179
The Small Wa : T ophic in e ac ions be ween small-bodied
non-na i e and na i e cha acoids in he ma ginal zones o
Neo opical ese oi s
João Daniel Fe az1,2 , A mando Césa Rod igues Casimi o1,2 , Diego Aze edo Zoccal Ga cia2,3 ,
Ana Paula Vido o-Magnoni1,4 , And é Lincoln Ba oso de Magalhães5, Ali Se han Ta kan6,7 ,
John Robe B i on7, Má io Luís O si1,2
1 Uni e sidade Es adual de Lond ina, P og ama de Pós-G aduação em Ciências Biológicas, Depa amen o de Biologia Animal e Vege al, Rodo ia Celso Ga cia
Cid, 86057-970, Lond ina, PR, B azil
2 Uni e sidade Es adual de Lond ina, Cen o de Ciências Biológicas, Depa amen o de Biologia Animal e Vege al, Labo a ó io de Ecologia de Peixes e In asões
Biológicas, Rodo ia Celso Ga cia Cid, 86057-970, Lond ina, PR, B azil
3 Uni e sidade Fede al de Ma o G osso do Sul, Faculdade de Medicina, A enida Cos a e Sil a, s/n, 79070-900, Campo G ande, MS, B azil
4 Labo a ó io de Ecologia e Compo amen o Animal, Depa amen o de Biologia Animal e Vege al, Uni e sidade Es adual de Lond ina, Lond ina, Pa aná, B asil
5 Rua P o esso A duíno Boli a , 80, Belo Ho izon e, Minas Ge ais 30350-140, B azil
6 Depa men o Ecology and Ve eb a e Zoology, Facul y o Biology and En i onmen al P o ec ion, Uni e si y o Lodz, Lodz, Poland
7 Depa men o Li e and En i onmen al Sciences, Facul y o Science and Technology, Bou nemou h Uni e si y, Poole, UK
Co esponding au ho s: João Daniel Fe az ([email p o ec ed]); Ali Se han Ta kan ([email p o ec ed])
Copy igh : © João Daniel Fe az e al.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Small-bodied cha acoids ep esen a signi ican po ion o B azil’s eshwa e ish auna, ye hey
emain unde s udied. In ese oi s, hese species a e highly abundan , colonise he ma ginal
zone and a e impo an in he aqua ic ood webs. In he lowe Pa anapanema Ri e , he neigh-
bou ing ese oi s Rosana and Taqua uçu di e in hei en i onmen al condi ions, bu bo h
hos na i e and non-na i e cha acoids. This s udy desc ibes he die , eeding ecology and in e -
ac ions o be ween co-exis ing na i e and non-na i e small-bodied cha acoids in bo h ese oi s,
as well as hei ophic esponses o seasonal clima ic a ia ions (d y s. we season). Samples
we e collec ed qua e ly be ween Sep embe 2018 and Sep embe 2020, wi h awls, sie es and
cas ne s collec ing ish in ma ginal zones and aqua ic mac ophy e beds. The s omach con en s
o 416 indi iduals om se en na i e and non-na i e cha acoids we e analysed, wi h hei p ey
classi ied in o se e al ood ca ego ies and esou ces. In he Rosana Rese oi , bo h na i e and
non-na i e cha acoids we e s ongly elian on alloch honous esou ces and hei die composi-
ion showed simila shi s be ween he we and d y season. Con e sely, in he Taqua uçu Res-
e oi , he cha acoids we e mo e elian on au och honous esou ces and showed weak die a y
shi s be ween seasons. Niche b ead h and ophic o e lap indices indica ed die specialisa ion
and seg ega ion in he cha acoids o he Rosana Rese oi , while, in Taqua uçu, hey o e lapped
signi ican ly, especially in he d y season. These esul s e eal conside able di e ences in he
ophic in e ac ions be ween na i e and non-na i e cha acoids in wo Neo opical ese oi s,
sugges ing high con ex dependency in he ecological implica ions o in oducing non-na i e
species o hese small-bodied ishes.
Key wo ds: Biological in asion, damming, ood esou ces, niche b ead h, Pa aná Ri e , ophic ecology.
Academic edi o : Josie Sou h
Recei ed:
30 July 2025
Accep ed:
28 Oc obe 2025
Published:
24 No embe 2025
Ci a ion: Fe az JD, Casimi o ACR,
Ga cia DAZ, Vido o-Magnoni AP,
de Magalhães ALB, Ta kan AS,
B i on JR, O si ML (2025) The Small
Wa : T ophic in e ac ions be ween
small-bodied non-na i e and na i e
cha acoids in he ma ginal zones o
Neo opical ese oi s. NeoBio a 104:
179–201. h ps://doi.o g/10.3897/
neobio a.104.167161
NeoBio a 104: 179–201 (2025)
DOI: 10.3897/neobio a.104.167161
Ad ancing esea ch on alien species and biological in asions
A pee - e iewed open-access jou nal
NeoBio a
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NeoBio a 104: 179–201 (2025), DOI: 10.3897/neobio a.104.167161
João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
In oduc ion
B azil has nume ous hyd opowe ese oi s, pa icula ly in he sou heas and sou h
egions (Agos inho e al. 2016), o en a anged in cascading sequences along majo
i e sys ems (Ga cia e al. 2018a). O e ime, hese a i icial ecosys ems ha e sim-
pli ied ish communi ies axonomically and unc ionally, leading o bio ic homo-
genisa ion (Vi ule e al. 2012; Daga e al. 2020; Fe az e al. 2021a). Amongs he
main d i e s o hese changes is he es ablishmen and p oli e a ion o non-na i e
ish species, which o en ou compe e o displace na i e species, al e ing ophic
dynamics and ecosys em unc ions (Pelicice and Agos inho 2006; Ganassin e al.
