113
An e idence-based p o ocol o de eloping lis s o ee plan ing
Luke J. Po gie e 1,2 , Ma c W. Cado e2, Sab ina Kumschick1,3, T udy Paap4, F ancois Roe s5,
John R.U. Wilson1,3 , Da id M. Richa dson1
1 Cen e o In asion Biology, Depa men o Bo any and Zoology, S ellenbosch Uni e si y, S ellenbosch, Sou h A ica
2 Depa men o Biological Sciences, Uni e si y o To on o Sca bo ough, 1265 Mili a y T ail, To on o, ON, M1C 1A4, Canada
3 Ki s enbosch Resea ch Cen e, Sou h A ican Na ional Biodi e si y Ins i u e, Cape Town, Sou h A ica
4 Depa men o Biochemis y, Gene ics and Mic obiology, Fo es y and Ag icul u al Bio echnology Ins i u e, Uni e si y o P e o ia, P e o ia, Sou h A ica
5 Cen e o In asion Biology, Depa men o Conse a ion Ecology and En omology, S ellenbosch Uni e si y, S ellenbosch, Sou h A ica
Co esponding au ho : Luke J. Po gie e ([email p o ec ed])
Copy igh : © Luke J. Po gie e 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
T ee-plan ing is inc easingly being p omo ed o u ban g eening, ca bon seques a ion, and o enhance
biodi e si y. Howe e , poo ly planned and execu ed ee-plan ing schemes can inad e en ly con ibu e
o biological in asions wi h de imen al e ec s on local ecosys ems, economies, and human well-being.
The e o e, sus ainable, igo ous, epea able, and anspa en species selec ion s a egies a e needed.
We de eloped a s a egic decision p o ocol o iden i ying ee axa sui able o plan ing schemes,
using a mul i-c i e ion app oach ha in eg a es na ional lis s o egula ed in asi e plan species, glob-
al e idence o in asi eness, and suscep ibili y o key pes s. Using he Polyphagous Sho Hole Bo e
(PSHB) in asion in he Ci y o Cape Town, Sou h A ica as a case s udy, we illus a e he p o ocol’s
applica ion and po en ial o in o ming plan ing decisions.
444 ee axa cu en ly plan ed in Cape Town we e assessed. O hese, 85 a e egula ed na ionally
as in asi e species (and a e p ohibi ed om use), while 49 me all sui abili y c i e ia and we e iden i-
ied as candida es o a plan ing lis (i.e., a sa e lis ).
This p o ocol p o ides e idence-based guidance o ee-plan ing o mi iga e he isk o ee in a-
sions and o educe he sp ead and impac o associa ed pes s and pa hogens. This p o ocol is epli-
cable and adap able o use in o he egions and can suppo en i onmen al planne s and manage s
in making in o med decisions o sa egua d ecosys ems and op imise ecosys em se ices (e.g., which
ees o plan in es o a ion ini ia i es).
Key wo ds: Alien species, biological in asions, ecosys em se ices, Euwallacea o nica us, g een
in as uc u e, Neocosmospo a euwallaceae, pes managemen , eescapes, ee in asions, ee plan ing,
u ban g eening
In oduc ion
T ee-plan ing ini ia i es ha e gained widesp ead a en ion as a na u e-based solu-
ion o global en i onmen al challenges, including u ban g eening, ca bon seques-
a ion, biodi e si y conse a ion, and clima e-change mi iga ion (Roy e al. 2012;
Bas in e al. 2019; Seddon e al. 2020). Ac oss he wo ld, go e nmen s, non-p o i
o ganisa ions, and communi ies a e in es ing in la ge-scale ee-plan ing p og ams
o es o e ecosys ems, imp o e ai and wa e quali y, and c ea e g een spaces ha
bene i bo h human heal h and wildli e (e.g., Bonn Challenge, he G ea G een
Academic edi o : Anibal Paucha d
Recei ed:
23 Ap il 2025
Accep ed:
20 Oc obe 2025
Published:
19 No embe 2025
Ci a ion: Po gie e LJ, Cado e
MW, Kumschick S, Paap T, Roe s F,
Wilson JRU, Richa dson DM (2025)
An e idence-based p o ocol o
de eloping lis s o ee plan ing.
NeoBio a 104: 113–137. h ps://doi.
o g/10.3897/neobio a.104.156206
NeoBio a 104: 113–137 (2025)
DOI: 10.3897/neobio a.104.156206
This a icle is pa o :
De eloping lis s o alien axa
in he Global Sou h: wo k lows,
p o ocols, p ocesses, and expe iences
Edi ed by John Wilson, Michele Dechoum,
Ka elyn Faulkne , Ba ba a Langdon,
Shyama Pagad, Aníbal Paucha d, Hanno
Seebens, Tsungai Zengeya, Sil ía Zille
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 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Wall Ini ia i e, T illion T ees Campaign). Many o hese ee-plan ing p og ams
ocus on apid plan ing e o s o mee ambi ious plan ing a ge s and o en o e -
look majo ecosys em impac s (Bond e al. 2019; Moyano e al. 2024) and o he
c i ical conside a ions such as app op ia e species selec ion, si e sui abili y, and he
isks o u u e pes and pa hogen in asions (Holl and B ancalion 2020).
As ee-plan ing ini ia i es gain momen um – o en wi h conside able i-
nancial, poli ical, and socie al suppo – he e is g owing conce n ha poo -
ly planned and execu ed e o s will inad e en ly in oduce new in asi e
non-na i e species o exace ba e exis ing in asions (B undu and Richa dson
2016; Blancha d e al. 2017). Such in asions, o en acili a ed by he plan ing
o non-na i e ee species, can ha e de as a ing e ec s on ecosys ems, econo-
mies, and human heal h (Richa dson and Rejmánek 2011). The plan ing o
na i e species, i done wi hou ega d o local pes dynamics, can also p o ide
sui able hos s o in asi e pes s and pa hogens (Paap e al. 2018). In some
cases, he e y ees in ended o enhance biodi e si y and p o ide ecosys em
se ices can become ec o s o pes p oli e a ion, unde mining he long- e m
goals o he ini ia i e. These o e sigh s can lead o educed biodi e si y, and
comp omised ecosys em se ices, c ea ing he need o cos ly managemen in-
e en ions (Holl and B ancalion 2020; Bindewald e al. 2021).
Sou h A ica has a long his o y o ee in oduc ions, beginning wi h Eu opean
colonisa ion in he mid-17 h cen u y (Poyn on 1984; Richa dson e al. 2003). The
mo i a ions o ee-plan ing ha e e ol ed o e ime – om mee ing basic needs
such as ood, uel, shade, and soil s abilisa ion o bols e ing he o es y sec o
and imp o ing ag icul u al sys ems (Richa dson e al. 2003), and, mo e ecen -
ly, enhancing u ban en i onmen s (Po gie e e al. 2022). Despi e he bene i s o
ee-plan ing, his p ac ice has gene a ed conside able con o e sy. Fo ins ance,
many plan ed non-na i e ee species in oduced o cul i a ion ha e become in-
asi e, sp eading beyond cul i a ion a eas and causing majo nega i e impac s ( an
Wilgen e al. 2022). A subs an ial in asion deb has accumula ed, as many species
ha e ye o ully ealise hei in asi e po en ial o cause maximum impac (Rouge
e al. 2016). Shi ing en i onmen al and socio-poli ical condi ions a e in luencing
ee-plan ing policies and p ac ices in Sou h A ica. Rapid u banisa ion is d i ing
he de elopmen o new and expanding u ban ecosys ems, wi h e o s also unde -
way o e o i exis ing ci ies o enhance sus ainabili y and li eabili y ( an S aden
and S o be g 2021). Inc easing ee co e has been deemed a key componen o
s a egies o build esilience o global change.
The g owing globalisa ion o ade, inc eased mo emen o people and goods,
and changing clima e pa e ns ha e con ibu ed o a d ama ic inc ease in he
sp ead o pes s and pa hogens a ound he wo ld (Hulme 2009; Raum e al.
2023). One o he mos ecen examples is he Polyphagous Sho Hole Bo e
(PSHB, Euwallacea o nica us), an in asi e amb osia bee le, na i e o Sou h-
eas Asia, ha has caused widesp ead damage a ound he wo ld (S ou hame
e al. 2017; Van Rooyen e al. 2021). PSHB, along wi h i s symbio ic ungus
Neocosmospo a euwallaceae L. Lomba d & C ous, 2019 (p e iously Fusa ium eu-
wallaceae, Lomba d e al. 2019), in es s hund eds o axonomically di e se ee
species, such as Ace negundo L., Que cus obu L., and Pla anus × hispanica Mill.
ex Münchh., leading o signi ican ee mo ali y in bo h u ban and na u al en-
i onmen s (F eeman e al. 2016; Townsend e al. 2025). In asi e species like
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NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
PSHB h i e in u ban landscapes, whe e a high densi y o suscep ible hos ees,
coupled wi h human-media ed mo emen , accele a e he in asion p ocess (Paap
e al. 2020; Raum e al. 2023; Po gie e e al. 2024).
