269
Seek and you shall ind: De ec ion o alien b yozoans along
he Chilean SE Paci ic coas wi h a simple and cos -e icien
me hodology
Sabine Rech1,2,3 , Leand o M. Viei a4,5 , And ea I. Va ela3,6 , Ma ía Soledad Rome o Bas ías2,7 ,
Juan Pablo Fuen es8, Dennis P. Go don9, Ma ina Cacciu olo-Said2, Ku Paschke10 ,
Ja ie Sellanes2,3 , Ma in Thiel2,3,11
1 MARE - Ma ine and En i onmen al Sciences Cen e, Regional Agency o he De elopmen o Resea ch, Technology and Inno a ion (ARDITI), Edi . Madei a
Tecnopolo, Caminho da Pen eada, Piso 2, 9020-105 Funchal, Po ugal
2 Depa amen o de Biologia Ma ina, Facul ad de Ciencias del Ma , Uni e sidad Ca ólica del No e, Coquimbo, Chile
3 Cen e o Ecology and Sus ainable Managemen o Oceanic Islands (ESMOI), Facul ad de Ciencias del Ma , Uni e sidad Ca ólica del No e, La ondo 1281,
Coquimbo, Chile
4 Labo a ó io de Es udos de B yozoa (LAEB y), Depa amen o de Zoologia, Cen o de Biociências, Uni e sidade Fede al de Pe nambuco, A . P o . Mo aes Rego
1235, Cidade Uni e si á ia, 50670-901 Reci e, PE, B azil
5 Depa men o Sciences, Na u al His o y Museum, C omwell Road, London SW7 5BD, UK
6 Depa amen odeOceanog a ía,Facul addeCienciasNa u alesyOceanog á icas,Uni e sidaddeConcepción,Ba ioUni e si a ioS/N,Concepción,Chile
7 Labo a o io de Mic oscopía Elec ónica, Dp o. de Biología Ma ina, Uni e sidad Ca ólica del No e, La ondo 1281, Coquimbo, Chile
8 Es ación Expe imen al Quempillén, Ins i u o de Ciencias Ma inas y Limnológicas, Uni e sidad Aus al de Chile, Valdi ia, Chile
9 New Zealand Ins i u e o Ea h Science (NZIES), P i a e Bag 14901, Kilbi nie, Welling on 6241, New Zealand
10 Ins i u odeAcuicul u a,Uni e sidadAus aldeChile,LosPinosS/N,Pelluco,Pue oMon ,Chile
11 Ma ineGEO, Smi hsonian En i onmen al Resea ch Cen e , 647 Con ees Wha Road, Edgewa e , MD 21037, USA
Co esponding au ho : Sabine Rech ([email p o ec ed])
Copy igh : © Sabine Rech 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
In asions by non-indigenous species (NIS) a e among he majo p oblems ha coas al ecosys ems
a e acing globally and a e d i en by an h opogenic ac i i ies, such as in e na ional essel a ic and
aquacul u e. B yozoans a e ypical cons i uen s o essel hull- ouling communi ies, and he phylum
con ains se e al no o ious globally in asi e species. Few NIS, including b yozoan species, a e e-
po ed along he Chilean Sou heas Paci ic coas , which has adi ionally been a ibu ed o he local
oceanog aphic and ecologic condi ions. Howe e , he low numbe s may also be he consequence o
insu icien moni o ing, as well as he small size and mo phologic simila i y o many species, which
hinde s hei de ec ion. He e, we deployed a i icial loa ing se lemen pla es in s a egic loca ions
(po s/ma inas o aquacul u e si es) along he Chilean coas ( om 29°S o 41°S), o de ec non-in-
digenous b yozoans. Wi h his me hod, we ound a o al o 14 species, including i e NIS and ou
c yp ogenic species. Addi ionally, we epo wo NIS om oppo unis ic indings in he ocky in e -
idal zone and om a piece o plas ic li e . Th ee species ep esen new eco ds om Chile: Bugulina
c . ul a, Bugulina s oloni e a, and Wa e sipo a a cua a. These a e desc ibed in de ail in he p esen
pape . The new eco ds impo an ly enhance he numbe o known (b yozoan) in asions along he
Chilean coas and co obo a e he need o egula moni o ing. The ins alla ion o a i icial se le-
men pla es a high- isk si es (especially po s and ma inas) has p o en o be a success ul and e icien
low-cos app oach, which can help o de ec NIS in oduc ions in hei ea ly s ages, be o e sp eading
o na u al en i onmen s. Regula moni o ing o hese si es, as well as compulso y legisla ion o hull
cleaning, bo h o long-dis ance (na ionally and in e na ionally) a elling and local essels, could
g ea ly educe he p oblem.
Academic edi o : Paula Chainho
Recei ed:
17 Decembe 2024
Accep ed:
14 May 2025
Published:
7 Oc obe 2025
Ci a ion: Rech S, Viei a LM, Va ela
AI, Bas ías MSR, Fuen es JP, Go don
DP, Cacciu olo-Said M, Paschke K,
Sellanes J, Thiel M (2025) Seek and
you shall ind: De ec ion o alien
b yozoans along he Chilean SE
Paci ic coas wi h a simple and cos -
e icien me hodology. In: Anas ácio P,
Ribei o F, Chainho P (Eds) In asions
in Aqua ic Sys ems. NeoBio a 102:
269–294. h ps://doi.o g/10.3897/
neobio a.102.144725
NeoBio a 102: 269–294 (2025)
DOI: 10.3897/neobio a.102.144725
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: 269–294 (2025), DOI: 10.3897/neobio a.102.144725
Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
Key wo ds: B yozoa, hull ouling, ma ine in asions, non-indigenous species, se lemen pla es,
Sou heas Paci ic
In oduc ion
Non-indigenous species (NIS) a e a global conce n due o hei e ec s on na i e
communi ies, global biodi e si y, and human li elihoods (Pyšek e al. 2020). Se -
e al an h opogenic ec o s anspo hese species ac oss he wo ld’s oceans o new
coas al habi a s. Amongs hose, essel a ic seems o be he quickes and mos
success ul o m o anspo o sessile ouling species (e.g., Seebens e al. 2013;
Ruiz e al. 2015; Ca l on e al. 2019), in oducing hem o po a eas a ound he
wo ld. Ou side he main shipping ou es, loa ing ma ine li e is also impo -
an , as i dispe ses wi h oceanic cu en s and eaches e en e y emo e loca ions
(Kiessling e al. 2015; Rech e al. 2021, 2023).
B yozoans a e among he mos equen g oups o ma i ime hull- ouling o gan-
isms (e.g., McCann e al. 2019; Rech e al. 2024a). Some b yozoan NIS a e al eady
widesp ead and epo ed om many pa s o he wo ld, whe e hei abili y o oul
la ge su aces makes hem a nuisance and a h ea (e.g., Nascimen o e al. 2022).
