A global synthesis of naturalised and invasive plants in aquatic habitats

473
A global syn hesis o na u alised and in asi e plan s
in aqua ic habi a s
Alessand a Ko z1, Ma in Hejda1, Jan Čuda1, Za ah Pa ison2, Jose B ůna3, Ana No oa1,4 ,
Jan Pe gl1, Pa el Pipek1,5 , Ka eřina Š aje o á1, Paulina Anas asiu6, Michael Ansong7,
Ma ga i a A ianou sou8, Julie F. Ba celona9, Sunee a Bha a1,5 , Fa zaneh Bo dba 10 ,
Is ael Bo okini11 , Lau a Celes i-G apow12 , Edua do Chacón-Mad igal13,14 , Wayne Dawson15 ,
Do jee16 , F anz Essl17 , Lilian Fe u ino-Acos a18 , Es ela Figuei edo19 , Rodol o Flo es20 ,
Guillaume F ied21 , Nicol Fuen es22 , Pablo Galán23 , Ch is ian Gilli24 , Michael Glase 17 ,
José Ramón G ande Allende25 , Zigman as Gudžinskas26 , Rachael Holmes27 , Philip E. Hulme28 ,
Inde ji 29 , Eun Su Kang30 , Holge K e 31 , Dan W. K ix32 , Ingol Kühn33 , Oma Lopez34,35 ,
AnaLu MacVean36 , T objon Makhkamo 37,38 , Elizabe e Ma chan e39 , Hélia Ma chan e40 ,
Al ed Ma oyi41 , Rachid Meddou 42 , Pie e Mee s43,44 , Sha i A. Mukul45,46,47 , B ad R. Mu ay48 ,
Megan L. Mu ay48 , Daniel L. Nick en 49 , P ince E. No man50 , Ali Ome 17,51 , Anne e Pa zel 52 ,
Pie e B. Pelse 9, Joan Pino53,54 , Ma c Rie a53 , Dagobe o Rod íguez Delcid55 ,
Julissa Rojas-Sando al56 , Rose Ro chés-Ribal a53,57 , José Yade Sage h Ruiz-C uz58,59 ,
S epan Sena o 60 , Alexande N. Senniko 61 , Bha a Babu Sh es ha62 , Gideon F. Smi h19 ,
Sima Soh abi63 , Ba ba a Toka ska-Guzik64 , Ma k an Kleunen65,66,67 , Mon se a Vilà68,69 ,
Vik o ia Wagne 70 , Pa ick Weigel 31,71 , Ma en Win e 72 , Ayşe Yazlık73 , Elena Zyko a74 , Pe Pyšek1,5
1 Depa men o In asionEcology,Ins i u eo Bo any,CzechAcademyo Sciences,P ůhonice,CzechRepublic
2 Facul y o Na u al Sciences, Biological and En i onmen al Sciences, Uni e si y o S i ling, S i ling, Sco land, UK
3 Depa men o Geoecology,Ins i u eo Bo any,CzechAcademyo Sciences,P ůhonice,CzechRepublic
4 Es aciónExpe imen aldeZonasÁ idas,ConsejoSupe io deIn es igacionesCien í icas(EEZA-CSIC),Alme ía,Spain
5 Depa men o Ecology,Facul yo Science,Cha lesUni e si y,P ague,CzechRepublic
6 Facul yo Biology&Bo anicalGa denD.B andza,Uni e si yo Bucha es ,Bucha es ,Romania
7 Depa men o Sil icul u eandFo es Managemen ,Facul yo RenewableNa u alResou ces,KwameNk umahUni e si yo ScienceandTechnology,Kumasi,Ghana
8 Depa men o Ecology and Sys ema ics, Facul y o Biology, Na ional and Kapodis ian Uni e si y o A hens, A hens, G eece
9 Schoolo BiologicalSciences,Uni e si yo Can e bu y,Ch is chu ch,NewZealand
10 Uni e si éLib edeB uxelles,He ba ium,B ussels,Belgium
11 Depa men o Ecology, Mon ana S a e Uni e si y, Bozeman, USA
12 Depa men o En i onmen alBiology,SapienzaUni e si y,Rome,I aly
13 He ba ioNacional,MuseoNacionaldeCos aRica,SanJosé,Cos aRica
14 Cen odeIn es igaciónenBiodi e sidadyEcologíaT opical,Uni e sidaddeCos aRica,SanJosé,Cos aRica
15 Depa men o E olu ion, Ecology and Beha iou , Ins i u e o In ec ion, Ve e ina y and Ecological Sciences, Uni e si y o Li e pool, Li e pool, UK
16 Na ionalPlan P o ec ionCen e,Depa men o Ag icul u e,Thimphu,Bhu an
17 Di isiono BioIn asions,GlobalChange&Mac oecology,Depa men o Bo anyandBiodi e si yResea ch,Uni e si yo Vienna,Vienna,Aus ia
18 He ba ioTEFH,EscueladeBiología,Facul addeCiencias,Uni e sidadNacionalAu ónomadeHondu as,Tegucigalpa,Hondu as
19 RiaOli ie He ba ium,Depa men o Bo any,NelsonMandelaUni e si y,Gqebe ha,Sou hA ica
20 Depa amen odeBo ánicayHe ba ioPMA,Facul addeCienciasNa u ales,Exac asyTecnología,Uni e sidaddePanamá,Panamáci y,Panama
21 Plan Heal hLabo a o y,ANSES,Mon e ie -su -Lez,F ance
22 Depa amen odeBo ánica,Facul addeCienciasNa u alesyOceanog á icas,Uni e sidaddeConcepción,Concepción,Chile
23 LaLibe ad,ElSal ado
24 Di isiono Sys ema icandE olu iona yBo any,Depa men o Bo anyandBiodi e si yResea ch,Uni e si yo Vienna,Vienna,Aus ia
25 Depa amen o de Bo ánica, Facul ad de Ciencias, Uni e sidad de G anada, G anada, Spain
26 Labo a o yo Flo aandGeobo any,S a eScien i icResea chIns i u eNa u eResea chCen e,Vilnius,Li huania
Copy igh : © Alessand a Ko z e al.