Al hough Sou h A ica has aced ex ensi e challenges om in asi e species,
pa icula ly ees ( an Wilgen e al. 2020, 2022), he coun y has un il now
a oided la ge-scale ecological dis up ions om ee disease epidemics (Wilson e
al. 2020) – hough his is changing. The PSHB in es a ion in Sou h A ica is pa
o a b oade pa e n whe e hund eds o ee species, many o which a e c i ical
o ecosys em se ices, a e inc easingly ulne able o pes s and pa hogens (Paap
e al. 2017; Ra a e al. 2023). This ala ming end unde sco es he impo ance o
de eloping p oac i e measu es o mi iga e he isk o biological in asions in ee-
plan ing e o s. While ee-plan ing emains a aluable s a egy o conse a ion
and clima e change mi iga ion (Made 2020; Mo an e al. 2020), he e is a p essing
need o shi om ad hoc species selec ion o mo e s a egic and e idence-based
app oaches ha conside ecological isks.
Speci ically, in he case o he PSHB in es a ion in Sou h A ica, esul ing u -
ban ee mo ali y necessi a es igo ous and anspa en guidelines o eplacemen
plan ings ha do no exace ba e plan o pes in asions in Sou h A ica. B undu
e al. (2020) p o ide global guidelines o he sus ainable use o non-na i e ees,
ou lining he p ecau ions ha should be aken when in oducing and plan ing
non-na i e ees. Thei guidelines include eigh key ecommenda ions, including
p io i ising na i e o non-in asi e non-na i e species, complying wi h egula ions,
assessing in asion isks unde global change scena ios, implemen ing ailo ed sil i-
cul u al p ac ices, and p omo ing ea ly de ec ion, s akeholde engagemen , and in-
e na ional collabo a ion. These guidelines aim o suppo sus ainable o es y, bio-
di e si y conse a ion, and global en i onmen al objec i es. Fu he , Kumschick
e al. (2024) ou line conside a ions o de eloping and implemen ing a sa e lis o
non-na i e axa including c i e ia o isk assessmen s, s akeholde in ol emen ,
and egula o y mechanisms o ensu e e ec i e applica ion.
Ou goal is o ope a ionalise hese guidelines in o a s a egic p o ocol o ee
selec ion ha mi iga es he in asion isk o bo h he plan ed ees and hei associ-
a ed pes s and pa hogens, using he PSHB in asion in Sou h A ica as a case s udy.
Iden i ying ee axa sui able o plan ing
To suppo e idence-based decision-making in ee-plan ing ini ia i es, we p esen
a s uc u ed wo k low ha ou lines he key s eps equi ed o iden i y ee axa
sui able o plan ing (Fig. 1). This wo k low is designed o be adap able o a i-
ous egional con ex s and ecological p io i ies, p o iding a eplicable me hod o
sys ema ically sc eening candida e species lis s using mul iple isk- ela ed c i e ia.
In ou example, we apply his wo k low o ee plan ing in he Ci y o Cape
Town Me opoli an Municipali y, Sou h A ica, a egion acing signi ican
challenges due o widesp ead ee in asions and he de as a ing impac s o
PSHB on bo h na u al and u ban o es s (de Wi e al. 2022). Cape Town was
selec ed as a case s udy due o he a ailabili y o comp ehensi e da a on plan -
ed ees and he u gency o mi iga ing pes and in asion isks. Howe e , he
s eps p esen ed he e a e b oadly applicable and can be adap ed o o he ci ies
o egions acing simila ecological p essu es.
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NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
S ep 1. Compile lis o all ee axa
The i s s ep in ol es compiling a comp ehensi e, baseline lis o all ee axa
al eady p esen in he s udy egion, including bo h plan ed and non-plan ed
ees. This b oad lis should cap u e he ull di e si y o ee species cu en ly
es ablished in na u al, semi-na u al, u ban, and cul i a ed se ings. Da a can
be sou ced om global and egional biodi e si y da abases such as he Global
Biodi e si y In o ma ion Facili y (GBIF, h ps://www.gbi .o g/), GlobalT e-
eSea ch (h ps:// ools.bgci.o g/global_ ee_sea ch.php), iNa u alis (h ps://
www.ina u alis .o g/), and na ional bo anical da abases. Ga dening books,
nu se y ca alogues (especially hose specialising on ees), species lis s om a -
bo e a, and a bo is s and ee en husias s can also be consul ed. Collabo a ion
wi h local bo anical ga dens, o es y depa men s, and esea ch ins i u ions
can ensu e he lis is exhaus i e and egionally app op ia e. To main ain axo-
nomic accu acy, names should be c oss-checked agains s anda dised axonom-
ic e e ences such as he Wo ld Checklis o Vascula Plan s (Go ae s e al.
2024). This p ocess ensu es consis ency in species iden i ica ion.
S ep 2. Compile lis o plan ed ee axa
Once he ull lis o ee axa p esen in he s udy egion has been compiled, he
nex s ep in ol es na owing his lis o include only hose axa ha a e cu en ly
plan ed wi hin he s udy egion. This subse e lec s axa ha a e al eady in use in
u ban landscapes, pa ks, ga dens, s ee scapes, o o he managed se ings wi hin
he s udy egion. The a ionale is ha hese axa ha e al eady been demons a ed
o su i e local condi ions. This s ep is b oadly applicable and can be implemen -
ed using municipal plan ing eco ds, ho icul u al ca alogues, o c owd-sou ced
Figu e 1. A gene ic wo k low o iden i ying ee axa sui able o plan ing. The p ocess begins wi h
compiling comp ehensi e lis s o all ee axa o he s udy egion and hose sui able o plan ing.
Fil e ing and inclusion c i e ia a e hen de ined, ollowed by da a colla ion o each c i e ion. Rel-
e an da a sou ces a e c oss- e e enced wi h he sui able axa lis , which is subsequen ly applied o
a decision p o ocol o de e mine he inal se o axa ecommended o plan ing. The wo k low is
i e a i e and adap i e (shown by he dashed a ow), wi h ce ain s eps subjec o ongoing e inemen
as new da a and insigh s become a ailable.
1.
2.
3.
4.
5.
6.
Compile lis o all ee axa
Compile lis o plan ed ee axa
Selec inclusion c i e ia
Colla e da a o each c i e ion
C oss- e e ence da a sou ces wi h plan ed ee axa lis
Apply plan ed ee axa o decision p o ocol
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NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
biodi e si y pla o ms such as iNa u alis . In addi ion o axa al eady plan ed in
he s udy egion, he lis can be expanded o include species no ye plan ed local-
ly bu iden i ied as sui able h ough es ablished isk assessmen s, clima e-adap ed
species guidelines, o p o enance ials. These candida e axa should be suppo ed
by obus e idence and aligned wi h local ecological and egula o y con ex s.
Fo his s udy, we used he na ional plan ed ee in en o y o Sou h A ica, which
comp ises o e 35,000 eco ds o he 805 ee axa cu en ly plan ed ac oss he coun-
y (he eina e e e ed o as he “ ee in en o y”; Richa dson and Po gie e 2024).
Only ee axa al eady p esen in Sou h A ica we e conside ed o plan ing o a oid
p omo ing new in oduc ions. In oducing new species ca ies inhe en ecological
isks, including un o eseen in asion po en ial, and equi es igo ous sc eening be-
o e impo s a e app o ed [ o de ails o he p ocess in Sou h A ica see Wilson and
Kumschick (2024)]. The esul ing lis o egionally es ablished ee axa cons i u es
he candida e pool o u he e alua ion agains in asion and pes isk c i e ia.
S ep 3. Selec inclusion c i e ia
A key componen o he p o ocol in ol es es ablishing clea c i e ia o sys ema i-
cally assess he po en ial isks associa ed wi h plan ing pa icula ee axa. This s ep
ensu es ha species selec ion is no based on ad hoc o subjec i e p e e ences. The
c i e ia de ine which a ibu es a e ele an o de e mining whe he a gi en ee axon
should be ecommended o o excluded om plan ing ini ia i es. These c i e ia can
be ailo ed o he speci ic goals, con ex , and managemen p io i ies o a gi en egion.
In his s udy, hese c i e ia include key isk ac o s associa ed wi h bo h ee
in asi eness and suscep ibili y o PSHB. The i s c i e ion ocused on he egula-
o y s a us o ee axa, speci ically iden i ying hose axa lis ed as in asi e unde
Sou h A ica’s Na ional En i onmen al Managemen : Biodi e si y Ac 10 o 2004
Alien and In asi e Species Regula ions [NEM:BA A&IS Regula ions, hese we e
mos ecen ly upda ed in 2020, o de ails see Wilson and Kumschick (2024)].
Howe e , o icial lis s migh no cap u e eme ging o localised in asions, especially
in egions wi h in equen ly upda ed in asi e species lis s. To add ess his, collab-
o a ion wi h local p ac i ione s can help iden i y incipien in asions and ensu e
plan ing decisions e lec he mos cu en on- he-g ound knowledge.
The second c i e ion conside ed PSHB hos s a us, dis inguishing be ween
Neocosmospo a-colonised, compe en , and kill-compe en hos s (Townsend e
al. 2025). Neocosmospo a-colonised hos s a e hose in which ungal ansmission
om PSHB is possible, bu which do no suppo bee le ep oduc ion. Compe-
en hos s a e hose in which he bee le success ully es ablishes a na al galle y and
p oduces o sp ing. Kill-compe en hos s included hose ee axa o which a
leas one indi idual has been documen ed as succumbing o he combined e -
ec s o he bee le and N. euwallaceae in es a ions. To implemen hese c i e ia,
mul iple da a sou ces we e consul ed.