Howe e , in e -species pheno ypic simila i y, in a-species pheno ypic plas ici y,
and his o ic axonomic con usion hinde he de ec ion and iden i ica ion o many
b yozoan NIS. Some o hem can be dis inguished only h ough elabo a e ex-si u
analyses, like scanning elec on mic oscope (SEM) imaging o gene ic ba coding,
and hey a e easily o e looked o con ounded wi h simila species in ield su eys.
Mo eo e , mos o he wo ld’s coas al egions a e se e ely unde s udied and ha e
no egula moni o ing in place. Recen s udies in o he wise poo ly sampled loca-
ions ound ha non-na i e b yozoans and e en undesc ibed species a e ela i ely
common in such egions. Fo example, Viei a e al. (2012) desc ibed 9 new species
o Bugulidae om ou unde s udied egions o he B azilian coas , based on man-
ual samplings. Simila ly, Rech e al. (2024a) ecen ly published he i s eco d o
he widesp ead ouling species Wa e sipo a sub o qua a on emo e Rapa Nui (Eas e
Island), based on a ou -mon h se lemen pla e-based su ey.
In he pas , he Sou h Ame ican Paci ic coas has been p oposed o be ela i ely
“mo e p is ine” o mo e esis an o bioin asions han o he global egions, due
o he small numbe o high-impac NIS ound he e (Cas illa and Neill 2009).
Howe e , a eliable es ima ion o NIS equency along he SE Paci ic coas line is
no possible due o a lack o sampling and egula moni o ing, meaning ha ac ual
numbe s o NIS a e p obably much highe han hose epo ed (Ca l on 2009;
S owhas e al. 2023). Cas illa e al. (2005) and Cas illa and Neill (2009) lis ed only
wo b yozoan NIS in he Eas e n Sou h Paci ic: Bugulina labella a and Bugula ne -
i ina. Ano he NIS, C yp osula pallasiana, was epo ed om aquacul u e buoys in
he no he n-cen al Coquimbo egion (As udillo e al. 2009). Mo eo e , Lecle c
e al. (2018, 2020a, b) ecen ly epo ed se en addi ional non-indigenous o c yp-
ogenic species (Alcyonidioides my ilii, Ama hia c . g acilis, Ama hia c . imb ica a,
Caulo amphus spini e um, Conopeum e iculum, Elec a monos achys, and Exochella
n. sp.) om su eys in Sou he n Chilean po s, and Rech e al. (2024a) epo ed
he globally in ading NIS W. sub o qua a om Rapa Nui (Eas e Island).
In con as o many o he axa, he Chilean b yozoan auna is well in es iga ed
and desc ibed, la gely hanks o he wo k o Hugo Moyano, who s udied he phy-
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
lum o mo e han 3 decades ( om he ea ly 1970s o he ea ly 2000s). Despi e his
ex ensi e s udies, he au ho epo ed only wo b yozoan NIS, B. ne i ina and B.
labella a (Moyano 1982, 1991), om Chile. On he one hand, his could sugges
ha mos b yozoan in asions along he Chilean coas a e indeed ecen (i.e., om
he las 20 yea s). On he o he hand, Moyano’s s udies ocused on na u al ma ine
habi a s, whe eas ecen ly epo ed NIS we e ound in po s and/o on a i icial
su aces (Rech e al. 2018, 2024a; Lecle c e al 2018, 2020a, b), highligh ing he
impo ance o hese loca ions and subs a a as en y poin s and p opagule ese -
oi s o ma ine NIS. Mo eo e , Moyano´s comp ehensi e desc ip ion o he Chil-
ean B yozoa is based on he s udy o speci ic habi a s and si es a speci ic poin s o
ime and does no ep esen a egula moni o ing o he 2000 km o Chilean coas -
line, which comp ises a la ge a ie y o coas al habi a s, bo h na u al and a i icial.
Chile has a long his o y o in e na ional essel a ic, wi h 39 in e na ional
po s and mo e han 5000 ship a i als pe yea (S owhas e al. 2023), as well
as abou 20 ma inas (h ps://www.ha bou maps.com/en/ha bou s/ egion/chile).
Such local ha bou s a e known o play an impo an ole in seconda y NIS dispe -
sal (e.g., Ash on e al. 2022a). Se e al o he non-indigenous b yozoans epo ed
om Chile so a a e known o ouling ship hulls (e.g., Gollasch 2002; Ramalhosa
e al. 2017). Mo eo e , some o hem a e known o a ach o loa ing deb is, which
may also be impo an o seconda y dispe sal along he Chilean coas (Rech e al.
2023). Ano he ec o o NIS in oduc ions is he in en ional o unin en ional e-
lease om aquacul u e acili ies. While egula ion o his sec o has led o dec eases
in NIS in oduc ions in, o example, Eu ope and Cali o nia (Ka sane akis e al.
2013; Williams e al. 2013), exis ing policies seem o be insu icien o ine ec i e
in Chile (Camus 2005; S owhas e al. 2023). In addi ion o he o ganisms di ec -
ly eleased om aquacul u e si es, NIS can also be dispe sed when aquacul u e
equipmen and gea is in en ionally mo ed (e.g., Iaca ella e al. 2019) o acciden-
ally los om he a ming si es (e.g., As udillo e al. 2009; Campbell e al. 2017;
Bee mann e al. 2025).
Conside ing all he abo e, we hypo hesized ha unde ec ed o o e looked b yo-
zoan NIS a e p obably p esen in high- isk si es associa ed wi h in e na ional, na-
ional, and local essel a ic and/o aquacul u e along he Chilean coas . The e-
o e, he goal o he p esen s udy was he de ec ion and iden i ica ion o such NIS
wi h a simple and well- es ed se lemen pla e-based app oach (e.g., Tambu ini e
al. 2021). Apa om his obse a ional app oach, we epo addi ional new ind-
ings o B yozoa NIS om Chilean coas al si es.
Me hods
S udy egion
The s udy was conduc ed along he Sou heas Paci ic Ocean, co e ing he Chilean
con inen al coas om 29°S o 41°S and Rapa Nui (Eas e Island, 27°S, 109°W).
Rapa Nui is si ua ed abou 4000 km o sho e and is connec ed o he con inen ia
he cu en s o he Sou h Paci ic Sub opical Gy e and egula essel a ic (Fig.