2021). While local en i onmen al ac o s such as ese oi mo phology, low e-
gime, wa e e en ion ime and land use also shape ish assemblages (Noguei a e
al. 2012; Fe a eze e al. 2014; Koushlesh e al. 2023), non-na i e species in o-
duc ions emain one o he mos pe asi e ecological dis u bances in Neo opical
ese oi s (Vi ule e al. 2012; Daga e al. 2020).
Small-bodied ish species make up he la ges po ion o he B azilian ich hyo-
auna, ye hey emain la gely “in isible” o he public and a e, he e o e, highly
h ea ened and poo ly s udied (Cas o and Polaz 2020). Many o hese species
ha e adap ed o ese oi s h ough speci ic ep oduc i e s a egies (e.g. sho li e
cycles, high ecundi y, non-mig a o y beha iou and ex ended spawning e en s)
and plas ic eeding habi s, including gene alis , omni o ous o oppo unis ic die s
(Agos inho e al. 2016). While na i e small-bodied species a e well adap ed o luc-
ua ing en i onmen al condi ions, non-na i e species wi h simila ai s can suc-
cess ully in ade and es ablish popula ions, o en leading o compe i ion o ood
and habi a (Ja duli e al. 2021; Fe az e al. 2024). Ma ginal zones, which p o ide
c i ical shel e , spawning g ounds and eeding a eas (Casa i e al. 2003; Pelicice
and Agos inho 2006), a e pa icula ly ulne able o in asions. Habi a s uc u e,
such as mac ophy e beds, d i wood and lea -li e ed o ocky subs a es, in luenc-
es species co-exis ence and ophic in e ac ions (Fonseca e al. 2022; San iago e al.
2022). Howe e , when non-na i e species exploi hese habi a s, hey may displace
na i e species, al e esou ce a ailabili y and dis up na u al eeding dynamics.
Al hough small-bodied species a e gene alis s capable o colonising ese oi s
(Agos inho e al. 2016), hei die s a e in luenced by seasonal a ia ions in p ey
a ailabili y (Qui ino e al. 2015, 2017). Consequen ly, die composi ion shi
h oughou he yea (Ne es e al. 2018; Fonseca e al. 2022; San iago e al. 2022)
is d i en by en i onmen al condi ions and esou ce a ailabili y (Vido o-Magnoni
and Ca alho 2009; Bennemann e al. 2011). The p esence o non-na i e species
u he complica es hese dynamics by in ensi ying esou ce compe i ion, o cing
na i e species o b oaden o specialise hei die a y niches (Ba os e al. 2017; Sán-
chez-He nández e al. 2017). Unde s anding how non-na i e and na i e small-bod-
ied ish in e ac in ese oi ecosys ems is essen ial o p edic ing po en ial ecologi-
cal consequences, pa icula ly in egions acing inc easing biological in asions.
The Pa anapanema Ri e , one o he main ibu a ies o he Uppe Pa aná Ri e
(Sampaio 1944), has been hea ily agmen ed by 11 cascading ese oi s (Ga cia e
al. 2018a). I s cou se is di ided in o h ee main sec ions: Uppe , Middle and Lowe
Pa anapanema (Sampaio 1944). The cons uc ion o he Rosana Rese oi (Lowe
s e ch) in 1986 c ea ed a 190 km s e ch wi h di e se habi a s including ma ginal la-
goons, o es ed ma gins and nume ous ibu a ies. Howe e , his s e ch was u he
agmen ed in 1991 by he Taqua uçu Dam (Casimi o e al. 2017). Al hough Rosana
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
and Taqua uçu Rese oi s a e pa o he Pa anapanema cascade (Ga cia e al. 2018b)
and sha e he same clima ic condi ions (Al a es e al. 2013; Te assi e al. 2018), hei
dis inc habi a s uc u es may lead o di e en ish communi y esponses.
Rosana Rese oi ea u es ma ginal lagoons wi h mac ophy es beds and o es -
ed su oundings (Cassa i e al. 2003; Pelicice and Agos inho 2006) and ecei es
in low om impo an ibu a ies (Agos inho e al. 2007). These habi a s p o ide
abundan ood esou ces o small-bodied ishes and a e closely linked o seasonal
a ia ions, being s ongly in luenced by ain all, looding and empe a u e (Qui i-
no e al. 2015; Fonseca e al. 2022; San iago e al. 2022). In con as , Taqua uçu
Rese oi lacks ma ginal lagoons and o es ed ma gins and has ew, human-im-
pac ed ibu a ies, o e ing ewe ood esou ces o ma ginal-zone species e en un-
de seasonal we -d y a ia ion (Vido o-Magnoni e al. 2015; Ga cia e al. 2018b).
