191
Epiphy ic algae mi iga e he inhibi o y e ec s o wo aqua ic in asi e
plan s, Pon ede ia c assipes and Pis ia s a io es, on a subme ged
plan communi y
Yi-Luan Shen1,2, Jing-Fang Cai1,2, Jing-Jing Xue1,2, Xiao-Li Yang1,2, Da-Zhi Wang1,2, Hong-Li Li1,2
1 School o Ecology and Na u e Conse a ion, Beijing Fo es y Uni e si y, Beijing 100083, China
2 Key Labo a o y o S a e Fo es y and G assland Adminis a ion on Ecological P o ec ion in he Yellow Ri e Basin, Beijing Fo es y Uni e si y, Beijing 100083, China
Co esponding au ho : Hong-Li Li ([email p o ec ed])
Copy igh : © Yi-Luan Shen 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
Exo ic plan in asions and epiphy ic algae po en ially in luence he g ow h o subme ged plan s. How-
e e , he e ec o epiphy ic algae on he esis ance o subme ged na i e plan s o plan in asion is elu-
si e. The e o e, we conduc ed his s udy o explo e he e ec s o in asion scena ios and epiphy ic algae
on a subme ged plan communi y (Hyd illa e icilla a, Po amoge on w igh ii, and Vallisne ia na ans)
and wa e quali y. We se up a con ol expe imen wi h ou in asion scena ios (no in asion, Pon -
ede ia c assipes in asion, Pis ia s a io es in asion, and co-in asion o P. c assipes and P. s a io es) and
wo epiphy ic algal s a es (wi h and wi hou epiphy ic algae). The p esence o in asi e plan s esul ed
in signi ican ly lowe biomass (P < 0.01) o he subme ged plan communi y wi hou epiphy ic algae
han ha wi h epiphy ic algae, and he communi y biomass was lowes in he case o co-in asion. The
p esence o epiphy ic algae esul ed in na i e plan s exhibi ing compe i i e in e ac ions only du ing
co-in asion, whe eas na i e plan s exhibi ed compe i i e in e ac ions a all imes in he absence o ep-
iphy ic algae. To al ni ogen and phospho us con en s in wa e we e lowes in he absence o in asion.
Addi ionally, o al phospho us con en s was highes in he p esence o epiphy ic algae and co-in asion.
These esul s demons a e ha co-in asion s ongly inhibi s he g ow h o subme ged plan communi-
ies mo e han single in asions, and epiphy ic algae po en ially o se his supp ession. These indings
will p o ide a scien i ic basis o subme ged na i e plan es o a ion in we land ecosys em.
Key wo ds: Co-in asion, na i e plan s, plan in asion, plan s es o a ion, wa e quali y
In oduc ion
Subme ged plan s a e p ima y p oduce s, p o iding abundan ood and habi a o
aqua ic bio a and aiding in wa e pu i ica ion (Jeppesen e al. 1997; Li e al. 2023;
Liao e al. 2024). The e o e, subme ged plan s a e c i ical in main aining he biodi e -
si y and s abili y o shallow lake ecosys ems (Liu 2021). Howe e , in asions con ibu e
signi ican ly o he decline and ex inc ion o subme ged plan communi ies (Phillips
e al. 2016). In asi e plan s ep oduce apidly and occupy c ucial ecological niches,
loa ing in asi e plan s can o m a dense meadow on he wa e su ace, compe ing wi h
subme ged plan s o ligh , nu ien s, and o he esou ces (Fleming and Dibble 2014).
Plan in asions a e equen ly accompanied by changes in he aqua ic en i onmen , u -
he deg ading he subme ged plan communi y (Zhou e al. 2017; Yuan e al. 2023).
