artigo eucilene

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Acta Limnol. Bras., 2009, vol. 21, no. 3, p. 359-366.

Abelha et al., 2001), and can provide data about habitat,

ood availability, and even behavioral eatures (Hahn et al.,1997).

Concepts or guilds or trophic groups established vary inaccordance with the purpose o studies. Root (1967) denestrophic groups or guilds, as representing a part o a commu-nity eeding on the same class o environmental resources.

According to Simberlo and Dayan (1991), the conceptor ecological guild includes subsets within species group,having high potential or competition. Austen et al. (1994)emphasized also that may provide a means to identiy spe-cies with similar responses to environmental variation. Ina review o the application o the guild concept in sheries

1. Introduction

A ood plain normally comprises various environmen-

tal types: rivers, temporary and permanent lakes, channels,and ressaccos, with unique characteristics that distinguishthem rom the other types. Considered semi-lentic (Fonsecaand Rodrigues, 2005), these areas - located in uvial islandsin the Paran River have been studied since the end o thelast century by PELD, a Brazilian program dedicated tolong-term research.

Te knowledge o sh diets allows not only the identi-cation o trophic categories but also inerences about theirstructure. Furthermore, it provides a basis or understandingthe relationships between ichthyoauna and other organ-isms present in the community (Gaspar da Luz et al., 2001;

Trophic organization the ichthyoauna o two semi-lenticenvironments in a food plain on the upper Paran River, Brazil

Estrutura trca da ictioauna em dois ambientes semi-lnticosda plancie de inundao do Alto Rio Paran, Brasil

Santana-Porto, EA.1 and Andrian, IF.2

1Ps-Graduao em Ecologia de Ambientes Aquticos Continentais, Universidade Estadual de Maring UEM,Av. Colombo, 5790, Zona 7, CEP 87020900, Maringa, PR, Brazil

e-mail: [email protected]; [email protected]

2Departamento de Biologia, Centro de Cincias Biolgicas,Instituto de Biologia e Ps-Graduao em Ecologia de Ambientes Aquticos Continentais,

Universidade Estadual de Maring UEM, Av. Colombo, 5790, Zona 7, CEP 87020900, Maringa, PR, Brazile-mail: [email protected]; [email protected]

Abstract: Aim: Te purpose o this study was to identiy, using the guild concept, the trophicgroups o ichthyoauna in little known environments called ressaccos(riverine inlets subject to periodicisolation depending on oodplain conditions) located on the ood plain o the upper Paran River.Methods: Individuals were caught by nets and aterwards separated according to their stomach contents. oidentiy trophic groups, the unweighted pair-group method (UPGMA) was adopted. Result: By analyzingsh diets, we classied 13 sh species distributed within ve trophic groups in the Manezinho ressacco,and 20 species o eight trophic groups in Bile ressacco. O the total the species identied, 11 occurred inboth environments. Conclusion: Even though Cladocera was the preerred ood o the majority o thespecies ound, an ampler quarrel o the use o the guild term is necessary since the variety o item ingestedor the species.

Keywords: ood, shes, trophic groups, ressaccos.

Resumo: Objetivos: Identicar grupos trcos da ictioauna de ambientes pouco conhecidos comoressacos, na plancie de inundao do alto rio Paran utilizando o conceito de guilda. Mtodos: Osexemplares de peixes oram capturados pelo mtodo de captura ativa analisados posteriormente quandoa preerncia alimentar presente no contedo gstrico. Para a ormao dos grupos trcos, utilizou-se

o mtodo de agrupamento pareado igualmente ponderado (UPGMA). Resultados: A identicao docontedo estomacal possibilitou a classicao de 13 espcies de peixes em cinco grupos trcos no ressacodo Manezinho e, das 20 espcies do Bile em oito categorias trcas. Das 22 espcies examinadas, 11 oramcomuns aos dois ambientes. Concluso: Apesar do itm preerencial ser Cladocera para a maioria dasespcies nos dois ambientes analisados, necessria uma discusso mais ampla da utilizao do termoguilda, devido variedade de itns ingeridos pelas espcies.

Palavras-chave: ictioauna, grupos trcos, guilda, ressaccos.


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360 Santana-Porto, EA. and Andrian, IF.


management, Austen et al. (1994), distinguished betweenstructural guilds (groups o species that use similar resourc-es) versus guilds that unctions as a super specie (groups ospecies that collectively respond to environmental variationin a more or less consistent manner). Species within guildsbased on dietary similarity (use similar resources) responddierently to key abiotic impacts such as ow alteration

(Welcomme et al., 2006) limited the application o theconcept. Furthermore, Regier et al. (1989) proposed theterm environmental guild, or identiying sh speciesthat respond in similar manner to changing hidrology andgeomorphology o river ecosystems.

