Using models and connectivity analysis to the predict the current and future patterns of invasion by Silver-wattle (Acacia dealbata Link) in the Northwest Iberian Peninsula
Full text
Using models and
connec i i y analysis
o p edic he cu en
and u u e pa e ns o
in asion by Sil e -
wa le (Acacia dealba a
Link) in No hwes
Ibe ian Peninsula
Alice Lucas Roxo
Mes ado em Ecologia, Ambien e e Te i ó io
Depa amen o de Biologia
2013
O ien ado
P o esso Dou o João Hon ado
P o esso Auxilia na Faculdade de Ciências da Uni e sidade do Po o e
In es igado no Cen o de In es igação em Biodi e sidade e
Recu sos Gené icos CIBIO -UP
Faculdade de Ciências da Uni e sidade do Po o
Coo ien ado
Dou o a Joana Vicen e
In es igado a no Cen o de In es igação em Biodi e sidade e
Recu sos Gené icos CIBIO –UP
Faculdade de Ciências da Uni e sidade do Po o
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Using models and connec i i y analysis o p edic he cu en and u u e pa e ns o in asion by Sil e -wa le
(Acacia dealba a Link) in No hwes Ibe ian Peninsula
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Acknowledgmen s
Fi s o all, I would like o hank P o esso João Hon ado o he oppo uni y, he
suppo and encou agemen he ga e me h oughou his yea .
I also wan o hank Joana Vicen e, o all he a ailabili y, suppo , guidance and
encou agemen p o ided his pas yea .
To Rui Fe nandes, o all he help, a ailabili y, suppo , companionship and iendship
h oughou his p ojec .
To all he people om he esea ch g oup o Biodi e si y and Conse a ion Ecology,
specially Angela Lomba, Paulo Al es, B uno Ma cos, João Gonçal es, An ónio
Mon ei o, So ia Vaz and Ana Ri a Sil a, o he way hey ecei ed me and o all he
assis ance hey p o ided.
To F ede ico San a ém and Ve onica Ono e, o he la e 6y iendship, o being he
bes iends you can ha e, and always being he e o me, o all he ollies and
because wi hou hem my jou ney h ough his mas e would no be he same hing.
To all my iends, especially Ismael Macaia, Po i io Cos a, Diogo Sil é io, Rica do
Felicio, And é Gonçal es, Joana Rocha, Ma ga ida Hen ique, João Mou a, João
San os, Sa a Mou inho and all Bio ocas o all he memo able momen s and all he
laughs.
To Miguel F iães, o all he suppo , help, dedica ion and company his pas yea , o
being p esen in he wo s days, and o making my days be e .
Finally, bu mos impo an ly o my amily, especially my pa en s, Ana and Paulo
Sob al, and my b o he Tomas, o all hey ha e done o me, o all he suppo ,
dedica ion and ca e, because wi hou hem I wouldn’ be who I am oday and o be,
abo e all, he bes iends you can possibly ha e.
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This ese ach was de eloped a FCUP wi hin p ojec “Biodi e sidad Vege al
Amenazada Galicia-No e de Po ugal. Conoce , ges iona e implica ”
(0479_BIODIV_GNP_1_E), unded by "2º Eixo p io i á io – Coope ação e Ges ão
Conjun a e Meio Ambien e, Pa imónio e Ges ão de Riscos" o he "P og ama
Ope acional de Coope ação T ans on ei iça Espanha – Po ugal (POCTEP) 2007-
2013".
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Abs ac
Biological in asions a e one o he majo p omo e s o biodi e si y loss wo ldwide.
In asions by alien species a e hough enhanced by changes in clima e and
dis u bance egimes, as well as by o he en i onmen al shi s. P ese ing na i e
biodi e si y and ecosys ems om in asion by alien species equi es comp ehensi e
s udies and measu es o an icipa e impac s, and o p o ec species and habi a s o high
conse a ion alue. This can be achie ed using species dis ibu ion models (SDM) o
p edic he dis ibu ion o in asi e alien species (IAS) in a egion o in e es , bo h o
cu en condi ions and unde scena ios o u u e en i onmen al changes.
In his esea ch, we modeled he dis ibu ion o Acacia dealba a Link (an agg essi e
IAS) in he c oss-bo de con ex o Galicia and No h o Po ugal, in o de o p edic and
explain he cu en dis ibu ion o Acacia dealba a and o o ecas possible impac s o
u u e clima e change scena ios on ha dis ibu ion. We applied a combined p edic i e
modeling (CPM) app oach, whe e we i ed species dis ibu ion models using subse s
o p edic o s p e iously classi ied as ac ing a egional o local scales, ha we e
spa ially combined o ob ain he inal p ojec ions. These combined models p edic a
wide a ie y o po en ial species esponses, p o iding mo e in o ma i e p ojec ions o
species dis ibu ions and dynamics han adi ional models. This app oach was
complemen ed by connec i i y analyses o he dis ibu ion o he in ade , o e alua e
cu en and u u e con lic s be ween his IAS and high conse a ion alue a eas.
Model p ojec ions sugges ha he dis ibu ion o Acacia dealba a in he s udy a ea will
inc ease in he u u e, i clima e changes scena ios a e con i med, and ha hese
changes in en i onmen will inc ease he connec i i y o he species dis ibu ion in he
a ea. The e o e, using CMPs and connec i i y analysis i is possible o suppo
esou ce p io i iza ion o an icipa ion and moni o ing o IAS impac s. CMPs can
suppo measu es o p e en in asions in his egion as well as c oss-bo de
managemen in which bo h bo de s will wo k oge he and ocus hei e o s o p o ec
and conse e speci ic impo an habi a s.
Key wo ds: Acacia; Clima e change; P o ec ed a eas; Combined p edic i e modelling;
Species dis ibu ion models.
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Resumo
As in asões biológicas são um dos p incipais p omo o es da pe da de biodi e sidade a
ní el global. Conside a-se que a in asão po pa e de espécies exó icas pode se
es imulada po al e ações climá icas e dos egimes de pe u bação, bem como po
ou as mudanças ambien ais. P ese a a biodi e sidade na i a e os ecossis emas das
in asões po espécies exó icas eque es udos exaus i os e medidas pa a an ecipa
os impac os das mesmas e p o ege as espécies e habi a s de ele ado alo de
conse ação. A u ilização de modelos de dis ibuição de espécies (SDMs) pe mi e
p e e a dis ibuição de espécies exó icas in aso as (EEI) numa ce a egião, an o
pa a as condições a uais como pa a cená ios u u os de mudanças ambien ais.
