139
Di e si y and dis ibu ion pa e ns o in asi e alien plan species
in mainland Po ugal
Raquel Fe nandes1, And y Cas o1, Hélia Ma chan e2,3 , Elizabe e Ma chan e3,4 , Césa Capinha1,3
1 Cen e o Geog aphical S udies, Ins i u e o Geog aphy and Spa ial Planning, Uni e si y o Lisbon, Rua B anca Edmée Ma ques, 1600-276 Lisboa, Po ugal
2 Resea ch Cen e o Na u al Resou ces, En i onmen and Socie y (CERNAS), Poly echnic Uni e si y o Coimb a, Coimb a Ag icul u e School, Bencan a, 3045-
601 Coimb a, Po ugal
3 Associa e Labo a o y Te a, Lisboa, Po ugal
4 Cen e o Func ional Ecology - Science o People & he Plane , Depa men o Li e Sciences, Uni e si y o Coimb a, Calçada Ma im de F ei as, 3000-456
Coimb a, Po ugal
Co esponding au ho : Raquel Fe nandes ( [email p o ec ed])
Copy igh : © Raquel Fe nandes 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
Unde s anding he dis ibu ion and di e si y o in asi e alien species is inc easingly c ucial o mee legal
equi emen s and guide e ec i e managemen . Po ugal cu en ly hos s a la ge numbe o in asi e alien
plan species, wi h signi ican en i onmen al and socio-economic impac s. Howe e , he dis ibu ion o
mos o hese species ac oss he e i o y and he ac o s d i ing hei sp ead and di e si y emain la gely
unexplo ed. To add ess his, we p esen he i s a las o in asi e alien plan s in mainland Po ugal. A o al
o 96 e es ial and aqua ic in asi e alien plan s a e p esen ed, encompassing all species lis ed unde Po -
uguese na ional legisla ion and he Eu opean Union’s lis o in asi e species o conce n. Occu ence da a
we e collec ed om a b oad a ay o sou ces, including na ional and in e na ional biodi e si y obse a ion
da abases, ci izen science da a, li e a u e, and da a collec ions owned by manage s and esea che s— o al-
ing app oxima ely 85,000 eco ds, wi h a mean o 879 eco ds pe species ( ange: 1–9,190). Each species
was cha ac e ized based on mul iple dis ibu ional pa ame e s, and a k-means analysis was used o g oup
species wi h simila dis ibu ion pa e ns. The ichness o in asi e plan s was mapped a he municipali y
le el. The d i e s o hei spa ial a ia ion we e in es iga ed using a comp ehensi e se o 30 a iables
ep esen ing mul iple en i onmen al and human ac o s. Resul s show ha in asi e plan s a e p esen in
all mainland Po uguese municipali ies bu wi h high a iabili y. Fou main pa e ns o dis ibu ion we e
iden i ied: species p ima ily loca ed along he coas (e.g., Acacia saligna) and a g adien o species wi h
na ow (e.g., Reynou ia japonica), mode a e (e.g., Ipomoea indica), and wide dis ibu ion anges (e.g.,
Co ade ia selloana). In asi e species ichness was signi ican ly highe in coas al and la ge municipali ies,
pa icula ly hose close o majo u ban cen e s and wi h a highe densi y o powe lines. Ou esul s p o-
ide he i s comp ehensi e assessmen o he dis ibu ion o in asi e alien plan s in mainland Po ugal,
es ablishing a much-needed baseline o u u e in asion p e en ion and managemen e o s.
Key wo ds: Baseline dis ibu ion, d i e s o in asion, in asion a las, in asi e alien plan s, in asi e
species ichness, occupancy
In oduc ion
Human ac i i ies a e acili a ing he global mo emen and in oduc ion o species be-
yond hei na i e anges (Pyšek e al. 2020; Lopez e al. 2022; Capinha e al. 2023).
Cu en ly, o e 37,000 es ablished alien species ha e been documen ed wo ldwide
Academic edi o : Inde ji
Recei ed:
26 June 2025
Accep ed:
21 Oc obe 2025
Published:
19 No embe 2025
Ci a ion: Fe nandes R, Cas o A,
Ma chan e H, Ma chan e E, Capinha
C (2025) Di e si y and dis ibu ion
pa e ns o in asi e alien plan
species in mainland Po ugal.
