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First insights into the scale of invasions in African marine protected areas: leveraging global databases and citizen science data

Author: Ackland, Sarah J.; Richardson, David M.; Robinson, Tamara B.
Publisher: Zenodo
DOI: 10.3897/neobiota.102.149275
Source: https://zenodo.org/records/17301601/files/NB_article_149275.pdf
399
Fi s insigh s in o he scale o in asions in A ican ma ine p o ec ed
a eas: le e aging global da abases and ci izen science da a
Sa ah J. Ackland1, Da id M. Richa dson1, Tama a B. Robinson1
1 Cen e o In asion Biology, Depa men o Bo any and Zoology, S ellenbosch Uni e si y, Ma ieland, 7609, Sou h A ica
Co esponding au ho : Tama a B. Robinson ([email p o ec ed])
Copy igh : © Sa ah J. Ackland 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
Up- o-da e in o ma ion on he dis ibu ion o alien species is essen ial o e idence-based managemen .
Howe e , ou ine moni o ing is esou ce in ensi e. This has led o egional biases in he a ailabili y o
in o ma ion on in asions, wi h li le unde s anding o in asions in many de eloping egions, including
A ica. This knowledge gap, pa icula ly p oblema ic o ma ine p o ec ed a eas (MPAs), challenges
he abili y o A ican s a es o mee hei obliga ions unde he Global Biodi e si y F amewo k (GBF)
Ta ge 6. This s udy d ew on eely a ailable global da abases (P o ec ed Plane ; Wo ld Regis e o
In oduced Ma ine Species; Wo ld Regis e o Ma ine Species) and ci izen science da a (iNa u alis ) o
p o ide no el insigh s in o ma ine alien species in A ican MPAs. A o al o 27 species we e eco ded
wi hin 17 MPAs ac oss se en coun ies. The po en ial h ea posed o hese MPAs was assessed using
he EICAT and SEICAT impac schemes. Wo yingly, species known o ha e impac s o massi e o
majo magni ude we e documen ed in nine MPAs (Table Moun ain Na ional Pa k, Robben Island, Six-
een Mile Beach, Langebaan Lagoon, Namaqua Na ional Pa k – all in Sou h A ica); Namibian Islands
(Namibia); Ilhas Fo mosa, Nago and Tchedia (U ok) (Guinea-Bissau); Banc d’A guin Na ional Pa k
(Mau i ania); Massa (Mo occo)). When used in conjunc ion wi h cu a ed da abases, iNa u alis o e ed
cos -e ec i e and e i iable eco ds o some alien species in unde -su eyed MPAs. By le e aging da a
om dispa a e da abases, his s udy imp o es ou knowledge o he scale o in asions in A ican MPAs
and p o ides a ounda ion upon which S a es can p io i ise moni o ing wi hin hei MPA ne wo ks.
Ul ima ely, hese da a could suppo he de elopmen o a ge ed ou ine moni o ing p og ammes
which could assis he managemen o ma ine in asions and he a ainmen o GBF Ta ge 6.
Key wo ds: A ica, biological in asions, communi y science, da a acquisi ion, impac s, iNa u alis ,
managemen , p o ec ed a eas
In oduc ion
Biological in asions a e a majo d i e o global en i onmen al change (Riccia di
e al. 2021) and p o ec ed a eas a e no immune o he impac s o in asi e alien
species (Shackle on e al. 2019, 2020). The Kunming-Mon eal Global Biodi e -
si y F amewo k (GBF), de eloped unde he Con en ion on Biological Di e si y
(CBD), p oposes 23 a ge s o suppo and guide he achie emen o he Sus ain-
able De elopmen Goals and he global ision o educe an h opogenic impac s on
biodi e si y by 2030 (Robinson e al. 2025). Ta ge 6 aims o educe he a e o in-
oduc ions and es ablishmen o known o po en ially in asi e alien species by a
leas 50% by 2030 (CBD/COP/DEC/15/4). The aim o his a ge is o elimina e,
Academic edi o : Paula Chainho
Recei ed:
10 Feb ua y 2025
Accep ed:
3 June 2025
Published:
7 Oc obe 2025
Ci a ion: Ackland SJ, Richa dson
DM, Robinson TB (2025) Fi s
insigh s in o he scale o in asions
in A ican ma ine p o ec ed a eas:
le e aging global da abases and
ci izen science da a. In: Anas ácio P,
Ribei o F, Chainho P (Eds) In asions
in Aqua ic Sys ems. NeoBio a 102:
399–418. h ps://doi.o g/10.3897/
neobio a.102.149275
NeoBio a 102: 399–418 (2025)
DOI: 10.3897/neobio a.102.149275
Ad ancing esea ch on alien species and biological in asions
A pee - e iewed open-access jou nal
NeoBio a
400
NeoBio a 102: 399–418 (2025), DOI: 10.3897/neobio a.102.149275
Sa ah J. Ackland e al.: In asions in A ican MPAs
minimise, educe and mi iga e he o e all impac s o alien species on biodi e si y
and ecosys em se ices (CBD/COP/DEC/15/5). The a ge also equi es he e adi-
ca ion o con ol o alien species in p io i y si es (e.g. p o ec ed a eas) as hese a eas
allow o conse a ion objec i es o be me while p o iding economic, poli ical
and social bene i s (Busch 2008).
Al hough he a ailabili y o global da a on alien species and hei dis ibu ion
has imp o ed g ea ly o e he las ew decades (Pyšek e al. 2020), subs an ial
geog aphical biases s ill emain, o en e lec ing he de elopmen al and socioeco-
nomical s a us o egions (Pyšek e al. 2008, 2020; Nuñez and Paucha d 2010).
This can be seen, o example, in he ocus o esea ch e o in he Ame icas and
Eu ope (Hulme e al. 2014). Inconsis encies in he a ailabili y o in o ma ion
ac oss con inen s has also been highligh ed. Fo example, as o 2006, wo k in
Sou h A ica accoun ed o app oxima ely wo- hi ds o esea ch e o on bio-
logical in asions on he A ican con inen (Pyšek e al. 2008). Dedica ed e o s
o p o ide up- o-da e alien species lis s (e.g. Zengeya and Wilson (2023) o
Sou h A ica)) has imp o ed he unde s anding o alien species in some coun-
ies, bu many s ill lag behind. Fo example, he e is a need o comp ehensi e
in asi e plan species lis s o sub-Saha an A ica (Wi e al. 2018) and no da a
a e a ailable o many egions (Richa dson e al. 2022). The same si ua ion exis s
o o he axa. While he esponsibili y o he p o ision o such in o ma ion
usually alls on o esea che s, he ou pu s o such s udies a e o p ac ical use
o manage s (Oja ee e al. 2015). The lack o ounda ional knowledge wi h-
in unde s udied egions has he po en ial o pe pe ua e knowledge deb . This
challenge is especially p e alen in de eloping egions whe e he managemen o
alien axa is al eady limi ed (Kuma Rai and Singh 2020).
