E Fundam. Appl. Limnol. Vol.

183/2 (2013), 163–175Article

Stuttgart, September 2013

Trophic ecology of two benthivorous fishes in relation

to drift and benthos composition in a pristine Serra do
Mar stream (Rio de Janeiro, Brazil)

Carla Ferreira Rezende 1, *, Javier Lobón-Cerviá 2, Érica Pellegrini Caramaschi 3 and

Rosana Mazzoni 4

With 3 figures, 4 tables and 2 appendices

Abstract: The objective was to explore how and when prey availability in the form of drift and benthos are
exploited by two phylo-genetically distant but morphologically similar fish species namely Characidium vidali
and Pimelodella lateristriga, the only two benthivorous species co-occurring in a pristine Serra do Mar stream
(Southeastern Brazil). We simultaneously quantified drift, benthos and the food of the two fish species. Benthos was
quantified once a day whereas drift and fish diets were quantified over diel cycles (six times per day) in December
2006 and February, April, July and October 2007. The major components of drift and benthos were Simuliidae,
Chironomidae and Baetidae with an additional contribution of Elmidae to benthos. Drift and benthos components
showed little differences in size across samples with 2–4 mm long invertebrates representing ≥ 80% of the total.
Both drift rates and benthos densities were surprisingly low and showed practically no difference between day and
night (drift) across months. Pimelodella fed more actively during the night whereas Characidium fed more actively
during the day but the two species fed essentially on the dominant, small sized Simuliidae, Chironomidae and
Baetidae. However, Pimelodella showed a markedly broader feeding niche including a higher prey diversity (i.e.,
21–30 taxa across months relative to only 12–15 taxa fed upon by Characidium). Apparently, eco-morphological
constraints and species-specific foraging behaviors eventually induce species-specific feeding patterns and as a
consequence, feeding overlap and therefore potential competition between these two species even if benthos den-
sity was low, appeared weak all year round.

Key words: Trophic ecology, feeding patterns, diel rhythms, drift rates, benthos density, Serra do Mar, Mata
Atlântica.

Introduction Ross (1986) suggested that fish assemblages segregate

resources use across three major dimensions, namely
Resource partitioning has long been recognized to be habitat, trophic and temporal, and emphasized that par-
a major mechanism underlying species co-occurrence titioning levels may be stronger among species with
in fish assemblages (Gerking 1994). In his review, a higher degree of kinship than among phylo-geneti-
Authors’ addresses:
1  Universidade Federal do Ceará, Campus do Pici, Centro de Ciências Bloco 909, Laboratório de Ecologia de Rios do Semiárido,
Departamento de Biologia. Fortaleza 6021 Brasil
* Corresponding author: [email protected]
2  Museo Nacional de Ciencias Naturales (CSIC), C/ José Gutiérrez Abascal, 2. Madrid 28006 Spain; [email protected]

3  Laboratório de Ecologia de Peixes, Instituto de Biologia, Departamento de Ecologia, Universidade Federal do Rio de Janeiro,

21941-902 Brasil
4  Laboratório de Ecologia de Peixes, Instituto de Biologia Roberto Alcantara Gomes, Departamento de Ecologia, Universidade do

Estado do Rio de Janeiro, 20.550-013 Brasil

© 2013 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de

DOI: 10.1127/1863-9135/2013/0430 eschweizerbart_XXX

1863-9135/13/0430 $ 3.25
164 Carla Ferreira Rezende et al.