NeoBio a 104: 139–162. h ps://doi.
o g/10.3897/neobio a.104.163291
NeoBio a 104: 139–162 (2025)
DOI: 10.3897/neobio a.104.163291
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: 139–162 (2025), DOI: 10.3897/neobio a.104.163291
Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
(IPBES 2023), wi h app oxima ely 200 new alien species (i.e., “exo ic” o “non-na i e”)
eco ded annually. Among hese, alien plan s cons i u e a signi ican p opo ion, wi h
mo e han 13,000 species es ablished ou side hei na u al anges (Seebens e al. 2023).
A mino po ion o alien plan s become in asi e, causing signi ican and di e se
nega i e impac s in egions o in oduc ion. These impac s include educ ions in
na i e species di e si y and abundance (Powell e al. 2011; Hansen e al. 2020;
Beau y e al. 2023), dis up ion o key ecosys em se ices (Viei es-Blanco and
González-P ie o 2020; Fe ei a e al. 2021; Ren e al. 2021; Galla do e al. 2024),
and majo economic losses a ec ing mul iple ac i i ies, including he loss o c ops
and p oduc i e land (Richa dson and Van Wilgen 2004; Le Mai e e al. 2011; Mi-
lano ić e al. 2020; Reynolds e al. 2020; Si bu e al. 2022). Addi ionally, se e al
in asi e plan s pose en i onmen al isks o human heal h (Rod iguez e al. 2021).
Mos o hese impac ul species ha e been epo ed in he Ame ican con inen , he
Asia–Paci ic egion, Cen al Asia, and Eu ope (IPBES 2023). In Eu ope, a leas
7,335 alien ascula plan species ha e been eco ded, o which 1,037 (abou 14%)
a e conside ed in asi e in a leas one Eu opean e i o y (Kaluso á e al. 2024).
Gi en he equen se e i y o he obse ed and po en ial impac s, many na ion-
al and egional go e nmen s now p io i ize managing biological in asions, includ-
ing such species in dedica ed legisla ion. Po ugal was one o he i s Eu opean
coun ies o adop legisla ion on in asi e alien species (he ea e in asi e species),
in oducing in 1999 a lis o species subjec o speci ic es ic ions, including bans
on comme cial use and possession (Minis é io do Ambien e 1999). This lis in-
cluded o e 40 plan species, wi h se e al lis ed as in asi e, such as Acacia dealba a,
Ca pob o us edulis, and Oxalis pes-cap ae, and o he s conside ed o ha e known
ecological isk (ei he in he ea ly s ages o in asion o ha ing a high po en ial
o become in asi e in he u u e), such as Ludwigia peploides, R. japonica, and
Senecio inaequidens. This legisla ion and associa ed species lis we e upda ed by
Dec ee-Law No. 92/2019 (Minis é io do Ambien e e T ansição Ene gé ica 2019),
which now includes mo e han 100 plan species lis ed as in asi e o mainland
Po ugal. The upda ed lis includes con i med and po en ial in asi e species known
o occu in he e i o y, as well as o he s ha a e p esumed absen bu expec ed o
become in asi e i in oduced. Addi ionally, he Eu opean Union Regula ion No.
1143/2014 (O icial Jou nal o he Eu opean Union 2014) includes he “Union
lis ,” which speci ically lis s he in asi e species o Union conce n. This egula ion
aims o p e en and mi iga e nega i e en i onmen al impac s on Eu opean Union
coun ies. Species lis ed a e au oma ically pa o he Po uguese na ional lis , be-
ing subjec o Dec ee-Law No. 92/2019. As o he hi d upda e o he Union lis
in 2022, 41 plan species a e lis ed (O icial Jou nal o he Eu opean Union 2022).
O e he yea s, a ele an numbe o s udies ha e in es iga ed plan in asions in
Po ugal (Sousa e al. 2018). Fo example, se e al ha e lis ed he di e si y o alien
plan s (Almeida and F ei as 2000; Almeida and F ei as 2012; Ma chan e e al.
2014), dis ibu ion, o po en ial in asion isk (Rod íguez-Me ino 2023). O he s
ha e analyzed indi idual species such as Baccha is spica a (Ve loo e e al. 2017),
C. edulis (Mal ez-Mou o e al. 2009; Che aoui and Chozas 2019), Co ula co o-
nopi olia (Cos a e al. 2009), o Pon ede ia c assipes (Pádua e al. 2022a, 2022b;
Mou a e al. 2023). S udies ha e also ocused on Acacia species (Ma chan e e al.