This is an open access a icle dis ibu ed unde
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License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
NeoBio a 102: 473–494 (2025)
DOI: 10.3897/neobio a.102.151156
Ad ancing esea ch on alien species and biological in asions
A pee - e iewed open-access jou nal
NeoBio a
474
NeoBio a 102: 473–494 (2025), DOI: 10.3897/neobio a.102.151156
Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
27 Depa men o Geog aphy, Geology and he En i onmen , Uni e si y o Leices e , Leices e , UK
28 TheCen e o OneBiosecu i yResea ch,AnalysisandSyn hesis,Depa men o Pes -Managemen andConse a ion,LincolnUni e si y,Can e bu y,
NewZealand
29 Depa men o En i onmen alS udies,Cen e o En i onmen alManagemen o Deg adedEcosys ems(CEMDE),Uni e si yo Delhi,Delhi,India
30 Ko eaNa ionalA bo e um,Pocheon,Sou hKo ea
31 Depa men o Biodi e si y, Mac oecology & Biogeog aphy, Uni e si y o Gö ingen, Gö ingen, Ge many
32 NewSou hWalesRu alFi eSe ice,Sydney,Aus alia
33 Depa men o Communi yEcology,Helmhol zCen e o En i onmen alResea ch-UFZ,Halle,Ge many
34 Smi hsonianT opicalResea chIns i u e,PanamaCi y,Panama
35 In e -Ame icanIns i u e o GlobalChangeResea ch,PanamaCi y,Panama
36 En i onmen alHo icul u eDepa men ,Yo kCollegeo Pennsyl ania,Yo k,USA
37 Depa men o Bo anyandGene ics,Na ionalUni e si yo Uzbekis an,Tashken ,Uzbekis an
38 Depa men o Fo es yandLandscapeDesign,Tashken S a eAg a ianUni e si y,Tashken ,Uzbekis an
39 Cen e o Func ionalEcology–Science o People& hePlane ,Associa eLabo a o yTERRA,Depa men o Li eSciences,Uni e si yo Coimb a,
Coimb a,Po ugal
40 Resea chCen e o Na u alResou cesEn i onmen andSocie y(CERNAS),Poly echnicUni e si yo Coimb a,Coimb aAg icul u eSchool(ESAC),
Coimb a,Po ugal
41 Depa men o Bo any,Uni e si yo Fo Ha e,Alice,Sou hA ica
42 Depa men o Ag onomicSciences,FSBSA,MouloudMamme iUni e si y,TiziOuzou,Alge ia
43 Bo anic Ga den Meise, Meise, Belgium
44 Uni e si éLib edeB uxelles,B ussels,Belgium
45 Depa men o En i onmen andDe elopmen S udies,Uni edIn e na ionalUni e si y,Dhaka,Bangladesh
46 T opicalFo es sandPeopleResea chCen e,Uni e si yo  heSunshineCoas ,Ma oochydo e,Aus alia
47 Depa men o Ea h and En i onmen , Flo ida In e na ional Uni e si y, Miami, USA
48 Schoolo Li eSciences,Uni e si yo TechnologySydney,Ul imo,Aus alia
49 Plan Biology Sec ion, School o In eg a i e Plan Science, College o Ag icul u e and Li e Science, Co nell Uni e si y, I haca, USA
50 Sie aLeoneAg icul u alResea chIns i u e(SLARI),F ee own,Sie aLeone
51 Depa men o Fo es Managemen , Facul y o Fo es y, Uni e si y o Kha oum, No h Kha oum, Sudan
52 Vege a ionEcology,LandscapeA chi ec u e,Uni e si yo AppliedSciencesWeihens ephan,F eising,Ge many
53 Cen edeRece caEcològicaiAplicacionsFo es als(CREAF),Bella e a,Spain
54 BABVE,Uni e si a Au ònomadeBa celona,Bella e a,Spain
55 AsociaciónJa dínBo ánicoLalaguna,He ba ioLAGU,LaLibe ad,ElSal ado
56Ins i u eo  heEn i onmen &Depa men o Geog aphy,Sus ainabili y,Communi y,andU banS udies,Uni e si yo Connec icu ,S o s,
Connec icu , USA
57 Uni e si a deVic-Uni e si a Cen aldeCa alunya,Vic,Spain
58 Uni e sidaddeElSal ado ,Facul adMul idisciplina iaO ien al,Depa amen odeCienciasNa u alesyMa emá ica,SeccióndeBiología,SanMiguel,
El Sal ado
59 Fundación Na u aleza, San Sal ado , El Sal ado
60 127273,Moscow,Russia
61 Bo anicalMuseum,FinnishMuseumo Na u alHis o y,Uni e si yo Helsinki,Helsinki,Finland
62 Cen alDepa men o Bo any,T ibhu anUni e si y,Ka hmandu,Nepal
63 RiceResea chIns i u eo I an(RRII),Rash ,I an
64 Ins i u eo Biology,Bio echnologyandEn i onmen alP o ec ion,Facul yo Na u alSciences,Uni e si yo SilesiainKa owice,Ka owice,Poland
65 Ecology, Depa men o Biology, Uni e si y o Kons anz, Kons anz, Ge many
66 ZhejiangP o incialKeyLabo a o yo Plan E olu iona yEcologyandConse a ion,TaizhouUni e si y,Taizhou,China
67 ZhejiangKeyLabo a o y o Res o a iono DamagedCoas alEcosys ems,Schoolo Li eSciences,TaizhouUni e si y,Taizhou,China
68 Es aciónBiológicadeDoñana,EBD-CSIC,ConsejoSupe io deIn es igacionesCien í icas,Se illa,Spain
69 Depa amen o de Biología Vege al y Ecología, Uni e sidad de Se illa, Se illa, Spain
