Effects of disturbance and alien plants on the phylogenetic structure of riverine communities

J Veg Sci. 2021;32:e12933.  
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1 o 11
h ps://doi.o g/10.1111/j s.12933
Jou nal o Vege a ion Science
wileyonlinelib a y.com/jou nal/j s
Recei ed:16May2018
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Re ised:25July2020
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Accep ed:27July2020
DOI: 10.1111/j s.12933
RESEARCH ARTICLE
E ec s o dis u bance and alien plan s on he phylogene ic
s uc u e o i e ine communi ies
Diego Liendo1 | Idoia Biu un1 | Juan An onio Campos1 |
I zia Ga cía-Mijangos1 | Pe e B. Pea man1,2,3
©2020In e na ionalAssocia ion o Vege a ionScience
1Depa men o Plan BiologyandEcology,
Uni e si yo  heBasqueCoun yUPV/EHU,
Bilbao,Spain
2IKERBASQUEBasqueFounda ion o 
Science,Bilbao,Spain
3BC3BasqueCen e o Clima eChange,
Leioa,Spain
Co espondence
DiegoLiendo,Depa men o Plan Biology
andEcology,Uni e si yo  heBasque
Coun yUPV/EHU,Bilbao,Spain.
Email:[email protected]
Funding in o ma ion
EuskoJau la i za,G an /Awa dNumbe :
IT936-16
Co-o dina ing Edi o :Ingol Kühn
Abs ac
Ques ions: Phylogene icanalysesp o ideimpo an insigh sin hes udyo biologi-
calin asions.P e iouss udiesha eshowncon as inge ec so alienspecieson
hephylogene ics uc u eo  ecipien communi ies.In hiss udy,we ocuson wo
ipa ianplan communi ieswi hcon as ingna u aldis u bance egimes: ipa ian o -
es sand i e ba communi ies.Weaskwhe he  hesecommuni iesdi e inalpha
di e si yanddeg eeo plan in asion.Fu he ,a e hephylogene icdi e si y(PD)
ands uc u eo  hesehabi a s ela ed o hele elo plan in asion?
Loca ion: No he nSpain.
Me hods: Wede e minedle elo plan in asionin o es and i e ba  ege a ion
plo s, and calcula ed PD using mean pai wise dis ance (MPD) and mean nea es 
axondis ance(MNTD).Weappliednullmodels oanalyse hephylogene ics uc-
u eo plo s,andwe es edwhe he inclusiono alienplan specieswasassocia ed
wi hdi e encesinobse edphylogene ics uc u e.
Resul s: Ri e ba plo sexpe iencedmo ein asion han o es plo s,bu  hele elo 
in asionwasonly ela ed o hePDo na i especiesin o es s.Mos plo shad an-
domphylogene ics uc u e,wi hamino i yo plo s ending ophylogene ico e d-
ispe sionin o es sand ophylogene icclus e ingin i e ba s.MPDinc easedwi h
heinclusiono alienplan speciesin o es plo s,sugges ingphylogene ico e dis-
pe sion,whilenosuchpa e nwasde ec edwi hMNTD.MPDinc easedsligh lywi h
inc easingin asionin i e ba plo s,sugges ing educedclus e ing,whileMNTD al-
uesdec easedwi hinc easingin asion,sugges ing heopposi e end.
Conclusions: In asionbyalienplan sisdi e en iallyassocia edwi hphylogene ic
s uc u ein ipa ianhabi a swi hdi e en dis u bancele els.Ou  esul scon as 
wi h hoseo p e iouss udieso plan in asiono  ipa iancommuni ies,whichsug-
ges s geog aphic and ecological a ia ion in he ela ionship be ween in asion and
phylogene iccommuni ys uc u ein ipa iansys ems.Resea chisneeded oiden i y
hecausal ac o sunde lying his a ia ion.
