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Non-na i e eshwa e ishes in India: exis ing e idence and
knowledge gaps
Lohi h Kuma 1,2,3 , Flo ian Ruland1,2,4 , Jona han M. Jeschke1,2
1 Leibniz Ins i u e o F eshwa e Ecology and Inland Fishe ies (IGB), Müggelseedamm 310, 12587 Be lin, Ge many
2 Ins i u e o Biology, F eie Uni e si ä Be lin, Königin-Luise-S . 1-3, 14195 Be lin, Ge many
3 ICAR-Cen al Inland Fishe ies Resea ch Ins i u e, Ba ackpo e, India
4 Wes Iceland Na u e Resea ch Cen e, Ha na go u 3, 340 S ykkisholmu , Iceland
Co esponding au ho : Lohi h Kuma ([email p o ec ed])
Copy igh : © Lohi h Kuma 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).
Re iew A icle
Abs ac
India has expe ienced he in oduc ion o nume ous non-na i e ish species (NNF), some o which
ha e caused ecological and economic impac s. This sys ema ic e iew p o ides a cu en ly lacking
o e iew o NNF esea ch in India, po en ial biases, a ailable e idence, and knowledge gaps. Using
he P e e ed Repo ing I ems o Sys ema ic Re iews and Me a-Analyses (PRISMA) amewo k, we
iden i ied a o al o 332 eco ds, documen ing he p esence o 58 NNF dis ibu ed ac oss 17 basins,
and 19 ansloca ed species in India. The Ganga was he mos s udied basin (113 s udies), ollowed by
he Wes Flowing Ri e s Tad i o Kanyakuma i basin (37 s udies), howe e , wi h 30 NNF epo ed
om each o hese basins. We demons a e how hese esul s can be due o sa u a ed sampling in he
Ganga and iden i y which basins migh be cu en ly unde s udied. We also illus a e how ex eme
loods p ecipi a ed an inc ease in NNF in o i e s and lakes om con inemen in he Wes Flowing
Ri e s Tad i o Kanyakuma i basin. The common ca p, Cyp inus ca pio, was he mos equen ly e-
po ed NNF (160 imes), while Mozambique ilapia, O eoch omis mossambicus, was he mos widely
dis ibu ed NNF (13 basins). We ound he e is a g owing numbe o publica ions in he ield, bu
ha up o 40% o s udies ha e appea ed in po en ially p eda o y jou nals. A mino i y o s udies
(44%) in es iga ed NNF impac s, mos o which used da a om he li e a u e (58%) and epo ed
only quali a i e impac s (69%). Mos documen ed impac s we e ecological (79%), while some we e
socio-economic (11%) o bo h (10%). Only 18% o he s udies add essed NNF managemen . The
knowledge syn hesized and he gaps iden i ied in his s udy migh se e as a basis o u u e s udies
and be use ul o e icien ly alloca ing limi ed esou ces o in es iga ing NNF in India.
Key wo ds: Asia, eshwa e ecosys ems, in ade impac s, in asi e alien ishes, managemen o
in asi e alien species, spa ial dis ibu ions
In oduc ion
While he esea ch ield o in asion biology has la gely ocused on e es ial ax-
onomic g oups (Pyšek e al. 2008; Jeschke and Hege 2018), eshwa e ishes a e
one o he mos ex ensi ely in oduced and impac ul g oups (Lep ieu e al. 2008;
Cuche ousse and Olden 2011; Ma colin e al. 2025). A o al o 1451 non-na i e
ish species (NNF) a e known o ha e es ablished globally in no el en i onmen s
(Seebens e al. 2023), and many o hese ha e become in asi e, i.e., ha e caused
Academic edi o : Filipe Ribei o
Recei ed:
10 Janua y 2025
Accep ed:
25 Ap il 2025
Published:
7 Oc obe 2025
Ci a ion: Kuma L, Ruland F, Jeschke
JM (2025) Non-na i e eshwa e
ishes in India: exis ing e idence and
knowledge gaps. In: Anas ácio P,
Ribei o F, Chainho P (Eds) In asions
in Aqua ic Sys ems. NeoBio a 102:
419–440. h ps://doi.o g/10.3897/
neobio a.102.146421
NeoBio a 102: 419–440 (2025)
DOI: 10.3897/neobio a.102.146421
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 102: 419–440 (2025), DOI: 10.3897/neobio a.102.146421
Lohi h Kuma e al.: Non-na i e ishes in India
dele e ious impac s on na i e biodi e si y, ecosys em se ices o human economy
and well-being ( ollowing he de ini ion o he In e na ional Union o Conse a-
ion o Na u e; IUCN 2000). Fo ins ance, he in asi e Nile pe ch, La es nilo icus,
caused he ex inc ion o 200 na i e endemic ish species in Lake Vic o ia, A ica,
esul ing in i e e sible damage o he lake’s ecosys em and a ec ing he li elihoods
o ishe men ( his example and o he s we e e iewed by Be ne y e al. 2022; Bach-
e e al. 2023). Fu he , an es ima ed 37 billion USD in economic losses ha e been
caused globally by 27 in asi e non-na i e ish species (Haub ock e al. 2022).
