RELATIVE EFFECT OF SEASONALITY AND BODY SIZE ON THE DIET OF JUVENILE
Lutjanus synagris (PERCIFORMES: LUTJANIDAE) AT A SANDY BEACH IN
SOUTHEASTERN BRAZIL
Daniel de Melo ROSA1; Ciro Colodetti VILAR1,2; Joelson MUSIELLO-FERNANDES1,3
ABSTRACT
The relative effect of seasonality and body size on the diet composition of juvenile Lutjanus synagris
was investigated at a sheltered sandy beach (20º18’S; 40º16’W) on the coast of Espírito Santo state,
Brazil. Monthly collections were carried out during the daytime from May 2004 to April 2005, using
a beach seine. The specimens captured in each season (e.g. cold/dry, May to October 2004;
hot/rainy, November 2004 to April 2005) were separated into three total length (TL) classes (≤ 40;
41–50; ≥ 51 mm). The stomach contents of 540 specimens with TL ranging from 22 to 135 mm were
analyzed. Among the 13 food items consumed by L. synagris, the most important were Amphipoda
(Index of Relative Importance, IRI = 81.2), Isopoda (IRI = 13.8) and Mysidacea (IRI = 4.4).
Significant variations in the diet composition were observed between the seasons (PERMANOVA;
P < 0.001), but not between the size classes (P = 0.170). There was, however, a significant interaction
between these two factors (P<0.001), indicating that the differences between the seasons did not
occur in every class. The most important items in the cold/dry and hot/rainy seasons were,
respectively, Mysidacea and Amphipoda, for every class. The average length of the preys varied
significantly (P<0.05) between the size classes, increasing progressively with the TL of the fishes.
Overall, the results indicate a carnivore-opportunist food habit for L. synagris, with significant
seasonal changes in the relative importance of the food items.
Keywords: food habit; trophic dynamics; Actinopterygii; sheltered beach; Atlantic ocean
EFEITOS RELATIVOS DA SAZONALIDADE E DO TAMANHO DO CORPO NA DIETA DE
JUVENIS DE Lutjanus synagris (PERCIFORMES: LUTJANIDAE) EM UMA PRAIA
ARENOSA NO SUDESTE DO BRASIL
RESUMO
O efeito relativo da sazonalidade e do tamanho do corpo na composição da dieta de juvenis de
Lutjanus synagris foi avaliado em uma praia arenosa abrigada (20º18’S; 40º16’W) no litoral do estado
do Espírito Santo, Brasil. Coletas mensais diurnas foram conduzidas no período de maio/04 a
abril/05, usando uma rede de arrasto de praia. Os espécimes capturados em cada estação
(fria/seca, maio a outubro de 2004; quente/chuvosa, novembro de 2004 a abril de 2005) foram
separados em três classes de comprimento total (CT) (≤ 40; 41–50; ≥ 51 mm). O conteúdo estomacal
de 540 espécimes com CT variando de 22 a 135 mm foi analisado. Entre os treze itens alimentares
descritos para a dieta de L. synagris, os mais importantes foram Amphipoda (Índice de Importância
Relativa, IIR = 81,2), Isopoda (IIR = 13,8) e Mysidacea (IIR = 4,4). Variações significativas na dieta
foram observadas entre as estações do ano (PERMANOVA; P<0,001), mas não entre as classes de
tamanho (P = 0,170). Houve, porém, uma interação significativa entre estes dois fatores (P<0,001),
indicando que as diferenças entre as estações não ocorreram em todas as classes. Os itens mais
importantes nas estações fria/seca e quente/chuvosa foram, respectivamente, Mysidacea e
Amphipoda, para todas as classes. O tamanho médio das presas variou significativamente (P<0,05)
entre as classes de tamanho, aumentando progressivamente com o CT dos peixes. No geral, os
resultados indicam que L. synagris têm um hábito alimentar carnívoro-oportunista e muda sua
dieta ao longo do ano.