Academic edi o : Ped o Anas ácio
Recei ed:
10 Decembe 2024
Accep ed:
21 Ap il 2025
Published:
7 Oc obe 2025
Ci a ion: Shen Y-L, Cai J-F, Xue J-J,
Yang X-L, Wang D-Z, Li H-L (2025)
Epiphy ic algae mi iga e he inhibi o y
e ec s o wo aqua ic in asi e
plan s, Pon ede ia c assipes and
Pis ia s a io es, on a subme ged
plan communi y. In: Anas ácio P,
Ribei o F, Chainho P (Eds) In asions
in Aqua ic Sys ems. NeoBio a 102:
191–207. h ps://doi.o g/10.3897/
neobio a.102.144004
NeoBio a 102: 191–207 (2025)
DOI: 10.3897/neobio a.102.144004
Ad ancing esea ch on alien species and biological in asions
A pee - e iewed open-access jou nal
NeoBio a
192
NeoBio a 102: 191–207 (2025), DOI: 10.3897/neobio a.102.144004
Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
Plan in asions a e p e alen , and ecological niches o ce ain in asi e plan s
o e lap such ha ecosys ems a e co-in aded by wo o mo e plan s (Wang e
al. 2020; Wei e al. 2020). The equen in e ac ion be ween in asi e plan s
impac s na i e plan communi ies in hese co-in aded ecosys ems (Zhong e
al. 2022; Li e al. 2024). Nega i e in e ac ions be ween in ade s can mode a e
hei impac s on na i e plan communi ies (Wang 2016). On he o he hand,
in asi e species can mu ually con ibu e o hei es ablishmen , u he accele -
a ing he in asion p ocess and causing mo e se e e impac s on na i e plan s, a
phenomenon known as in asional mel down (Simbe lo and Von Holle 1999;
Wang 2016). A s udy showed ha he co-in asion o wo in asi e plan s E ig-
e on annuus (L.) Pe s. and Solidago canadensis L. can impose syne gis ic impac s
on soil bac e ial ichness, he eby acili a ing hei subsequen in asion (Wei
e al. 2020). No only will he co-in asion o plan s cause mo e se ious dam-
age, bu he co-in asion o plan s and he bi o es will also ha e simila e ec s;
he bi o y by he in asi e snail Pomacea canalicula a may enhance he in asi e-
ness o My iophyllum aqua icum (Vell.) Ve dc. (Yan e al. 2022). Compelling
e idence indica es ha single-species in asions a e de imen al o na i e plan
communi ies, and he in asional mel down hypo hesis con inues o be alida -
ed a he same ime (Kuehne e al. 2016; Yan e al. 2022). Howe e , he in e ac-
ions be ween co-in ading plan s in aqua ic ecosys ems and hei e ec s unde
ex eme in asion on na i e subme ged plan communi ies emain elusi e.
Nume ous biological and abio ic ac o s egula e he e ec s o plan in asion
on subme ged plan communi ies, including epiphy ic algae, wa e nu ien s,
empe a u e, and wa e dep h (Huang e al. 2023; Peng e al. 2023; Yuan e
al. 2023). Among hese egula o y ac o s, epiphy ic algae, as a bio ic ac o ,
also po en ially impac he in asion esis ance o subme ged plan communi ies
(Song e al. 2017; L e al. 2023). Epiphy ic algae o m a bio ilm on he su ace
o subme ged plan s ems and lea es, o ming complex in e ac ions, compe i-
i e o symbio ic, wi h subme ged plan s (Ca aneo e al. 1998; Zhang e al.
2023; Alena e al. 2025). On he one hand, excessi e epiphy ic algae inhibi he
g ow h o subme ged plan s by compe ing o ligh and nu ien s, ul ima ely
causing he collapse o subme ged plan s and u ning he wa e om clea o
u bid (Asaeda e al. 2004; Min e al. 2017). On he o he hand, epiphy ic al-
gae can p omo e plan g ow h by aiding in ni ogen ixa ion (Dai e al. 2023),
complica ing he e ec s o epiphy ic algae on subme ged plan s. Howe e , in
aqua ic ecosys ems whe e plan s co-exis , he e ec o epiphy ic algae on he
esis ance o communi ies o subme ged plan s o in asion emains unknown.