2003; Ma ins e al. 2016; Vicen e e al. 2019; Raposo e al. 2021; G oße-S ol en-
be g e al. 2023), a highly p oblema ic g oup o axa due o hei in asi e beha -
io , wi h nine species al eady es ablished in he e i o y (Ma chan e e al. 2023).
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
Speci ic egions and ecosys ems ha e also ecei ed pa icula a en ion, including
p o ec ed a eas (Gu ie es e al. 2011; Dua e e al. 2023), dune habi a s (Dua e e
al. 2023), o es s (Fe ei a e al. 2021), ipa ian sys ems (Be nez e al. 2005; Aguia
and Fe ei a 2013; Pabs e al. 2022), and inland wa e s (O icialdegui e al. 2023).
Despi e he nume ous aluable con ibu ions and he g owing numbe o ci izen
science dis ibu ion eco ds on di e en pla o ms (e.g., he p ojec “In aso as.p ”
a iNa u alis ), he o e all dis ibu ion o in asi e plan s ac oss mainland Po ugal
emains la gely unknown. This gap pe sis s despi e na ional in en o ies o in asi e
species being essen ial ools o moni o ing in asions, guiding policy, and o mula -
ing managemen plans (La ombe e al. 2017; Pabs e al. 2022; Si bu e al. 2022).
The lack o comp ehensi e dis ibu ion da a is likely due, in pa , o in o ma ion
on indi idual axa being dispe sed ac oss nume ous sou ces, making colla ion and
mapping a esou ce-in ensi e and challenging ask. Addi ionally, o many species,
in o ma ion abou dis ibu ion a local scales is incomple e o missing.
He e, we p esen a comp ehensi e a las o he dis ibu ion o in asi e plan
ichness in mainland Po ugal. This a las was c ea ed by compiling and ha mo-
nizing da a om a wide ange o sou ces, including na ional and in e na ional
biodi e si y obse a ion da abases, scien i ic li e a u e, and da a collec ions om
manage s and esea che s. In Po ugal, municipali ies play a cen al ole in e i-
o ial managemen , en i onmen al planning, and decision-making ega ding ield
in e en ions. This is also he case o local- and na ional-le el esou ce alloca ion
o in asi e species p e en ion and managemen , ci izen science ini ia i es, and
public awa eness campaigns. Hence, municipali ies we e selec ed as e e ence sam-
pling uni s. Wi h he in o ma ion collec ed, we aimed o: i) map he eco ded
dis ibu ion o each in asi e alien plan species in he e i o y; ii) iden i y species
sha ing simila dis ibu ion pa e ns; iii) assess spa ial pa e ns o in asi e species
ichness; and i ) e alua e he en i onmen al and socio-economic d i e s o spa ial
a ia ion in in asi e species ichness alues. Ou esul s a e expec ed o p o ide
subs an ial suppo o decision-making in local- o-na ional-scale p e en ion and
managemen e o s o alien plan in asions in Po ugal.
Ma e ials and me hods
S udy a ea
Mainland Po ugal is loca ed in he wes e n Ibe ian Peninsula, a he sou hwes e n
edge o Eu ope (Fig. 1). I s e i o y spans bo h he A lan ic and Medi e anean
biogeog aphical egions (Roekae s 2002). The cen al and he sou he n egions
(Alen ejo and Alga e) a e cha ac e ized by a Medi e anean clima e wi h d y, ho
summe s, while he no he n egions expe ience a empe a e oceanic clima e wi h
d y, mild summe s (Beck e al. 2023). The sou he n and coas al a eas a e gene ally
low-lying, whe eas he no he n and cen al egions a e mo e moun ainous, wi h
se e al peaks exceeding 1,000 m in ele a ion (Ramos and Ramos-Pe ei a 2020).
The e i o y is di ided in o 278 municipali ies. The popula ion is p ima ily con-
cen a ed in coas al municipali ies, pa icula ly in he me opoli an a eas o Lisbon
and Opo o (Ins i u o Nacional de Es a ís ica 2022). Fo es s occupy la ge a eas in
all Nomencla u e o Te i o ial Uni s o S a is ics II (NUTS II) egions, especially
in he Cen e egion, while ag icul u al a eas a e mos ep esen ed in Alen ejo and
he Lisbon Me opoli an A ea (Al es e al. 2022).
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
Lis o in asi e plan s
The mapping e o ocused on he plan species lis ed in Po uguese Dec ee-Law
No. 92/2019 (Minis é io do Ambien e e T ansição Ene gé ica 2019) and Eu opean
Regula ion No. 1143/2014 (O icial Jou nal o he Eu opean Union 2014). Since
ou ocus is on mainland Po ugal, species men ioned in na ional legisla ion ha
pe ain only o he insula egions, such as he Madei a a chipelago, we e excluded.