Palavras chave: hábito alimentar; dinâmica trófica; Actinopterygii; praia abrigada; oceano Atlântico
Artigo Científico: Recebido em 09/01/2014 – Aprovado em 23/10/2014
Núcleo de Atividades Ambientais – NATIVA. Av. Hugo Musso, 1333, apto 1106 – CEP: 29101-280 – Vila Velha – ES –
Brazil. e-mail: daniel@nativa.org.br (corresponding author)
2 Programa de Pós-Graduação em Ecologia e Conservação, Universidade Federal do Paraná (UFPR). Caixa Postal 19031 –
CEP: 81531-980 – Curitiba – PR – Brazil. e-mail: cirovilar@nativa.org.br
3 Programa de Pós Graduação em Oceanografia Ambiental, Universidade Federal do Espírito Santo (UFES). Av. Fernando
Ferrari, 514 – CEP: 29075-900 – Vitória – ES – Brazil. e-mail: joelson.pesca@gmail.com
1
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
20
ROSA et al.
INTRODUCTION
The first year of life is a period of rapid
growth for fish species and is commonly marked
by alterations in food habits (WOOTTON, 1990).
Such alterations generally occur due to
morphological changes that accompany growth,
as well as modifications in the availability of prey
during the daily or seasonal cycle (GERKING,
1994; CORRIGAN et al., 2011; DAVIS et al., 2012).
Studies of the diet of juveniles may help in the
understanding of the relative importance of
factors acting as drivers of such changes in the
food spectrum and also the trophic role of these
fish in nursery habitats. In addition, ecological
information may even apply to the culture of
economically important fish species, for instance,
in the production of different types of food for
specific sizes of fish (ZAVALA-CAMIN, 1996).
Lutjanids are important fishery resources in
tropical and subtropical regions (MOURA et al.,
2011; CAVALCANTE et al., 2012), and occupy
high trophic levels in the marine ecosystems
(DUARTE and GARCIA, 1999). The young live
mainly in shallow areas, while adults look for
deeper waters off the coast where they usually
spawn (FREITAS et al., 2014). The lane snapper
Lutjanus synagris (Linnaeus, 1758) is distributed
from North Carolina (USA) to southeastern Brazil,
but is more abundant in the Antilles, Campeche
Bank and on the northern coast of South America
(ALLEN, 1985). This species inhabits coral reefs,
seagrass banks, estuaries and dark brackish
waters over muddy bottoms (MCEACHRAN and
FECHHELM, 2005; GARCIA-JÚNIOR et al., 2010).
It is considered important in the commercial and
recreational fisheries in the northern region of the
Gulf of Mexico (GMFMC, 1981) and Brazil, where
it occupies the first position in the artisanal fishery
of Abrolhos Bank (FONSECA, 2009).
Despite its economic importance in Brazilian
waters, studies on the diet of L. synagris have been
carried out manly in the Caribbean Sea (e.g.
DUARTE and GARCIA, 1999; FRANKS and
VANDERKOOY, 2000). The limited information
available coming from the Brazilian coast suggests
this species acts as an important predator in the
food webs of coastal habitats, such as reefs and
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
estuaries (FONSECA, 2009; PIMENTEL and
JOYEUX, 2010). Therefore, the functional role of
juvenile snappers at alternative nursery grounds
such as sheltered sandy beaches is unknown,
although they can figure among the most
abundant species in these environments (e.g.
ARAUJO et al., 2008). This study aims to fill this
gap by investigating if seasonality and body size
are important drivers of changes in the diet
composition of L. synagris inhabiting a sheltered
sandy beach in southeastern Brazil.
MATERIAL AND METHODS
Study area
The samplings were carried out at a sheltered
beach on Ilha do Frade (20º18’S; 40º16’W), located
about 380 m from the coast, between the north
(Canal da Passagem) and south (Baía de Vitória)
inlets of the estuary that surrounds Vitória (capital
of Espírito Santo state, southeastern Brazil)
(Figure 1). Its coastline is formed predominantly
by granitic rocks, with small sandy beaches
between the rocky formations. These beaches are
occupied in some periods of the year by
macroalgae (e.g. Ulva Linnaeus, 1753 and
Sargassum C. Agardh, 1820) detached from
adjacent rocky shores by wave action. As in other
metropolitan
regions,
diverse
potentially
impacting activities occur around the study area,
including the extensive urbanization of the island
and its surroundings, the presence of three large
ports (i.e. Vitória, Capuaba and Tubarão) and the
entry into Baía de Vitória of part of the industrial
and domestic sewage produced by 1.02 million
inhabitants. However, a relatively diversified fish
fauna composed of more than 159 species has
been recorded around Ilha do Frade (HELMER
and PERRONE, 1991; ARAUJO et al., 2008).