1). Pla e-based se lemen su eys we e conduc ed a six s a egic loca ions (po s/
yach ing clubs and/o aquacul u e si es; Suppl. ma e ial 1: able S1). On Rapa
Nui, we chose he main po , Hanga Piko (27.15°S, 109.43°W) ha ha bou s
local ishing boa s and ba ges, used o e ch goods om ships a he o sho e an-
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
Figu e 1. S udy egion wi h densi y map o o al essel a ic (yea 2023), anging om 1 (blue)
o > 5000 (da k ed) ou es * 9.78 km-2 yea -1. Sou ce: h ps://www.ma ine a ic.com/. S udy si es
a e indica ed wi h le e s: a. Rapa Nui (Eas e Island), Hanga Piko ha bou . Pho o c edi : Pamela
A e ill. b. Coquimbo Bay. Pho o c edi : Ma ina Cacciu olo-Said. c. San An onio In e na ional
Po . Pho o c edi : Sabine Rech. d. Alga obo, Co adía Náu ica del Paci ico. Pho o c edi : Ma iano
Galdames Beckdo . e. Pue o Mon , Yach ing Club Relonca í. Pho o c edi : Sabine Rech. . Ancud,
Quempillén Resea ch S a ion. Pho o c edi : Sabine Rech. Black a ows poin o loa ing ame wi h
se lemen pla es (no isible in pic u es b, d and ). Whi e a ow indica es he si e whe e Wa e sipo a
a cua a was ound in ading he ocky sea loo a he Coquimbo si e.
cho age si e o c uise ships and comme cial essels, loca ed abou 500 m o sho e.
On he con inen al coas , ou no he nmos s udy si e was a small aquacul u e
concession (29.97°S, 71.35°W), belonging o he campus o Uni e sidad Ca ólica
del No e in he p o ec ed Coquimbo Bay. Adjacen o he si e is he local ishing
po o Guayacán and he ancho age si e o la ge comme cial essels associa ed
wi h he local i on mining indus y ( essel can be seen in Fig. 1b). In he cen al
egion o he Chilean coas , se lemen pla es we e ins alled in he In e na ional
Po o San An onio (33.58°S, 71.62°W) and a a local yach ing club (33.36°S,
71.69°W), bo h o which a e si ua ed di ec ly on he exposed coas line, p o ec ed
273
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
only by b eakwa e s. In he sou he n egion, ou s udy si es we e in Pue o Mon
and Ancud and we e no di ec ly exposed o he sea. Pue o Mon is loca ed on
he coas o he Inne Sea o Chiloé. The e, se lemen su eys we e conduc ed a a
yach ing club (41.49°S, 72.99°W) loca ed in a na ow jo d. In Ancud, se lemen
pla es we e ins alled close o a small aquacul u e concession, loca ed in he Quem-
pillén Ri e es ua y (41.87°S, 73.78°W).
Sampling app oach
Se lemen pla es we e ins alled be ween Feb ua y and No embe 2022. I possi-
ble, se lemen su eys we e conduc ed wice, once in he aus al sp ing/summe
and once in he aus al au umn/win e , a each si e. Howe e , due o limi a ions
ela ed o he COVID19 pandemic, hey could only be ca ied ou in one season
a Hanga Piko and San An onio. Se lemen pla es we e deployed o 4 mon hs pe
season a each si e. The exac da es o ins alla ion and e ie al a e speci ied in Ta-
ble S1. A each loca ion, wo loa ing uni s, each con aining ou 13.5 × 13.5 cm2
poly inylchlo ide (PVC) se lemen pla es inside o a PVC ame, we e ins alled
a a p o ec ed si e (ma inas/po s o aquacul u e ins alla ions; see p e ious sec ion
“s udy egion”). A e 4 mon hs o imme sion, pla es we e e ie ed swimming o
om a boa . Fo a mo e de ailed desc ip ion o he loa ing uni s and sampling p o-
ocols, please see Rech e al. (2024a). In he po o San An onio, only one loa ing
s uc u e (= 4 se lemen pla es) could be e ie ed, while he second s uc u e was
los in a s o m e en . A all o he si es, bo h s uc u es (= 8 se lemen pla es) we e
success ully e ie ed. A each s udy si e, a i s analysis, including aking pho os
and c ea ing a p elimina y species/ axon lis , was pe o med in a nea by ield lab-
o a o y. In ag eemen wi h o he pla e-based se lemen s udies (e.g., Ash on e al.
(2022b), all analyses we e only conduc ed on he lowe (= sea loo - acing) side o
he pla es, as he ouling auna was almos exclusi ely concen a ed on his side.
A e he p elimina y analysis, pla es and hei a ached epi auna we e p ese ed in
95% e hanol and anspo ed o he ma ine biology labo a o y a he Ma ine Sci-
ence Facul y o Uni e sidad Ca ólica del No e in Coquimbo. The e, pla es we e
immedia ely ozen and s o ed a -20 °C.
Fo subsequen analyses, pla es we e hawed, and he a ached ouling communi-
y, which was o en e y abundan and consis ed o se e al phyla (Suppl. ma e ial 1:
ig. S1, da a no published), was analysed and iden i ied o he ines possible le el
wi h a dissec ing mic oscope while s ill a ached. The ex ensi e communi ies had o
be analysed and cha ac e ised wi hin a limi ed ime ame, due o he apid decom-
posi ion o he la ge so -bodied animals (e.g., soli a y ascidians). In consequence,
smalle specimens o simila -looking species like Bugulina c . labella a and Buguli-
na s oloni e a could no always be dis inguished immedia ely and we e eco ded as
Bugulidae spp. I he ouling co e on se lemen pla es was e y abundan , which
was he case a se e al con inen al s udy si es, he dense uppe laye s o bio ouling
we e analysed i s and ca e ully emo ed be o e species/specimens on he g ound o
he se lemen pla e could be de ec ed. Samples o each dis inguishable axon we e
aken o each s udy si e and season (bu no om each pla e a each si e) and s o ed
in 95% e hanol. Samples con aining b yozoans we e hen sea ched o p e iously
uniden i ied species, which we e sepa a ed and iden i ied as desc ibed below.
In addi ion o he species ound on se lemen pla es, wo addi ional species o
in asi e b yozoans we e ound oppo unis ically: One species was ound in he
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
local in e idal o he uni e si y campus o Uni e sidad Ca ólica del No e in Co-
quimbo and ano he one was iden i ied on a piece o plas ic li e om an ea lie
s udy (Rech e al. 2023).
All samples we e analysed ia ligh mic oscope (LM), scanning elec on mic o-
scope (SEM), and/o DNA ba coding (see below). Species we e i s en a i ely
iden i ied mo phologically, using a dissec ing mic oscope, and om LM images,
and hei iden i y was con i med based on SEM images and DNA ba coding. The
emaining e hanol-p ese ed colony agmen s we e deposi ed in he zoological
collec ions o he Facul y o Ma ine Sciences a Uni e sidad Ca ólica del No e in
Coquimbo, Chile (SCBUCN; see Suppl. ma e ial 1: able S2 o de ails and mu-
seum codes). Measu emen s we e made om SEM o LM images, using he Im-
ageJ so wa e (Schneide e al. 2012). Specimens om he b yozoan collec ion a
he Na u al His o y Museum, London (NHMUK) and he Smi hsonian Na ional
Museum o Na u al His o y (USNM) we e used o compa ison wi h ou samples.