While mos esea ch on NNF has ocused on weal hy egions like No h Ame -
ica, Eu ope and Aus alia (Olden e al. 2008; Be ne y e al. 2022; Ma colin e al.
2025), we he e ocus on India, which has also expe ienced many NNF in oduc-
ions. I is a as coun y wi h 4% o he ea h’s eshwa e esou ces sp ead ac oss
25 i e basins (India-WRIS 2012) and hos ing abou 936 na i e eshwa e ish
species (AqGRISI 2024), many o which a e endemic (Dahanuka e al. 2004;
Singh and Sa ma 2015). The i s documen ed NNF in India was he b own ou ,
Salmo u a, which was in oduced in 1863 as a spo ish o angling (Singh and
Lak a 2011). Subsequen ly, he e we e in en ional and unin en ional in oduc-
ions o many NNF h ough a ious pa hways, such as o namen al ade, aqua-
cul u e and biological con ol (in oduc ion pa hways o India we e e iewed by
Singh and Lak a 2011), whose es ima ed numbe s oday a y be ween 288 and
633 (Ghosh e al. 2003; Singh 2021a; Sandilyan 2022). Fou een NNF ha e been
classi ied as in asi e in India, ollowing he c i e ia sugges ed by he Con en ion
on Biological Di e si y 2002 (Sandilyan e al. 2019).
Va ious aspec s o eshwa e NNF ha e been ex ensi ely s udied in India,
encompassing hei p esence (Ku up e al. 2006; Das e al. 2013; Gu umayum
2021), popula ion ends (Kuma 2000; Ragha an e al. 2008), biology (Raj e
al. 2021b; Singh e al. 2021), impac s (e.g. decline in na i e ish ca ch in i e
Ganga; Singh e al. 2013), and s a egies o manage hem (e.g. decla ing Hima-
layan i e s’ headwa e as in asion e ugia o na i e ishes; Sha ma e al. 2021).
The e has also been esea ch on NNF as po en ial disease ec o s (T ipa hi e al.
2022) and on hea y me als in NNF including con amina ion, bioaccumula ion
and use o NNF in oxicological s udies (Singh e al. 2012). O he s udies ex-
amined he pe o mance o NNF in aquacul u e (e.g. common ca p, Cyp inus
ca pio; Manna and Som 1982; Das e al. 2021) and o biological con ol (e.g.
wes e n mosqui o ish, Gambusia a inis; Chak abo y e al. 2008; Ghosh e al.
2011). Mo eo e , se e al e iews ha e p o ided insigh s in o a ious aspec s o
NNF (Sa ka e al. 2012; Singh 2021a; Sandilyan 2022). Despi e he ex ensi e
esea ch conduc ed in hese a eas, a comp ehensi e syn hesis in eg a ing hese
s udies is cu en ly lacking which can suppo e idence-based decision-making
and help p io i ize managemen ac ions (Cook e al. 2013).
This s udy aims o p o ide such a e iew on NNF in India, compiling s udies
on NNF ac oss all eshwa e basins in India. Speci ically, we p o ide an o e iew
o he a ailable ypes o s udies ha di ec ly o indi ec ly in es iga ed NNF in In-
dia, hei de elopmen o e ime and in which scien i ic jou nals (non-p eda o y
s. po en ially p eda o y) hey we e published. Fu he , we iden i y he NNF o
which mos esea ch s udies we e ca ied ou in India and p o ide a map showing
he dis ibu ion o NNF co ec ed o basin a ea and esea ch e o in India. We
also analyze de e minan s o he numbe o NNF and show how hese numbe s
ha e changed h ough ime o di e en basins and India as a whole. Addi ionally,
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Lohi h Kuma e al.: Non-na i e ishes in India
we epo on he p opo ion o s udies add essing di e en ypes o NNF impac s
and he le el o collabo a ion be ween execu ing agencies, as well as he in luence
o unding on he assessmen o po en ial impac s. Finally, we compa e ecom-
mended managemen s a egies and hei comp ehensi eness.