70 Depa men o BiologicalSciences,Uni e si yo Albe a,Edmon on,Canada
71Depa men o En i onmen alScience,RadboudIns i u e o BiologicalandEn i onmen alSciences(RIBES),RadboudUni e si y,Nijmegen,
Ne he lands
72 Ge manCen e o In eg a i eBiodi e si yResea ch(iDi )Halle-Jena-Leipzig,Leipzig,Ge many
73 Depa men o Plan P o ec ion,Facul yo Ag icul u e,DüzceUni e si y,Düzce,Tu kiye
74 No osibi sk,Russia
Co espondingau ho :Alessand aKo z([email p o ec ed])
475
NeoBio a 102: 473–494 (2025), DOI: 10.3897/neobio a.102.151156
Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
Abs ac
Global da abases ha e con ibu ed o ou unde s anding o alien, na u alised and in asi e plan species
dis ibu ions. S ill, he ole o species in asions in habi a s, speci ically in aqua ic habi a s, emains unde -
explo ed a he global scale. Acco dingly, a comp ehensi e global syn hesis o he s a us o plan in asions
in aqua ic habi a s has been missing. He e, we ocus on mac oecological pa e ns o na u alised non-in-
asi e and in asi e plan s in aqua ic habi a s using he ecen ly buil SynHab da abase. Amongs all he
plan eco ds compiled in SynHab, 592 a e assigned o aqua ic habi a s, o which 183 a e unique plan
axa ( u he e med ‘species’) belonging o 49 amilies. O he o al numbe o eco ds, 462 e e o axa
wi h na u alised non-in asi e occu ences and 130 o in asi e occu ences. The species pool analysed
he e e e s o 78 egions dis ibu ed ac oss all bo anical con inen s as de ined by he Wo ld Geog aphical
Scheme o Reco ding Plan Dis ibu ions. The numbe o na u alised non-in asi e aqua ic species is
simila ac oss di e en con inen s and biomes, bu T opical Asia had mo e and he Medi e anean zono-
biome had ewe in asi e species han expec ed. T opical Asia, Tempe a e Asia and A ica ha e he high-
es p opo ions o na u alised species ha ha e become in asi e, while ac oss con inen s, in asi e p opo -
ions we e highes o opical and sub opical zonobiomes. New Zealand, I aly and Cali o nia con ained
disp opo iona ely mo e na u alised species han expec ed, gi en he a ea co e ed by aqua ic habi a in
hose egions, whe eas Sou h Sudan, Papua New Guinea and Ky gyzs an had disp opo iona ely ewe
species. In pai wise dissimila i y compa isons, all con inen s had dis inc species composi ions ( om
0.73 o 0.92 o he Jacca d dissimila i y index) and so did zonobiomes (0.69 o 1.00). The high p opo -
ion o in asi e species in T opical Asia in compa ison wi h e es ial in asions in his egion, indica es
a g ea e suscep ibili y o wa me egions o aqua ic plan in asions. This may be exace ba ed by u he
na u alisa ions in he u u e, as da a om empe a e egions sugges a la ge pool o a ailable species.
Key wo ds: Mac ophy e in asion, plan in asion pa e ns, SynHab da abase
In oduc ion
Aqua ic habi a s a e pa icula ly p one o in asion by alien species due o se e al ac-
o s, including na u al dis u bance egimes (e.g. i e low a ia ion and lake wa e
le el luc ua ions), high le els o connec i i y and an h opogenic ac i i ies (e.g. dam-
ming o hyd opowe and wa e ex ac ion o ag icul u e) (Richa dson e al. 2007).
High le els o connec i i y, bo h na u al and an h opogenic (such as aw wa e ans-
e schemes and canals), make moni o ing and managing pa hways o in asion in
aqua ic habi a s di icul (Waine e al. 2025). In asi e aqua ic plan s (mac ophy es)
o en ac as au ogenic enginee s and p ima y p oduce s (Riccia di and MacIsaac
2010), i.e. hey ans o m eshwa e sys ems, change habi a s uc u e (Valley and
B emigan 2002; Ceschin e al. 2020) and educe wa e quali y by al e ing low and
p ima y p oduc i i y (Pe na and Bu ows 2005; Galla do e al. 2016; Sou h e al.
2016), esul ing in changes in local na i e di e si y (Schoole e al. 2006). In addi-
ion, p ojec ed accumula ions o na u alised alien plan species sugges as e inc eas-
es in aqua ic sys ems han in e es ial habi a s (Seebens e al. 2021), indica ing ha
u u e impac s a e likely o in ensi y. The se e i y o impac s caused by alien plan s
is magni ied because aqua ic habi a s a e ex emely di e se ela i e o he a ea hey
occupy, compa ed o o he habi a s (Román-Palacios e al. 2022).