As common in asi e species in aqua ic ecosys ems, Pon ede ia c assipes and
Pis ia s a io es L. cause e iden damage and ecological pollu ion o subme ged
plan communi ies, and he e a e nume ous s udies on hei in asion mechanisms
(Zhou e al. 2017; Qian e al. 2022). Howe e , he e is ela i ely limi ed esea ch
on he e ec s o epiphy ic algae on he esis ance o subme ged plan communi ies
o he co-in asion o P. c assipes and P. s a io es. The e o e, in his expe imen , we
de eloped a subme ged plan communi y wi h ou in asion scena ios (no in a-
sion, P. c assipes in asion, P. s a io es in asion, and co-in asion o P. c assipes and
P. s a io es) and wo epiphy ic algae (wi h and wi hou epiphy ic algae) ea men s
o es ou p oposed hypo heses: (1) Co-in asion supp esses he subme ged plan
communi y mo e han single in asions; (2) Epiphy ic algae inc ease he esis ance
o subme ged plan communi y o co-in asion.
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NeoBio a 102: 191–207 (2025), DOI: 10.3897/neobio a.102.144004
Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
Ma e ials and me hods
Expe imen al design
The expe imen was conduc ed in he expe imen al base o Sanqing Pa k, Bei-
jing Fo es y Uni e si y, Haidian Dis ic , Beijing, China (40°0'28.17"N,
116°20'17.64"E), om June 10 o July 28, 2023. The subme ged and in asi e
plan s used in he expe imen we e collec ed om an aqua ic plan b eeding base
in Suqian Ci y, Jiangsu P o ince, and we e cul u ed in wa e o wo weeks be o e
he commencemen o he expe imen .
Acco ding o p e ious esea ch (Fle che e al. 2023; Ma e al. 2023), we se
he expe imen al pe iod as 48 days, and used 25 L plas ic bucke s (38 cm high,
32.5 cm diame e a he uppe mou h, 27.5 cm diame e a he lowe mou h) as he
expe imen al con aine , wi h he bo om illed wi h 15 cm o subs a e (sand, soil
a io 2: 3, soil was collec ed om he ields o Sanqing Pa k). Subme ged plan s wi h
a o able and compa able g ow h condi ions we e selec ed and plan ed uni o mly in
he expe imen al bucke s, each o which included ou indi iduals o each subme ged
plan species (Hyd illa e icilla a (L. .) Royle, Po amoge on w igh ii Mo ong, and
Vallisne ia na ans (Lou .) H. Ha a. Ini ial leng h o H. e icilla a and P. w igh ii was
oughly 15 cm, ini ial leng h o V. na ans was oughly 10 cm).
The expe imen al se up comp ised g oups o epiphy ic algae and non-epiphy ic al-
gae. Subme ged plan s wi h he epiphy ic algae g oup we e plan ed di ec ly in he expe -
imen al bucke s, while subme ged plan s wi hou epiphy ic algae g oup we e ca e ully
b ushed wi h a so -b is led b ush in clean wa e o emo e epiphy ic algae on he su -
ace o he lea es be o e plan ing in he expe imen al bucke s. In addi ion, ou di e en
in asion scena ios we e es ablished, namely, no in asion, P. c assipes in asion wi h ou
indi iduals, P. s a io es in asion wi h ou indi iduals, and co-in asion o P. c assipes
and P. s a io es, wi h wo indi iduals o each species (Fig. 1). In asi e plan s co e ed
abou 95% o each expe imen al bucke . The e we e 8 ea men s, each wi h 5 epli-
ca es, and plan s we e wa e ed a egula in e als o main ain he wa e dep h a 20 cm.
Sampling and pa ame e de ec ion
All subme ged and in asi e plan s we e ha es ed a he end o he expe imen , and
wa e samples we e collec ed om each expe imen al bucke . The biomass o sub-
me ged and in asi e plan s was measu ed by d ying hem in an o en a 70 °C o
Figu e 1. Expe imen al design.