Me hods
Da a collec ion
We ga he ed eco ds o NNF in India up o he end o 2022 ollowing he P e-
e ed Repo ing I ems o Sys ema ic Re iews and Me a-Analyses (PRISMA) o
ecology and e olu iona y biology (O’Dea e al. 2021). We scanned li e a u e om
he Google Schola and Web o Science da abases using he English sea ch s ing:
“In asi e” OR “Exo ic” OR “Alien” OR “In oduced” OR “Non-na i e” OR
“Nonna i e” OR “Non-indigenous” OR “Nonindigenous”) AND “India” AND
“Fish*”. A o al o 2305 s udies we e iden i ied o which 332 s udies we e consid-
e ed ele an o ou s udy (see Suppl. ma e ial 1: able S1 o de ails). A s udy was
conside ed ele an i i deal wi h eshwa e NNF (excluding s udies on ma ine
and es ua ine NNF) in he geog aphical ex en o India.
Fo each o he 332 iden i ied ele an s udies, we compiled he ollowing in o -
ma ion: (1) au ho name(s); (2) yea o publica ion; (3) name o jou nal o o he
ype o publica ion; (4) ype o a icle (pee - e iewed o po en ially p eda o y a i-
cles in scien i ic jou nal, book/book chap e , semina p oceedings o magazine a i-
cle); (5) he ocal s udy a ea (“checklis ”: documen ed di e si y o ish, “biology”: bi-
ology o NNF, “ ish heal h”: disease/pa hogens and hei sp ead, “aquacul u e”: ish
p oduc ion and enhancemen including nu i ion, “ e iew”: e iew pape s, “im-
pac ”: impac o NNF in ecipien ecosys em, “managemen ”: managemen o NNF,
“ ishe y”: ishe y o NNF, “ oxici y”: bioaccumula ion and e ec o pes icides and
hea y me als, “ epo s”: epo ing o p esence o ange ex ension o NNF, “s a us”:
popula ion end o NNF; “biocon ol e iciency”: es ing e iciency o NNF o dis-
ease con ol; and “o he s”: o namen al ade/da abase/supply chain managemen /
use o DNA in axonomy and pos -ha es /p ocessing); (6) in which i e basin he
s udy was ca ied ou ( ield s udies) o he ishes we e collec ed (labo a o y s ud-
ies); (7) he ocal NNF and/o ansloca ed species; (8) quali a i e o quan i a i e
impac ; (9) ype o impac (expe imen al, obse a ional o in e ed); (10) di ec ion
o impac ; (11) any acknowledged unding; (12) au ho a ilia ions; and (13) any
sugges ed managemen measu es (see below and Suppl. ma e ials 3, 4 o de ails).
We consul ed se e al sou ces o de e mine i an a icle was published in a po-
en ially p eda o y jou nal. Beall’s lis con ains po en ially p eda o y jou nals o
publishe s (h ps://beallslis .ne /), whe eas Cla i a e’s Jou nal Ci a ion Repo
(h ps://jc .cla i a e.com/jc /b owse-jou nals) and Na ional Academy o Ag icul-
u al Science (NAAS), India (h ps://naas.o g.in/NJS/jou nals2022.pd ), publish
lis s o pee - e iewed jou nals e e y yea . The e o e, we ca ego ized a s udy as po-
en ially p eda o y i he espec i e jou nal was lis ed in Beall’s lis and i i was no
lis ed in he epo s o Cla i a e’s Jou nal Ci a ion Repo o NAAS.