Global declines in ma ine biodi e si y and h ea s posed by an h opogenic ac-
ions ha e led o he inc easing implemen a ion o ma ine p o ec ed a eas (MPAs)
(Ban e al. 2015; Chukwuka e al. 2025). De ined as geog aphical spaces dedica -
ed o legally manage long- e m conse a ion o ecosys em unc ions, se ices and
cul u al alues (Day e al. 2012), MPAs a e impo an ma ine conse a ion ools.
Gi en he mul i ude o h ea s (including biological in asions) acing ma ine sys-
ems (E a is e e al. 2018; Ho wi z e al. 2020; Pyšek e al. 2020; Renchen e al.
2021) and he ac ha conse a ion agencies ha e limi ed esou ces (Delaney e
al. 2008; Be ic and MacIsaac 2015), manage s mus p io i ise how and whe e o
spend sca ce esou ces. Howe e , despi e much e o , cu en managemen and
conse a ion e o s a e inadequa e in mos p o ec ed a eas (C ain e al. 2009; Bo -
ja e al. 2016; Chukwuka e al. 2025). Ma ine p o ec ed a eas se e a numbe o
economic, ecological and socie al aims, all o which a e unde mined by biological
in asions. The need o add ess alien species and quan i y he scale o in asions in
MPAs is hus c ucial o ensu e ha MPAs mee hei in ended pu pose.
Moni o ing p o ides c i ical in o ma ion on he changing scale o in asions and
is c ucial o e idence-based managemen (Leh iniemi e al. 2020; Lou ei o e
al. 2021). When collec ed sys ema ically o e space and ime, such in o ma ion
suppo s comp ehensi e da abases ha p o ide empo al and spa ial insigh s in o
in asions (Delaney e al. 2008). Howe e , moni o ing is challenging in ma ine
en i onmen s because o o hei complex na u e (D’Amen and Azzu o 2020).
Thei unde wa e landscapes and in e connec edness make in asions in he ma ine
ealm di icul o documen and manage (A nd e al. 2018). Addi ionally, la ge-
scale, long- e m moni o ing a ely occu s due o esou ce limi a ions (Delaney e
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NeoBio a 102: 399–418 (2025), DOI: 10.3897/neobio a.102.149275
Sa ah J. Ackland e al.: In asions in A ican MPAs
al. 2008; Be ic and MacIsaac 2015). Cos -e ec i e me hods o moni o ing a e
needed o b idge he gap be ween da a equi emen s and scien i ic unde s and-
ing (Leh iniemi e al. 2020; Howa d e al. 2022). One way o achie ing his is
o u ilise c owd-sou ced da a p o ided by ci izen science ini ia i es (Ho wi z e
al. 2020; Howa d e al. 2022). The ad ancemen o echnology and he ecen
p oli e a ion o online pla o ms has enhanced he po en ial o moni o ing by
using de ices and sma phone applica ions (Daniels e al. 2022; Howa d e al.
2022). Applica ions, such as iNa u alis , enable obse e s o upload pho og aphic
e idence o indi idual sigh ings and sugges axonomic iden i ica ions (Pimm e
al. 2015; Howa d e al. 2022). Reco ds collec ed by obse e s ha e he po en ial
o p o ide a la ge olume o da a o e a wide a ea. I u ilised wi h due a en ion
o inhe en limi a ions, ci izen science o e s a complemen a y sou ce o da a o
suppo e ec i e moni o ing o alien species (Howa d e al. 2022; Po gie e e al.
2024). iNa u alis has acili a ed many use ul insigh s wi hin he ma ine ealm, in-
cluding imp o ed unde s anding o ee ish species ichness (Robe s e al. 2022),
biodi e si y pa e ns (Rocha e al. 2024) and in asions (Compagnone e al. 2024).
Fo A ican coun ies o mee hei obliga ions unde GBF Ta ge 6, be e
ounda ional knowledge on he p esence o alien species wi hin MPAs is essen ial.
Unde de eloped egions a e al eady expe iencing a high economic bu den which
is exace ba ed by he inancial demands associa ed wi h managing in asi e spe-
cies (B adshaw e al. 2024). The moni o ing equi ed o p o ide in o ma ion o
ou ine managemen o alien species has he po en ial o u he in ensi y inan-
cial s ain, challenging he acquisi ion o ounda ional knowledge and he imple-
men a ion o e ec i e managemen . Acco dingly, his s udy aimed o: (1) p o ide
insigh s in o he scale o alien species in A ican MPAs h ough he use o global
da abases and ci izen science da a om iNa u alis ; and (2) assess he po en ial
en i onmen al and socio-economic impac s posed by hese axa.
Me hods
Ma ine p o ec ed a eas (MPAs) along he A ican coas we e iden i ied using he
open-access P o ec ed Plane da abase (h ps://www.p o ec edplane .ne /en/ he-
ma ic-a eas/ma ine-p o ec ed-a eas; accessed: 9 Janua y 2024). This da abase, he
mos comp ehensi e and up- o-da e sou ce o da a on p o ec ed a eas, colla es
submissions om go e nmen agencies, non-go e nmen al o ganisa ions and local
communi ies. Ma ine P o ec ed A eas we e de ined using he il e s lis ed in Suppl.
ma e ial 1. Da a, ex ac ed o e a single day (9 Janua y 2024), co e ed all A ican
MPAs decla ed p io o Janua y 2024.
Reco ds o all species epo ed as alien [i.e. species whose p esence in a egion is
a ibu able o human ac ions enabling c ossing o biogeog aphical ba ie s sensu
Richa dson e al. (2011)] along he A ican coas we e ex ac ed om he Wo ld
Regis e o In oduced Ma ine Species (WRiMS; Cos ello e al. (2024); h ps://
www.ma inespecies.o g/in oduced/aphia.php?p=checklis ; accessed: 10 Sep em-
be 2024). Da a included bo h occu ence eco ds o alien species as well as e-
co ds o es ablished popula ions. This da abase was selec ed as i colla es da a om
pee - e iewed a icles o p o ide he mos comp ehensi e and s anda dised glob-
al-scale da abase o ma ine in oduced o ganisms (Cos ello e al. 2021). All species
eco ds ha we e ex ac ed we e inspec ed and a h ee-s ep cleaning p ocedu e
was applied. As da a ex ac ed om WRiMS include bo h accep ed names and
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Sa ah J. Ackland e al.: In asions in A ican MPAs
synonyms wi h no op ion o il e ou hese edundancies, all axonomic duplica es
we e checked and emo ed o a oid o e es ima ions o he numbe o alien species.
Scien i ic names we e hen upda ed acco ding o he mos ecen nomencla u e
colla ed in he Wo ld Regis e o Ma ine Species (WoRMS; Ahyong e al. (2024);
h ps://www.ma inespecies.o g; accessed: 10 Sep embe 2024). Since WRiMS
con ains bo h ma ine and es ua ine da a, all oligohaline species we e inspec ed.
Using cu en li e a u e o in es iga e salini y ole ances, only species wi h a ole -
ance o ma ine en i onmen s we e e ained.