Nes e abalho, modelámos a dis ibuição de Acacia dealba a Link (uma in aso a
ag essi a) num con ex o ans on ei iço (Galiza e No e de Po ugal), de o ma a
explica a a ual dis ibuição da Acacia dealba a e a an ecipa os possí eis e ei os das
al e ações climá icas nes a egião. Desen ol emos um modelo p edi i o combinado
(MPC), em que os modelos de dis ibuição o am ajus ados usando subconjun os de
a iá eis an e io men e classi icadas como impo an es à escala egional ou à escala
local, endo os dois submodelos sido espacialmen e combinados de o ma a ob e as
p ojeções inais. Es es modelos combinados o necem uma maio a iedade de
po enciais espos as das espécies, p opo cionando p ojeções da dis ibuição e
dinâmica de espécies mais in o ma i as que os modelos. Es a abo dagem oi
complemen ada com análises de conec i idade da dis ibuições da espécie in aso a,
pa a a alia os con li os a uais e u u os en e a sua dis ibuição e á eas de ele ado
alo de conse ação.
Os esul ados mos am que a dis ibuição de Acacia dealba a na á ea de es udo i á
aumen a no u u o, se as p e isões de al e ações climá icas se con i ma em, e que
es as al e ações no ambien e i ão aumen a a conec i idade da dis ibuição da espécie
na á ea. Des a o ma, a a és da u ilização de MPCs e análises de conec i idade
iden i icámos as á eas com maio p obabilidade de se em a e adas po es a in aso a,
com pa icula des aque pa a a ede egional de á eas p o egidas, e é possí el apoia
a p io ização de ecu sos pa a an ecipação e moni o ização dos impac os das EEIs.
Es a abo dagem pode apoia medidas de p e enção de in asões nes a egião e pode
ambém auxilia a ges ão ans on ei iça das in asões biológicas e de ou as ameaças
a uais e u u as à in eg idade ecológica dos habi a s e à conse ação da sua
biodi e sidade.
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Pala as – cha e: Acacia; Al e ações climá icas; Á eas p o egidas; Modelos p edi i os
combinados; Modelos de dis ibuição de espécies.
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1. In oduc ion
1.1. Clima ic change and h ea s o biodi e si y
The Ea h's biosphe e is acing an inc easing deg ada ion and loss o di e si y due o
human ac i i ies and o he an h opogenic p essu e on he global en i onmen
(Vi ousek e al., 1977). To unde s and hese changes i ’s impo an o be awa e o he
concep o biodi e si y. Biodi e si y is he a iabili y among li ing o ganisms om all
sou ces and he ecological complexes o which hey a e pa , which includes di e si y
wi hin species, be ween species as well as o ecosys ems (Millenium
Ecosys em Assessmen , 2005). Biodi e si y is undamen al o ecosys em s uc u e and
unc ioning, and i suppo s he b oad spec um o goods and se ices ha humans
de i e om na u al sys ems (S audinge e al., 2012).
Human al e a ion o he global en i onmen has induced he six h ex inc ion e en in
his o y o li e (Di zo and Ra en, 2003; Chapin e al., 2000) p omo ing changes in he
global dis ibu ion o o ganisms (Chapin e al., 2000). Modi ica ions in species
dis ibu ion can change bo h ecosys em p ocesses, and he esilience o ecosys ems o
en i onmen al changes (S audinge e al., 2012, Chapin e al., 2000), wi h hough ul
consequences in ecosys em se ices (Chapin e al., 2000). The majo causes o
biodi e si y decline a e: land use changes, in asi e species, pollu ion, clima e change
and o e exploi a ion esul ing om economic, sociopoli ical, cul u al, demog aphic,
echnological, and o he indi ec d i e s (Millennium Ecosys em Assessmen , 2005).
1.2. Biological in asions as a d i e o biodi e si y change
Biological in asions can be de ined as he in oduc ion and sp ead o exo ic o ganisms
in o egions ou side o hei na i e ange (Méndez e al., 2011), and hey a e among he
mos impo an human-d i en agen s o change in na u al en i onmen s wo ldwide
(Na ščius e al., 2012). The a e o biological in asions is quickly inc easing las yea s
(Essl e al., 2011) and in asi e species a e causing nega i e e ec s on bo h
biodi e si y and human well-being (Essl e al., 2011). Impac s o in ade species
include: modi ica ions in composi ion, s uc u e and unc ioning o he ecosys ems
(Te e eai e al., 2013), habi a changes by elimina ion and/o educ ion o na i e
species (Na ščius e al., 2012), bio ic homogeniza ion, loss o biodi e si y and loss o
ecosys em se ices in many pa s o he wo ld (Cas o-Díez e al., 2011).
Plan s in asions ha e become ecognized as a majo h ea o bo h na u al and
human-exploi ed ecosys ems wo ldwide (Hobbs and Humph ies, 1995), as hey a ec
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ecosys em s uc u e and unc ion (Lake and Leishman, 2004). Human ac i i ies helped
(in en ionally and acciden ally) ha many plan species g ow ou side hei na i e
anges, some o hese non-na i e plan s o e come se e al ba ie s, becoming in asi e
(B adley e al. 2010). To a success ul in asion many di e en mechanisms a e in ol ed
(Gulezian and Nybe g, 2010) and many di e en s ages/ecological p ocesses need o
be o e come a di e en spa ial scales (B own e al., 2008; Richa dson e al., 2000).
The i s ba ie co esponds o a majo geog aphical ba ie ha he plan o e comes
h ough human help (Figu e 1, Ba ie A); a e , he species is called casual. The
second ba ie co esponds o an en i onmen al il e , in ol ing bio ic and abio ic
ac o s (Figu e 1; Ba ie B). Then, he species needs o o e come he ep oduc ion
ba ie (Figu e 1; Ba ie C). A e o e coming hese h ee i s ba ie s, a axon is
conside ed success ully na u alized. A species o popula ion can be called in asi e i
o e comes he local/ egional dispe sal ba ie s (Figu e 1; Ba ie D), he en i onmen al
ba ie (s) in human modi ied o alien-domina ed ege a ion (Figu e 1; Ba ie E), and
he en i onmen al ba ie s in na u al o semi-na u al ege a ions (Figu e 1; Ba ie E;
Richa dson e al., 2000). Mo eo e , he ex en o in asion by alien species is a unc ion
o ecosys em-le el p ope ies (in asibili y), p opagule p essu e, cha ac e is ics o he
in asi e species (in asi eness), and he p ope ies o he indi idual na i e species in
he ecosys em (Wes phal e al., 2008).
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Figu e 1 - S ages o in asion p ocess and hei ep esen a i e ba ie s ( om Richa dson e al., 2000).
