J. Appl. Ichthyol.
25 (2009), 335–342
 2009 The Authors
Journal compilation  2009 Blackwell Verlag, Berlin
ISSN 0175–8659
Size-related shifts in dietary composition of Centropomus parallelus (Perciformes:
Centropomidae) in an estuarine ecosystem of the southeastern coast of Brazil
By R. Feltrin Contente1,2, M. Freitas Stefanoni1,2 and O. B. F. Gadig1
1
Campus Experimental do Litoral Paulista, Universidade Estadual Paulista (UNESP), São Vicente, Brazil; 2Programa de
Pós-graduação em Zoologia, Universidade Federal do Paraná (UFPR), Curitiba, Brazil
Summary
Size-related and seasonal evaluation of the dietary composi-
tion of fat snook (Centropomus parallelus Poey 1860) in the
upper sector of an estuary of the southeastern coast of Brazil
were carried out based on stomach analyses of specimens
ranging from 40 to 170 mm standard length. Results reveal
that C. parallelus is a carnivorous species feeding mainly on
benthic crustaceans. Relatively high stomach replenishment
suggests that this environment is an important feeding ground
for fat snook juveniles. Multivariate analyses indicated that
predator size effect is significantly more important than
seasonal variation in determining dietary composition. Pred-
ator length was associated with increased consumption of
palaemonid shrimps (Macrobrachium spp.) and grapsid crabs,
and decreased foraging on tanaids (Kalliapseudes schubarti),
thus showing a preference shift from smaller to larger prey.
Predator length was also positively associated with an increase
in the stomach repletion index. Additionally, allometric
growth of both gape and head were consistently correlated
with this ontogenetic dietary transition, suggesting that such
changes might be related to an individualÕs ability to capture
and consume larger, more elusive prey. The digestive tube is
short and grows isometrically, which is in accordance with the
carnivorous habit of this estuarine fish and its maintenance
through ontogeny.
Introduction
Fat snook, Centropomus parallelus Poey 1860, is an important
predator fish in near-shore habitats of the tropical and
subtropical western Atlantic, occurring from Florida to the
southern coast of Brazil (Rivas, 1986; Orrel, 2002). This
medium-sized centropomid inhabits mostly oligohaline and
mesohaline habitats of estuaries (Rivas, 1986; Aliaume et al.,
1997; Itagaki, 2005). Along the southeastern coast of Brazil,
fat snook are traditionally exploited by commercial and
recreational fishing (Mendonça and Katsuragawa, 2001).
Furthermore, populations of fat snook have become increas-
ingly threatened by aquatic pollution and habitat loss, namely,
the deforestation of mangrove nursery areas. Due to the high
market value of this species, several studies have been carried
out in order to improve its culture (e.g. Alvarez-Lajonchère
et al., 2002). However, comprehensive accounts on its feeding
habits in southeastern Brazilian estuaries are scarce. Although
Della-Patrona (1984) indicated that juveniles are zooplanktiv-
orous and that adults are carcinophagous-piscivorous in an
estuary of the central coast of Rio de Janeiro, he did not
identified the principal consumed prey taxa or statistically
U.S. Copyright Clearance Centre Code Statement: 0175–8659/2009/2503–0335$15.00/0
Received: August 8, 2007
Accepted: June 20, 2008
doi: 10.1111/j.1439-0426.2008.01179.x
evaluate seasonal and ontogenetic dietary patterns. Another
study performed in the Cananéia estuarine system (São Paulo)
focused only on diet descriptions of fat snook larval and early
juvenile stages (i.e. 7–35 mm SL) (Itagaki, 2005). Comprehen-
sive information on seasonal and size-related dietary patterns
of a commercial species is important to achieve better fisheries
management and conservation plans (Blaber, 2000).
Morphological attributes related to the capture and intake
of prey have been successfully used to explain dietary patterns
in fishes (Norton, 1995; Wainwright and Richard, 1995; Ward-
Campbell et al., 2005). The predatorÕs gape size is an impor-
tant factor in setting the upper size limit for potential prey
(Wainwright and Richard, 1995), while the digestive tube
length associates consistently with predator (either carnivorous
or herbivorous) feeding habits and dietary physiology (Ger-
king, 1994). By potentially changing the spectra of size and
nutritional quality of ingested prey, size-related shifts of these
characters are important in differentiating ontogenetic prey-
use patterns that exert a direct impact on partitioning of the
food resource in size-structured fish populations (Werner and
Gilliam, 1984).
In order to provide further information on the trophic
ecology of C. parallelus, the aim of this study was to describe
its dietary composition regarding predator size and seasonal
effects at the upper sector of an estuary in southeastern Brazil.
Due to its high abundance, this species likely acts as an
important trophic component in low salinity estuarine reaches
(Itagaki, 2005). Additionally, morphological change of oral
and body structures was evaluated in light of the prey-use data.
Materials and methods
Study area, sampling and laboratory procedures
The study was performed in the upper estuarine sector of the
Perequê-Mirim ⁄ Itaguaré riverine system, situated on the
coastal plain of Bertioga, northern coast of São Paulo State
(Fig. 1). This coastal plain is covered by the Atlantic rainforest
with extensive mangrove forests fringing the river margins.
Several pools are formed along the margins due to erosion.
These macro-habitats are relatively deep (‡1.0 m), slow
flowing and structurally complex (containing wood debris,
mangrove roots and some marginal grass patches), constitut-
ing a typical habitat for snook prey that include swimming and
walking crustaceans and small fishes. Large groups of
prey-foraging snooks have often been observed in pools
(R. F. Contente, pers. obs.). From the upper estuarine reaches,
four pools were randomly selected for sampling (Fig. 1).
Fishery surveys were conducted monthly from November 2004
336 R. Feltrin Contente, M. Freitas Stefanoni and O. B. F. Gadig