SEM analyses
Fo SEM analysis, he samples we e d ied in an o en a 40 °C o 24 hou s, moun -
ed on slide glasses using double-sided ca bon ape and coa ed wi h gold in a JEOL
JFC-100 e apo a o . The specimens we e pho og aphed unde a JEOL IT300LV
SEM in he Mic oscopy Labo a o y o Uni e sidad de Chile, San iago, Chile.
DNA ba coding
DNA was ex ac ed using DNeasy Blood & Tissue Ki (Qiagen®). A segmen o
he cy och ome c oxidase subuni I (COI) gene was ampli ied using he uni e sal
p ime s HCO2198 & LCO1490 (Folme e al. 1994). Polyme ase chain eac ion
(PCR) was pe o med wi h he ollowing eagen s and quan i ies: ~ 50 ng o DNA,
1X PCR bu e (The moPol bu e , New England BioLabs), 2 mM MgCl2, 0.6µM
o each p ime , 0.25 mM o each dNTPs, 1U o Taq polyme ase (New England
BioLabs), and 0.06 mg mL-1 o Bo ine Se um Albumin (BSA). PCR cycles we e
pe o med on an DLAB (TC1000-G) The mal Cycle , unde he ollowing condi-
ions: 94 °C o 3 min, ollowed by 40 cycles o 94 °C o 30 s, 55 °C o 30 s, 72 °C
o 60 s, and a inal ex ension a 72 °C o 10 min. PCR ampli ica ion was e i ied
using 1% aga ose gels. Sange sequencing was pe o med by Mac ogen in San iago,
Chile, using he same p ime s. DNA sequences we e analysed wi h Geneious P ime®
2023.2.1 and compa ed agains he Genbank da abase, using he online BLAST
unc ion (h ps://blas .ncbi.nlm.nih.go /Blas .cgi). Ma ching sequences we e ali-
da ed, and only sequences co esponding o a icles published in scien i ic jou nals,
iden i ied by in e na ionally enowned axonomis s, and wi h a specimen ouche
we e conside ed. The DNA ba codes we e deposi ed in he Genbank da abase (ac-
cession numbe s PP109395–PP109397) and can also be ound in he supplemen .
S a is ical analyses
Species ichness and communi y composi ion in se lemen pla e-based su eys we e
calcula ed based on he p esence/absence o each o 12 dis inguishable b yozoan
axa: Bugulidae spp. (including Bugulina c . labella a and B. s oloni e a), C enos-
oma ida sp. (including Ama hia c . g acilis), Bugula ne i ina, C yp osula pallasiana,
275
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
Cheilos oma ida sp., Alcyonidiidae sp., Sc upa ia ambigua, Cellepo ella c . hyalina,
Conopeum sp., Exochella sp., Lagenicella a iabilis, and Wa e sipo a sub o qua a. Spe-
cies ichness was calcula ed as he o al numbe o species pe independen sampling
uni (= loa ing ame, n = 2 ames pe combina ion o si e and season, excep o
San An onio, whe e n = 1, see sec ion “sampling app oach” abo e). Species compo-
si ion was de ined by he pe cen age o se lemen pla es ouled by each species pe
sampling uni a each combina ion o si e and season.
Species ichness and composi ion we e compa ed be ween seasons (summe ,
win e ; ixed ac o ) and si es (Coquimbo, Alga obo, San An onio, Pue o Mon ,
Ancud, Rapa Nui; ixed ac o ), using pe mu a ional ANOVA (PERMANOVA).
The es was based on an Euclidean dis ance ma ix o ichness. Fo species compo-
si ion, i was based on a B ay-Cu is simila i y ma ix, a e excluding bo h samples
om he Ancud si e in win e , as hey ca ied no b yozoans. PERMANOVA was
un wi h a pe mu a ion o esiduals unde a educed model as pe mu a ion me h-
od, sums o squa es ype III, and 9999 pe mu a ions, using he PERMANOVA+
unc ion (Ande son e al. 2008) o he so wa e P ime 6 (Cla ke and Go ley 2006).
Non-me ic mul idimensional scaling was applied o a B ay Cu is simila i y ma-
ix, based on he a e age equency o each species o each combina ion o s udy
si e and season, a e excluding samples om he Ancud si e in win e (no b yozo-
ans p esen ), and om Rapa Nui (no common species wi h any o he s udy si e).
Resul s
Se lemen pla e-based su eys a s a egic si es along he Chilean coas esul ed in he
de ec ion o 14 species o B yozoa, including i e non-indigenous, ou c yp ogenic,
one na i e, and se e al uniden i ied species (Table 1). Non-indigenous b yozoans
we e ound a all s udy si es, excep o he es ua ine loca ion on he sou he n Chil-
ean inne coas (Ancud; see Table 1). The p e iously un epo ed B. s oloni e a was
iden i ied om h ee s udy si es, eaching om he no he n o he sou he n egion
(Table 1). Ano he p e iously un epo ed species, Bugulina c . ul a, was ound a -
ached o some e hanol-p ese ed samples o o he Bugulidae om pla es a he Co-
quimbo si e, du ing a dedica ed examina ion o hose samples. Gi en he e y small
size o he specimens (~5 mm) and hei o en epibion ic g ow h on simila -looking
bugulid species, i canno be excluded ha his species has also been p esen in o he
loca ions. In e es ingly, he e was no conco dance in species occu ence be ween he
Chilean con inen al coas and oceanic Rapa Nui, whe e only one species, he NIS W.
sub o qua a, was ound on se lemen pla es ( esul s published in Rech e al. 2024a).
Species ichness was highes a he yach ing club o Pue o Mon in bo h sea-
sons (6.5 ± 0.7 and 6.0 ± 0.0 species in win e s. summe ; Suppl. ma e ial 1:
ig. S2). In win e , he lowes species ichness was ound a he es ua ine si e in
Ancud, whe e no b yozoans had a ached o he pla es. In summe , he lowes
species ichness was eco ded on pla es in he Hanga Piko ha bou o Rapa Nui,
whe e only one b yozoan species was ound. Richness was s ongly in luenced by
si e, and wi h a weake e ec by season, and he in e ac ion o bo h ac o s (PER-
MANOVA, p ≤ 0.0003 o each ac o , see Suppl. ma e ial 1: able S3), bu no
o e all la i udinal pa e n was obse ed (Suppl. ma e ial 1: ig. S2). Communi y
composi ion a ied be ween s udy si es and, o a much lesse ex en , due o he
in e ac i e e ec o s udy si e and season (PERMANOVA, p < 0.05 o each ac o ;
Suppl. ma e ial 1: able S3), bu season alone had no o e all e ec . While oceanic
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
Rapa Nui sha ed no species wi h any o he si e, he es ua ine si e in Ancud had
he mos dissimila communi y composi ion among he con inen al si es (Suppl.
ma e ial 1: ig. S3, able S4).