To ensu e ha no published eco ds o alien species we e inad e en ly ex-
cluded, a sea ch was conduc ed on he ISI Web o Science on 10 Sep embe
2024. A sys ema ic li e a u e e iew was conduc ed using he ollowing sea ch
s ing “ma ine” AND “alien” OR “in asi e” OR “in oduced” OR “nonna i e”
AND “coun y name”. As he e minology used o desc ibe alien species is in-
consis en and highly a iable in he li e a u e (Lockwood e al. 2013), b oad
sea ch e ms (i.e. “alien”, “in asi e”, “nonna i e” and “in oduced”) we e used
o cap u e as much o he ele an li e a u e as possible. As no addi ional e-
co ds o ma ine alien species we e de ec ed in he li e a u e sea ch, he da ase
ex ac ed om WRiMS was accep ed as comple e.
U ilising he explo e page on iNa u alis (h ps://www.ina u alis .o g; ac-
cessed: 18–19 Sep embe 2024), sea ches we e made o each species lis ed as
alien in any A ican coun y on WRiMS. Reco ds o each species ac oss all
quali y g ades we e ex ac ed om iNa u alis o he pe iod 1 Ap il 2008 ( he
da e o es ablishmen o he iNa u alis pla o m) o 18 Sep embe 2024. In o -
ma ion on he species iden i ica ion (scien i ic name and associa ed media), he
da e o obse a ion, he da e in which he obse a ion was uploaded o he pla -
o m, he coo dina es o he obse a ions and posi ional accu acy we e ex ac ed
o each obse a ion. As some species may ha e a na i e ange in one A ican
coun y, bu be alien in ano he , he na i e ange o each species was con i med
using he Wo ld Regis e o Ma ine Species (WoRMS; Ahyong e al. (2024);
h ps://www.ma inespecies.o g; accessed 20–21 Sep embe 2024) and published
a icles. To in e oga e he geog aphic dis ibu ion o iNa u alis obse a ions
and ensu e ha he obse a ions we e, in ac , obse ed wi hin an MPA, he
coo dina es o each obse a ion we e plo ed using he Quan um Geog aphic
In o ma ion Sys em (QGIS; e sion 3.22 LTR). Each obse a ion was plo ed
on o a map o A ica wi h he MPAs dema ca ed [MPA shape iles om P o-
ec ed Plane (h ps://www.p o ec edplane .ne /en/ hema ic-a eas/ma ine-p o-
ec ed-a eas; accessed: 9 Janua y 2024]). All poin s we e isually inspec ed and
any eco ds om ou side MPAs we e excluded om u he analysis. iNa u alis
obse a ions occu ing wi hin he na i e ange o a species we e also excluded.
To add ess unce ain y in iNa u alis da a and imp o e i s accu acy and eli-
abili y, con idence o each obse a ion was sco ed ollowing he p o ocol p o-
posed by Ackland e al. (2024). This p ocess included h ee s eps: assessing
he species, he image and geoloca ion o each eco d. Only eco ds ecei ing
medium o high con idence sco es we e e ained o u he analysis. A Kend-
all’s ank co ela ion was used o assess he ela ionship be ween he numbe o
eco ds o alien species as de ec ed in A ican MPAs h ough ime. E o was
quan i ied as he numbe o ma ine alien species eco ds pe 100 eco ds wi hin
A ican MPAs. All s a is ical analyses we e conduc ed in he R s a is ical en i-
onmen ( e sion 4.3.2 – “Eye Holes”, R Co e Team (2023)).
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NeoBio a 102: 399–418 (2025), DOI: 10.3897/neobio a.102.149275
Sa ah J. Ackland e al.: In asions in A ican MPAs
To de e mine he h ea ha hese species may pose o MPAs, a li e a u e e iew
was ca ied ou using ISI Web o Science o iden i y all p e iously published e-
po s o impac o each species a any global loca ion. The sea ch s ing “scien i ic
name” AND “impac *” OR “e ec *”, was applied along wi h each species name
and all p e ious synonyms. Sea ches we e comple ed be ween 25 and 27 Sep em-
be 2024. En i onmen al impac was ca ego ised acco ding o he mechanisms
h ough which hey mani es [i.e. (1) compe i ion; (2) p eda ion; (3) hyb idisa-
ion; (4) disease ansmission; (5) pa asi ism; (6) poisoning/ oxici y; (7) bio- oul-
ing; (8) g azing/he bi o y/b owsing; (9) chemical; (10) physical o (11) s uc u al
impac on an ecosys em; and (12) in e ac ion wi h o he alien species] using he
En i onmen al Impac Classi ica ion o Alien Taxa scheme (EICAT; Blackbu n e
al. (2014)). Socio-economic impac s we e classi ied acco ding o he cons i uen s
o human well-being hey a ec [i.e. (1) sa e y; (2) ma e ial and imma e ial asse s;
(3) heal h; and (4) social, spi i ual and cul u al ela ions] using he Socio-econom-
ic Impac Classi ica ion o Alien Taxa scheme (SEICAT; Bache e al. (2017)).
Las ly, he magni ude o impac (i.e. massi e, majo , mode a e, mino , minimal o
da a de icien ) epo ed p e iously in li e a u e was assigned o each species. Clas-
si ica ion o magni ude was based on he maximum impac no ed o he species,
ega dless o loca ion, ollowing Kumschick e al. (2017).
Resul s
Se en y-nine A ican ma ine p o ec ed a eas (MPAs) we e iden i ied ac oss 14
coun ies (Fig. 1, Suppl. ma e ial 2: able S2). A o al o 497 ma ine species we e
epo ed as alien in coas al A ican coun ies (Suppl. ma e ial 2: able S3) on he
Wo ld Regis e o In oduced Species (WRiMS), wi h a o al o 440 iNa u alis e-
co ds o 34 o hose species epo ed om wi hin A ican MPAs. A e excluding
eco ds ha we e no sco able and hose wi h low con idence sco es, 307 eco ds
(70%) o 27 species we e e ained o u he analysis (Table 1, Suppl. ma e ial 2:
able S4). O e wo- hi ds o hese obse a ions [211 eco ds (72%)] ecei ed a
maximum sco e o image cla i y. Examples o such eco ds a e p o ided in Fig. 2.
O he 27 species eco ded on iNa u alis , ou we e obse ed wi hin coun ies
wi h no p e ious eco ds acco ding o WRiMS. These species included he ba na-
cle Balanus igonus (Mau i ania), he b yozoan Ama hia e icilla a (Mau i ania
and Sou h A ica), he colonial unica e Symplegma b akenhielmi (Senegal) and he
c ab Callinec es sapidus (Guinea-Bissau and Mau i ania).
A gene al end o inc eased engagemen on iNa u alis was obse ed, wi h e-
co ds o ma ine alien species in A ican MPAs inc easing signi ican ly o e ime
(Kendall’s ank co ela ion: τ = 0.51, p < 0.05; Fig. 3). Reco ds we e sp ead ac oss
17 MPAs om se en coun ies (Fig. 4). Two MPAs accoun ed o 85.7% o all e-
co ds (i.e. Table Moun ain Na ional Pa k MPA (Sou h A ica) and Banc d’A guin
Na ional Pa k (Mau i ania)], wi h many o he MPAs ha ing ewe han i e e-
co ds. Table Moun ain Na ional Pa k MPA had he mos eco ds o alien axa [i.e.