Alien in ade s a e o en “passenge s” p i ileged (Gae ne and Richa dson, 2011) by
o he en i onmen al changes (Gae ne and Richa dson, 2011), such as ising
empe a u e, inc eased a mosphe ic ca bon dioxide (CO2), ex eme e en s in
p ecipi a ion, ni ogen (N) deposi ion, and dis u bances associa ed wi h changes in land
use o land co e (B adley e al., 2009). Fo example, changes in clima ic condi ions
migh di ec ly in luence he likelihood o alien species in a e i o y, and also a ec he
na u aliza ion success (Wal he e al., 2009). Syne gic e ec s be ween alien in ade s
and global changes di icul he indi idualiza ion be ween bo h he e ec s o alien
species on he na i e ecosys ems, and he e ec s o he dis u bance ha lead o he
ini ial plan in asion (Gae ne and Richa dson, 2011).
1.3. The speci ic case o in asi e acacias
The in asi e alien plan s belonging o genus Acacia a e among he mos common
in ade s in many pa s o he wo ld (González-Muñoz e al., 2012), a ec ing
ecosys ems s uc u e and unc ioning wo ldwide (Lo enzo e al., 2010), and a e al eady
p esen in sensi i e habi a s in Eu ope (coas al dunes, i e cou ses, na u al pa ks and
biosphe e ese es; Lo enzo e al., 2010). The e o e in asions by Acacia genus
Ba ie s: A B C DE F
1. Geog aphic
2. En i onmen al (local)
3. Rep oduc ion
4. Dispe sal
5. En i onmen al (Dis u bed habi a s)
6. En i onmen al (Na u al habi a s)
Alien
Na u alizedCasual
In asi e
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(wa les) ep esen s a h ea o na u al habi a s h ough compe i ion and eplacemen o
na i e species, homogeniza ion o he communi y, and loss o na i e biodi e si y
(Fuen es-Ramí ez e al., 2011). Some cha ac e is ics o his genus p omo e he h ea
o na i e ecosys ems and species, pa icula ly he high coloniza ion capaci y, e.g. in
places dis u bed by i e, ha es ing and o he ypes o human dis u bance (Fuen es-
Ramí ez e al., 2011). Some key ai s a e p esen in his genus p omo ing he
dominance in compe i i e in e ac ions wi h na i e species as: apid g ow h a es and
abili y o ou -compe e na i e plan s; capaci y o accumula e high olumes o biomass;
la ge, pe sis en seed banks, and he capaci y o ix ni ogen (Le Mai e e al., 2011).
The e a e a leas eigh wa le species na u alized and po en ial in asi e plan s in he
sou h o Eu ope (Lo enzo e al., 2010): A. dealba a, A. melanoxylon, A. e inodes, A.
saligna, A. ka oo, A. pycnan ha, A. mea nsii, and A. longi olia. Th ee o he Acacia
species: A. dealba a; A. melanoxylon, and A. longi olia a e he mos p oblema ic
in ade s in F ance, Po ugal, Spain and I aly, pa icula ly in conse a ion a eas, being
A. dealba a he mos widesp ead species (Lo enzo e al., 2010).
1.4. P o ec ed a eas
Acco ding o In e na ional Union Conse a ion o Na u e (IUCN, 2008), a p o ec ed
a ea is “a clea ly de ined geog aphical space, ecognized, and managed, h ough legal
o o he e ec i e means, o achie e he long e m conse a ion o na u e wi h
associa ed ecosys em se ices and cul u al alues” (h p://www.iucn.o g/). P o ec ed
a eas comp ises mo e han 12.7% o he plane ’s land su ace (Geldmann e al., 2013)
being an impo an ool and s a egy o main ain habi a in eg i y and species di e si y,
ensu ing he main enance o na u al p ocesses ac oss landscapes and ecosys ems
(h p://www.cbd.in /p o ec ed/o e iew/). The designa ion and main enance o
p o ec ed a eas is he mos impo an and e icien conse a ion s a egy wo ldwide
(Kleinbaue e al., 2010), none heless he impo ance o na u e ese es a ies
depending he egions o he wo ld, ela ed o he deg ee o an h opic ans o ma ion o
he e i o y (Pysek e al., 2002). Al hough p o ec ed a eas emain a c ucial key o
global di e si y conse a ion s a egies, hey emain suscep ible o an h opogenic
changes and consequences (Spea e al., 2013), especially o alien plan in asion.
Alien in ade species a e e y p oblema ic o conse a ion alue a eas due o he
po en ially h ea o he pe sis ence o na i e species (Uddin e al., 2013). The deg ee
o a ese e in asion can be ela ed in some cases wi h he o al numbe o isi o s
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(Pysek e al., 2002). Recen ly Pysek e al. (2002) shown ha na u e ese es all o e
he wo ld a e in aded abou hal as o en as si es ou sides ese es.
1.5. Using models o achie e and an icipa e pa e ns o in asion
Species dis ibu ion models (SDM) a e empi ical models ha ela e ield obse a ions
o en i onmen al p edic o s a iables cen e ed on he s a is ical o heo e ical de i ed
esponse su aces (Guisan and Thuille , 2005). The use o SDMs has g own in he las
wo decades due o he ecognized applica ion alue in applied scien i ic ields as
ecosys em managemen and biodi e si y conse a ion (Rod íguez-Reya e al., 2013).
SDMs can also be applied in undamen al ecological ques ions as, such as he
ecological impac s o clima e and land-use changes in biological in asions (Vicen e e
al., 2011; Guisan and Thuille , 2005). Many scien i ic s udies in in asions applied
SDMs as a ool, o example: o assess species in asion and p oli e a ion, o model
species assemblages (biodi e si y, composi ion), o quan i y he en i onmen al niche o
he species, o suppo app op ia e managemen plans o species eco e y (mapping
sui able si es o species ein oduc ion), o suppo conse a ion planning and ese e
selec ion (sugges ing su eyed si es o high po en ial o occu ence o a e species;
Guisan and Thuille , 2005). Despi e many ecen applied pu poses o SDMs ela ed o
clima e change and conse a ion planning, he use o hese ools in heo e ical ecology
and e olu ion is e-eme ging (Guisan and Thuille , 2005).