In this study we analyze relationships between ood-chain consumers or species shes, aggregate entities bygrouping o trophically similar species that are sometimescalled guild with objective o the identiy trophic groupsin shes ound in the ressaccoso the ood plain o theupper Paran River, an ecosystem with high spatial and

temporal variations.

1.1 Study area

Te Paran River, which is next to the Municipality oPorto Rico, is divided into two main branches and a smallcanal by two islands, the Mutum and the Porto Rico. Botho these contain several lakes and resaccos(Cunico et al.,2002).

Te studied area includes two resaccos, a popular termor riverine inlets, called Manezinho and Bile, both locatedon Mutum Island (Figure 1). In the years in which thisstudy took place, these ressaccos were connected to theParan River only by a channel, which is not visible owingto prevailing morphological conditions.

Te 582.6 m long Bile resacco (22 45 13.56 S and53 17 9.48 W) had an average depth o 1.3 m duringthe period studied. Cyperacae and Leguminosae, such as

Mimosa pigra e Inga uruguensis, are present in its margins.Te 100 m long Manezinho resacco (22 4644.7 S

and 53 20 56.76 W) occupies a 1 ha area. In the stud-

ied period, its average depth was 2.1 m, and the unique

BrazilBrazil

Paran

R.

Paran

R.

Mato Grosso do Sul

State

Mato Grosso do Sul

State

Paran

River

Paran

River

ParanRiv

er

ParanRiv

er

Porto

RicoIs

land

Porto

RicoIs

land

Mutum

Island

Mutum

Island

Paran

State

Paran

State

Ressacco BilRessacco Bil

Ressacco ManezinhoRessacco Manezinho

BaiaRiver

BaiaRiver

Carioca

Island

Carioca

Island

00 11 22 33 44 5 km5 kmScale graphicalScale graphical

NN

SS

WW EE

53 21 W53 21 W

22 40 S22 40 S

53 12 W53 12 W

53 21 W53 21 W 53 12 W53 12 W

22 40 S22 40 S

22 48 S22 48 S 22 48 S22 48 S

StreamC

aracu

StreamC

aracu

Str

eam

gua

Do

is

Str

eam

gua

Do

is

Stream

Curutu

ba

Stream

Curutu

ba

NUPELIA

Advanced Base

NUPELIA

Advanced Base

*

Rib

.Baile

Rib

.Baile

Rib

.SoPedro

Rib

.SoPedro

Figure 1. Location o Manezinho and Bile ressaccoson the ood plain o the Paran River.


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rophic organization the ichthyoauna o two semi-lentic environments in a ood plain... 361


connection with the river was through a channel 1 m longand 3 m wide. Te margins o Manezinho are covered byarboreous vegetation, with a predominance o Cecropiaand Inga spp.

In this resacco, ichthyoauna sampling was done quaterly,in 2000 (February, May, August and October) and 2001(February, May, August and November). Atypical years arenot uncommon on ood plains because inundation can bedelayed, e.g., at March/00, and at the end o January/01.

According to Agostinho and Zalewski (1995), seasonalchanges usually occur rom November through March..

2. Material and Methods

Active capture with three attached drag nets (5 mmmesh, 50 m long, 2.8 m high) was used to collect shes,

which were measured, weighed, and dissected. Stomachsand respective contents were xed in 4% ormaldehyde.

Stomach content was analyzed with a stereoscopic mi-croscope; items ound were identied at the lowest possibletaxonomic level. Calculations o volumetric requency andoccurrence were obtained by methods proposed by Hynes(1950) and Hyslop (1980). Using a graduated test tube(1, 2, and 5 mL), volumes were determined rom liquiddisplaced.

Occurrence (Fo) and volumetric (Fv) requencies werecombined in the alimentary index (IAi) (Kawakami andVazzoler, 1980). Te IAi values were converted into per-centages and then into cumulative sums. Based on thesedata, the preerred item and eating habits o a given species

were indicated by the IAi percentage (50%) (Gaspar daLuz et al., 2001). Subsequently the cluster analysis was

applied, based on the unweighted pair-group method(UPGMA), using Statistica.5.5. Tis analytical techniqueis useul in determining signicant groups o individuals,or objects (Hair, 1987), or trophospecies. Te IAi percent-ages were used or each ressacco in the analysis. Accordingto Pinto-Coelho (2000), the percentages had a classica-tory property, in this case signiying that all species wereorganized in distinct subgroups, in an ordered sequence ohierarchical levels.