The ecological impac s o in asi e alien plan species and managemen e o s
impose subs an ial economic cos s. F om 1975 o 2040, he documen ed global
cos s o in asi e mac ophy es a e p ojec ed o each US$ 32.8 billion (in 2017
dolla s, Macêdo e al. 2024). Howe e , his igu e likely ep esen s a signi ican
unde es ima ion due o da a limi a ions and geog aphical gaps in cu en esea ch
Academic edi o :
Ped o Anas ácio
Recei ed:
28 Feb ua y 2025
Accep ed:
8 July 2025
Published:
7 Oc obe 2025
Ci a ion: Ko z A, Hejda M, Čuda
J, Pa ison Z, B ůna J, No oa
A, Pe gl J, Pipek P, Š aje o á
K, Anas asiu P, Ansong M,
A ianou sou M, Ba celona JF,
Bha a S, Bo dba F, Bo okini
I, Celes i-G apow L, Chacón-
Mad igal E, Dawson W, Do jee,
Essl F, Fe u ino-Acos a L,
Figuei edo E, Flo es R, F ied G,
Fuen es N, Galán P, Gilli C, Glase
M, G ande Allende JR, Gudžinskas
Z, Holmes R, Hulme PE, Inde ji ,
Kang ES, K e H, K ix DW, Kühn I,
Lopez O, MacVean A, Makhkamo
T, Ma chan e E, Ma chan e H,
Ma oyi A, Meddou R, Mee s P,
Mukul SA, Mu ay BR, Mu ay ML,
Nick en DL, No man PE, Ome
A, Pa zel A, Pelse PB, Pino J,
Rie a M, Rod íguez Delcid D,
Rojas-Sando al J, Ro chés-Ribal a
R, Ruiz-C uz JYS, Sena o S,
Senniko AN, Sh es ha BB, Smi h
GF, Soh abi S, Toka ska-Guzik B,
an Kleunen M, Vilà M, Wagne
V, Weigel P, Win e M, Yazlık
A, Zyko a E, Pyšek P (2025) A
global syn hesis o na u alised
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habi a s. In: Anas ácio P, Ribei o
F, Chainho P (Eds) In asions in
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neobio a.102.151156
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NeoBio a 102: 473–494 (2025), DOI: 10.3897/neobio a.102.151156
Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
(Macêdo e al. 2024), c ucially conce ning he unde es ima ed cos s o in asi e
plan s (No oa e al. 2021). This aises addi ional conce ns abou he ex en o
which he economic cos s o in asi e plan s in aqua ic habi a s may be eliably
cap u ed. As such, upda ed knowledge o egional le els o in asion is c ucial o a
be e unde s anding o impac s and cos es ima es.
The Wo ld’s 100 wo s in ade s lis (Lowe e al. 2000) p o ides examples o
he ha m ul e ec s ha plan in asions can ha e on aqua ic habi a s. One o he
globally mos p ominen in ade s is wa e hyacin h (Pon ede ia c assipes Ma .,
synonym Eichho nia c assipes (Ma .) Solms), which is na i e o Sou h Ame ica
and has sp ead o o e 70 coun ies (Gezie e al. 2018). Pon ede ia c assipes is a
as -g owing loa ing mac ophy e ha o ms dense ma s on he wa e su ace, al-
e s habi a s uc u e and dis up s he ecological unc ioning o aqua ic ecosys ems
(Tobias e al. 2019). This species also has di ec socioeconomic impac s, including
educed na igabili y and ec ea ional quali y, es ic ed wa e supply o ag icul-
u e due o damaged pipe sys ems and educed access o ishing g ounds and boa -
ing (Ka e egga and S e ne 2009; Villamagna and Mu phy 2010). Simila ly, Azolla
iliculoides Lam., o en ound alongside Pon ede ia c assipes and Pis ia s a io es L.,
is a small (< 25 mm) loa ing mac ophy e ha impac s physical and chemical wa-
e ea u es, educes he ichness and biomass o mac ophy es, al e s zooplank on
composi ion and educes he su i al o amphibian la ae (Pine o-Rod íguez e al.
2021). Laga osiphon majo (Ridl.) Moss is a subme ged mac ophy e (Howa d-Wil-
liams and Da ies 1988) ha o ms dense unde wa e beds, ou compe ing na i e
mac ophy es in he ange i has in aded (Ma in e al. 2018).
Speci ic inhe en biological a ibu es (such as unc ional ai s) de e mine he
in asi eness o species and inc ease hei po en ial o become in asi e (Gio ia e
al. 2023) by in e ac ing wi h na i e bio a and he en i onmen (Pyšek e al. 2020).
Addi ionally, one o he mos impo an ac o s explaining he success o plan in-
asions is p opagule p essu e (e.g. Von Holle and Simbe lo 2005; Colau i e al.
2006; Cassey e al. 2018), which also applies o in asi e mac ophy es (Chadwell
and Engelha d 2008; Xie e al. 2013). High le els o connec i i y can acili a e he
sp ead o p opagules, inc easing dispe sal dis ances o in asi e plan s wi hin and be-
ween wa e bodies in aqua ic habi a s (Richa dson e al. 2007; Leu en e al. 2009).
Fo example, na u al lood e en s accele a e he dispe sal p ocess by acili a ing he
mo emen o p opagules wi hin ca chmen s (Gu nell e al. 2008; Čuda e al. 2017).
Clima e- ela ed changes o he low egime u he acili a e he sp ead o p opagules
by inc easing he equency o ex eme loods (Pa ison e al. 2017). In addi ion,
dams c ea e len ic habi a s wi h slow-mo ing wa e s, which a e p e e ed by Azolla,
Pis ia and Pon ede ia species. Gi en he expec ed inc ease in dam cons uc ion o
mi iga e clima e-d i en ain all educ ions, his may esul in enhanced p oli e a ion
o hese mac ophy es. Raw wa e ans e schemes (wa e di e sions) mo e la ge
olumes o eshwa e be ween ca chmen s ia complex in as uc u e ne wo ks, a-
cili a ing he long-dis ance sp ead o in asi e species (Waine e al. 2024a, b).