While he wo p ima y c i e ia ou lined he e a e cen al o ou case s udy, he
p o ocol allows o addi ional c i e ia o be used. Fo example, in o he egions,
il e s based on clima e sui abili y, i e suscep ibili y, o social and cul u al alues
migh be ele an . The selec ion and p io i isa ion o c i e ia should be adap -
ed acco ding o local ecological con ex s, egula o y amewo ks, and s ake-
holde needs, ensu ing ha he p o ocol emains lexible and applicable ac oss
di e se plan ing scena ios.
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Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
S ep 4. Colla e da a o each c i e ion
Wi h he c i e ia es ablished, he nex s ep in ol es colla ing axa lis s o each c i-
e ion. Mul iple independen da ase s should be used whe e e possible o ensu e
a obus and anspa en assessmen . App op ia e da a sou ces will a y depending
on he ocus o he assessmen , he a ailabili y o egion-speci ic in o ma ion, and
he geog aphic scope o he p o ocol’s applica ion.
We used 1) he NEM:BA A&IS Lis s o 2020 (Wilson 2024) o de e mine
he egula o y s a us and legal implica ions o each axon, 2) he biogeog aphical
s a us (na i e o non-na i e) using he ca ego isa ion o Richa dson and Po gie e
(2024), 3) he Global Na u alised Alien Flo a (GloNAF, upda ed ollowing an
Kleunen e al. 2019) o de e mine whe he each axon is na u alised in and/o
ou side o Sou h A ica, 4) he PSHB global hos lis (DPIRD 2024), which was
compiled and alida ed using molecula eco ds con i ming PSHB iden i y om
mul iple sou ces, and 5) Lynch e al. (2025) o de e mine he hos s a us o each
ee axon. We also iden i ied axa po en ially suscep ible o PSHB in es a ion by
e alua ing whe he hey belong o he same genus as con i med PSHB hos s, gi en
he inc eased likelihood o suscep ibili y among closely ela ed species (Lynch e
al. 2025); no ing such axa migh ha e no ye been exposed, in ec ions no o -
mally eco ded ye , o could become suscep ible o e ime. To accoun o di e -
ing Neocosmospo a suscep ibili ies wi hin gene a, axa wi h unknown s a uses we e
conse a i ely assigned he highes known isk le el among hei congene s. This
p ecau iona y app oach aims o ensu e ha po en ial isks we e no unde es ima -
ed, i.e., i is a conse a i e assessmen o hos suscep ibili y.
S ep 5. C oss- e e encing da a sou ces wi h plan ed axa lis
Once all ele an da a ha e been colla ed o each c i e ion, he nex s ep in ol es
in eg a ing hese da ase s wi h he cu a ed lis o plan ed ee axa. This s ep is
he co e analy ical phase o he p o ocol, as i in eg a es he mul iple isk ac o s
assessed in he p e ious s ep. By sys ema ically c oss- e e encing each ee axon
agains he selec ed da a sou ces, comp ehensi e, e idence-based classi ica ion can
be de eloped ha iden i ies axa sui able o plan ing, hose ha should be exclud-
ed, and hose equi ing u he moni o ing.
We c oss- e e enced ou cu a ed lis o plan ed ee axa wi h mul iple key da a
sou ces (desc ibed abo e). This p ocess in ol ed ex ac ing and assigning a ibu es
o each axon, including i s lis ing in he NEM:BA A&IS Regula ions, in asion s a-
us, and PSHB hos s a us. This app oach ensu ed ha each axon was e alua ed
using mul iple, independen c i e ia, allowing o a obus and e idence-based classi-
ica ion o ee axa in ela ion o bo h in asion po en ial and suscep ibili y o PSHB.
S ep 6. Apply plan ed ee axa o decision p o ocol
We p esen a s uc u ed, e idence-based decision p o ocol o suppo he iden i i-
ca ion o ee axa sui able o plan ing (Fig. 2). Designed o guide planne s, land
manage s, and policymake s, he p o ocol ollows a sequence o il e ing and as-
sessmen s eps ha inco po a e bo h gene al ecological isks, such as na u alisa ion
po en ial, and con ex -speci ic h ea s, including pes o pa hogen suscep ibili y.
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Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
The decision p o ocol in eg a es he ou pu s o S eps 3–5 in o a anspa en ,
epea able amewo k o de e mining which ee axa a e sui able o plan ing
o should be excluded. I consis s o eigh a ge ed ecommenda ions ( a ious-
ly add essing ee plan ing, emo al, and moni o ing). In cases o unce ain y,
he p o ocol encou ages consul a ion o global o egion-speci ic da a sou ces
o e ine ecommenda ions.
The p o ocol is in ended o a wide ange o s akeholde s, including go e nmen
agencies, non-go e nmen al o ganisa ions, en i onmen al consul an s, landscape
a chi ec s, and es o a ion p ac i ione s. While pa icula ly ele an o u ban and
municipal planning, i is equally applicable o b oade o es and land-use con-
ex s. I s lexible, mul i-c i e ion s uc u e enables use s o iden i y low- isk ee
axa ailo ed o local ecological condi ions and egula o y amewo ks, he eby
educing he likelihood o biological in asions and pes ou b eaks. Al hough some
amilia i y wi h ee ecology and access o ele an da a a e bene icial, he p o-
ocol is designed o be anspa en , adap able, and accessible o use s wi h a y-
ing le els o expe ise. Whe e needed, implemen a ion can be suppo ed h ough
collabo a ion wi h expe s o local au ho i ies.
Figu e 2. Decision p o ocol o ee selec ion o mi iga e he isk o Polyphagous Sho Hole Bo e (PSHB, Euwallacea o nica us)
in es a ion and ee in asions. The p o ocol consis s o eigh ecommenda ions ca ego ised in o h ee ypes: ee-plan ing, ee e-
mo al, and ee moni o ing. These ecommenda ions a e u he g ouped in o scena ios, ep esen ed by a colou g adien om da k
b own o g een. Scena io B is when plan ing he ee axon can esul in a high likelihood o bo h PSHB in es a ion and in asion
o he ee axon i sel , necessi a ing he exclusion o ce ain ee axa om plan ing ini ia i es. P og essi ely ligh e shades ep e-
sen dec easing likelihood o PSHB in es a ion and/o ee in asion, wi h g een deno ing ee axa posing a low likelihood o bo h
PSHB in es a ion and ee in asion and which a e hus sui able o plan ing. Should he e be any unce ain y ega ding decisions,
he global da a sou ces highligh ed in he Me hods sec ion (o any egion-speci ic da a sou ces ele an o he local con ex ) should
be consul ed.
No
No
No
No
Yes
Yes
Yes
Yes
Yes
Is i lis ed unde he
Alien and In asi e
Species egula ions?
Is i a kill-compe en hos ?
Is i a compe en hos ?
No
Is i in he same genus as a
compe en hos ?
Is i a Neocosmospo a-colonised
hos ?
Is i na u alised in o
ou side o he coun y?
Is i in he same genus as
Neocosmospo a -colonised hos ?
Is i na u alised in o
ou side o he coun y?
Is i na u alised in o
ou side o he coun y?
Is i na u alised in o
ou side o he coun y?
A oid plan ing he axon
Remo e ees o he same species
in PSHB-in es ed and ecologically
sensi i e a eas
Moni o su ounding ees o
signs o PSHB in es a ion and
na u alisa ion o he ee axon
Remo e ees o he same species
in PSHB-in es ed and ecologically
sensi i e a eas
Moni o su ounding ees o
signs o PSHB in es a ion and
na u alisa ion o he ee axon
I plan ed, moni o he ee axon
and su ounding ees o signs o
PSHB in es a ion and na u alisa ion
o he ee axon
I plan ed, moni o he ee axon
and su ounding ees o signs o
PSHB in es a ion and
na u alisa ion o he ee axon
A oid la ge-scale plan ing in
hea ily PSHB-in es ed a eas
I plan ed, moni o he ee axon
and su ounding ees o signs o
PSHB in es a ion and
na u alisa ion o he ee axon
I plan ed, moni o he ee axon
o signs o na u alisa ion
Plan ing Remo al Moni o ing
RECOMMENDATIONS
CRITERIA
PSHB In asion s a us
Yes
Yes
No
Yes
Yes
Yes
Yes
Is i na u alised in o
ou side o he coun y?
No
A
B
C
D
E
F
G
Do no plan unless unde pe mi o
exemp ion
Remo e exis ing ees i specified
unde he egula ions
Plan ings o be eco ded and
epo ed o acili a e egula o y
compliance
No
Yes
Is i na u alised in o
ou side o he coun y?
H
I plan ed, moni o he ee axon
in PSHB-in es ed a eas
Is i non-na i e?
Is i non-na i e?
Is i non-na i e?
Is i non-na i e?
Is i non-na i e?
Is i non-na i e?