Gi en he inc easing p e alence o non-na i e ish in Neo opical ese oi s, his
s udy aims o: (i) examine he die and eeding ecology o small-bodied species,
wi h a ocus on in e ac ions be ween na i e and non-na i e species ac oss ese oi
and seasons (we s. d y) and (ii) compa e eeding pa e ns o assess how habi-
a condi ions in luence ophic dynamics. We posi ha : (i) in Rosana Rese oi ,
whe e habi a complexi y and ood esou ce a ailabili y a e high, na i e species will
display na owe die a y niches and lowe o e lap wi h non-na i e species, main-
aining s onge seasonal esponses in p ey selec ion and (ii) in Taqua uçu Rese -
oi , whe e he habi a complexi y is lowe and esou ces a e sca ce , bo h na i e
and non-na i e species will exhibi b oade die a y niches and highe die o e lap,
wi h weake seasonal a ia ion in die . By highligh ing ophic in e ac ions be ween
na i e and non-na i e small-bodied ish, ou s udy p o ides insigh in o he ecolog-
ical consequences o species in oduc ions in ese oi ecosys ems and con ibu es
o a be e unde s anding o ood web al e a ions in agmen ed Neo opical i e s.
Ma e ials and me hods
S udy a ea
The Pa anapanema Ri e ises in he A lan ic Pla eau, in he Municipali y o Capão
Boni o, S a e o São Paulo (SP) (Sampaio 1944). As one o he main le -bank ib-
u a ies o he Uppe Pa aná Ri e Basin, i lows o 930 km, wi h 330 km o i s
main channel o ming he bo de s be ween sou h-eas e n SP and no he n Pa aná
(PR) (Maack 1981). The Lowe Pa anapanema Ri e begins downs eam o Sal o
G ande Falls, which is now subme ged by he Sal o G ande Rese oi . F om he e,
i lows o a se ies o ese oi s, including Sal o G ande, Canoas II, Canoas I,
Capi a a, Taqua uçu and Rosana (Duke Ene gy 2008). The las wo ese oi s, he
Taqua uçu and Rosana, a e he ocus o his s udy (Fig. 1).
The Rosana Hyd oelec ic Powe Plan (Rosana Dam) is loca ed be ween he mu-
nicipali ies o Diaman e do No e (PR) and P ima e a (SP). The Rese oi ope a es as
a un-o - i e low, wi h a leng h o 110 km, a maximum dep h o 26 m and wi h a
looded a ea o 220 km2 (Fe a eze e al. 2014). I s sinuous cou se closely esembles he
o iginal loodplain shape o he Uppe Pa aná Ri e Basin, acili a ing he o ma ion
o ma ginal lagoons along i s su oundings (Agos inho e al. 2007). Subme ged and
loa ing mac ophy es, including Eichho nia spp., Elodea sp., Sagi a ia sp. and Sal inia
sp., a e widely dis ibu ed h oughou he Rese oi (Casa i e al. 2003; Pelicice and
Agos inho 2006). Two conse a ion uni s bo de he Rese oi : Mo o do Diabo S a e
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
Pa k (SP) on he igh and he Caiuá Ecologic S a ion (PR) on he le . The Rese oi
ecei es in lows om he Pi apó Ri e (le bank), Pi apozinho Ri e ( igh bank) and
h ee main s eams - Iancã, Cuiabá and Boni o - all on i s igh bank (Fig. 1).
The Escola Poli écnica Hyd oelec ic Powe Plan (Taqua uçu Dam) is loca ed
be ween he municipali ies o I aguajé (PR) and Sando alina (SP). The Rese oi
has a un-o - i e sys em, wi h a leng h o 80 km, maximum dep h o 18 m and a
looded a ea o 105.5 km2 (B i o and Ca alho 2006). Unlike Rosana Rese oi ,
Taqua uçu ollows a ela i ely s aigh cou se wi h minimal meande ing, limi ing
he o ma ion o ma ginal lagoons and educing looding po en ial. As a esul ,
subme ged mac ophy es a e spa se (Vido o-Magnoni e al. 2015). The main ib-
u a ies o Taqua uçu Rese oi include he Capim, Cen ená io and Tenen e Ri e s
on he le bank and he Anhumas Ri e on he igh bank, he la e being he
only one o unde go a e o es a ion p ocess (Mosqui o Fo es ) (Leme e al. 2015)
(Fig. 1). Despi e his, mos o he Rese oi ’s su oundings a e domina ed by ag i-
cul u al and pas u e land (Rod igues e al. 2019), while i s ibu a ies ace mul iple
an h opogenic p essu es, including ipa ian de o es a ion, u banisa ion and he
discha ge o domes ic and indus ial e luen s (Vido o-Magnoni e al. 2015).
The Pa anapanema Ri e expe iences wo p onounced seasons, a d y season (Ap il
o Sep embe ) and a we season (Oc obe o Ma ch), The annual mean empe a u e
is 17 °C ( ange 13 o 22 °C and 1550 mm as annual p ecipi a ion ha is ocused in
he we season (Al a es e al. 2013; Te assi e al. 2018). Ai empe a u e da a om he
da abase o Na ional Ins i u e o Me eo ology (INMET) e ealed he Rosana Rese -
oi anged om 19 °C (Augus 2019) o 27.2 °C (Decembe 2020). Simila ai em-
pe a u e da a we e una ailable o he Taqua uçu Rese oi . P ecipi a ion da a em-
phasised he di e ences be ween he d y and we season, whe e da a ex ac ed om
he da abase o he Ins i u e o Wa e s o Pa ana S a e (ANA) e ealed he Rosana
Rese oi had no p ecipi a ion in July 2018 and Augus 2019, bu a mon hly maxi-
mum o 250 mm in Decembe 2019. The Taqua ucu Rese oi had no p ecipi a ion
in Augus 2019 and a mon hly maximum o 315 mm in Decembe 2019 (Fig. 2).
Figu e 1. Sampling si es in he Rosana and Taqua uçu Rese oi s, lowe Pa anapanema Ri e . Hy-
d oelec ic powe plan s: 1 = Rosana; 2 = Taqua uçu; 3 = Capi a a. MS = S a e o Ma o G osso do
Sul; PR = S a e o Pa aná; SP = S a e o São Paulo.