Resea ch on plan in asions in aqua ic habi a s is unde - ep esen ed compa ed
o e es ial habi a s, pa icula ly in he T opics (E angelis a e al. 2014; Ha el e
al. 2015; McKnigh e al. 2017), wi h ce ain species o axonomic g oups domi-
na ing he li e a u e (S e enson e al. 2023). He e, we u ilise he newly-de eloped
SynHab da abase (h ps://www.synhab.com/ he-p ojec ), which con ains in o -
ma ion on plan na u alisa ions and in asions in speci ic habi a ypes wo ldwide
(Pyšek e al. 2022; Dawson e al. 2025), o achie e a mo e balanced pe spec i e o
477
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Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
he o e all di e si y o na u alised plan s in aqua ic habi a s. We add ess his knowl-
edge gap by asking he ollowing ques ions: (i) Which con inen s and biogeog aph-
ical egions (zonobiomes) ha e he highes le els o na u alisa ion and in asion o
plan s in aqua ic habi a s? (ii) Which aqua ic na u alised non-in asi e and in asi e
plan s a e mos widely dis ibu ed? (iii) Does he ulne abili y o aqua ic habi a s o
na u alisa ions and in asions o plan s di e in empe a e s. opical egions? (i )
Which egions ( egions o s a es) ha bou mo e na u alised plan species in aqua ic
habi a s han expec ed? ( ) Which egions a e mo e simila in e ms o na u alised
plan species composi ion in aqua ic habi a s? ( i) Which na u alised aqua ic plan
species a e indica i e o con inen s and zonobiomes?
Ma e ials and me hods
Habi a classi ica ion and mac ophy e de ini ion
Habi a classi ica ion, adap ed om Hejda e al. (2015) o he SynHab da abase,
esul ed in 14 ca ego ies (Pyšek e al. 2022). The aqua ic habi a is a ca ego y de-
ined as “wa e bodies and s eams wi h subme ged and loa ing plan species”.
Mac ophy es can be de ined in se e al ways, wi h some de ini ions including mac-
oalgae (e.g. Chambe s e al. 2008); howe e , he e we es ic ou ocus o aqua ic
ascula plan s. Species we e assigned o he aqua ic habi a i hey exhibi ed g ow h
in wa e as subme ged o loa ing, including li o al species ha can cope wi h
long- e m looding, wi h oo s ancho ed below he wa e le el. Some species, such
as I is pseudaco us L., Eleocha is palus is (L.) Roem. & Schul ., Juncus e usus L.
and Jacobaea aqua ica (Hill) G.Gae n., B.Mey. & Sche b., can g ow in he li o al
zones o eshwa e o b ackish wa e bodies, as well as in mud. In such cases, hey
we e assigned o bo h aqua ic and we land habi a s in SynHab, o which only he
o me is deal wi h in he p esen pape . Species inhabi ing mud, bu in ole an o
p olonged looding, such as Lysimachia nummula ia L. we e excluded. Dimo phic
species, such as Al e nan he a philoxe oides (Ma .) G iseb., exhibi ing mud-dwell-
ing and li o al o ms, we e also a ilia ed wi h bo h aqua ic habi a s and we lands.
In such cases, species we e assigned o he aqua ic habi a i he habi a desc ip ions
explici ly con i med g ow h in a subme ged en i onmen o eshwa e lakes, i e s,
unning wa e s, coas al basins o s eams. Species ha g ow along i e s, bu no
in he wa e we e no included. We hus included also aqua ic ascula plan s ha
a e no exclusi ely aqua ic, bu ha a e impo an in ade s in he aqua ic habi a .
Da a acquisi ion
The da a on habi a a ilia ions o he SynHab da abase we e ex ac ed om li -
e a u e sou ces ( egional checklis s o alien lo as), whe e he e bal desc ip ion o
habi a s allowed unequi ocal assignmen o a gi en species (see Suppl. ma e ial 1:
able S1 o he comple e e e ences). I published sou ces lacked in o ma ion on
habi a a ilia ions, we in i ed he au ho s o o iginal publica ions o collabo a e
and p o ide habi a a ilia ions o SynHab. The SynHab p ojec leade s (PP, MH,
AK) assessed he ecei ed in o ma ion o ha monise he habi a classi ica ion wi h
o he da ase s and hen he inal habi a a ilia ions we e ag eed upon in collabo-
a ion wi h da a p o ide s. In o al, 78 egions (coun ies o s a es) om all con i-
nen s we e included (Suppl. ma e ial 1: able S1).

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Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
Each plan species was assigned he in asion s a us as ‘na u alised’ o ‘in asi e’
in he egion, ollowing he de ini ions o Richa dson e al. (2000) and Blackbu n
e al. (2011) ca; no e ha he species’ in asion s a us can a y ac oss di e en e-
gions. In asion s a us was assigned, based on in o ma ion in he GloNAF (Global
Na u alized Alien Flo a) da abase ( an Kleunen e al. 2015, 2019; Pyšek e al.
2017; Da is e al. 2025) o in e ed om he desc ip ion in he o iginal sou ce o
p o ided by da a con ibu o s o hei egions. In he con ex o he analyses de-
sc ibed below, he e m ‘na u alised’ e e s o all na u alised axa and is composed
o wo sepa a e subse s: ‘na u alised non-in asi e’ and ‘in asi e.’
Each axon name, as gi en in he o iginal sou ce, was s anda dised acco ding
o The Wo ld Checklis o Vascula Plan s (WCVP) da abase (h ps://powo.sci-
ence.kew.o g/abou -wc p; Go ae s e al. 2021) using he WCVP package o R
(B own e al. 2023). Accep ed names o he species we e used in he analyses o
enable species compa isons amongs egions. Fou axa we e iden i ied a he sub-
species le el, bu we e me ged wi h he co esponding species le el o analyses.