Yes
No
No
No
No
No
No
No
No
No
No
Yes
Yes
Yes
Yes
Yes
A oid la ge-scale plan ing in
ecologically sensi i e a eas
Sa e o plan ing
A oid la ge-scale plan ing in
ecologically sensi i e a eas
Sa e o plan ing
A oid la ge-scale plan ing in
ecologically sensi i e a eas
A oid la ge-scale plan ing in
PSHB-in es ed a eas
A oid plan ing in ecologically
sensi i e a eas
A oid plan ing in PSHB-in es ed
a eas
A oid plan ing in ecologically
sensi i e a eas
A oid plan ing in PSHB-
in es ed a eas
Remo e ees o he same species in
ecologically sensi i e a eas
Remo e ees o he same species in
ecologically sensi i e a eas
Remo e ees o he same species in
ecologically sensi i e a eas
Remo e ees o he same species in
ecologically sensi i e a eas
S a he e
No applicable
120
NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
We applied his p o ocol o he Ci y o Cape Town Me opoli an Municipali y,
and ca ego ised ee axa om high o low likelihood o PSHB in es a ion and
in asion o he ee axon i sel . Ta ge ed ecommenda ions associa ed wi h each
ca ego y we e also p o ided. The ci y has he mos comp ehensi e plan ed ee da a
co e age o any municipali y in Sou h A ica (Richa dson and Po gie e 2024) and
has also ecen ly expe ienced PSHB in asions ( i s de ec ed in 2019) wi h o e
10,000 ees in es ed and ~4000 al eady emo ed om public land due o PSHB
in es a ion (Ci y o Cape Town In asi e Species Uni , pe s. comm., 14 Ma ch 2025;
Fig. 3a). The ci y has a ela i ely low ee canopy co e a jus six pe cen (Ci y o
Cape Town 2021), and his is likely o dec ease u he due o u ban expansion,
ee emo als o in as uc u e de elopmen , he impac s o clima e change, and
pes -pa hogen in es a ions. By aligning wi h global amewo ks such as he Uni ed
Na ions Sus ainable De elopmen Goals (Uni ed Na ions 2015), and ini ia i es
like he New U ban Agenda (Uni ed Na ions 2017), and he Global Co enan
o Mayo s o Clima e and Ene gy (GCoM 2022), Cape Town has commi ed o
de eloping policies ha aim o augmen i s g een in as uc u e o mee hese in-
e na ional a ge s while ensu ing a heal hie , mo e li eable u u e o i s esiden s.
Applying he p o ocol a he municipal le el allows o con ex -speci ic deci-
sions ha align wi h en i onmen al condi ions, socio-economic p io i ies, and
egula o y equi emen s. This scale also suppo s in eg a ion wi h u ban g eening,
biodi e si y conse a ion, and clima e esilience s a egies, and os e s collabo a-
ion wi h local s akeholde s. The p o ocol is b oadly ans e able and can be adap -
ed o add ess di e en h ea s (e.g., d ough mo ali y, lammabili y) and applied
ac oss spa ial scales, om municipali ies o p o inces o coun ies, depending on
a ailable da a and egional p io i ies.
The wo k low p oduces an e idence-based lis o ee axa sa e o plan ing,
p o iding a scien i ically in o med ounda ion o species selec ion in ee-plan ing
ini ia i es. In ou case s udy, he ee axa on his lis a e 1) no cu en ly lis ed as
in asi e unde na ional legisla ion, 2) no known o be na u alised in Sou h A ica
o elsewhe e, and 3) no known o be suscep ible o PSHB. These axa can he e-
o e cu en ly be ecommended as sui able o plan ing in he ci y. Depending on
he applica ion, lis s can be ailo ed o speci ic land-use con ex s (e.g., s ee ees
s. ipa ian bu e s) o demog aphic needs (e.g. alle gen- ee ees o schools).
The lis can also suppo nu se y indus y planning and public p ocu emen by
highligh ing demand o low- isk species. The lis s should be e iewed pe iodically
as new da a eme ge.
The a ailabili y o axa on his lis in nu se ies o online plan e aile s in Sou h
A ica was also assessed. To de e mine his, we sea ched a ep esen a i e selec ion
o nu se y ca alogues, including bo h p in ed ca alogues om majo Sou h A ican
wholesale and e ail nu se ies and publicly a ailable online pla o ms. Online plan
e ail s o es we e iden i ied h ough a ge ed keywo d sea ches (e.g., “buy [spe-
cies name] ee Sou h A ica”, “ ee nu se y Sou h A ica”, “o namen al ee sales
Sou h A ica”). Each axon om he inal il e ed lis was c oss-checked agains
hese sou ces o eco d i s a ailabili y in one o mo e li e s ages (seeds, seedlings,
saplings, o ma u e ees). A axon was conside ed accessible i i was a ailable o
sale a any li e s age (seeds, seedlings, saplings, o ma u e ees) h ough a leas
one nu se y o online plan e aile . This p ocess p o ided an indica ion o each
axon’s accessibili y wi hin he ho icul u al ade, which is ele an o p ac ical
implemen a ion o plan ing ecommenda ions.
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NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Case s udy: Ci y o Cape Town
Decision p o ocol
T ee axa lis ed unde he NEM:BA A&IS Regula ions mus no be plan ed
unless a pe mi o exemp ion has been g an ed by he ele an en i onmen al
au ho i y (scena io A, Fig. 3b). Unau ho ised plan ing can esul in legal conse-
quences and con ibu e o socio-ecological and economic impac s and con lic s.
Figu e 3. Pho o panel including. a. Polyphagous Sho Hole Bo e (PSHB, Euwallacea o nica us; pho o c edi : A de Villie s); b. A pop-
ula o namen al ee Me oside os excelsa Sol. ex Gae n. (New Zealand Ch is mas ee; pho o c edi : DM Richa dson) lis ed unde Sou h
A ica’s Na ional En i onmen al Managemen : Biodi e si y Ac 10 o 2004 Alien and In asi e Species Regula ions; c. Signs o PSHB in-
es a ion o Ace negundo L. (Box elde ; pho o c edi : LJ Po gie e ), a kill-compe en hos , and d. Dais co ini olia L. (Pom Pom ee; pho o
c edi : DM Richa dson) cu en ly deemed sa e o plan ing ac oss he Ci y o Cape Town Me opoli an Municipali y.
128
NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Ca negie and G an (2025) highligh he c i ical ole o social license in he suc-
cess o u ban biosecu i y esponses o in asi e ee pes s. They emphasise ha man-
agemen e o s, such as ee emo al o pes icide applica ions, o en ace public
esis ance due o di e se s akeholde alues associa ed wi h u ban ees. The e iew
iden i ies key ac ions o imp o e he social accep abili y o con ol measu es, in-
cluding ea ly s akeholde engagemen , us -building, pa icipa o y decision-mak-
ing, and e ec i e communica ion abou he ecological and economic isks o pes
incu sions. By p oac i ely add essing social conce ns, biosecu i y agencies can en-
hance public suppo o necessa y in e en ions, ul ima ely inc easing he likeli-
hood o success ul pes e adica ion.
Ou decision p o ocol aligns wi h hese p inciples by inco po a ing an e i-
dence-based assessmen o p omo e ee axa selec ion by minimising u u e con-
lic s be ween biosecu i y ac ions and u ban ee managemen . By p io i ising
non-in asi e and non-hos axa, he p o ocol educes he need o eac i e in-
e en ions, such as ee emo al, ha could o he wise igge public opposi ion.
Mo eo e , he ecommenda ion o moni o plan ings o signs o in es a ion sup-
po s ea ly de ec ion, a key ac o in bo h biosecu i y success and s akeholde us
(Liebhold e al. 2012). In eg a ing social licence conside a ions in o ee selec ion
and biosecu i y planning can hus enhance bo h ecological esilience and public
coope a ion in managing in asi e ee pes s like PSHB.
A key conside a ion in ee selec ion is p io i ising na i e axa and p omo ing
di e si y o enhance ecosys em esilience and educe suscep ibili y o pes -pa ho-
gen in asions (Vashis e al. 2025). Monocul u es o axa wi h simila ai s o
in es ed hos s p o ide li le esis ance, allowing pes -pa hogen popula ions o pe -
sis . A di e si y-based plan ing app oach limi s hos a ailabili y, educes he isk o
la ge-scale pes ou b eaks and aligns wi h b oade u ban o es esilience p inciples
(Paque e e al. 2021).
Shackle on and Gwedla (2021) explo ed how colonial and apa heid legacies
ha e shaped u ban g een in as uc u e in Sou h A ica, leading o pe sis en in-
equali ies in he dis ibu ion and composi ion o u ban g een spaces. They high-
ligh ha weal hie , his o ically whi e neighbou hoods e ain ex ensi e g een
spaces domina ed by non-na i e ee species in oduced du ing colonial ule,
whe eas lowe -income a eas, his o ically designa ed o ma ginalised communi-
ies, ha e ewe and lowe -quali y g een spaces. This colonial legacy has no only
in luenced u ban ecology bu has also esul ed in a signi ican bu den o in asi e
species managemen . The s udy also c i iques how public g een spaces con inue
o e lec colonial-e a aes he ics and ec ea ional no ms, o en neglec ing A i-
can cul u al pe spec i es and con empo a y u ban needs. The au ho s ad oca e
o a mo e inclusi e, pa icipa o y app oach o u ban g eening ha conside s
local iden i ies, wo ld iews, and ecological esilience. Ou p o ocol allows o
he selec ion o na i e ee axa which can suppo e o s o b eak om colo-
nial-e a plan ing p ac ices ha ha e con ibu ed o ongoing ecological and social
dispa i ies. P io i ising na i e species and a di e si y-based plan ing app oach
can suppo he c ea ion o mo e ecologically sus ainable and socially equi able
u ban o es s. Add essing he his o ical imbalance in g een space dis ibu ion,
coupled wi h he p oac i e selec ion o app op ia e ee axa, has he po en ial o
os e u ban landscapes ha a e bo h esilien o biological in asions and mo e
e lec i e o local cul u al and en i onmen al p io i ies.