Non-indigenous and c yp ogenic species domina ed he ouling B yozoan com-
muni ies: Bugulidae spp. (iden i ied in se e al samples as Bugulina c . labella a
and/o Bugulina s oloni e a) had he highes a e age abundances ac oss all si es and
seasons, ouling 69 ± 48% o se lemen pla es. They we e ollowed by he c yp o-
genic C enos oma ida sp. (iden i ied in se e al samples as Ama hia c . g acilis; a e -
age abundance: 55 ± 50%) and by Bugula ne i ina (a e age abundance: 40 ± 47%,
Suppl. ma e ial 1: able S4). The only na i e species, Lagenicella a iabilis, was only
iden i ied on one se lemen pla e a he Coquimbo si e in summe . In addi ion o
he species g owing on se lemen pla es, wo mo e NIS we e ound oppo unis i-
cally: Wa e sipo a a cua a was de ec ed by s uden s o he Ma ine Biology Facul y
in he in e idal zone a he uni e si y campus o Uni e sidad Ca ólica del No e
in Coquimbo. Elec a monos achys had been ound on wo pieces o plas ic li e
in a p e ious s udy ca ied ou a Ri oque beach (Rech e al. 2023), bu was only
ecen ly iden i ied a species le el.
O he ele en non-indigenous/c yp ogenic species epo ed in he p esen wo k,
wo (Bugula ne i ina and Bugulina c . labella a) a e commonly ound along he
Chilean coas and ha e been epo ed he e since he ea ly 1980s (Moyano 1982).
Fi e o he species, Ama hia c . g acilis, Cellepo ella c . hyalina, C yp osula pallasi-
ana, Elec a monos achys, and Sc upa ia ambigua ha e been epo ed in p e ious
Table 1. B yozoan species ound a each s udy si e. ✔ = species p esen . nd = no da a, oppo unis ic eco d. C yp = c yp ogenic, NIS =
non-indigenous species, na = na i e. P e ious eco ds: 1 = Lecle c e al. 2018; 2 = Moyano 1991; 3= As udillo e al. 2009; 4 = Rech e al.
2024a. * = classi ied as “na i e” in Lecle c e al. (2018). Bold w i ing = i s eco ds om Chile.
Hanga Piko
(27°S)
Coquimbo
(29°S)
Ri oque
(32°S)
Alga obo
(33°S)
San
An onio
(33°S)
Pue o
Mon
(41°S)
Ancud
(41°S)
a) S anda dized su eys: Se lemen pla es
Alcyonidiidae sp. nd ✔ ✔
Ama hia c . g acilis1c yp ✔nd ✔ ✔ ✔
Bugulina c . labella a2NIS ✔nd ✔ ✔ ✔
Bugulina c . ul a c yp ✔nd
Bugula ne i ina2NIS ✔nd ✔✔✔
Bugulina s oloni e a NIS ✔nd ✔ ✔
Cellepo ella c . hyalina1c yp nd ✔
Cheilos oma ida sp. nd ✔
Conopeum sp. nd ✔
C yp osula pallasiana3NIS ✔nd ✔ ✔
Exochella sp. nd ✔
Lagenicella a iabilis na ✔nd
Sc upa ia ambigua1* c yp nd ✔
Wa e sipo a sub o qua a4NIS ✔nd
b) Addi ional: Na u al in e idal
Wa e sipo a a cua a NIS nd ✔nd nd nd nd nd
c) Addi ional: S anded plas ic li e
Elec a monos achys1NIS nd nd ✔nd nd nd nd
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s udies om Chilean po s (Lecle c e al. 2018; 2020b). Sc upa ia ambigua had
been ea ed as na i e o Chile in Lecle c e al. (2018, 2020a, b) bu , gi en ha he
na i e ange o his cosmopoli an species is no unequi ocally esol ed, we ollow
he classi ica ion o Schwind e al. (2020), who classi y i as c yp ogenic e en in
he Sou hwes A lan ic, om whe e i was o iginally desc ibed (d´O bigny, 1841).
SEM images we e ob ained om E. monos achys and a e shown in Suppl. ma e ial
1: ig. S4. Finally, h ee o he species ound he e (Bugulina c . ul a, Bugulina
s oloni e a, and W. a cua a) ep esen i s eco ds o Chile. De ailed desc ip ions
and (SEM) images o hese species a e p esen ed below.
Sys ema ic Accoun
Family Bugulidae (G ay, 1848)
Genus Bugulina (G ay, 1848)
Bugulina s oloni e a (Ryland, 1960)
Bugula s oloni e a Ryland 1960: 78, ig. 6; P enan and Bobin 1966: 541, ig. 158
XII, 159 II, 187; Ryland and Haywa d 1977: 170, ig. 82; Haywa d and Ryland
1998: 228, ig. 72; Go don and Mawa a i 1992: 23, Pla e 6a; Scholz e al. 2003;
Chimenz Gusso e al. 2014.
Bugulina s oloni e a: Fehlaue -Ale e al. 2015; Ramalhosa e al. 2017: 9, ig. 7.
Ma e ial analyzed. Colonies om se lemen pla es ( ozen a -20 °C) on loa -
ing de ices. UCN uni e si y campus, Coquimbo, Chile (29.97°S, 71.35°W), SC-
BUCN-8176, Genbank PP109395, ID COQWINBX20230712_02 (Fig. 2), col-
lec ed 22 June 2022; SCBUCN-8177, Genbank PP109396, ID COQ1_13.09/
COQSUMBS (Fig. 2), collec ed 7 Feb ua y 2023. San An onio In e na ional po ,
Chile (33.58°S, 71.62°W), SCBUCN-8178, ID SANBX01 (Fig. 3), collec ed 18
Sep embe 2022. Club Náu ico Relonca í, Pue o Mon , Chile: SCBUCN-8179,
ID PMSUMBX20230712_01 (Fig. 3), collec ed 23 Janua y 2023.
Compa a i e ma e ial analyzed. NHMUK 1959.3.18.1, Bugula s oloni e a
Ryland, 1960, holo ype, Swansea Docks, UK.
Cha ac e iza ion. Colony e ec , u ed, whi ish in colou , bise ially b anching,
wi h bi u ca ion ype 4 (see Ryland 1960). Specimens a he small, maximum
heigh o 2.5 cm. Zooids long, wi h leng h and wid h a ying be ween specimens
om di e en loca ions ( o measu emen s see Suppl. ma e ial 1: able S5). Dis al
zooid wi h one inne and wo ou e spines. Peduncula e a icula ia dimo phic,
p esen a he ou e side o he zooids, sligh ly longe han zooid wid h, bu smalle
ones can be ound a bi u ca ions (Fig. 2c, d), wi h down-cu ed beak (Figs 2e,
3d), abou 1.5 o 2 imes as long as wide (Suppl. ma e ial 1: able S5). O icells sub-
globula , 0.6–0.7 imes as long as wide, cen e ed in midline, dis inc , wi h on al
memb anous a ea (Fig. 2c, d).