Fig. 1. Map of the Perequê-Mi-

rim ⁄ Itaguaré riverine system, north-
ern coast of the São Paulo State,
southeastern Brazil. Locations of
pools selected for sampling (L1, L2,
L3 and L4) are also showed

to August 2005, except in March 2005. Temporal variations in
temperature and salinity were similar among selected pools
(Fig. 2; R. F. Contente, unpubl. data). Higher temperatures
and freshwater conditions (salinity oscillating around 0)
predominated from November to February during the warm-
rainy season (precipitation of 286 mm in the season peak)
when continental discharge is higher (total water volume ca.
5.0 · 107 m3; Santos, 1965; Anonymous, 2007). The opposite
situation occurs during the cold-dry season (April–August),
when salinity rises, oscillating around 7 (rainfall of 78 mm and
total water volume ca. 2.0 · 107 m3; Santos, 1965; Anony-
mous, 2007). Overall, the mean annual rainfall for the region is
2600 mm. The mean flow rate of the riverine system is ca.
6 m3 s)1 (Anonymous, 2007). For the region, the intertidal

20
18
16
14
12
Salinity

10
8
6
4
2
0
30
Pool 1
28
Temperature (ºC)

Pool 2
26 Pool 3
Pool 4
24
22
20
18
16 Fig. 2. Seasonal water temperature
Nov/04 Dec/04 Jan/05 Feb/05 Apr/05 May/05 Jun/05 Jul/05 Aug/05 and salinity profiles from sampling
pools along the upper estuarine sector
Rainy season Dry season of the Perequê-Mirim ⁄ Itaguaré river-
Month and season ine system
Size-related shifts in dietary composition of Centropomus parallelus
range during neap and spring tides is near 0.5 and 1.0 m,
respectively. Low variations in the tide regime occur season-
ally.
Samples were obtained during daylight ebb tides during
neap periods, as there is evidence of increased feeding among
snooks during tidal currents (Fore and Schmidt, 1973). Fish
were captured by hook (ref. Mustad 37140, nº 8 and 14) and
line (a suitable method to capture snooks; Fore and Schmidt,
1973), using live or dead non-indigenous bait to avoid bias in
the stomach content analysis. Fish were stored in ice for
subsequent laboratory examination. Animals for each pool
were grouped for each fishery survey.
In the laboratory, (i) specimens were identified following
Rivas (1986) and Orrel (2002), (ii) morphological attributes
(see below) and wet weight (pt – nearest 0.01g) were taken from
each individual, (iii) stomachs were removed and weighed full
and empty (nearest 0.0001 g) to determine content weight (pe),
(iv) stomach fullness was estimated visually on a scale of 0
(empty) to 1 (fully distended), and (v) food contents were
preserved in 70% ethanol. Stomach contents were sorted to the
lowest possible taxonomic category. Volumetric contribution
of prey items (vi) was estimated by spreading all contents in a
counting cell chamber with uniform depth and calculating the
proportional area of prey i in relation to total prey area.
Statistical analysis of diet
It is assumed that dietary diversity increases with the increas-
ing number of stomachs analysed. Thus, the number of
stomachs in which the asymptote of a cumulative prey-type
curve is reached, based on random orders of non-empty
stomachs, can be considered as an adequate sample size for
describing the diet of a species. This procedure was employed
based on 999 random orders to assess sample size adequacy.
Fat snook diet was described by frequency of occurrence
(%FOi) and by mean percentage volumetric contribution
(%Vi), as follows (Hyslop, 1980; Tirasin and Jørgensen, 1999):
! "
X n
%FOi ¼ 100n 1
Nij
j¼1
and
! !1
X
n k X
X n
%Vi ¼ 100 vij vij
j¼1 i¼1 j¼1
th
where i = any prey item; j = j stomach; n = number of
non-empty stomachs analysed; Nij = stomach j in which prey
occurred i; and k = total number of prey items found in n
stomachs. Fishing surveys yielded a total sample composed
largely of juvenile specimens. Individuals captured throughout
the study period at all catch locations were pooled and then
sorted into four size-classes (Cm, in SL) to assess size-related
changes in diet: C1 = 40–60 mm; C2 = 60.1–80 mm;
C3 = 80.1–100 mm; and C4 = 100.1–170 mm. In order to
assess seasonal dietary changes, stomachs were pooled accord-