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NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
The impo ance o go e nance o imp o ing he sus ainabili y o ee-plan -
ing schemes is ecei ing inc easing a en ion (Yi ba ek e al. 2023). The p o ocol
desc ibed he e is essen ial backg ound o bo h p ojec ini ia ion and planning
phases, i.e., guiding he selec ion o sui able ees.
Moni o ing
Es ablishing a sys ema ic moni o ing p o ocol o ees in egions whe e pes s and
pa hogens a e p esen o pose a isk o in oduc ion is essen ial o ea ly de ec ion
and e ec i e managemen (Po gie e e al. 2024). Fo PSHB, ou ine isual in-
spec ions should be conduc ed o iden i y cha ac e is ic symp oms o in es a ion
pa icula ly on he unk and b anches, including s aining a ound he en y holes,
ass (sawdus -like deb is), gumming, and discolo a ion ( an Rooyen e al. 2021).
These indica o s can signal he p esence o PSHB be o e widesp ead damage occu s.
Inco po a ing bai ed aps in o moni o ing p og ams can u he enhance de ec ion
e o s and imp o e esponse imes. Communi y science ini ia i es can acili a e
b oad-scale su eillance, as local s akeholde s can assis in epo ing ea ly signs o
in es a ion (Po gie e e al. 2024). Collabo a ion wi h en i onmen al and o es y
agencies o es ablish a egional moni o ing ne wo k would u he s eng hen de ec-
ion capabili ies and ensu e he dissemina ion o bes p ac ices o ee managemen .
Educa ing esiden s, landscape manage s, and municipal au ho i ies abou he
isks o plan ing pes -suscep ible o in asi e species and he impo ance o p omp
pes epo ing can imp o e su eillance and apid esponse e o s. Whe e pes -sus-
cep ible ee species mus be e ained o cul u al, aes he ic, o ecological easons, an
In eg a ed Pes Managemen (IPM) app oach can be implemen ed. This may in ol e
egula moni o ing, physical con ol (e.g., emo al o in es ed ma e ial), and, whe e
necessa y, he ca e ul applica ion o chemical ea men s o high- alue ees o limi
pes -pa hogen sp ead while minimising unin ended ecological consequences.
Applicabili y beyond Cape Town
The wo k low (Fig. 1) and he mul i-c i e ion decision p o ocol (Fig. 2) can be
applied ac oss di e se ecological, geog aphic, and egula o y con ex s. Use s can
apply he wo k low by subs i u ing egion-speci ic da ase s, na ional o munici-
pal plan ing in en o ies, and pes -hos eco ds ele an o local pes h ea s. Fo
example, in a Medi e anean-clima e ci y such as Ba celona, S ep 3 (Selec inclu-
sion c i e ia) could inco po a e d ough ole ance alongside ee in asion s a us.
In empe a e o es es o a ion p ojec s in No h Ame ica, S ep 4 (Colla e da a
sou ces) migh p io i ise pes -hos suscep ibili y ela ed o Asian Long-ho ned bee-
le (Anoplopho a glab ipennis), using Uni ed S a es Depa men o Ag icul u e o
s a e-le el pes moni o ing lis s.
Al hough he p o ocol was applied o he Ci y o Cape Town as a case s udy,
i can be applied in o he egions and plan ing scena ios. I can accommoda e
new pes h ea s, clima e-adap ed species guidelines, o addi ional socio-ecological
il e s. Fo ins ance, in u ban o es y planning in Eu opean ci ies, he p o ocol
could in eg a e na ional in asi e species lis s alongside p o enance isk assess-
men s o clima e- esilien species selec ion. The p o ocol can also be applied o
guide plan ing ini ia i es o o he egions acing simila plan and pes -pa hogen
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NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
in asions. Fo ins ance, a municipali y in sou he n Eu ope expe iencing h ea s
om pes s such as Xylella as idiosa could adap he p o ocol by eplacing he
PSHB-speci ic hos lis s wi h da a on Xylella hos plan s, while using Eu opean
o es and u ban ee in en o ies alongside lis s o EU- egula ed in asi e species.
Simila ly, clima e-adap ed species guidelines o p o enance isk assessmen s could
be inco po a ed as addi ional il e ing c i e ia whe e ele an . This lexibili y en-
su es ha he p o ocol p o ides a s uc u ed ye adap able ool o suppo ing e -
idence-based ee selec ion in di e se se ings, helping s akeholde s align plan ing
ini ia i es wi h local ecological, egula o y, and biosecu i y p io i ies.
To u he e ine sui abili y o plan ing, posi i e selec ion c i e ia can be in-
co po a ed in he inal p io i isa ion o sa e ee candida es. These can include
ai s associa ed wi h clima e esilience (e.g. hea and d ough ole ance), ecosys-
em se ice p o ision (e.g. canopy co e , ca bon s o age, pollina o suppo ), and
low main enance equi emen s. These c i e ia can be applied o he il e ed species
pool ( hose no lagged by any exclusion c i e ia) o ensu e ha ecommended axa
a e no only low isk bu also ecologically and unc ionally aluable. The esul ing
candida e lis can o e a p ac ical esou ce o municipali ies seeking o balance
biodi e si y, esilience, and isk in ee plan ing ini ia i es.
Limi a ions
Compiling and cu a ing comp ehensi e species lis s can be esou ce-in ensi e,
pa icula ly in biodi e se, da a-poo , o unde - esou ced egions. In a eas wi h
high na i e and non-na i e species ichness, such as opical o Medi e anean-cli-
ma e egions, he ini ial ask o iden i ying all egionally ele an ee axa can be
ime-consuming and dependen on agmen ed o inconsis en da a sou ces. While
global and na ional biodi e si y da abases (e.g., GBIF, GlobalT eeSea ch, iNa u-
alis ) p o ide aluable s a ing poin s, hese eposi o ies o en lack comple eness,
s anda disa ion, o local e i ica ion. Fu he mo e, in many low- and middle-in-
come coun ies, ins i u ional capaci y o ee moni o ing, axonomic e i ica ion,
and pes /pa hogen su eillance is limi ed. This can hinde he ou ine upda ing and
e inemen o species lis s and educe he esponsi eness o he p o ocol o eme ging
h ea s. Add essing hese limi a ions will equi e sus ained coo dina ion be ween
p ac i ione s, esea ch ins i u ions, he ba ia and ci il socie y, as well as in es men
in open-access da a in as uc u e, egional axonomic expe ise, and aining. The
de elopmen o collabo a i e pla o ms o lis cu a ion, ancho ed in na ional o
municipal en i onmen al planning amewo ks, would help imp o e he easibili y,
anspa ency, and long- e m u ili y o he p o ocol ac oss di e se con ex s.
Richa dson and Po gie e ’s (2024) li ing ee in en o y p o ides a comp ehen-
si e lis o ee axa cu en ly plan ed in Sou h A ica bu does no necessa ily ac-
coun o all ee axa plan ed in he coun y. The in en o y also does no accoun
o ee axa ha ha e no ye been ialled o plan ing. This p esen s a knowledge
gap ha could exclude po en ially low- isk, sui able species om decision-making
simply due o a lack o widesp ead es ing.
While ou lis o ecommended ee axa p o ides a aluable guideline o selec -
ing axa wi h a lowe likelihood o in asion o PSHB suscep ibili y, i s p ac ical im-
plemen a ion is cons ained by he a ailabili y o hese axa in comme cial nu se ies.
Many nu se ies p io i ise species based on ma ke demand, aes he ic appeal, and
g ow h cha ac e is ics a he han ecological sui abili y, which can limi access o
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Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
ce ain ecommended species. To b idge his gap, collabo a ion be ween esea che s,
policymake s, and he ho icul u al indus y is essen ial o encou age he p opaga-
ion and dis ibu ion o hese ee axa. The lis can also se e no only as a deci-
sion-suppo ool o planne s bu also as a p io i y lis o nu se ies. By ocusing
p opaga ion e o s on axa iden i ied as low- isk, nu se ies can help inc ease he
a ailabili y o sa e species, suppo di e si ica ion o u ban and es o a ion plan ings,
and educe he likelihood o inad e en ly in oducing in asi e o pes -p one spe-
cies. In his way, he lis can guide bo h sho - e m plan ing choices and longe - e m
changes in nu se y p oduc ion p ac ices owa d mo e ecologically sound op ions.
