81
In asi e po en ial o Phymac is papillosa: assessing
en i onmen al ole ance and ecological impac on he
Po uguese in e idal ecosys ems
Ana M. Pe ei a1,2 , Nú ia Teixei a2, Tomás To ão2, Jade Fe bus2, Ma ga ida Sil a2, Joana I. Robalo1,2
1 MARE-ISPA, Rua Ja dim do Tabaco, 34 1149-041 Lisboa, Po ugal
2 ISPA, Rua Ja dim do Tabaco, 34 1149-041 Lisboa, Po ugal
Co esponding au ho : Ana M. Pe ei a (ana_pe ei [email p o ec ed])
Copy igh : © Ana M. Pe ei a 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
Human ac i i ies and en i onmen al changes ha e signi ican ly inc eased he in oduc ion o non-na-
i e species (NIS) in o ma ine habi a s, posing a challenge o biodi e si y and ecosys em manage-
men . This s udy p esen s a comp ehensi e app oach o assess he in asi eness o Phymac is papillosa,
a ecen ly in oduced sea anemone on he Po uguese in e idal coas . An expe imen was conduc ed
o e alua e he salini y and empe a u e ole ances o his species and p edic i s po en ial dis ibu ion
beyond i s na i e ange. A o al o 270 indi iduals we e used in a wo- ac o expe imen al design wi h
g oups o six indi iduals a i e empe a u e ea men s (15, 19, 23, 27, 31 °C) and h ee salini y con-
di ions (32, 35, 38 psu) eplica ed h ee imes. Mo ali y le els we e measu ed a e wo weeks. Re-
sul s showed ha salini y had no impac on mo ali y, bu empe a u e signi ican ly a ec ed su i al.
A 31 °C, all indi iduals died, a 27 °C mo ali y le el was in e media e, and no mo ali y occu ed a
23 °C o lowe . These indings sugges ha P. papillosa could po en ially in ade he Po uguese coas ,
he no he n Ibe ian Peninsula, and he No h Sea; howe e he high empe a u es o he Medi e a-
nean may be a limi ing ac o . Being con ined o he ocky in e idal zone, P. papillosa could compe e
wi h na i e species, pa icula ly wi h hose om he same unc ional g oup. Agg essi e in e ac ions
wi h six na i e sea anemone species we e induced, wi h P. papillosa domina ing h ee species om
he genus Ac inia and showing equi alen beha iou o A. sulca a. This beha iou al dominance may
nega i ely impac he spa ial dis ibu ion o na i e Ac inia species. The in eg a ion o hese indings
sugges s ha P. papillosa should be classi ied as an in asi e species wi h he po en ial o signi ican ly
impac ben hic communi ies along he Po uguese and no he n A lan ic in e idal coas s.
Key wo ds: Beha iou , Cnida ia, compe i ion, in asi e species, sea anemone
In oduc ion
Human ac ions and en i onmen al changes in ma ine habi a s ha e signi ican ly
inc eased he numbe o non-na i e species (NIS), in oduced and success ully es-
ablished ou side hei na i e dis ibu ion a ea (Glon e al. 2020). The consequenc-
es o hese species, pa icula ly in ecological aspec s, hold signi ican ele ance o
legisla o s, manage s, policymake s, and conse a ionis s when p io i ising man-
agemen ac ions and alloca ing unds (Oja ee e al. 2015). One o he p ima y
an icipa ed impac s is on na u al ben hic communi ies, leading o a educ ion in
biodi e si y and in he g ow h o o he species, po en ially caused by compe i ion
Academic edi o : Paula Chainho
Recei ed:
27 Janua y 2025
Accep ed:
14 Augus 2025
Published:
7 Oc obe 2025
Ci a ion: Pe ei a AM, Teixei a N,
To ão T, Fe bus J, Sil a M, Robalo JI
(2025) In asi e po en ial o Phymac is
papillosa: assessing en i onmen al
ole ance and ecological impac on
he Po uguese in e idal ecosys ems.
In: Anas ácio P, Ribei o F, Chainho P
(Eds) In asions in Aqua ic Sys ems.