Habi a a ea
To quan i y he a ea o aqua ic habi a s wi hin each egion, g idded global land-co e
da a, wi h a spa ial esolu ion o 300 m, we e acqui ed om De ou ny e al. (2023).
These da a comp ised 37 land-co e classes, con o ming o he Uni ed Na ions Land
Co e Classi ica ion Sys em (UN-LCCS) (Di G ego io and Jansen 2005). We con e -
ed he yea 2019 as e o a shape ile and in e sec ed i wi h SynHab egions o u he
analysis. We excluded pe manen snow and ice classes, which esul ed in 35 classes
included in he analyses. Wa e su ace a ea o classes was summed o use in analyses.
Biogeog aphical a iables: con inen s and zonobiomes
Each o he 78 egions o which da a on na u alised plan species in aqua ic hab-
i a s we e a ailable was assigned o a con inen and a s a e o coun y (le els 1 and
4, espec i ely, o he TDWG Wo ld Geog aphic Scheme o Reco ding Plan Dis-
ibu ions; B ummi 2001): No he n Ame ica (da a a ailable o n = 13 egions);
Sou he n Ame ica, n = 11; A ica, n = 16; Eu ope, n = 16; Tempe a e Asia, n = 9;
T opical Asia, n = 11; and Aus alasia, n = 2. Only one eco d was sampled in he Pa-
ci ic, which was no conside ed in he analyses. Fu he , egions we e assigned o one
o he ollowing zonobiomes: I. T opical (equa o ial), n = 18 egions; II. T opical (sa-
annah), n = 15; III. Sub opical (a id), n = 7; IV. Medi e anean, n = 8; VI. Tempe -
a e (nemo al), n = 17 egions; VII. A id empe a e (con inen al), n = 4; VIII. Cold
empe a e (bo eal), n = 1; and n = 8 egions we e assigned as “mul iple” zonobiomes
(Chile, China, Flo es, Faial and San a Ma ia, Azo es, Flo ida, New Sou h Wales,
Russia Kos oma, Russia Middle Volga and Russia No osibi sk based on Wal e and
B eckle (1991). While some egions, ca ego ised as ‘mul iple’, may encompass he V.
Wa m Tempe a e Zonobiome, no egions we e exclusi ely assigned o i .
S a is ical analyses
To es whe he na u alisa ion and in asion in aqua ic habi a s di e amongs
con inen s and zonobiomes, we used gene alised linea models (GLM) wi h
Poisson dis ibu ion. Using s anda dized esiduals om gene alised linea
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Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
models (GLMs), we de e mined i obse ed species ichness was signi ican ly
highe o lowe han expec ed ac oss zonobiomes and con inen s. S a is ical
signi icance was assessed by compa ing hese esiduals o c i ical alues co e-
sponding o P < 0.05 (± 1.96), P < 0.01 (± 2.58) and P < 0.001 (± 3.29). Posi-
i e alues indica e highe and nega i e alues indica e lowe na u alised species
ichness han expec ed by chance.
The species-a ea ela ionship was calcula ed by i ing a linea model o he
numbe o all na u alised species eco ded and he a ea o aqua ic habi a in
each egion on a log-log scale. To compa e he dissimila i y o he egions in
e ms o hei na u alised species composi ion in aqua ic habi a s, we calcula ed
he Jacca d dissimila i y index using he “be apa ” package (Baselga and O me
2012; Baselga e al. 2023). To iden i y which species we e signi ican indica-
o s o each con inen and zonobiome, we used he “mul ipa ” unc ion o
he indica o alue analysis (IndVal) in he R package “indicspecies” (Du êne
and Legend e 1997; De Cáce es and Legend e 2009), calcula ed conside ing
g oups o egions wi hin each con inen o zonobiome. Complemen a y o he
species composi ion analysis, he indica o alue analysis de e mines g oups o
indica o species by measu ing hei associa ion wi h each g oup (con inen o
zonobiome) o combina ion o g oups and i does no explici ly conside cli-
ma e, local en i onmen o in oduc ion his o y.
All s a is ical analyses we e pe o med in R e sion 4.4.1 (R Co e Team 2024).
Resul s
Na u alised and in asi e species numbe s
O he 585 eco ds in aqua ic habi a s, 458 e e ed o na u alised, non-in asi e
occu ences and 127 o in asi e occu ences. The same species can be na u alised,
non-in asi e in one egion and in asi e in ano he ; hus, he ocus was on he
numbe o occu ences a he han he numbe o species. In o al, we eco ded
183 na u alised plan axa ( u he e e ed o as ‘species’ o simplici y) belonging
o 46 amilies and 88 gene a; o hese 183, he e we e 52 classi ied as in asi e in a
leas one egion. The 10 amilies wi h he la ges numbe o species we e Hyd o-
cha i aceae, Plan aginaceae, Cype aceae, Poaceae, A aceae, Alisma aceae, Onag a-
ceae, Nymphaeaceae, Pon ede iaceae and Sal iniaceae.
