93
Nu ien en ichmen and a i icial ligh a nigh syne gis ically con e
a compe i i e ad an age o alien aqua ic species o e na i es
Jin Zhang1, Yilin Nie1, Fukang Li1, Yidan Zhang1, Haihao Yu1, Chunhua Liu1
1 The Na ional Field S a ion o F eshwa e Ecosys em o Liangzi Lake, College o Li e Science, Wuhan Uni e si y, Wuhan 430072, China
Co esponding au ho s: Chunhua Liu ([email p o ec ed]); Haihao Yu ([email p o ec ed])
Copy igh : © Jin Zhang 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
Many eshwa e ecosys ems a e cu en ly acing he dual challenges o a i icial ligh a nigh (ALAN)
and nu ien en ichmen . How hese simul aneously ac ing en i onmen al ac o s a ec compe i i e
ou comes be ween alien and na i e aqua ic plan s has no been ho oughly in es iga ed. He e, we
conduc ed a g eenhouse expe imen o quan i y he compe i i e ou comes and hei de e minan s o
nine pai s o alien and na i e aqua ic plan species in China unde ully c ossed ac o ial combina ions
o ALAN (Ambien s. ALAN) and nu ien en ichmen (Low s. High). Nu ien en ichmen signi i-
can ly a ec ed compe i i e ou comes. Speci ically, na i e species ended o ha e mo e biomass unde
low-nu ien condi ions, while he e e se was ue unde high-nu ien , al hough he ends we e
no signi ican . This pa e n was e en mo e p onounced in he p esence o ALAN. Di e en ial e ec s
o nu ien en ichmen and ALAN on in e speci ic compe i ion o alien and na i e species may be
esponsible o a ia ion in compe i i e ou comes. We ound ha nu ien en ichmen ended o in-
c ease he in e speci ic compe i ion coe icien s o na i e species, whe eas an opposi e end o alien
species was obse ed, his e ec being mos p ominen in he p esence o ALAN. Ou indings sugges
ha nu ien en ichmen and ALAN could syne gis ically con e a compe i i e ad an age o alien
species o e na i e species. The e o e, in e en ion measu es a e necessa y o minimise nu ien en ich-
men o eshwa e habi a s, especially in habi a s exposed o ligh pollu ion esul ing om ALAN.
Key wo ds: A i icial ligh a nigh , compe i i e ou come, eshwa e ecosys em, in a- and in e -
speci ic compe i ion, in insic g ow h a es, nu ien a ailabili y, plan in asion
In oduc ion
The An h opocene is cha ac e ised by he widesp ead in asion o alien plan s, pos-
ing h ea s o local biodi e si y, dis up ing ecosys em se ices and causing sub-
s an ial economic losses (Vilà e al. 2011; Scha ne e al. 2020; Diagne e al.
2021; Zhang e al. 2023b). The success o hese in asions hinges on compe i i e
in e ac ions be ween alien and na i e species (Funk and Vi ousek 2007; Pea se and
Al e ma 2013; Wang e al. 2017). I is hypo hesised ha alien species a e gen-
e ally mo e compe i i e han na i e species, a no ion suppo ed by me a-analyses
p ima ily ocusing on in e speci ic compe i ion in ensi y (Kuebbing and Nuñez
2016; Goli e s and Wallin 2018). Howe e , in insic g ow h a es and in aspeci ic
compe i ion in ensi y also signi ican ly a ec he compe i i e ou comes (Chesson
2000; Ha e al. 2018). Thus, p edic ing co-exis ence dynamics be ween na i e
and alien species equi es conside a ion o all h ee de e minan s: in insic g ow h
Academic edi o : Ped o Anas ácio
Recei ed:
26 No embe 2024
Accep ed:
21 Ap il 2025
Published:
7 Oc obe 2025
Ci a ion: Zhang J, Nie Y, Li F, Zhang
Y, Yu H, Liu C (2025) Nu ien
en ichmen and a i icial ligh a nigh
syne gis ically con e a compe i i e
ad an age o alien aqua ic species
o e na i es. In: Anas ácio P, Ribei o
F, Chainho P (Eds) In asions in
Aqua ic Sys ems. NeoBio a 102:
93–108. h ps://doi.o g/10.3897/
neobio a.102.142791
NeoBio a 102: 93–108 (2025)
DOI: 10.3897/neobio a.102.142791
Ad ancing esea ch on alien species and biological in asions
A pee - e iewed open-access jou nal
NeoBio a
94
NeoBio a 102: 93–108 (2025), DOI: 10.3897/neobio a.102.142791
Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
a es, in a- and in e speci ic compe i ion. Ye , s udies in eg a ing hese ac o s
comp ehensi ely emain limi ed (bu see Godoy e al. (2014); Godoy and Le ine
(2014); Chu and Adle (2015); Zhang and an Kleunen (2019)).
