw, 1984).
favourable period for each species. It is known that Examining the spatial and temporal patterns in the
most of the species have characteristic distribution and abundance of the ichthyoplankton
reproductive strategies, adjusting their in relation to the oceanographic conditions may
reproductive patterns with the environmental provide insight into the adaptation of the spawning
factors, such as the surface current. For example, strategies to the prevailing physical and biological
in the northeast Pacific, many coastal species processes as well as into the effect of the
which have pelagic stages of the larvae, tend to variability in these processes on year – class
spawn during the winter, when wind – induced strength (Somarakis et al., 2002). Among the
surface transport directs onshore and not during causes that influence the distribution of the
the intensive upwelling season when there is a ichthyoplankton is the trophic relations of the
strong offshore transport (Katsuragawa and spawning adult populations and the transport of
Matsuura, 1992). the ichthyoplankton by chains (Arosemena et al.,
Favourable larval habitats have been defined by 2000), while the initial phases study of the fish
both their biological (e.g. high abundance of food, life cycle permit an adequate and more profitable
low abundance of predators) and physical (e.g. use of fishery resources (Lasker, 1987).
circulation patterns promoting retention or In conclusion, the results this study demonstrated
transport to nursery area) characteristics (Heath, that the studied oceanographic factors
1992). However, the environmental conditions at (temperature, salinity, primary biomass and
the time of egg and larval development may differ secondary biomass) appeared to influence the
from year to year, due to variation in the abundance of the Carangidae species, allowing
environmental characteristics, changes in the that each species had a period and place more
timing of emergence of eggs and larvae or a favourable to spawn in the Brazil Northeast
combination of both (Page and Frank, 1989). Exclusive Economic Zone.
T. lathami larvae had the most densities in less
saline waters in comparision to the D. punctatus
and C. chrysurus. Larvae of C. chrysurus occurred ACKNOWLEDGEMENTS
in higher temperature comparing with D.
punctatus and T. lathami. There was spatial and The authors express their thank to the Ministry of
temporal variation in the distribution and Environment, Direction of Hydrography and
abundance, and possibly in the reproductive cycle Navigation and ANTARES oceanographical ship
of the Carangidae species. Ditty et al. (2004) for supporting this study though the program
found the larvae of several species of the carangids REVIZEE.
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
1276 Souza, C. S. and Mafalda Júnior, P.
Braz. arch. biol. technol. v.51 n.6: pp. 1267-1278, Nov/Dec 2008
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