NeoBio a 102: 81–91. h ps://doi.
o g/10.3897/neobio a.102.148042
NeoBio a 102: 81–91 (2025)
DOI: 10.3897/neobio a.102.148042
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: 81–91 (2025), DOI: 10.3897/neobio a.102.148042
Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
(Blackbu n e al. 2014; Oja ee e al. 2015). When hese impac s a e measu able
in ecological and/o socio-economic e ms, and he e is a capabili y o sp ead, he
species is e med in asi e (Ca deccia e al. 2018).
Biological in asions ypically un old h ough ou undamen al s ages (Black-
bu n e al. 2011): anspo , in oduc ion, es ablishmen , and sp ead. The eco-
logical and biological ai s in luence he likelihood o success ul in oduc ion o
ce ain species wi hin speci ic axonomic g oups. Cnida ians ha e se e al cha -
ac e is ics ha a e pa icula ly sui able o a success ul in asion. These include a
di e se ange o li e his o y s a egies, sexual and asexual ep oduc ion, high dis-
pe sal, and ole ance o abio ic ac o s like empe a u e and salini y (Shick 1991).
Many sea anemone species display a gene al p edisposi ion owa ds ha diness and
po en ially apid p oli e a ion, making hem well-adap ed o in oduc ion (Glon
e al. 2020). Al hough asexual ep oduc ion appea s o be he mos impo an in
in asion success in sea anemones, he abili y o ep oduce h ough bo h asexual
and sexual modes in he non-na i e ange, along wi h he lack o senescence, con-
ibu es o a species’ abili y o be du able in ade s (Glon e al. 2020). Wo ldwide,
ele en species o sea anemones ha e al eady been desc ibed as in asi e, wi h mea-
su able en i onmen al impac s (Glon e al. 2020).
The i s in asion s age ( anspo ) is al eady accomplished when a NIS species is
iden i ied. A e his ini ial in oduc ion, ac o s con ibu ing o he success o he
es ablishmen and sp ead include he abili y o acclima e o local en i onmen al con-
di ions, high ep oduc i e a e, and he ou come o ecological in e ac ions wi h he
esiden communi y (Podbielski e al. 2016; Ca deccia e al. 2018; Glon e al. 2020).
In 2020, ou esea ch eam iden i ied he p esence o Phymac is papillosa (Les-
son, 1830) on he Po uguese ocky in e idal coas (Pe ei a e al. 2022), a Paci -
ic Sou h Ame ican in e idal sea anemone species wi h no o he known eco ds
ou side i s na i e ange (Fig. 1). Th ee o he ou desc ibed mo pho ypes o his
species we e eco ded in Cascais, Po ugal, being he ed and g een he mos com-
mon ones. Thei abundances ha e been inc easing in Po ugal (Pe ei a e al. 2022),
bu he e is no in o ma ion abou he impac o in asi eness o his species. Be-
ing in e idal, in e speci ic compe i ion is expec ed o occu , mainly due o space
esou ce limi a ions (B an e e al. 2019) and sea anemones a e known o display
in a and in e speci ic agg essi e beha iou s, which modula e hei spa ial dis ibu-
ion (e.g., Esc ibano-Ál a ez and López-González 2018).
Figu e 1. G een and ed mo pho ypes o Phymac is papillosa. Pho og aphs aken in June 2025 a Cabo Raso, Cascais, Po ugal
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NeoBio a 102: 81–91 (2025), DOI: 10.3897/neobio a.102.148042
Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
In his s udy, we assessed he in asi e po en ial o P. papillosa using wo comple-
men a y app oaches: i s , we e alua ed i s ole ance o abio ic ac o s o p edic
i s po en ial dis ibu ion along he Eu opean coas ; second, we examined i s be-
ha iou al in e ac ions - pa icula ly compe i ion - wi h na i e in e idal anemone
species o gauge i s impac on he ecosys em and cla i y i s in asi e s a us.