cally distant species. Matthews (1998) also suggested dynamics in two Neotropical fish species which are
that niche overlap among species may vary over time morphologically similar but phylo-genetically diver-
and that species tend to exploit the seasonally most gent namely, Characidium vidali Travassos 1967,
abundant resources. and Pimelodella lateristriga (Lichtenstein 1823), two
Stream-living fishes exhibit a bewildering diversity benthivorous species co-occurring in a pristine Serra
of feeding patterns (Gerking 1994). Studies on food do Mar stream (South-east Brazil). The objective was
partitioning among members of stream fish assem- to determine how and when these two species use the
blages have described a variety of interactions includ- available drift and benthos and further quantified lev-
ing instances in which segregation of food partitioning els of trophic niche overlap or feeding similarity over
among species is nearly absent (Moyle & Senanayake diel cycles across seasons.
1984), instances where morphology influences feed-
ing overlap among species more than phylo-genetic
relationships (Pouilly et al. 2003) and, at the opposite Study area, Material and methods
extreme, instances where gross morphological differ-
ences among species do not preclude high levels of The Mato Grosso Stream
niche overlap (Gabler & Amundsen 1999).
The role of prey availability for food partitioning The Serra do Mar is a 2000-km long mountainous
among co-occurring fish species has been documented range over the Brazilian coast from northern Bahia to
for fish populations and assemblages of Paleartic Rio Grande do Sul. It is drained by a complex web
streams (Dahl & Greenberg 1996, Rincón & Lobón- of streams flowing east, through Mata Atlântica for-
Cerviá 1999, Giroux et al. 2000). With practically no est, towards the Atlantic Ocean. The Mata Atlântica
exception (but see Uieda & Pinto 2011) studies on Ne- forest was the second largest Neotropical rainforest.
otropical fishes have failed to quantify the relative im- However, during recent decades all this region has
portance of prey availability for the feeding patterns been deforested to the extent that only 8% of its origi-
of fish assemblages’ members. Nevertheless, dietary nal distribution remains (Mazzoni & Lobón-Cerviá
studies on Neotropical fishes highlighted patterns 2000). Moreover, the fish fauna of this coastal region
similar to those elucidated for other geographical/cli- is characterized by high levels of endemic fish species
matic regions where feeding overlap may vary widely (Böhlke et al. 1978, Abell et al. 2008).
ranging from a total overlap in prey use by two or Pristine and well-preserved streams still remain on
more species (Knöppel 1970, Esteves & Galetti 1995, the upper and mid altitudes of the Serra do Mar. For
Esteves et al. 2008), to very low or negligible overlap the purpose of this study, we selected a pristine reach
(Zaret & Rand 1971, Power 1983, Herder & Freyhof of Mato Grosso, a stream located ≈70 km from Rio de
2006). The majority of those studies also emphasized Janeiro (22º55’S, 42º35’W). From its sources, at about
some level of segregation across the aforementioned 800m above sea level to its mouth at Saquarema la-
three major dimensions, trophic (Esteves & Lobón- goons, the stream drains a 30km2 drainage area over
Cerviá 2001, Deus & Petrere 2003, Novakowski et the northwestern area of Saquarema municipality (Rio
al. 2008), spatial (Esteves & Lobón-Cerviá 2001, de Janeiro). Stream depth ranged from 2.0 to 60.0 cm,
Benneman et al. 2005) and temporal (Lobón-Cerviá width from 0.9 to 3.3 cm and discharge from 0.1 to
& Benneman 2000) and further suggested that when 1.2 m s–1. The wet season lasts from October to March
overlapping over one of the three dimensions is high, and dry season from April to September. The Mato
then segregation along other dimensions is present Grosso fish assemblage is composed of five species, of
(Lobón-Cerviá & Benneman 2000, Esteves & Lobón- which Characidium vidali (henceforth Characidium)
Cerviá 2001). and Pimelodella lateristriga (henceforth Pimelodella)
Consequently, although feeding patterns have been comprised 95% of the total fish density at the study
reported for numerous Neotropical fishes, the relative site.
importance of prey availability for their feeding pat-
terns and the conditions under which those patterns Data collection and sampling scheme
between two or more species actually overlap and/or
the potential for inter-specific competition in Neotrop- For the purpose of this study, we selected a 250 m
ical stream fish assemblages are most unclear. In this long stream section located at the uppermost stream
context, we attempted to identify the relative impor- reaches. This site was fully covered by riparian vege-
tance of food availability as determinant of the trophic tation and abundant canopy with practically no illumi-

eschweizerbart_XXX
 Trophic ecology of two benthivorous fishes 165

nation on the stream water and a substratum composed each prey consumed, through a graduated plate with
of stones, gravel, pebbles and sand along successive 1 mm high and 1 mm width. In turn, the volume fre-
riffles, runs and pools. To prevent sampling-induced quency was calculated by the total volume of each sin-
perturbations, the study site was divided into sub- gle prey divided by the total volume of all prey. The
sections as follows: the uppermost 50 m of the study occurrence frequency was calculated by presence or
site was used to quantify benthos composition and the absence of each prey in each fish divided by the total
next 10m downstream to quantify drift composition. fishes sampled. Finally, benthos was quantified once a
The fishes were collected in a 150 m long sub-section day during daylight hours with a Surber (20 × 20 cm,
located 20 m below the drift site. 250 mm mesh size) with a 250 mm mesh screen at the
Sampling of drift, benthos and fishes were con- upper part of the frame to prevent the entry of drifting
ducted simultaneously on five occasions, in December organisms. Benthos samples were collected after com-
2006 and February, April, July and October 2007. The pletion of the 24-h drift and fish samples. We collected
drift and benthos data analyzed in this study is an ex- 30 stratified samples of benthos, 10 samples per type
tension (i.e., includes additional data on February) of of substratum (pebble, sand and leaf litter), with 150
the data set previously analyzed by Lobón-Cerviá et samples in total. Benthos densities are expressed as
al. (2012) in a comparison among Neotropcial streams. the number of individuals per m–2.
Physico-chemical characteristics and substrate com- After every collection, invertebrates were fixed in
position of the study sites are detailed in Lobón-Cerviá 90% alcohol. Drift and benthos samples were screened
et al. (2012) and are summarized below. in the lab and organisms were identified to the low-
Sampling hours in April, July and October in- est possible taxonomical level (Carvalho et al. 2001,
cluded daylight at 7:00, 11:00, 15:00 and 19:00 hours Melo 2003, Olifiers et al. 2004, Salles et al. 2004, Pas-
and night included 23:00 and 3:00 hours. In December sos et al. 2007), counted and measured (in mm). In
and February, day light included 6:00, 10:00, 14:00, addition, all organisms were grouped into sets of 1mm
18:00 hours and night at 22:00 and 2:00 hours. Due long. In the field, fishes were fixed in 10% formalin
to differences in day length, the sampling time of day and were transferred to 70% alcohol in the lab. Each
did not exactly coincide in summer and winter and individual fish was measured (standard length, SL,
the number of light and dark hours varied somewhat cm) and weighed (Tw, g). After dissection, full stom-
among seasons with a maximum of 14 light hours in achs contents were weighed (Ws, g) and subsequently
the summer (December-March) and a minimum of 11 prey were identified, counted and when possible meas-
light hours in winter (June-September). Water temper- ured (in mm).
atures varied little over the study period ranging from
26.4 °C in April to 30 °C in January. Data analysis
Drift and fishes were collected every four hours
within 24-hour diel cycles (i.e., six samples a day). Feeding diel activities for each single species over diel
Drift was quantified with two nets (20 × 20 cm, 1.2 m cycles and month are described with the Gut Fullness
long, 250 µm mesh and 250 ml glass collector) posi- Index (GFI) (Santos 1978), in the form:
tioned side by side, near the substratum at the end of a
GFI = 100 Ws Tw –1
run and remained operative for 30 minutes every four
hours, totaling 12 samples per day and 60 samples in where Ws is the weight of the stomach with contents
total. Drift rates are expressed as the number of indi- and Tw = weight of fish. To detect effects of the time
viduals per water volume filtered during the sampling of day, month and species on the Gut Fullness Index,
period (ind. m3 s–1) (Lobón-Cerviá et al. 2012). Simul- GFI, we applied a standard Analysis of the Variance
taneously, fishes were collected with electric fishing (ANOVA). Normality and homogeneity of variance
(Mazzoni & Lobón-Cerviá 2000). The 150 m long were tested with Kolmogorov-Smirnov and Levene
study site was divided into three 50 m sub-sections. tests.
For each single sample, each sub-section was fished Diet overlap or similarity among the feeding pat-
upstream until 15 individuals of Pimelodella and terns of the fish species and among those patterns and
Characidium were collected. We repeated this proce- the drift and benthos composition were examined with
dure at dawn and at dusk. Every sample yielded ≥90 the Horn’ Index (Djk, Horn 1966) for abundance data
fish per day. We followed Hyslop (1980) to describe (volume) where Nj = Σ Xji and Nk = Σ Xki (where X
fish diets in volumetric and occurrence frequencies. = prey item), and the overlapping or similarity Djk is
Volume was calculated by measuring the volume of given by:

eschweizerbart_XXX
166 Carla Ferreira Rezende et al.

Σi [(Xji + Xki) Ln (Xji + Xki)] – ΣiXji LnXji – ΣiXki LnXki

Djk =
(Nj + Nk) Ln (NjNk) – NjLn Nj – Nk LnNk

Moreover, patterns of prey size selection were ex- occurrence of these size classes in the drift, benthos
plored for the three most abundant size classes across and in the food of fishes.
samples and associations were examined among the

Table 1. Total and relative density of organisms in the benthos and drift quantified in December 2006 and February, April, July and
October 2007, in Mato Grosso, a pristine stream of the Serra do Mar (Rio de Janeiro, Southeastern Brazil). Except for February,
more detailed information on drift and benthos have been reported by Lobón-Cerviá et al. (2012).
Benthos Benthos Drift Drift
(ind. m–2) % (ind * m3 * s–1) %
Diptera
Ceratopogonidae 1.1 0.1
Chironomidae 194.8 23.1 0.4 12.0
Simuliidae 406.4 48.2 1.1 34.1
Empididae 2.1 0.2 0.02 0.5
Tipulidae 0.4 0.1 0.01 0.1
Simuliidae Pupae 0.6 0.1
Dixidae 0.2 0.02 0.03 1.2
Psychodidae 0.04 0.01 0.02 0.5
Ephemeroptera
Baetidae 75.8 9.0 0.9 28.9
Leptohyphidae 8.7 1.0 0.2 5.2
Leptophlebiidae 21.8 2.6 0.1 2.0
Trichoptera
Philopotamidae 0.8 0.1 0.3 0.9
Leptoceridae 14.0 1.7 0.01 0.4
Hydropsychidae 12.4 1.5 0.05 1.8
Hydroptilidae 3.4 0.4 0.01 0.05
Helicopsychidae 0.6 0.1
Calamoceratidae 0.9 0.1 0.01
Hydrobiosidae 0.4 0.04 0.01 0.06
Glossosomatidae 0.04 0.0
Plecoptera
Gripopterygidae 3.4 0.4 0.03 1.1
Perlidae 12.1 1.4 0.01 0.2
Coleoptera
Elmidae larvae 50.3 6.0 0.1 4.0
Elmidae adult 27.1 3.2 0.2 5.1
Odonata
Anisoptera 2.1 0.3 0.01 0.05
Zygoptera 0.6 0.1 0.003 0.1
Lepidoptera
Pyralidae 0.4 0.1
Hymenoptera
Formicidae 0.02
Hemiptera
Hemiptera terrestrial 0.01
Veliidae 0.2 0.02 0.05 1.6
Decapoda
Palaemonidae (Macrobrachium spp.) 1.6 0.2 0.02 0.1
Trichodactylidae (Trichodactylus sp.) 0.6 0.1
Total density rate 843 100 3 100
Total of Individuals 9825 4352
Total number of samples 150 60

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 Trophic ecology of two benthivorous fishes 167