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
Fish sampling
Samples o ish we e collec ed qua e ly be ween Sep embe 2018 and Sep embe
2020 a se en si es ac oss bo h Rese oi s (Fig. 1). The samples we e collec ed us-
ing awls, sie es and cas ne s om he ege a ion in he ma ginal zone and along
aqua ic mac ophy e beds, when p esen . A each si e, a sampling e o o wo
hou s was used, co e ing a s e ch o 100 m and ho oughly explo ing he a ail-
able mic ohabi a s wi hin he a ea. All cap u ed indi iduals we e anaes he ised and
eu hanised by o e exposu e o 1 g ml-1 Eugenol, hen ixed in 10% o maldehyde
o 48 hou s be o e being s o ed in 70% alcohol. Sampling was conduc ed unde
app o al o The Animal E hics Commi ee o he Uni e sidade Es adual de Lon-
d ina (CEUA N° 21149.2012.53; CEUA N° 24310.2017.78). In he labo a o y,
ish species we e iden i ied using keys in li e a u e (O a e al. 2018), wi h ouche
specimens (Suppl. ma e ial 1: able S1) and we e deposi ed a he Museu de Zoo-
logia da Uni e sidade Es adual de Lond ina (MZUEL).
Die analysis
In labo a o y, indi iduals we e dissec ed and hei s omachs emo ed, wi h he
s omach con en s analysed unde a s e eomic oscope. P ey i ems we e iden i ied
o he lowes possible axonomic le el wi h he help o speci ic li e a u e (Thomaz
e al. 2002; Mugnai e al. 2010; Biolo and Rod igues 2011). The olume (V) o
Figu e 2. Tempe a u e (Rosana Rese oi ) and p ecipi a ion om Rosana and Taqua ucu Rese oi
om he s udy pe iod. A. Pa anapoema Me eo ological S a ion (Rosana Rese oi ); B. Po eca u Me-
eo ological S a ion (Taqua uçu Rese oi ).
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
each o he s omach con en s was measu ed by comp ession in a millime e Pe i
dish, wi h he olume eco ded in mm3 and la e con e ed o ml (Hellawell and
Abel 1971). The p ey i ems we e g ouped in o 10 ca ego ies and ep esen ed in
pe cen age (%), as ollows: Algae (Filamen ous Algae); Aqua ic Insec s (Aqua ic
Insec F agmen , Coleop e a La ae, Coleop e a Pupa, Dip e a La ae, Dip e a
Pupe, Epheme op e a Nymph, Odona a Nymph and T ichop e a Nymph); De-
i us (Ino ganic and O ganic De i us); Fish (Fish, Fish Scale); Mac oc us acean
(C ab, Sh imp); Mic oc us acean (Cladoce a, Copepoda, Mic oc us acean F ag-
men , Os acoda); Mollusc (Bi al ia, Gas opoda); O he (Mic oplas ic); Te es-
ial Plan (F ui , Lea , S ick, Seed) and Te es ial In e eb a es (Aca i, A anae,
Bla odea (Isop e a), Bla odea Adul , Coleop e a Adul , Diplopoda, Dip e a
Adul , Epheme op e a Adul , Hemip e a Adul , Hymenop e a Adul , Lepidop e a
Adul , Te es ial Insec F agmen and Thysanop e a Adul . The p ey ca ego ies
we e o ganised in o ood esou ces, based on hei o igin and we e la e sepa a ed
by ese oi and season, wi h bo h being ep esen ed as pe cen age (%). The p ey
ca ego ies we e classi ied in o wo g oups: Au och honous (Aqua ic Insec ; Mic o-
c us acean; Mac oc us acean; Mollusc; Fish, Algae and De i us) and alloch ho-
nous (Te es ial in e eb a es; Te es ial Plan ; and O he s).
P ey ca ego ies we e ep esen ed ac oss ese oi s and la e sepa a ed by season
acco ding o he ‘ equency o occu ence’ (FO) me hod, wi h adap a ions om
Hamidan e al. (2016). To calcula e he equency o occu ence (de ined as he
pe cen age o s omachs in which a pa icula p ey ca ego y occu ed), he ollowing
o mula was used: %Fi = (ni/n) x 100, whe e: ni = numbe o s omachs con aining
p ey ca ego y and n = o al numbe on s omachs analysed. Nex , we calcula ed he
ampli ude and die a y niche o e lap o he species be ween he we and d y sea-
sons. To assess he niche ampli ude o species, we used he Shannon-Wiene Index
(Shannon 1948): H’ = -∑ pk x ln pk, whe e H’ means he Shannon-Wiene niche
wid h measu e, pk is he p opo ion o indi iduals collec ed using he k esou ce
and ln is he nepe ian loga i hm o he pk alue. Die a y niche o e lap be ween
species was calcula ed using he Pianka Index: Ojk = (n∑i=j|Pij - Pik|) / n∑i=j Pij
2. n∑i=j
Pik
2, whe e Ojk = measu e o Pianka die a y niche o e lap be ween species j and
species k; pij = p opo ion o p ey ca ego y i in he o al o p ey ca ego ies used by
species j; pik = p opo ion o p ey ca ego y i in he o al o p ey ca ego ies used by
species k, n = o al numbe o p ey ca ego ies. The esul s o Ojk we e conside ed as
ollows: low o e lap (< 0.4), in e media e o e lap (0.4–0.6) o high o e lap (> 0.6)
(G ossman 1986). Due o he sample size limi a ions, i was no possible o include
he species H. ma gina us in he seasonal analyses o Rosana Rese oi .