Acco ding o Nen wig e al. 2008, he main esea ch ields o biological in asions can
be classi ied as: pa hways o biological in asions, ai s o a good in ade , pa e ns o
in asion and in asibili y, ecological impac s o biological in asions, economy and
socio-economy o biological in asions, and inally, p e en ion and managemen o
biological in asions. One o he majo goals in biological in asion s udies is o
unde s and which and how bio ic and abio ic ac o s cons ain he sp ead o in asi e
species, de e mining cu en and u u e po en ial dis ibu ions (Wang and Jackson,
2011; Fa ashi e al., 2013). Resea ch e o s mus be di ec ed o ind ways o an icipa e
in asions in o de o p o ec global biodi e si y om he e ec s o alien species
(Vicen e e al., 2011; Jones, 2012). This goal can be pa ially achie ed de e mining he
po en ial geog aphic ex en o in asions, equi ing p edic i e ools o assess he
geog aphic dis ibu ion o in asi e species unde cu en and u u e en i onmen al
condi ions (Vicen e e al., 2011; Fa ashi e al., 2013). These p edic ions can simpli y
he ea ly de ec ion o in asi e species and maximize moni o ing e iciency (Jones,
2012).
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Ongoing clima e and land-use changes a e o ecas ed o boos he in asion o alien
species in se e al habi a s (Vicen e e al., 2011) being impo an o ha e a mul i-scale
app oaches due o mos o he pa hways o species in oduc ions esul om human
ac i i ies a se e al spa ial and empo al scales (Vicen e e al., 2011; Razgou e al.,
2011). Al hough mos o he s udies using SDMs conside ed en i onmen al p edic o s
a a single g ain size and ixed spa ial ex en , ac o s d i ing he dis ibu ion and
abundance o o ganisms can ac a di e en scales (Vicen e e al., 2011; Guisan and
Thuille , 2005). Fo example: opog aphy, geomo phology, human land-use o bio ic
in e ac ions a e usually conside ed o ope a e a egional and local scales while clima e
ope a e a he global scale (Vicen e e al., 2011; Eli h and Lea hwicK, 2009).
The e a e se e al a ailable modelling ools (Thuille e al., 2009), being Biomod2 one
o he newes and accu a e s a is ical package. Biomod2 is implemen ed in he R
s a is ical so wa e (Geo ges and Thuille , 2013), allowing he calib a ion o ensemble
o ecas ing species dis ibu ions, o e coming a ange o unce ain ies and assump ions
in he me hodological models. This package allows he calib a ion o a model applying
10 di e en modelling echniques, and o o ecas he po en ial species dis ibu ion
unde di e en en i onmen al condi ions (clima e and land use change scena ios;
Thuille e al. 2009). Due o he complex na u e o species ange dynamics and he
ela ed ac o s ac ing a mul iple spa ial scales, modelling species dis ibu ion is a
challenging p ocedu e (Vicen e e al., 2011; Guisan and Thuille , 2005).
1.6. Objec i es
In his wo k we applied a combined p edic i e modelling (CPM) amewo k o: (1)
iden i y he a eas ha a e oday po en ially in aded by Sil e -wa le (Acacia dealba a
Link) in No hwes Ibe ian Peninsula (Galicia and No h o Po ugal); and (2) o ecas
he e ec s o clima e changes on he dis ibu ion o his species by p ojec ing he
models o u u e condi ions. Ou a ionale was ha he applica ion o CMPs o p edic
he cu en and u u e pa e ns o in asion can suppo measu es o p e en in asions
in he s udy egion and suppo c oss-bo de managemen o p o ec impo an habi a s
om in asion.
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2. Me hods
2.1. S udy a ea
The s udy a ea is loca ed in he No hwes Ibe ian Peninsula (Fig. 2), co e ing he
No h Po ugal and Galicia. This a ea p esen s a high plan biodi e si y, wi h mo e han
3000 species and subspecies, many o hem endemic o his egion
(h p://www.biodi e sidade.eu/p /in o/p oxec o/). This egion is expe iencing a s ong
en i onmen al dis u bance p omo ed by many ac o s, like in asi e species, c ea ing a
se ious h ea o conse a ion o biodi e si y
(h p://www.biodi e sidade.eu/p /in o/p oxec o/).
Figu e 2 – Loca ion o he s udy a ea in Eu ope (a) and Ibe ian Peninsula (b) wi h digi al ele a ion model de ail (c).
The a ea ex en is 35017 km², co e ing he ansi ion be ween he Medi e anean and
A lan ic biogeog aphic egions (Rí as-Ma ínez e al., 2004). The s udy a ea is e y
he e ogeneous, wi h ele a ion anging om he sea le el (wes ) o 2059 m in he
eas e n moun ains, wi h he majo i e s, as Dou o, Tua, and Cenza, unning om eas
o wes . The geology o he moun ains is o e whelmed by acid igneous and
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me amo phic ocks ha o m acid soils (Roucoux e al., 2005). The s udy a ea p esen s
a e y complex land co e (mo e han 11 classes o land co e , acco ding o he
Eu opean En i onmen Agency) and a la ge ne wo k o p o ec ed si es: Pa que
Nacional Peneda-Ge es, Pa que Na u al de Mon esinho and Pa que Na u al de Al ão
in Po ugal and Se a Candán and Pena T e inca in Spain. Cu en ly his egion has a
empe a e clima e (Roucoux e al., 2005) and ca ies he ull o ce o he wes e ly winds
b inging cyclones om he A lan ic (Roucoux e al., 2005). The p ecipi a ion le els in
he a ea a e he highes in he whole peninsula (Roucoux e al., 2005) (up o 3000mm
pe yea in he eas e n moun ain summi s). The annual mean empe a u es anges
om ca. 5°C o ca. 17°C, whe e he summe s a e cool (18-22ºC (Roucoux e al., 2005)
and win e s a e mild and os - ee (10-13ºC a ound he coas ) (Roucoux e al., 2005).
2.2. Tes species and occu ence da a
The es species is an alien in ade plan om he Leguminosae amily, Acacia
dealba a Link (Fig. 3). This wa le species is o iginal om he sou heas o Aus alia
and ound in a wide ange o di e en habi a s, om coas al o subalpine egions, and
om high ain all o a id inland a eas, g owing in opical, sub opical and wa m
empe a e egions (Lo enzo e al., 2009).
Figu e 3 – Acacia dealba a Link.
This species can be ound in a eas wi h o e 500 mm ain all, ypically a al i udes om
350-1000m abo e he sea le el (Lo enzo e al., 2009). In oduced in Eu ope in he
1820s (Ca ballei a and Reigosa, 1999), and plan ed as an o namen al plan in sou he n
Eu ope, which o e ed a o able clima es o his de elopmen , wi h su icien sun
exposu e an li le os s, becoming widely na u alized in his a ea (Lo enzo e al.,
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2009). In he Sou hwes Eu ope his species occu s in ipa ian zones, wa e cou ses
and sunny edges o pinewoods, o on he sou h and wes acing slopes, whe e i o ms
dense s ands ha choke in he na u al ege a ion and i o en in ades a ea unde
in ensi e ag icul u e uses and u he away om he sea han o he Acacia species
p esen in hese egions (Lo enzo e al, 2009). Acacia dealba a was in oduced in he
Ibe ian Peninsula in he second hal o he 19 h cen u y (Sanz Elo za e al., 2004) and
became a p oblema ic species in Po ugal and Spain, whe e i is h ea ening he na i e
lo a and becoming a se ious en i onmen al p oblem. In addi ion o i s g ea colonizing
capaci y i leads o a e y low co e ing o unde g ow h species caused by i s
alellopa hic abili y (Vicen e e al, 2011).