As i e basins (also known as ca chmen a eas o wa e sheds) o m he na u al
biogeog aphical ba ie s o eshwa e ishes (Lep ieu e al. 2008), we no ed in
which i e basins each s udy was ca ied ou . To do so, we ex ac ed bounda ies o
25 i e basins p o ided by Hyd oBASINS (Lehne and G ill 2013) based on he
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Lohi h Kuma e al.: Non-na i e ishes in India
i e basin bounda ies es ablished by he Go e nmen o India (India-WRIS 2012)
(h ps://indiaw is.go .in/w is/#/a las). Depending on he geog aphic loca ion ha
was in es iga ed in a s udy, i was assigned o he espec i e i e basin. Labo a o y
s udies we e assigned o i e basins i he in o ma ion on he loca ion o sample
collec ion was a ailable; i no , eco ds we e classi ied as labo a o y s udies wi hou
in o ma ion on i e basin. Reco ds whe e he s udy a eas we e men ioned as po-
li ical bounda ies (s a es o coun y) o eco- egions (Indian No heas e n Region,
Indian Himalaya, Wes e n Gha s) we e classi ied sepa a ely, and he same was ue
o s udies co e ing mul iple i e basins.
We no ed which non-na i e o ansloca ed ish species we e men ioned in each
s udy. We de ined non-na i e ish species (NNF) as hose ha ha e been in o-
duced o India by human ac ion, ei he in en ionally o unin en ionally ( ollowing
Blackbu n e al. 2011; Jeschke e al. 2022; synonyms o “non-na i e” a e “alien”,
“exo ic” o “non-indigenous”); and we de ined ansloca ed ish species as hose
ha a e na i e o India bu ha e been in oduced wi hin India by human ac ion,
ei he in en ionally o unin en ionally, o one o mo e i e basins ou side o hei
na i e ange. Since bo h La in and common English species names change o e
ime and synonyms exis , we s anda dized species names using Eschmeye ’s Ca a-
log o Fishes (F icke e al. 2024).
Rega ding NNF impac s, we disc imina ed quali a i e om quan i a i e im-
pac assessmen s. S udies ha epo ed impac s non-nume ically o desc ip i ely
( o example, non-na i e ishes g adually eplacing na i e ish auna in he middle
s e ch o he Cau e y i e ; Roshi h e al. 2022) we e labelled as quali a i e im-
pac s; whe eas s udies ha epo ed impac s nume ically ( o example, es ima es o
annual ish landings o non-na i e common ca p, Cyp inus ca pio and na i e ish
species in he Ri e Ganga, and i s compa ison wi h his o ical ish landings; Ray e
al. 2021) we e conside ed as quan i a i e impac s. Impac s we e desc ibed as eco-
logical, socio-economic, o bo h, and he di ec ion o impac as dele e ious, ben-
e icial, o bo h ( ollowing Jeschke e al. 2014). Finally, we classi ied he mode o
impac assessmen as ei he (i) obse a ional, impac s we e obse ed by he au ho s
( o example, bu ows dug by he Amazon sail in ca ish, P e ygoplic hys pa dalis, in
canal ma gins; (Raj e al. 2021b) o empo al a ia ion in ish ca ch composi ion
(Sa ka e al. 2012); (ii) expe imen al, impac s we e assessed by conduc ing expe -
imen s as in compa ing he e icacy o wo non-na i e la i o ous ish (Ghosh e
al. 2011); o (iii) in e ed, impac deduced om he li e a u e, based on he species
known o ha e impac s elsewhe e (Raj e al. 2020).
We also ga he ed in o ma ion on sugges ed managemen measu es. Robe son
e al. (2020) p oposed a amewo k o he managemen o biological in asions
wi h en key managemen e ms ha b oadly co e all ac i e managemen objec-
i es including he suppo o ac i e managemen (moni o ing, isk assessmen
e c.). Using hese e ms, we ca ego ized he managemen measu es sugges ed in
ou s udies (see Suppl. ma e ial 4 o de ails).
Wi h he esul ing da ase , we in es iga ed ends in NNF esea ch, p opo ion
o po en ially p eda o y a icles, numbe s o NNF, mos s udied NNF, mos s ud-
ied i e basins, i e basin wi h mos NNF. We also map he spa ial dis ibu ion o
NNF ac oss basins (co ec ed o basin a ea) and o he basins Ganga, B ahmapu-
a, EFR (Eas Flowing Ri e s) Penna o Kanyakuma i, and WFR (Wes Flowing
Ri e s) Tad i o Kanyakuma i, as well as collec i ely o India, we ch onologically
analyzed he ela ionship be ween he cumula i e numbe o NNF species and he
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Lohi h Kuma e al.: Non-na i e ishes in India
numbe o s udies conduc ed he e. Fu he , we examined pa e ns in impac as-
sessmen s, p opo ion o s udies in di e en ocal a eas, collabo a ion in esea ch,
and sugges ed managemen measu es.