The climate of the region is tropical hot and
humid, with an average annual temperature of
about 24ºC. Two seasons may be clearly identified
over the year: one relatively hot (average monthly
temperature from 24.8 to 26.8 ºC) and rainy, which
extends from November to April; and one
relatively cold (average monthly temperature
from 21.5 to 24.3 ºC) and dry, which extends from
May to October (Figure 2).
Relative effect of seasonality and body size on the diet of juvenile Lutjanus synagris…
21
Figure 1. Location of the sampling site (smaller arrow) on the coast of Ilha do Frade, Vitória, Espírito Santo
state, Brazil.
Figure 2. Monthly precipitation (mm) and average temperature (ºC) between May 2004 and April 2005 in
the region of Ilha do Frade. The precipitation values are presented for the period of study (2004–2005) and
for the ten years immediately previous to this period (1994–2003). The continuous horizontal lines represent
the annual averages. Source: INCAPER (meteorological station of Vitória, Espírito Santo state, at 20º30’00”S
and 40º31’07”W; altitude: 36 m).
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
22
ROSA et al.
Sampling methods
Monthly collections were carried out during
the daytime from May 2004 to April 2005, using a
beach seine (6 × 2 m, with a mesh of 1 cm between
opposite knots). Every month, eight 10-meter
trawls lasting approximately five minutes each
were carried out at low tide. The trawls were
conducted parallel to the beach to a maximum
depth of 1.5 m. All of the captured fish were
immediately fixed in a 10% formaldehyde
solution to interrupt the digestive process. The
individuals of L. synagris were later identified
(following MENEZES and FIGUEIREDO, 1980),
counted, measured [total length (TL) in mm] and
had their stomachs removed for stomach content
analysis. The food items in each stomach were
spread on a Petri dish and identified under a
stereoscopic microscope (following BRUSCA and
BRUSCA, 2003; MENEZES and FIGUEIREDO,
1980) major item groups. After the removal of
excess humidity using paper towels, the number
of individuals and the weight (± 0.001 g precision)
were recorded for each taxon identified in the
stomach contents. The length of the largest prey
found in each stomach was measured using a
caliper (± 0.01 mm precision), considering the
longer axis.
Data analysis
The percent frequency of occurrence (%F), the
numeric proportion (%N) and the mass
proportion (%M) of the food items found in the
stomachs were used to describe the diet of the
species (HYSLOP, 1980). These values were
combined to identify the main food items through
the Index of Relative Importance (IRI) proposed
by PINKAS et al. (1971): IRI = (%N + %M) × %F.
The values of this index were expressed in
percentages using the following equation: %IRI =
(IRI/Ʃ IRI) × 100.
In order to analyze if seasonal and sizerelated changes in the diet of the species occurred,
the fishes captured in each season were grouped
into three TL classes with at least 20 individuals
each: ≤ 40 mm; 41–50 mm; ≥ 51 mm. The %IRI of
the prey was calculated separately for each size
class in each season.
A similarity matrix was constructed using the
Bray-Curtis coefficient based on the values of the
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
%IRI. This matrix was submitted to cluster
analysis applying the Unweighted Pair Group
Method with Arithmetic Mean (UPGMA). In
order to test if there were significant differences in
the diet composition between seasons and size
classes (independent variables), a permutational
multivariate analysis of variance was used
(PERMANOVA; ANDERSON et al., 2008), where
all of the prey were analyzed simultaneously
(dependent variables). This analysis was based on
a similarity matrix constructed using the BrayCurtis coefficient calculated from the %M data of
the prey in each stomach, after being log(%M + 1)transformed. The values of P were obtained using
5000 permutations.
Differences in the average length of the
largest food items between size classes were
investigated using a one-way ANOVA and the a
posteriori test of Tukey. Before these analyses, the
length data of the items were log transformed to
reach homogeneity of the variances (based on the
Levene test) and normality (based on the
Kolmogorov-Smirnov test), as suggested by
SOKAL and ROHLF (1981).
