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Neotropical Ichthyology

versão impressa ISSN 1679-6225versão On-line ISSN 1982-0224

Neotrop. ichthyol. vol.13 no.2 Maringá abr./jun. 2015  Epub 02-Jun-2015 


Feeding ecology of immature Lithodoras dorsalis (Valenciennes, 1840) (Siluriformes: Doradidae) in a tidal environment, estuary of the rio Amazonas

Thiago Augusto Pedroso Barbosa1 

Ronaldo Borges Barthem2 

Luciano Fogaça de Assis Montag1 

1Laboratório de Ecologia e Conservação, Instituto de Ciências Biológicas, Universidade Federal do Pará, Rua Augusto Corrêa, 01, Guamá, Caixa Postal 479, 66075-110 Belém, Pará, Brazil. (TAPB) (corresponding author), (LFAM)

2Museu Paraense Emílio Goeldi, Departamento de Zoologia (CZO), Ictiologia, Av. Perimetral, 1901, Terra Firme, 66077-530 Belém, PA, Brazil.


Studies of feeding ecology are important for the evaluation of interactive processes in fish communities. This study evaluated the feeding ecology of Lithodoras dorsalis (Doradidae) from streams within the Amazon estuary delta (Brazil), a macro-tidal area, on different pluviometric periods. A total of 371 young specimens was collected during 12 months of sampling (July 2010 to June 2011). The species diet was composed of 28 food items analyzed by Repletion Index, Alimentary Index and Niche Breadth. Young L. dorsalis was classified as herbivore with a frugivory tendency due to the high importance of fruit and seeds in its diet. Food intake varied among sampled months, with the lowest intake being recorded during the rainy-dry season transition period, and the highest at the beginning of the dry season. The importance of food items and the composition of the diet were different throughout the year, probably due to the daily tides that allow fish to access new environments and the pluviometric periods. These results provide important data on the feeding ecology of Amazonian doradids. The study also emphasized the importance of allochthonous resources, derived from the riparian forest, which reinforces the importance of this habitat for the conservation of Neotropical freshwater fishes.

Key words: Diet; Doradidae; Feeding Index; Rock-bacu; Thorny catfish


O estudo da ecologia alimentar de peixes é uma abordagem consistente na avaliação dos processos interativos dentro das comunidades. Dessa forma, este trabalho teve como objetivo investigar a ecologia alimentar do bacu-pedra Lithodoras dorsalis em furos próximos no delta do estuário Amazônico (Brasil), uma área sobre influência de macro-marés, em diferentes períodos pluviométricos. Durante 12 meses de coletas (julho de 2010 a junho de 2011), foram coligidos 371 espécimes jovens, sendo que a dieta da espécie foi composta por 28 itens alimentares analisados pelos seguintes índices: Índice de Repleção Estomacal, Índice de Importância Alimentar e Amplitude de Nicho. Lithodoras dorsalis quando jovem foi classificada como herbívora com tendência à frugivoria, devido aos altos valores de importância de frutos e sementes em sua dieta. A intensidade de obtenção de alimento por L. dorsalis diferiu entre os meses de coleta, onde o final do período de transição chuva-estiagem e o início da estiagem foram os períodos de menor e maior atividade alimentar, respectivamente. Também houve diferença na importância alimentar dos itens entre os períodos pluviométricos. Estes resultados fornecem informações importantes sobre a ecologia alimentar de doradídeos na Amazônia. Além disso, percebeu-se o alto consumo de material alóctone pelo bacu-pedra, sendo estes itens alimentares provenientes da floresta ripária, o que reforça a importância deste ambiente para a conservação da ictiofauna neotropical.


Studies of feeding ecology are of fundamental importance for the understanding of interactive processes in fish communities (Winemiller, 1989; Hahn et al., 1997; Abelha et al., 2001), including those involving habitat features. In the Neotropics, aquatic environments provide fishes with an ample variety of food items, ranging from invertebrates to fruit and fishes (Goulding, 1980; Lowe-McConnell, 1999). However, the abundance of these resources may vary considerably over the course of the year, reflecting seasonal fluctuations in rainfall levels or hydrological parameters (Junk, 1980). This limits the degree of ecological specialization of most fish species for a single type of food item. Given this, most Neotropical fish species are dietary generalists or opportunists (Lowe-McConnell, 1999), and present considerable versatility in response to fluctuations in food availability (Abelha et al., 2001; Correa & Winemiller, 2014).