ing to the season of catch (i.e. rainy or dry season). Two
approaches were used to identify size-related trophic units
(Gy), based on volumetric data: (i) a group average hierarchical
cluster analysis using the Bray–Curtis similarity (BSA), by
considering a Gy clusters of size classes situated above cut level,
which was defined by means of the average of all groups within
the distance matrix (Arancibia, 1987), and (ii) discriminant
analysis using WilksÕ criterion (k), according to Legendre and
337
Legendre (1983) and Castillo-Rivera et al. (2000). The k-values
show the dietary overlap, which range from 1 (no dispersion
between trophic groups, i.e. total similarity of trophic spectra)
to 0 (maximum dispersion of centroids, i.e. total distinction
between trophic spectra), and identifies the most important
prey item in differentiating diets of the size classes. The v2
statistic was employed to evaluate the significance of k. When
there was no significant difference between comparable diets of
the size classes, data were pooled forming a Gy. Prior to both
analyses, data were square-root transformed. The concordance
between BSA and k-values was assessed by the Spearman rank
correlation coefficient rsp (Zar, 1996). Finally, seasonal shift in
the dietary composition was evaluated by k.
The trophic niche breadth for each Gy and season was
calculated using the normalised version of the LevinÕs index
(Ba) (Krebs, 1989; Raymundo-Hiuzar et al., 2005) such that:
2 !1 3
Xn
Ba ¼ 4 2
pj 15ðk  1Þ1
j¼1
where
! !1
X
n k X
X n
pj ¼ Nij Nij
j¼1 i¼1 j¼1
pj gives an appropriated distribution function for this index,
ensuring statistical independence of the prey consumed
(Raymundo-Hiuzar et al., 2005). Ba values ranged from 0
(feeding specialization) to 1 (feeding generalization) (Krebs,
1989). To assess the uncertainty associated with Vi and Ba,
non-parametric confidence intervals (CI95%) were calculated
using the bootstrap method (based on re-sampling 5000 times),
considering each stomach to be a sampling unit (Tirasin and
Jørgensen, 1999).
The feeding intensity for each size class and season was
quantified by: (i) the relative fullness (RF), the mean of
stomach fullness values, and (ii) the repletion index (RI):
#
Xn
RI ¼ 100pe ðpt Þ1 n1
u¼1
Values of size-related and seasonal RI and RF were compared
using the Kruskal–Wallis (KW) test (Zar, 1996).
Morphometric and morphological-dietary relationship analysis
The morphological attributes, relevant to prey capture perfor-
mance and fish diet (Gatz, 1979; Wainwright and Richard,
1995; Ward-Campbell et al., 2005), were: standard length (SL;
nearest 0.1 mm), head length (HL), digestive tube length
(DTL), and mouth width (MW) and height (MH; nearest
0.05 mm) in order to estimate the mouth area (MA), such as
MA = 0.25p(MWMH). The logged form of the allometric
growth model, ln Y = ln a + b (ln SL), was used to evaluate
variations in the morphology of feeding structures with body
length. Potential allometric growth was identified using a t-test
to compare observed (b) and expected (bÕ) slope values, and the
significance of the model was assessed using a one-way ANOVA
(Zar, 1996).
The relationship between the most important prey items (i.e.
with Vi > 10% in the overall diet) and the morphological
attributes was described with the aid of a canonical correlation
analysis (CCA) (Miles and Ricklefs, 1984). Squared canonical
correlation coefficient (R2), which indicates the proportion of
338
common variation between morphological attributes and diet
components roots, was tested by running a v2 test, considering
as a null hypothesis that the lth canonical correlation and all
that follow are zero (Miles and Ricklefs,p1984).
ffi Prior to CCA,
morphological and dietary data were ½4x transformed to
provide an approximation of normal distribution.
Results
Diet composition
The asymptote of the cumulative number of prey items was
reached at 70–80 inspected stomachs, thus indicating a
sufficient sample size to describe the diet of the fat snook.
The tanaid Kalliapseudes schubarti Mañe-Garzon volumetri-
cally decreased whereas palaemonid shrimps Macrobrachium