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Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
48 hou s un il a cons an weigh was achie ed. Wa e quali y es s o collec ed wa-
e samples included o al ni ogen con en (TN), o al phospho us con en (TP),
anspa ency, pH, and dissol ed oxygen (DO). The wa e TN con en was e alua ed
using ul a iole spec opho ome y wi h alkaline po assium pe sul a e diges ion,
whe eas he wa e TP con en was assessed by ammonium molybda e spec opho-
ome y. T anspa ency was e alua ed using he Secchi disk me hod, and wa e pH
was de e mined using a wa e quali y de ec ion pen (TDS-PH-EC). The DO con-
en was measu ed using a po able dissol ed oxygen (JPB-607A) analyze . O hese,
TN and TP measu emen s we e done in he labo a o y, and DO, pH and ans-
pa ency we e measu ed a 10:00 a.m. on he day plan s we e ha es ed. The abo e
measu emen me hods all e e o ou p e ious esea ch me hods (Shen e al. 2023).
S a is ical analysis
The compe i ion e ec s (CE) o in asi e plan s on subme ged plan s we e de e -
mined om he biomass o each plan acqui ed by d ying and weighing using he
ollowing o mula (Wang e al. 2023):
CE = ln(R0/Rs) (1)
Whe e R0 deno es he o al biomass o he subme ged plan communi y in he
absence o in asi e plan s, and Rs deno es he o al biomass o he subme ged plan
communi y in he p esence o in asi e plan s. A posi i e alue o he calcula ed
compe i ion e ec alue implied ha he in asi e plan s inhibi ed he g ow h o
he subme ged plan communi y, and he la ge he alue, he s onge he inhi-
bi ion e ec . A nega i e alue o he compe i ion e ec implied ha he in asi e
plan s p omo ed he g ow h o he subme ged plan communi y.
The ela i e dominance index (RDI) o in asi e plan s in ea men s wi h he
p esence o in asi e plan s was calcula ed as ollows (Yuan e al. 2013):
RDI = A/(A+B) (2)
Whe e A deno es he biomass o in asi e plan s in he ea men in he p esence o
in asi e plan s, and B deno es he o al biomass o he subme ged plan commu-
ni y in he same ea men .
The di e si y index o he communi y (Shannon-Wiene di e si y index) was
calcula ed acco ding o he ollowing o mula:
H = −∑Pi ln (Pi) (i = 1, 2, 3 …, S) (3)
Whe e S deno es he numbe o species in he communi y, and Pi deno es he
biomass o species i di ided by he sum o he biomass o all communi y species.
The e enness index (Pielou e enness index) was calcula ed om he di e si y
index using he ollowing o mula:
E = H/ln (S) (4)
Whe e S deno es he numbe o species in he communi y, and H deno es he
Shannon-Wiene di e si y index.
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Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
Da a analysis
A wo-way ANOVA was applied o de e mine he e ec s o in asion scena ios,
epiphy ic algae, and he wo-way in e ac ion on communi y indexes (including
di e si y index, e enness index, o al communi y biomass, and compe i ion e ec ),
he biomass and ela i e dominance index o in asion plan s, as well as he TN, TP,
anspa ency, pH, and DO o he wa e . A one-way analysis o a iance (ANOVA)
was employed o e alua e signi ican di e ences be ween ea men s. Be o e ANO-
VA was pe o med, all da a we e es ed o no mali y and homogenei y o a iances.
Pea son co ela ion analysis was employed o e alua e associa ions be ween in asi e
plan s, physical and chemical p ope ies o wa e , and subme ged plan communi ies.
S uc u al equa ion modeling was used wi h he piecewiseSEM R package o assess he
e ec s o in asi e plan s’ and wa e ’s physical and chemical p ope ies on subme ged
plan communi ies. S uc u al equa ion modeling in ol es ea ing each in luencing
ac o as an independen obse ed a iable. In his iew, we i s cons uc ed a model
ha inco po a ed all possible pa hs and hen used eg essions o e ine he model, yield-
ing he inal s uc u al equa ion model. The s anda d eg ession coe icien , o pa h
coe icien be ween indica o s, was used o quan i y he e ec s o in ading plan s and
physical and chemical p ope ies o wa e on he subme ged plan communi y. Fu -
he mo e, he model i was e alua ed using AIC, P-Value, Fishe ’s C, and Chi-Squa e.