Me hods
Tempe a u e and salini y ole ance assessmen
A o al o 300 indi iduals o Phymac is papillosa we e collec ed in h ee ield ips
du ing he mon hs o Oc obe 2023 o Janua y 2024 in Cabo Raso, Cascais, Po -
ugal (38.70954°N, 9.4863°W) whe e he species is locally abundan . Indi iduals
o a ious pedal sizes we e selec ed, anging om 0.6 o 3.5 cm. Hal o he indi-
iduals belong o he g een mo pho ype and hal o he ed mo pho ype. The indi-
iduals we e andomly selec ed, ca e ully emo ed om he ocky subs a e, placed
in a con aine wi h seawa e , and subsequen ly anspo ed back o he labo a o y
(i.e. ISPA Bio e ium). P io o any expe imen , he o ganisms we e kep o wo
weeks in a qua an ine aqua ium a oom empe a u e (app oxima ely 16 °C) and
35 psu salini y, simila o he salini y whe e hey we e ound.
A e acclima ion, he anemones we e andomly di ided in o g oups o six indi id-
uals, hal o each mo pho ype (i.e., ed mo pho ype = 3 indi iduals; g een mo pho-
ype = 3 indi iduals), and placed in i een 22 L il e ed (wi h a biological il e wi h
a 10 mic a po e size) and ae a ed aqua ia. To e alua e he ole ance o P. papillosa o
di e en wa e empe a u es and salini ies, a wo- ac o expe imen al design was used.
Fi een aqua ia we e kep a i e di e en empe a u e ea men s (15, 19, 23, 27,
31°C) and h ee salini y condi ions (32, 35, 38 psu). The salini y and empe a u e
alues e lec he a ia ion obse ed in coas al en i onmen s in Po ugal, including
coas al lagoons (Dias e al. 1999; New on and Mudge 2003; Samma ino e al. 2022).
Animals we e placed in i een ae a ed aqua ia wi h 22 L o il e ed wa e wi hou s ep-
wise acclima ion, due o challenges in empe a u e egula ion, which equi ed ime o
s abilize. Tempe a u e and salini y we e e alua ed and co ec ed daily, and su i al was
e alua ed daily o a wo weeks’ pe iod. Fluc ua ions o up o 1.5 °C we e allowed o
he selec ed empe a u es. Due o a ia ions in salini y alues, he a e age salini y o e
he cou se o he expe imen was conside ed. Du ing acclima ion and h oughou
he expe imen al p ocedu e, sea anemones we e no ed and we e main ained unde
a ligh /da k cycle ha mimicked na u al condi ions. This expe imen was eplica ed
h ee imes wi h new collec ed indi iduals subjec ed o he same p ocedu e.
S a is ical analysis was pe o med using SPSS S a is ics e sion 29 – IBM Co p.
A non-pa ame ic Wilcoxon signed- ank es was used o compa e su i al a es
be ween sea anemone mo pho ypes, using he su i al a e pe aqua ium pe mo -
pho ype as he s a is ical uni . This es assesses whe he signi ican di e ences in
su i al exis be ween mo pho ypes, ea ing hem as pai ed wi hin each aqua -
ium. Non-linea eg essions we e pe o med o assess he e ec s o empe a u e
and salini y on he su i al a e o anemones (pe aqua ium). The mos signi ican
models we e selec ed based on highe R squa e alues. Subsequen ly, he in e ac-
ion e m be ween a iables was added o he model, and i s signi icance was e al-
ua ed using an F-change es and he co esponding p obabili y. Finally, su i al
cu e analysis was conduc ed using he Kaplan-Meie me hodology.