In heo y, species wi h highe in insic g ow h a es and lowe in a- and in e speci -
ic compe i ion in ensi y a e expec ed o ou compe e o he s (Zhang and an Kleunen
2019). Howe e , in eali y, hese de e minan s may no ope a e independen ly. Alien
species o en exhibi highe in insic g ow h a es, bu his can coincide wi h s onge
in aspeci ic compe i ion (Dos ál e al. 2019). Fu he mo e, he e is a ade-o be-
ween in a- and in e speci ic compe i i e abili ies, obse ed in some alien plan spe-
cies (Lankau and S auss 2007; Lankau 2009). Fo ins ance, in en i onmen s whe e
an alien species aces in ense in aspeci ic compe i ion due o high abundance, indi-
iduals may p io i ise enhancing in aspeci ic compe i i e abili y, po en ially a he
expense o educed compe i i e abili y agains o he species (Lankau 2008). The e-
o e, accu a ely p edic ing species co-exis ence emains a signi ican challenge.
Ecosys ems wo ldwide a e cu en ly unde going unp eceden ed en i onmen al
changes d i en by ac o s such as nu ien en ichmen (Sinha e al. 2017). Nu ien
en ichmen can po en ially ampli y he compe i i e ad an age o alien plan species
o e na i e ones. Fo example, ga den expe imen s wi h indi idual plan s ha e demon-
s a ed ha alien species de i e g ea e bene i s om inc eased esou ce a ailabili y
compa ed o na i e species (Liu and an Kleunen 2017; Kawawa Abonyo and Oduo
2024; Shan e al. 2024). The e o e, nu ien en ichmen may al e compe i i e ou -
comes by impac ing he dispa i y in in insic g ow h a es be ween alien and na i e
species. S udies also indica e ha nu ien en ichmen educes he compe i i e e ec s
o na i e species on aliens (Ba ge e al. 2003; Elle and Oli ei a 2017; Sun e al. 2023),
al hough his o en e lec s changes in ela i e di e ences be ween in a- and in e -
speci ic compe i ion (bu see Sla e e al. (2022)). Addi ionally, nu ien en ichmen
signi ican ly educes in aspeci ic compe i ion in ensi y o alien species, while inc eas-
ing i o na i e species (Ba and Facelli 2022). I emains unclea whe he his pa e n
pe sis s in he p esence o in e speci ic compe i ion. In eg a ing in insic g ow h a es,
in a- and in e speci ic compe i ion p o ides deepe insigh s in o how nu ien en ich-
men in luences compe i i e ou comes be ween alien and na i e species. In addi ion,
he cu en unde s anding o hese e ec s o nu ien en ichmen is mainly om e -
es ial ecosys ems, while li le is known abou he si ua ion in aqua ic habi a s.
A i icial ligh a nigh (ALAN) imposes signi ican an h opogenic p essu e on
na u al biological sys ems (Falchi e al. 2016; Kyba e al. 2017; Sande s e al.
2021). ALAN a ec s no only animal beha iou , physiology and li e his o ies, bu
also al e s plan communi y composi ion and phenology (Bennie e al. 2016; Liu
e al. 2022). Recen ield su eys by Mu phy e al. (2021) ha e ound a posi i e
co ela ion be ween he p esence o he in asi e plan B omus ec o um L. and
s ee ligh s, sugges ing ha ALAN may di e en ly in luence alien and na i e spe-
cies, he eby a ec ing hei compe i i e ou comes. Howe e , Speiße e al. (2021)
ound no e idence ha ALAN enhances he dominance o alien species in com-
muni ies. Mo eo e , hey obse ed a end o dec eased i ness o alien species
unde ALAN compa ed o ambien ligh condi ions when g own indi idually.