The vegetation along the stream and rivers provide food and habitat for fish during the flooding period. The flooding may occur irregularly or seasonaly by the intense rainfall or daily by the tide (Junk, 1980; Barthem & Schwassmann, 1994; Claro-Jr. et al., 2004; Junk et al., 2012). The rainy period influences also the phenology of growth and reproduction of plants, which may produce fruits in wet or dry season. Fruit and seeds may provide fishes with a valuable source of energy to overcome periods of reduced resource availability and specially to produce their lipid-rich eggs (Goulding, 1980). Fishes may change their diet in response to fluctuations in the abundance of specific resources between climatic seasons (Abelha et al., 2001; Novakowski et al., 2008). Species adapted to frugivory, for example, may be forced to shift the diet and consume leaves, flowers or other plants parts during certain periods, depending on plant phenology patterns and the characteristics of the fish species, such as its energetic demands and body size, among others (Chick et al., 2003; Anderson et al., 2011; Horn et al., 2011). Species such as açai berry (Euterpe oleracea Mart.), aninga (Montrichardia linifera (Arruda) Schott) and buriti (Mauritia flexuosa L.) are abundant on the tidal floodplain of the rio Amazonas mouth. They play a prominent role in the feeding ecology of frugivorous fishes due primarily to the fact that they have asynchronous fruiting periods, and thus provide resources throughout most of the year (Maia & Chalco, 2002; Guimarães et al., 2004; Leão & Carvalho, 2005).

The rock-bacu or bacu-pedra Lithodoras dorsalis (Valenciennes, 1840) is one of the largest thorny catfish (Siluriformes, Doradidae), reaching at least 90 cm fork length and 12 kg in weight (Goulding, 1980). The species occurs in northern South America in the Amazon estuary and neighboring coastal areas of French Guiana (Sabaj & Ferraris-Jr., 2003) and it is exploited by the commercial and local fisheries in the Amazon mouth. Lithodoras dorsails eats fruits and seeds and plays a role in the seed dispersal processes (Goulding, 1980; Souza-Stevaux et al., 1994; Waldhoff et al., 1996; Pilati et al., 1999; Maia & Chalco, 2002). This species was classified as omnivorous by Santos et al. (2004) and herbivorous by Santos et al. (2006), and consumes molluscs, aquatic insect larvae, fruits and seeds (Ringuelet et al., 1967; Santos et al., 2004, 2006).

The goal of this study was to investigate the diet shift throughout the year and the relative importance of food types in the diet of young Lithodoras dorsalis in the rio Amazonas estuary, an environment with daily tidal influence and marked pluviometric periods. The species foraging activity was expected to be higher during the rainy period due to the increase in habitats availability because of the greater range of variation in the river levels. This allows the fish to access new environments and resources daily, which does not occur in the dry season.

Material and Methods

Study Area. Data were collected in the municipality of Abaetetuba, in the confluence of the rio Tocantins and rio Pará, in Pará State, Brazil. Rio Pará receives part of the rio Amazonas discharge, wich contributes with suspended solids and high turbidity, changes in water level and current direction are directly related to the tidal effect (Sioli, 1984; Barthem & Schwassmann, 1994). The average daily tidal range at Abaetetuba is less than 3 m, but during the equinoxial spring tides and the full or new phases of the moon the tide amplitude reaches 4.0 m (Hida et al., 1999). The limnological caracteritics of the water change with the variation of the rio Tocantins discharge. The water is rather transparent during the high discharge period of the rio Tocantins and become turbid in the low discharge period, due to the influence of rio Amazonas waters (Sioli, 1984; Barthem & Schwassmann, 1994).

The vegetation is defined as tidal floodplain vegetation with ombrophilous, broadleaved species, merged with palm trees like buriti tree (M. flexuosa) and açai berry tree (E. oleracea), the latter being a species of great economic importance for local populations (Machado, 2008). The fruiting period of E. oleracea is from July to November (Guimarães et al., 2004), while M. flexuosa fruits from March to August (Sampaio & Carrazza, 2012).