3. Results

Examination o 230 gastric contents the twenty-twospecies, 13 rom Manezinho and 20 rom Bile was madeor the determination o trophic organization in thetwo ressaccos. O the 22 species, 11 were ound in bothenvironments.

In Manezinho ressaco, ive trophic groups wereregistered (able 1), o which zooplanktivory predomi-

Table 1. Number o stomachs analyzed; minimum and average length (L)); trophic characterization; preerred ood item; and IAi(Index o Alimentary Importance) o sh species ound in the Manezinho ressacco on Mutum Island in the Paran River, Brazil.

Species Number of

stomachs

LT

minimum

(cm)

LT

average

lengt (cm)

Preferred food item %IAi Tropic caracterization

Astyanax altiparanae

(Garutti and Britski, 200)

1 7.2 Insects (Hymenoptera,

Ortopthera and Diptera)

Vegetable (fruit the macrophytes,

Algae and plants remnants)

48.67 Omnivorus

Aphyocharaxsp. 13 2.9 3.2 Cladoceran 51.33 (Begon et al., 1990)

Aphyocharax anisitsi

(Eigenmann andKennedy, 1903)

1 4.5 Cladoceran 61.56 Zooplanktophagous

Bryconameicus stramineus

(Eigenmann, 1908)

16 2.7 4.1 Cladoceran 78.80 (Lansac-Tha & Alves, 1994;

Gaspar da Luz & Okada, 1999)

Serrapinnus notomelas 2 2.4 3.0 Cladoceran 69.25 Zooplanktophagous

Hyphessobrycon sp. 19 3.0 3.4 Cladoceran 83.33 Zooplanktophagous

Hemigrammus marginatus

(Ellis, 1911)

9 4.1 4.4 Insects

(Diptera and Hymenoptera)

61.14 Zooplanktophagous

Hoplias aff. malabaricus

(Bloch, 1794)

1 9.6 Fish rest 59.80 Piscivores

(Agotinho et al.,1997)

Moenkhausia intermedia

(Eigenmann, 1908)

4 5.3 6.0 Cladoceran 100.00 Zooplanktophagous

Moenkhausia

sanctaelomenae

(Steidachner, 1907)

2 3.3 3.7 Coleoptera 83.33 Insectivorous (Adrian,

Lansac-Tha & Alves, 1994;Gaspar da Luz & Okada, 1999)

Odontostilbe sp. 11 2.4 2.8 Cladoceran 71.67 Zooplanktophagous

Steindachnerina insculpta

(Fernandes-Ypez, 1948)

3 3.5 3.5 Detritus 98.37 Detrivorous

Satanoperca pappaterra

(Heckel, 1840)



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nated in eight species, representing 57% o total spe-cies ound in this environment. In Aphyocharax sp.,

Aphyocharax anisitsi, Bryconamericus stramineus, Serrapinnusnotomelas, Hyphessobrycon sp., Moenkhausia intermedia,Satonoperca pappaterra and Odontostilbe sp., cladoceranpredominated.

Groups shown in the dendogram (Figure 2) comprisetwo sets. Te rst o these groups includes B. stramineusandM. intermedia because o high IAi rankings o cladoceranand diptera in stomach contents. Group 2, which presentedIAi values above 60% or cladoceran, consisted o theprevious two species plusAphyocharaxsp.,Aphyocharax an-isitsi, Odontostilbesp., Hyphessobrycon sp., S. notomela, andS. pappaterra. Te Hemigrammus marginatusisolated specieso the two main groups shows no trophic similarity with theother groups because o its preerence or cladoceran.

Te sh assemblage o Bile ressaccowas distributed in

eight trophic groups (able 2), which six species (30% othe total) were identied in the zooplanktivorous trophicgroups: Apareiodon ainis, Bryconamerius stramineus,Serrapinnus notomelas, Hyphessobrycon eques, Hemigrammus

marginatus, and Moenkhausia intermedia. O these, three

species (B. stramineus, S.notomelasand M. intermedia) wereound in both ressacco environments.

Te dendogram (Figure 3) shows our groups. Te rstincludes the speciesM. intermedia, A. anis, H. marginatus,S.notomelas, H. eques, and B.stramineus, classied togetherbecause their preerred dietary item (cladoceran) ranks on

the IAi above 50%, and in some cases reaches 100%.Group two comprised two species: Serrasalmusmarginatusand Metynnis c. maculatusbecause o the com-mon presence in stomach content o algae, macrophytes,detritus, and scales.