Con e sely, ano he s udy sugges ed ha ALAN ended o p omo e he g ow h o
alien species unde monocul u es (Liu e al. 2022). In aspeci ic in e ac ions may
pa ially explain he disc epancies be ween hese s udies’ esul s. O e all, ou un-
de s anding o how ALAN impac s he pe o mance o in asi e and na i e species,
as well as hei in a- and in e speci ic in e ac ions, emains limi ed.
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NeoBio a 102: 93–108 (2025), DOI: 10.3897/neobio a.102.142791
Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
A la ge numbe o exo ic aqua ic plan s ha e been consciously in oduced o
ecological es o a ion and landscape c ea ion and many aqua ic habi a s ha e su -
e ed se e e plan in asions as a esul (Hussne e al. 2021; Wang e al. 2024).
In addi ion, eshwa e ecosys ems a e pa icula ly suscep ible o ALAN because
ligh ing oad ne wo ks, u ban de elopmen and indus ial in as uc u e a e o en
loca ed along i e s, lakes and coas s (Reid e al. 2019; Seg es in e al. 2021), while
indus ial was ewa e and s o m-wa e uno om u ban a eas simul aneously
con ibu e o nu ien en ichmen (Maha jan e al. 2022; Kawawa Abonyo and
Oduo 2024). Thus, aqua ic habi a s p o ide an ideal si e o us o es he in e ac-
i e e ec s o ALAN and nu ien en ichmen .
He e, we conduc ed expe imen s using a eshwa e model sys em wi h se en
alien and six na i e species. Employing a esponse su ace design (Inouye 2001) in
which we a ied he densi y o bo h species o a pai independen ly, we plan ed 4320
indi iduals, desc ibing compe i i e ela ionships be ween nine pai s o alien and
na i e aqua ic species unde ully c ossed ac o ial combina ions o ALAN and nu-
ien en ichmen ea men s, each a wo le els. We es ed: (1) whe he alien aqua ic
species a e mo e compe i i e han na i e species and (2) how nu ien en ichmen
and ALAN a ec compe i i e ou comes be ween alien and na i e aqua ic species.
Ma e ials and me hods
S udy species
To examine he e ec s o ALAN and nu ien en ichmen on compe i i e ou -
comes be ween alien and na i e aqua ic plan s, we selec ed se en alien and six
na i e species o c ea e nine species pai s (Suppl. ma e ial 1: able S1). The alien
species we used in he expe imen included all o China’s in asi e subme ged and
loa ing plan s, excep o Azolla iliculoides Lam. (Wang e al. 2019; Wu and Ding
2019), which inc eased he gene alisabili y o he esul s ( an Kleunen e al. 2014).
Species pai s we e chosen, based on hei co-occu ence in habi a s and di e ences
in g ow h o ms. Hyd ocha is dubia (Blume) Backe , Pis ia s a io es L., Cabomba
ca oliniana A.G ay, Ege ia densa (Planch.) Casp., Elodea nu allii (Planch.) H.S .
John and Pon ede ia c assipes Ma . we e sou ced om comme cial plan ade s,
while he emaining species we e collec ed om pools o wa e om cemen anks
a he Na ional Field S a ion o F eshwa e Ecosys ems o Liangzi Lake, Hubei
P o ince, China (30°15'N, 114°33'E)(he ea e e e ed o as he “Liangzi Lake
S a ion”). Be o e s a ing he expe imen , we selec ed 240 indi iduals o simila
size o each species. As h ee species we e used in mul iple pai s, he ac ual num-
be o indi iduals was 240*n, whe e n ep esen s he numbe o epe i ions.
Expe imen al se -up
The expe imen was conduc ed in he g eenhouse a he Liangzi Lake S a ion, China.
Fo each alien-na i e species pai , we ansplan ed indi idual plan s acco ding o a e-
sponse-su ace expe imen al design (Inouye 2001). This design included 15 di e en
densi y combina ions, wi h o al densi ies o 4, 8 o 12 plan s pe po and he equency
o each species om 0 ia 0.25, 0.5 and 0.75 o 1. Each densi y combina ion o each
species pai was eplica ed 4 imes and hen andomly assigned o a ully c ossed ac o ial
combina ion o ALAN and nu ien en ichmen ea men s a wo le els each (Fig. 1).