Na u alised aqua ic species occu ed in 78 egions on all con inen s (Suppl.
ma e ial 1: able S1). O he sampled egions, 34 (i.e. 43.6%) con ained in o -
ma ion on in asi e species. New Zealand had he la ges numbe o na u alised
species (n = 37), ollowed by Cali o nia and I aly (n = 29 each), F ance (n = 27),
Flo ida (n = 23), Chile and Vi ginia (n = 20). Species eco ded as na u alised in
a leas one egion we e na i e o Tempe a e Asia (n = 94), No he n Ame ica
(n = 87), T opical Asia (n = 81), A ica (n = 79), Sou he n Ame ica (n = 77), Eu-
ope (n = 58), Aus alasia (n = 45) and he Paci ic (n = 21). The mos widesp ead
in ade , occu ing in 55.7% o he sampled egions (n = 43), was Pon ede ia
c assipes ( his species was conside ed in asi e in 15 o he 43 egions), ollowed
by Pis ia s a io es and Elodea canadensis Michx., Azolla iliculoides, Elodea densa
(Planch.) Casp. and My iophyllum aqua icum (Vell.) Ve dc., all occu ing in a
leas 25% o he sampled egions (Table 1). In con as , 105 species (56.6% o
all sampled species) we e ound in only one egion.
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Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
Biogeog aphical pa e ns
The numbe s o na u alised aqua ic species (conside ing na u alised non-in asi e
and in asi e species oge he ) pe con inen and zonobiome we e no signi ican ly
di e en om he alues expec ed by chance. Howe e , a di e en pa e n was e i-
den o in asi e species. Aqua ic habi a s in T opical Asia ha bou ed mo e in asi e
species han expec ed; he opposi e was ound o he Medi e anean zonobiome,
whe e in asi e aqua ic species we e unde - ep esen ed (Table 2). Using he p o-
po ion o na u alised species ha ha e become in asi e as a measu e, T opical
Asia, A ica and T opical Asia anked he highes amongs con inen s, wi h 58.6%,
42.3% and 40.7%, espec i ely (Fig. 1). T opical zonobiomes, bo h Sa annah
(44.4%) and Equa o ial (43.5%) and he Sub opical A id zonobiome (38.7%)
had he g ea es p opo ions o na u alised species eco ded as in asi e (Fig. 1). No
species we e sampled in he Wa m Tempe a e zonobiome.
A con inen al-scale compa ison o na u alised species ichness in aqua ic e sus
e es ial habi a s, using a combined da ase o all na u alised species, demon-
s a ed ha in A ica and Tempe a e Asia, he obse ed p opo ion o aqua ic na -
u alised species was signi ican ly lowe han expec ed by chance (Table 3). Fo
zonobiomes, he aqua ic habi a s in he Medi e anean ha bou ed disp opo ion-
ally mo e and Wa m Tempe a e ewe (none) na u alised species han e es ial
habi a s in his egion (Table 3, Fig. 2).
Fo in asi e species, he numbe o aqua ic compa ed o e es ial was
highe han expec ed by chance in T opical Asia and No he n Ame ica and
lowe in Tempe a e Asia (Table 3). Rega ding zonobiomes, he ep esen a ion
Table 1. The op 21 aqua ic species eco ded in he la ges numbe o SynHab egions p esen ed by
in asion s a us. These ep esen 11.3% o species eco ded and 50.3% o all eco ds. Am-S = Sou h-
e n Ame ica, Am-N = No he n Ame ica, EU = Eu ope, AUS = Aus alasia, AS- emp = Tempe a e
Asia, AS- op = T opical Asia, AF = A ica; I = numbe o eco ds as in asi e, N = numbe o eco ds
as na u alised non-in asi e.
Species I N To al Con inen o o igin
Pon ede ia c assipes 15 28 43 Am-S
Pis ia s a io es 7 16 23 AF, Am-N, Am-S
Elodea canadensis 7 15 22 Am-N
Azolla iliculoides 7 14 21 Am-N, Am-S, An a c ica
Elodea densa 5 15 20 Am-S
My iophyllum aqua icum 6 14 20 Am-S
Nas u ium o icinale 2 17 19 AF, AS- emp, AS- op, EU
Sal inia moles a 10 7 17 Am-S
Hyd illa e icilla a 5 11 16 EU, AF, AS- emp, AS- op, AUS
Al e nan he a philoxe oides 6 7 13 Am-S
Po amoge on c ispus 1 9 10 EU, AF, AS- emp, AS- op, AUS
My iophyllum spica um 3 7 10 EU, AF, AS- emp, AS- op, Am-N
Pon ede ia aginalis 1 8 9 AS- emp, AS- op, AUS
Elodea nu allii 2 6 8 Am-N
Lemna minu a 2 7 9 Am-N, Am-S
Ve onica anagallis-aqua ica 8 8 AF, AS- emp, AS- op, EU
Nymphoides pel a a 1 6 7 AF, AS- emp, AS- op, EU
Aco us calamus 7 7 AS- emp, AS- op, Am-N
Typha angus i olia 1 5 6 EU, AF, AS- emp, AS- op, Am-N
Nelumbo nuci e a 6 6 EU, AS- emp, AS- op, AUS
Sagi a ia la i olia 6 6 Am-N, Am-S
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Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
Figu e 1. P opo ion o in asi e species amongs na u alised species in con inen s (a) and in zono-
biomes (b).
A ica
Tempe a e Asia
T opical Asia
Aus alasia
Eu ope
No he n Ame ica Sou he n Ame ica
0
1
2
3
4
0102030
Na u alised aqua ics (%)
a
Bo eal
Con inen al
Medi e anean
Mul iple
Nemo al
Sub opical A id
T opical Equa o ial
T opical Sa anna
Wa m Tempe a e
0
1
2
3
4
0102
03
0
A e age na u alised e es ials (%)
b
Figu e 2. Na u alisa ion in aqua ic habi a compa ed o e es ial habi a s (mean o all o he habi-
a s) by con inen (a) and zonobiome (b). The blue line shows he heo e ical linea inc ease in mean
na u alisa ion in bo h aqua ic habi a and e es ial habi a s, connec ing ze o and mean na u alisa-
ion in each habi a (ma ked by a blue poin ).