Rema ks. The specimens o en co-occu ed wi h he congene Bugulina c . la-
bella a and may ha e been con ounded wi h he p io species in pas apid assess-
men su eys in he ield. Bugulina s oloni e a had no been epo ed om he
Sou heas Paci ic coas so a ; i is known, howe e , om he Galapagos Islands
(McCann e al. 2019). COI sequences o ou specimens (Suppl. ma e ial 1: able
S6) om Coquimbo ma ch e i ied eco ds om Spain (Fehlaue -Ale e al. 2015),
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
in he Sou heas Paci ic. W. a cua a di e s om bo h in he p esence o pai ed dis al
in azooidal sep ula. Mo eo e , W. sub o qua a and W. nig a a e espec i ely cha -
ac e ised by a U-shaped and squa ed sinus (see Rech e al. 2024a o W. sub o qua a
and Suppl. ma e ial 1: ig. S5 o W. nig a), while W. a cua a has a mo e ec angula
sinus (Fig. 8 c, d). Visual iden i ica ion o ou specimens as W. a cua a is co obo-
a ed by gene ic analysis. Ou ma e ial is highly simila (98 – 100% simila i y) o
published COI sequences om he No heas and Sou hwes Paci ic (Cali o nia,
Hawaii, Aus alia; see Suppl. ma e ial 1: able S10).
Discussion
Pa e ns o species ichness and communi y composi ion
We de ec ed a o al o six een b yozoan species, including ele en non-indigenous
o c yp ogenic species, along he Chilean coas using a combined app oach o s an-
da dised se lemen pla e-based su eys and addi ional eco ds. This includes h ee
new species eco ds o Chile. Conside ing he low numbe o known NIS on he
Chilean con inen al coas (e.g., S owhas e al. 2023), hese addi ions ep esen a
signi ican upda e o he local non-indigenous auna. Despi e si e and season- e-
la ed di e ences in species ichness, he e was no o e all geog aphic o seasonal
pa e n. Each s udy si e had a cha ac e is ic indi idual composi ional p o ile, and
Figu e 8. Wa e sipo a a cua a. samples SCBUCN8181 (a, c), and sample SCBUCN10539 (b, d). a, b. Colony o e iew; c, d. O i ice
wi h in azooidal sep ula: dis = dis al, pos = pos e io , e = e ical.
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
he highes en i onmen al and geog aphic dissimila i y coincided wi h he high-
es dissimila i y in communi y composi ion. This was seen o oceanic Rapa Nui,
which sha ed no species wi h he con inen al si es and o he only es ua ine si e,
Ancud, whe e species composi ion had he leas simila i y wi h all o he con inen-
al si es. These dissimila i ies may be based on geog aphic ac o s, like dis ance o
he mainland ( o Rapa Nui), o en i onmen al condi ions, like salini y ( o An-
cud) o wa e empe a u e. O he ac o s in luencing (in asi e o c yp ogenic) spe-
cies ichness and composi ion may be he in ensi y o long-dis ance and local essel
a ic, as well as he p esence and abundance o ouling auna al eady p esen a he
espec i e si e (see Lecle c e al. 2020a). Despi e hese conside a ions, he b yozoan
species analysed he e we e pa o a mo e di e se ouling communi y g owing on
se lemen pla es a each sampling si e, which may in luence he ichness and abun-
dance o ou a ge species h ough ecologic in e ac ions. The e o e, he p esen
s udy, which is ocussed on a axonomic desc ip ion o non-indigenous B yozoa,
was no designed o and canno disce n he causes o he obse ed di e ences.
NIS in oduc ion
Vessel a ic is one o he mos common ec o s o ma ine in asions (e.g., Cas o
e al. 2020; Bailey e al. 2020), wi h po s se ing as key si es o NIS in oduc ion
(e.g., Lecle c e al. 2018; Pinoche e al. 2023; Rech e al. 2024a). In ac , 30–40%
o Chilean NIS, including he b yozoans Bugula ne i ina and Bugulina c . labella-
a, a e conside ed o be ship-bo ne in oduc ions (Cas illa and Neill 2009). This is
likely also he case o he newly epo ed non-indigenous and c yp ogenic species
in ou s udy, which a e ypical cons i uen s o ma ine ouling communi ies and
we e ound in po s and ma inas. Howe e , iden i ying he ec o and geog aphic
sou ce o an in oduc ion can be challenging, pa icula ly when i is unclea how
long a NIS has been p esen a a si e be o e de ec ion. This is ue o he wo
newly eco ded species, Bugulina c . ul a and Bugulina s oloni e a, bo h o which
a e easily con used wi h he widesp ead Bugulina c . labella a in apid ield su eys
wi hou mic oscopic analysis and may ha e gone unde ec ed o ex ended pe iods.
A no ewo hy inding in ou s udy is he disco e y o Wa e sipo a a cua a in he
ocky in e idal zone a he Coquimbo si e. This si e is equen ed by s uden s om
he adjacen Facul y o Ma ine Sciences and was also isi ed du ing a ma ine bio-
in asions wo kshop in 2019. Howe e , W. a cua a was only de ec ed in Sep embe
2023, when s uden s iden i ied dis inc i e black pa ches wi h eddish edges co e ing
he in e idal ocks and s ones. The colonies we e abundan and easily isible a low
ide, sugges ing ha his is a ecen in oduc ion (likely wi hin he las i e yea s).
The in aded si e is loca ed nea he local mining po o in e na ional ca go essels,
whe e an a e age o 23 essels a i ed annually be ween 2019 and 2023. Mos o
hese essels came om Asian coun ies (66%), pa icula ly China and Sou h Ko ea
(Suppl. ma e ial 1: able S11), and s ayed in po o an a e age o 7 days. Vessels
om Sou h Ame ica and Cen al Ame ica also a i ed, hough wi h lowe equen-
cies (24% and 10%, espec i ely), and sho e s ays (Suppl. ma e ial 1: able S11).
Conside ing he na u e o W. a cua a as a ypical ouling species and he loca-
ion o he colonies, i seems likely ha his species was in oduced ia in e na-
ionally a elling essels a i ing a he mining po . Howe e , he na i e ange o
W. a cua a is unknown, and i has no been eco ded om Asia, mainland Sou h
Ame ica, and Cen al Ame ica in he scien i ic li e a u e. While he e a e eco ds
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
o W. a cua a om he US Eas Coas (Viei a e al. 2014), only one o he 115
essels a i ing a he Guayacán e minal be ween 2019 and 2023 came om ha
egion. The e o e, gi en he lack o o e lap be ween he known dis ibu ion o W.
a cua a and he o igin o he essels a i ing a he Guayacán e minal, we canno
con iden ly in e he geog aphic o igin o he colonies ound in Coquimbo, Chile.