spp. (mainly Macrobrachium acanthurus Wiegmann) and
grapsid crabs [mainly adult forms of Sesarma rectum Randall
and Chasmagnathus granulata (Dana)] increased with the
increase in body size of C. parallelus (Fig. 3), each contributing
more than 10% in Vi for the overall diet (Table 1). Results of
multivariate analyses yielded a similar conclusion
(rsp = 0.771; P < 0.001) and both identified two size-related
trophic units (Gy) (G1 = C1 and G2 = C2, C3 and C4), since
BSA isolated C1 from other size classes (Fig. 4) and discrimi-
nant analysis showed significant differences between C1 and
remaining size classes (Table 2). Considering k-values, the diet
of C1 was distinguished from that of C2 by K. schubarti (0.580),
followed by Macrobrachium spp. (0.501); between the diets
of C1 and C3 the principal discriminant components were
K. schubarti (0.835), followed by brachyuran megalopae
(0.786); between those of C1 and C4 were K. schubarti
(0.570), followed by grapsid crabs (0.566) and Macrobrachium
spp. (0.517). Regarding Gy diet, the smallest fat snooks (G1)
fed mainly on K. schubarti and secondly on Macrobrachium
spp., both in terms of Vi and FOi (Table 1). In contrast to G1,
the former prey item decreased 29% in FOi and approximately
25% in Vi, and the latter prey item increased about 10% in FOi
and ca. 18% in Vi in the diet of G2. In this group significantly
more grapsids were consumed (Table 1).
Prey items that contributed <10% volumetrically for the
overall diet were considered minor components, although for
42 48 35 45
70
60
50
40 Macrobrachium spp.
V (%)
K. schubarti
Grapsid crabs
30 Other prey
20
10
0 items. Stomach data from all study
C1 C2 C3 C4
Size class
R. Feltrin Contente, M. Freitas Stefanoni and O. B. F. Gadig
some FOi these were substantial (for example, sphaeromatids
had FOi = ca. 20%, Table 1). Minor components were
chironomid larvae, brachyuran megalopae, sphaeromatid
isopods and teleosts (Table 1). Sediment and vegetal frag-
ments found in stomach contents probably were ingested
accidentally.
There was no significant difference in dietary composition
between seasons, as revealed by discriminant analysis
(Table 2). Principal prey items exhibited similar volumetric
proportions between seasons (Table 1). As a whole, CI-values
for Vi were narrower in G2 than in G1, probably because the
former had a larger number of non-empty stomachs than the
latter (Table 1).
Trophic niche breadth
Mean values of trophic niche breadth (Ba), for size classes as
well as seasons, were larger than 0.5 (Figs 5 and 6), suggesting
a slight trend towards feeding generalization. The highest
feeding generalization occurred in C2, while the smallest was in
C4. No differences of trophic niche breadth were apparent with
regard to predator size or season because there was a
pronounced overlap among CI-values (Figs 5 and 6).
Feeding intensity
The total number of empty stomachs was relatively low (17%).
Size-related and seasonal variations of repletion index (RI) and
relative fullness (RF) are shown in Figs 5 and 6, respectively.
The KW test detected size-dependent differences in RI
(P < 0.05) except between C1 and C2. Non-significant sea-
sonal differences of RI (P > 0.05) were observed. Size-related
and seasonal RF values were larger or equal to 0.5. No size or
seasonal trends were detected for RF (P > 0.05).
Allometric growth and relationships between diet and morphology
All allometric models were positive and significant (ANOVA,
F1.204, P < 0.001; Fig. 7). Positively allometric growth (i.e.
b statistically >bÕ) occurred only for HL (HL t = 3.798,
P < 0.001), whereas the remaining morphological attributes
Fig. 3. Mean percentage volumetric
contributions (%Vi) of the main
discriminant prey items among size
class diets of Centropomus parallelus.
Size classes (Cm, mm): C1 = 40–60;
C2 = 60.1–80; C3 = 80.1–100 and
C4 = 100.1–170. Bars = confidence
intervals (CI95%). n – sample of each
size class is given above each bar.
ÔOther preyÕ include minor dietary
period from all catch locations were
sorted into the mentioned size classes
Size-related shifts in dietary composition of Centropomus parallelus 339