KEYWORDS
aliene ec ,alienplan s,dis u bance,MNTD,MPD,no he nIbe ianPeninsula,phylogene ic
di e si y, ipa ian o es s, i e ba communi ies, ege a ionplo s
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1 | INTRODUCTION
Biodi e si y esea chonplan communi ieso en ocusesondi-
e si y a  he le el o  species, gene ally by quan i ying species
ichness and indices ha inco po a e species ela i e abundances
(Pee ,1974).Ne e heless, hebiodi e si yconcep includesaddi-
ionalcomponen sbesidesspecies ichness(Ca monae al.,2012;
Dainesee al.,2015).Among hese,phylogene icdi e si y(PD,i.e.,
heex en o e olu iona yhis o y ep esen edbyspecieswi hin
plan assemblages)isakeydesc ip o o plan communi ydi e si y
andisnowwidelyused(P ocheşe al.,2008;Benne e al.,2014;
B unbje ge al.,2014;Kusumae al.,2018).Inpa icula ,PDcan
p o ideinsigh sin oecologicalp ocesses ha a ec  heassem-
bly o  plan communi ies(Ca ende -Ba es e  al.,2009;Cado e
e  al.,2010; Pu schkee  al., 2013).Acco ding o hisapp oach,
co-occu ingspeciesinplan communi iescanexp ess andom,
o e dispe sedo clus e edphylogene icpa e ns,dependingon
whe he communi iesa e andomlyassembled,o con ainmo e
o  ewe dis an  ela i es hanexpec edunde  andomassembly.
Clus e edphylogene icpa e nsa ein e p e ed oindica e ha 
en i onmen al il e ingselec sspecies ha sha esimila  ai sand
ecological equi emen s(B unbje ge al.,2014;Lososo áe al.,
2015a), while o e dispe sed pa e ns sugges  he exis ence o 
compe i i eexclusion ha limi s hecoexis enceo closely ela ed
species(Webbe al.,2002).
Phylogene icanalysisp o ides impo an insigh sin he s udy
o biologicalin asion,whichisap inciplean h opogeniccauseo 
biodi e si y loss globally (Vi ousek e  al., 1997; Vilà e  al., 2006).
Speci ically, unde s anding he deg ee o which in ading species
ep esen no ele olu iona yo iginsiskey oin e ing hemecha-
nismso plan in asion(Bezenge al.,2013;Lie al.,2015;Vilàe al.,
2015). Empi ical s udies a  a wide ange o  spa ial scales p o ide
mixed e idence o  phylogene ic consequences o  plan  in asion,
wi h s udies suppo ing he associa ion o  species in asion wi h
phylogene icclus e ing(Win e e al.,2009;Lososo áe al.,2015a;
Loiola e  al., 2018), phylogene ic o e dispe sion (Ge hold e  al.,
2011;Bezenge al.,2013),o noappa en associa ion(Ca alloand
Cas o, 2017). Thesecon as ing esul sindica e heneed o  e-
sea chon heassocia iono alienplan swi h hePDands uc u e
o na i ecommuni ies,andonhowongoingan h opogenicandna -
u aldis u bancemayal e  hese ela ionships(Liendoe al.,2015;
Lososo áe al.,2015a).
Ripa ian habi a s b oadly expe ience high le els o  in asion
(Richa dson e  al.,2007; Pyšeke  al., 2010),wi h ipa ianplan 
communi ies( o es s,g a elba s, all-he bcommuni ies, i e ine
ma shes, e c.) di e ing in equency and co e  o  alien species
(Campose al.,2013;Liendoe al.,2015).Ripa iancommuni ies
di e  ina numbe o  ways ha  may in luence bo h he deg ee
o  in asion and communi y PD. Communi ies on i e  ba s a e
looded egula ly,in enselyando e longpe iodsasa esul o 
hei p oximalposi ionon he i e bed(Kalníko áe al.,2018).In
con as , ipa ian o es sa ehighe  han i e ba sinele a ion
ela i e o he channel cen e and, consequen ly, a e looded
du ing sho e  pe iods, and expe ience lowe  low and physical
dis u bance.