241 eco ds (78.5%)] and he highes numbe o species de ec ed on iNa u alis
(15 species). The MPA wi h he second highes numbe o eco ds was Banc d’A -
guin Na ional Pa k in Mau i ania (22 eco ds; 7.2%), he second highes numbe
o species (8 species) was also de ec ed in his MPA.
O he 27 alien species de ec ed in A ican MPAs, en i onmen al impac had
been epo ed o 23 species and socio-economic impac o 14 species (Suppl.

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Sa ah J. Ackland e al.: In asions in A ican MPAs
ma e ial 2: able S5) and ou species we e lis ed as da a de icien . Namibian
Islands (Namibia) and Langebaan Lagoon (Sou h A ica) we e he only MPAs
ha suppo ed species wi h massi e (i.e. he oys e Magallana gigas and he Med-
i e anean mussel My ilus gallop o incialis) and majo impac (i.e. he mussel
Semimy ilus pa agonicus, he ascidian Ciona obus a and he b yozoan Ama hia
e icilla a). Table Moun ain Na ional Pa k MPA in Sou h A ica had he highes
numbe o species wi h p e iously documen ed impac s. This MPA had a single
species known o massi e impac (i.e. he Medi e anean mussel My ilus gallo-
p o incialis) and h ee species o which majo impac had been epo ed (i.e. he
Eu opean g een c ab Ca cinus maenas, he mussel Semimy ilus pa agonicus and
he ascidian Ciona obus a). Banc d’A guin Na ional Pa k in Mau i ania had he
second highes numbe o species wi h p e iously documen ed impac s. While
some species displayed only en i onmen al mechanisms o impac , a gene al pa -
e n was ha species ha exe ed en i onmen al impac h ough mul iple mech-
anisms also had many mechanisms o socio-economic impac and ul ima ely ex-
e ed high impac s (Pea son’s p oduc -momen co ela ion: = 0.79, p < 0.001;
Fig. 5, Suppl. ma e ial 2: able S5). Th ee species had a high numbe o en i-
onmen al and socio-economic mechanisms o impac (i.e. he oys e Magallana
gigas and he mussels My ilus gallop o incialis and Semimy ilus pa agonicus). The
Paci ic oys e , Magallana gigas, had he highes numbe o mechanisms h ough
which i has caused en i onmen al and socio-economic impac s. This species was
only obse ed wi hin Namibian Islands MPA in Namibia.
Figu e 1. Loca ion o he 14 A ican coun ies and he numbe o ma ine p o ec ed a eas decla ed wi hin hose coun ies on he P o ec ed
Plane da abase.
Mo occo
Mau i ania
Senegal
Guinea-Bissau
Sie a-Leone
Angola
Namibia
Sou h A ica
Tanzania
Kenya
Seychelles
Egyp
Sudan
E i ea
1
1
1
7
1
11
41
3
1
5
2
3
1
1
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Sa ah J. Ackland e al.: In asions in A ican MPAs
Discussion
In asi e species a e majo d i e s o global change (Pyšek e al. 2020) and in a-
sions di ec ly unde mine he conse a ion objec i es o p o ec ed a eas (Ren e al.
2021). I is, he e o e, c i ical o iden i y he p o ec ed a eas ha a e mos a isk
o in asions and he species ha may pose he g ea es h ea o hem (Pyšek e al.
2020). Howe e , he cu en lack o baseline da a on alien species in MPAs hinde s
his ask, since in o ma ion on he scale o in asions in MPAs is lacking in mos
Table 1. The species de ec ed on iNa u alis wi hin A ican ma ine p o ec ed a eas and he magni ude o hei en i onmen al impac (as
pe Blackbu n e al. 2014) and socio-economic impac (as pe Bache e al. 2017). Ma ine p o ec ed a ea codes a e as ollows: iSimangaliso
(iSi); Aliwal (Ali); P o ea Banks (P o); Ama hole (Ama); Addo Elephan (Add); Be y’s Bay (Be ); Table Moun ain Na ional Pa k Ma ine
P o ec ed A ea (Tmn); Robben Island (Rob); Six een Mile Beach (Smb); Langebaan Lagoon (Lan); Namaqua Na ional Pa k (Nam); Na-
mibian Islands (Nim); Namibe Ma ine (Nmc); Ihas Fo mosa, Nago & Tchedia (Iha); Go ée (Go ); Banc d’A guin Na ional Pa k (Ban)
and Massa (Mas). Numbe s (#) assigned o species co espond o he numbe s in Figs 4, 5.
# Scien i ic name Coun y Ma ine p o ec ed a ea En i onmen al impac Socio-economic impac Impac li e a u e
1Ec opleu a c ocea Sou h A ica Tmn Mode a e Mino Fi idge and Keough 2013; Yan e
al. 2021
2Obelia dicho oma Sou h A ica Tmn Mode a e Da a de icien Wya e al. 2005
3T iden a a ma gina a Mau i ania Ban Da a de icien Da a de icien
4Anisolabis ma i ima Sou h A ica Tmn Da a de icien Da a de icien
5Amphibalanus enus us Sou h A ica Ali; Ama Da a de icien Da a de icien
6Balanus glandula Sou h A ica Ali; Tmn Mode a e Da a de icien Kado 2003; Valla ino and Elias 2008
7Balanus igonus Angola Nmc Minimal Minimal Lozano-Co ès and Zapa a 2014;
Abelouah e al. 2024
Mau i ania Ban
8Megabalanus in innabulum Sou h A ica iSi, Tmn Mode a e Da a de icien P a e al. 2022
9Callinec es sapidus Guinea-Bissau Iha Majo Mino Cla e o e al. 2022; Mancinelli e
al. 2017
Mau i ania Ban
Mo occo Mas
10 Ca cinus maenas Sou h A ica Tmn Majo Mino G osholz e al. 2011; B een and Me axas
2008 Whi low 2010; de Ri e a e al.
2011; Mabin e al. 2022
11 Ama hia e icilla a Mau i ania Ban Majo Mino McCann e al. 2015
Sou h A ica Lan
12 Bugula ne i ina Sou h A ica Be Mino Mino Yu e al. 2007; Bae e al. 2022
13 Vi ididen ula den a a Sou h A ica iSi; Tmn Minimal Da a de icien Micael e al. 2016
14 Wa e sipo a sub o qua a Sou h A ica Add; Tmn Mino Mino Sellheim e al. 2010; McKenzie e
al. 2011
15 Magallana gigas Namibia Nam Massi e Majo T oos 2010; Kelly e al. 2008
16 My ilus gallop o incialis Mau i ania Ban Massi e Majo B anch and S e ani 2004; S e ani and
B anch 2005; Alexande e al. 2015
Namibia Nim
Sou h A ica Lan; Nam; Rob; Smb; Tmn
17 Semimy ilus pa agonicus Namibia Nam Majo Majo Skein e al. 2018
Sou h A ica Lan; Tmn
18 Siphona ia pec ina a Mau i ania Ban Mino Da a de icien Boukhicha e al. 2015
19 Thecace a pennige a Sou h A ica Tmn Da a de icien Da a de icien
20 Ophiac is sa ignyi Guinea-Bissau Iha Mino Da a de icien Çina e al. 2019
Sou h A ica P o
21 Bo yllus schlosse i Sou h A ica Tmn Mode a e Da a de icien Wong and Ve caeme 2012
22 Ciona obus a Sou h A ica Lan; Tmn Majo Mino Robinson e al. 2017
23 Cla elina lepadi o mis Sou h A ica Tmn Minimal Da a de icien Casso e al. 2018
24 Diplosoma lis e ianum Sou h A ica Tmn Mino Minimal Ald ed and Cla e 2014
25 Symplegma b akenhielmi Senegal Go Mino Minimal Spyksma e al. 2024
26 Acan hopho a spici e a Mau i ania Ban Mode a e Da a de icien Da idson e al. 2015
27 Hypnea musci o mis Guinea-Bissau Iha Mode a e Minimal Okuha a e al. 2023
Mau i ania Ban
Senegal Go
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(a) (b)
(c) (d)
© Lau en an Noo ©Te y Gosline
© Ca el an de Col © ca olina_dlhs
Figu e 2. iNa u alis eco ds ha con ibu ed owa ds he indings o his s udy. a. Cla elina lepadi o mis (iNa u alis ID 152956163); b. Ciona
obus a (iNa u alis ID 69033891); c. Thecace a pennige a (iNa u alis ID 23247918); d. Callinec es sapidus (iNa u alis ID 174999050).
