31
Simila d i e s bu dis inc pa e ns o woody and he baceous
alien plan in asion
Da id G ego y1, Ma Whi e2, Jane A. Ca o d1,3,4
1 Depa men o Geog aphy, King’s College London, 40 Aldwych, London, WC2B 4BG, UK
2 Depa men o Ene gy, En i onmen and Clima e Ac ion, A hu Rylah Ins i u e o En i onmen al Resea ch, Vic o ia, Aus alia
3 School o Ag icul u e, Food & Ecosys em Sciences, The Uni e si y o Melbou ne, Melbou ne, Vic 3010, Aus alia
4 Fenne School o En i onmen & Socie y, The Aus alian Na ional Uni e si y, Canbe a, ACT 2601, Aus alia
Co esponding au ho : Jane A. Ca o d ([email p o ec ed])
Copy igh : © Da id G ego y 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
The ex en o alien plan in asion and numbe s o in asi e species a e inc easing, exace ba ing
in asion impac s. E ec i e and e icien managemen equi es unde s anding he d i e s and
dis ibu ion o plan in asions a he landscape scale. In his s udy, we used a species dis ibu ion
modelling app oach o de e mine whe he he pa e ns and co ela es o alien in asion a y by
plan g ow h o m. Focusing on he occupancy and p opo ional co e o o bs, g aminoids and
woody ege a ion, we used boos ed eg ession ees (BRTs) o cha ac e ise alien plan in asion
isk in wo majo ca chmen egions in Vic o ia, Aus alia. O 7,630 quad a s su eyed be ween
1970 and 2019, 69% con ained alien plan s, wi h o bs being he mos p e alen g ow h o m.
Alien plan s cons i u ed 22% o he o al numbe o plan species eco ded. Alien species co e
a ied widely, wi h alien species con ibu ing be ween 0.2% o 100% o ege a ion co e in
a plo . Alien o bs and g aminoids had highe mean co e compa ed o alien woody plan s.
Abio ic condi ions, pa icula ly empe a u e and p ecipi a ion, had he g ea es in luence on
alien plan in asion o e all, explaining 41–76% o obse ed a ia ion. Summe mean maximum
empe a u e was a s ong p edic o ac oss all g ow h o ms. A eas wi h highe ege a ion co e
(na i e + alien species) we e p edic ed o ha e highe occupancy, bu lowe p opo ional co e ,
o alien o bs and g aminoids. In con as , alien woody plan s had a nega i e ela ionship wi h
woody ege a ion co e . High le els o in asion we e p edic ed in a eas wi h in ensi e land use,
such as u ban and ag icul u al zones. Fo bs had a high p obabili y o occupancy h oughou he
egion, e en in highe ele a ions, while g aminoids and woody ege a ion we e mo e es ic ed
o lowe ele a ions and a eas cha ac e ized by human ac i i y. The s udy highligh s ha alien
plan in asion is in luenced by a complex in e play o abio ic ac o s, p opagule p essu e, human
ac i i y and bio ic condi ions. The indings unde sco e ha , while he e a e common d i e s
ac oss g ow h o ms, speci ic pa e ns and in luences a y. Fo ins ance, alien o bs we e mo e
widesp ead bu less dominan in a eas wi h high ege a ion co e , while alien woody plan s we e
less common and mo e cons ained by ege a ion han he o he wo g ow h o ms. Manage-
men s a egies should p io i ise main aining and es o ing na i e ege a ion o limi he domi-
nance o alien species and con olling in asi e plan s a e dis u bance. Al hough single-species
models emain aluable, ou s udy shows ha species dis ibu ion models based on g ow h o m
o e a p ac ical app oach o assessing plan in asions ac oss di e se landscapes.
Key wo ds: Alien ege a ion managemen , Boos ed Reg ession T ees, ecosys em in asibili y,
habi a sui abili y models, in asi e plan species isk, plan g ow h o m, species dis ibu ion models
Academic edi o : Milan Chy ý
Recei ed:
18 July 2025
Accep ed:
17 Augus 2025
Published:
9 Oc obe 2025
Ci a ion: G ego y D, Whi e M, Ca o d
JA (2025) Simila d i e s bu dis inc
pa e ns o woody and he baceous
alien plan in asion. NeoBio a 103:
31–52. h ps://doi.o g/10.3897/
neobio a.103.164914
NeoBio a 103: 31–52 (2025)
DOI: 10.3897/neobio a.103.164914
Ad ancing esea ch on alien species and biological in asions
A pee - e iewed open-access jou nal
NeoBio a
32
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
In oduc ion
Inc eases in ade and a el ha e inc eased incidence o human-media ed alien
species in oduc ion and human-induced en i onmen al change has acili a ed
highe a es o alien plan es ablishmen (Seebens e al. 2018; Pyšek e al. 2020b).