Results the drift. Simuliidae, Chirontomidae and Baetidae (es-

sentially Baetis) accounted for 80% of the total ben-
Size and abundance of drift and benthos thos composition. Concurrently, larvae and adults of
Elmidae also showed a relatively high contribution
In the drift, we collected 4352 individuals belonging
with 9% of the total benthos density, similar to that
to 24 families. Simuliidae and Chironomidae predom-
recorded in the drift (Table 1).
inated in the drift composition over diel cycles and
Comparisons among the size distributions of drift
seasons. Also Baetidae (genus Baetis) contributed, at
and benthos organisms highlighted great similarities.
least, 75% and Elmidae (larvae and adults) 9.1% to the
With the exception of several 7 to 10mm long organ-
total drift with all other taxa showing minor contribu-
isms mostly Crustacea which contributed < 6% to the
tions. Only a few, scattered terrestrial organisms were
recorded in the drift (Table 1). Overall, the drift rates benthos composition (Fig. 1), the size of > 80% of the
were extremely low (mean across samples = 0.11 ind. individuals in both drift and benthos fell in the range
m–3, maximum in July = 0.18 ind. m–3 and minimum 2 to 4 mm. Thus, < 15% of the drift were in the range
in April = 0.03 ind. m–3). Moreover, the drift showed 1–2 mm whereas the range 2–4 mm long organisms
no consistent difference between day and night or appeared over-represented in the size distributions of
among months. Given such similarity and for the sake both drift and benthos across months (Fig. 1).
of simplicity, drift rates for single taxa were pooled for
Fish feeding activity
day and night across months in Table 1.
A markedly higher number and diversity of inver- In total, we examined the stomach contents of 323
tebrates were recorded in the benthos with 9825 in- Characidium and 358 Pimelodella. A visual inspection
dividuals belonging to 32 families. However, benthos of the gut fullness index averaged across individuals
composition was also dominated by taxa dominant in for each single species plotted against time of day and

Fig. 1. Size frequency distributions of drift and benthos organisms and of the prey determined in the feeding patterns of Characi­
dium and Pimellodella in Mato Grosso, a pristine stream of the Serra do Mar (Rio de Janiero, Southeast Brazil).

eschweizerbart_XXX
168 Carla Ferreira Rezende et al.

Fig. 2. Mean values (with standard

deviations) of the Gut Fullness Index
(GFI) averaged across individuals
for each sampling month (indicated
by the figure) for Characidium and
Pimelodella in Mato Grosso, a pris-
tine Serra of the Mar stream (Rio de
Janeiro, Southeast Brazil). Dark lines
indicated night hours.

month revealed that in all months, the feeding activity during the night (Fig. 2). Consistent with this pattern,
of Characidium attained greatest values during the day an ANOVA for log-transformed GFI values revealed
whereas Pimelodella showed greater feeding intensity significant differences between the two species in the

eschweizerbart_XXX
 Trophic ecology of two benthivorous fishes 169

Table 2. Results of ANOVA for the effects of species (two), sampling month (five) and time of day (six) on the gut content index
(GFI) estimated for Pimelodella and Characidium in Mato Grosso, a pristine stream of the Serra do Mar (Rio de Janeiro, Brazil).
DF are the degrees of freedom; F – the F-value and p – the significance level.
Sample DF F p
Species 1 215 < 0.01
Month 4 5 < 0.01
Time of day 5 23 < 0.01
Species × Month 4 0.60 0.66
Species × Time of day 5 23 < 0.01
Month × Time of Day 20 0.80 0.71
Species × Month × Time of Day 40 2 < 0.01

Table 3. Estimates of pairwase Morisita overlap index between Pimelodella and Characidium over time of day and month in Mato
Grosso, a pristine stream of the Serra do Mar (Rio de Janeiro, Brazil).
Pimelodella/ Characidium
Months / Time of day 6:00/7:00 10:00/11:00 14:00/15:00 18:00/19:00 22:00/23:00 2:00/3:00
April 12 0 0 20 0 0
July 8 8 7 50 0 0
October 11 1.5 0 7 0 0
December 11 1 0 7 0 0
February 59 63 0 50 0 0

GFIs (p < 0.01) with higher values in Pimelodella was almost double (i.e., 40 taxa, Appendix 2) the num-
(mean GFI = 0.02) than in Characidium (mean GFI ber of taxa fed upon by Characidium. Yet, Chironomi-
= 0.01). The effects of time of day (p < 0.01), month dae and Simuliidae were major components all year
(p < 0.01) and the interaction species*time of day (p round though their relative contributions were lower
< 0.01) were highly significant but all other interac- than those recorded for Characidium. These dominant
tions including species*month (p = 0.6), month*time prey were also followed by a temporally variable, but
of day (p = 0.7) and species*month*time of day (p = substantial contribution of Baetidae added to minor
0.4) were not significant (Table 2). but yet significant contributions of Elmidae, Trichop-
tera (Fam. Leptoceridae, Hydropsychidae and Hy-
Fish feeding patterns droptilidae) and Lepidoptera (Fig. 3). Moreover, a few
Appendix 1 and 2 summarized the monthly feeding large-sized crustaceans (Palaemonidae and Trichodac-
patterns of the two fish species. Overall, there were no tylidae), scattered contributions of fish fragments and
obvious feeding shifts related to ontogenetic develop- scales and oligochaeta with an additional contribution
ment (i.e., fish size) in either species nor were differ- of terrestrial insects occurred in the Pimelodella diet
ences detected over the diel cycles. (Apendix 2).
The two species fed heavily on the most abundant Although dominant prey were the same in the two
and dominant aquatic insects. The Characidium diet species, niche breadth or prey diversity was substan-
showed definitively no terrestrial insects but fed upon tially higher in Pimelodella. In every month, the num-
24 prey taxa. In this, Simuliidae and Chironomidae ber of taxa in Pimelodella diet (range from 21 taxa in
showed the higher frequencies of occurrence all year December to 30 taxa in July) was almost double that
round with 100% contribution of Chironomidae in of Characidium (range from 12 taxa in December and
April, October and February and 55.9% and 67.6% in February to 15 taxa in April, July and October). An es-
July and December, respectively. These dominant prey timation of the feeding overlap between the two fishes
were followed by temporally variable contributions of with Horn index (in %) over diel cycles and months
Baetidae ranging from 11.5% in April to an extremely highlighted low levels all year round. Moreover, be-
high 92.7% in February (Appendix 1). cause Characidium does not feed during dark hours,
Unlike Characidum, the Pimelodella diet was the overlapping levels were practically zero during the
characterized by a markedly high diversity of prey that night across months (Table 3). All other values of this