Figu e 3. The numbe o alien species wi h eco ds on iNa u alis in A ican ma ine p o ec ed a eas (MPAs) o e ime. Numbe s abo e
ba s indica e he numbe o alien species eco ds pe 100 eco ds wi hin A ican MPAs.
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Figu e 4. The numbe o ma ine alien species wi h eco ds in iNa u alis ac oss ma ine p o ec ed a eas (MPAs) and he coun ies in which
hey occu ed. Species a e ca ego ised acco ding o he highes magni ude o impac no ed o he species. Species codes (numbe s in he
boxes) a e explained in Table 1. MPA names (below x axis): iSimangaliso (iSi); Aliwal (Ali); P o ea Banks (P o); Ama hole (Ama); Addo
Elephan (Add); Be y’s Bay (Be ); Table Moun ain Na ional Pa k Ma ine P o ec ed A ea (Tmn); Robben Island (Rob); Six een Mile Beach
(Smb); Langebaan Lagoon (Lan); Namaqua Na ional Pa k (Nam); Namibian Islands (Nim); Namibe Ma ine (Nmc); Ihas Fo mosa, Nago
& Tchedia (Iha); Go ée (Go ); Banc d’A guin Na ional Pa k (Ban) and Massa (Mas).
Sou h A ica
Namibia
Angola
Senegal
Mau i ania
Mo occo
13
8
5
6
20 5 14 12
22
4
6
21
10
23
24
1
8
16
17
2
14
19
13
2516 16
16
11
22
17
16
16
17
15
720
9
27 27
9
11
27
7
18
26
3
16
9
Figu e 5. The ela ionship be ween he en i onmen al and socio-economic mechanisms epo ed o each species eco ded wi hin an
A ican ma ine p o ec ed a ea. The magni ude o impac o each species is indica ed by he colou s; he size o he poin s ep esen s he
numbe o species. The numbe s alongside poin s indica e species as lis ed in Table 1.
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al e o aging beha iou , locomo ion, and me abolic a e in a keys one ma ine mollusc. Scien i ic
Repo s 10: 5461. h ps://doi.o g/10.1038/s41598-020-62304-4
Howa d L, an Rees CB, Dahlquis Z, Luika G, Hand BK (2022) A e iew o in asi e species e-
po ing apps o ci izen science and oppo uni ies o inno a ion. NeoBio a 71: 165–188. h ps://
doi.o g/10.3897/neobio a.71.79597
Hulme PE, Pyšek P, Pe gl J, Ja ošík V, Scha ne U, Vilà M (2014) G ea e ocus needed on alien plan
impac s in p o ec ed a eas. Conse a ion Le e s 7: 459–466. h ps://doi.o g/10.1111/conl.12061
Hulme PE, Lieu ance D, Richa dson DM, Robinson TB (2025) Mul iple a ge s o he Global
Biodi e si y F amewo k mus be add essed o manage in asi e alien species in p o ec ed a eas.
NeoBio a 99: 149–170. h ps://doi.o g/10.3897/neobio a.99.152680
Iaca ella JC, Mankiewicz PS, Riccia di A (2015) Nega i e compe i i e e ec s o in asi e plan s
change wi h ime since in asion. Ecosphe e 6: 1–14. h ps://doi.o g/10.1890/ES15-00147.1
Kado R (2003) In asion o Japanese sho es by he NE Paci ic ba nacle Balanus glandula and i s eco-
logical and biogeog aphical impac . Ma ine Ecology P og ess Se ies 249: 199–206. h ps://doi.
o g/10.3354/meps249199
Kang B, Vi ule JRS, Li S, Shuai F, Huang L, Huang X, Fang J, Shi X, Zhu Y, Xu D, Yan Y, Lou F
(2022) In oduc ion o non‐na i e ish o aquacul u e in China: A sys ema ic e iew. Re iews in
Aquacul u e 15: 676–703. h ps://doi.o g/10.1111/ aq.12751
Kelly JR, P oc o H, Volpe JP (2008) In e idal communi y s uc u e di e s signi ican ly be ween
subs a es domina ed by na i e eelg ass (Zos e a ma ina L.) and adjacen o he in oduced oys e
C assos ea gigas (Thunbe g) in B i ish Columbia, Canada. Hyd obiologia 596: 57–66. h ps://
doi.o g/10.1007/s10750-007-9057-6
Kock AA, S anb idge D, B ink R, e al. (2022) Table Moun ain Na ional Pa k Ma ine P o ec ed
A ea - S a e o Knowledge. In e nal Repo , Sou h A ican Na ional Pa ks, Scien i ic Se ices,
P e o ia, 47 pp.
Kuma Rai P, Singh JS (2020) In asi e alien plan species: Thei impac on en i onmen , ecosys-
em se ices and human heal h. Ecological Indica o s 111: 106020. h ps://doi.o g/10.1016/j.
ecolind.2019.106020
Kumschick S, Measey JG, Vime ca i G, de Villie s FA, Mokha la MM, Da ies SJ, Tho p CJ, Rebelo
AD, Blackbu n TM, K aus F (2017) How epea able is he en i onmen al impac classi ica ion o
alien axa (EICAT)? Compa ing independen global impac assessmen s o amphibians. Ecology
and E olu ion 7: 2661–2670. h ps://doi.o g/10.1002/ece3.2877
Leh iniemi M, Ou inen O, Pun ila-Dodd R (2020) Ci izen science p o ides added alue in he mon-
i o ing o coas al non-indigenous species. Jou nal o En i onmen al Managemen 267: 110608.
h ps://doi.o g/10.1016/j.jen man.2020.110608
Lina es MS, Macedo DR, Ma ques JC, Callis o M (2022) An h opogenically physically changed
habi a s enable an easie p opaga ion o in asi e bi al e in neo opical headwa e s eams. Biolog-
ical In asions 24: 3855–3863. h ps://doi.o g/10.1007/s10530-022-02886-4
Lins DM, Zbawicka M, Wenne R, Poćwie z-Ko us A, JMolina RA, Al es LP, Rocha RM (2021)
Ecology and gene ics o My ilus gallop o incialis: A h ea o bi al e aquacul u e in sou he n
B azil. Aquacul u e (Ams e dam, Ne he lands) 540: 736753. h ps://doi.o g/10.1016/j.aquacul-
u e.2021.736753
Lockwood JL, Hoopes MF, Ma che i MP (2013) In asion ecology (2nd edn.) Wiley-Blackwell, Ox o d.