Po uguese Dec ee-Law No. 92/2019 lis s he genus Acacia collec i ely, unlike o he
axa, which a e lis ed a he species le el. The e o e, we consul ed ele an li e a u e
(Ma chan e e al. 2014, 2023) o iden i y he Acacia species known o occu in he
e i o y. Addi ionally, we included Wa sonia me iana in ou mapping, as he e-
sea ch communi y ecognizes i s in asi e beha io in se e al a eas o mainland Po -
ugal (Ma chan e e al. 2014; Mo ais e al. 2017). Dis ibu ion da a we e collec ed
o a o al o 96 species om 64 di e en gene a (Suppl. ma e ial 1: able S1.1).
Figu e 1. Loca ion o mainland Po ugal showing i s adminis a i e di isions, including NUTS II egions (ou lined in black) and he
278 municipali ies (ou lined in g ey), used as uni s o analysis in his s udy. Popula ion densi y (pe sons/km2) is highes in he Lisbon and
Opo o Me opoli an A eas, ep esen ed by da k ed egions.
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
Occu ence da abase
We ga he ed occu ence eco ds o each in asi e plan species om mul iple sou c-
es. The online da abases consul ed included he Global Biodi e si y In o ma ion
Facili y (GBIF) (GBIF.o g 2024); Flo a-On (Flo a de Po ugal In e ac i a 2024), an
expe -led p ojec aimed a sys ema izing in o ma ion o all ascula plan species
in Po ugal, including hei dis ibu ion; and he ci izen-science pla o ms iNa u al-
is (iNa u alis 2024) (only esea ch-g ade eco ds we e conside ed) and Pl@n Ne
(Pl@n Ne 2024). Reco ds we e also collec ed om nume ous esea ch publica ions
iden i ied in Web o Science and Scopus using he sea ch e ms “Po ugal” and each
species name, as well as om mas e ’s and doc o al heses iden i ied in RENATES,
a Po uguese pla o m con aining o icial in o ma ion on disse a ions p oduced in
Po ugal (DGEEC 2024). Addi ionally, we inco po a ed unpublished da a om
he au ho s, collabo a o s, and companies in ol ed in in asi e plan managemen
ac ions (see Acknowledgmen s sec ion). Da a up o May 2024 we e conside ed.
The axonomy o species was de ined and ha monized acco ding o he GBIF
Backbone Taxonomy (GBIF Sec e a ia 2023). Occu ence eco ds we e collec ed
o all a ailable yea s and included subspecies (i p esen ). Reco ds iden i ied as
ha ing coo dina es o s o ed specimens (e.g., he ba ium loca ions) a he han he
ac ual place o obse a ion in he wild we e emo ed. Addi ionally, eco ds wi h a
spa ial p ecision o less han 100 m we e excluded.
An excep ion was made o da a om Flo a-On. This sou ce is c ucial o he
comp ehensi eness o he mapping bu p o ides da a as cen oids o g id squa es
a a 10 × 10 km esolu ion. Despi e i s con ibu ion o spa ial comp ehensi eness,
he lowe geog aphical p ecision may in oduce some spa ial e o s in a ibu ing
occu ences o municipali ies— he spa ial uni s o mapping and analysis used (see
below). The e o e, we pe o med wo mapping exe cises and associa ed analyses:
one including Flo a-On da a and ano he wi hou i . The esul s om bo h da a-
se s we e highly cong uen (see Resul s sec ion). Thus, we p ima ily desc ibe he
esul s, including he en i e se o occu ence da a, while hose excluding Flo a-On
da a a e p o ided in Suppl. ma e ial 3.
Gi en he b ead h o consul ed sou ces, we expec o ha e ob ained a ep esen-
a i e iden i ica ion o he species occu ing in each municipali y. Howe e , as wi h
any simila exe cise, ou da a may s ill exhibi some gaps and biases, pa icula ly
owa d egions wi h highe eco ding e o s and species ha a e mo e conspicuous
and easie o iden i y (see Discussion sec ion o de ails).