Es ablishmen and local dispe sal
Po s no only se e as impo an en y poin s o NIS, bu hey also p o ide ideal
habi a s o hese species a e hei in oduc ion. In ac , Lecle c e al. (2020a)
obse ed ha he high colonisa ion success o NIS on se lemen pla es was in-
dependen o in e na ional essel a ic, indica ing ha hese species had al eady
es ablished in he espec i e po s. This is likely due o he abundance o a i icial
in as uc u e, pa icula ly loa ing s uc u es, which o e se lemen oppo uni-
ies and p o ec ion om ben hic p eda ion o newcome s (Dumon e al. 2011;
Lecle c e al. 2020b; Rech e al. 2024b). In con as , NIS colonisa ion o ben hic
se lemen pla es (Lecle c e al. 2020a) o e en adjacen na u al ben hic en i on-
men s (e.g., Fo es e al. 2013; Simkanin e al. 2013) ends o be less equen .
The disco e y o Wa e sipo a a cua a in he in e idal zone a he Coquimbo
si e is conce ning. Unlike many o he NIS, Wa e sipo a species ha e in aded ocky
in e idal and sub idal habi a s, as well as coas al ee s in No heas Paci ic Cali-
o nia (Zabin e al. 2018; Page e al. 2019), likely dispe sing om local po s and
ma inas. O e sho e dis ances (< 4.5 km), la al connec i i y wi h he nea es
po has been sugges ed as a dispe sal mechanism (Page e al. 2019). Howe e , o e
longe dis ances, seconda y dispe sal o NIS is o en acili a ed by locally ope a ing
ec ea ional boa s (e.g., Da idson e al. 2010; Ash on e al. 2022a). In he small
Bay o Coquimbo, wi h a maximum wid h o 2 km, la ae could easily sp ead
na u ally. Addi ionally, a isanal ishing boa s and o he leisu e c a a e equen ly
ancho ed nea he in aded si e, posing a signi ican isk o u he dispe sing he
species along he coas .
A ou s udy si e in he empe a e SE Paci ic, W. a cua a was p esen on he
sea loo , whe e p eda ion p essu e is s ong, bu no on se lemen pla es loa ing
in he wa e column (a a dis ance o only 100 m om he in aded si e), whe e p e-
da ion is i ually absen (see Dumon e al. 2011; Rech e al. 2024b). This inding
is mos likely explained by a acili a ing e ec o ben hic p eda ion on Wa e sipo a
es ablishmen . On he loa ing se lemen pla es, whe e he e is no p eda ion p es-
su e, a dense and di e se ouling communi y, la gely domina ed by Ciona obus a
and o he ascidians (unpublished da a, bu see pho o in Suppl. ma e ial 1: ig. S1),
exe s bio ic p essu e h ough compe i ion and may inhibi Wa e sipo a es ablish-
men . On he sea loo , on he o he hand, high p eda ion p essu e h ough (main-
ly) ben hic c abs and ishes (see Mus i e al. 2019) impedes he g ow h o so -bod-
ied species wi h no s uc u al de ence mechanisms like Ciona (e.g., Dumon e al.
2011; Giache i e al. 2020). Wa e sipo a a cua a, in con as , is p o ec ed om
p eda o s due o i s la , enc us ing o m and calca eous skele on, and can sp ead
mo e e ec i ely when compe ing species a e emo ed by p eda o s (O icchio and
Munis Diaz 2020; Chebaane e al. 2023). Bio ic acili a ion o Wa e sipo a species
h ough p eda ion has also been documen ed in o he s udies: Needles e al. (2015)
obse ed inc eased abundances o W. sub o qua a in he p esence o sea o e and sea
s a p eda o s ha emo ed he compe ing ounda ion species My ilus cali o nianus
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
in a Cali o nian bay. Simila ly, Chebaane e al. (2023) ound ha abundances o
W. suba a on se lemen pla es we e highe in open (= p eda o -accessible) han in
caged (= p eda o -excluding) ea men s in wo Po uguese ma inas.
Ano he signi ican inding o ou s udy was he de ec ion o Bugulina s oloni e a
a h ee di e en si es, spanning a o al dis ance o 1,300 km om Coquimbo o
Pue o Mon . This sugges s ha he species may al eady be well es ablished in Chile.
Howe e , we canno ye de e mine i he e is any connec i i y be ween he colonies
ac oss hese di e en s udy egions o how many indi idual in oduc ions ha e con-
ibu ed o hei sp ead. Popula ion genomics ha e been used o assess in e -popu-
la ional ela ionships o ma ine NIS in p e ious s udies (e.g., Page e al. 2019) and
could be an impo an ool o elucida e he in asion his o y o B. s oloni e a in Chile.
Recommenda ions o moni o ing and p e en ion
Two o he h ee newly epo ed species we e de ec ed h ough se lemen pla es
deployed a s a egically chosen high- isk si es o e a ela i ely sho pe iod o jus
ou mon hs. Se lemen pla e-based su eys a e widely used wo ldwide o de ec -
ing NIS (e.g., Tambu ini e al. 2021) and ha e been success ully implemen ed in
Chilean con inen al po s (Lecle c e al. 2018, 2020a,b), as well as on he emo e
island o Rapa Nui (Eas e Island; Rech e al. 2024a). The ela i ely high numbe
o p e iously un epo ed species ound in ou s udy, despi e including only se en
si es, sugges s ha many o he NIS may emain unde ec ed in o he loca ions, bo h
wi hin Chile and in o he unde s udied egions globally. This highligh s he po en-
ial o se lemen pla es as a p omising ool o u u e NIS de ec ion in such a eas.
As no ed in his s udy, isual iden i ica ion o b yozoans a he species le el is
challenging and o en necessi a es scanning elec on mic oscope (SEM) images.
Visual iden i ica ion in he ield is equen ly ambiguous, as simila species can eas-
ily be con used o o e looked. This is likely he case o B. s oloni e a and Bugulina
c . ul a, bo h o which may ha e been misiden i ied as he congene Bugulina c .
labella a in p e ious s udies. Gi en hese new indings, a comp ehensi e gene ic
and mo phological analysis o he genus Bugulina in Chile will be c ucial o cla i y
which species a e p esen and how hey a e dis ibu ed. Unambiguous iden i ica-
ion o NIS b yozoans in Chile is essen ial o unde s anding hei in oduc ion
pa hways and implemen ing e ec i e p e en ion and con ol measu es.