Subme ged plan communi y me ics we e calcula ed in Excel, while co ela ion
analyses o in asion scena ios, epiphy ic algae, and subme ged plan communi y
g ow h we e pe o med in R 4.3.3. All plo s we e gene a ed in SigmaPlo 14.0.
Resul s
Cha ac e is ics o he subme ged plan communi y
The in asion scena ios and epiphy ic algae had signi ican e ec s on he biomass
o H. e icilla a and P. w igh ii. The in e ac ion be ween he wo ac o s had a sig-
ni ican e ec on H. e icilla a, while he in asion scena ios, epiphy ic algae, and
hei in e ac ion had no signi ican e ec on he biomass o V. na ans (P < 0.05,
Table 1). The in asion scena ios had signi ican e ec s on all communi y indica-
o s, including o al biomass, Shannon-Wiene di e si y index, Pielou e enness
index, and compe i ion e ec (P < 0.05, Table 1). Bu epiphy ic algae only signi i-
can ly a ec ed o al biomass and compe i ion e ec . The in e ac ion be ween he
wo ac o s was only signi ican o o al biomass (P < 0.05, Table 1).
Table 1. ANOVA esul s on he e ec s o in asion scena ios (I) and epiphy ic algae (E) on he indi-
ca o s o subme ged plan communi ies.
Indica o s In asion scena ios (I) Epiphy ic algae (E) I × E
Biomass o Hyd illa e icilla a 11.135*** 12.377** 4.378*
Biomass o Po amoge on w igh ii 3.296* 16.839*** 2.066
Biomass o Vallisne ia na ans 0.747 2.667 2.025
To al biomass 4.479** 24.798*** 5.051**
Shannon-Wiene di e si y index 14.341*** 0.952 0.154
Pielou e enness index 16.058*** 0.087 1.172
Compe i ion e ec 3.492* 41.014*** 0.706
†: Signi ican alues o F a e shown in bold; *** deno es P < 0.001, and ** deno es P < 0.01.
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Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
The end o changes in he biomass o H. e icilla a was consis en wi h he
changes in he o al biomass o communi y. Only when he e was no in asion, he
biomass o H. e icilla a wi hou epiphy ic algae was highe han ha wi h epiphy ic
algae (Fig. 2A). Excep o co-in asion, he biomass o P. w igh ii wi h epiphy ic algae
unde o he condi ions was highe han ha wi hou epiphy ic algae (Fig. 2B). Fo
V. na ans, only when P. c assipes in aded alone, he biomass wi h epiphy ic algae was
highe han ha wi hou epiphy ic algae, while unde o he ea men condi ions,
Pon ede ia c assipes
Pis ia s a io es
Co-in asion
Con ol
Biomass o
Hyd illa e icilla a
(g)
0.0
0.4
0.8
1.2
1.6
Wi h epiphy ic algaeWi hou epiphy ic alga
e
E:**
I:***
E×I:*
Biomass o Po amoge on w igh ii (g)
0.0
0.1
0.2
0.3
0.4
0.5 E:***
I:*
E×I:ns
In asion scena ios
Biomass o
Vallisne ia na ans
(g)
0.00
0.05
0.10
0.15
0.20 E:ns
I:ns
E×I:ns
A
B
C
a
a
a
ab ab
bc
cc
bc
d
ab
cd
a
ccbc
ab
b
a
b
ab
ab
b
ab
Figu e 2. E ec s o in asion scena ios (I) and epiphy ic algae (E) on biomass o h ee subme ged
plan s (mean + SE, n = 5). Con ol deno es “no in asion”; Pon ede ia c assipes deno es “Pon ede ia
c assipes in asion”; Pis ia s a io es deno es “Pis ia s a io es in asion”; co-in asion deno e “co-in asion
o Pon ede ia c assipes and Pis ia s a io es”; * deno es “P < 0.05”; ** deno es “P < 0.01”; *** deno es
“P < 0.001.” Di e en le e s indica e signi ican di e ences among ea men s (P < 0.05).