96
NeoBio a 102: 93–108 (2025), DOI: 10.3897/neobio a.102.142791
Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
Due o di e ences in g ow h pe iods and g ow h o ms amongs di e en spe-
cies, he expe imen al du a ion and he ype o con aine s we e a ied. On 12, 13
Ma ch 2024, we ansplan ed he indi iduals o he i s six species pai s in o soil-
illed plas ic po s ( op diame e : 18 cm, bo om diame e : 13 cm, heigh : 12.5 cm)
acco ding o he expe imen al design. The soil (TN: 0.60 ± 0.005 g kg−1, TP: 0.58
± 0.03 g kg−1, mean ± s anda d e o (SE), n = 3) was ob ained om he Liangzi
Lake S a ion and me iculously sie ed o elimina e s ones, plan s and animal em-
nan s. We op ed o e es ial opsoil as opposed o lake-bo om sil , in o de o
mi iga e he po en ial o he eme gence o unwan ed mac ophy es s emming om
an aqua ic plan seed bank (Yan e al. 2024). We hen placed six po s o he same
densi y combina ion and ea men in a speci ic o de in a ec angula aqua ium
(leng h × wid h × heigh : 105 × 50 × 60 cm), a anged in h ee ows and wo
columns and illed he aqua ium wi h lake wa e o a dep h o 50 cm. On 4 Ap il
2024, we placed he indi iduals o he 7 h and 8 h species pai s in cylind ical s ain-
less-s eel anks (50 cm diame e , 60 cm heigh ) wi h lake wa e dep h o 50 cm
acco ding o he expe imen al design. On he same day, we placed he indi iduals
o he 9 h species pai in ci cula lowe po s (70 cm op diame e , 50 cm bo om
diame e , 40 cm heigh ) wi h a wa e dep h o 30 cm.
A e h ee days o ansplan ing he indi iduals, we s a ed di e en expe imen al
ea men s. We used LED s ips ixed abo e he h ee ypes o con aine s o simula e
ALAN. The LED s ips (6 W/m, IP 180 lm/m, wa m whi e 3300K-5000K; NVC,
Figu e 1. G aphical illus a ion o he esponse-su ace design used o each pai o alien and na i e aqua ic plan species. Each pai is
ea ed wi h di e en nu ien (N− s. N+) and ALAN (ALAN− s. ALAN+) condi ions.
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NeoBio a 102: 93–108 (2025), DOI: 10.3897/neobio a.102.142791
Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
China) we e posi ioned 10 cm abo e he wa e su ace o he ec angula aqua i-
um, 56 cm o he cylind ical s ainless-s eel anks and 40 cm o he ci cula lowe
po s. The con aine s we e a anged in ows, wi h he plas ic po s in he ec angula
aqua ium also a anged in ows. The e o e, he LED s ips ixed in he middle o
he wo ows o cylind ical s ainless-s eel anks, he wo ows o ci cula lowe po s
and he single ow o ec angula aqua iums p o ided uni o m illumina ion o each
con aine . A nigh , he con ol g oup emained in da kness, while he ALAN g oup
was con inuously exposed o ligh a an in ensi y o 16 ± 3.5 lux. Du ing he expe -
imen , he LED ligh s we e swi ched on a 6:30 pm and o a 6:30 am he nex day.
We con olled he nu ien le els by a ying he amoun o slow- elease e ilise
(Osmoco e Plus 15N-9P-12K+2MgO+TE, E e is). Speci ically, we added 2.3 g
o e ilise o hal o he ec angula aqua iums o high nu ien ea men , lea -
ing he o he hal wi hou e ilise o low nu ien ea men . Fo he cylind ical
s ainless-s eel anks and ci cula lowe po s, he high nu ien ea men in ol ed
adding 7 g and 1.6 g o e ilise , espec i ely, while he low nu ien ea men
in ol ed adding 2.5 g and 0.7 g, espec i ely. The con olled elease e ilise was
placed in a mesh bag a he bo om o he con aine . Fo he ec angula aqua i-
ums, he nu ien concen a ions in he wa e column unde low and high nu ien
ea men s we e 0.758 ± 0.025 mg l−1 and 0.967 ± 0.022 mg l−1 o TN, 0.014
± 0.001 mg l−1 and 0.030 ± 0.001 mg l−1 o TP, espec i ely. Fo he cylind ical
s ainless-s eel anks and ci cula lowe po s, he nu ien concen a ions in he
wa e column unde low and high nu ien ea men s we e 1.236 ± 0.077 mg l−1
and 1.476 ± 0.080 mg l−1 o TN, 0.141 ± 0.016 mg l−1 and 0.222 ± 0.030 mg
l−1 o TP, espec i ely. Da a ep esen mean alues ± s anda d e o , n = 18. These
wo wa e quali y indices we e signi ican ly di e en be ween he wo nu ien le -
els. The concen a ion o nu ien s in he high nu ien ea men was wi hin he
ange o nu ien en ichmen in eshwa e lakes ac oss China (Huo e al. 2013).