In asi e plan ichness mapping and occupancy
To analyze species dis ibu ion pa e ns, we agg ega ed occu ence eco ds a he
municipali y le el, ollowing he bounda ies de ined by he Ca a Adminis a i a
O icial de Po ugal (CAOP) (Di eção-Ge al do Te i ó io 2024). Fo each mu-
nicipali y, duplica e eco ds o he same species we e emo ed, e aining only one
occu ence pe species. In asi e species ichness was hen calcula ed as he o al
numbe o unique species eco ded pe municipali y. Species occupancy pa e ns
we e assessed by quan i ying, wi hin each municipali y, he numbe o 1 × 1 km
g id cells in which he species was eco ded. The mean pe cen age o occupied g id
cells ac oss municipali ies was subsequen ly calcula ed o ep esen o e all species
occupancy. Analyses we e pe o med in R 4.3.2 (R Co e Team 2022).
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
Assessing pa e ns o species dis ibu ions
To iden i y simila i ies in species-le el pa e ns o dis ibu ion ac oss he e i o y,
we i s cha ac e ized he dis ibu ion o each species using i e a iables: he num-
be o municipali ies whe e occu s; he median and in e qua ile ange o dis ance
o he coas ( om he cen oid o each municipali y, in kilome e s); and he medi-
an and in e qua ile ange o la i ude (in decimal deg ees).
We hen pe o med a clus e analysis based on hese a iables. To accoun o
mul icollinea i y among he a iables, we used p incipal componen analysis
(PCA), which explained 72.3% o he a iance wi h he i s wo componen s
(Suppl. ma e ial 1: ig. S1.1). Nex , we pe o med a k-means clus e ing analysis
based on he sco es o each species in he wo-dimensional PCA space. The PCA
was pe o med using he “p comp” unc ion om he s a s R package. The k-means
p ocedu e employed he “kmeans” unc ion om he same package, and he elbow
me hod (Bholowalia and Kuma 2014) was used o iden i y he op imal numbe
o clus e s (Suppl. ma e ial 1: ig. S1.2).
We also examined whe he he numbe o municipali ies whe e a species
occu s e lec s i s o e all occupancy ac oss he e i o y. To do his, we i ed a
linea model ela ing he mean occupancy o species (i.e., he a e age pe cen -
age o occupied 1 × 1 km g id cells pe municipali y) o he o al numbe o
municipali ies whe e i was eco ded.
Spa ial d i e s o in asi e plan ichness
Explana o y a iables
To iden i y he ac o s d i ing he spa ial a ia ion in he ichness o in asi e plan s
ac oss he e i o y, we selec ed a comp ehensi e se o 30 a iables (Table 1) ep-
esen ing dis inc en i onmen al and human- ela ed ac o s, including p oxies o
eco ding e o . These a iables we e mapped a he municipali y le el and p o ide
a b oad ep esen a ion o he hypo heses on d i ing ac o s conside ed in p e ious
s udies (Vicen e e al. 2010; San os e al. 2011; Vicen e e al. 2019; Viei es-Blanco
and González-P ie o 2020; Pabs e al. 2022).
The a iables ep esen ing clima ic condi ions, opog aphy, con inen ali y,
socioeconomy (g oss domes ic p oduc [GDP] and a el ime o ci ies, i.e., he
ime i akes o a el o he nea es u ban cen e ia su ace anspo ) (Weiss e
al. 2018), and soil g oups consis ed o mean alues o as e cell da a wi hin he
bounda ies o each municipali y. The land-use a iables used ec o - ype base da a
and we e ob ained by di iding he o al a ea occupied by each land-use class by
he municipali y’s o al a ea. Addi ionally, a iables ep esen ing dispe sal co ido s
we e calcula ed by summing he ex en o hese en i ies (hyd og aphic ne wo k,
oad in as uc u e, ailways, and powe lines) wi hin each municipali y and di-
iding by i s a ea. The minimum dis ance o he coas line was de e mined as he
sho es dis ance om he cen oid o each municipali y.
The a ailabili y o species dis ibu ion da a can be s ongly in luenced by
a ying le els o eco ding e o (Tiago e al. 2017). The e o e, we conside ed
a se o a iables expec ed o ep esen spa ial pa e ns in his e o ac oss
he e i o y. These a iables we e he a e age numbe o highe educa ion
s uden s pe municipali y, he numbe o highe educa ion ins i u ions pe
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
municipali y, and he numbe o iNa u alis p ojec s wi h in asi e plan e-
co ds pe municipali y. To a oid edundancy in hei spa ial a ia ion, we p o-
jec ed hese a iables in o a PCA. The i s p incipal componen , explaining
a ound 75% o he a iance, was e ained, ep esen ing mainly he join a ia-
ion o he numbe o highe educa ion s uden s and ins i u ions. The sco es o
his i s composi e we e ex ac ed and used as a new a iable named “academic
sampling e o ” (Table 1). Because he numbe o iNa u alis p ojec s con ib-
u ed li le o his componen , we used he o iginal alues o his a iable as a
p oxy o “ci izen-science sampling e o ”.