Group three included the species R. paranensis andAphyocharaxsp., linked by the common consumption oAmphipoda, a ood item restricted to this group.

he species Aphyocharax anisitsiand Moenkhausiasanctaelomenaecomprised group our, whose diet includedcladoceran, diptera, detritus, Hemiptera, Conchostraca,

and Ephemeroptera (able 3).

Insectivorous

Insectivorous

Detrivorous

Piscivores

ZooplanktivorousZooplanktivorous


II

I

OmnivorousOmnivorous

0 20 40 60 80 100 120

(Dlink/Dmax)*100

A.al

A.an

S.n

S.p

A.sp

B.s

M.i

H.sp

O.sp

H.ma

M.s

S.i

H.m

Figure 2. Dendogram grouping o IAi (Index o AlimentaryImportance) values or 13 sh species ound in the Manezinhoressacco, located on the oodplain o the upper Paran River.Caption= (I, II, III) = separate groups; sh species (H.malaba= Hoplias af.malabaricus; S.inscul = Steidachenerina insculpta;M.sancta = Moenkhausia sanctaeilomenae; H.margin =Hemigrammus marginatus; Odontos = Odontostilbesp.; Hyphesso =Hyphessobrycon sp.; M.inter =Moenkhausia intermedia ; Bstramin =Bryconamericus stramineus; Aphyocha=Aphyocharaxsp.; S.pappat =Satonoperca pappaterra; S.notome = Serrapinnus notomelas;A.anists =Aphyocharax anisitsi; A.altipa =Astyanax altiparanae).

0 20 40 60 80 100 120

(Dlink/Dmax)*100

A.alI.i

A.anM.saS.ma

S.pS.iM.iA.af

H.maS.nB.sH.eR.p

A.spM.mS.mH.mL.p

C.m


CarnivorousnesssCarnivorousnesss

Algvorous

II

III

I

EurifgicIV

Figure 3. Dendogram grouping o IAi (Index o AlimentaryImportance) values or 20 sh species ound in the Marezinho

ressacco, located on the oodplain o the upper Paran River.Caption = (I, II, III) = separate groups; sh species (C.monoculu =Cichla monoculus; L.platym = Loricariichtys platymetopon;H.malaba = Hoplias af.malabaricus; S.macula = Serrassalmusmaculatus; M.macula = Metynnis c.maculatus; Aphyocha =Aphyocharax sp.; R.parane = Roeboide paranensis; H.eques =Hyphessobrycon eques; B.strami = Bryconamericus stramineus;S.notome = Serrapinnus notomelas; H.margin = Hemigrammusmarginatus; A.aini = Apareidon ainis; M.intermdia = Moenkhausia intermedia; S.inscul = Steidachenerina insculpta;S.pappat = Satonoperca pappaterra; M.sancta = Moenkhausiasanctaeilomenae; A.anists = Aphyocharax anisitsi; I.labros =Iheringichthys labrosus; A.altipa =Astyanax altiparanae).


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4. Discussion

Te variety o ood items available avors predominanceo opportunists or generalists rather than specialists on theood plain (Gaspar da Luz, 2000). Sudden environmentalchanges explain ood item variations, particularly insectsand other invertebrates ound in stomachs o detrivorousspecies in the two ressaccos(Leo et al., 1991).

Te studied period occurred in atypical years withsevere droughts and late ooding on the ood plain o theupper Paran River (Agostinho and Zalewski, 1995). Such

conditions complicate distinguishing in guilds. However,Junk (1980) suggested that in environments with varyinghydrological conditions, studies must be carried out underextreme conditions, which produce the greatest dierencesin available ood items and which thereore orce sh toutilize their adaptive capacities.

Another actor that makes guild delineation difcult isood plasticity. According to Gaspar da Luz et al. (2001), itconsists in the interaction o quantity and quality o avail-able ood. Is is outstanding in tropical riverine ichthyoauna(Goulding, 1980; Hahn et al., 1997; Lowe-McConnell,

Table 2. Number o stomachs analyzed; minimum and average length (L)); trophic characterization; preerred ood item; and IAi(Index o Alimentary Importance) o sh species ound in the Bile ressacco on Mutum Island in the Paran River, Brazil.