eschweizerbart_XXX
170 Carla Ferreira Rezende et al.

Fig. 3. Month-to-month variation in the volumetric frequency of the dominant prey (Chironomidae, Simuliidae and Baetidae,
and other organisms) in the diet of Characidium and Pimelodella in Mato Grosso, a pristine Serra do Mar stream (Rio de Janiero,
Southeast Brazil).

index over the diel cycles indicated < 20% overlap. position between drift and benthos, the feeding pat-
The only exception occurred during the rainy Febru- terns of Characidium and Pimelodella were compared
ary when this index was surprisingly low at 15:00h with benthos composition only. These comparisons
(i.e., practically 0%) but reached 59%, 63% and 50% with the Horn index (in %) highlighted high values for
during light hours and at sunset (Table 3). Actually, the two species. For Characidium the mean similar-
the two time periods when overlapping maximized at ity ranged from 51% in April to 94% in December.
down (i.e., 6:00 h) and dusk (i.e., 18:00 h) occurred at For Pimelodella the similarity levels were still higher
the beginning and completion of the feeding activities ranging from 66% in February to 93% in April (Table
of the two species: nocturnal feeding by Pimellodella 4). These values indicate that during the dry season
and diurnal feeding by Characidium. (April and July), the similarity between the feeding
Given the extremely low drift rates recorded dur- patterns and benthos composition maximized in Pime­
ing the study period and the similarity in species com- lodella and minimized in Characidium whereas the

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 Trophic ecology of two benthivorous fishes 171

Table 4. Similarity index (%) between the feeding patterns of essentially composed of Simuliidae, Chironomidae
Characidium and Pimelodella and benthos composition for the and Baetidae, with an additional contribution of Elmi-
five study months (daily data pooled) in Mato Grosso, a pristine
dae to the benthos composition. That is, only small-
stream of the Serra do Mar (Rio de Janeiro, Brazil).
sized aquatic invertebrates in low abundance with a
Characidium Pimelodella remarkably low contribution of terrestrial insects pre-
April 51 93 dominate in the drift and benthos composition that in
July 58 80 turn, were quite similar in size and composition over
October 78 74 diel cycles and seasons. Thus, temporally persistent
December 94 72
small size organisms in the drift and benthos form the
February 71 66
bulk of the prey availability for the two studied benthi-
vorous fishes over time.
The relative contribution of the dominant organ-
opposite occurred during the rainy season (Table 4). isms to the feeding patterns of the two fishes (2–4 mm
Moreover, during the dry season the two fishes also long Simuliidae, Chironomidae and Baetidae) matches
maximized the diversity of prey fed upon relative to their relative contribution to the drift and benthos.
other months. Prey diversity minimized in the two Expectedly, such pattern may be indicative of high
species during the rainy season. feeding overlaps underlying inter-species competition
We further explored for the occurrence of size- particularly given the low benthos densities that typify
selective predation by Characidium and Pimelodella our pristine stream. However, over the diel cycles, not
on drift and benthos. Occasionally, Pimelodella fed only the feeding intensity differed between the two
on large-sized crustaceans and on a few terrestrial in- fishes but also Pimellodella fed on a much greater
sects but these were poorly represented in both drift diversity of prey. Pimellodella fed intensively during
and benthos. Once these large-sized organisms were the night and appears to be able to feed on practically
excluded, the three temporally persistent size classes all prey available in the benthos to the extent that, in
in the drift and benthos namely, 2, 3 and 4 mm long every month, the Pimellodella diet practically matches
invertebrates also predominated in the diet of the two the total benthos composition. In contrast, Characi­
fish species. Because of the small seasonal variation, dium fed during the day and appears to be much more
we pooled all data. In this pooled data set these size specialized on Chironomidae, Simuliidae and Baeti-
classes represented ≥ 80% of the drift and benthos dae. Additional prey in the Characidium diet appears
organisms and showed no significant differences be- scattered and very limited suggesting that a number of
tween the size of drift and benthos organisms and prey relatively abundant in the benthos such as Lep-
fish prey for either single or combined species (G- toceridae, Coleoptera, and Lepidoptera may not be
test = 8.7, p = 0.3, a comparison among size distri- available for Characidium.
bution in the drift, benthos and fish diets is included Critical differences in the functional behaviour
in Fig. 1). and eco-morphological attributes between Pime­
lodella and Characidium are apparently responsible
for the tendency to exhibit species-specific feed-
Discussion ing patterns. Both Pimelodella and Characidium
are small-sized, benthic feeders that typically share
Because no previous study has assessed the relative similar but not identical micro-habitats in high water
importance of prey availability for the feeding pat- current stream reaches associated with stone, pebble
terns of Neotropical stream-living fishes, this study and gravel substratum and to some extent also sand
explored the role of prey availability as components (Costa 1987, Uieda & Pinto 2011). Pimelodella pos-
of the drift and benthos for the feeding patterns of two sesses chemo-sensory barbels used to detect prey and
phylo-genetically divergent but morphologically simi- a slightly larger mouth and moreover, Pimelodella
lar benthivorous fishes co-occurring in a pristine Serra exhibits a markedly different foraging behavior in
do Mar stream. A striking result of this study was the which individuals patrol small stream areas during
remarkably low drift rates and benthos densities re- the night in continuous movements in search of prey.
corded all year round. Values recorded appeared to be These attributes might facilitate Pimelodella to ex-
among the lowest ever recorded across streams world- ploit a broader range of spatial meso-habitat result-
wide (Lobón-Cerviá et al. 2012). Also striking was the ing in higher diversity of prey. In contrast, the strictly
predominance of small sized (2–4 mm) dominant taxa sedentary Characidium is a visual hunter feeding dur-