S a is ical analyses
We analyzed di e ences in esea ch e o among i e basins. Some basins
ha e no been s udied a all. The e o e, we i s used a binomial Gene alised
Linea Model (GLM) o es i he p obabili y o a leas one s udy being con-
duc ed in a basin is dependen on i s size. We used he na u al loga i hm o
a ea (in km2) o each basin in he model:
[1] s udyConduc ed ~ log(basinA ea)
Then we used a linea model o see i , among he s udied basins, la ge basins
we e p io i ized. To ensu e no mal dis ibu ion o esiduals, bo h he numbe o
s udies and he basin a ea we e log- ans o med. The da a we e scanned o ou lie s
and he ollowing model was un wi h all basins as well as wi hou he Ganga:
[2] log(nS udies) ~ log(basinA ea)
A hi d model was used o de ec any possible ela ionship be ween basin a ea,
numbe o s udies and he numbe o de ec ed NNF in a basin. He e again, he
educed da ase was used co e ing only s udied basins and a ea and numbe o
s udies we e log- ans o med. The ull model consequen ly is:
[3] nNonNa i eFishSpecies ~ log(nS udies) + log(a ea) +log(nS udies):log(a ea)
All models wi h all pa ame e combina ions we e compa ed using he d edge
unc ion o he MuMIn package (Ba oń 2023). All pa ame e s ha appea ed in
he bes models wi h and accumula ed model weigh o > 0.95 we e conside ed.
The esul ing mos pa simonious model is gi en in he Resul s sec ion.
We also in es iga ed o he po en ial biases in ou s udies. Fo his, we pe o med
pai wise Chi-squa e es s wi h 100’000 boo s ap simula ions o es i he unding
sou ce (public o p i a e) has an in luence on he p edic ed impac o he in ade
(dele e ious o bene icial). Whe e no explici ly men ioned o he wise, all analyses
we e conduc ed using he R base package (R Co e Team 2020).
Resul s
The e has been an exponen ial g ow h in he numbe o publica ions on NNF in
India o e ime. The i s o he 332 iden i ied s udies is om 1957; howe e , he e
we e no publica ions in he nex decade (1961–70) and ewe han 10 publica ions
pe decade un il 2000. In he ollowing decades, he numbe o s udies accele a ed,
wi h a ound 50 publica ions om 2001 o 2010 and almos 190 om 2011 o 2020.
The e we e 73 publica ions in he i s wo yea s o he cu en decade (2021–2022).
P ojec ing his numbe o all 10 yea s o he cu en decade means he e will be an
es ima ed 365 publica ions om 2021 o 2030 (Fig. 1). This p ojec ion is conse a-
i e, as i does no assume a u he inc ease in he numbe s o publica ions pe yea .
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Lohi h Kuma e al.: Non-na i e ishes in India
Mos eco ds a e publica ions in scien i ic jou nals (309 eco ds o 93%), bu
41% o hose we e published in po en ially p eda o y jou nals. Fo s udies pub-
lished by au ho s a ilia ed wi h uni e si ies, e en he majo i y (54%) o publica-
ions in scien i ic jou nals we e published in po en ially p eda o y jou nals, whe e-
as i was abou a qua e (24%) o publica ions om non-uni e si y esea ch
ins i u es (see Suppl. ma e ial 2: ig. S1). Fo s udies wi h bo h uni e si ies and
non-uni e si y esea ch ins i u es as a ilia ions, he p opo ion o publica ions in
po en ially p eda o y jou nals was in be ween (35%).
Abou 38% o he eco ds examined NNF di ec ly, in es iga ing ei he hei
biology, impac , managemen , ishe y, epo , s a us o bio-con ol (see Suppl. ma-
e ial 2: ig. S2). A high numbe o s udies (31% o 103 eco ds) ocused on docu-
men ing ish di e si y (checklis ) whe e NNF we e oppo unis ic de ec ions, while
only 3% o he s udies p ima ily epo ed NNF (11 eco ds). Fish heal h (9%)
and aquacul u e (7%) a e wo o he impo an opics o he s udies in ou da ase ,
which also includes se e al e iews (6%).