In hiss udy,weadd esshowplan in asionandna u aldis-
u banceco a ywi hPDandspeciescomposi ioninplan com-
muni iesin wophysicallyclose ipa ianhabi a s, ipa ian o es 
andni i ied i e ba .These wohabi a  ypesexpe iencena u al
loodingo di e ingin ensi iesanddu a ionsdue o hei dis inc 
opog aphic loca ions and p oximi y o channel cen e. We exam-
ine he ela ionshipsamonghabi a  ype,alienin asion andPD
usingplo da aonplan communi ycomposi ion.Weask:(a)do i-
pa ian o es sand i e ba plan communi ies,wi h hei con as -
ingna u aldis u bance egimes,di e inalphadi e si ypa e ns
anddeg eeo plan in asion?and(b)a e hePDands uc u eo 
hesehabi a s ela ed o hele elo plan in asion?Weexpec 
ha  hemo e equen andin ense loodingexpe iencedby i e 
ba s leads communi ies in his habi a o exhibi g ea e phyloge-
ne icclus e ing han ipa ian o es sasa esul o en i onmen al
il e ing, a ou ingclosely ela edspecies ha a eadap ed o he
dis u bance egime(Helmuse al.,2010).Asa esul o  his,we
expec ha he alien plan s colonising i e ba s will be closely
ela ed o he esiden communi y, husinc easinga e agephy-
logene icclus e ing(Lososo áe al.,2015a).Incon as ,en i on-
men al il e ingdue odis u bancemaybelessin ensein ipa ian
o es sand, hus,compe i ionmaybemo eimpo an  o species
assemblyin hishabi a .He e,weexpec  ha alienspecieswill
also include dis an ly ela ed species. Inc easing in asion in ipa -
ian o es shouldei he ha enoassocia ionwi hobse edPDo 
accompany inc eased o e dispe sion.
2 | METHODS
2.1 | S udy a ea and da a collec ion
This s udy was conduc ed in he ipa ian o es s and he ophy e
communi ies o  ni i ied i e  ba s in se e al Can ab ian i e s o 
no he nSpain(Figu e1a).These i e sa esho ands eep,wi h
as  unningwa e sandmode a e low educ ioninsumme when
i e ba seme ge(Liendoe al.,2015;Liendoe al.,2016).Ripa ian
o es s include alde  (Alnus glu inosa) and ash (F axinus excelsio )
o es so  heassocia ionsHype ico and osaemi-Alne um glu inosae
and S egnog ammo pozoi-Alne um glu inosae (Biu un e  al., 2016;
Figu e1b).Ri e ba communi iesincludeni ophilousmac o e o-
phy ecommuni ieso  heassocia ionBiden i ondosi-Polygone um
lapa hi olii(Campos,2010;Figu e1c).Wecons uc ed heda ase 
by conside ing da a on ege a ion plo s ha we e p e iously sam-
pled ollowing hephy osociologicalme hod(B aun-Blanque ,1951)
ands o edin heVege a ion-Plo Da abaseo  heUni e si yo  he
Basque Coun y (BIOVEG; GIVD code EU-00-011; Biu un e  al.,
2012).Whenmo e hanoneplo o ei he habi a wasa ailablein
hesameUni e salT ans e seMe ca o g idcello 1km×1km,we
selec edoneo  hem andomly.Plo size anged om100 o200m2
o  ipa ian o es sand10–50m2 o  i e ba s(AppendixS1).Plo s
16541103, 2021, 1, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1111/j s.12933 by Readcube (Lab i a Inc.), Wiley Online Lib a y on [13/06/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
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wi hou  alien species we e excluded om he analyses. The inal
da ase included60 o es plo sand38 i e ba plo s om hepe-
iod1985–2015(AppendixS1),wi ha o alo 421species.Species
nomencla u e ollowsEu o+Med(2019).
2.2 | Di e si y pa e ns and le el o in asion o
o es s and i e ba s
Wecalcula ed he o alnumbe o na i eandalien axaa di e en 
axonomicle elsineachhabi a and hemeannumbe andco e o 
na i eandalien axaa plo le el.Wequan i ied hele elo plan 
in asionwi h h eeindices: henumbe o alienspeciesineachplo 
(absolu ealienspecies ichness,AR), hep opo iono alienspecies
inaplo ( ela i ealien ichness,RAR)and hep opo iono alien
speciesco e in ela ion o he o alplan co e inaplo ( ela i e
alienco e ,RAC;Ca o de al.,2012;Liendoe al.,2016).Allalien
plan species,i espec i eo  hei in asions a us,we eincludedin
he analyses. Wede e mined hes a is ical signi icance o  di e -
ences in di e si y measu es and le els o  plan  in asion be ween
ipa ian o es sand i e ba communi ieswi hS uden 's es s.