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Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
he e was no signi ican di e ence (Fig. 2C). To al communi y biomass was highe
in he absence o in asion wi hou epiphy ic algae han wi h epiphy ic algae, bu he
opposi e e ec was obse ed in he p esence o in asi e plan s, wi h o al communi y
biomass being lowes in he co-in asion scena io (Fig. 3A). The di e si y and e en-
ness indices o plan communi ies we e lowes in he absence o in asion (Fig. 3B, C).
Pon ede ia c assipes
Pis ia s a io es
Co-in asion
Con ol
To al biomass (g)
0
1
2
3
4
5
6
Wi h epiphy ic algae Wi hou epiphy ic algae
E:***
I:**
E×I:**
Shannon–Wiene di e si y index
0.0
0.2
0.4
0.6
0.8
1.0
1.2 E:ns
I:***
E×I:ns
In asion scena ios
Pielou e enness index
0.0
0.2
0.4
0.6
0.8
1.0
1.2 E:ns
I:***
E×I:ns
A
B
C
a
a
a
a
a
bb
b
b
b
aa
aa
aa
b
b
aa
aa
a
a
Figu e 3. E ec s o in asion scena ios (I) and epiphy ic algae (E) on indica o s o subme ged
plan communi ies (mean + SE, n = 5). Con ol deno es “no in asion”; Pon ede ia c assipes deno es
“Pon ede ia c assipes in asion”; Pis ia s a io es deno es “Pis ia s a io es in asion”; co-in asion deno e
“co-in asion o Pon ede ia c assipes and Pis ia s a io es”; ** deno es “P < 0.01”; *** deno es “P <
0.001.” Di e en le e s indica e signi ican di e ences among ea men s (P < 0.05).
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Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
The p esence o epiphy ic algae caused a nega i e compe i i e e ec be ween he sub-
me ged plan communi y and in asion plan s unde he in asion o P. c assipes alone
and P. s a io es alone, bu posi i e unde he co-in asion (Fig. 4A). On he o he
hand, he absence o epiphy ic algae esul ed in a posi i e compe i ion e ec in all
in asion scena ios, wi h co-in asion exe ing he highes e ec (Fig. 4B).
Cha ac e is ics o in asi e plan s
The indings e ealed ha epiphy ic algae exe ed a signi ican e ec on he ela i e
dominance index o in asi e plan s, bu nei he in asion scena ios no epiphy ic
algae had a signi ican e ec on in asi e plan biomass (Table 2). In asi e plan s
had a highe biomass in he absence o epiphy ic algae han in he p esence o
epiphy ic algae (Fig. 5A). The ela i e dominance index o in asi e plan s was sig-
ni ican ly highe in he absence o epiphy ic algae han in he p esence o epiphy ic
algae, wi h he highes alue o co-in asions (Fig. 5B).
Table 2. ANOVA esul s on he e ec s o in asion scena ios (I) and epiphy ic algae (E) on he indi-
ca o s o he in asion plan s.
Indica o s In asion scena ios (I) Epiphy ic algae (E) I × E
In asion plan s biomass 0.623 2.030 0.706
Rela i e dominance index 2.342 23.601*** 0.442
†: Signi ican alues o F a e shown in bold; *** deno es P < 0.001.