Table 1
Overall seasonal and size-related trophic unit dietary composition of Centropomus parallelus

G1 G2 Wet season Dry season Overall diet

FOi Vi FOi Vi FOi Vi FOi Vi FOi Vi

Prey items (%) (%) CI95% (%) (%) CI95% (%) (%) CI95% (%) (%) CI95% (%) (%) CI95%

Decapoda 47.4 35.8 (21.4–47.6) 70.8 64.4 (57.0–72.7) 60.6 44.5 (34.5–52.2) 63.8 55.8 (37.9–69.0) 66.9 50.1 (42.4–56.5)
Macrobrachium spp. 42.1 28.0 (16.7–38.1) 52.1 46.1 (38.3–53.1) 50.7 40.0 (30.0–47.8) 46.8 34.1 (23.4–42.9) 52.3 37.1 (30.0–43.1)
Grapsid crabs 5.3 2.1 (0.0–4.2) 20.8 18.2 (11.7–24.2) 12.7 4.5 (1.1–7.9) 23.4 15.8 (7.8–22.1) 18.4 10.1 (5.9–14.1)
Brachyuran megalopa 5.3 5.7 (0.0–9.5) 2.1 0.1 (0.0–3.6) 6.4 5.8 (1.3–9.1) 2.8 2.9 (0.6–4.8)
Peracarida 68.4 50.6 (35.7–61.9) 47.9 23.7 (15.6–29.0) 40.9 42.8 (33.3–50.0) 51.6 31.5 (20.8–40.3) 48.6 37.2 (30.0–43.0)
Kalliapseudes schubarti 63.2 43.4 (28.6–54.8) 34.2 18.7 (11.7–25.0) 33.8 37.5 (27.8–45.6) 36.2 24.6 (15.6–32.5) 36.7 31.1 (24.7–36.6)
Sphaeromatid isopods 21.1 7.0 (0.0–11.9) 20.8 4.8 (1.6–8.6) 15.5 5.2 (1.1–9.0) 23.4 6.6 (1.4–11.8) 20.2 5.9 (2.4–8.8)
Gammarid amphipods 5.3 0.2 (0.0–3.6) 2.1 0.3 (0.0–4.1) 1.4 0.1 (0.0–4.0) 4.3 0.3 (0.0–4.1) 2.8 0.2 (0.0–4.1)
Crustaceans not 31.6 5.5 (0.0–9.5) 19.8 5.7 (1.6–9.4) 21.1 7.4 (2.2–12.2) 21.3 5.1 (1.3–7.8) 20.2 5.6 (2.4–8.2)
identified
Chironomid larvae 21.1 8.1 (0.0–16.7) 11.5 2.1 (0.0–4.1) 9.9 5.3 (1.2–8.9) 10.6 4.9 (1.3–7.9) 11.9 5.1 (2.4–7.8)
Teleosts 4.2 4.1 (0.8–6.3) 2.8 1.4 (0.0–3.9) 4.3 2.6 (0.0–4.5) 3.7 2.0 (0.1–4.1)
Vegetal fragments 57.9 – 27.1 – 21.1 – 36.2 – 29.6 –
Sediment 10.5 8.3 – 5.6 – 8.5 – 7.4 –
Size 69.0 ± 9.0 99.3 ± 22.0 94.9 ± 21.4 85.0 ± 17.3 90.9 ± 23.5
(mean ± SD mm SL)
n-sample 42 128 78 92 170

Confidence intervals (CI95%) refer to %Vi values. –, not quantified. According to multivariate analysis, G1 = C1 and G2 = (C2, C3 and C4). Size
classes (Cm, mm): C1 = 40–60; C2 = 60.1–80; C3 = 80.1–100 and C4 = 100.1–170. For the ontogenetic diet description, stomach data from all
study periods from all catch locations were sorted into the mentioned size classes. For the seasonal diet description, stomach data were pooled by
caught season, irrespective of fish size and catch pools.
%FOi, frequency of occurrence; %Vi, mean percentage volumetric contribution; Gy, size-related trophic unit.