While his p o ocol ocuses on species-le el isk sc eening, b oade conside a ions
such as p o enance selec ion, seed sou cing, and phy osani a y p ac ices a e also c i ical
o esponsible ee-plan ing. Economic and logis ical ba ie s, such as limi ed na i e
ee p opaga ion in as uc u e and eliance on impo ed plan ing ma e ial, can con-
s ain implemen a ion. These ac o s should be add essed h ough he in eg a ion o
complemen a y amewo ks, including p o enance isk assessmen s and clima e-adap -
ed species guidelines, wi hin b oade u ban and o es planning p ocesses.
I is also impo an o ecognise he dynamic na u e o PSHB hos suscep ibili y
and he ongoing e olu ion o sui able hos lis s. As PSHB is a ela i ely ecen
in ade in egions like Sou h A ica and Aus alia, no all ee axa wi hin hese
in aded anges ha e had equal exposu e o he bee le and he ungus N. euwalla-
ceae, sugges ing a “hos deb ”. The cu en li ing ee in en o y in Sou h A ica
and exis ing hos lis s a e based on in es a ions obse ed since he bee le’s a i al.
Consequen ly, ee axa ha a e no ye widely plan ed o ha occu in a eas no
ye signi ican ly in aded migh no ha e been adequa ely e alua ed. As PSHB con-
inues o sp ead, i is likely o encoun e and colonise new ee axa, leading o
u he expansion o he known hos ange. Ou p esen unde s anding o PSHB
hos s hus ep esen s a empo al snapsho ; his will change as he in asion un olds.
Addi ional in o ma ion
Con lic o in e es
JW was pa o he edi o ial eam o he special issue 'De eloping lis s o alien axa in he Global
Sou h: wo k lows, p o ocols, p ocesses, and expe iences', bu ook no pa in he pee e iew and
decision-making p ocess o his pape .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
This s udy was suppo ed by he Cen e o In asion Biology and he Depa men o Bo any and
Zoology, S ellenbosch Uni e si y. LJP and MWC ecei ed suppo om he Na u al Sciences and
Enginee ing Resea ch Council o Canada (g an 386151). JRUW and SK hank he Sou h A ican
Depa men o Fo es y, Fishe ies and he En i onmen (DFFE) o unding, no ing ha his publica-
ion does no necessa ily ep esen he iews o opinions o DFFE o i s employees. FR acknowledges
inancial suppo om Ve gelegen and Lou ens o d Wine Es a es, and Ho g o Science (V-19-USE-
PM06). TP acknowledges suppo om he Uni e si y o P e o ia.
132
NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Au ho con ibu ions
All au ho s con ibu ed o he s udy concep ion and design. Ma e ial p epa a ion, da a collec-
ion and analysis we e pe o med by LJP. The i s d a o he manusc ip was w i en by LJP, and
all au ho s commen ed on p e ious e sions o he manusc ip . All au ho s ead and app o ed
he inal manusc ip .
Au ho ORCIDs
Luke J. Po gie e h ps://o cid.o g/0000-0001-7790-2721
Ma c W. Cado e h ps://o cid.o g/0000-0002-5816-7693
T udy Paap h ps://o cid.o g/0000-0003-1364-4350
F ancois Roe s h ps://o cid.o g/0000-0003-3849-9057
John R.U. Wilson h ps://o cid.o g/0000-0003-0174-3239
Da id M. Richa dson h ps://o cid.o g/0000-0001-9574-8297
Da a a ailabili y
The inal il e ed da ase gene a ed and analysed du ing he s udy has been a chi ed on Zenodo:
h ps://doi.o g/10.5281/zenodo.15259591 (Po gie e 2025).
Re e ences
A onson MF, Handel SN, Cleman s SE (2007) F ui ype, li e o m and o igin de e mine he success
o woody plan in ade s in an u ban landscape. Biological In asions 9: 465–475. h ps://doi.
o g/10.1007/s10530-006-9053-1
Baa d JA, K aaij T (2014) Alien lo a o he Ga den Rou e Na ional Pa k, Sou h A ica. Sou h A i-
can Jou nal o Bo any 94: 51–63. h ps://doi.o g/10.1016/j.sajb.2014.05.010
Bas in JF, Finegold Y, Ga cia C, Mollicone D, Rezende M, Rou h D, Zohne CM, C ow he TW (2019)
The global ee es o a ion po en ial. Science 365: 76–79. h ps://doi.o g/10.1126/science.aax0848
Bindewald A, B undu G, Schuele S, S a inge U, Bauhus J, Lapin K (2021) Si e‐speci ic isk as-
sessmen enables ade‐o analysis o non‐na i e ee species in Eu opean o es s. Ecology and
E olu ion 11: 18089–18110. h ps://doi.o g/10.1002/ece3.8407
Blancha d R, Kumschick S, Richa dson DM (2017) Bio uel plan s as po en ial in asi e species:
En i onmen al conce ns and p og ess owa ds objec i e isk assessmen . In: Ruppel O, Dix H
(Eds) Roadmap o Sus ainable Bio uels in Sou he n A ica: Regula o y F amewo ks o Im-
p o ed De elopmen ? Nomos Ve lagsgesellscha , Baden-Baden, Ge many, 47–59. h ps://doi.
o g/10.5771/9783845277585-47
Bond WJ, S e ens N, Midgley GF, Lehmann CE (2019) The ouble wi h ees: A o es a ion plans o
A ica. T ends in Ecology & E olu ion 34: 963–935. h ps://doi.o g/10.1016/j. ee.2019.08.003
B ancalion PHS, Holl KD (2020) Guidance o success ul ee plan ing ini ia i es. Jou nal o Ap-
plied Ecology 57: 2349–2361. h ps://doi.o g/10.1111/1365-2664.13725
B undu G, Richa dson DM (2016) Plan ed o es s and in asi e alien ees in Eu ope: A code o
managing exis ing and u u e plan ings o mi iga e he isk o nega i e impac s om in asions.
NeoBio a 30: 5–47. h ps://doi.o g/10.3897/neobio a.30.7015
B undu G, Paucha d A, Pyšek P, Pe gl J, Bindewald AM, B uno i A, Cana an S, Campagna o T, Ce-
les i-G apow L, Dechoum MDS, Du ou -D o JM (2020) Global guidelines o he sus ainable
use o non-na i e ees o p e en ee in asions and mi iga e hei nega i e impac s. NeoBio a 61:
65–116. h ps://doi.o g/10.3897/neobio a.61.58380
Ca negie AJ, G an A (2025) Social licence o e adica e in asi e pes s o ees in u ban landscapes:
A e iew o p epa e o he nex incu sion. U ban Fo es y & U ban G eening 106: 128722.
h ps://doi.o g/10.1016/j.u ug.2025.128722
133
NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Ci y o Cape Town (2021) U ban Fo es Policy. h ps://www.cape own.go .za/councilonline/_layou s/
OpenDocumen /OpenDocumen .aspx?Documen Id=a466936a-2 99-4833-a6 a-38de3889d9ad
de Wi M, C ooks D, Blignau J, De Bee W, Paap T, Roe s F, an de Me we C, an Wilgen BW,
Richa dson DM (2022) An Assessmen o he Po en ial Economic Impac s o he In asi e Po-
lyphagous Sho Hole Bo e (Coleop e a: Cu culionidae) in Sou h A ica. Jou nal o Economic
En omology 115: 1076–1086. h ps://doi.o g/10.1093/jee/ oac061
Donaldson JE, Hui C, Richa dson DM, Wilson JRU, Robe son MP, Webbe BL (2014) In asion
ajec o y o alien ees: The ole o in oduc ion pa hway and plan ing his o y. Global Change
Biology 20: 1527–1537. h ps://doi.o g/10.1111/gcb.12486
DPIRD (2024) Polyphagous sho -hole bo e (PSHB) Aus alian Hos Lis . Ve sion 22.0, 30 June
2024. Depa men o P ima y Indus ies and Regional De elopmen , Wes e n Aus alia. h ps://
www.ag ic.wa.go .au/si es/ga eway/ iles/PSHB-Global-Hos -Lis .pd
F eeman S, Sha on M, Do i-Bachash M, Maymon M, Belauso E, Maoz Y, Ma gali O, P o aso
A, Mendel Z (2016) Symbio ic associa ion o h ee ungal species h oughou he li e cycle o
he amb osia bee le Euwallacea n . o nica us. Symbiosis 68: 115–128. h ps://doi.o g/10.1007/
s13199-015-0356-9
Global Co enan o Mayo s o Clima e & Ene gy (GCoM) (2022) 2022 Impac Repo : Powe ing
collec i e clima e ac ion. h ps://www.globalco enan o mayo s.o g
Go ae s R, Nic Lughadha E, Black N, Tu ne R, Pa on A (2024) The Wo ld Checklis o Vascula
Plan s, a con inuously upda ed esou ce o explo ing global plan di e si y. Scien i ic Da a 8:
215. h ps://doi.o g/10.1038/s41597-021-00997-6
Holl KD, B ancalion PHS (2020) T ee plan ing is no a simple solu ion. Science 368: 580–581.
h ps://doi.o g/10.1126/science.aba8232
Hulme PE (2009) T ade, anspo and ouble: Managing in asi e species pa hways in an e a
o globaliza ion. Jou nal o Applied Ecology 46: 10–18. h ps://doi.o g/10.1111/j.1365-
2664.2008.01600.x
Kinlock NL, Dehnen‐Schmu z K, Essl F, Pe gl J, Pyšek P, K e H, Weigel P, Yang Q, an Kleunen
M (2022) In oduc ion his o y media es na u aliza ion and in asi eness o cul i a ed plan s.