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NeoBio a 102: 81–91 (2025), DOI: 10.3897/neobio a.102.148042
Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
Beha iou al in e ac ions wi h he na i e in e idal anemone species
A o al o 120 indi iduals o Phymac is papillosa and app oxima ely 20 indi idu-
als o each o six na i e sea anemone species – Ac inia equina (Linnaeus, 1758),
Ac inia medi e anea Schmid , 1971, Ac inia agacea Tugwell, 1856, Anemonia
sulca a (Pennan , 1777), Ac ino hoe sphy ode a (Gosse, 1858) and Bunodac is e -
ucosa (Pennan , 1777) we e collec ed in se e al ield ips be ween Oc obe 2022
and Ma ch 2024 in Cabo Raso, Cascais, Po ugal (38.70954°N, 9.4863°W) and
Ba u ei a beach, Cascais, Po ugal (38.69206°N, 9.36588°W). Na i e species we e
selec ed based on hei local abundance and when mul iple mo pho ypes we e
p esen , indi iduals we e andomly chosen. The indi iduals we e andomly selec -
ed, ca e ully emo ed om he ocky subs a e, placed in a con aine wi h seawa-
e , and subsequen ly anspo ed o he labo a o y (i.e. ISPA Bio e ium), whe e
hey we e main ained in 20 L o 100 L il e ed and ae a ed aqua ia. Each indi id-
ual species/mo pho ype was kep in a di e en aqua ium o minimize in e speci ic
agg essi e in e ac ions be ween indi iduals. Wa e empe a u e was 17–18 °C and
salini y 35 psu and indi iduals we e ed wice a week wi h ozen mussels. P io o
any expe imen , he sea anemones we e le o a mon h o acclima ize.
A e his pe iod, indi idual es s be ween P. papillosa and each o he na i e sea
anemone species we e pe o med in 5 L aqua ia illed wi h new wa e a he same
salini y and empe a u e condi ions. Each anemone was le o a small pe iod o
a ach o a small s one. The wo s ones we e hen mo ed oge he , o allow physical
con ac be ween anemones. The in e ac ion be ween he sea anemones was ilmed
o one hou , using cell phones came as. Each indi idual was only used in one es .
The ilms we e sped up 12 imes and analysed by wo di e en obse e s using BO-
RIS (F ia d and Gamba 2016). An upda ed e hog am o sea anemone agg ession
was made (Table 1 and Fig. 2). The ou come o P. papillosa in each con es was ca -
ego ized as ollows: ‘win’ i P. papillosa e ained i s posi ion and he na i e sea anem-
one wi hd ew, ‘lose’ i P. papillosa wi hd ew and he na i e sea anemone e ained i s
posi ion, and ‘ ie’ i bo h sea anemones ei he wi hd ew o emained in hei espec-
i e posi ions (a e Esc ibano-Ál a ez and López-González 2018). A chi-squa e es
o independence was conduc ed using SPSS S a is ics e sion 29 (IBM Co p.) o
assess he ela ionship be ween es ou comes and opponen species.
Table 1. E hog am o he sea anemone species obse ed.
Beha iou Desc ip ion
Ten acula ouch A sea anemone applies i s en acles o o he sea anemone
Column expansion A sea anemone in la ed i s column in o he sea anemone di ec ion, e en ually ouching i .
O al disc applica ion A sea anemone bends in di ec ion o o he , ouching i wi h i s o al disc.
Li opponen A sea anemone uses i s en acles o li he opponen and keep i su ounded by i s en acles nea i s o al disc.
Scapus ele a ion A sea anemone ele a es he uppe pa o i s column in he opponen ’s di ec ion.
Ac o hagi use A sea anemone in la ed i s ac o hagi, bends i s body o i s opponen and e en ually ouches i wi h hese s uc u es,
which may s ay a ached o i s body.
Ten acle e ac ion A sea anemone e ac s pa o all i s en acles apidly.
Acon ia applica ion A sea anemone p ojec s acon ia om i s en acles, mou h o small po es in i s collum ha adhe e o i s opponen .
App oach A sea anemone mo es in di ec ion o i s opponen , by con ac ion and dis en ion o i s pedal disc.
Wi hd aw A sea anemone mo es away om i s opponen , by con ac ion and dis en ion o i s pedal disc.
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NeoBio a 102: 81–91 (2025), DOI: 10.3897/neobio a.102.148042
Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
Resul s
Tempe a u e and salini y ole ance assessmen
Su i al a es by empe a u e and salini y a e p esen ed in Table 2. They we e high
a empe a u es anging om 15 °C o 23 °C, dec eased sligh ly a 27 °C, and we e
ma kedly low a 31 °C (Table 2). No no iceable di e ences we e obse ed be ween
salini y le els a he same empe a u es. Mo pho ype did no ha e a signi ican
e ec on su i al (T = 11, p = 0.291).