Lou ei o TG, Pe e s K, Robinson TB (2021) D opping pla es o pick up aliens: Towa ds a s an-
da dised app oach o moni o ing alien ouling species. A ican Jou nal o Ma ine Science 43:
483–497. h ps://doi.o g/10.2989/1814232X.2021.1989488
Lozano-Co és DF, Zapa a FA (2014) In e eb a e coloniza ion on a i icial subs a es in a co al ee
a Go gona Island, Colombian Paci ic Ocean. Re is a de Biología T opical 62: 161–168. h ps://
doi.o g/10.15517/ b . 62i0.16273

415
NeoBio a 102: 399–418 (2025), DOI: 10.3897/neobio a.102.149275
Sa ah J. Ackland e al.: In asions in A ican MPAs
Mabin CA, Robinson TB, Wilson JRU, Hi sch H, Cas illo ML, Joos e M, Roux JJL (2022) Molecu-
la insigh s in o he in asion dynamics o Ca cinus c abs in Sou h A ica. Biological In asions 24:
3597–3613. h ps://doi.o g/10.1007/s10530-022-02865-9
Mancinelli G, Chainho P, Cilen i L, Falco S, Kapi is K, Ka selis G, Ribei o F (2017) The A lan-
ic blue c ab Callinec es sapidus in sou he n Eu opean coas al wa e s: Dis ibu ion, impac and
p ospec i e in asion managemen s a egies. Ma ine Pollu ion Bulle in 119: 5–11. h ps://doi.
o g/10.1016/j.ma polbul.2017.02.050
McCann L, Kei h I, Ca l on JT, Ruiz GM, Dawson TP, Collins K (2015) Fi s eco d o he non-na-
i e b yozoan Ama hia (= Zoobo yon) e icilla a (delle Chiaje, 1822) (C enos oma a) in he
Galápagos Islands. BioIn asions Reco ds 4: 255–260. h ps://doi.o g/10.3391/bi .2015.4.4.04
McKenzie LA, B ooks R, Johns on EL (2011) He i able pollu ion ole ance in a ma ine in ade .
En i onmen al Resea ch 111: 926–932. h ps://doi.o g/10.1016/j.en es.2010.12.007
Meeus S, Sil a-Rocha I, Ad iaens T, B own PMJ, Cha osia N, Cla amun -López B, Ma inou AF,
MJPocock O, P eda C, Roy HE, T ica ico E, G oom QJ (2023) Mo e han a bi o un: The
mul iple ou comes o a biobli z. Bioscience 73: 168–181. h ps://doi.o g/10.1093/biosci/biac100
Mghili B, Lamine I, Laam aoui MR, Aksissou M, Galanidi M (2024) Upda ing he na ional lis
o ma ine alien species in Mo occo. Medi e anean Ma ine Science 25: 231–249. h ps://doi.
o g/10.12681/mms.35846
Micael J, Ja dim N, Núñez C, Occhipin i-Amb ogi A, Cos a AC (2016) Some B yozoa species e-
cen ly in oduced in o he Azo es: Rep oduc i e s a egies as a p oxy o u u e sp ead. Helgoland
Ma ine Resea ch 70: 7. h ps://doi.o g/10.1186/s10152-016-0458-7
Munzi S, Isoc ono D, Ra e a S (2023) Can we us iNa u alis in lichenology? E alua ing he e -
ec i eness and eliabili y o a i icial in elligence in lichen iden i ica ion. Lichenologis 55: 193–
201. h ps://doi.o g/10.1017/S0024282923000403
Nuñez MA, Paucha d A (2010) Biological in asions in de eloping and de eloped coun ies: Does
one model i all? Biological In asions 12: 707–714. h ps://doi.o g/10.1007/s10530-009-9517-1
Oja ee H, Galil BS, Campbell ML, Ca l on JT, Canning-Clode J, Cook EJ, Da idson AD, Hewi
CL, Jelme A, Ma chini A, McKenzie CH, Minchin D, Occhipin i-Amb ogi A, Olenin S, Ruiz
G (2015) Classi ica ion o non-indigenous species based on hei impac s: Conside a ions o ap-
plica ion in ma ine managemen . PLoS Biology 13: e1002130. h ps://doi.o g/10.1371/jou nal.
pbio.1002130
Okuha a BK, Dele aux JMS, Richa ds Donà A, Smi h CM, Gibson VL, Dulai H, El-Kadi AI, S a-
moulis K, Bu ne KM, Wada CA, B eme LL (2023) E ec s o mul iple d i e s o en i onmen al
change on na i e and in asi e mac oalgae in nea sho e g oundwa e dependen ecosys ems. Wa e
Resou ces Resea ch 59: e2023WR034593. h ps://doi.o g/10.1029/2023WR034593
Oya zún PA, To o JE, Nuñez JJ, e al. (2024) The Medi e anean mussel My ilus gallop o incialis
(Mollusca: Bi al ia) in Chile: dis ibu ion and gene ic s uc u e o a ecen ly in oduced in asi e
ma ine species. Animals 14: 823. h ps://doi.o g/10.3390/ani14060823
Pa isi C, Sandonnini J, Coppola MR, Madonna A, Abdel-Gawad FK, Si ie i EM, Gue ie o G
(2022) Biocide s. eco- iendly an i oulan s: Role o he an ioxida i e de ence and se lemen
in My ilus gallop o incialis. Jou nal o Ma ine Science and Enginee ing 10: 792. h ps://doi.
o g/10.3390/jmse10060792
P a MC, Bicca d A, M ula PE, Olbe s J, Mushanganyisi K, Macdonald A, Samaai T (2022) Gian s
and i ans: Fi s eco ds o he in asi e aco n ba nacles Megabalanus in innabulum (Linnaeus,
1758) and Megabalanus coccopoma (Da win, 1854) on in e idal ocky sho es o Sou h. BioIn a-
sions Reco ds 11: 721–737. h ps://doi.o g/10.3391/bi .2022.11.3.14
Pimm SL, Alibhai S, Be gl R, Dehgan A, Gi i C, Jewell Z, Joppa L, Kays R, Loa ie S (2015) Eme g-
ing echnologies o conse e biodi e si y. T ends in Ecology & E olu ion 30: 685–696. h ps://
doi.o g/10.1016/j. ee.2015.08.008
416
NeoBio a 102: 399–418 (2025), DOI: 10.3897/neobio a.102.149275
Sa ah J. Ackland e al.: In asions in A ican MPAs
Pos les M, Ba le M (2018) The ise o biobli z: E alua ing a popula e en o ma o public en-
gagemen and wildli e eco ding in he Uni ed Kingdom. Applied En i onmen al Educa ion and
Communica ion 17: 365–379. h ps://doi.o g/10.1080/1533015X.2018.1427010
Po gie e LJ, e Huu ne MB, Richa dson DM (2024) Communi y science can in o m in asi e spe-
cies managemen : Melaleuca (My aceae) in Sou h A ica. Ecological Solu ions and E idence 5:
e12391. h ps://besjou nals.onlinelib a y.wiley.com/doi/10.1002/2688-8319.12391
Pyšek P, Richa dson DM, Pe gl J, Ja ošík V, Six o á Z, Webe E (2008) Geog aphical and axo-
nomic biases in in asion ecology. T ends in Ecology & E olu ion 23: 237–244. h ps://doi.