RESULTS
A total of 540 specimens of L. synagris were
caught during samplings, all with TL (range: 22 to
135 mm; mode = 30 mm; s2 = 12.3 mm) below the
minimum size at first maturity (i.e. TL = 187 mm,
sensu FREITAS et al., 2011a). Considering all of
the species caught (45), L. synagris was dominant
in number of individuals (20.1% of the total
catch) and occurred in all of the sampling
months, except June 2004 (cold/dry season).
Approximately 86% of the individuals of L. synagris
were captured in the hot/rainy season. About
78% belonged to the ≤ 40 mm class, 10% to the 41–
50 mm class and 11% to the ≥ 51 mm class.
Eighty-seven percent (n = 472) of the
specimens presented food items in their stomachs.
Five large groups of food items were found in the
stomach contents: Teleostei, Crustacea, Mollusca,
Echinodermata and Annelida. The diet was
composed of thirteen taxa, with small crustaceans
as the most important food items in the diet of L.
synagris. Among them, Amphipoda was the main
item, followed by Isopoda, Mysidacea and
Penaeidae. The other items presented the %IRI
below 0.01 (Table 1).
Relative effect of seasonality and body size on the diet of juvenile Lutjanus synagris…
23
Table 1. Percentage values of frequency of occurrence (%F), numeric proportion (%N), mass proportion
(%M) and Index of Relative Importance (%IRI) of each food item identified in the stomachs of Lutjanus
synagris on the cost of Ilha do Frade, southeastern Brazil. N = 472 stomachs.
Items
TELEOSTEI
Syngnathidae
Teleostei
CRUSTACEA
Amphipoda
Brachyura
Euphausiacea
Isopoda
Mysidacea
Penaeidae
Larvae of Decapoda
Crustacea
MOLLUSCA
Gastropoda
ECHINODERMATA
Larvae of Echiuroidea
ANNELIDA
Polychaeta
The cluster analysis based on the values of
the %IRI differentiated the diet primarily as
regards seasonality, separating the samples
from each season into two groups, with a
similarity of approximately 46% (Figure 3).
Similarity between the size classes varied
substantially in each group (relatively higher in
%F
%N
%M
%IRI
0.42
1.06
0.03
0.09
0.33
3.51
< 0.01
< 0.01
64.41
1.69
1.06
46.82
17.58
11.65
1.69
2.54
67.47
0.10
0.08
15.77
13.39
2.58
0.26
0.16
38.57
1.51
0.28
24.07
14.67
15.36
0.13
1.40
81.20
< 0.01
< 0.01
13.81
4.40
0.56
< 0.01
< 0.01
0.21
0.01
0.01
< 0.01
0.21
0.01
0.01
< 0.01
0.64
0.04
0.13
< 0.01
the cold/dry season than in the hot/rainy
season). In the group comprising the hot/rainy
season, the ≥ 51 mm and the 41–50 mm classes
were more similar (78%), while in the cold/dry
season, the higher similarity occurred between
the 41–50 mm and the ≤ 40 mm classes (92%)
(Figure 3).
Figure 3. Dendrogram based on the Bray-Curtis similarity between the values of Index of Relative
Importance (%IRI) of the food items consumed by Lutjanus synagris from three size classes in the cold/dry
and hot/rainy seasons at a sandy beach in southeastern Brazil.
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
24
ROSA et al.
In the cold/dry season, Mysidacea were
relatively more important than in the hot/rainy
season for all size classes (the opposite for
Amphipoda). For individuals from the ≤ 40 mm
class, Mysidacea (%IRI = 86.1) were more
important in the cold/dry season and Amphipoda
(%IRI = 84.7) in the hot/rainy season. For fish
from the 41–50 mm class, a diet similar to the one
presented by the ≤ 40 mm class was observed in
the cold/dry season, with Mysidacea (%IRI = 76.6)
dominating; but in the hot/rainy season,
Amphipoda (%IRI = 45.5) and Isopoda (%IRI = 45.2)
were more important. A similar food pattern
was observed for individuals from the ≥ 51 mm
class in the cold/dry season, with Mysidacea
(%IRI = 86.3) dominating again. However, in
the hot/rainy season, in addition to Amphipoda
(%IRI = 39.8) and Isopoda (%IRI = 16.3),
Penaeidae (%IRI = 33.4) were important items.