eschweizerbart_XXX
172 Carla Ferreira Rezende et al.

ing the day whose foraging behavior rather resembles References

a sit-and-wait predator (Sazima 1986) that may con- Abell, R., Thieme, M.L., Revenga, C., Bryer, M., Kottelat, M.,
strain the use of the spatial meso-habitat resulting in Bogutskaya, B., Coad B, Mandrak N., Balderas, S.C., Bus-
a more specialized feeding tactic. Thus, the two fishes sing, W., Stiassny, M.L. J., Skelton, P., Allen, G.R., Unmack,
fed intensively on the most common prey namely, P., Naseka, A., Ng R., Sindorf, N., Robertson, J., Armijo, E.,
Chironomidae, Simuliidae and Baetidae. However, Higgins, J.V., Heibel, T.J., Wikramanayake, E., Olson, D.,
López, H.L., Reis, R.E., Lundberg, J.G., Pérez, M.H.S. &
species-specific eco-morphological and behavioral
Petry, P., 2008: Freshwater ecoregions of the world: a new
traits may not only prevent the simultaneous use of map of biogeographic units for freshwater biodiversity con-
the same prey over day and night but also enhances servation. – BioScience 58: 403–414.
Pimelodella ability to feed on a broader diversity of Aranha, J.M.R., Gomes, J.H.C. & Fogaca, F.N.O., 2000: Feed-
prey that eventually results in low levels of feeding ing of two sympatric species of Characidium, C. lanei and C.
pterostictum (Characidiinae) in a coastal stream of Atlantic
overlap over the seasons.
Forest (Southern Brazil). – Braz. Arch. Biol. Technol. 43:
Studies focused on Neotropical streams have em- 527–531.
phasized that Simuliidae, Chironomidae and Bae- Boyero, L. & Bosch, J., 2002: Spatial and temporal variation
tidae predominate in the drift and benthos (Ramírez of macroinvertebrate drift in two neotropical streams. – Bio-
& Pringle 1998, Ramírez & Pringle 2001, Boyero tropica 34: 567–574.
& Bosh 2002, March et al. 2003, Callisto & Goulart Böhlke, J.E., Weitzman, S.H. & Menezes, N.A., 1978: Estado
atual da sistemática dos peixes de água doce da América do
2005, Nessimian et al. 2008, Lobón-Cerviá et al 2012). Sul. – Acta Amazonica 8: 657–677.
Moreover, a high similarity between drift and benthos Callisto, M. & Goulart, M., 2005: Invertebrate drift along a lon-
composition appears the rule rather than the exception gitudinal gradient in a neotropical stream in Serra do Cipo
(Silveira et al. 2006, Nessimian et al. 2008). Independ- National Park, Brazil – Hydrobiologia 539: 47–56.
ent studies focused on the feeding patterns of fish – but Carvalho, A. L, Werneck-de-Carvalho, P. L. & Calil, E. R.,
2001: Description of the larvae of two species of Dasythemis
not on prey availability – also emphasized that Sim- Karsch, with a key to the genera of Libellulidae occurring in
uliidae, Chironomidae and Baetidae are major feed- the states of Rio de Janeiro and São Paulo, Brazil (Anisop-
ing resources for many members of fish assemblages tera). – Odonatologica 31: 23–33.
across Neotropical streams (Aranha et al. 2000, Uieda Costa, W.J.E.M., 1987: Feeding habits of a fish community in
& Pinto 2011). Thus, our study is the first to recon- a tropical coastal stream, Rio Mato Grosso, Brazil. – Stud.
Neotr. Faun. & Environ. 22: 145–153.
cile the results of the two sets of independent studies
Dahl, J. & Greenberg, L., 1996: Impact on stream benthic prey
by illustrating direct relationships between drift and by benthic vs. drift feeding predators: A meta-analysis. –
benthos composition and the feeding patterns of fish Oikos 77: 177–181.
and further emphasizes the importance of Simuliidae, Deus, C.P. & Petrere-Junior, M., 2003: Seasonal diet shifts on
Chironomidae and Baetidae in the drift and benthos seven fish species in an Atlantic Rainforest stream in south-
eastern Brazil. – Braz. J. Biol. 63: 579–588.
and in feeding patterns of benthivorous stream-living
Esteves, K. E. & Galleti, P. M., 1995: Food partitioning among
fishes. Further studies on other Neotropical streams some characids of a small Brazilian floodplain lake From the
with different invertebrates’ assemblages may help to Paraná river system – Environ. Biol. Fish. 42: 375–389.
address several questions that still remain unresolved Esteves, K. E. & Lóbon-Cerviá, J., 2001: Composition and
as, for example, whether or not the feeding patterns trophic structure of a fish community of a clear water Atlan-
elucidated in our study prevail in the high diversity tic rainforest stream in southeastern Brazil. – Environ. Biol.
Fish. 62: 429–440.
fish assemblages that typify Neotropical fish assem- Esteves, K. E., Lobo, P. A. V. & Faria, M. D. R., 2008: Trophic
blages. Alternatively, whether increased fish diversity structure of a fish community along environmental gradients
may cause inter-specific competition and subsequent of a subtropical river (Paraitinga River, Upper Tietê River
trophic niche shifts among members of these high di- Basin, Brazil). – Hydrobiologia 598: 373–387.
versity fish assemblages. Gabler, H.M. & Amundsen, P.A., 1999: Resource partitioning
between Siberian sculpin (Cottus poecilopus Heckel) and
Atlantic salmon parr (Salmo salar L.) in a sub-Arctic river,
Acknowledgements northern Norway. – Ecol. Freshw. Fish. 8: 201–208.
This study is part of the PhD Thesis of the senior author Gerking, S.D., 1994: Feeding Ecology of Fish. 1st ed. – Aca-
(CFR) financially supported by FAPERJ project n. demic Press, San Diego, California, pp. 1–416.
E-26/170.578/2006 and by several successive grants to CFR Giroux, F., Ovidio, M., Phillippart, J.C. & Baras, E., 2000: Re-
(CNPq 140928/2005-7; CAPES-PDEE 012008-01; Fundación lationship between the drift of macroinvertebrates and the
Carolina Spain), E.P.C. (CNPq 470587/2004-0) and R.M. activity of brown trout in a small stream. – J. Fish Biol. 56:
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ratório de Ecologia de Peixes/UERJ and UFRJ for help in the Herder, F. & Freyhof, J., 2006: Resource partitioning in a tropi-
field and laboratory. cal stream fish assemblage. – J. Fish Biol. 69: 571–589.