The occu ence da ase o Po ugal was collec ed om p e ious scien i ic wo ks
(Vicen e e al., 2013) and Spain da a we e and p o ided by Di eccion Xe al de
Conse acion da Na u eza da Xun a da Galicia. Sampling si es we e selec ed using a
andom-s a i ied s a egy (Vicen e e al., 2013), employing clima e (mean annual
empe a u e) and bed ock as s a i ying laye . Sampling uni s we e cells o 1km2 in
which p esen s whe e collec ed be ween Ma ch and Ap il 2011.
2.3. Selec ion and classi ica ion o en i onmen al p edic o s
We selec ed 53 p edic o s ha , by expe knowledge and acco ding o p e ious
epo ing in scien i ic li e a u e, ope a e as de e minan s o ecology and dis ibu ion o
he a ge species (see annex 1). To a oid co ela ion, we applied he Spea man ank
co ela ion coe icien (non-pa ame ic es ), o selec he p edic o s wi h lowes
co ela ion be ween each o he . We use he so wa e STATISTICA o selec he
p edic o s and only hose wi h co ela ion lowe han 0.5 we e conside ed (Vicen e e
al., 2011). The e o e we selec 17 en i onmen al p edic o s o calib a e he models
(Table 1 desc ibes he p edic o s chosen). Clima e p edic o s we e ob ained using
in o ma ion om he Wo lClim da abase in as e o ma wi h a spa ial esolu ion o
1km2 while he emaining en i onmen al p edic o s we e ob ained om de ailed
en i onmen al maps.
The en i onmen al p edic o s we e classi ied a p io i as “ egional” o “local” based on
ecological heo y and hei spa ial scale o a ia ion using a s a is ical p edic o
classi ica ion based on spa ial au oco ela ion (Vicen e e al., 2011).Using poin pa e n
s a is ic (PPSA—spdep R package; a ailable a h p://c an. -p ojec .o g/web/
packages/spdep) we calcula ed Gea y’s c au oco ela ion measu e o all p edic o
a iables o inc easing neighbou hood dis ances. A leas , we pe o med a hie a chical
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clus e ing based on a Euclidean dis ance ma ix o all Gea y’s alues and he wo
g oups in he inal classi ica ion ee we e e y well sepa a ed co esponding o
p edic o s wi h local e sus egional in luence (Vicen e e al., 2011).
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Figu e 5 – Possible combina ions o species dis ibu ions dynamics using combined models (Vicen e e al., 2011). The
le e s A, B, C and D e e s o he di e en possible combina ions (A- egional and local sui abili y, B- only egional
sui abili y. C- only local sui abili y and D- no sui abili y).
2.7. Cu en and u u e con lic s wi h p o ec ed a eas
The models o he species we e used o o ecas he cu en and u u e species
dis ibu ions and hei con lic wi h he p o ec ed a eas p esen in he s udy a ea, by
spa ially o e lapping he po en ial dis ibu ion maps wi h a map o he p o ec ed a ea,
co esponding o Na u a 2000 ne wo ks. Wi h his p ojec ion i 's possible o de e mine
he po en ial h ea s o he p o ec ed a eas in he nea u u e, as well as he possible
impac s ha may a ise om hem. By o e lapping he models o he species wi h a
map o p o ec ed a eas (co esponding o Na u e Ne wo k 2000 – ICNF o Po ugal
and o Espacios Na u ales P o egidos en España o Spain) he dis ibu ion maps o
combined models allow a de ec ion o p esen and u u e in asion a eas.
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2.8. Connec i i y o he dis ibu ion o he in asi e species
Fi s , we calcula ed he empo al changes in spa ial connec i i y o p edic ed sui able
a eas o he es species ac oss he es a ea, using he connec i i y index de eloped
by Randy e al., 2009 (Vicen e e al., 2013). We conside ed alue o species p esence:
p obabili y o 1 o he esponse ype A, p obabili y o 0.5 o esponse ype B and C,
and p obabili y o 0 o esponse ype D. The connec i i y index a ains a maximum
alue o 1 when all he cells su ounding a ocal sui able cell a e also sui able (Vicen e
e al., 2013). We quan i ied he changes in he spa ial ela ionship be ween p o ec ed
a eas and he connec i i y o p edic ed a eas using he as e calcula o in A cGIS 9.3
(ESRI, 2010). Then, we analysed he end o connec i i y o he specie in space ( ull
a ea and inside p o ec ed a eas) and ime (2020 and 2050). A las , a Spea man ank
co ela ion es was used o iden i y signi ican ela ions be ween connec i i y and
p o ec ion alue.
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3. Resul s
3.1. P edic ions and de e minan s o cu en dis ibu ions
The po en ial dis ibu ion o he species Acacia dealba a unde cu en condi ions is
illus a ed in Figu e 6. The combined p edic i e model was calib a ed using 17
en i onmen al a iables "a p io i" classi ied as " egional" and "local" (see Table 2).
Figu e 6 - Spa ial p ojec ion o Acacia dealba a po en ial dis ibu ion using combined models, unde cu en condi ions
(2000).
.
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Table 2 – En i onmen al p edic o s selec ed o calib a e he combined models.
The pa ial egional model was calib a ed wi h 7 en i onmen al a iables, and an AUC
alue o 0.849 was ob ained, while he pa ial local model was calib a ed wi h 10
en i onmen al a iables wi h a inal AUC alue o 0.887. F om he 7 egional
en i onmen al p edic o s, he mos impo an o pa ial egional model calib a ion we e
Minimum Tempe a u e o Coldes Mon h (MTCM, wi h a alue o 0.394), Annual
P ecipi a ion (AP, wi h a alue o 0.282) and Tempe a u e Annual Range (TAR, wi h a
alue o 0.238). Conside ing he pa ial local model, be ween he o al o 10 local
p edic o s, he mos impo an we e Densi y o local oad ne wo k (DensRoad, wi h a
alue o 0.392), pe cen age o a able land (pA L, wi h a alue o 0.173 )and pe cen age
co e o mixed o es (pMixFo, wi h a alue o 0.197).