The local climate can be classified as Af following the Köppen-Geiger classification, corresponding to the typical conditions of tropical rainforest ecosystems (Peel et al., 2007). Annual precipitation is approximately 2000 mm (Machado, 2008). The rainy season lasts from February to April, rain-drought from May to July, the dry season from August to November and drought-rain from December to January. These periods will be analyzed and related to the feeding of L. dorsalis. Mean temperature is 27ºC, ranging from 20ºC to 35ºC over the year. Relative humidity is high, around 85%, varying normally between 81% and 90% (Machado, 2008).

Sampling. Lithodoras dorsalis specimens were collected monthly over a year, between July 2010 and June 2011. Specimens were collected in the streams of Sirituba Island in Abaetetuba, Pará, Brazil (01º41'13.6"S 48º52' 48.8"W; Figure 1).

Fig. 1. Location of study area near the mouth of the rio Amazonas in Brazil. A, Location of Brazil on South America; B, rio Amazonas mouth with the sampling site; C, Study area (specimens were collected within the shaded area). 

Weir fishing nets of aproximately 10 m in lenght, 3 m in height (30 m2 of area), with a between-knots mesh size of 3-6 cm, were used to capture specimens. These nets were set in the stream mouth at dusk (between 17:00 h and 19:00 h), depending on the tide, and removed at dawn (05:00-07:00 h). This period of the day was adopted to ensure the capture of individuals moving from the smaller rivers to the main channel during the low tide. Specimens were removed each two hours from the area using seine nets (5 m x 1 m), and hand- or dip-nets.

The specimens captured were analised in laboratory conditions in the Federal Institute of Pará (Instituto Federal do Pará - IFPA), where they were weighed (total weight in grams - Wtotal) and measured from the anterior extremity to the end of the spine (standard length in centimeters - Lstd). A ventral-longitudinal incision was then made from the urogenital opening to the head for the removal of the stomach, which was weighed (grams) and emptied for the collection of its contents. The contents were weighed separately (grams) and then sorted in a Petri dish. Each distinct food item was identified using a stereomicroscope (40 x), and weighted separetely. The gonads of each specimen were also removed for the definition of maturation phases.

Following evisceration, the specimens were fixed in 10% formaldehyde, conserved in 70% alcohol, and then incorporated into the ichthyological collection of Museu Paraense Emílio Goeldi (MCT/MPEG), under the following catalog numbers: MPEG19134; MPEG19202; MPEG19203; MPEG19610; MPEG19611; MPEG21668-MPEG21681.

Data analysis. The Lithodoras dorsalis diet composition was inferred by analyzing stomach contents. Each food item was initially classified as to its origin (allochthonous or autochthonous) and taxonomy. Prey was identified to the lowest possible taxonomic level, depending on the current digestion degree, with the assistance of specialists and appropriate literature (Ruppert et al., 2005; Costa et al., 2006).

The Repletion Index (RI%; Zavala-Camin, 1996) was calculated to verify feeding activity levels. This parameter is obtained by RI% = (Wi / Wt) * 100, where: Wi represents total items weight and Wt total specimen weight. We used a Kruskal-Wallis nonparametric analysis of variance (H), with a 5% significance level, to assess differences in food ingestion rates (RI%) between pluviometric periods.

The relative contribution of different food items to the species' diet was assessed based on the Frequency of Occurrence (FOi%; Hyslop, 1980), the porcentage of stomachs in which each item occurred relative to total number of items, and on the Relative Weight (W%; Hynes, 1950), the weight percentage of each item relative to the weight of all analysed food items (Hyslop, 1980; Zavala-Camin, 1996). FOi% and W% values were combined to estimate the Alimentary Index (AIi%, modified from Kawakami & Vazzoler, 1980), which provides a measure of the importance of each food item. This index is calculated as AIi% = (FOi% * W% / ∑ FOi% * W%) * 100, where: AIi% represents the food item i alimentary index, FOi% the item i frequency of ocurrence and W% the relative weight of item i. Empty stomachs were not considered for this analysis.

To assess the diet shift of L. dorsalis among pluviometric periods, the AIi% values recorded for each item in the different seasons were log (X +1) transformed. The data were then analyzed using Non-metric Multidimensional Scaling (NMDS) based on the Bray-curtis similarity index of the transformed data (Clarke & Warwick, 2001). AIi% values were evaluated using an Analysis of Similarity (ANOSIM), with a 5% significance level. A Similarity Percentage Analysis (SIMPER; Clarke & Warwick, 2001) was then performed to identify which items predominated in the different seasons.