Measu emen s
On 5 May 2024, we ha es ed all plan s om he ec angula aqua ium, d ied
each indi idual (including oo s) o a cons an weigh a 70 °C and weighed hem
o he nea es millig am. On 20 May 2024, we ha es ed, d ied and weighed he
emaining plan s in he same manne .
S a is ical analyses
In insic g ow h a es and compe i ion coe icien s
To desc ibe he compe i ion be ween alien and na i e species pai s, we i ed ou da a
on he biomass o indi idual plan s o a linea ised e sion o a modi ied Ricke com-
pe i ion model (Ricke 1954) by aking he na u al log o bo h sides o he equa ion:
ln(Bi, ) = ln(λi) – αii Ni, – αij Nj,
In he equa ion, Bi, is he indi idual biomass o species i in yea . Ni, and Nj,
a e he numbe o indi iduals o species i and j in yea , espec i ely. λi is he
in insic g ow h a e o species i, which ep esen s he expec ed biomass o an in-
di idual in he absence o conspeci ic and he e ospeci ic neighbou s. αii ep esen s
98
NeoBio a 102: 93–108 (2025), DOI: 10.3897/neobio a.102.142791
Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
he pe capi a e ec o species i on i sel ( he in aspeci ic compe i ion coe icien )
and αij ep esen s he pe capi a e ec o species j on species i ( he in e speci ic
compe i ion coe icien ). Posi i e alues o compe i ion coe icien s indica e com-
pe i ion amongs indi iduals and nega i e alues indica e acili a ion. Bi, ,Ni, and
Nj, we e measu ed di ec ly du ing he expe imen , while λi, αii and αij we e es i-
ma ed by i ing he linea ised model. Al hough he subsc ip ypically deno es a
ime s ep (e.g. in yea s), he e i e e s only o he single ime poin o ha es a he
end o he expe imen and does no ep esen in e -annual dynamics.
Fo each species pai unde he ou di e en expe imen al condi ions, we i ed
a linea mixed-e ec s model. We used o al biomass o each indi idual a he end
o he expe imen . We included he numbe o conspeci ic and he e ospeci ic com-
pe i o s as ixed e ec s, wi h con aine as a andom e ec o accoun o non-in-
dependence o indi iduals wi hin he same con aine . We used he numbe o in-
di iduals a he end o expe imen a he han a he beginning o he expe imen .
Nex , we used linea mixed-e ec s models o es he e ec s o species s a us, nu-
ien en ichmen and ALAN on in insic g ow h a es and in aspeci ic compe i-
ion coe icien s. We included species s a us, nu ien en ichmen , ALAN and hei
in e ac ions as ixed ac o s, wi h iden i y (nes ed wi hin amily) o a ge species as
andom e ec s. We also es ed he e ec s o species s a us, nu ien en ichmen and
ALAN on in e speci ic compe i ion coe icien s. In his mixed-e ec s linea model,
we included species s a us, nu ien en ichmen , ALAN and hei in e ac ions as ixed
ac o s, wi h a ge species and compe i o species (nes ed wi hin hei espec i e am-
ilies) as andom e ec s. In all models, we included he in e se o a iance as weigh s.