In asi e alien species, including plan s, a e conside ed one o he majo h ea s o
global biodi e si y and ha e he po en ial o cause subs an ial en i onmen al and
economic damage (Diagne e al. 2021; No oa e al. 2021; IPBES 2023). In Aus a-
lia alone, es ima es o he cos o alien plan in asions exceed AU$13.6 billion pe
yea (Ho mann and B oadhu s 2016), wi h o als o a leas AU$299 billion since
he 1960s (B adshaw e al. 2021). The e icien and e ec i e managemen o alien
species is o high economic, social and ecological impo ance. Howe e , o manage
alien plan species e ec i ely, i is i al o unde s and wha d i es in asion, how
hese d i e s migh di e ac oss di e en g oups o plan s and how his in o ma-
ion can be used o spa ially p edic he occupancy and abundance o alien plan s.
Many ecosys ems a e in aded by mul iple alien plan species (Kuebbing e al.
2013) and ege a ion managemen o ganisa ions a e o en manda ed o con ol
mul iple in asi e species ac oss a landscape (B and e al. 2023). In asi e species
managemen is mo e cos e ec i e when mul iple species a e managed in conce ,
a he han indi idually (Januchowski-Ha ley e al. 2011; Loh e al. 2017). E -
ec i e spa ial p io i isa ion o in asi e species con ol equi es in o ma ion abou
whe e alien plan s occu in he landscape, enabling ho spo s o in asion o be a -
ge ed. Gi en he limi ed a ailabili y o ege a ion su eys and he impe ec da a on
in asi e species dis ibu ions, he e is a need o p edic i e models ha can es ima e
a eas likely o expe ience high le els o in asion. Such app oaches mus be applica-
ble o a wide ange o plan species and habi a s. Howe e , unde s anding abou he
ac o s ha d i e alien plan in asions ac oss a ious landscapes emains limi ed and
e en less is known abou how hese d i e s may di e ac oss plan g ow h o ms.
Species dis ibu ion models (SDMs) a e commonly used o p edic he spa-
ial dis ibu ion o indi idual in asi e species (Fice ola e al. 2007; S i as-
a a e al. 2019). Howe e , SDMs can be da a- and esou ce-demanding and,
when buil o indi idual species, many SDMs would be equi ed o p o ide
a gene al pic u e o alien plan in asion. Fo example, based on he numbe
o eco ded alien g aminoid species, a leas 175 indi idual SDMs would be
equi ed o es ima e he spa ial dis ibu ion o alien g aminoids ha occu in
a 42,000 km2 egion o Vic o ia, Aus alia. Inc easing he numbe o SDMs
can inc ease he amoun o ime spen p epa ing he models and in e p e ing
and in eg a ing hei esul s, and he lack o empo ally and spa ially unbiased
da a o indi idual species could esul in un eliable p edic ions (Co die e al.
2020). T ai -based s udies ha examine cha ac e is ics associa ed wi h species
in asi eness p o ide insigh in o gene al d i e s o plan in asion, combining
da a om ens o hund eds o species (Ca o d e al. 2016; F is oe e al. 2021;
Palma e al. 2022). Howe e , hese s udies also demand conside able amoun s
o da a and mos ai -based s udies do no p o ide spa ially-explici p edic-
ions (Ca o d e al. 2019; Junaedi e al. 2021; Palma e al. 2021).
A majo axis o a ia ion amongs plan s is g ow h o m, wi h di e en g ow h
o ms esponding o en i onmen al g adien s and an h opogenic p essu es in
dis inc ways (Gio gis e al. 2016; Šímo á e al. 2018; Ba le e al. 2023). In
in asion ecology, classi ying alien plan species by g ow h o ms – such as o bs,
33
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
g aminoids and woody plan s – can acili a e unde s anding o hei esponses o
socioecological ac o s. Species wi hin he same g ow h o m o en ha e simila
unc ional ai s o in asions his o ies (e.g. pa hways o in oduc ion; Ba le e
al. (2023)) ha in luence hei in asi eness and in e ac ions wi h he en i on-
men (Gio gis e al. 2016; Šímo á e al. 2018). The e o e, g ouping alien plan
species by g ow h o ms can be a use ul app oach o p edic ing hei ela ion-
ships wi h en i onmen al a iables, as i can accoun o sha ed unc ional ai s
ha in luence in asion success.
G ow h o m in o ma ion is eadily a ailable o all known axa (e.g. ees,
sh ubs, o bs, g aminoids). When combined wi h species occupancy and abun-
dance eco ds, his enables g oup-based dis ibu ion models o in asion isk on
a landscape scale (Ca o d e al. 2011). G ow h o m-based dis ibu ion models
could enable spa ial p io i isa ion o mul i-species alien ege a ion managemen
wi hou he need o build hund eds o indi idual SDMs and could p o ide esul s
ha a e widely unde s ood by esea che s, p ac i ione s and he gene al public.