Trophic niche breadth was calculated using Levin's standardized index in order to describe changes in the level of dietary specialization relative to pluviometric periods. This index varies from zero, when a species consumes a single type of food category, to one, when it consumes all available foods in equal amounts (Hulbert, 1978). The index is calculated by Bi = [(∑jP2ij)-1] (n-1)-1, where: Bi is the standardized trophic niche; Pij is the ratio of the food category j on species' i diet; and n, the total number of food categories. To improve graphic visualization, the trophic niche breadth was calculated using month data, however the pluviometric periods were shown.


During the 12 months of the study period, 371 specimens of sexually immature Lithodoras dorsalis were captured. The mean standard length of these specimens was 15.40 cm (SD± 4.87cm) and mean weight was 94±149.45 g. Four of the specimens had empty stomachs, and were not included in the AIi% analyses. The stomachs sample size per month and season are shown in Table 1.

Table 1. Number of stomachs sampled, per month and pluviometric periods, of Lithodoras dorsalis at the rio Amazonas mouth, Brazil, from July 2010 to June 2011. 

Month/Period Number of stomachs sampled
July/10 20
August/10 40
September/10 40
October/10 30
November/10 30
December/10 30
January/11 31
February/11 25
March/11 27
April/11 37
May/11 27
June/11 34
Rain-Drought 81
Dry 140
Drought-Rain 61
Rainy 89
Total 371

Foraging intensity, as measured by the Repletion Index (RI%), varied significantly between seasons (H3,368 = 17.81; p < 0.05). The dry season returned the lowest and the highest value RI% (Median = 7.899; Max. = 23.084 and Min. = 0; Figure 2). Significant differences were also identified in the post-hoc multiple comparisons conducted for the Kruskal-Wallis test (Table 2). In particular, the dry season was significantly different from rain-drought and rainy seasons, showing higher values.

Fig. 2. Box-plot of Lithodoras dorsalis Repletion Index (RI%) at the rio Amazonas mouth, Brazil, from July 2010 to June 2011. 

Table 2. Probability values for pair-wise comparisons of the Repletion Index (RI%) among pluviometric periods for the rock-bacu Lithodoras dorsalis, sampled in the rio Amazonas mouth region, Brazil. (*) Significant values. 

Rain-drought Dry Drought-rain Rainy
Dry 0.001*
Drought-rain 0.195 1.000
Rainy 1.000 0.010* 0.543

A total of 28 food items were identified (Table 3), of which 16 were allochthonous and 12 autochthonous. In general, açaí fruit (E. oleracea) was the most important item in Lithodoras dorsalis diet (AIi% = 48.681%), followed by plant fragments (AIi% = 14.157%) and aninga (M. linifera) (AIi% = 13.179). Per pluviometric periods, M. linifera was the main food item consumed in the drought-rain period (AIi% = 37.677%), E. oleracea in dry and drought-rain periods (AIi% = 79.041 and 44.871, respectively) and plant fragments in the rainy season (AIi% = 24.715). According to the Alimentary Index (AIi%), L. dorsalis feeds mainly on plant material, in particular fruit and seeds. However, items such as brachyurans, decapods (Paleomonidae), and gastropods were also consumed (see Table 3).

Table 3. Alimentary Index (AIi%) per pluviometric period of food resources consumed by the rock-bacu Lithodoras dorsalis on the rio Amazonas mouth region, Brazil. (-) Represents the absence of the food item. 