Global Ecology and Biogeog aphy 31: 1104–1119. h ps://doi.o g/10.1111/geb.13486
Klimešo á J, Ma ínko á J (2022) Clonal g ow h, esp ou ing, and ege a i e p opaga ion o weeds.
In: Upadhyaya M, Clemen s DR, Sh es ha A (Eds) Pe sis ence s a egies o weeds, 200–218.
h ps://doi.o g/10.1002/9781119525622.ch11
Kumschick S, Wilson JR, Foxc o LC (2020) A amewo k o suppo alien species egula ion: The Risk
Analysis o Alien Taxa (RAAT). NeoBio a 62: 213–239. h ps://doi.o g/10.3897/neobio a.62.51031
Kumschick S, Fe nandez Winze L, McCulloch-Jones EJ, Che y D, F ied J, Go ende T, Po gie e
LJ, Rape soa MC, Richa dson DM, an Velden J, Van de Col D, Miza S, Wilson JRU (2024)
Conside a ions o de eloping and implemen ing a sa e lis o alien axa. Bioscience 74: 97–108.
h ps://doi.o g/10.1093/biosci/biad118
Liebhold AM, B ocke ho EG, Ga e LJ, Pa ke JL, B i on KO (2012) Li e plan impo s: The
majo pa hway o o es insec and pa hogen in asions o he US. F on ie s in Ecology and he
En i onmen 10: 135–143. h ps://doi.o g/10.1890/110198
Lockwood JL, Cassey P, Blackbu n TM (2005) The ole o p opagule p essu e in explaining species in-
asions. T ends in Ecology & E olu ion 20: 223–228. h ps://doi.o g/10.1016/j. ee.2005.02.004
Lomba d L, Sando al-Denis M, Cai L, C ous PW (2019) Changing he game: Resol ing sys ema ic
issues in key Fusa ium species complexes. Pe soonia-Molecula Phylogeny and E olu ion o Fungi
43: i–ii. h ps://doi.o g/10.3767/pe soonia.2019.43.00
Lynch SC, Reyes-Gonzalez E, Bossa d EL, Ala con KS, Lo e NL, Hollande AD, Nobua-Beh mann BE,
Gilbe GS (2025) A phylogene ic epidemiology app oach o p edic ing he es ablishmen o mul i-
hos plan pes s. Communica ions Biology 8: 117. h ps://doi.o g/10.1038/s42003-025-07540-y
134
NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Made S (2020) Plan ees o he plane : The po en ial o o es s o clima e change mi iga ion and
he majo d i e s o na ional o es a ea. Mi iga ion and Adap a ion S a egies o Global Change
25: 519–536. h ps://doi.o g/10.1007/s11027-019-09875-4
Mo an R, Allison T, Bell B, B own E, Dews M, Fos e S, Kidane L, Olukani Z, Phillips T, Reyes-Men-
dez L, Roy N, Scho ne L, S ancell K, Wa son J, W igh T (2020) U ban o es and he ee can-
opy: A pa hway o clima e esilience. The In e na ional Unde g adua e Jou nal o Se ice-Lea n-
ing, Leade ship, and Social Change 10: 10–25.
Moyano J, Dima co RD, Pa i sis J, Pe e son T, Pel ze DA, C aw o d KM, McCa y MA, Da is KT,
Paucha d A, Nunez MA (2024) Unin ended consequences o plan ing na i e and non-na i e
ees in eeless ecosys ems o mi iga e clima e change. Jou nal o Ecology 112(2): 480–2491.
h ps://doi.o g/10.1111/1365-2745.14300
Nelu ule T, Robe son MP, Wilson JR, Faulkne KT (2022) Na i e-alien popula ions-An appa -
en oxymo on ha equi es speci ic conse a ion a en ion. NeoBio a 74: 57–74. h ps://doi.
o g/10.3897/neobio a.74.81671
Nunez‐Mi GC, Guo Q, Rejmánek M, Iannone BV III, Fei S (2019) P edic ing in asi eness o exo -
ic woody species using a ai s‐based amewo k. Ecology 100: e02797. h ps://doi.o g/10.1002/
ecy.2797
Paap T, Bu gess TI, Wing ield MJ (2017) U ban ees: B idge-heads o o es pes in asions and sen-
inels o ea ly de ec ion. Biological In asions 19: 3515–3526. h ps://doi.o g/10.1007/s10530-
017-1595-x
Paap T, De Bee ZW, Miglio ini D, Nel WJ, Wing ield MJ (2018) The polyphagous sho hole bo e
(PSHB) and i s ungal symbion Fusa ium euwallaceae: A new in asion in Sou h A ica. Aus al-
asian Plan Pa hology 47: 231–237. h ps://doi.o g/10.1007/s13313-018-0545-0
Paap T, Wing ield MJ, De Bee ZW, Roe s F (2020) Lessons om a majo pes in asion: The polyph-
agous sho hole bo e in Sou h A ica. Sou h A ican Jou nal o Science 116: 8757. h ps://doi.
o g/10.17159/sajs.2020/8757
Paap T, Wing ield MJ, Bu gess TI, Wilson JRU, Richa dson DM, San ini A (2022) In asion
amewo ks: A o es pa hogen pe spec i e. Cu en Fo es y Repo s 8: 74–89. h ps://doi.
o g/10.1007/s40725-021-00157-4
Paque e A, Sousa-Sil a R, Mau e F, Came on E, Belluau M, Messie C (2021) P aise o di e si y:
A unc ional app oach o educe isks in u ban o es s. U ban Fo es y & U ban G eening 62:
127157. h ps://doi.o g/10.1016/j.u ug.2021.127157
Pe i pie e B, Kue e C, B oennimann O, Randin C, Daehle C, Guisan A (2012) Clima ic niche
shi s a e a e among e es ial plan in ade s. Science 335(6074): 1344–1348. h ps://doi.
o g/10.1126/science.1215933
Po gie e L (2025) T ee plan ing guidance o Sou h A ica [Da a se ]. Zenodo. h ps://doi.
o g/10.5281/zenodo.17473434
Po gie e LJ, Douwes E, Gae ne M, Measey J, Paap T, Richa dson DM (2022) Biological in asions
in Sou h A ica’s u ban ecosys ems: pa e ns, p ocesses, impac s and managemen . In: an Wilgen
BW, Measey J, Richa dson DM, Wilson JR, Zengeya TA (Eds) Biological In asions in Sou h
A ica, Sp inge Open, Cham, 275–309. h ps://doi.o g/10.1007/978-3-030-32394-3_11
Po gie e LJ, Cado e MW, Roe s F, Richa dson DM (2024) Moni o ing u ban biological in asions
using ci izen science: The polyphagous sho hole bo e (Euwallacea o nica us). Jou nal o Pes
Science 97: 2073–2085. h ps://doi.o g/10.1007/s10340-024-01744-7
Poyn on RJ (1984) Cha ac e is ics and uses o selec ed ees and sh ubs cul i a ed in Sou h A ica.
Di ec o a e o Fo es y, P e o ia.
Pyšek P, Kři ánek M, Ja ošík V (2009) Plan ing in ensi y, esidence ime, and species ai s de e mine
in asion success o alien woody species. Ecology, 2734–2744. h ps://doi.o g/10.1890/08-0857.1
135
NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Ra a KF, B ocke ho EG, G égoi e J-C, Hamelin RC, Liebhold AM, San ini A, Vene e RC, Wing-
ield MJ (2023) App oaches o o ecas ing damage by in asi e o es insec s and pa hogens: A
c oss-assessmen . Bioscience 73: 85–111. h ps://doi.o g/10.1093/biosci/biac108
Raum S, Collins CM, U quha J, Po e C, Paulei S, Ege e M (2023) T ee insec pes s and pa ho-
gens: A global sys ema ic e iew o hei impac s in u ban a eas. U ban Ecosys ems 26: 587–604.
h ps://doi.o g/10.1007/s11252-022-01317-5
Rejmánek M, Richa dson DM (2013) T ees and sh ubs as in asi e alien species - 2013 upda e o he
global da abase. Di e si y and dis ibu ions 19: 1093–1094. h ps://doi.o g/10.1111/ddi.12075
Richa dson DM, Po gie e LJ (2024) A li ing in en o y o plan ed ees in Sou h A ica de i ed om iNa -
u alis . Sou h A ican Jou nal o Bo any 173: 365–379. h ps://doi.o g/10.1016/j.sajb.2024.08.006
Richa dson DM, Pyšek P (2012) Na u aliza ion o in oduced plan s: Ecological d i e s o bio-
geog aphical pa e ns. The New Phy ologis 196: 383–396. h ps://doi.o g/10.1111/j.1469-
8137.2012.04292.x
Richa dson DM, Rejmánek M (2011) T ees and sh ubs as in asi e alien species-a global e iew.