Unde cu en clima e condi ions (2000), he po en ial pe cen age o p edic ed a ea o
Acacia dealba a in a eas wi h egional and local sui abili y, ype A, was 18.3%. Fo
a eas wi h only- egional sui abili y, ype B, he pe cen age o p edic ed a ea 27% while
in he a eas wi h only local sui abili y, ype C, he pe cen age o p edic ed a ea was
P edic o s
Desc ip ion
Scale o a ia ion
AP
Annual P ecipi a ion
Regional
Dis Ri
Dis an e o main i e
Regional
GPP
Mean g oss annual p ima y p oduc i i yMM
Regional
MTCM
Min Tempe a u e o Coldes Mon
Regional
pCamb
% o cambissoils (pe 1km2)
Regional
PS
P ecipi a ion Seasonali y
Regional
TAR
Tempe a u e Annual Range
Regional
DensRi
Densi y o local hyd og aphic ne wo k
Local
DensRoad
Densi y o local oad ne wo k
Local
Dis oad
Dis ance o oads
Local
MPAR
Mean pe ime e -a ea a io
Local
MSI
Mean shape index
Local
PA L
% co e o a able land (pe 1km2)
Local
PA S
% co e o a i icial s ands (pe 1km2)
Local
pBiFo
% co e o b oad-lea o es (pe 1km2)
Local
pCoFo
% co e o coni e o es (pe 1km2)
Local
pMixFo
% co e o mix o es (pe 1km2)
Loca
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14%. Finally, o a eas p edic ed o be unsui able o he species (unsui able local and
egional condi ions), ype D, he pe cen age o p edic ed a ea was 40.7% (Table 3).
Table 3 – P edic ed pe cen age o he s udy a ea occupied by each “occupancy ype” in he combined models, o he
es species, unde cu en condi ions (2000).
Occupancy ypes
Pe cen age o p edic ed á ea
Type A - egional and local sui abili y
18.3%
Type B - only egional sui abili y
27.0%
Type C - only local sui abili y
14.0%
Type D - no sui abili y
40.7%
3.2. Fo ecas o u u e dis ibu ions
The spa ial dis ibu ion o he es species unde u u e clima e condi ions o he yea
2020 is illus a ed in Figu e 7. The combined models o he A1 scena io (Fig. 7-b) and
o he B2 scena io (Fig. 7-d) we e ob ained by o e lapping he spa ial p ojec ions o
he local and he egional pa ial models.
Figu e 7- Spa ial p ojec ions unde cu en condi ions (2000 - a) and u u e condi ions - 2020 A1 scena io (b) and Bb2
scena io (d) and 2050 A1 scena io (c) and B2 scena io (e).
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Unde u u e condi ions (2020) conside ing A1 clima ic scena io, p edic ed a ea wi h
local and egional sui abili y, ype A, was 22.3%.The pe cen age o a eas wi h only
egional sui abili y, ype B, was 33.3%, and o a eas wi h only local sui abili y, ype C
was 10%. Finally, unsui able egional and local a eas, ype D, we e p edic ed as
34.4%. Fo B2 clima ic scena io, ype A (bo h egional and local sui abili y) p edic ed
a ea was 20%, ype B ( egional sui abili y) was 28.9%, o ype C (local sui abili y) was
12.3% and o ype D (bo h egional and local unsui abili y) was 38.8% (Table 4).
Table 4 - P edic ed pe cen age o he s udy a ea occupied by each “occupancy ype” in he combined models, o he
es species, unde u u e condi ions (2020), o he a1 and b2 scena ios.
Pe cen age o p edic ed a ea
Occupancy ypes
A1
B1
Type A - egional and local sui abili y
22.3%
20.0%
Type B - only egional sui abili y
33.3%
28.9%
Type C - only local sui abili y
10.0%
12.3%
Type D - no sui abili y
34.4%
38.8%
A spa ial dis ibu ion o he species Acacia dealba a, unde u u e clima e condi ions
o he yea 2050 is p esen ed in he igu e 7, whe e he combined model o he A1
scena io ( ig. 7- c) and o he B2 scena io( ig 7- e).
In able 5 is p esen ed he es ima ed pe cen age o he s udy a ea occupied by each
“occupancy ypes”. A eas p edic ed wi h bo h egional and local sui abili y (Type A) was
26,5% in A1 scena io and 22.8% in B2 scena io. The pe cen age o a ea p edic ed o
be ype B (only egional sui abili y) was 46% in he a1 scena io and 36.4% in he b2
scena io, and he pe cen age o a eas p edic ed as ype C (only local sui abili y) was
5.8% in he a1 scena io and 9,4% in b2. A eas p edic ed as unsui able local and
egional condi ions (Type D) was 21.7% and 31.4% in he a1 and b2 scena ios,
espec i ely.
Table 5 - P edic ed pe cen age o he s udy a ea occupied by each “occupancy ype” in he combined models, o he
es species, unde u u e condi ions (2050), o he a1 and b2 scena ios.
Pe cen age o p edic ed a ea
Occupancy ypes
A1
B2
Type A - egional and local sui abili y
26.5%
22.8%
Type B - only egional sui abili y
46.0%
36.4%
Type C - only local sui abili y
5.8%
9.4%
Type D - no sui abili y
21.7%
31.4%
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3.3. Range dynamics be ween yea s 2000 and 2050
Based on he po en ial dis ibu ion o he species p edic ed o he yea s o 2000, 2020
and 2050, we analysed he po en ial ange dynamics o he species Acacia dealba a
be ween he ime pe iods: 2000-2020; 2000-2050; and 2020-2050. P edic ed changes
in ange dynamics o he species Acacia dealba a be ween 2000-2020, 2000-2050
and 2020-2050 (unde clima e change scena ios) a e shown in Figu e 8.
Fo he h ee ime pe iods, p edomina e dynamic ob ained was he esponse ype “no
change” (see Table 6). In bo h scena ios, o he esponses ype “Imp o emen o he
condi ions” and “Coloniza ion”, om he pe iods 2000-2020 o 2000-2050 an inc ease
o he esponse was obse ed, and o he pe iod o 2020-2050 a dec ease was
obse ed compa ed o he ime pe iod 2000-2050. Fo he esponse ype “De e io a ion
o he condi ions”, o bo h scena ios, a dec ease end was obse ed om he pe iod
o 2000-2020 o he pe iod o 2000-2050. F om ime pe iod 2000-2050 o 2020-2050
he ob ained alues we e he same. Finally, o he esponse ype “ex inc ion”, a
dec easing o he alues was obse ed om he ime pe iod 2000-2020 o 2000-2050
and also om he ime pe iod o 2000-2050 o 2020-2050.