grew isometrically (i.e. b statistically = bÕ) in C. parallelus system. Also, our study suggests that this environment
(MA t = )1.780, DTL t = 0.646, P < 0.001). R2 of the first constitutes an important feeding ground for juvenile fat snook
two canonical roots (root-1 and root-2) were found to be due to high feeding activity, evidenced by the average large
significant (P < 0.001). Root-1 and root-2 explained 88.0 and replenishment of stomachs (generally >50% full) throughout
9.2% of the variance, respectively. Root-2, even though size classes and times of year, and the low overall occurrence of
significant, explains little of the data variance and therefore empty guts (17%). In fact, shallow estuarine environments
was not considered for the interpretation of results. Root-1 have been recognized as important nursery habitats for this
indicated considerable relationship between morphological species (Aliaume et al., 1997; Itagaki, 2005) and other
and dietary roots (R2 = 0.92; Fig. 8), suggesting a strong centropomids (Fore and Schmidt, 1973; Gilmore et al., 1983;
effect of body size, head size, mouth area and, to a lesser Aliaume et al., 1997).
extent, gut size on consumption of Macrobrachium spp., Fat snook did not vary significantly in diet composition in
followed by grapsids, and on a concurrent reduction in seasonal terms, concentrating on the same pool of prey
consumption of K. schubarti (Table 3). throughout the study period. This does not support the
findings of Della-Patrona (1984), who reported that individ-
uals of similar size range exhibit some preference for shrimps
Discussion during summer and for fishes during winter in the Rio de
The present research reveals that C. parallelus is a carnivorous Janeiro coast. Despite lacking a seasonal evaluation, the study
species that feeds mainly on benthic crustaceans in the upper
estuarine sector of the Perequê-Mirim ⁄ Itaguaré riverine
Table 2
Discriminant analysis results comparing diets of Centropomus paral-
lelus among size classes and between seasons
C2
Pairwise WilksÕ
G2 comparisons lambda (k) d.f. v2 P
C3
C1 vs C2 0.697 8 21.73 0.017
C4 C3 vs C1 0.699 8 20.57 0.015
C4 vs C1 0.450 8 28.32 0.001
G1
C1 C2 vs C3 0.862 8 8.52 0.482
C2 vs C4 0.800 8 7.92 0.542
C3 vs C4 0.817 8 14.48 0.106
100 77.12 Rainy season 0.927 8 7.58 0.577
vs dry season
Bray-Curtis similarity
Fig. 4. Dendrogram comparing size class diets of Centropomus paral- Size classes (Cm, mm) (and n of stomachs containing food): C1 = 40–
lelus using Bray–Curtis similarity coefficient. Size classes (Cm, mm) 60 (42); C2 = 60.1–80 (48); C3 = 80.1–100 (35); and C4 = 100.1–170
(and n of stomachs containing food): C1 = 40–60 (42); C2 = 60.1–80 (45). For the ontogenetic analysis, stomach data from all study periods
(48); C3 = 80.1–100 (35); and C4 = 100.1–170 (45). Cut line defines from all catch locations were sorted into the mentioned size classes.
size-related trophic units (Gy). Stomach data from all study period For seasonal analysis, stomach data were pooled by caught season,
from all catch locations were sorted into the mentioned size classes irrespective of fish size and catch pools.
340 R. Feltrin Contente, M. Freitas Stefanoni and O. B. F. Gadig