Compe i i e ou comes
Based on popula ion-le el biomass, we calcula ed lnRRk, he compe i i e ou come
be ween alien and na i e plan s in each o he bicul u e con aine s (k) as:
ln RRk = ln(TBk_a) – ln (TBk_n)
whe e TBk_a and TBk_n a e he o al biomass o alien and na i e species, espec i ely, in
he bicul u e con aine k. Subsequen ly, o each pai o species in di e en condi ions,
we used a linea model wi h lnRRk as he esponse a iable and an in e cep as he only
explana o y a iable. The e o e, a posi i e in e cep alue i ed o a pai signi ies ha ,
on a e age, he alien species in ha pai exhibi ed highe biomass han he na i e spe-
cies ac oss a b oad ange o densi y combina ions (indica ing g ea e compe i i eness
o he alien species), whe eas a nega i e in e cep alue indica es he opposi e.
To es how nu ien en ichmen and ALAN a ec compe i i e ou comes be-
ween alien and na i e species, we i ed a linea mixed-e ec s model. The model
included nu ien en ichmen , ALAN and hei in e ac ions as ixed ac o s, wi h
iden i y (nes ed wi hin amily) o na i e species as andom e ec s. We also includ-
ed iden i y o alien species as andom e ec s and we allowed each alien species o
espond di e en ly o he nu ien ea men s (i.e. we included andom slopes).
We included he in e se o a iance o compe i ion ou comes as weigh s.
Conside ing po en ial di e ences be ween species combina ions, we also i ed
a linea model using he gls unc ion. This model included nu ien en ichmen ,
ALAN, iden i y o species pai and hei in e ac ions as ixed ac o s. Addi ionally,
i allowed o di e en a iances o each species pai .
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NeoBio a 102: 93–108 (2025), DOI: 10.3897/neobio a.102.142791
Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
All analyses we e conduc ed using he lme4 package (Ba es e al. 2015) and nlme
package (Pinhei o e al. 2023) in R 4.2.0 (Team 2022). Resul s we e isualised using
he plo unc ion o he ggplo unc ion om he ggplo 2 package (Wickham 2016).
Resul s
Compa ed wi h na i e aqua ic species, alien species ended o ha e highe in insic
g ow h a es, bu he di e ence was no s a is ically signi ican (Table 1, Fig. 2a).
The in insic g ow h a e was also no a ec ed by he in e ac ion be ween s a us and
nu ien en ichmen o he in e ac ion be ween s a us and ALAN (Table 1, Fig. 2a).
Alien species ended o ha e lowe in aspeci ic compe i ion coe icien s compa ed
wi h na i e aqua ic species, bu he di e ence was no s a is ically signi ican (Table
1, Fig. 2b). The in aspeci ic compe i ion coe icien s we e also no a ec ed by he
in e ac ion be ween s a us and nu ien en ichmen o he in e ac ion be ween s a-
us and ALAN (Table 1, Fig. 2b). O e all, he e was no di e ence in in e speci ic
compe i ion coe icien s be ween alien and na i e species (Table 1, Fig. 2c). How-
e e , nu ien en ichmen a ec ed he in e speci ic compe i ion coe icien s o alien
and na i e species signi ican ly di e en ly (Table 1). Speci ically, nu ien en ich-
men ended o inc ease he in e speci ic compe i ion coe icien s o na i e species,
whe eas an opposi e end was obse ed o alien species (Suppl. ma e ial 1: ig. S1),
his e ec being mos p onounced in he p esence o ALAN (a ma ginally signi i-
can h ee-way in e ac ion be ween s a us, nu ien and ALAN in Table 1, Fig. 2c).
O e all, when g own oge he , alien species and na i e species exhibi ed simila
biomass (Fig. 3). Compe i i e ou comes be ween alien and na i e species we e sig-
ni ican ly a ec ed by nu ien en ichmen (Table 2). Speci ically, unde low-nu i-
en condi ions, na i e species ended o ha e highe biomass, while unde high-nu-
ien condi ions, alien species ended o ha e highe biomass (Fig. 3). Howe e ,
hese ends we e no s a is ically signi ican . This pa e n was mo e p onounced
Table 1. E ec s o s a us o a ge species (na i e s. alien), nu ien en ichmen (low-nu ien s.
high-nu ien le els), ALAN ea men (ambien s. ALAN) and hei in e ac ion on in insic g ow h
a es (λi) and in a- and in e speci ic compe i ion coe icien s (αii and αij). Signi ican (P < 0.05) e ec s a e
in bold. Ma ginally signi ican (0.05 ≤ P < 0.10) e ec s a e shown in i alics. The signi icance o ixed e -
ec s was assessed wi h likelihood- a io es s by compa ing models wi h and wi hou he e ec o in e es .