Da a analysis
A pe mu a ional mul i a ia e analysis o a iance (PERMANOVA) (Ande son
e al. 2008) was used o es o di e ences in ish die s wi hin each ese oi ,
based on a B ay-Cu is simila i y ma ix o olume da a (log ans o med, x + 1).
The esul ing pseudo-F s a is ic was es ed using he Mon e Ca lo me hod wi h
999 andomisa ions. Species pai s ha di e ed signi ican ly we e iden i ied using
pai wise pos -hoc compa isons, based on he adonis2() unc ion ( egan package;
Oksanen e al. (2020)). Species pai s ha di e ed signi ican ly we e iden i ied us-
ing a pai wise adonis es . Nex , we isualised he o dina ion o species acco ding
o hei die composi ion (niche b ead h) using a p incipal coo dina e analysis
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
(PCoA), based on a B ay-Cu is dissimila i y ma ix (B ay and Cu is 1957). To
es o seasonal di e ences in species die s, we applied PERMANOVA again, us-
ing a B ay-Cu is simila i y ma ix o log ans o med olume da a (x + 1) and he
pseudo-F s a is ic was es ed by he Mon e Ca lo me hod using 999 andomisa-
ions. Pos -hoc pai wise compa isons we e ca ied ou using he adonis2() ame-
wo k o iden i y signi ican di e ences amongs species. A PCoA was also used o
isualise he o de ing o popula ions acco ding o die composi ion, based on he
B ay-Cu is dissimila i y ma ix (B ay and Cu is 1957). Due o he sample size
limi a ions, i was no possible o include he species H. ma gina us in he seasonal
analyses o Rosana Rese oi . Ins ead, only he o al abundance o his species in
he Rese oi was used. All he analyses we e pe o med using R P og amming
so wa e e sion 3.5.3 (R Co e Team 2024), wi h he “ egan” package (Oksanen e
al. 2020 and he “ggplo 2” package (Wickham 2016). Fo each species g oup pe
ese oi , he ollowing we e applied: he comple ed analyses o s omach con en s
( o al and be ween seasons), equency o occu ence o p ey, niche b ead h and
niche o e lap. Addi ionally, di e ences be ween species die in he same ese oi
and be ween seasons we e e i ied using PERMANOVA.
Resul s
Sample composi ion by species
A o al o 416 indi idual ish we e sampled om se en species wi hin he ami-
lies Aces o hamphidae and Cha acidae (Cha aci o mes) ac oss bo h Rese oi s,
including h ee non-na i e and ou na i e species. O he non-na i e species, 96
we e Megalamphodus eques (S eindachne , 1882), 60 we e Aphyocha ax den a us
Eigenmann & Kennedy and 36 we e Roeboides descal adensis Fowle , 1937. The
na i e anges o hese non-na i e species a e Pa aguay and lowe Pa aná Ri e
Basins o A. den a us and R. descal adensis and Amazon, Guapo é and Pa aguay
Ri e Basins o M. eques. The in oduc ions o A. den a us and R. descal adensis
likely ollowed he looding o he Sal o das Se e Quedas a e he cons uc ion o
he I aipu Hyd oelec ic Powe Plan , which allowed ups eam dispe sal o Lowe
Pa aná ich hyo auna, while M. eques was p obably in oduced h ough aqua ium
eleases (Ga cia e al. 2018a). Amongs he na i e species, 75 we e As yanax lacus-
is (Lü ken, 1875), 57 we e Hemig ammus ma gina us Ellis, 1911 and 58 we e
Se apinnus no omelas (Eigenmann, 1915).
In Rosana Rese oi , A. den a us, A. lacus is, M. eques, H. ma gina us and S. no-
omelas we e analysed. In Taqua uçu Rese oi , he same i e species we e eco ded,
oge he wi h M. in e media and R. descal adensis, o alling 12 popula ions ac oss
bo h Rese oi s; ep esen a i e images o each species a e p esen ed in Fig. 3.
Fish die by ese oi and season
Amongs all iden i ied p ey i ems, 20 we e au och honous and 18 we e alloch-
honous. In Rosana Rese oi , he s omach con en s e ealed a p edominance
o e es ial in e eb a es (alloch honous esou ces), complemen ed by o he
die a y componen s (Suppl. ma e ial 1: able S2). Con e sely, in Taqua uçu
Rese oi , die s we e cha ac e ised mainly by aqua ic insec s, mic oc us aceans
and ish (Suppl. ma e ial 1: able S3).
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Rega ding seasonal pa e ns, species in Rosana Rese oi showed consis en
seasonal shi s, eeding mo e on e es ial insec s du ing he we season and
inc easing consump ion o aqua ic insec s, mic oc us aceans and de i us (all
au och honous) in he d y season (Suppl. ma e ial 1: able S2). In con as ,
species in Taqua uçu Rese oi displayed less consis en seasonal pa e ns, wi h
die s du ing he we season including aqua ic insec s, algae (au och honous)
and e es ial plan s (alloch honous), while in he d y season, e es ial in-
sec s, mic oc us aceans and ish we e mo e equen p ey i ems.