42 48 35 45 78 92
1.0 1.0
C1C2C3C4
0.9 0.9
0.8 0.8
0.7 0.7
RF

RF
0.6 0.6
0.5 0.5
0.4 0.4
0.3 0.3

C1C2C3C4
1.4
1.2
1.3
1.2 1.1
1.1
RI

RI
1.0
1.0
0.9 0.9
0.8
0.8
0.7

0.75
0.70 0.7
0.65
0.60 0.6
Ba
Ba

0.55 0.5
0.50
0.45 0.4
0.40
0.35 0.3
C1 C2 C3 C4 Rainy Dry
Size class Season

Fig. 5. Size-related variations of trophic niche breadth (Ba), repletion
index (RI) and relative fullness (RF) of Centropomus parallelus. Size
classes (Cm, mm): C1 = 40–60; C2 = 60.1–80; C3 = 80.1–100; and
C4 = 100.1–170. For Ba, vertical bars refer to confidence intervals
(CI95%) and for RI and RF, standard error (±SE). Horizontal bars
below size class for RF and RI represent non-significant difference
(P > 0.05) according to Kruskal–Wallis test. n-sample of each size
class is given at the top of the figure. Stomach data from study period
from all pools were sorted into the mentioned size classes
of Tonini et al. (2007) performed in a lagoon system at
the northeastern Brazilian coast reported a piscivorous diet for
C. parallelus, again in contrast to our results. However, the
mentioned studies, as well as several others (Fore and Schmidt,
1973; Gilmore et al., 1983; Aliaume et al., 1997; Teixeira,
1997), have demonstrated that juveniles of Centropomus
species are opportunistic feeders, which exploit temporally
Fig. 6. Seasonal variations of trophic niche breadth (Ba), repletion
index (RI) and relative fullness (RF) of C. parallelus. For Ba, vertical
bars refer to confidence intervals (CI95%) and for RI and RF, standard
error (±SE). RF and RI did not vary seasonally (Kruskal–Wallis test;
P > 0.05). n-sample of each season is given at the top of the figure.
Stomach data were pooled by caught season, irrespective of fish size
and catch pools
and locally abundant prey. Our results also suggest this feeding
behavior for C. parallelus because K. schubarti and Macro-
brachium spp., the major consumed prey, are abundant elements
of the macrofauna at the study estuarine system (Leite et al.,
2003; Souza and Moulton, 2005). The lack of seasonal change in
the diet may probably be due to their constancy and abundance
throughout the year (R. F. Contente, pers. obs.; Leite et al.,
2003; Souza and Moulton, 2005).

ln(HL) = 1.052ln(SL) - 1.266, r2 = 0.94 (a SE ± 0.089, b SE ± 0.019)

ln(DTL) = 1.041ln(SL) - 0.864, r2 = 0.69 (a SE ± 0.239, b SE ± 0.053)
ln(MA) = 1.835ln(SL) - 3.152, r2 = 0.92 (a SE ± 0.187, b SE ± 0.042)
7.0
6.5
ln(HL)mm, ln(DTL)mm and