Origin Food Items Rain-drought Dry Drought-rain Rainy
Allochthonous Formicidae 0.001 <0.0001 < 0.0001 0.001
Isoptera 0.014 - - 0.003
Diptera - - - 0.001
Diplopoda 0.001 - < 0.0001 < 0.0001
Arthropod Fragment 0.002 < 0.0001 0.001 0.011
Euterpe oleracea Mart. 18.274 79.041 44.871 16.539
Mauritia flexuosa Mart. 0.846 1.187 11.617 13.029
Montrichardia linifera (Arruda) Schott 37.677 0.108 0.078 24.391
Fruit fragments 1.590 1.688 0.712 3.150
Zea mays - < 0.0001 - -
Anacardium occidentale L. - 0.001 - -
Spondias mombin L. - - 0.090 0.019
Mangifera indica L. - - 0.216 -
Other plant fragments 8.561 9.471 13.744 24.715
Flowers 0.455 0.107 0.038 0.102
Leaves 0.026 0.151 - -
Autochthonous Ucarapax and Armases benedict 11.125 3.196 11.125 10.928
Macrobrachyum amazonicum 0.127 0.183 0.048 0.133
Gastropoda 0.197 0.001 10.725 0.480
Paxyodon syrmatophorus and Triplodon corrugatus 0.304 0.004 1.273 0.343
Sponges (Demospongiae) - - - < 0.0001
Chironomidae Larvae < 0.0001 < 0.0001 - -
Coleoptera Larvae - < 0.0001 0.030 0.002
Ciclorapha Larvae - < 0.0001 - -
Ephemeroptera Nymph - < 0.0001 - -
Oligochaeta - < 0.0001 - -
Fish fragments 0.001 0.022 0.001 0.009
Algae 2.174 0.120 0.581 1.131
Digested organic material 12.587 4.660 1.977 1.201
Substrate 6.038 0.001 0.261 2.192
Minerals < 0.0001 0.059 2.612 1.620

The NMDS and ANOSIM analyses based on the Alimentary Index (AIi%) of all items in the diet showed differences in the importance of food items in relation to pluviometric periods (ANOSIM: R = 0.38; p < 0.05) (Figure 3), with clear separation between dry and rainy seasons. The SIMPER analysis revealed that items such as M. linifera, E. oleracea, M. flexuosa and brachyura were responsible for differences among seasons (Table 4).

Fig. 3. Non-metric Multidimensional Scaling based on the Alimentary Index (AIi%) of all food items consumed by the rock-bacu Lithodoras dorsalis among pluviometric periods, rio Amazonas mouth region, Brazil. 

Table 4. Percentage contribution of the main resources consumed by Lithodoras dorsalis at Abaetetuba, south of the rio Amazonas estuary (Brazil), during the different pluviometric seasons, based on the results of the SIMPER analysis. 

Period Items % of Contribution
Rain-drought Montrichardia linifera (Arruda) Schott (Aninga) 35.1%
Brachyura 20.6%
Plant Fragment 20.1%
Dry Euterpe oleracea Mart. (Açai Berry) 50.2%
Plant Fragment 26.3%
Brachyura 10.2%
Drought-rain Euterpe oleracea Mart. (Açai Berry) 27.3%
Mauritia flexuosa Mart. (Buriti) 23.6%
Plant Fragment 23.0%
Rainy Montrichardia linifera (Arruda) Schott (Aninga) 37.6%
Plant Fragment 28.2%
Mauritia flexuosa Mart. (Buriti) 21.6%

The trophic niche breadth showed low seasonal variation in resource use by Lithodoras dorsalis (Figure 4), since this index ranges from 0 to 1 and the highest value was 0.13 in January. This species has a specialist behavior tendency, with narrow niche breadth, mainly in the beginning of dry season, on 2010, and at the end of the drought-rain transition period, on 2011, due to the high consumption of acai E. oleracea (AIi% = 89.89%) and M. linifera (AIi% = 84.41%), respectively.

Fig. 4. Trophic niche breadth recorded for Lithodoras dorsalis on the rio Amazonas mouth, Pará, Brazil, from July 2010 to June 2011. The solid line represents the niche breadth values and the dashed line indicates mean pluviosity registered for the studied region. 


The dominance of young individuals at the rio Amazonas mouth region is because the rock-bacu is a migratory fish with different feeding and reproductive sites. The rio Amazonas mouth is its nursery or growth area, and upstream sections of the rio Amazonas (including tributaries), the reproduction area (Goulding, 1980; Goulding et al., 1996; Barthem et al., 1991). Growth areas, essential for migrating species such as L. dorsalis, are sites where young individuals meet food and shelter needs all year long and are able to invest on body growth until attain sexual maturity, when they migrate to other regions (Goulding, 1980; Barthem et al., 1991). The rio Amazonas mouth has a high food availability, because most plant species fructify on different times, providing fruits and seeds for L. dorsalis almost all year long (Guimarães et al., 2004; Leão & Carvalho, 2005; Sampaio & Carrazza, 2012). In addition, because the water variation in the region is determined by the tides, fruits are accessible daily as they fall on the water surface (Barthem & Schwassmann, 1994; Hida et al., 1999).