F om ou eco ds, we de ec ed 58 NNF dis ibu ed ac oss 17 basins and 19
ansloca ed ishes in India (Tables 1, 2). O hese, 55 NNF we e epo ed om
open wa e s (na u al i e s and lakes as well as man-made ese oi s) in a leas
one basin. We we e able o assign 250 s udies (75%) o 17 i e basins, whe eas no
s udies we e ound o he emaining eigh i e basins (Table 3). The o he 25%
o he s udies e e ed o poli ical bounda ies a he han i e basins as s udy a ea
(13%), we e labo a o y s udies wi h no in o ma ion on whe e he ish sampling
loca ion (10%) o ocused on eco- egions o mul iple i e basins (3% combined).
Abou hal o he 250 s udies we e conduc ed in he Ganga basin (46% o 113
eco ds), making i by a he mos s udied i e basin in India ollowed dis an ly
by he basin Wes Flowing Ri e s Tad i o Kanyakuma i (15% o 37 eco ds). Sig-
ni ican ly, despi e his esea ch bias he e we e 30 NNF epo ed o each o hese
Figu e 1. Decadal end o publica ions in esea ch abou non-na i e ishes in India. The g ay box o he cu en decade indica es he
conse a i ely es ima ed numbe o publica ions om 2021 o 2030, assuming ha he publica ion a e obse ed in his decade’s i s wo
yea s (73 publica ions in 2021 and 2022) will be cons an un il he end o he decade.
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Lohi h Kuma e al.: Non-na i e ishes in India
Table 1. Lis o non-na i e ish species (NNF) in India epo ed in he s udies included in ou da ase .
Non-na i e ish species Non-na i e ish species
Os eoglossi o mes: A apaimidae Silu i o mes: Pangasiidae
A apaima gigas Pangasianodon hypoph halmus
Os eoglossi o mes: Os eoglossidae Cha aci o mes: Se asalmidae
Os eoglossum bici hosum Colossoma mac opomum
Lepisos ei o mes: Lepisos eidae Pia ac us b achypomus
A ac os eus spa ula Pia ac us mesopo amicus
Salmoni o mes: Salmonidae Pygocen us na e e i*
Onco hynchus mykiss † Beloni o mes: Ad ianich hyidae
Onco hynchus ne ka ‡‡ O yzias ja anicus
Salmo sala ‡‡ Cyp inodon i o mes: Poeciliidae
Salmo u a † Gambusia a inis* †
Sal elinus on inalis ‡‡ Gambusia holb ooki* †
Sal elinus namaycush ‡‡ Poecilia mexicana
Cyp ini o mes: Cyp inidae Poecilia e icula a
Ba bonymus al us Xiphopho us helle ii
Ba bonymus goniono us Xiphopho us macula us
Ca assius au a us Cichli o mes: Cichlidae
Ca assius ca assius Amphilophus imacula us
C enopha yngodon idella Andinoaca a i ula us §
Cyp inus ca pio* † As ono us ocella us
Cyp inus ub o uscus Cop odon zillii
Pun ius b e is Hemich omis bimacula us
Cyp ini o mes: Xenocyp ididae Mayahe os u oph halmus
Hypoph halmich hys moli ix O eoch omis au eus
Hypoph halmich hys nobilis* O eoch omis mossambicus* †
Mylopha yngodon piceus* O eoch omis nilo icus*
Cyp ini o mes: Tincidae O eoch omis u olepis §
Tinca inca Tho ich hys meeki §
Silu i o mes: Cla iidae Anaban i o mes: Channidae
Cla ias ga iepinus* Channa lucius
Silu i o mes: Ic alu idae Anaban i o mes: Helos oma idae
Ic alu us punc a us Helos oma emminckii
Silu i o mes: Lo ica iidae Anaban i o mes: Osph onemidae
Hypos omus plecos omus Mac opodus ope cula is
P e ygoplich hys amb ose ii* Osph onemus go amy
P e ygoplich hys disjunc i us* T ichopodus mic olepis
P e ygoplich hys mul i adia us* T ichopodus ichop e us
P e ygoplich hys pa dalis* T ichopsis i a a
* In asi e in India based on Sandilyan e al. (2019) who ollowed he de ini ion o he Con en ion o Biological
Di e si y (2002).