By conside ing hese g ow h o ms, esea che s can be e p edic and manage he
spa ial pa e ns o alien plan in asions, as each g oup may espond di e en ly
o en i onmen al p edic o s such as clima e, soil ype and dis u bance egimes.
Func ional g ouping can hus enhance he accu acy and applicabili y o p edic i e
models ac oss di e se landscapes and plan species.
In his s udy, we use a species dis ibu ion modelling app oach o de e mine
whe he he pa e ns and co ela es o alien in asion a y by plan g ow h
o m. Focusing on he occupancy and p opo ional co e abundance o o bs,
g aminoids and woody ege a ion, we use boos ed eg ession ees (BRTs) o
cha ac e ise alien plan in asion isk in wo majo ca chmen egions in Vic o-
ia, Aus alia o add ess wo ques ions:
1) How does he ela i e in luence o a iables linked o p opagule p essu e, hu-
man dis u bance, bio ic and abio ic cha ac e is ics di e amongs alien o bs,
g aminoids and woody ege a ion?
2) How do spa ial pa e ns in alien plan in asion di e amongs he h ee
g ow h o ms?
We ocus on he p opo ion o ege a ion co e ha is made up o alien plan s
as his indica es ela i e dominance (Ca o d e al. 2012; Seabloom e al. 2013,
2015). To p o ide con ex , we also examine he obse ed le el o in asion in he
s udy egion (including alien species ichness, % o su eyed plo s ha we e oc-
cupied by alien plan s, p opo ion o ege a ion co e made up o alien plan s).
Iden i ying how landscape ulne abili y o in asion a ies be ween di e en
plan g ow h o ms will enable mo e e icien mapping and moni o ing ech-
niques, acili a ing managemen p io i isa ion (Foxc o e al. 2017)
Ma e ials and me hods
S udy egion
This s udy ocuses on a egion encompassing wo Ca chmen Managemen Au-
ho i ies wi hin Vic o ia, sou h-eas e n Aus alia: Wes Gippsland and Goulbu n
B oken (Fig. 1). The Ca chmen Managemen Au ho i ies a e esponsible o
34
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
planning and coo dina ion o en i onmen al managemen wi hin hei ca ch-
men s. Wes Gippsland (~ 17,700 km2) s e ches om he Bass S ai coas in he
sou h o he G ea Di iding Range whe e i bo de s wi h Goulbu n B oken (~
24,300 km2), which ex ends o he ag icul u al loodplains o he Ri e Mu ay in
he no h. The s udy a ea co e s a la i ude ange o app oxima ely -39.16 o -35.8
and a longi ude ange o 144.7 o 148.0.
The sou he n ca chmen managemen a ea in he s udy egion (Wes Gipps-
land) expe iences d y ho summe s wi h he majo i y o he ~ 850 mm an-
nual ain occu ing in win e mon hs, wi h he cen al uplands o en ecei -
ing iple ha o he sou he n lowlands (Bu eau o Me eo ology and CSIRO
2019b).The his o ical mean daily minimum July empe a u e o Mo well, a
cen al Wes Gippsland own, is 3.7°C, while he his o ical mean daily maxi-
mum Janua y empe a u e is 26.7°C. The clima e o he no he n ca chmen
a ea (Goulbu n B oken) a ies wi h opog aphy esul ing in annual ain all
anging om 1600 mm a Lake Moun ain o 460 mm in Kyab am in he
Figu e 1. A map o he s udy egion in Vic o ia, sou h-eas Aus alia. The map illus a es a eas o he wo Ca chmen Managemen Au-
ho i ies: Goulbu n B oken (ou lined in ed) and Wes Gippsland (ou lined in yellow), which we e combined o c ea e he s udy egion.
The map highligh s he main cha ac e is ics o he s udy egion including majo owns and ci ies, oads, na ional pa ks, land o ms and
loca ions o he 7,630 ege a ion quad a su ey si es.
35
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
no h (Bu eau o Me eo ology and CSIRO 2019a). The his o ical mean daily
minimum July empe a u e o Sheppa on, a cen al Goulbu n B oken own,
is 3.4°C, while he his o ical mean daily maximum Janua y empe a u e is
32.0°C (Bu eau o Me eo ology 2025).
Geology ac oss he s udy egion a ies om clay, o g ani e, sil s one, mud-
s one, sil , sand and sands one (Geoscience Aus alia 2021). The s udy egion
has a o al popula ion o app oxima ely 415,000 people who mainly eside in
egional ci ies and u al owns (WGCMA 2019). Mo e han 50% o land is un-
de p i a e owne ship and is mainly used o ag icul u e esul ing in habi a ag-
men a ion. Public lands associa ed wi h he G ea Di iding Range expe ience
signi ican ly lowe le els o human ac i i y (GBCMA 2013; WGCMA 2019).
The s udy egion con ains a a ie y o habi a s anging om coas al sal ma sh-
es, mang o es and hea hlands in he sou h o alpine and hea ily o es ed egions
in he cen e o loodplain o es s and semi-a id woodlands in he no h. The
s udy egion was selec ed based on i s wide a ie y o en i onmen al and an-
h opogenic condi ions, high le els o in asion and in asi e plan managemen
in e es om managemen agencies.