Table 2. Numbe o na u alised non-in asi e and in asi e aqua ic species in con inen s and zonobi-
omes. Ligh o ange cells indica e mo e and blue ewe alien species han expec ed by chance based on
he gene alised linea models. Signi icance indica ed by as e isks (* P < 0.05).
Con inen Na u alised non-in asi e In asi e
A ica 20 11
Tempe a e Asia 18 11
T opical Asia 21 17*
Aus alasia 39 6
Eu ope 67 14
No he n Ame ica 61 15
Sou he n Ame ica 45 10
Zonobiome
Bo eal 3 0
Con inen al 8 1
Medi e anean 69 10*
Mul iple 50 18
Nemo al 79 19
Sub opical A id 22 12
T opical Equa o ial 33 17
T opical Sa annah 27 16
Wa m Tempe a e 0 0
488
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Alessand a Ko z e al.: A global syn hesis o plan in asions in aqua ic habi a s
Pa el Pipek h ps://o cid.o g/0000-0003-1116-1013
Ka eřina Š aje o á h ps://o cid.o g/0000-0001-7824-1793
Paulina Anas asiu h ps://o cid.o g/0000-0001-6355-2126
Michael Ansong h ps://o cid.o g/0000-0003-3811-3230
Ma ga i a A ianou sou h ps://o cid.o g/0000-0002-6743-9240
Julie F. Ba celona h ps://o cid.o g/0000-0001-5087-8637
Sunee a Bha a h ps://o cid.o g/0000-0002-2993-7193
Fa zaneh Bo dba h ps://o cid.o g/0000-0002-3042-8329
Is ael Bo okini h ps://o cid.o g/0000-0002-1258-7932
Lau a Celes i-G apow h ps://o cid.o g/0000-0002-9533-6919
Edua do Chacón-Mad igal h ps://o cid.o g/0000-0002-8328-5456
Wayne Dawson h ps://o cid.o g/0000-0003-3402-0774
Do jee h ps://o cid.o g/0000-0001-6866-9278
F anz Essl h ps://o cid.o g/0000-0001-8253-2112
Lilian Fe u ino-Acos a h ps://o cid.o g/0000-0002-2065-9174
Es ela Figuei edo h ps://o cid.o g/0000-0002-8511-8213
Rodol o Flo es h ps://o cid.o g/0000-0002-7911-9228
Guillaume F ied h ps://o cid.o g/0000-0002-3653-195X
Nicol Fuen es h ps://o cid.o g/0000-0002-3773-9832
Pablo Galán h ps://o cid.o g/0000-0002-7037-7853
Ch is ian Gilli h ps://o cid.o g/0000-0002-6679-4654
Michael Glase h ps://o cid.o g/0000-0002-4695-6150
José Ramón G ande Allende h ps://o cid.o g/0000-0002-7066-0608
Zigman as Gudžinskas h ps://o cid.o g/0000-0001-6230-5924
Rachael Holmes h ps://o cid.o g/0000-0002-6045-8705
Philip E. Hulme h ps://o cid.o g/0000-0001-5712-0474
Inde ji h ps://o cid.o g/0000-0002-4142-1392
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Supplemen a y ma e ial 1
Comple e esul s o he indica o species analysis
Au ho s: Alessand a Ko z, Ma in Hejda, Jan Čuda, Za ah Pa ison, Jose B ůna, Ana No oa, Jan Pe gl,
Pa el Pipek, Ka eřina Š aje o á, Paulina Anas asiu, Michael Ansong, Ma ga i a A ianou sou, Julie
F. Ba celona, Sunee a Bha a, Fa zaneh Bo dba , Is ael Bo okini, Lau a Celes i-G apow, Edua do
Chacón-Mad igal, Wayne Dawson, Do jee, F anz Essl, Lilian Fe u ino-Acos a, Es ela Figuei edo,
Rodol o Flo es, Guillaume F ied, Nicol Fuen es, Pablo Galán, Ch is ian Gilli, Michael Glase , José
Ramón G ande Allende, Zigman as Gudžinskas, Rachael Holmes, Philip E. Hulme, Inde ji , Eun Su
Kang, Holge K e , Dan W. K ix, Ingol Kühn, Oma Lopez, AnaLu MacVean, T objon Makhkam-
o , Elizabe e Ma chan e, Hélia Ma chan e, Al ed Ma oyi, Rachid Meddou , Pie e Mee s, Sha i A.
Mukul, B ad R. Mu ay, Megan L. Mu ay, Daniel L. Nick en , P ince E. No man, Ali Ome , Anne e
Pa zel , Pie e B. Pelse , Joan Pino, Ma c Rie a, Dagobe o Rod íguez Delcid, Julissa Rojas-Sando al,
Rose Ro chés-Ribal a, José Yade Sage h Ruiz-C uz, S epan Sena o , Alexande N. Senniko , Bha a
Babu Sh es ha, Gideon F. Smi h, Ba ba a Toka ska-Guzik, Sima Soh abi, Ma k an Kleunen, Mon -
se a Vilà, Vik o ia Wagne , Pa ick Weigel , Ma en Win e , Ayşe Yazlık, Elena Zyko a, Pe Pyšek
Da a ype: docx
Explana ion no e: able S1. Lis o egions wi h da a ( egion, con inen , ype o da a ex ac ion = li -
e a u e/collabo a ion; da a sou ces; da a p o ide in case o collabo a ion). able S2. Indica o
species selec ed o a con inen o a g oup o con inen s. able S3. Indica o species selec ed o a
zonobiome o a g oup o zonobiomes.
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.102.151156.suppl1