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Submitted: 11 October 2012; accepted: 31 July 2013.

eschweizerbart_XXX
174 Carla Ferreira Rezende et al.

Appendix 1. Monthly feeding patterns as expressed by volumetric and occurrence frequency of each prey in Characidium over
five study months in a pristine Serra do Mar stream (Mato Grosso, Saquarema, Southeast Brazil). N and Size are the number of
individuals analyzed and their corresponding sizes (in mm).
April July October December February
FN% FO% FV% FO% FV% FO% FV% FO% FV% FO%
N (size range) 86 (3.1–6.0) 77 (3.0–5.8) 59 (3.1–5.8) 50 (3.5–7.3) 51 (3.4–7.0)
Diptera
Ceratopogonidae 0.06 1.9
Chironomidae 14.8 100 3.6 55.9 0.3 100.0 23.0 67.6 0.1 100
Chironomidae Pupae
Simuliidae 66.6 100 67.9 45.8 72.4 43.3 53.8 44.1 3.6 100
Simuliidae Pupae 4.8 16.9 0.1 15 2.2 14.7
Empididae 0.2 5.8 0.2 3.4 0.04 13.3 0.8 8.8 0.03 22.0
Tipulidae 0.1 1.9
Diptera unidentified
Dixidae
Psychodidae
Muscidae
Ephemeroptera
Baetidae 0.8 11.5 6.4 27.1 0.6 45 12.1 64.7 0.3 92.7
Leptohyphidae 1.9 3.8 9.9 5.1 9.5 9.8
Leptophebiidae 0.4 1.9 19.8 13.3 0.2 2.9 0.2 12.2
Ephemeroptera Nymph
Baetidae Nymph 84.8 7.3
Trichoptera
Philopotamidae 0.2 2.9
Leptoceridae 0.4 1.9 0.3 6.8 0.1 15 0.005 9.8
Hydropsychidae 1.1 5.8 0.3 5.1 0.04 3.3 0.2 5.9
Hydroptilidae 0.1 3.8 0.5 16.9 5.4 30 0.4 11.8 0.0 22.0
Helicopsychidae
Plecoptera
Gripopterygidae 0.2 1.7
Perlidae
Coleoptera
Elmidae larvae 9.4 3.8 0.02 3.3
Elmidae adult 0.3 3.8 0.4 1.7 0.03 8.3
Psephenidae
Odonata
Anisoptera 0.2 2.9
Zygoptera
Odonata unidentified 1.7 1.7 0.0 1.7 0.01 2.4
Lepidoptera
Pyralidae 1.9 3.8 0.5 1.7 0.1 6.7 0.5 2.9 0.3 17.1
Neuroptera
Corydalidae
Terrestrial insects
Lepidoptera
Hymenoptera Formicidae 0.009 1.7 0.2 2.9
Hemiptera
Odonata
Diptera 0.04 1.7
Fragment of adult insects
Coleoptera
Blattodae
Decapoda
Palaemonidae (Macrobrachium spp.)
Trichodactylidae (Trichodactylus sp.)
Vegetal matter
Plant material
Seed
Others ítems
Scales 0.09 1.7
Fragment Crustacean
Aranae
Acarina 0.2 7.7
Molluscs Fish Oligochaeta
Insects larvae fragments 1.8 15.4 3.2 15.3 1.0 31.7 6.2 26.5 1.2 46.3
Total of items 15 15 15 12 12