Response a iables
We used ege a ion su ey da a om 7,630 quad a s be ween 1970 and 2019 by
he Vic o ian Depa men o Ene gy, En i onmen and Clima e Ac ion (DEECA).
These da a p o ide in o ma ion on plan species occupancy (p esence/absence)
and species co e abundance o all na i e and alien (non-na i e o Aus alia) plan
species in he s udy egion.
The coo dina es o he cen e o each 30 m × 30 m quad a we e eco ded wi h
ei he a map (p e-1993: ± 100 m accu acy) o Geog aphical Posi ioning Sys em
(pos 1993 ± 7 m accu acy). Foliage co e o all plan axa ound wi hin he
quad a s was es ima ed using he B aun-Blanque scale (Ken and Coke 1992).
Fo s a is ical analysis, B aun-Blanque scale alues we e con e ed o pe cen ag-
es on a p oximal o dinal scale (+: 1%, 1: 2%, 2: 10%, 3: 30%, 4: 60%, 5: 80%)
o es ima e g ow h o m and o al ege a ion co e abundance o each quad a .
To al co e abundance alues pe quad a could exceed 100% whe e mul iple
laye s o ege a ion (e.g. g ound and ae ial co e ) we e p esen .
The co e abundance o combined alien axa was hen calcula ed as a pe cen age
o o al (i.e. na i e and alien) plan co e . Alien o b, g aminoid (Poaceae, Juncaceae,
Cype ceae) and woody ( ees, sh ubs) plan co e was exp essed as a pe cen age o
he o al co e o hei espec i e g ow h o m. We classi ied alien plan species ac-
co ding o hei dominan g ow h o m (i.e. o bs, g aminoids o woody), a widely
used unc ional ca ego isa ion ha cap u es key ecological and phylogene ic di e -
ences amongs species (Gio gis e al. 2016; Šímo á e al. 2018; Ba le e al. 2023).
G ow h o m is a ac able and widely a ailable ai , o e ing high da a co e age
ac oss la ge species pools, and is equen ly used in in asion ecology o p o ide gene -
ali y beyond species-speci ic s udies. Mo eo e , g ow h o ms a e eadily unde s ood
by p ac i ione s and land manage s, enabling di ec applica ion o indings o policy
and managemen con ex s. While s udies using con inuous ai s can o e ine-scale
esolu ion, hey a e o en limi ed by da a a ailabili y and complexi y (Palma e al.
2021, 2022). Ou g ow h- o m-based app oach hus balances ecological in e p e -
abili y, me hodological accessibili y and ele ance o applied conse a ion.
36
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
By p oducing alien co e abundance as a p opo ion o o al co e , alien spe-
cies con ibu ion o he su ounding lo a communi y can be be e unde s ood
(Ca o d e al. 2011, 2012). As has been discussed and demons a ed in p e i-
ous s udies (e.g. Ca o d e al. (2012); Seabloom e al. (2013); Seabloom e al.
(2015)), p opo ional alien co e can e ec i ely indica e ela i e dominance o
ege a ion by exo ic species. Al hough exo ic species ichness can be co ela ed
wi h exo ic species abundance, ha is no always he case (Seabloom e al. 2013;
Seabloom e al. 2015); o example, one in asi e alien species could po en ially
con ibu e mo e co e han 10 non-in asi e alien species. We hus ocus on alien
co e a he han alien species ichness.
En i onmen al a iables
We assembled a ange o en i onmen al a iables as p oxies o p opagule p essu e,
human ac i i y, abio ic and bio ic condi ions ha a e likely o in luence alien plan
in asion (Ca o d e al. 2009; Ca o d e al. 2011; Szymu a e al. 2018; Pyšek e
al. 2020a). Raw da a o o e 30 a iables was sou ced om go e nmen al da a-
se s (i.e. om DEECA and he opensou ce h ps://www.da a. ic.go .au/da abase)
be o e being compiled in QGIS 3.14 whe e a iable da a we e sampled o each
quad a su ey loca ion. Al hough boos ed eg ession ees (BRTs) a e obus o
e ec s o mode a e mul icollinea i y amongs independen a iables (Eli h e al.
2008), we used co ela ion analysis and he li e a u e o educe he numbe o
a iables o a oid model o e i ing (De Ma co and Nób ega 2018). Co ela ion
analysis highligh ed a iables wi h high le els o co ela ion (Pea son co ela ion
coe icien > |0.7|). When mul icollinea i y was ound, he mo e dis al a iable(s)
we e emo ed in a ou o he mo e p oximal one(s). Fo his eason, empe a u e
and ain all a iables we e chosen ahead o la i ude and longi ude due o hei mo e
p oximal in luence on plan g ow h. The model o combined alien axa included
19 a iables and he models o woody, o b and g aminoids included 20 en i on-
men al a iables ollowing he addi ion o a a iable o each speci ic g ow h o m
(Table 1). Fo indi idual g ow h o m models, we included he addi ional speci ic
g ow h o m a iable o unde s and he in luence on g ow h o m-speci ic habi a
sui abili y and compe i ion on he alien g ow h o m occupancy and co e le els.