λi; χ2Pαii; χ2Pαij; χ2P
S a us (S) 1.521 0.217 1.902 0.168 0.613 0.433
Nu ien (N) 0.542 0.462 1.137 0.286 1.688 0.193
ALAN (L) 1.305 0.253 0.014 0.906 0.765 0.381
S × N 1.439 0.23 1.479 0.224 5.503 0.018
S × L 0.821 0.364 0.063 0.802 0.578 0.446
N × L 4.653 0.031 2.831 0.093 0.389 0.532
S × N × L 1.031 0.309 1.399 0.237 3.385 0.065
Random e ec s SD SD SD
Species o a ge 0.906 0.000 0.009
Family o a ge 0.503 0.019 0.013
Species o compe i o - - 0.019
Family o compe i o - - 0.002
Residual 0.407 0.450 0.280
R2
mR2
cR2
mR2
cR2
mR2
c
0.080 0.877 0.001 0.003 0.003 0.011
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Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
Figu e 2. Mean in insic g ow h a es (a λi) and in a- and in e speci ic compe i ion coe icien s (b αii; c αij) o alien and na i e aqua ic
plan species unde di e en expe imen al ea men s. Posi i e alues o αii and αij indica e compe i i e in e ac ions. E o ba s ep esen
95% con idence in e als. In insic g ow h a es (λi), whose uni is millig am, was na u al log- ans o med.
Figu e 3. Mean alues o compe i i e ou comes be ween alien and na i e plan species (i.e. mean alues o lnRRk) unde di e en expe -
imen al ea men s. E o ba s ep esen 95% con idence in e als.
Table 2. E ec s o nu ien en ichmen (low-nu ien s. high-nu ien le els), ALAN ea men
(ambien s. ALAN) and hei in e ac ion on compe i i e ou comes (i.e. which species ha e highe
biomass in bicul u es ac oss di e en densi y combina ions). Signi ican (P < 0.05) e ec s a e in bold.
Ma ginally signi ican (0.05 ≤ P < 0.10) e ec s a e shown in i alics. The signi icance o ixed e ec s
was assessed wi h likelihood- a io es s by compa ing models wi h and wi hou he e ec o in e es .
χ2P
Nu ien (N) 4.524 0.033
ALAN (L) 0.255 0.614
N × L 4.008 0.045
Random e ec s SD
Alien species 1.050 (0.260)a
Na i e species 0.457
Family o na i e 1.231
Residual 0.211
R2
mR2
c
0.022 0.984
aHe e, he s anda d de ia ions (SD) a e gi en o he andom in e cep s (co esponding o he alues in he high
nu ien ea men ) and andom slopes (co esponding o he e ec s o he low nu ien ea men ) o species o
alien. The alues in b acke s ep esen he SDs o he andom slopes.
101
NeoBio a 102: 93–108 (2025), DOI: 10.3897/neobio a.102.142791
Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
in he p esence o ALAN, demons a ed by a signi ican ALAN × nu ien en ich-
men in e ac ion (Table 2, Fig. 3). We also ound ha compe i i e ou comes we e
signi ican ly a ec ed by species combina ion (Suppl. ma e ial 1: able S2, ig. S2).
Mo eo e , he e ec (Suppl. ma e ial 1: able S2, ig. S2) and de e minan s (Suppl.
ma e ial 1: igs S3–S5) o nu ien en ichmen on compe i i e ou comes a ied
amongs di e en species combina ions.