Die a y pa e ns o na i e and non-na i e species
The non-na i e A. den a us consumed mainly e es ial in e eb a es and mic o-
c us aceans in bo h Rese oi s, wi h addi ional algae and e es ial plan s du ing
he we season in Taqua uçu (Suppl. ma e ial 1: ables S2, S3). The na i e A. la-
cus is p ima ily ed on e es ial plan s in Rosana. Non-na i e M. eques exhibi -
ed a di e se die , domina ed by e es ial insec s, aqua ic in e eb a es and mi-
c oc us aceans, inco po a ing algae and de i us seasonally in Rosana. The na i e
H. ma gina us mainly consumed e es ial in e eb a es and mic oc us aceans in
bo h Rese oi s and added plan s du ing he we season in Taqua uçu. The na-
i e, M. in e media, had a b oad die , eeding on e es ial in e eb a es, aqua ic
insec s and mic oc us aceans and expanded i s die o algae and plan s in he d y
Figu e 3. A. Aphyocha ax den a us 45 mm (MZUEL20735); B. As yanax lacus is 43 mm (MZU-
EL20735); C. Hemig ammus ma gina us 26 mm (MZUEL20773); D. Megalamphodus eques 28 mm
(MZUEL20732); E. Moenkhausia in e media 55 mm (MZUEL20790); F. Roeboides descal adensis 60
mm (MZUEL20772); G. Se apinnus no omelas 41 mm (MZUEL20736). Ex ac ed om Gelle (2021).
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
season o Taqua uçu (Suppl. ma e ial 1: able S3). The non-na i e R. descal aden-
sis p e e ed ish (mainly scales), bu also consumed e es ial in e eb a es and
aqua ic insec s ac oss seasons in Taqua uçu (Suppl. ma e ial 1: able S3). The na-
i e S. no omelas p ima ily ed on de i us in Rosana, showing seasonal a ia ion,
wi h highe in ake o e es ial in e eb a es, aqua ic insec s, mic oc us aceans
and e es ial plan s in Taqua uçu (Suppl. ma e ial 1: ables S2, S3).
F equency o occu ence, niche b ead h and o e lap by ese oi and season
F equency o occu ence (FO) a ied be ween ese oi s and seasons. In Rosana
Rese oi , e es ial in e eb a es, e es ial plan s and de i us we e he mos e-
quen ly consumed esou ces. Te es ial i ems domina ed du ing he we season,
while aqua ic insec s and mic oc us aceans we e mo e common in he d y season
(Suppl. ma e ial 1: able S4). In Taqua uçu Rese oi , pa e ns we e mo e a iable,
wi h high FO alues o e es ial in e eb a es, aqua ic insec s, mic oc us aceans
and ish, bu no consis en seasonal end (Suppl. ma e ial 1: able S5).
Niche b ead h (H’) also di e ed amongs species, ese oi s and seasons.
In Rosana Rese oi , non-na i e A. den a us had he na owes niche in bo h
he we (H’ = 0.66) and d y (H’ = 0.67) seasons, indica ing die a y specialisa-
ion. In con as , non-na i e M. eques had he b oades niche in he we season
(H’ = 1.61), while he na i e A. lacus is showed he la ges b ead h in he d y sea-
son (H’ = 1.37). In Taqua uçu Rese oi , H. ma gina us had he na owes niche
in he we season (H’ = 0.63), whe eas S. no omelas had he b oades (H’ = 1.38).
Du ing he d y season, A. lacus is exhibi ed he smalles niche (H’ = 0.72), while
M. in e media had he la ges (H’ = 1.68).
The Pianka Index e ealed clea seasonal shi s in die a y o e lap. In Rosana,
o e lap was highe in he we season, whe eas in he Taqua uçu, i was g ea e
in he d y season (Table 1). Non-na i e species exhibi ed subs an ial o e lap
wi h na i es, pa icula ly in Taqua uçu. Fo ins ance, A. den a us o e lapped
s ongly wi h H. ma gina us (Ojk = 0.99) and M. in e media (Ojk = 0.68) in
he d y season, while M. eques o e lapped wi h M. in e media (Ojk = 0.75),
sugges ing po en ial in e speci ic compe i ion.
Die a y a ia ion amongs na i e and non-na i e species
PERMANOVA indica ed signi ican die a y di e ences amongs species in bo h
Rese oi s. In Rosana Rese oi , die composi ion a ied signi ican ly amongs he
i e species, while in he Taqua uçu, di e ences we e obse ed amongs all se -
en species. Pos -hoc pai wise adonis es con i med signi ican di e ences be ween
mos pai s o species, excep o A. den a us and H. ma gina us (Table 2).
PCoA o dina ions sepa a ed species acco ding o die a y composi ion, wi h Axis
1 and Axis 2 explaining 99.7% o o al a iance (Fig. 4). In Rosana, non-na i e
M. eques and na i e S. no omelas had b oade niche sizes (posi i e side o Axis 2),
while A. lacus is, H. ma gina us and A. den a us had na owe niches (nega i e
side o Axis 2) (Fig. 4). In Taqua uçu, A. den a us, M. in e media and S. no omelas
displayed a b oade niche (nega i e side o Axis 2), H. ma gina us was cen ally
dis ibu ed and M. eques occupied he nega i e side (Fig. 4).
When assessing seasonal e ec s, PERMANOVA de ec ed signi ican die a y di -
e ences be ween we and d y seasons o all species in Rosana (Table 3). In he
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
2003), as well as la al aqua ic insec s and algae (Pelicice and Agos inho 2006). I s
die composi ion ends o luc ua e seasonally in esponse o esou ce a ailabili y
(Qui ino e al. 2015). In ou s udy, M. eques main ained his b oad die a y niche
and equen ly o e lapped wi h o he species, including he mo e specialised A.
lacus is. Such o e lap could in ensi y compe i ion and po en ially h ea en he
pe sis ence o na i e species wi h mo e es ic ed die s.