o g/10.1016/j. ee.2008.02.002
Pyšek P, Hulme PE, Simbe lo D, Bache S, Blackbu n TM, Ca l on JT, Dawson W, Essl F, Foxc o
LC, Geno esi P, Jeschke JM, Kühn I, Liebhold AM, Mand ak NE, Meye son LA, Paucha d A,
Pe gl J, Roy HE, Seebens H, an Kleunen M, Vilà M, Wing ield MJ, Richa dson DM (2020)
Scien is s’ wa ning on in asi e alien species. Biological Re iews o he Camb idge Philosophical
Socie y 95: 1511–1534. h ps://doi.o g/10.1111/b .12627
R Co e Team (2023) R: A language and en i onmen al o s a is ical compu ing. R Founda ion o
S a is ical Compu ing, Vienna, Aus ia. h p://www.R-p ojec .o g/
Rees SE, Pi man SJ, Fos e N, Langmead O, G i i hs C, Fle che S, Johnson DE, A ill M (2018)
B idging he di ide: Social-ecological cohe ence in ma ine p o ec ed a ea ne wo k design. Aqua ic
Conse a ion 28: 754–763. h ps://doi.o g/10.1002/aqc.2885
Ren J, Chen J, Xu C, e al. (2021) An in asi e species e odes he pe o mance o coas al we land
p o ec ed a eas. Science Ad ances 7: eabi8943. h ps://doi.o g/10.1126.sciad .abi8943
Renchen GF, Bu le CB, Ma hews TR (2021) Ma ine deb is knows no bounda ies: Cha ac e is ics
o deb is accumula ion in ma ine p o ec ed a eas o he Flo ida Keys. Ma ine Pollu ion Bulle in
173: 112957. h ps://doi.o g/10.1016/j.ma polbul.2021.112957
Riccia di A, Iaca ella JC, Ald idge DC, Blackbu n TM, Ca l on JT, Ca o d JA, J Dick TA, Hulme
PE, Jeschke JM, Liebhold AM, Lockwood JL, MacIsaac HJ, Meye son LA, Pyšek P, Richa dson
DM, Ruiz GM, Simbe lo D, Vilà M, Wa dle DA (2021) Fou p io i y a eas o ad ance in asion
science in he ace o apid en i onmen al change. En i onmen al Re iews 29: 119–141. h ps://
doi.o g/10.1139/e -2020-0088
Richa dson DM, Pyšek P, Ca l on JT (2011) A compendium o essen ial concep s and e -
minology in in asion ecology. In: Richa dson DM (Ed.) Fi y yea s o in asion ecol-
ogy. The legacy o Cha les El on. Wiley-Blackwell, Ox o d, 409–420. h ps://doi.
o g/10.1002/9781444329988.ch30
Richa dson DM, Wi ABR, Pe gl J, Dawson W, Essl F, K e H, an Kleunen M, Weigel P, Win e
M, Pyšek P (2022) Plan in asions in A ica. In: Clemen s DR, Upadhyaya MK, Joshi S, Sh es ha
A (Eds) Global plan in asions. Sp inge In e na ional Publishing, Cham, 225–252. h ps://doi.
o g/10.1007/978-3-030-89684-3_11
Robe s CJ, Ve gés A, Callaghan CT, Poo e AGB (2022) Many came as make ligh wo k: Oppo -
unis ic pho og aphs o a e species in iNa u alis complemen s uc u ed su eys o ee ish o
be e unde s and species ichness. Biodi e si y and Conse a ion 31: 1407–1425. h ps://doi.
o g/10.1007/s10531-022-02398-6
Robinson TB, Ha enga B, Van De Me we M, Jackson S (2017) Mind he gap - con ex dependency
in in asi e species impac s: A case s udy o he ascidian Ciona obus a. NeoBio a 32: 127–141.
h ps://doi.o g/10.3897/neobio a.32.9373
Robinson TB, Hulme PE, Lieu ance D, Richa dson DM (2025) Managing in asions in p o ec ed
a eas: Seeking new s a egies o achie e global a ge s. Biological In asions 27: 118. h ps://link.
sp inge .com/a icle/10.1007/s10530-025-03581-w
Rocha RM, Aze edo F, Oli ei a U, Ca doso MNM, Cle ie PHB, Fo es RR, Lopes-Filho EAP,
Lo ini ML, Mi anda LS, Mou a RB, Senna AR, Sil a FM, S ampa SN, Venekey V (2024) Wes
417
NeoBio a 102: 399–418 (2025), DOI: 10.3897/neobio a.102.149275
Sa ah J. Ackland e al.: In asions in A ican MPAs
A lan ic coas al ma ine biodi e si y: The con ibu ion o he pla o m iNa u alis . Aqua ic Ecolo-
gy 58: 57–71. h ps://doi.o g/10.1007/s10452-023-10062-6
Sala E, Lubchenco J, G o ud-Col e K, No elli C, Robe s C, Sumaila UR (2018) Assessing eal
p og ess owa ds e ec i e ocean p o ec ion. Ma ine Policy 91: 11–13. h ps://doi.o g/10.1016/j.
ma pol.2018.02.004
Sellheim K, S achowicz JJ, Coa es R (2010) E ec s o a nonna i e habi a - o ming species on mo-
bile and sessile epi aunal communi ies. Ma ine Ecology P og ess Se ies 398: 69–80. h ps://doi.
o g/10.3354/meps08341
Shackle on RT, Shackle on CM, Kull CA (2019) The ole o in asi e alien species in shaping local
li elihoods and human well-being: A e iew. En i onmen al Managemen 229: 145–157. h ps://
doi.o g/10.1016/j.jen man.2018.05.007
Shackle on RT, Foxc o LC, Pyšek P, Wood LE, Richa dson DM (2020) Assessing biological in a-
sions in p o ec ed a eas a e 30 yea s: Re isi ing na u e ese es a ge ed by he 1980s SCOPE p o-
g amme. Biological Conse a ion 243: 108424. h ps://doi.o g/10.1016/j.biocon.2020.108424
Simbe lo D (2021) Main enance managemen and e adica ion o es ablished aqua ic in ade s. Hy-
d obiologia 848: 2399–2420. h ps://doi.o g/10.1007/s10750-020-04352-5
Skein L, Robinson TB, Alexande ME (2018) Impac s o mussel in asions on he p ey p e e ence o
wo na i e p eda o s. Beha iou al Ecology 29: 353–359. h ps://doi.o g/10.1093/beheco/a x172
Spyksma AJP, Mille KI, Shea s NT (2024) Ma ine hea wa e p omo es popula ion expansion o
he in asi e ascidian Symplegma b akemhielmi. Biological In asions 26: 1987–1995. h ps://doi.