Table 6 – Dynamic o en i onmen al sui abili y o he es species om combined modes, unde clima e
change scena ios, o he pe iods 2000-2020, 2000-2050 and 2020-2050.
Scena ios
Dynamics
No change
Imp o emen
De e io a ion
Ex inc ion
Coloniza ion
A1B
2000-2020
77.9%
5.1%
1.0%
4.9%
11.1%
2000-2050
68.1%
8.6%
0.3%
2.0%
2.1%
2020-2050
80.0%
4.5%
0.3%
1.3%
13.9%
B2
2000-2020
89.9%
2.7%
0.9%
4.3%
6.2%
2000-2050
79.1%
5.2%
0.6%
2.9%
12.2%
2020-2050
84.2%
3.4%
0.6%
2.2%
9.6%
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Figu e 8 – Changes in he po en ial and dynamic o he es species be ween 2000-2020 ( o he A1 scena io – a- and
he B2 scena io - d), 2000-2050 ( o he A1 scena io – b- and he B2 scena io - e) and 2020-2050 ( o he A1 scena io –
c - and he B2 scena io - ).
3.4. Cu en and u u e con lic s wi h p o ec ed a eas
In Table 7 we p esen he pe cen age o a ea o Na u a 2000 si es occupied by he
di e en esponse ypes o he es species Acacia dealba a, o cu en (2000) and
u u e (2020 and 2050) en i onmen al condi ions (Figu e 9).Fo he clima ic scena io
A1, he pe cen age o p edic ed a ea wi h he esponse ypes A ( egional and local
sui abili y o he species) wi hin Na u a 2000 si es inc eases in he yea 2020 and
2050. Conside ing p edic ed ype B ( egional sui abili y) and D (bo h unsui able
egional and local) inside he Na u a 2000 si es, i ’s p edic ed no changes o he yea
2020 ollowed by an inc ease o p edic ed pe cen age o ype B ( egional sui abili y),
and a dec ease in he ype D (bo h egional and local unsui abili y), o 2050. The
pe cen age o occupancy o he ype C (local sui abili y) inside he Na u a 2000 si es
dec eases bo h yea s.
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Fo he clima ic scena io B2, a dec ease o he pe cen age o occupancy o he species
inside Na u a 2000 si es can be obse ed o he yea o 2020 o esponse ypes A
( egional and local sui abili y) and B ( egional sui abili y), ollowed by an inc ease in he
yea 2050. An inc ease o he pe cen age is obse ed o esponse ypes C (local
sui abili y) and D (bo h egional and local unsui abili y) o he yea 2020, while in 2050
he pe cen age dec eases.
Table 7– Pe cen age o occupancy in Na u a 2000 o each occu ence ype ( egional and local sui abili y ( ype A), only
egional sui abili y ( ype B), only local sui abili y ( ype C), and no sui abili y ( ype D), o he es species, o cu en
(2000) and u u e (2000, 2050) condi ions.
Pe cen age o p edic ed a ea
Occupancy ype
2000
2020 a1
2020b2
2050a1
2050b2
Type A - egional and local sui abili y
9.0%
10.6%
8.6%
13.9%
11.0%
Type B - only egional sui abili y
18.9%
18.9%
15.4%
34.5%
22.2%
Type C - only local sui abili y
11.8%
10.2%
12.2%
6.9%
9.8%
Type D - no sui abili y
60.3%
60.3%
63.8%
44.7%
57.0%
Figu e 9- Spa ial p ojec ions o in asion wi hin Na u a 2000 si es (p o ec ed a eas o combined models, unde cu en
(2000) and u u e condi ions (2020 and 2050).
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P edomina e dynamic ype p edic ed by he combined models in each ime pe iod
wi hin Na u a 2000, was “No change” (see Fig 10 and Table 8). Fo bo h clima ic
scena ios, esponses ypes “de e io a ion o he condi ions” and “ex inc ion” dec eased
om he ime pe iod 2000-2020 o 2000-2050 and om he ime pe iod 2000-2050 o
2020-2050. Conside ing he A1 clima ic scena io, he esponse ypes “Imp o emen o
he condi ions” and “Coloniza ion”, om he ime pe iod 2000-2020 o 2000-2050, was
inc easing and hen, om he ime pe iod 2000-2050 o 2020-2050, was dec eased.
Conside ing he B2 clima ic scena io, om he ime pe iod 2000-2020 o 2000-2050
and 2000-2050 o 2020-2050, an inc easing was obse ed.
Table 8 – Pe cen age o occupancy in Na u a 2000 o each dynamic ype (no change, imp o emen o he condi ions,
de e io a ion o he condi ions, ex inc ion and coloniza ion), o he es species, o he pe iods 2000-2020, 2000-2050
and 2020-2050.
Scena ios
Dynamics
No change
Imp o emen
De e io a ion
Ex inc ion
Coloniza ion
A1B
2000-2020
82.2%
2.8%
1.2%
6.9%
6.9%
2000-2050
70.4%
5.5%
0.6%
3.9%
19.6%
2020-2050
79.3%
3.5%
0.1%
0.7%
16.4%
B2
2000-2020
84.4%
1.3%
1.6%
8.1%
4.6%
2000-2050
84.3%
2.8%
0.8%
4.4%
7.7%
2020-2050
85.9%
2.9%
0.5%
2.0%
8.7%
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4.4. CPMs, connec i i y o in ade dis ibu ions, and p o ec ed a eas
Connec i i y analysis is use ul in ecology esea ch (Vicen e e al., 2013), and i
elucida es pa e ns, he eby p o iding impo an insigh s in o he p ocesses ha d i e
plan in asions in habi a s (Mino e al., 2009). Su aces wi h high connec i i y
ep esen po en ial dispe sal co ido s and a eas wi h high connec i i y will be he ones
whe e managemen may be less e ec i e, as local con ol measu es may be
coun e ac ed by immig a ion and p opagule p essu e om neighbou ing in aded a eas
(Vicen e e al., 2013).
In his s udy, we used a connec i i y index based on he nea es neighbou s ha is
conside ed a c ude connec i i y me ic, al hough simple o ob ain. Howe e , combined
p edic i e models p ojec ions allowed o weigh he po en ial sui able a eas o he
Acacia dealba a species by e alua ing and quan i ica ion o he a ailabili y o bo h
egional condi ions and sui able local habi a .