Discussion
While alien species a e o en pe cei ed as mo e compe i i e han na i e coun e -
pa s (Vilà and Weine 2004; Pea se e al. 2019), ou s udy did no ind signi -
ican di e ences in compe i i e abili y be ween hem (Fig. 3). Simila ly, Zhang
and an Kleunen (2019) ound in hei s udy quan i ying compe i ion ou comes
amongs 48 pai s o na i e and na u alised alien plan s in Ge many ha alien
and na i e species gene ally exhibi ed compa able compe i i e abili ies. Howe e ,
hei esea ch indica ed ha common alien plan s we e mo e compe i i e han a e
na i es. P e ious s udies in o alien-na i e compe i ion may ha e been skewed o-
wa ds using a e na i e species (Vilà and Weine 2004), po en ially con ibu ing o
a iable ou comes. The na i e aqua ic species in ou s udy a e widely dis ibu ed in
China and some a e ecognised as in asi e in o he egions wo ldwide (e.g. My io-
phyllum spica um L. (Ma ko e al. 2008; Ma in and Coe zee 2014), Po amoge on
c ispus L. (Fleming e al. 2015), Hyd illa e icilla a (L. .) Royle (Louback-F anco
e al. 2020; Mo mul e al. 2020). A me a-analysis p e iously indica ed no signi i-
can ai dispa i ies be ween in asi e alien and na i e species known o be in asi e
elsewhe e (Van Kleunen e al. 2010). Thus, ou indings, alongside exis ing e-
sea ch, sugges ha alien species do no inhe en ly possess a compe i i e ad an age
o e commonly encoun e ed na i es.
G ow h and in aspeci ic compe i ion may exhibi a ade-o ela ionship, whe e
la ge and as e -g owing species ypically ace s onge esou ce compe i ion and
g ea e nega i e densi y dependence (Dos ál e al. 2019). Howe e , ou s udy did
no obse e his ade-o pa e n. Ins ead, we ound ha alien species gene ally
displayed highe in insic g ow h a es coupled wi h lowe , a he han highe , le -
els o in aspeci ic compe i ion (Fig. 2a, b). Fu u e esea ch should in es iga e he
p e alence and mechanisms unde lying his ela ionship be ween g ow h and in a-
speci ic compe i ion. In con as o indings om se e al me a-analyses (Kuebbing
and Nuñez 2016; Goli e s and Wallin 2018), we did no ind e idence ha alien
species consis en ly exhibi lowe in e speci ic compe i ion coe icien s compa ed
o na i es (Fig. 2c). Po en ial explana ions include he smalle sample size o species
pai s in ou s udy compa ed o me a-analyses, esul ing in educed s a is ical powe .
Mo eo e , he in ensi y o in e speci ic compe i ion be ween alien and na i e spe-
cies may a y depending on en i onmen al condi ions (e.g. nu ien a ailabili y), a
ac o ha ou s udy’s limi ed en i onmen al scope may no ully cap u e. Las , da a
om se e al me a-analyses may ha e been de i ed p ima ily om e es ial ecosys-
ems, whe eas he subjec s o ou s udy we e subme ged and loa ing aqua ic plan s.
P e ious empi ical s udies examining he e ec s o nu ien en ichmen on com-
pe i i e ou comes ha e p oduced mixed esul s. While some s udies indica e posi i e
e ec s (Zhang e al. 2017; Uddin and Robinson 2018; Ren e al. 2019), o he s sug-
ges ha in he p esence o common na i es, nu ien en ichmen may no a ec o
could e en diminish he compe i i e ad an age o alien species (Going e al. 2009;
108
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Jin Zhang e al.: Nu ien -ALAN syne gy a o s alien species
Zhang Z, an Kleunen M (2019) Common alien plan s a e mo e compe i i e han a e na i es bu
no han common na i es. Ecology Le e s 22(9): 1378–1386. h ps://doi.o g/10.1111/ele.13320
Zhang H, Chang R, Guo X, Liang X, Wang R, Liu J (2017) Shi s in g ow h and compe i i e domi-
nance o he in asi e plan Al e nan he a philoxe oides unde di e en ni ogen and phospho us
supply. En i onmen al and Expe imen al Bo any 135: 118–125. h ps://doi.o g/10.1016/j.en -
expbo .2016.12.014
Zhang X, an Kleunen M, Chang C, Liu Y (2023a) Soil mic obes media e he e ec s o esou ce
a iabili y on plan in asion. Ecology 104(10): e4154. h ps://doi.o g/10.1002/ecy.4154
Zhang X, Zhang T, Liu Y (2023b) E ec s o a buscula myco hizal ungi on plan in asion success
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Supplemen a y ma e ial 1
Supplemen a y in o ma ion
Au ho s: Jin Zhang, Yilin Nie, Fukang Li, Yidan Zhang, Haihao Yu, Chunhua Liu
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