o g/10.1007/s10530-024-03296-4
S e ani CN, B anch GM (2005) Mechanisms and consequences o compe i ion be ween an alien
mussel, My ilus gallop o incialis, and an indigenous limpe , Scu ellas a a gen illei. Jou nal
o Expe imen al Ma ine Biology and Ecology 317: 127–142. h ps://doi.o g/10.1016/j.jem-
be.2004.11.022
Tho aldsen T, Holmen IM, Moe HK (2015) The escape o ish om No wegian ish a ms:
Causes, isks and he in luence o o ganisa ional aspec s. Ma ine Policy 55: 33–38. h ps://doi.
o g/10.1016/j.ma pol.2015.01.008
T oos K (2010) Causes and e ec s o a highly success ul ma ine in asion: Case-s udy o he in-
oduced Paci ic oys e C assos ea gigas in con inen al NW Eu opean es ua ies. Jou nal o Sea
Resea ch 64: 145–165. h ps://doi.o g/10.1016/j.sea es.2010.02.004
Tsiamis K, Azzu o E, Ba iche M, Çina ME, C oce a F, Cle ck OD, Galil B, Gómez F, Ho man R,
Jensen KR, Kambu ska L, Langeneck J, Lange MR, Ya'a i Le i -Ba ma s, Lezzi M, Ma chini
A, Occhipin i-Amb ogi A, Oja ee H, Pi aino S, Shenka N, Yanko a M, Zene os A, Žulje ić
A, Ca doso AC (2020) P io i izing ma ine in asi e alien species in he Eu opean Union h ough
ho izon scanning. Aqua ic Conse a ion 30: 794–845. h ps://doi.o g/10.1002/aqc.3267
Valla ino EA, Elias R (2008) The dynamics o an in oduced: Balanus glandula popula ion in he
Sou hwes e n A lan ic ocky sho es. The consequences on he in e idal communi y. Ma ine
Ecology (Be lin) 18: 319–335. h ps://doi.o g/10.1111/j.1439-0485.1997. b00445.x
Veiga P, Ramos-Oli ei a C, Sampaio L, Rubal M (2020) The ole o u banisa ion in a ec ing My ilus
gallop o incialis. PLoS One 15: e0232797. h ps://doi.o g/10.1371/jou nal.pone.0232797
Wa kins HV, Yan HF, Dunic JC, Cô é IM (2021) Resea ch biases c ea e o e ep esen ed “pos e chil-
d en” o ma ine in asion ecology. Conse a ion Le e s 14: e12802. h ps://doi.o g/10.1111.12802
Whi low WL (2010) Changes in su i o ship, beha iou , and mo phology in na i e so ‐shell clams
induced by in asi e g een c ab p eda o s. Ma ine Ecology (Be lin) 31: 418–430. h ps://doi.
o g/10.1111/j.1439-0485.2009.00350.x
Wijsman JWM, T oos K, Fang J Ronca a i (2019) Global p oduc ion o ma ine bi al es. T ends and
challenges. In: Smaal AC, Fe ei a JG, G an J, e al. (Eds) Goods and Se ices o Ma ine Bi al es.
Sp inge , Cham, 7–26. h ps://doi.o g/10.1007/978-3-319-96776-9_2
418
NeoBio a 102: 399–418 (2025), DOI: 10.3897/neobio a.102.149275
Sa ah J. Ackland e al.: In asions in A ican MPAs
Wi A, Beale T, Van Wilgen BW (2018) An assessmen o he dis ibu ion and po en ial ecological
impac s o in asi e alien plan species in eas e n A ica. T ansac ions o he Royal Socie y o Sou h
A ica 73: 217–236. h ps://doi.o g/10.1080/0035919X.2018.1529003
Wong MC, Ve caeme B (2012) E ec s o in asi e colonial unica es and a na i e sponge on he
g ow h, su i al, and ligh a enua ion o eelg ass (Zos e a ma ina). Aqua ic In asions 7: 315–
326. h ps://doi.o g/10.3391/ai.2012.7.3.003
Wya ASJ, Hewi CL, Walke DI, Wa d TJ (2005) Ma ine in oduc ions in he Sha k Bay Wo ld
He i age P ope y, Wes e n Aus alia: A p elimina y assessmen . Di e si y & Dis ibu ions 11:
33–44. h ps://doi.o g/10.1111/j.1366-9516.2005.00109.x
Yan T, Lin M, Cao W, Han S, Song X (2021) Fouling cha ac e is ics o cnida ians (Hyd ozoa and
An hozoa) along he coas o China. Jou nal o Oceanology and Limnology 39: 2220–2236.
h ps://doi.o g/10.1007/s00343-021-0242-7
Yu X, Yan Y, Gu J (2007) A achmen o he bio ouling b yozoan Bugula ne i ina la ae a ec ed
by ino ganic and o ganic chemical cues. In e na ional Biode e io a ion & Biodeg ada ion 60:
81–89. h ps://doi.o g/10.1016/j.ibiod.2006.12.993
Zengeya TA, Wilson JR [Eds] (2023) The s a us o biological in asions and hei managemen in
Sou h A ica in 2022. Ki s enbosch and DSI-NRF Cen e o Excellence o In asion Biology,
S ellenbosch, Sou h A ican Na ional Biodi e si y Ins i u e.
Supplemen a y ma e ial 1
Fil e s used o iden i y ma ine p o ec ed a eas along he coas o A ica on he
P o ec ed Plane da abase
Au ho s: Sa ah J. Ackland, Da id M. Richa dson, Tama a B. Robinson
Da a ype: docx
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License (h p://openda a-
commons.o g/licenses/odbl/1.0/). The Open Da abase License (ODbL) is a license ag eemen
in ended o allow use s o eely sha e, modi y, and use his Da ase while main aining his same
eedom o o he s, p o ided ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/neobio a.102.149275.suppl1
Supplemen a y ma e ial 2
Supplemen a y in o ma ion
Au ho s: Sa ah J. Ackland, Da id M. Richa dson, Tama a B. Robinson
Da a ype: xlsx
Explana ion no e: able S2. Ma ine p o ec ed a eas iden i ied om he P o ec ed Plane da abase
and he coun ies in which hey ha e been decla ed. able S3. Ma ine alien species as iden i ied
on WRiMs and he coun ies in which hey ha e been epo ed. able S4. Species epo ed wi h-
in A ican ma ine p o ec ed a eas on iNa u alis . able S5. The assessmen o he magni ude o
impac o species acco ding o he EICAT (Blackbu n e al. 2014) and SEICAT (Bache e al.
2017) schemes.
Copy igh no ice: This da ase is made a ailable unde he Open Da abase License (h p://openda a-
commons.o g/licenses/odbl/1.0/). The Open Da abase License (ODbL) is a license ag eemen
in ended o allow use s o eely sha e, modi y, and use his Da ase while main aining his same
eedom o o he s, p o ided ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/neobio a.102.149275.suppl2