The land-co e classi ica ions used in his s udy we e spli in o i e-yea pe iods
be ween 1985–2019 and con ained 19 land-co e ca ego ies (Table 1; Suppl. ma-
e ial 1: able S1). By selec ing he land-co e pe iod closes o ha o he su ey
da e, land-co e classi ica ion accu acy a each quad a si e was maximised. To en-
su e he in asion models can be p edic ed o e la ge egions, no jus small speci ic
a eas, he land-co e ca ego ies used we e b oad enough o be used in e na ionally
ac oss a ious clima ic en i onmen s (e.g. “u ban”, “i iga ed ho icul u e”, “na i e
ees”; see Suppl. ma e ial 1: able S1).
Gap analysis
We unde ook gap analysis o ensu e he aining da a on which he model was buil
(i.e. quad a su ey loca ions) encompassed he ull ex en o en i onmen al condi-
ions p esen ac oss he s udy egion. We compa ed he maximum, minimum and
mean alues o en i onmen al a iables in he quad a su ey da a o hose o he
b oade s udy egion o ensu e ep esen a i eness. The en i onmen al condi ions
37
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
obse ed in he quad a s ep esen ed up o 100% o he en i onmen al a ia ion
in he s udy egion. No a iables exceeded 100%, con i ming ha he models did
no ex apola e beyond he condi ions ound in he aining da a.
Modelling
We an eigh models, using boos ed eg ession ees (BRTs), o model ela ionships
be ween he en i onmen al a iables and he occupancy and co e abundance o
combined alien axa, and alien o bs, g aminoids and woody ege a ion g ow h o ms.
BRTs use a wo-s age modelling me hod, which i s a se ies o simple eg ession ee
Table 1. Independen a iables used in he s udy and hei co esponding numbe ID and desc ip i e s a is ics. Fac o : B indica es bio ic
condi ions; A indica es abio ic condi ions; P indica es p opagule p essu e; H indica es human ac i i y. Because ege a ion laye s can o e -
lap, ege a ion co e can exceed 100%. Longe desc ip ions o he independen a iables can be ound in Suppl. ma e ial 1: able S1 and
hei co ela ions in Suppl. ma e ial 1: ig S1.
ID Independen Va iable B ie Desc ip ion Fac o Mean (SE) Range
1 Absolu e o b co e To al o b o e lapping co e in quad a
(na i e + alien species)
B 25.7% (0.3) 1–290%
2 Absolu e g aminoid co e To al g aminoid o e lapping co e in quad a
(na i e + alien species)
B 32.2% (0.4) 1–317%
3 Absolu e woody ege a ion
co e
To al woody o e lapping co e in quad a
(na i e + alien species)
B 57.7% (0.5) 1–425%
4 Janua y mean maximum
empe a u e
Mean maximum empe a u e in Janua y A 24.2°C (0.04) 15.7–30.9°C
5 Land-use in ensi y Pa cel size is a commonly used p oxy o land-use
in ensi y. A ea (ha) o each pa cel in Vic o ia, Aus alia
P, H 2332.0 ha (47) 0.643–26526.5 ha
6 Su ey yea Yea in which 30 m × 30 m quad a su ey was
unde aken
P 1996 (0.1) 1972–2019
7 Geology The su ace geology o he s udy a ea A Sands one (Mode) See Suppl. ma e ial 1:
able S1
8 Land co e Land-co e ca ego ies ha mos closely co espond o
su ey yea
P, A, H Na i e T ees (Mode) See Suppl. ma e ial 1:
able S1
9 Absolu e ege a ion co e To al ege a ion o e lapping co e in quad a (na i e
+ alien species)
B 120% (0.6) 1–665%
10 Wind exposi ion Wind e ec index. Re lec s a loca ions gene al
exposu e o wind. Uni less
P, A 1.1 (0.002) 0.8–1.4
11 Janua y mean ain all The mean ain all in Janua y A 58.3 mm (0.2) 28–103 mm
12 Radio-elemen Tho ium Radiome ic da a: Gamma adia ion o Tho ium
(p oxy o soil ex u e, Read e al. (2018))
A 8.6 ppm (0.06) -0.26–38.7 ppm
13 = Heigh Abo e Ri e (HAR) Log Ve ical heigh abo e i e in me es. Uni less A 0.79 (0.002) 0–1.0
14 No malised Di e ence
Vege a ion Index (NDVI)
The median No malised Di e ence Vege a ion Index.