† Among he ‘100 o he Wo ld’s Wo s In asi e Alien Species’ (GISD 2024).
§ Repo ed om ish a ms bu no eco d om he wild in ou da ase .
‡‡ Non-na i e popula ion po en ially ex inc (Joshi and Lal 2017).
Table 2. Lis o ansloca ed ish species in India epo ed in he s udies included in ou da ase .
Basin(s) Repo ed ansloca ed ish species
5, 7, 9, 12, 19, 24, 25 Ci hinus m igala, Labeo ca la, Labeo ohi a
1Anabas cobojius, Anabas es udineus, Channa punc a a, Channa s ia a, Ci hinus m igala, Cla ias magu , Danio e io, Esomus
dan ica, He e opneus es ossilis, Labeo ca la, Labeo ohi a, No op e us no op e us, Pun ius sopho e
7Badis badis, Lepidocephalich hys gun ea, Os eob ama co io, Pe hia gelius, Pe hia phu unio
5, 9, 17, 24, 25 To khud ee
Basins: (1) Andaman Nicoba Islands, (5) Cau e y, (7) EFR Penna and Kanyakuma i, (9) Goda a i, (12) K ishna, (17) Na mada, (19) Penna , (24) WFR
Tad i o Kanyakuma i, (25) WFR Tapi o Tad i.
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Lohi h Kuma e al.: Non-na i e ishes in India
wo basins (Fig. 2A, B). The common ca p, Cyp inus ca pio, was he mos equen -
ly epo ed NNF (160 imes) ollowed by he Mozambique ilapia, O eoch omis
mossambicus (108 imes) (see Suppl. ma e ial 2: ig. S3A). Howe e , he la e was
he mos widely dis ibu ed NNF (13 basins) ollowed by Cyp inus ca pio (12 ba-
sins) (see Suppl. ma e ial 2: ig. S3B). Addi ionally, he e we e 24 NNF ha we e
epo ed om only one basin each (see Suppl. ma e ial 5 o de ails).
The log(n) o he basin a ea had no signi ican ela ionship wi h he p obabili y
o a leas one s udy being conduc ed in a basin (binomial GLM wi h p = 0.15).
Fu he , he numbe o s udies o s udied basins was co ela ed o he basin a ea
when using log-log ans o ma ion (linea model wi h adj. R2 = 0.18, p = 0.049),
bu his ela ionship was exclusi ely d i en by he la ges and mos s udied i -
e basin – he Ganga – and disappea s when i is emo ed (linea model wi h
adj. R2 = 0.02, p = 0.27). Resul s o he ull model (model [3] in he Ma e ial
and Me hods sec ion) show ha he numbe o de ec ed non-na i e ish species
is mainly ela ed o he numbe o s udies conduc ed in he espec i e basin, no
o i s a ea o he in e ac ion o s udies and a ea (see Suppl. ma e ial 1: able S6).
Resul s o he mos pa simonious model – he numbe o ish species in ela ion o
he log o he numbe o s udies (adjus ed R2 = 0.69, p < 0.001) – a e shown in Fig.
2C, D. This model is obus o emo ing he Ganga basin (R2 = 0.56, p < 0.001).
The i e basins in India displayed a a ying ex en o (appa en ) sa u a ion in
e ms o NNF de ec ions. The sa u a ion cu e almos la ened in he WFR Tad i o
Table 3. Lis o i e basins in India wi h he numbe o non-na i e ish species (NNF) epo ed o
each basin and he numbe o s udies.
Numbe Basin name A ea (km2) # NNF # S udies
1 Andaman Nicoba Islands 8249 12 4
2 Ba ak and O he s 41723 0 0
3 B ahmani and Bai a ani 51822 0 0
4 B ahmapu a 194413 12 17
5 Cau e y 81155 11 10
6 EFR Mahanadi and Penna 86643 0 0
7 EFR Penna and Kanyakuma i 100139 19 11
8 Ganga 861452 30 113
9 Goda a i 312812 10 7
10 Indus 321289 7 16
11 Inland D ainage in Rajas han 139917 0 0
12 K ishna 258948 11 16
13 Lakshadweep Islands 32 0 0
14 Mahanadi 141589 7 5
15 Mahi 34842 1 4
16 Mino Ri e s D aining in o Myanma and Bangladesh 36202 0 0
17 Na mada 98796 4 2
18 No h Ladakh no d aining in o Indus 26018 0 0
19 Penna 55213 6 1
20 Saba ma i 21674 2 1
21 Sube na ekha 29196 0 0
22 Tapi 65145 1 1
23 WFR Ku ch and Sau ash a 321851 2 1
24 WFR Tad i o Kanyakuma i 55177 30 37
25 WFR Tapi o Tad i 55940 8 4
NNF: Non-na i e ish species, EFR: Eas lowing i e , WFR: Wes lowing i e .