Species Number

of

stomachs

LT

minimum

(cm)

LT

average

lengt (cm)

Preferred

food

item

%IAi Tropic

caracterization

Apareiodon afnis

(Steindachener, 1879)

1 2.1 Cladoceran 100.00

Astyanaxaltiparanae(Garutti and Britski, 200) 19 2.7 7.5 Macrophytes 57.80 Herbiorous

Aphyocharaxsp. 1 4.6 Amphipod 100.00

Aphyocharaxanisitsi

(Eigenmann and Kennedy, 1903)

6 3.1 4.0 Diptera 72.00 Insectivorous

Bryconamericus stramineus

(Eigenmann, 1908)


Cichla monoculus (Spix, 1831) 1 13.2 S. pappaterra 100.00 Psicivores (Agostinho et al. , 1997)

Serrapinnus notomelas

(Eigenmann, 1915)


Hyphessobrycon eques

(Steindachner, 1882)


Hemigrammus marginatus

(Bloch, 1794)

5 4 4.8 Cladoceran 79.20 Zooplanktophagous

Hoplias aff. malabaricus

(Bloch, 1794)

11 2.7 31.0 Decapod 85.68 Carnivorousness

Lheringicthys labrosus(Ltken, 1874)

4 17.0 23.0 Bivalvia 60.13 Benthivorous

Loricariichthys platymetopon

(Isbeker and Nijssen, 1979)

6 2.8 16.1 Detritus 84.54 Detrivrous

Moenkhausia intermedia

(Eigenman, 1908)


Metynnis cf. maculatus

(Knerr, 1858)

4 9.9 15.7 Algae 95.04 Algivorous

Moenkhausia sanctaelomenae

(Eigenmann, 1908)

19 3.2 4.9 Insects 73.74 Insectivorous

Roeboides paranensis(Pignalberi, 1975)

11 2.6 3.9 Amphipod 68.74 Carnivorousness

Steindachnerina insculpta

(Fernndez-Ypez, 1948)

10 3.3 7.9 Detritus 70.74 Detrivorous

Serrasalmus marginatus(Valenciennes, 1836)

5 2.3 17.6 Fish rest 56.33 Piscivores

Satanoperca pappaterra

(Heckel, 1840)

3 2.5 18.8 Psocoptera 76.34 Insectivorous

Serrasalmus maculatus

(Kner, 1858)

1 15.9 Algae 79.74 Pscivorous (Agostinho et al., 1997)


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Table 3. Food items consumed by sh species in the ressaccoso Bile () and Manezinho () on the ood plain o the Paran Riverin 2000-2001.

Foo

d

resources

Apareiodonafnis

Aphyo

charaxanisitsi

Aphyocharaxsp.

Astyan

axaltiparanae

Bryconam

ericusstramineus

Cichlamonoculus

Serrapinnusnotomelas

Hyphessobryconeques

Hoplias

aff.

malabaricus

Hemigram

musmarginatus

Iheringichthyslabrosus

Loricariich

tysplatymetopon

Moenkhausiaintermedia

Metynniscf.maculatus

Moenkhausiasanctaeflomenae

Roeboid

esparanaensis

Serrassa

lmusmarginatus

Satanop

ercapappaterra

Serrassa

lmusmaculatus

Steindachnerinainsculpta

Hyphe

ssobryconsp.

Odo

ntostilbesp.

Trichoptera

Copepoda

Cladocera

Diptera

Hymnoptera

Detritus

Hydracarina

Hemiptera

Decapoda

Parts insects

Parts vegetable

Acarina

Fragments fshes

Nematoda

Conchostraca

Ortophtera

Fruit the

macrophytes

Algae

Scale fsh

Amphipoda

Coleoptera

Lepidoptera

Gastropoda

Bivalvia

Neuroptera

Macrophytes

Matter ingested

Tecameba

Ephemeroptera Odonata

Satanoperca.

pappaterra

Homoptera

Annelida

Culicidae

Collembola

Ceratopogonidae

Psocoptera

Parts seed


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1999), because it allows species to alter ood item consump-tion in accordance with prevailing relative abundances.

In the present study, plasticity was observed inR. paranensis, which although classied as insectivorouson the ood plain (Hahn et al., 2002; Agostinho et al.,1997), the sh tends towards carnivorousness in the Bileressacco, in which the predominant ood item consumed

was Amphipoda.In spite o the prolonged drought occurring in the

studied period, it appears that the environment neverreached the destabilization level, since omnivorous species

were limited toA. altiparanaein the Manezinho ressacco.According to Begon et al. (1990), trophic web theoryviews omnivorousness as a rare phenomenon because it isa destabilizing actor, i.e., species so characterized competemore intensely within their own trophic level, as well asbeing preyed upon by sh o higher trophic levels (Pimm1982; 1991).