Non-linea eg essions indica e ha only empe a u e has a signi ican impac
on P. papillosa su i al a es (Suppl. ma e ial 1). This signi ican e ec emains
consis en ega dless o he model used; howe e , he cubic model demons a ed
he bes i o he da a (z(2,42) = 142.4, p < 0.001; 2 = 0.871) (Fig. 3). The inclu-
sion o in e ac ion e ms be ween salini y and empe a u e ( empe a u e2 and em-
pe a u e3) did no signi ican ly imp o e he explained a iance, as i changes om
0.871 o 0.875 (F change2;40 = 0.638, p = 0.534).
Kaplan-Meie su i al cu es illus a e he p obabili y o anemone su i al o e
ime (Fig. 4). A 31 °C, anemone su i al begins o dec ease om he hi d day
onwa d. By he ou h day he p obabili y o su i al is less han 50%, a his em-
pe a u e. Anemones exposed o empe a u es o 15 °C, 19 °C, and 23 °C show a
highe p obabili y o su i al un il he end o he expe imen , while hose a 27 °C
exhibi a su i al p obabili y compa able o he lowe empe a u es, wi h a sligh
dec ease owa d he end o he expe imen . The Log-Rank es conduc ed showed
signi ican di e ences in su i al p obabili ies among he empe a u e g oups (X2=
345.408, p < 0.001).
Figu e 2. Obse ed agg essi e and de ensi e beha iou s. Desc ip ions o each beha iou in Table 1. The sea anemone ha displayed his
beha iou is ma ked wi h a poin benea h. a. Ten acula ouch; b. Column expansion; c. O al disc applica ion; d. Li opponen ; e. Scapus
ele a ion; . Ac o hagi use; g. Ten acle e ac ion; h. Acon ia applica ion; i. App oach; j. Wi hd aw.
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Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
Table 2. Su i al a e o P. papillosa pe aqua ia by empe a u e and salini y ea men (N = 3).
Tempe a u e (°C) Salini y (mean ± s anda d de ia ion) Su i al a e (mean ± s anda d de ia ion)
15 37.2 ± 0.41 100 ± 0
34.4 ± 0.37 100 ± 0
32.0 ± 0.28 88.9 ± 19.2
Pooled 96.3 ± 11.1
19 37.8 ± 0.53 100 ± 0
35.2 ± 0.20 94.4 ± 9.6
31.7 ± 0.68 88.9 ± 19.2
Pooled 94.4 ± 11.8
23 38.2 ± 0.37 94.4 ± 9.6
35.5 ± 0.47 94.4 ± 9.6
32.3 ± 0.29 100 ± 0
Pooled 96.3 ± 7.3
27 38.7 ± 1.05 72.2 ± 19.2
35.9 ± 0.19 94.4 ± 9.6
32.2 ± 0.67 94.4 ± 9.6
Pooled 87.0 ± 16.2
31 38.7 ± 0.33 0 ± 0
36.2 ± 1.58 0 ± 0
33.1 ± 0.77 0 ± 0
Pooled 0 ± 0
Figu e 3. Rela ionship be ween pe cen age su i al o P. papillosa and empe a u e exposu e ollowing
14 days. A signi ican cubic model is p esen ed as a black line (Su i al = 9.688 + 0.688 Tempe a u e2
– 0.022 Tempe a u e3, R2 = 0.871).
Su i al (%)
Tempe a u e (ºC)
= 9.688 + 0.688 −0.022
The ou come o all in e ac ions is summa ized in Table 3. A chi-squa e es
showed ha he ou come is signi ican ly dependen on he opponen species
(X2
10= 32.168, p < 0.001). The analysis o adjus ed s anda dized esiduals shows
ha P. papillosa wins signi ican ly mo e o en agains Ac inia equina and A. agacea
and less o en wi h B. e ucosa, and loses mo e o en wi h Anemonia sulca a and A.
sphy ode a and less o en wi h Ac inia equina.