Di e si y & Dis ibu ions 17: 788–809. h ps://doi.o g/10.1111/j.1472-4642.2011.00782.x
Richa dson DM, Thuille W (2007) Home away om home-objec i e mapping o high‐ isk sou ce
a eas o plan in oduc ions. Di e si y & Dis ibu ions 13: 299–312. h ps://doi.o g/10.1111/
j.1472-4642.2007.00337.x
Richa dson DM, Camb ay JA, Chapman RA, Dean WRJ, G i i hs CL, Le Mai e DC, New on DJ,
Wins anley TJ (2003) Vec o s and pa hways o biological in asions in Sou h A ica - Pas , p esen
and u u e. In: Ruiz GM, Ca l on JT (Eds) In asi e Species: Vec o s and Managemen S a egies.
Island P ess, Washing on, D.C., 292–349.
Richa dson DM, Le Roux JJ, Ma chan e EM (Eds) (2023) Wa les: Aus alian Acacia species a ound
he wo ld. CABI, Walling o d. h ps://doi.o g/10.1079/9781800622197.0000
Rouge M, Robe son MP, Wilson JRU, Hui C, Essl F, Ren e ia JL, Richa dson DM (2016) In asion
deb – Quan i ying u u e biological in asions. Di e si y and Dis ibu ions 22: 445–456. h ps://
onlinelib a y.wiley.com/doi/10.1111/ddi.12408
Roy S, By ne J, Picke ing C (2012) A sys ema ic quan i a i e e iew o u ban ee bene i s, cos s, and
assessmen me hods ac oss ci ies in di e en clima ic zones. U ban Fo es y & U ban G eening
11: 351–363. h ps://doi.o g/10.1016/j.u ug.2012.06.006
Seddon N, Chausson A, Be y P, Gi a din CAJ, Smi h A, Tu ne B (2020) Unde s anding he alue
and limi s o na u e-based solu ions o clima e change and o he global challenges. Philosoph-
ical T ansac ions o he Royal Socie y o London. Se ies B, Biological Sciences 375: 20190120.
h ps://doi.o g/10.1098/ s b.2019.0120
Shackle on CM, Gwedla N (2021) The legacy e ec s o colonial and Apa heid imp in s on u ban
g eening in Sou h A ica: Spaces, species, and sui abili y. F on ie s in Ecology and E olu ion 8:
579813. h ps://doi.o g/10.3389/ e o.2020.579813
Sousa-Sil a R, Du los M, Ba ona CO, Paque e A (2023) Keys o be e planning and in eg a -
ing u ban ee plan ing ini ia i es. Landscape and U ban Planning 231: 104649. h ps://doi.
o g/10.1016/j.landu bplan.2022.104649
S ou hame R, Rugman-Jones P, Thu PQ, Eskalen A, Thibaul T, Hulc J, Wang L-J, Jo dal BH,
Chen C-Y, Coope band M, Lin C-S, Kama a N, Lu S-S, Masuya H, Mendel Z, Rabaglia R,
Sanguansub S, Shih H-H, Si ichaya W, Zong S (2017) T acing he o igin o a c yp ic in ad-
e : phylogeog aphy o he Euwallacea o nica us (Coleop e a: Cu culionidae: Scoly inae) species
complex. Ag icul u al and Fo es En omology 19: 366–375. h ps://doi.o g/10.1111/a e.12215
Thuille W, Richa dson DM, Pyšek P, Midgley GF, Hughes GO, Rouge M (2005) Niche-based
modelling as a ool o p edic ing he isk o alien plan in asions a a global scale. Global Change
Biology 11(12): 2234–2250. h ps://doi.o g/10.1111/j.1365-2486.2005.001018.x
136
NeoBio a 104: 113–137 (2025), DOI: 10.3897/neobio a.104.156206
Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
Townsend G, Hill M, Hu ley BP, Roe s F (2025) Escala ing h ea : Inc easing impac o he polyph-
agous sho hole bo e bee le, Euwallacea o nica us, in nea ly all majo Sou h A ican o es ypes.
Biological In asions 27: 88. h ps://doi.o g/10.1007/s10530-025-03551-2
Uni ed Na ions (2015) T ans o ming ou wo ld: The 2030 Agenda o Sus ainable De elopmen .
Uni ed Na ions Gene al Assembly. h ps://sdgs.un.o g/2030agenda
Uni ed Na ions (2017) New U ban Agenda. Uni ed Na ions Human Se lemen s P og amme
(UN-Habi a ). h ps://habi a 3.o g/ he-new-u ban-agenda
an Kleunen M, Webe E, Fische M (2010) A me a‐analysis o ai di e ences be ween in asi e
and non‐in asi e plan species. Ecology Le e s 13: 235–245. h ps://doi.o g/10.1111/j.1461-
0248.2009.01418.x
an Kleunen M, Pyšek P, Dawson W, Essl F, K e H, Pe gl J, Weigel P, S ein A, Dullinge S, König
C, Lenzne B, Mau el N, Mose D, Seebens H, Ka esz J, Nishino M, Aleksanyan A, Ansong M,
An ono a LA, Ba celona JF, B eckle SW, B undu G, Cabezas FJ, Cá denas D, Cá denas-To o J,
Cas año N, Chacón E, Cha elain C, Conn B, de Sá Dechoum M, Du ou -D o JM, Ebel AL,
Figuei edo E, F agman-Sapi O, Fuen es N, G oom QJ, Hende son L, Inde ji , Jogan N, K es o
P, Kup iyano A, Masciad i S, Mee man J, Mo ozo a O, Nick en D, Nowak A, Pa zel A, Pelse
PB, Shu WS, Thomas J, Uludag A, Velayos M, Ve khosina A, Villaseño JL, Webe E, Wie inga
JJ, Yazlık A, Zeddam A, Zyko a E, Win e M (2019) The Global Na u alised Alien Flo a (GloN-
AF) da abase. Ecology 100: 1–2. h ps://doi.o g/10.1002/ecy.2542
an Rooyen E, Paap T, De Bee W, Townsend G, Fell S, Nel WJ, Mo gan S, Hill M, Gonzalez A, Ro-
e s F (2021) The polyphagous sho hole bo e bee le: Cu en s a us o a pe ec in ade in Sou h
A ica. Sou h A ican Jou nal o Science 117: 11–12. h ps://doi.o g/10.17159/sajs.2021/9736
an S aden EM, S o be g GH (2021) The G eening Sowe o ee-plan ing p ojec in Sou h A i-
ca–Elimina ing he “g een di ide” legacy o apa heid. U ban Fo es y & U ban G eening 65:
127371. h ps://doi.o g/10.1016/j.u ug.2021.127371
an Wilgen BW, Measey J, Richa dson DM, Wilson JR, Zengeya TA (Eds) (2020) Biological In-
asions in Sou h A ica. Sp inge Open, Cham, 975 pp. h ps://doi.o g/10.1007/978-3-030-
32394-3
an Wilgen BW, Zengeya TA, Richa dson DM (2022) A e iew o he impac s o biological in asions
in Sou h A ica. Biological In asions 24: 27–50. h ps://doi.o g/10.1007/s10530-021-02623-3
Vashis M, Singh SK, Kuma TV (2025) Enhancing esilience o sus ainable ci ies: A e iew o
h ea s o u ban ees. Biodi e si y and Conse a ion 34: 1231–1258. h ps://doi.o g/10.1007/
s10531-025-03025-w
Wilson JR (2024) A lis o axa cu en ly and his o ically egula ed unde Sou h A ica’s Na ion-
al En i onmen al Managemen : Biodi e si y Ac , Alien & In asi e Species Regula ions. 1.0
(20240331). h ps://doi.o g/10.5281/zenodo.10809766
Wilson JRU, Kumschick S (2024) The egula ion o alien species in Sou h A ica. Sou h A ican
Jou nal o Science 120: 1–14. h ps://doi.o g/10.17159/sajs.2024/17002
Wilson JRU, Richa dson DM, Rouge M, P ocheş Ş, Amis MA, Hende son L, Thuille W (2007)
Residence ime and po en ial ange: C ucial conside a ions in modelling plan in asions. Di e si-
y & Dis ibu ions 13: 11–22. h ps://doi.o g/10.1111/j.1366-9516.2006.00302.x
Wilson JR, Measey J, Richa dson DM, an Wilgen BW, Zengeya TA (2020) Po en ial u u es o
biological in asions in Sou h A ica. In: an Wilgen BW, Measey J, Richa dson DM, Wilson
JR, Zengeya TA (Eds) Biological In asions in Sou h A ica. Sp inge Open, Cham, 917–946.
h ps://doi.o g/10.1007/978-3-030-32394-3_31
Yi ba ek TW, Wilson JRU, Dehnen-Schmu z K (2023) A go e nance amewo k o he design
and e alua ion o ee plan ing schemes. Fo es Policy and Economics 152: 102980. h ps://doi.
o g/10.1016/j. o pol.2023.102980
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Luke J. Po gie e e al.: P o ocol o de eloping lis s o ee plan ing
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Addi ional si e-speci ic c i e ia o conside when selec ing ee axa o plan ing
Au ho s: Luke J. Po gie e , Ma c W. Cado e, Sab ina Kumschick, T udy Paap, F ancois Roe s, John
R.U. Wilson, Da id M. Richa dson
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