In asion scena ios
Compe i ion e ec
-2
-1
0
1
2
3
P. c assipesP. s a io es
Co-in asion
Pon ede ia c assipes
Pis ia s a io es
Co-in asion
Pon ede ia c assipes
Pis ia s a io es
Co-in asi
on
E:***
I:*
E×I:ns
cc
c
bb
a
Wi h epiphy ic algaeWi hou epiphy ic algae
Figu e 4. E ec s o in asion scena ios (I) and epiphy ic algae (E) on compe i ion e ec o sub-
me ged plan communi ies (mean + SE, n = 5). Pon ede ia c assipes deno es “Pon ede ia c assipes
in asion”; Pis ia s a io es deno es “Pis ia s a io es in asion”; co-in asion deno e “co-in asion o Pon -
ede ia c assipes and Pis ia s a io es”; * deno es “P < 0.05”; *** deno es “P < 0.001.” Di e en le e s
indica e signi ican di e ences among ea men s (P < 0.05).
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Yi-Luan Shen e al.: Epiphy ic algae mi iga e he inhibi o y e ec s
Physical and chemical p ope ies o wa e
In asi e scena ios had signi ican e ec s on TN, DO, pH, and anspa ency, and
epiphy ic algae exe ed signi ican e ec s on TP and DO. Howe e , he in e ac ion
be ween he wo ac o s had no signi ican e ec on any o he wa e quali y indica-
o s (Table 3). TN and TP we e lowes in he absence o in asion. In he p esence o
epiphy ic algae, TN was highes unde he in asion o P. s a io es alone, while in he
absence o epiphy ic algae, TN was highes unde he in asion o P. c assipes alone
(Fig. 6A). In he p esence o epiphy ic algae, TP inc eased wi h he p esence o he
in asi e plan s and was he highes le el unde co-in asion. Mo eo e , TP was sig-
ni ican ly highe in he p esence o epiphy ic algae han in he absence o epiphy ic
algae (Fig. 6B). On he o he hand, pH and DO we e highes in he absence o in a-
si e plan s and signi ican ly dec eased in he p esence o in asi e plan s, and ans-
pa ency was lowes in he p esence o P. s a io es alone. The p esence o absence o
epiphy ic algae had no signi ican e ec on pH and anspa ency (Fig. 6C–E).
Rele ance o in asi e plan s and wa e quali y o subme ged plan
communi y
The Pea son co ela ion analysis e ealed ha o al communi y biomass was nega-
i ely co ela ed wi h in asi e plan biomass (IB), whe eas he communi y di e si-
y index (SI) and e enness index (PI) we e nega i ely co ela ed wi h DO and pH
Pon ede ia c assipes
Pis ia s a io es
Co-in asion
In asion scena ios
Pon ede ia c assipes
Pis ia s a io es
Co-in asion
Biomass o in asion plan s (g)
0
5
10
15
20
25
Wi h epiphy ic algaeWi hou epiphy ic algae
A
E:ns
I:ns
E×I:ns
Rela i e dominance index
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4 E:***
I:ns
E×I:ns
B
a
a
a
aaa
dcd bc
ab ab a
Figu e 5. E ec s o in asion scena ios (I) and epiphy ic algae (E) on biomass and ela i e dominance index o in asion plan s (mean + SE, n = 5).
Pon ede ia c assipes deno es “Pon ede ia c assipes in asion”; Pis ia s a io es deno es “Pis ia s a io es in asion”; co-in asion deno es “co-in asion o
Pon ede ia c assipes and Pis ia s a io es”; *** deno es “P < 0.001.” Di e en le e s indica e signi ican di e ences among ea men s (P < 0.05).
Table 3 ANOVA esul s on he e ec s o in asion scena ios (I) and epiphy ic algae (E) on he phys-
ical and chemical wa e p ope ies.
Indica o s In asion scena ios (I) Epiphy ic algae (E) I × E
To al ni ogen con en 8.529*** 0.751 1.152
To al phospho us con en 1.220 4.748* 0.990
Dissol ed oxygen 72.212*** 6.766* 2.340
pH 131.515*** 0.945 0.996
T anspa ency 19.438*** 0.271 1.754
†: Signi ican alues o F a e shown in bold; *** deno es P < 0.001, and * deno es P < 0.05.
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