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Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
Figu e 4. P. papillosa Kaplan-Meie su i al cu es unde di e en empe a u e ea men s.
Table 3. Ou come absolu e equency o 108 con es s be ween P. papillosa and six na i e ocky in e idal sea anemone species. Adjus ed
s anda dized chi-squa e es esiduals a e p esen ed be ween squa e b acke s (signi ican alues in bold).
Ou come esul o
P. papillosa
Opponen species
Ac inia equina Ac inia
medi e anea Ac inia agacea Anemonia sulca a Bunodac is
e ucosa
Ac ino hoe
sphy ode a
Win 10 [2.2] 6 [0.4] 11 [3.0] 3 [-1.5] 1[-2.6] 1 [-1.8]
Tie 10 [-0.6] 11 [0.5] 7 [-1.8] 10 [-0.3] 15 [2.3] 7 [-0.1]
Loose 0 [-2.1] 1 [-1.2] 1 [-1.3] 6 [2.3] 3 [0.1] 5 [2.6]
Discussion
When a non-indigenous species (NIS) is in oduced in o a new a ea, he i s s age
o in asion, i.e., anspo , is al eady comple ed. Following his ini ial in oduc-
ion, se e al ac o s con ibu e o he species’ success ul es ablishmen and sp ead,
and so o i s in asion success. One o hem is i s capaci y o acclima e o local
en i onmen al condi ions. Ou wo k showed ha empe a u e has a signi ican
e ec on he su i al o his species. A empe a u es o 15 °C, 19 °C, and 23 °C,
he specimens o P. papillosa s udied su i ed, while a 27 °C, he e was a sligh de-
c ease in su i al le els. Only a 31 °C was he e e idence o comp omised su i al
o his species, pa icula ly a e h ee days o exposu e o his empe a u e. This
esul sugges s ha he maximum ole ance limi o his species is 31 °C o less.
In ac , acco ding o he non-linea eg ession equa ion es ima ed, a 29 °C only
52% o he anemones su i e. These esul s sugges a high po en ial o his species
o colonize he Po uguese and no he n Ibe ian coas . In i s na i e habi a , his
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Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
species is exposed o a ange o empe a u es consis en wi h he esul s ob ained
(see Hausse mann 2004).The simila i y be ween empe a u es in he na i e ange
o P. papillosa and hose along he Po uguese coas likely acili a es i s success-
ul es ablishmen and po en ial in asion. Howe e , he coloniza ion o he Med-
i e anean, whe e wa e empe a u es expe ience signi ican luc ua ions be ween
win e and summe (Simolo e al. 2014), is highly comp omised du ing he sum-
me mon hs. While his species can ole a e lowe win e empe a u es, he ise in
summe empe a u es would lead o e y low su i al a es, se e ely hinde ing he
es ablishmen o he species in his a ea. Rega ding he salini y s udy, we ound no
signi ican e ec on he su i al o P. papillosa, indica ing ha his species is ole -
an o he ange o salini ies s udied (32–38 psu). The e o e, we can conclude ha
his species is capable o colonizing bo h oceanic en i onmen s and es ua ine a eas.
Despi e p o iding insigh s in o he salini y and empe a u e ole ance o P. pa-
pillosa, he expe imen al design used in his s udy p esen s se e al limi a ions. In
na u al in e idal en i onmen s, indi iduals o his species a e egula ly exposed o
apid and o en unp edic able luc ua ions in bo h empe a u e and salini y due
o idal cycles, sola adia ion, e apo a ion, and eshwa e inpu s. These dynamic
condi ions we e no eplica ed in ou expe imen , whe e mo e s able and con-
olled pa ame e s we e applied. Addi ionally, he expe imen al animals we e no
ed h oughou he ial. We adop ed his app oach o a oid con ounding a iabil-
i y in ood in ake among indi iduals o di e en sizes, bu i can in luence su i al
ou comes, pa icula ly a highe empe a u es whe e me abolic a es a e expec ed
o be ele a ed (Seebache e al. 2014). The combina ion o ene ge ic s ess om
inc eased me abolic demands and he absence o nu i ional inpu could ha e ex-
agge a ed mo ali y o physiological s ess. Gi en hese limi a ions, he esul s o
his s udy should be in e p e ed as indica i e a he han conclusi e.