Table 1. Explana o y a iables conside ed as p edic o s o spa ial a ia ion pa e ns o in asi e plan ichness. The inal se o a iables (in
bold) was de e mined a e assessing mul icollinea i y.
G oup Va iables T ans o ma ion Sou ce VIF
Clima e Annual p ecipi a ion amoun (kg m-2) Chelsa (1981–2010)
(Ka ge e al. 2017, 2018)
>5
P ecipi a ion amoun o he we es mon h (kg m-2) >5
P ecipi a ion amoun o he d ies mon h (kg m-2) >5
Annual mean empe a u e (°C) >5
Daily mean ai empe a u es o he d ies qua e (°C) >5
Daily mean ai empe a u es o he we es qua e
(°C)
3.44
Daily mean ai empe a u es o he coldes qua e (°C) >5
Mean daily maximum ai empe a u e o he
wa mes mon h (°C)
2.38
Mean daily minimum ai empe a u e o he coldes
mon h (°C)
>5
Topog aphy Slope (deg ees) Ea h Resou ces Obse a ion and Science
(EROS) Cen e (2018)
>5
Ele a ion (me e s) >5
Ruggedness Index 3.06
Geog aphy Minimum dis ance o coas line (km) 2.58
Municipali y a ea (km2)Sq Di eção-Ge al do Te i ó io (2024) 2.55
Land use Densi y o a i icialized e i o ies (%) Log(x+1) COS, Ve sion 2018 (Di eção Ge al do
Te i ó io (DGT), 2020)
>5
Densi y o ag icul u al a eas (%) 2.02
Densi y o o es a eas (%) 2.70
Densi y o p o ec ed a eas (%) Log(x+1) Ins i u o da Conse ação da Na u eza e das
Flo es as (ICNF) (2022, 2023a, 2023b,
2023c, 2023d)
1.89
Bu n a ea (2007–2022) (%) Ins i u o da Conse ação da Na u eza e das
Flo es as (ICNF) (2024)
2.65
Soil A e age soil pH
(15–30 cm laye )
Log Poggio e al. (2021) 2.32
Densi y o
dispe sal co ido s
Densi y o local hyd og aphic ne wo k (km/km2)
(2024)
OpenS ee Map. Geo ab ik GmbH, and
OpenS ee Map Con ibu o s (2018)
2.13
Densi y o oads (km/km2) (2024) Log(x+1) >5
Densi y o ailways (km/km2) (2024) Log(x+1) 1.92
Densi y o powe lines (km/km2) (2024) Log(x+1) REN—Redes Ene gé icas Nacionais, SA (see
acknowledgmen s)
1.26
Socioeconomic Mean popula ion densi y acco ding o he na ional
census (mean alue o 2001, 2011, and 2021; numbe
o pe sons/km2)
Log INE, PORDATA (2024) >5
GDP (1990–2015; 2011 in e na ional US dolla s) Log Kummu e al. (2015) >5
T a el ime o ci ies (minu es) Weiss e al. (2018) 2.97
Sampling e o (academic and ci izen-science
sampling e o )
DGEEC/ME-MCTE, PORDATA (2021,
2024); INE, PORDATA (2024)
3.21
Numbe o iNa u alis p ojec s wi h IAP eco ds Log(x+1) iNa u alis (2024) 2.51
Numbe o plan nu se ies and aqua ium s o es Log(x+1) Nunes e al. (unpublished) 1.69
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
Tes ing o d i e s o in asi e plan ichness
To assess he ela ionship be ween he ichness o in asi e plan s ac oss he e i-
o y (dependen a iable) and he explana o y a iables (Table 1), we used a gen-
e alized leas squa es (GLS) eg ession model (Do mann e al. 2007). P io o
model i ing, squa e oo and loga i hmic ans o ma ions we e applied o se e al
explana o y a iables (Table 1). The esponse a iable was also log- ans o med o
mee model expec a ions o a Gaussian dis ibu ion o e o (I es 2015).
Po en ial mul icollinea i y among he explana o y a iables was assessed using
he a iance in la ion ac o (VIF) (Alin 2010), calcula ed wi h he “ i s ep” unc-
ion om he usdm R package (Naimi 2022). Only he se o explana o y a iables
wi h a VIF < 5 was conside ed o he GLS model (Table 1).