The na i e H. ma gina us was ano he species ha showed a ia ions in niche
b ead h be ween ese oi s, while main aining a consis en p e e ence o e es ial
insec s and mic oc us aceans in bo h sys ems. Species o en become mo e special-
ised when hei p e e ed ood esou ces a e eadily a ailable (Sánchez-He nández
e al. 2017). Fo ins ance, in he Mogi-Guaçu Ri e (sou h-eas e n B azil), H. ma -
gina us has been desc ibed as p ima ily insec i o ous (F agoso-Mou a e al. 2017),
al hough i may shi owa ds omni o y in mo e deg aded habi a s (Ba e o e al.
2018). These indings align wi h ou esul s: in Rosana Rese oi , H. ma gina us
showed a na ow niche cen ed on e es ial in e eb a es and mic oc us aceans,
while in Taqua uçu Rese oi , i b oadened i s die o include aqua ic insec s and
e es ial plan s, likely as a esponse o seasonal esou ce luc ua ions. The o e -
lap in die a y composi ion wi h non-na i es, such as A. den a us and M. eques,
as well as he na i e A. lacus is, sugges po en ial o ophic in e ac ions ha
could lead o compe i i e p essu e – especially in Taqua uçu Rese oi , whe e ood
esou ce limi a ions a e mo e p onounced. Despi e his, a p e ious s udy epo -
ed good body condi ion amongs hese popula ions in bo h Rese oi s (Fe az e
al. 2021b), indica ing ha , a leas o now, ood esou ces may be su icien o
suppo co-occu ence. Howe e , sus ained o e lap wi h compe i i ely dominan
non-na i e species could pose longe - e m isks o na i e popula ions.
The na i e M. in e media and he non-na i e R. descal adensis we e only
su icien ly abundan o analysis in he Taqua uçu Rese oi , which limi s
u he in e p e a ion. Moenkhausia in e media demons a ed a b oad die a y
niche composed o mul iple p ey ypes, wi h a p e e ence o e es ial in e e-
b a es and aqua ic insec s – simila o Tie ê Ri e (Vido o-Magnoni and Ca -
alho 2009; Smi h e al. 2018). Howe e , he species also consumed no able
amoun s o mic oc us aceans, algae and e es ial plan s ac oss seasons, a gen-
e alis and oppo unis ic eeding s a egy p e iously documen ed in ese oi
en i onmen s (Bennemann e al. 2011), posi ioning M. in e media as a s ong
compe i o - pa icula ly wi h co-occu ing non-na i es like A. den a us and
R. descal adensis, as well as wi h he b oadly o e lapping M. eques. In con as ,
R. descal adensis displayed a na ow, specialis niche ocused on ish scales, a
beha iou ypical o lepidophagous species. While membe o his genus a e
usually scale-ea e s a ge ing la ge ish, hey may also oppo unis ically con-
sume o he a ailable esou ces (Alb ech e al. 2013). Ou esul s a e simila o
hose o Casa i e al. (2003), who epo ed consump ion o e es ial in e -
eb a es and aqua ic insec s, bu di e ge om Pelicice and Agos inho (2006),
who desc ibed a mic oc us acean-based die . These disc epancies could be a -
ibu ed o spa ial a ia ion, di e en ial use o mic ohabi a s (e.g. mac ophy e
beds wi hin ma ginal lagoons) and pa icula ly he noc u nal eeding habi s
o R. descal adensis (Pelicice and Agos inho 2006), making die o e lap wi h
o he species o lesse conce n, gi en he empo al di ision o eeding ac i i y.
The na i e S. no omelas exhibi ed an unusual die a y pa e n, cha ac e ised by
a b oad niche and a die domina ed by de i us in he Rosana Rese oi , which
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
con as s wi h p e ious s udies in he a ea (which epo ed a specialised algi -
o ous die ) (Casa i e al. 2003; Pelicice and Agos inho 2006). In Taqua uçu
Rese oi , S. no omelas showed eeding pa e ns mo e aligned wi h p e ious
s udies, including he consump ion o e es ial in e eb a es, aqua ic insec s,
mic oc us aceans, algae and plan s ac oss he seasons (Casa i e al. 2003; Pe-
licice and Agos inho 2006; San iago e al. 2022). I is impo an o no e ha
so ma e ials, such as algae and animal issue, deg ade apidly du ing diges ion
(Ga cia e al. 2018b), po en ially leading o an o e - ep esen a ion o mo e
esilien i ems such as de i us. Addi ionally, he species’ ben hic eeding be-
ha iou may con ibu e o he inciden al inges ion o sedimen s (San iago e
al. 2022). The species exhibi ed signi ican die a y o e lap wi h he non-na i e
A. den a us and M. eques, as well as wi h he na i e M. in e media. This o e lap
highligh s he po en ial o in e speci ic compe i ion, pa icula ly wi h non-na-
i e species ha display high ophic plas ici y and compe i i e ad an age.
Ou esul s ha e implica ions o he ongoing global eshwa e biodi e si y c isis,
which is d i en by habi a al e a ion, in asi e species, hyd ological egula ion and
clima e change (Ha ison e al. 2018; Albe e al. 2020). Small-bodied ishes, such
as cha acoids in his s udy, a e o en igno ed in conse a ion p io i ies; howe e , hey
play c i ical oles in ulne able opical eshwa e ecosys ems. They ha e speci ic
ea u es, such as high unc ional di e si y and sho gene a ion imes and hei sen-
si i i y o en i onmen al change makes hem ea ly indica o s o ecosys em deg ada-
ion and bio ic homogenisa ion (Da wall e al. 2018; Dudgeon and S aye 2025).