These esul s, associa ed wi h he high ep oduc i e a e al eady obse ed in his
species in Po ugal (Pe ei a e al. 2022), indica e ha i can success ully es ablish and
sp ead along he Po uguese coas . In his con ex , P. papillosa is expec ed o compe e
o space, pa icula ly wi h o ganisms belonging o he same unc ional g oups ha
sha e simila needs, such as a ious species o anemones and co als (Esc ibano-Ál a-
ez and López-González 2018). Es ablishing spa ial dominance in hese species wi h-
in a habi a whe e a achmen space is limi ed inc eases he likelihood o be e de el-
oping o he essen ial biological unc ions, such as g ow h, eeding, o ep oduc ion.
In sea anemones, compe i ion o space is closely ied o compe i ion o ood,
as hese o ganisms a e slow-mo ing, emain a ached o he subs a e, and ely
on passi e mechanisms o eeding. In seden a y species ha eed by il a ion,
in e speci ic compe i ion is o en epo ed in he in e idal zone o ood and space
esou ces (B an e e al. 2019). In he Po uguese in e idal ocky coas , se e al
species o sea anemone occu , wi h some o he mos widesp ead being hose o
he genus Ac inia some o he mos widesp ead (Pe ei a e al. 2014, 2021). Ou
esul s show ha P. papillosa is pa icula ly dominan ega ding Ac inia equina and
Ac inia agacea, wo o he mos equen sea anemone species in he Po uguese
in e idal. These wo species, as P. papillosa, possess ac o hagi, speci ic s uc u es
used in in e and in aspeci ic agonis ic in e ac ions. No no iceable di e ences a e
obse ed when he cnidae cell ypes a e compa ed (desc ibed by Hausse mann
2004 and Wa s e al. 2000). Anyway, when in con ac wi h wo hese na i e spe-
cies, P. papillosa only wi hd aws in 2.5% o he es s pe o med, and globally only
in less han 15% o he es s. This means ha in 85% o he es s, P. papillosa can
89
NeoBio a 102: 81–91 (2025), DOI: 10.3897/neobio a.102.148042
Ana M. Pe ei a e al.: In asi e po en ial o Phymac is papillosa
ad e sely impac na i e species, ei he by causing hei displacemen o by compe -
ing di ec ly o esou ces wi hin he same a ea. Based on hese indings, P. papillosa
may be classi ied as an in asi e species in Po ugal, as i ul ils he c i e ia com-
monly used o de ine biological in asi eness: i is a non-na i e species capable o
es ablishing and ep oducing beyond i s na u al ange (Pe ei a e al. 2022), and i
exe s ecological impac s, pa icula ly on na i e sea anemone popula ions.
In conclusion, P. papillosa shows a wide empe a u e and salini y ole ances, ag-
g essi e dominance o e na i e sea anemones, and s ong in asi e po en ial ac oss
he Po uguese and no he n A lan ic in e idal coas s. These indings unde sco e
he u gen need o moni o ing and managemen o sa egua d ben hic communi-
ies and main ain coas al biodi e si y.
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
This s udy was unded by he Fundação pa a a Ciência e Tecnologia (FCT) Po ugal, h ough he s a egic
p ojec s UIDB/04292 and UIDP/04292 awa ded o MARE and LA/P/0069/2020 g an ed o ARNET.
Au ho con ibu ions
Concep ualiza ion: JIR, AMP. Fo mal analysis: AMP, TT, NT. In es iga ion: JF, AMP, NT, TT. Su-
pe ision: JIR, AMP. W i ing – o iginal d a : TT, NT, AMP. W i ing – e iew and edi ing: TT, JF,
AMP, JIR, NT.
Au ho ORCIDs
Ana M. Pe ei a h ps://o cid.o g/0000-0001-7616-4683
Nú ia Teixei a h ps://o cid.o g/0009-0009-1016-087X
Joana I. Robalo h ps://o cid.o g/0000-0002-7470-0574
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.
Re e ences
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