A eas wi h high connec i i y o he species will be he ones wi h less e ec i e
managemen , as local con ol measu es may be coun e ac ed and p opagule p essu e
om neighbo ing in aded a eas (Vicen e el al., 2013). Fo he es species, connec i i y
was p edic ed o inc ease in he u u e. Al hough connec i i y o sui able a eas o he
species inside p o ec ed a eas is lowe han ac oss he ull a eas (because p o ec ed
a eas coincide wi h a eas o high ele a ion) he changes in en i onmen al condi ions
p edic ed o he u u e may change he connec i i y o he species (Vicen e e al.,
2013). In conclusion, he connec i i y analysis showed ha p o ec ed a eas will
po en ially su e a high p essu e om Acacia dealba a unde u u e en i onmen al
condi ions.
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5. Conclusions
Using a combined p edic i e modeling app oach o o ecas he cu en and u u e
dis ibu ion o Acacia dealba a in No hwes Ibe ian Peninsula, as well as assessing i s
cu en and u u e con lic s wi h p o ec ed a eas, we conclude ha :
Models p edic ha he a ea wi h sui able condi ions o clima e ( egional) and
habi a /landscape (local) will inc ease in he u u e, i he clima e change
scena ios a e con i med;
The species cu en ly in ades na u e ese es in he s udy a ea, and models
p edic ha he pe cen age o occupancy will inc ease in he u u e; and
Connec i i y analysis showed ha changes in u u e en i onmen may inc ease
he connec i i y o Acacia dealba a dis ibu ion in he es a ea.
F om an applied pe spec i e, ou esul s sugges ha :
Spa ial p edic ions o he models p o ide an impo an basis o he e icien
con ol o his in asi e species in in aded a eas and o he ea ly de ec ion o
new a eas o in asion;
CMP p ojec ions can suppo measu es o p e en in asions in his egion and
p omo e c oss-bo de managemen o in asions; and
Connec i i y analyses a e impo an in in asion ecology, as hey can con ibu e
o p io i ize managemen ac ions conside ing egional dispe sal co ido s and
he p obabili y o ecoloniza ion a e local con ol measu es ha e been
unde aken.
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Annex
Annex 1: En i onmen al p edic o s selec o apply he Spea man ank co ela ion
coe icien .
Va iable
ype
P edic o s
Desc ip ion
Re e ences
ANT
Annual Mean Tempe a u e
B oennimann and Guisan, 2008
Clima e
AP
Annual P ecipi a ion
Ba ows and Mu phy- Maiscal, 2012
I
Iso he mali y
Cas o-Díez e al., 2011
MDR
Mean Diu nal Range
C ossman and Cooke, 2011
MTCM
Min Tempe a u e o Coldes Mon h
E angelis a e al., 2011
MTCQ
Min Tempe a u e o Coldes Qau e
Fé nandez e al., 2012 ; Fische e al.,
MTDQ
Mean Tempe a u e o D ies Qua e
Fische e al., 2011
MTWM
Max Tempe a u e o Wa mes Mon h
Gaikwad e al., 2011; Gallien e al., 2012
MTWaQ
Mean Tempe a u e o Wa mes Qua e
Godoy e al., 2008
MTWeQ
Mean Tempe a u e o We es Qua e
Ohlemulle e al., 2006; Pino e al., 2005
PCQ
P ecipi a ion o Coldes Qua e
T e howan e al., 2011; Thuille e al. 2005
PDM
P ecipi a ion o D ies Mon h
Vicen e e al., 2010; Vicen e e al., 2011
PDQ
P ecipi a ion o D ies Qua e
Vicen e e al., 2013a; Vicen e e al., 2013b
PS
P ecipi a ion Seasonali y
Vicen e e al., 2013c; Vicen e e al., 2013d
PWM
P ecipi a ion o We es Mon h
Vicen e e al., 2013ª
PWaQ
P ecipi a ion o Wa mes Qua e
PWeQ
P ecipi a ion o We es Qua e
TAR
Tempe a u e Annual Range
TS
Tempe a u e Seasonali y
Land co e
pA L
% co e o a able land (pe 1 km²)
(Landscape
pA S
% co e o a i icial s ands (pe 1 km²)
Chy y e al., 2008
composi ion)
pBiFo
% co e o b oad-lea o es (pe 1 km²)
h p://www.eu ope-aliens.o g
pCoFo
% co e o coni e o es (pe 1 km²)
g/pd /Acacia_dealba a.pd
pHAa
% co e o he e ogeneous ag icul u al a eas (pe 1 km²)
Ohlemulle e al., 2006
pMiFo
% co e o mix o es (pe 1 km²)
Pino e al., 2005
pOs
% co e o open spacie wi h small o no ege a ion (pe 1 km²)
Vicen e e al., 2013a
pPas
% co e o pas u es land (pe 1 km²)
Vicen e e al., 2013d
pPeC
% co e o pe manen c ops (pe 1 km²)
pSH
% co e o sc ub and/o he baceous ege a ion associa ions (pe 1
km²)
Pwa
% co e o wa e (pe 1 km²)
T anspo
DensRail
Densi y o local ail ne wo k
Ca o d e al. 2011
and
DensRi
Densi y o local hyd og aphic ne wo k
Vicen e e al., 2010; Vicen e e al., 2011
hyd og aphic
DensRoad
Densi y o local oad ne wo k
Vicen e e al., 2013a; Vicen e e al., 2013b
ne wo k
Dis Rail
Dis ance o ails
Vicen e e al., 2013c; Vicen e e al., 2013d
Dis Ri
Dis ance o i e s
Dis Road
Dis ance o oads
Soil
pCamb
% o cambissoils (pe 1 km²)
FCUP
Using models and connec i i y analysis o p edic he cu en and u u e pa e ns o in asion by Sil e -wa le
(Acacia dealba a Link) in No hwes Ibe ian Peninsula
54
pFlu i
% o lu issoils (pe 1 km²)
pHis
% o his ossoils (pe 1 km²)
pLi
% o li hossoils (pe 1 km²)
pLu i
% o lu issoils (pe 1 km²)
Vicen e e al., 2013a
pPodz
% o podzossoils (pe 1 km²)
Vicen e e al., 2013b
pRank
% o anke ssoils (pe 1 km²)
pRego
% o egossoils (pe 1 km²)
Lanscape
Ed
Edge Densi y
S uc u e
GPP
Mean g oss annual p ima y p oduc i i y
Lomba e al., 2010
MPAR
Mean pe ime e -a ea a io
Vicen e e al., 2010
MPFD
Mean pa ch ac al dimensions
Vicen e e al., 2013a; Vicen e e al., 2013b
MPS
Mean pa ch size
Vicen e e al., 2013c; Vicen e e al., 2013d
MSI
Mean shape index
Vicen e e al., 2013ª
NumP
Numbe o pa ches
PSSD
Pa ch size s anda de ia ion
TE
To al Edge