Uni less
B 0.67 (0.002) -0.33–0.91
15 Road-demog aphic cos
dis ance analysis
Summa ion o 5 cos -dis ance a iables. Uni less P, H 11.2 (0.02) 0–15.1
16 July mean ain all The mean ain all in July A 101 mm (0.6) 38–223 mm
17 Dis ance o i e Dis ance o nea es wa e cou se P, A 0.43 km (0.01) 0–11.2 km
18 Time since i e Time since las wild i e A 35 yea s (0.4) 0–98 yea s
19 Diu nal aniso opic hea ing
x uggedness index
Diu nal Aniso opic Hea ing mul iplied by a
Ruggedness Index. Uni less
A 0.01 (0.0001) 0.3 ×10-5–0.09
20 July mean minimum
empe a u e
Mean minimum empe a u e in July A 2.9°C (0.02) -2.7–6.7°C
21 His o ical land use 1888 Land use in 1888 (Sinclai e al. 2012) P, A, H D yland Ag icul u e
(Mode)
See Suppl. ma e ial 1:
able S1
22 Topog aphic we ness index Soil mois u e index. Uni less A 0.62 (0.002) 0.18–1.0
38
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Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
analyses and hen combines hei esul s in a o wa d s age-wise manne using i e a-
i e machine-lea ning algo i hms o maximise p edic i e powe wi h high accu acy
(De’a h 2007). BRTs a e obus when modelling ecological da a due o hei abili y
o handle a ious da a ypes (con inuous and disc e e), missing da a and non-linea
ela ionships, whils p o iding s ong pe o ming p edic i e and explana o y models
(Eli h e al. 2008). BRTs also allow o easy g aphical and nume ical in e p e a ion
o complex ela ionships be ween independen a iables, making hem pa icula ly
sui able o s udying alien in asion (De’a h 2007; Eli h e al. 2008).
This s udy ollowed he BRTs me hods se ou by Eli h e al. (2008) and Eli h
and Lea hwick (2017) o model he ela ionships be ween independen a iables
and alien plan in asion. BRTs o alien occupancy and co e abundance we e un
sepa a ely, wi h he co e abundance models only using quad a s whe e alien species
we e p esen . The sepa a ion o occupancy and co e abundance models o e comes
issues associa ed wi h da a con aining a high ze o coun , allows easy in e p e a ion
and also p o ides insigh in o how in asion d i e s di e amongs in asion me ics
(Ca o d e al. 2012). Occupancy models p o ide insigh in o alien species abili y o
occupy si es and hus ole a e hei bio ic and abio ic condi ions, while co e abun-
dance models indica e when an alien species becomes dominan a a si e, esul ing in
inc eased impac s on he na i e ecosys em s uc u e and unc ion.
BRTs we e un in R ( e sion 1.3.1056) using packages “gbm” ( e sion 2.1.8; Ridge-
way (2019)) and “dismo” ( e sion 1.1–4; Hijmans e al. (2017)) and code p o id-
ed in Eli h e al. (2008) and Eli h and Lea hwick (2017). Be noulli (binomial) and
Gaussian e o dis ibu ion models we e i ed o occupancy and co e abundance
models espec i ely. To imp o e model pe o mance, co e abundance models o
combined alien axa and alien g ow h o ms ( o bs, g aminoids and woody ege a ion)
we e log- ans o med o no malise esiduals. BRT models we e adjus ed o maximise
pe o mance wi h alien plan occupancy models un using a lea ning a e o 0.01, a
ee complexi y o ou and a bag ac ion o 0.7 o maximise p edic i e pe o mance,
ollowing Ca o d e al. (2011). The pa ame e s emained he same o he alien p o-
po ional co e models excep o ee complexi y, which was inc eased o i e.
We conduc ed a Mo an’s I es (Mo an 1950), using he “spdep” package in R
(Bi and e al. 2013), o examine o spa ial au oco ela ion in he models (see Sup-
pl. ma e ial 1: appendix S1). To ensu e ha he e we e no ze o-neighbou loca ions
o disconnec ed subg aphs ac oss all models, he nea es -neighbou dis ance was
se o 0.2 deg ees (~ 22 km). Al hough s a is ically signi ican in se e al occupancy
models, he magni ude o Mo an’s I was consis en ly low (all < 0.04), indica ing
ha , while spa ial s uc u e exis s, i is unlikely o meaning ully bias p edic ions o
in alida e model assump ions (Legend e and Fo in 1989; Do mann e al. 2007).
No co ec i e spa ial modelling (e.g. spa ial eigen ec o il e ing, spa ially-explici
models) was he e o e necessa y, hough we no e ha he e was some weak spa ial
s uc u e in he occupancy models.