6.0
5.5
ln(MA)mm2

5.0
4.5
Fig. 7. Allometric growth models for
4.0 head (HL), digestive tube (DTL) and
mouth area (MA) of Centropomus pa-
3.5 HL rallelus. Slopes (b), linear coefficients
MA
3.0 (a) and respective standard errors
DTL
(±SE) are given. Morphological mea-
2.5 surements obtained from individuals
3.6 3.8 4.0 4.2 4.4 4.6 4.8 5.0 5.2 5.4
caught throughout the study period at
ln(SL)mm all catch locations. Total n = 204
Size-related shifts in dietary composition of Centropomus parallelus
Canonical scores of diet
components
Fig. 8. First root canonical correla-
tion relating morphological and
dietary components of Centropomus
parallelus. Chi-square test indicates
that this root is statistically significant.
Morphological and dietary data
obtained from individuals caught
throughout the study period at all
catch locations. Total n = 204
Table 3
First root canonical coefficients from canonical correlation analysis,
based on morphological and dietary data of Centropomus parallelus
Morphological variables Coefficients
SL )0.99
HL )0.97
MA )0.93
DTL )0.75
Diet components
Macrobrachium spp. )0.97
Kalliapseudes schubarti 0.81
Grapsid crabs )0.73
Morphological and dietary data obtained from individuals caught
throughout the study period at all catch locations. Total n = 204.
SL, standard length; HL, head length; DTL, digestive tube length;
MA, mouth area.
Fat snook experienced a pronounced size-related shift in
dietary composition, as revealed by multivariate analysis.
Individuals 40–60 mm SL consumed mainly K. schubarti,
followed by Macrobrachium spp., while those between 61
and 170 mm SL changed their diet from K. schubarti to
mainly Macrobrachium spp. and, to a lesser extent, grapsids.
This transition meant a change of spectra size of swallowed
prey, i.e. from smaller (K. schubarti) to larger prey (Mac-
robrachium spp. and grapsids). Moreover, this dietary shift
coupled with an increase in the repletion index (RI) may be
related to the increased preference for large prey. Other
studies are consistent with this shift to large prey. For
similar-size fish, Della-Patrona (1984) found that individuals
<70 mm SL mainly prey upon crustacean larvae, and those
71–170 mm SL mainly prey on post-larval shrimps. In
Puerto Rico, fat snook shift from zooplankton to post-larval
shrimps and fishes at 30 mm SL (Aliaume et al., 1997).
Positive correlation between fish size and consumed prey size
has also been reported for other Centropomus species (Fore
and Schmidt, 1973; Gilmore et al., 1983; Aliaume et al.,
1997; Teixeira, 1997). Optimization of energy gained can
explain this ontogenetic dietary change. Macrobrachium
shrimp is a highly digestible food item with high energetic
content (Lemos et al., 2006), probably supporting more
effectively the particularly elevated metabolic demands of
fast-growing fat snook juveniles (Lemos et al., 2006) com-
pared to tanaids. This predator–prey relationship, therefore,
appears to be in accordance with the optimal foraging
theory, which states a maximum energetic return when a
predator selects and feeds on a particular prey type
(Gerking, 1994).
341
4
Root 1: χ 2 = 338.67; d.f. = 12; P < 0.001
3 R2 = 0.92
2
1
0
–1
–2
–1.5 –1.0 –0.5 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5
Canonical scores of morphological variables
Fat snooks of 61–170 mm SL showed high overlap of
their trophic niches (Table 2). This overlap, however, may
be reduced by the trend towards the generalized and
opportunist feeding behavior (Fig. 5) or due to the prey
availability not being a limiting factor for these sizes.
Further studies are needed to evaluate this intra-specific
trophic relationship by considering prey abundance and
dynamic patterns. Moreover, for a complete knowledge of
the functional role of C. parallelus in the structure and
dynamic of estuarine trophic webs, future studies should
focus on the description of ontogenetic and temporal
patterns beyond the limits examined in this study.
Relationships between shape and diet
Large ram-feeding percoid fishes generally feed on larger
and ⁄ or more elusive prey; this is partially related to the
increase in gape and head sizes (Gatz, 1979; Norton, 1995;
Wainwright and Richard, 1995). Wainwright et al. (2006)
observed experimentally that the larger the volume and area of
cross-section (approximately mouth area) of the oral cavity of
Centropomus undecimalis, the larger the volume of water
displaced into the mouth during prey capture. These dimen-
sional modifications improve the ability of this snook to
capture faster, larger prey as well as increase its reaction
distance that further aids in catching more elusive prey
(Norton, 1995; Wainwright et al., 2006). Assuming similarities
in gross morphology and feeding behavior with C. undecimalis,
our findings can be explained by this morphological ontoge-
netic process, since C. parallelus, a ram-feeder predator
(Luczkovich et al., 1995), exhibited a dietary transition from
smaller to larger prey, which consistently correlated with
increasing body, head and mouth area dimensions, as evi-
denced by the first canonical root (Table 3; Fig. 8). Moreover,
palaemonid shrimps are considerably more elusive than
tanaids, as the former is a fast swimmer (e.g. tail-flip) (Norton,
1995) and the latter is slow-moving and exhibits a tubicolous
life (Leite et al., 2003). Although grapsids are less elusive than
palaemonids (Norton, 1995), they are relatively larger than
tanaids. In addition, head growth is positively allometric in the
juvenile phase examined, rending an improved ability to
capture larger prey (Gatz, 1979), probably by increasing
disproportionately the oral cavity volume.
The digestive tube length, relative to SL, is relatively low (ca.
half of SL; mean = 0.04 ± 0.945 SD) and grows isometri-
cally. These findings are consistent with the carnivorous habit
of C. parallelus, maintained through an ontogenetic progress.
Overall, a carnivorous fish possesses a gut shorter than its own
length (Ward-Campbell et al., 2005). Animal food items,
342
which have high protein contents, are processed efficiently in
shorter guts, whereas plant items and detritus, which have a
relatively poor protein content, require a greater digestion time
to absorb protein suitably in a longer digestive tube (Ward-
Campbell et al., 2005).
Acknowledgements
We thank Matthew T. Wilson, Augusto V. Flores and two
anonymous referees for considerably improving the manu-
script and E. Mumtaz Tirasin for providing us with a
bootstrap method software. We thank Célia Freitas and
Janete Contente for logistic support, CNPq for financial
support and IBAMA for the research license.
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AuthorÕs address: Riguel Feltrin Contente, Experimental do Litoral
Paulista, Universidade Estadual Paulista (UNESP),
São Vicente, Brazil.
E-mail: riguel.contente@gmail.com