This model was i ed using he “gls” unc ion om he nlme R package (Pin-
hei o e al. 2023), assuming an exponen ial decay in he spa ial au oco ela ion o
ichness alues (Do mann e al. 2007). Signi ican ela ionships be ween depen-
den and explana o y a iables we e de e mined a α = 0.05, and he goodness o i
o he model was measu ed using he pseudo-R2 o Nakagawa e al. (2017).
Resul s
In asi e plan s o e Po uguese municipali ies
O he 96 in asi e plan s wi h dis ibu ion eco ds in mainland Po ugal (Sup-
pl. ma e ial 1: able S1.1), 86% a e e es ial, wi h a smalle po ion being
aqua ic o semi-aqua ic (14%), including plan s such as Al e nan he a philox-
e oides and Ludwigia spp. They include g asses and he bs (47%), ees and
sh ubs (29%), and only a ew climbing plan s (5%) and o he g ow h o ms
(Suppl. ma e ial 1: ig. S1.3). These species belong o 36 plan amilies, mainly
As e aceae (16%), Ama an haceae (14%), Fabaceae (14%), and Poaceae (9%)
(Suppl. ma e ial 1: ig. S1.4).
O he 10 species wi h eco ds in he g ea es numbe o municipali ies (Table 2),
he majo i y a e g asses and he bs (50%; e.g., Da u a s amonium, C. selloana) and
ees and sh ubs (40%; e.g., A. dealba a, Ailan hus al issima) (Table 2). Six a e
al eady p esen in mo e han 200 municipali ies (Table 2). The species eco ded
in only a ew municipali ies (up o 10; Suppl. ma e ial 1: able S1.2) a e mos ly
aqua ic o semi-aqua ic (40%; e.g., Nymphaea mexicana o Ludwigia g andi lo a).
Table 2. The 10 in asi e plan s eco ded in he g ea es numbe o Po uguese municipali ies.
Species Numbe o municipali ies wi h eco ds o he species
Acacia dealba a 249
Da u a s amonium 233
A undo donax 232
Phy olacca ame icana 224
Oxalis pes-cap ae 213
Co ade ia selloana 208
Ailan hus al issima 198
Acacia melanoxylon 190
Aga e ame icana 187
Robinia pseudoacacia 186
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
Fou species ha e been eco ded in a single municipali y (A. philoxe oides, Ama-
an hus hypochond iacus, Ama an hus mu ica us, and Baccha is halimi olia; Sup-
pl. ma e ial 1: able S1.2). The dis ibu ion o each species pe municipali y is
p o ided in Suppl. ma e ial 4.
The h ee species wi h he highes mean pe cen age o occupancy in main-
land Po ugal—O. pes-cap ae (7.53%), A undo donax (6.90%), and C. selloana
(6.87%)—a e also among he mos widely dis ibu ed, occu ing in mo e han
200 municipali ies (Table 2, Suppl. ma e ial 1: able S1.3). Thei b oad geog aphic
ange e lec s bo h high local p esence and widesp ead dispe sal ac oss he coun-
y. In con as , A c o heca calendula (5.88%) and Acacia longi olia (5.58%) ank
ou h and i h in mean occupancy bu ha e a mo e es ic ed na ional dis i-
bu ion. Thei p esence is concen a ed in coas al egions, sugges ing high local
abundance bu limi ed expansion a a b oade scale (Suppl. ma e ial 1: able S1.3).
Simila ly, he eg ession model assessing he ela ionship be ween he numbe
o municipali ies in which a species was eco ded and i s mean occupancy e ealed
a s a is ically signi ican posi i e associa ion (p < 0.05), explaining 42% o he
obse ed a ia ion. This sugges s ha species occu ing in a g ea e numbe o mu-
nicipali ies also end o occupy la ge a eas wi hin hose municipali ies. Howe e ,
he subs an ial unexplained a ia ion indica es ha addi ional ac o s in luence
species occupancy pa e ns (Fig. 2).
Figu e 2. Linea eg ession model showing he ela ionship be ween he o al numbe o municipali ies whe e each species was eco ded
and i s a e age occupancy (%). Each do ep esen s a species. The dashed line indica es he i ed linea ela ionship.