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 Trophic ecology of two benthivorous fishes 175

Appendix 2. Monthly feeding patterns as expressed by volumetric and occurrence frequency of each prey in Pimelodella over five
study months in a pristine Serra do Mar stream (Mato Grosso, Saquarema, Southeast Brazil). N and Size are the number of individu-
als analyzed and their corresponding sizes (in mm).
April July October December February
FN% FO% FV% FO% FV% FO% FV% FO% FV% VO%
N (size range) 83 (5.3–11.8) 72 (5.3–11.0) 47 (5.3–10.8) 68 (5.3–10.9) 88 (5.4–11.0)
Diptera
Ceratopogonidae 0.1 6.3 0.1 4.6
Chironomidae 1.2 29.2 3.5 53.8 1.07 44.2 22.5 56.8 2.2 52.6
Chironomidae Pupae 0.02 2.1 1.5
Simuliidae 1.5 35.4 2.9 47.7 4.8 50 5.9 47.7 19.2 62.8
Simuliidae Pupae 0.07 2.1 1.5 21.5 0.5 11.5 0.8 13.6 0.6 11.5
Empididae 0.1 6.25 0.8 20 2.04 25 3.1 34.1 1.4 15.4
Tipulidae 0.8 3.8 0.1 2.3 0 0
Odonata unidentified
Dixidae
Psychodidae 0.01 2.08
Muscidae 0.5 1.5
Ephemeroptera
Baetidae 0.4 10.4 2.4 33.8 3.5 36.5 21.0 56.8 4.2 20.5
Leptohyphidae 0.5 3.8
Leptophebiidae 2.5 5.8
Ephemeroptera Nymph 0.01 2.1
Baetidae Nymph 0.3 4.6
Trichoptera
Philopotamidae 1.03 6.2 0.4 2.3 0.6 2.6
Leptoceridae 1.0 10.4 1.4 21.5 0.8 11.5 0.8 20.5 4.0 32.1
Hydropsychidae 0.8 7.7 0.9 9.6 0.1 4.5 2.6 7.7
Hydroptilidae 0.4 9.2 0.7 7.7 0.7 11.4 0.3 9.0
Helicopsychidae 0.1 2.1 1.1 7.7 0.7 5.8 0.3 2.3 0.8 2.6
Plecoptera
Gripopterygidae 0.6 4.6
Perlidae 0.2 3.1 0.3 1.9 0.2 1.3
Coleoptera
Elmidae larvae 0.1 4.2 0.4 7.7 0.8 21.2 2.2 18.2 1.3 17.9
Elmidae adult 0.1 4.2 0.3 9.2 0.3 3.8 0.2 2.3 0.3 6.4
Psephenidae 0.07 1.92
Odonata
Anisoptera 0.4 1.5 0.7 1.9 0.1 1.3
Zygoptera 0.9 1.9
Odonata unidentified 0.09 1.5 1.7 3.8
Lepidoptera
Pyralidae 2.15 12.50 2.65 12.31 0.82 7.69 0.05 2.27 1.60 7.69
Neuroptera
Corydalidae 1.2 2.1 5.4 1.5
Terrestrial insects
Lepidoptera 0.6 4.5
Hymenoptera Formicidae 0.01 2.1 0.1 3.1 0.2 1.9 1.5 3.8
Hemiptera 1.2 2.1
Odonata
Diptera 0.010 2.08
Fragment of adult insects 44.5 6.3 1.3 6.2 15.3 40.4 10.6 11.4 7.5 30.8
Coleoptera
Blattodae 4.7 1.5
Decapoda
Palaemonidae (Macrobrachium spp.)
Trichodactylidae (Trichodactylus sp.) 18 12.5 11.7 7.7 5.6 5.8 0.6 2.3 2.4 12.8
Vegetal matter
Plant material 4.1 12.5 0.3 1.5 0.4 1.9 0.3 3.8
Seed 0.0 2.1 0.0 1.5 1.3 1.9 0.1 2.6
Others items
Scales 4.7 39.6 3.1 27.7 16.0 44.2 0.3 11.4 0.8 6.4
Fragment Crustacean
Aranae 0.7 2.08
Acarina 0.01 2.08
Molluscs 0.19 3.08
Fish 6.92 2.08 4.7 2.3 2.9 1.3
Oligochaeta 0.6 2.1 8.0 9.6 3.8 2.3 5.1 2.6
Insects larvae fragments 11.1 43.8 51.8 70.8 31.1 53.8 21.1 18.2 37.8 75.6
Total of items 28 30 25 21 27

eschweizerbart_XXX