In general, Isopoda and Penaeidae were
relatively more important to the larger size
classes (41–50 mm and ≥ 51 mm) and were
consumed, particularly, in the hot/rainy season
(Figure 4).
Figure 4. Histograms of seasonal variations in the Index of Relative Importance (%IRI) of food items
consumed by Lutjanus synagris from three size classes at a sandy beach in southeastern Brazil. Amp =
Amphipoda, Mys = Mysidacea, Iso = Isopoda, Pen = Penaeidae, Tel = Teleostei, Bra = Brachyura and Oth =
Others (e.g. unidentified crustaceans, larvae of Decapoda, Euphaseacea, Gastropoda, Echiuroidea and
Polychaeta).
Significant variations were found in the diet
of L. synagris between seasons (PERMANOVA:
F1, 466 = 24.15; P<0.001), but not between size
classes (PERMANOVA: F1, 466 = 1.54; P = 0.170).
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
There was, however, a significant interaction
between these two factors (PERMANOVA: F2, 466 =
4.48; P<0.001), indicating that differences between
seasons did not occur in all classes. In order to
Relative effect of seasonality and body size on the diet of juvenile Lutjanus synagris…
identify the source of this interaction, each class
was tested individually against the seasons. In
these analyses, only the ≥ 51 mm class did not vary
significantly (PERMANOVA: F1, 44 = 2.04; P = 0.101).
Were obtained lengths of 465 food items
(≤ 40 mm class, n = 371; 41–50 mm class, n = 47;
≥ 51 mm class, n = 47). The items measured were
25
Amphipoda, Isopoda and Mysidacea. The average
length of the captured items varied significantly
(P<0.05) between size classes, increasing
progressively with the TL of the fish. However,
the pairwise comparisons showed that these
differences were significant only between the
≤ 40 mm class and the other two (Figure 5).
Figure 5. Variation of the average length (± SE) of the larger food items consumed by Lutjanus synagris from
three size classes at a sandy beach in southeastern Brazil. The results of the ANOVA and the pairwise
comparisons using the Tukey test are also presented.
DISCUSSION
The diet composition of L. synagris changed,
firstly, as regards seasonality, indicating that
environmental factors may have influenced that
change. Seasonal variations in the diet were
also observed by DUARTE and GARCIA (1999)
for L. synagris, by MONTEIRO et al. (2009) for
Lutjanus jocu and by PIMENTEL and JOYEUX
(2010) for Lutjanus analis. These changes occur
mainly due to alterations in the availability of the
items, which are associated with reproductive
pulses and physical-chemical alterations in
aquatic ecosystems (PLATELL et al., 1997). In this
study, the season with higher temperatures
seems to be related to the substantial increase
in consumption of Amphipoda by juveniles of
L. synagris. DUBIASKI-SILVA and MASUNARI
(1995) mention that high temperatures should
favor the abundance and reproduction of this
small crustacean. Studies suggest that factors such
as temperature do not seem to influence the
distribution and abundance of Mysidacea (RAPPÉ
et al., 2011). However, the pluviometric historical
of last ten years (i.e. that preceded the study
period, see Figure 2) showed low rainfall in the
cold/dry season. Rainfalls influence the volume of
freshwater discharge on coastal ecosystems and
may increase or decrease the fluvial input at these
sites, interfering with the salinity and influencing
the biota (GONZÁLEZ-ORTEGÓN and DRAKE,
2012). The dominance of Mysidacea in the diet of
L. synagris coincided with the period of lowest
rainfall. According to SUZUKI et al. (2009), the
distribution patterns of Mysidacea can be linked
to salinity tolerance of the respective life stages.
Although seasonality can influence the diet
composition, it also undergoes continuous
alterations during the growth of the fish, either in
composition or item size (UCHIKAWA and
KIDOKORO, 2014). It was possible to detect
significant differences related to the size of the
items consumed for each size class, suggesting a
food pattern in which the size of the item
increases proportionally with the size of the
juveniles of L. synagris. Furthermore, small
deviations in the amplitude of items size (0.5 to
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
26
ROSA et al.
1.0 mm) indicate a strong choice of L. synagris for
items of specific sizes in the tree size classes.