2.3 | Spa ial pa e ns
We es ed o spa ials uc u ein heda ase s o  o es and i e ba 
plo ssepa a elybygene a ingma iceso Jacca ddis ancesusing
he“ egan”Rpackage.We henconduc edMan el es so  hese
ma ices agains  he ma ix o  s aigh -line geog aphic dis ances
wi h he“ade4”Rpackage(D ayandDu ou ,2007).Innocasewas
spa ials uc u ede ec ed( andomisa ionp >0.05,n = 60 in ipa ian
o es s; andomisa ionp >0.05,n =38in i e ba s).Addi ionally,
spa ialau oco ela ion(SAC)in eg ession esidualswase alua ed
wi hMo an'sIusing heco elog() unc iono  hepackage“nc ”.To
emo ee ec so SACp io  odecisionsons a is icalsigni icance,
spa ial eigen ec o s we e gene a ed wi h he dnea neigh() and
Spa ialFil e ing() unc ionso package“spdep”(Bi andandWong,
2018).These ec o swe eadded oo dina yleas -squa es eg es-
sions, esul inginadjus edF-s a is icsande o deg eeso  eedom
o  hee ec so in e es .
2.4 | Phylogene ic di e si y and da a analyses
Aphylogene ic eewasassembled oincludena i eandalienspe-
ciesinbo hhabi a s.Somep e iouss udiesha eexcludedp e ido-
phy esbecauseo  hei ea lydi e genceand esul inglongb anch
leng hs,whichmays onglyin luenceobse edPD(VanMee beek
e al.,2014;Čeplo áe al.,2015).Howe e ,p e idophy esa ean
impo an na u alcomponen o Can ab ian ipa ian o es s(Biu un
e  al., 2016) and, consequen ly, we e included in he phylogeny
(Table 1). None heless, phylogene ic indices we e also calcula ed
excludingp e idophy es oassess hee ec o  hei inclusiongi en
hepo en ialin luenceo beingdis an ly ela edwi h espec  o he
dominan  angiospe m lo a, hei  mino  con ibu ion in i e  ba s
and heabsenceo alienp e idophy esin heda ase .Thecomple e
phylogenyincludingp e idophy escon ained413speciesandwas
cons uc edwi h he “phyloma ic” unc iono  he Rpackage“b -
anching”,using he ime-calib a edzanne2014 ee(Zannee al.,
2014)as he e e ence owhichspeciesnameswe ema ched.Ten
FIGURE 1 S udya eashowing he
lu ialne wo kand hemainci ies(a)and
exampleso a ipa ian o es (b)anda
i e ba (c)
16541103, 2021, 1, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1111/j s.12933 by Readcube (Lab i a Inc.), Wiley Online Lib a y on [13/06/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
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p e idophy especieswe emissing om his ime-calib a ed ee,
so hey we e included as o he  congene ic o  con amilial spe-
cies. Addi ionally, we andomly esol ed 26 poly omies using he
“mul i2di” unc iono  heRpackage“ape” oob aina ully esol ed
ee(Pa adisandSchliep,2019).
Indiceso PDwe ecalcula edsepa a ely o  o es sand i e ba 
plo s o  hena i especiescomponen alone,andagainincluding
bo hna i eandalienspecies( ou communi yclassesin o al).Two
complemen a yPDindiceswe ecalcula ed:meanpai wisedis ance
(MPD)andmeannea es  axondis ance(MNTD).The i s o  hese,
MPD,is hea e agepai wisedis ancebe weenspeciesinaplo o 
communi yandisameasu eo  heo e allPD(Swenson,2014).In
con as ,MNTDis hemeandis ancebe weenaspeciesandi sclos-
es  ela i einaplo (Swenson,2014).Thismeasu e ocusesondi-
e si ya  he ipso  hecommuni yphylogeny,sinceclosely ela ed
speciesmaybeecologically simila and likely expe iences onge 
in e ac ions handis an  ela i es.SmallMNTD aluesindica e he
co-occu enceo closely ela edspecies.Theseindicescanbea -
ec edbydi e encesinspecies ichness(Vellende al.,2011).Fo 
his eason,wecalcula ed hes anda disede ec sizeso MPDand
MNTD,MPDSESandMNTDSES,whicha eequi alen  o henega i e
o  hene  ela ednessindexandnea es  axonindex(Webbe al.,
2002), espec i ely,andindependen o species ichness(Pa oine
and Bonsall, 2011). S anda disa ion was done by compa ing he
obse edMPDandMNTD alueswi h he alueso 999 andom
communi iesusing he“ axa.labels”nullmodelin he“pican e”pack-
age,whichshu leslabelso all axa in hephylogene icdis ance
ma ix(Kembele al.,2010), husholdingcons an species ichness
o plo sand hespecies u no e among hem.TheindicesMPDSES
and MNTDSES we e calcula ed as (MPDOBS–MPDRAND.MEAN)/SD
MPDRAND.MEAN,whe eMPDOBSis heobse edMPD,andMPDRAND.