Hull ouling plays a c i ical ole in he in oduc ion o NIS, p omp ing se e al
coun ies and egions, such as Cali o nia and New Zealand (Scianni e al. 2021), o
implemen egula ions a ge ing essels a i ing in hei e i o ies. Howe e , such
legisla ion is s ill lacking in mos pa s o he wo ld. In Chile, wi h o e 5,000 in e -
na ional essel a i als annually (S owhas e al. 2023), he only s udy conduc ed on
hull ouling so a has ound se e al NIS a ached o a i ing in e na ional essels (Pi-
noche e al. 2023), and Chilean po s a e ecognized as ho spo s o ma ine in asions
(e.g., Lecle c e al. 2018, 2020a,b). Despi e he ela i ely low numbe o epo ed
ma ine NIS o da e, models p edic ing shipping equencies sugges a signi ican in-
c ease o 15 o 35 NIS a i ing pe yea (Sa dain e al. 2019). In e es ingly, a numbe
o p e iously un epo ed NIS ha e been iden i ied in Chile du ing he pas decade,
including se e al species in Coquimbo (Häusse mann e al. 2015; Bee mann e al.
2025). While Chile has begun p epa ing measu es o mi iga e hull ouling- ela ed
bioin asions (S owhas e al. 2023), no legisla ion is ye in place. I is impe a i e ha
such egula ions be implemen ed o sa egua d Chile’s coas lines om u u e in asions.
288
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Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
Acknowledgemen s
We since ely hank he ep esen a i es and pe sonnel om he ma inas, po s,
and aquacul u e acili ies ha suppo ed and hos ed ou su eys. In pa icula ,
we ex end ou g a i ude o he ollowing indi iduals and en i ies o hei ac i e
assis ance: Cap . Pa icio González, F ancisco Loyola López, Daniel Aguile a
A aya, Ge man Alejand o Li a Godoy (Uni e sidad Ca ólica del No e, Campus
Guayacán), Ma iano Galdames Beckdo , Claudio González, and Fede ico Núñez
(Co adía Náu ica del Pací ico, Alga obo), Se gio Guzmán and Camilo Va gas
(Club de Ya es Relonca í, Pue o Mon ), Felipe Be ios (Emp esa po ua ia San
An onio), Capi anía del Pue o and he c ew o LSG San An onio (Pue o San
An onio), Ko o Nui O Te Vaika a, Comunidad Indígena Ma’u Henua, Albe o
He e e i, and Alcaldía del Ma de Hanga Piko, Rapa Nui.
We a e g a e ul o Jo ge A ilés om he Sala de Colecciones Biológicas (SC-
BUCN) a Uni e sidad Ca ólica del No e o p o iding logis ical and p ac ical sup-
po du ing sample analysis, and o D . Se gio A. Na a e e o his logis ical suppo
and o g an ing access o he labo a o y and esou ces a he Es ación Cos e a de
In es igaciones Ma inas o Pon i icia Uni e sidad Ca ólica in Las C uces, Chile.
We hank D . And ea Waeschenbach, D . Ka in Fehlaue -Ale, and D . Linda
McCann o hei aluable in o ma ion and guidance on he gene ic analysis o
b yozoan samples, and D . Tim Kiessling o gene ously sha ing his expe ise on
a ious s a is ical me hods. Mo eo e , we acknowledge he suppo o Capi anía
de Pue o de Coquimbo (A mada de Chile) by p o iding da a on essel a i als
a he CMP e minal in Guayacán, Coquimbo. Las ly, we hank wo anonymous
e iewe s o hei e alua ion and cons uc i e commen s.
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
All p ocedu es in ol ing animals we e in compliance wi h he Eu opean Communi y Coun-
cil Di ec i e o 24 No embe 1986, and e hical app o al was g an ed by he E hics Commi ee
o Uni e sidad Ca ólica del No e (CEC UCN N° 07/ 2020, Coquimbo, Chile). On Rapa Nui,
pe mi o he s udy was g an ed by Ko o Nui o e Vaika a o Rapa Nui (Consejo del Ma de
Rapa Nui; Pe mi N°3_2022).
Funding
The p ojec ha ga e ise o hese esul s ecei ed he suppo o a ellowship om he ”la Caixa”
Founda ion (ID 100010434). The ellowship code is LCF/BQ/PI24/12040006. SR ecei ed und-
ing om ANID (Agencia Nacional de In es igación y Desa ollo, Chile) in he p og am FON-
DECYT POSTDOCTORADO 2020, p ojec No. 3201074 un il Ap il 2022. LMV was sup-
po ed by he Na ional Council o Scien i ic and Technological De elopmen , B azil (Pq-CNPq
311523/2021–8) and he Coo denação de Ape eiçoamen o de Pessoal de Ní el Supe io – B asil
(CAPES) – Finance Code 001. Sala de Colecciones Biológicas o Uni e sidad Ca ólica del No e
(SCBUCN) has ecei ed suppo o Anillo – ANID ATE 220044 BiodUCCT and FONDECYT
1241386 (ANID, Chile). The unde s had no ole in s udy design, da a collec ion and analysis, deci-
sion o publish, o p epa a ion o he manusc ip .
289
NeoBio a 102: 269–294 (2025), DOI: 10.3897/neobio a.102.144725
Sabine Rech e al.: De ec ion o alien b yozoans along he Chilean SE Paci ic coas
Au ho con ibu ions
SR: Concep ualisa ion, Me hodology, Fo mal analysis, In es iga ion, Da a cu a ion, W i ing – o igi-
nal d a , Visualisa ion, P ojec adminis a ion, Funding acquisi ion. LMV: In es iga ion, Valida ion,
W i ing – e iew and edi ing. AIV: In es iga ion, Me hodology, W i ing – e iew and edi ing. MSRB:
In es iga ion, Me hodology, W i ing – e iew and edi ing. JPF: In es iga ion, Resou ces. DPG: Vali-
da ion, W i ing – e iew and edi ing. MCS: In es iga ion. KP: Resou ces. JS: Resou ces. MT: Concep-
ualisa ion, Me hodology, Resou ces, Supe ision, Funding acquisi ion, W i ing – e iew and edi ing.
Au ho ORCIDs
Sabine Rech h ps://o cid.o g/0000-0002-7500-6330
Leand o M. Viei a h ps://o cid.o g/0000-0001-8661-8861
And ea I. Va ela h ps://o cid.o g/0000-0002-9752-0816
Ma ía Soledad Rome o Bas ías h ps://o cid.o g/0000-0002-7291-7462
Juan Pablo Fuen es h ps://o cid.o g/0009-0002-3577-8579
Dennis P. Go don h ps://o cid.o g/0000-0001-9398-996X
Ma ina Cacciu olo Said h ps://o cid.o g/0009-0001-6434-9047
Ku Paschke h ps://o cid.o g/0000-0002-4785-8034
Ja ie Sellanes h ps://o cid.o g/0000-0002-6942-0762
Ma in Thiel h ps://o cid.o g/0000-0001-7535-3888
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex o Supplemen a y
In o ma ion.
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