The high presence of fruits and seeds, like E. oleracea, M. linifera, M. flexuosa, highlights the importance of riparian forests as a substantial food source for Amazonian fishes (Goulding, 1980). Many studies demonstrate the importance of these food items for the ichthyofauna, as a consequence of its high energy level (Hahn et al., 1992; Waldhoff et al., 1996; Winemiller & Jepsen, 1998; Claro-Jr. et al., 2004; Galetti et al., 2008; Pollux, 2011; Correa & Winemiller, 2014). However, the variation in resource availability among pluviometric periods may modify species diet, where most fish species may alter their feeding behaviors according to changes in resource relative abundance (Abelha et al., 2001). The fish fauna of the Amazon basin present a number of specific adaptations to maximize the ingestion of energy according to major seasonal fluctuations in resource availability, for instance the opportunistic feeding behavior (Junk, 1985; Lowe-McConnell, 1999). This pattern was recorded for young L. dorsalis, here considered herbivore with a high frugivory tendency. The species fed mainly on fruits and seeds (almost 85% of total AIi%), according to the phenology of these plants, but also consumed items such as Crustacea and Gastropoda on different times of the year, based on the availability of these invertebrates on the environment. The herbivorous behaviour with high frugivory tendency of adult L. dorsalis was also observed by Goulding (1980) in the rio Madeira .

The preference for a given food item due to its availability may be explained by the Optimal Foraging Theory (MacArthur & Pianka, 1966), which predicts that species will choose resources to combine high ease of capture with large amount of energy obtained. Therefore, during the year, species may present a specialist or generalist behaviour in obtaining food resources, depending on availability (Abelha et al., 2001; Wolff et al., 2009; Mazzoni et al., 2010; Masdeu et al., 2011; Correa & Winemiller, 2014). Considering the low taxonomic variety of items ingested, L. dorsalis exhibited a behariour tending to specialist, eating principally fruits and seeds from the riparian forest, varying the species consumed probably according to availability. Futhermore, the presence of sediments in the stomach of Lithodoras dorsalis specimens indicates that the species also forages on the bottom, where it obtains the bivalves (mussels and oysters) consumed (Beasley, 2001; Hohn & Costa, 2002). This type of foraging behavior has also been recorded in other doradid species (e.g., Hahn et al., 1997; Lowe-McConnell, 1999). Species of this family explores all environments searching for food, such as the surface to eat floating fruits, although it is adapted to live in river bottoms.

In view of the discussion above, food availability among pluviometric periods probably influenced L. dorsalis feeding intensity (RI%) and its diet composition along the year at the rio Amazonas mouth. The different fructification periods among plant species influence which fruit is consumed during pluviometric periods, changing its foraging behavior (more or less specialist). In addition, the daily variation of water level due to tides is a factor that influences the availability of usable areas by fish populations (Morton et al., 1987; Krumme et al.; 2004, Stevens et al., 2010), since the daily flooding of riverbanks allows fishes to reach resources outside the main channel everyday. It does not occurs in environments influenced only by seasonal flood pulses; in such areas, fruits are available for fishes only during flooding periods, few months of the year (Goulding, 1980), or in sites where trees are near the river (Anderson et al., 2011; Horn et al., 2011). In conclusion, allochthonous items, especially those of vegetal origin, are important for Lithodoras dorsalis as a food source. It highlights the importance of riparian forest conservation at the rio Amazonas mouth, a key source of food resources for Neotropical fishes.


We thank to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the scholarship (TAPB). The Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) provided LFAM with a research fellowship (process no. 301343/2012-8). The Instituto Federal do Pará, IFPA, Abaetetuba for the logistic support. We also thank the researches and professors M.Sc. Marta Coutinho (IFPA), Dr. José Birindeli (MZUSP), Dr. Cléverson Santos (MPEG), Dr. Rony Vieira (UFPA), Dr Roberta Valente (UFPA) and Dr Alba Lins (MPEG) for the help identifying the stomach contents of Lithodoras dorsalis, Dr. Leandro Juen (UFPA) and Dr. Bruno Spacek (UFPA) for the help with statistical analysis, Mr. Ilso, Jaime and Giovani for field assistance and Dr. Stephen F. Ferrari for his assistance with the english review.


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Received: January 22, 2013; Accepted: January 16, 2015

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