Dis ibu ion pa e ns o in asi e plan ichness
O e all, he species a e g ouped in o ou dis inc clus e s (Fig. 3, Suppl. ma e ials
1, 3: able S1.4, ig. S3.1). The i s clus e (Fig. 3) includes 26 species p edomi-
nan ly eco ded in coas al a eas, such as A. saligna (Fig. 4a) and C. co onopi olia.
154
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Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
Conclusion
This s udy p o ides insigh s in o he spa ial pa e ns and en i onmen al and
human d i e s o in asi e plan ichness in mainland Po uguese municipali ies.
Coas al u ban a eas show he highes in asi e species ichness, la gely d i en
by g ea e accessibili y. This sugges s ha municipali ies in hese egions should
p io i ize in asi e plan managemen due o hei highe le els o impac . A he
same ime, municipali ies wi h ewe in asi e species mus ac ea ly o p e en
hei sp ead, he eby inc easing he e ec i eness and sus ainabili y o manage-
men e o s na ionwide. Despi e elying on he o al numbe o in asi e plan s
in each municipali y, and no hei abundance, he esul s o e a aluable base-
line o iden i ying in asion ho spo s and guiding managemen p io i ies. Con-
inued moni o ing and sys ema ic da a collec ion a e essen ial o suppo e ec-
i e p e en ion and con ol o plan in asions ac oss he coun y.
Acknowledgmen s
We would like o exp ess ou hanks o he many expe s who con ibu ed by
p o iding use ul in o ma ion on occu ence eco ds: Ascendi, Al i, In aes u u as
de Po ugal, S.A., Liliana Dua e, Mónica Almeida, Síl ia Ma ins, MED–Medi-
e anean Ins i u e o Ag icul u e, En i onmen and De elopmen , and he Bo -
any Lab/Applied Ecology and Conse a ion Resea ch G oup o he Uni e si y o
É o a; he Na ional Elec ici y T ansmission Ne wo k shape ile p o ided by Da id
Almeida (Redes Ene gé icas Nacionais [REN]); and in o ma ion on plan nu se ies
and aqua ium s o e da a collec ed by Ana So ia Nunes (Nunes e al. unpublished).
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
We used AI o polish he manusc ip ex and help de elop he R code o he me hodology.
Funding
RF was suppo ed by a g an (PRT/BD/152100/2021 and h ps://doi.o g/10.54499/PRT/
BD/153505/2021) unded by he Po uguese Founda ion o Science and Technology (FCT) unde
he MIT Po ugal P og am. AC was suppo ed by a g an (PRT/BD/152100/2021 and h ps://
doi.o g/10.54499/PRT/BD/152100/2021) unded by he Po uguese FCT unde he MIT Po ugal
P og am. AC, RF, and CC acknowledge suppo om FCT h ough unding o he CEG/IGOT
Resea ch Uni (UID/295/2025 and h ps://doi.o g/10.54499/UID/00295/2025).
Au ho con ibu ions
RF: Concep ualiza ion, Da a cu a ion, Me hodology, W i ing—o iginal d a , W i ing— e iew
and edi ing. AC: Me hodology, W i ing—o iginal d a , W i ing— e iew and edi ing. HM: Con-
cep ualiza ion, Supe ision, W i ing— e iew and edi ing. EM: Concep ualiza ion, Supe ision,
155
NeoBio a 104: 139–162 (2025), DOI: 10.3897/neobio a.104.163291
Raquel Fe nandes e al.: Dis ibu ion pa e ns o in asi e alien plan species in mainland Po ugal
W i ing— e iew and edi ing. CC: Concep ualiza ion, Supe ision, Me hodology, W i ing—o iginal
d a , W i ing— e iew and edi ing.
Au ho ORCIDs
Raquel Fe nandes h ps://o cid.o g/0000-0002-3401-115X
And y Cas o h ps://o cid.o g/0000-0001-5635-5271
Hélia Ma chan e h ps://o cid.o g/0000-0002-3247-5663
Elizabe e Ma chan e h ps://o cid.o g/0000-0003-1303-7489
Césa Capinha h ps://o cid.o g/0000-0002-0666-9755
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex o Supplemen a y
In o ma ion.
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Au ho s: Raquel Fe nandes, And y Cas o, Hélia Ma chan e, Elizabe e Ma chan e, Césa Capinha
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.104.163291.suppl3
Supplemen a y ma e ial 4
A las o in asi e plan species
Au ho s: Raquel Fe nandes, And y Cas o, Hélia Ma chan e, Elizabe e Ma chan e, Césa Capinha
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.104.163291.suppl4