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Lohi h Kuma e al.: Non-na i e ishes in India
Kanyakuma i basin un il 2021, bu he e was an ab up ise in he numbe o de ec ed
NNF in 2022, which is also e iden o India as a whole (Fig. 3A, E). Wi h ewe addi-
ions o he NNF de ec ions in he B ahmapu a basin in he las 50% o s udies, i ap-
pea s ha mos NNF in his basin ha e been de ec ed (Fig. 3B). The de ec ion o new
NNF con inues o inc ease almos linea ly in he EFR Penna o Kanyakuma i basin
(Fig. 3C). In he Ganga basin, he de ec ion o NNF inc eased g adually and hen al-
mos pla eaued wi h ewe de ec ions o e e y addi ional s udy conduc ed (Fig. 3D).
Figu e 2. A numbe o s udies on non-na i e ish species (NNF) o each i e basin in India B numbe o epo ed NNF o each basin
C ela ionship be ween he numbe o epo ed NNF and he numbe o s udies o each basin (log- ans o med) (adjus ed R2 = 0.69,
p < 0.001) D map o esiduals om plo C: ed and blue colo s indica e highe and lowe numbe s o epo ed NNF o each basin con-
side ing he numbe o s udies, espec i ely. G ey colo indica es basins wi h no s udies (A), wi h no NNF epo ed (B) and ha we e no
conside ed o analysis due o he absence o s udies (D). Basins: (1) Andaman Nicoba Islands, (2) Ba ak and O he s, (3) B ahmani and
Bai a ani, (4) B ahmapu a, (5) Cau e y, (6) Eas Flowing Ri e (EFR) Mahanadi and Penna , (7) EFR Penna and Kanyakuma i, (8) Ganga,
(9) Goda a i, (10) Indus, (11) Inland D ainage in Rajas han, (12) K ishna, (13) Lakshadweep, (14) Mahanadi, (15) Mahi, (16) Mino Ri e s
D aining in o Myanma and Bangladesh, (17) Na mada, (18) No h Ladakh no d aining in o Indus, (19) Penna , (20) Saba ma i, (21) Su-
be na ekha, (22) Tapi, (23) Wes Flowing Ri e (WFR) Ku ch and Sau ash a, (24) WFR Tad i o Kanyakuma i, (25) WFR Tapi o Tad i.
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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.
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Supplemen a y ma e ial 1
PRISMA and models
Au ho s: Lohi h Kuma , Flo ian Ruland, Jona han M. Jeschke
Da a ype: docx
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License (h p://openda a-
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
eedom o o he s, p o ided ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/neobio a.102.146421.suppl1
Supplemen a y ma e ial 2
Supplemen a y images
Au ho s: Lohi h Kuma , Flo ian Ruland, Jona han M. Jeschke
Da a ype: zip
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License (h p://openda a-
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
eedom o o he s, p o ided ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/neobio a.102.146421.suppl2
440
NeoBio a 102: 419–440 (2025), DOI: 10.3897/neobio a.102.146421
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Supplemen a y ma e ial 3
Reco ds used and in o ma ion ex ac ed
Au ho s: Lohi h Kuma , Flo ian Ruland, Jona han M. Jeschke
Da a ype: xlsx
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in ended o allow use s o eely sha e, modi y, and use his Da ase while main aining his same
eedom o o he s, p o ided ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/neobio a.102.146421.suppl3
Supplemen a y ma e ial 4
Managemen measu es sugges ed in eco ds
Au ho s: Lohi h Kuma , Flo ian Ruland, Jona han M. Jeschke
Da a ype: xlsx
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eedom o o he s, p o ided ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/neobio a.102.146421.suppl4
Supplemen a y ma e ial 5
Spa ial dis ibu ion o NNF
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Da a ype: xlsx
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