Similar results were reported for the species by
FONSECA (2009) in Abrolhos bank, Bahia state.
According to PARRISH (1987), lutjanids have a
tendency for an increase in the rate of
consumption of larger items and a decrease in the
rate of consumption of smaller items, with the
increase in age. It is consistent with the optimal
foraging theory, which predicts the predators
consume preys with higher energetic value to
maximize their fitness (GERKING, 1994). Other
factors can also be important, including agespecific changes in the use of habitats (MARTINO
and HOUDE, 2012) like nursery-to-coral reef
migrations by snappers (MCMAHON et al., 2012).
The diet of L. synagris on Ilha do Frade is
consistent with their congeners and corroborates
studies carried out in other Brazilian states (e.g.
FONSECA, 2009; PIMENTEL and JOYEUX, 2010),
as well as in other countries in Central and North
America (e.g. DUARTE and GARCIA, 1999;
FRANKS and VANDERKOOY, 2000). Lutjanid
juveniles feed mainly on benthic and planktonic
invertebrates, while adults may invest mainly in
fish or in benthic invertebrates (DUARTE and
GARCIA, 1999). The use of crustaceans by fish of
the genus Lutjanus has been documented for a
number of species, such as L. analis (FREITAS
et al., 2011b), L. jocu (MONTEIRO et al., 2009),
Lutjanus argentiventris and Lutjanus guttatus
(ORTEGA et al., 2010), Lutjanus griseus (YEAGER
et al., 2014).
The relevance of peracarid crustaceans in the
diet composition of juvenile lane snappers is
associated with the opportunistic feeding habits of
the species and the environment investigated. In
intertidal and subtidal rock shores, peracarids,
manly Amphipoda, Isopoda and Mysidacea, have
an important trophic role as food for juveniles and
adults of several commercially important fish
species (HERRERA et al., 2014). Additionally, the
accumulation of macroalgae such as Sargassum
and Ulva on the sandy beach of Ilha do Frade
contributes to an increase in the availability of
items such as Amphipoda, which, according to
several studies (DUARTE et al., 2010; POORE and
GALLAGHER, 2013), are predominant in this
type of substrate. Nevertheless, the results
indicate that peracarids were important
Bol. Inst. Pesca, São Paulo, 41(1): 19 – 29, 2015
component in the environment studied playing a
key role in the food web as food items, thereby
contributing to the nursery function that Ilha do
Frade sandy beach provide for many fish species.
Although the result of the PERMANOVA for
size classes did not show significant alterations in
the relative biomass of the items, a significant
interaction between seasonality and body size was
observed, indicating that the differences between
the seasons did not occur in every class. Notably,
there was a greater availability of items in the
hot/rainy
period.
However,
possible
morphological restrictions present in the smaller
fish (≤ 40 mm) seemed to have made the capture
of Isopoda and Penaeidae (larger items) more
difficult. In this way, the factors availability and
morphology, combined to the recognized
opportunist character of L. synagris, may explain
the differentiated use of the food items among the
juveniles in the studied environment. Within a
species, differences in the diet composition at
different
life-history
stages
or
between
populations can be a result of morphological
differences (DAVIS et al., 2012). Such differences
have important consequences in recruitment
because they reduce the intensity of competitive
interactions (BRANSKI and DORN, 2013;
DUKOWSKA et al., 2013).
CONCLUSIONS
The results indicate that L. synagris exhibits a
carnivore-opportunist food habit, shifting their
diet based on the seasonality. Increased
consumption of Amphipoda shows the diet
flexibility of juveniles and suggests greater
availability of this small crustacean during the
hot/rainy season in the Ilha do Frade.
Furthermore, the results suggest that juvenile lane
snapper are of great importance in regulating
epifaunal populations. The information that has
been obtained highlights the important functional
role of L. synagris in the trophic dynamics of the
community
inhabiting
the
sandy
beach
investigated.
ACKNOWLEDGMENTS
The authors are grateful to Helena Ferraz
Ayres for her valuable suggestions and
contributions to the text, Leandro Vieira Ripoli
Relative effect of seasonality and body size on the diet of juvenile Lutjanus synagris…
for his help in field collections and Hélio Sá
Santos for his assistance in the identification of the
food items.
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