Ou indings documen ing ophic in e ac ions and esou ces pa i ioning be-
ween na i e and non-na i e cha acoids clea ly indica e how e en sub le shi s
a he lowe ophic le els can p opaga e h ough ood webs and al e ecosys em
s abili y. In an a emp o “bend he cu e” o eshwa e biodi e si y loss, hese
indings suppo in e na ional calls o imp o e in eg a i e, p ocess-based assess-
men s o in asion impac s (Tickne e al. 2020; O oni e al. 2023, 2025). E ec-
i e conse a ion and managemen mus ake in o accoun bo h he p esence o
non-na i e species and he ways in which local en i onmen al il e s shape hei
ecological e ec s. This is demons a ed by con ex -dependen ou comes, such as
hose ound be ween Rosana and Taqua uçu Rese oi s.
In conclusion, we epo no el insigh s in o he unde s udied in e ac ions be-
ween na i e and non-na i e small-bodied ishes co-exis ing in he ma ginal zones
o Neo opical ese oi s. By analysing he die s o se en na i e and non-na i e
om he Lowe Pa anapanema Ri e , we p o ide an impo an upda e on ish
ophic ecology in he egion. No able die a y di e ences we e obse ed be ween
species ac oss ese oi s and seasons, likely d i en by en i onmen al a iabili y and
eeding plas ici y. In he Rosana Rese oi , ish elied mo e hea ily on alloch ho-
nous esou ces and esponded consis en ly o seasonal changes – pa e ns likely
acili a ed by he p esence o ipa ian habi a s ha p omo e e es ial-aqua ic ex-
change. In con as , ish in he Taqua uçu Rese oi we e es ic ed o au och-
honous esou ces and exhibi ed uncoo dina ed seasonal esponses, likely due o
he lack o habi a he e ogenei y. The na owe niche b ead hs and lowe die a y
o e lap in Rosana sugges g ea e ophic specialisa ion and seg ega ion, whe eas
he b oade niches and highe o e lap obse ed in Taqua uçu –especially amongs
non-na i e species du ing he d y season – may e lec ood limi a ion, leading
o esou ce sha ing and gene alis s a egies. These indings a e pa icula ly con-
ce ning, as hey highligh he po en ial o nega i e in e ac ions in sou ce-poo
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
en i onmen s, po en ially esul ing in popula ion declines o local ex inc ions o
na i e species. Gi en he ecological impo ance o small-bodied ishes in media ing
ene gy low and nu ien cycling, we emphasise he need o ongoing moni o ing
o hese species wi hin ese oi ma ginal zones.
Acknowledgemen s
We would like o hank China Th ee Go ges Co po a ion (CTG B asil) o g an -
ing inancial assis ance o his wo k. Addi ionally, we would like o hank Apa-
ecido de Souza, Edson San ana da Sil a and Lucas Ribei o Ja duli o helping
wi h he ieldwo k and Thiago De uza Ga cia o helping wi h analysis and Iago
Vinicios Gelle o helping wi h species pho og aphs. Finally, we acknowledge he
cons uc i e and insigh ul commen s om he Academic Edi o Josie Sou h and
e iewe Felipe O oni, which subs an ially imp o ed he manusc ip .
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
This s udy was unded by Coo denação de Ape eiçoamen o de Pessoal de Ní el Supe io (CAPES),
B azil – inance code 001.
Au ho con ibu ions
JDF: concep ualisa ion, da a collec ion, da a analysis, w i ing; ACRC: da a collec ion, e iew;
DAZG: da a collec ion, e iew; APVM: da a analysis, e iew; ALBM: concep ualisa ion, e iew;
AST: concep ualisa ion, da a analysis, e iew; JRB: concep ualisa ion, da a analysis, e iew; MLO:
concep ualisa ion, da a analysis, e iew.
Au ho ORCIDs
João Daniel Fe az h ps://o cid.o g/0000-0003-1346-1642
A mando Césa Rod igues Casimi o h ps://o cid.o g/0000-0001-8826-5609
Diego Aze edo Zoccal Ga cia h ps://o cid.o g/0000-0001-5709-6347
Ana Paula Vido o-Magnoni h ps://o cid.o g/0000-0003-1819-7019
And é Lincoln Ba oso de Magalhães h ps://o cid.o g/0000-0002-9463-1836
Ali Se han Ta kan h ps://o cid.o g/0000-0001-8628-0514
John Robe B i on h ps://o cid.o g/0000-0003-1853-3086
Má io Luís O si h ps://o cid.o g/0000-0001-9545-4985
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex o Supplemen a y
In o ma ion.
197
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João Daniel Fe az e al.: In e ac ions be ween non-na i e and na i e cha acids
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Supplemen a y ma e ial 1
Supplemen a y in o ma ion
Au ho s: João Daniel Fe az, A mando Césa Rod igues Casimi o, Diego Aze edo Zoccal Ga cia,
Ana Paula Vido o-Magnoni, And é Lincoln Ba oso de Magalhães, Ali Se han Ta kan, John Rob-
e B i on, Má io Luís O si
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commons.o g/licenses/odbl/1.0/). The Open Da abase License (ODbL) is a license ag eemen
in ended o allow use s o eely sha e, modi y, and use his Da ase while main aining his same
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