An unusual characteristic o the ressaccoswas that onlyone herbivorous species, was recorded: A. altiparanaeinBile ressacco. Tis supports Junks theory (1980), whichpostulates that when water levels are low, even though agreat number o ood items are available to predators this isuntrue or herbivorous species, since macrophytes as well asvegetation in normally ooded areas have dried up.

Te presence o piscivores was attributed to H. af.malabaricus, which is common to the two environments,and C. monoculus, ound only in Bile ressacco. Accordingto Agotinho et al. (1997), in the ood plain o the ParanRiver, piscivory by species whose lie cycle occurs in lentic

environments (like H. af. malabaricus) is a permanentcondition. In addition, this trophic category tends to havegreater biomass in such environments, particularly when the

water level is low. Te same probably holds or ressaccos.In the Manezinho ressacco, detritivory was attributed

to S. insculpta, and, to L. platymetopon and S. insculpta inBile. In general, only a small percentage o species eed ondetritus. According to Bowen (1983), most shes belongto higher trophic levels and use invertebrates as a link tothe detrivorous base o the trophic chain.

Items only occasionally consumed by the speciesS. insculpta included cladoceran, nematodes, algae, insects,

terrestrial vegetation, acarids, and thecamoebas, but as theycomprised only a small percentage o items registered,species can be classied as detrivorous. Te same variety

was observed by Peretti and Andrian (2004) or the PauVeio. Even ressacco so, Agostinho et al. (1997) classiedS. insculpta as a mud-eater. Tese results show the alimen-tary plasticity o this sh species, i.e., the capacity to adaptto environmental circumstances aecting ood quality.

Te same variation showed that ressaccoshave the sameunction as lakes along the banks o the river in the oodplain, namely their importance in maintaining intact re-gional biodiversity. Because o their supply o ood items

and types o suitable habitats e. g., or aquatic macrophytes,they are the preerred environment o small, sedentaryspecies.

In the two Manezinho and Bile ressacos, the greatestnumber o sh species belonged to the zooplanktivoroustrophic category because o their preerred ood item:cladoceran. Tis may be associated with the availability o

this resource in 2001 on the ood plain environments, asveried by Lansac-ha et al. (2002). However, in spite othe predominance o Cladoceran, these species presenteda variety o ood items in their diets, suggesting that theyare actually opportunists, and thus cannot be grouped ina single guild, which would necessitate urther study as totheir oraging behavior.

Based on the ndings presented in this work, we con-clude that more study is needed on small sh (up to 5 cm)

whose eeding habits have not yet been completely identi-ed and, principally, their role (guild) within the trophic

web, an approach to which is exemplied in the work oHahn et al. (2002).

Acknowledgements

For granting us nancial support, we thank the Long-term Study Program (PELD) o the National Council orScientic and echnological Development (CNPq) and theCoordinating Agency or Furthering raining o Collegeand University eachers and Researchers (CAPES). We arealso grateul to the research group dedicated to limnology,ichtiology, and aquaculture (NUPELIA) o Maring StateUniversity or logistical, technical, and scientic support.

We express our gratitude to the proessors and administra-tive personnel o the post-graduate program in Ecology oContinental Aquatic Environments or their contributionsand support.

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AUSEN, DJ., BAYEL, PB. and MENZEL, BW. Importanceo the guild concept to sheries research and management.Fisheries, 1994, vol. 19, no. 6, p. 12-20.

AGOSINHO, AA., HAHN, NS., GOMES, LC. and BINI, LM.Estrutura trca. In VAZZOLER, AEAM., AGOSINHO,AA. and HAHN, NS. (Eds).A plancie de inundao do altorio Paran: aspectos sicos, biolgicos e socioeconmicos.Maring: EDUEM/NUPELIA, 1997. p. 229-248.

AGOSINHO, AA. and ZALEWSKI, M. Te dependence oshe community structure and dynamic on oodplain andriparian ecotone zone in Paran River, Brazil. Hydrobiology,1995, vol. 303, no. 2, p.141-148.

BEGON, MJL., HARPER, and OWNSEND, CR. Ecology:individuals, populations and communities. 2 ed. Oxord:Blackwell Sc. Publ., 1990. 943 p.


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Received: 08 December 2008Accepted: 15 September 2009

artigo eucilene

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