To assess whe he he use o a 49-yea da ase (1975–2023) in oduced empo-
al biases, we e‐ an all BRT models using a empo ally es ic ed da ase co e ing
1988–2003. This 16-yea subse cap u es he pe iod wi h he highes su ey den-
si y and ep esen s a b oad ange o en i onmen al a ia ion. The esul s om he
empo ally es ic ed da ase we e hen compa ed o he main model ou pu s (see
Suppl. ma e ial 1: ig S2). This compa ison con i med ha sampling bias was no
esponsible o d i ing he ela ionships be ween en i onmen al p edic o s and
alien plan occupancy and ela i e co e .
39
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
Model e alua ion
We used wo pe o mance me ics o selec he bes pe o ming models o each
dependen a iable:
1. Pe cen age de iance explained, exp essed as a pe cen age o he null de iance
o each esponse a iable (Lea hwick e al. 2008), p o ided a ‘goodness-o - i ’
me ic by p edic ing o da a excluded om he o iginal model aining da a.
2. A ea Unde he ecei e ope a ing Cha ac e is ic cu e (AUC) was used o
de e mine occupancy models’ abili y o disc imina e be ween quad a s wi h
o wi hou alien species. AUC is gene a ed du ing he andom c oss- ali-
da ion analysis. An AUC sco e o 1.0 indica es pe ec disc imina ion and
a sco e o 0.5 sugges s no disc imina ion accu acy (Lea hwick e al. 2008).
We also unde ook andom C oss- alida ion (CV) co ela ion analysis o measu e
he co ela ion be ween p edic ed alues and aw da a no used in model aining.
CV is unde aken by andomly spli ing he da a in o 10- olds and aining he mod-
el on nine olds, wi h he inal old being used o es ing. This es ing was epea ed a
o al o i e imes wi h he a e ages being used o display model pe o mance.
Spa ial p edic ions
We hen used hese models o p edic occupancy and co e abundance o h ee plan
g ow h o ms ( o bs, g aminoids and woody ege a ion) ac oss he s udy egion in
Vic o ia, Aus alia. The p edic ions we e mapped using he “ as e ” package in R ( e -
sion 3.3–13; Hijmans (2019)), p o iding spa ial p edic ions o he combined alien
axa and indi idual g ow h o ms’ p obabili y o occupancy and expec ed p opo -
ional co e . Spa ial esolu ion was limi ed o 50 m × 50 m due o he esolu ion o
he independen a iables. All independen a iables we e p ocessed in o as e laye s
clipped o he ex en o he s udy egion. Fo independen a iables whe e he e is no
o al co e age (e.g. o al ege a ion co e and g ow h o m ege a ion co e ), no da a
alues we e included o ensu e no da a alues we e no in e p e ed as ze os. Expec ed
co e was p oduced by mul iplying he p edic ed pe cen age co e by he p obabili y
o an alien species being p esen , ollowing he app oach o Ca o d e al. (2011).
Resul s
Le el o in asion
O he 7,630 quad a s su eyed, 69% con ained alien plan species and alien
species made up 22% o he 3,087 plan species eco ded (688 alien species,
o which 14.2% we e woody plan s, 25.4% g aminoids and 71.7% o bs). The
mean species ichness ac oss all quad a s was 28.4 (0.16 SE), anging om 1–115
species. To al ege a ion co e was weakly co ela ed wi h species ichness ( =
0.248; P < 0.001), bu no ela ionship was ound wi h o al ege a ion co e and
alien species ichness ( = 0.067; P < 0.001). When examining only he 5,239
quad a s ha con ained alien species, he mean alien p opo ional co e was
16.67% (0.31% SE), bu his anged om 0.19–100%, while he mean alien
species ichness pe quad a was 6.6 (0.09 SE) and anged om 1–45 species.
46
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
Despi e simila esponses o alien woody and he baceous ege a ion o human
ac i i ies, he e was a ia ion in he le els and spa ial dis ibu ions o he h ee
g ow h o ms examined (Fig. 6). Fo bs could be ound almos e e ywhe e in he
s udy egion, hough hei co e was lowe in he high ele a ion a eas. Compa ed
wi h alien o bs and woody ege a ion, g aminoid in asion was la gely con ined o
lowe ele a ion a eas domina ed by human ac i i ies and was much mo e limi ed
in a eas domina ed by na i e ege a ion, like in na ional pa ks and ese es and
in he G ea Di iding Range (Figs 1, 6). Gio gis e al. (2016) also ound lowe
g aminoid in asion in na i e woodlands. Woody in asion was much mo e limi ed
han he baceous in asion, which may e lec a ange o ac o s including lowe
colonisa ion and p opagule p essu e, lowe habi a sui abili y and sho e e ec i e
esidence imes (i.e. especially when conside ing species gene a ion imes).
The e a e some unusual spa ial pa e ns o alien g aminoid in asion in he
no h-wes o Goulbu n B oken ca chmen (Fig. 6e, ). This anomaly is likely
due o he lowe a ailabili y o ield obse a ion da a ac oss his egion due o
he in ensi e land-use p ac ices associa ed wi h he p o ision o i iga ion se ices.
This has a concomi an e ec o es ic ing access o land o unde ake su eys and
dampening obse e in e es . As such, he model is in o med by da a om he ew
la ge public land blocks in e ac ing wi h clima e splines, esul ing in an anoma-
lous pa e n o alien g aminoid in asion.
The p esence o weak, bu s a is ically signi ican spa ial au oco ela ion in ou
occupancy models (see Suppl. ma e ial 1: appendix S1) unde sco es he impo -
ance o conside ing spa ial p ocesses in ecological modelling, e en when using
obus machine-lea ning me hods like BRT. Howe e , he low Mo an’s I alues
sugges hese e ec s a e modes and unlikely o comp omise he o e all model
alidi y (Bini e al. 2006; Do mann e al. 2007). In con as , he lack o signi ican
au oco ela ion in co e models suppo s he spa ial adequacy o p edic o a iables
in explaining a ia ion in alien plan co e (see Suppl. ma e ial 1: appendix S1).
Managemen implica ions
Ou s udy illus a es ha in asion is d i en by a sui e o ac o s, many o hem sha ed
ac oss di e en g ow h o ms, highligh ing mul iple ulne abili ies and le e s o
managemen . Focusing on p opo ional abundance o alien axa, ou esul s sugges
ha u he na i e ege a ion loss will acili a e alien plan dominance in he s udy
egion. De o es a ion, wild i e and o he land-use changes ha dis u b na i e ege-
a ion all pose a isk o u he in asion. The s udy egion – and sou h-eas e n Aus-
alia mo e b oadly – is p one o wild i e, as he de as a ing bush i es o 2019–2020
demons a ed, a isk ha is se o ise as clima e change s eng hens (Tu belin and
Ca o d 2021). Ou esul s highligh he impo ance o main aining o es o ing
na i e ege a ion o limi alien plan dominance, especially o woody alien species
ha may o he wise s uggle o colonise. By ex ension, ou indings also con i m he
alue o ac i ely managing alien plan popula ions, while na i e ege a ion eco e s
om dis u bances like i e (Lindenmaye e al. 2015).
As well as educing na i e ege a ion and associa ed bio ic esis ance, u ban
and ag icul u al expansion will inc ease habi a agmen a ion and p opagule p es-
su e, likely inc easing in asion in a eas cu en ly expe iencing lowe in asion le els
(Boscu i e al. 2022). Based on ou s udy, alien g aminoids should bene i he mos
om his so o land-co e change (Fig. 6e, ). In conjunc ion wi h inc easing
47
NeoBio a 103: 31–52 (2025), DOI: 10.3897/neobio a.103.164914
Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
wild i e isk, clima e change is expec ed o acili a e u he in asion in he s udy
egion by aising empe a u es (Fig. 3, Ca o d and Jones (2019)). Coun e ac ing
hese changes (among o he s; Nolan e al. (2021)) h ough local and egional man-
agemen measu es, like s a egic land-use planning o ege a ion es o a ion, will
be impo an o limi ing u he in asion (O’Reilly-Nugen e al. 2024).
Iden i ying common d i e s o in asion, and egions highly suscep ible o in-
asion, can help p io i ise a eas o managemen ac ions. While use ul o examine
ends o all alien plan s combined, g ouping species by g ow h o m enables mo e
a ge ed science and managemen . Fo example, he e icacy o ce ain weed con-
ol me hods, including he bicide, can a y ac oss g ow h o ms. Single-species
models will emain a key ool in unde s anding and managing in asi e plan s, bu
mo e gene al models based on plan g ow h o m o e an e icien and in o ma i e
app oach o assessing plan in asion ac oss a iable landscapes.
Acknowledgemen s
We acknowledge he T adi ional Owne s o he land on which his wo k is
based. We hank hund eds o bo anis s who collec ed he ield da a, and Gab iele
Midolo, Milan Chy ý and an anonymous e iewe o commen s ha enabled
us o imp o e he manusc ip .
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
Funding was p o ided o JC by King’s College London Geog aphy Resea ch Impac Fund and he
Eu opean Resea ch Council (ERC) unde he Eu opean Union’s Ho izon 2020 Resea ch and Inno-
a ion P og amme (g an ag eemen No. [101002987]).
Au ho con ibu ions
JC, DG & MW concep ualised he s udy; MW p o ided he da a; DG unde ook he modelling;
DG w o e he i s d a o he manusc ip ; all au ho s con ibu ed o la e e sions o he manu-
sc ip ; JC acqui ed unding and supe ised he p ojec .
Au ho ORCIDs
Ma Whi e h ps://o cid.o g/0000-0003-2120-0071
Jane A. Ca o d h ps://o cid.o g/0000-0003-0582-5960
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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Da id G ego y e al.: Simila i ies and di e ences in woody and he baceous in asion
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Supplemen a y ma e ial 1
Simila d i e s bu dis inc pa e ns o woody and he baceous alien plan in asion
Au ho s: Da id G ego y, Ma Whi e, Jane A. Ca o d
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