MEAN and SDMPDRAND.MEANa e hemeanands anda dde ia iono 
expec edMPD om he andomisedcommuni ies, espec i ely.We
used hes anda disedindices o es  o non- andomphylogene ic
s uc u eo communi ies.Nea -ze o aluesindica e andomphylo-
gene ics uc u e,while aluesbelow−1.96o abo e+1.96 indica e
signi ican  clus e ing and o e dispe sion, espec i ely, wi h a 5%
Type-Ie o  a e(Lososo áe al.,2015a).Weusedspeciesp esence/
absence in calcula ing he aw and s anda dised indices in o de o
conside he en i e phylogeny and a oid bias caused by species dom-
inance(Loiolae al.,2018).
The s anda dised e ec  sizes o  MPD and MNTD we e used
o c ea e boxplo s o isualise he o e all phylogene ic s uc u e
a  hehabi a le elwi handwi hou inclusiono aliens,wi h hese
o e alldi e ences es edbymeanso  es s. These s anda dised
e ec sizeswe ealsousedin helinea  eg essionsassessing he
ela ionship be ween he PD o  he na i e communi ies and he
le elo plan in asion(AR,RARandRAC).Thele elo plan in a-
sionwaslog- ans o med(na u alloga i hm) oimp o eno mali yo 
esiduals.Finally,a  heplo le el hee ec o  heinclusiono alien
speciesonphylogene ics uc u e(i.e.,“aliene ec ”)wases ima ed
o eachplo sepa a elyas hedi e encebe ween hes anda dised
index calcula ed wi h na i e and alien species and he index calcu-
la edwi hna i especiesalone(Win e e al.,2009;QianandSandel,
2017).Thisaliene ec was eg essedagains  hes anda disedMPD
andMNTDo  hena i ecomponen o  ipa ian o es and i e ba 
plo s.Theassump ionso no mali yo  esidualsandhomoscedas-
ici ywe e es edwi h heShapi o-Wilkss a is icandplo so  e-
siduals.Allanalyseswe epe o medusingR e sion3.5.2(RCo e
Team,RFounda ion o S a is icalCompu ing,Vienna),andRcodeis
includedinAppendixS2.
3 | RESULTS
3.1 | Di e si y pa e ns and le el o in asion
A o alo 338 axa(includingsubspeciesand axaiden i ieda genus
le el)a e ep esen edin he60 o es plo s.O  hese axa,285a e
na i eand53a ealien.Asmanyas112 axa(33.1%)occu inonly
oneplo ,including85na i eand27alien axa.Themos  equen 
na i e species (occu ing in mo e han 50 plo s) a e A. glu inosa
(57),Hede a hibe nica(56),B achypodium syl a icum(55),Hype icum
and osaemum (54), Ca ex pendula (53) and F. excelsio (52).The
mos  equen alienspecies(occu inginmo e han20plo s)a e
C ocosmia × c ocosmii lo a(22),Pla anus hispanica(22)andRobinia
pseudoacacia (22). A o al o  238 axa (including subspecies and
hoseiden i ieda  hegene icle el)occu in he38 i e ba plo s,
including176na i eand62alien axa.Asmanyas78 axa(32.8%)
appea inonlyoneplo ,including58na i eand20alien axa.The
mos  equen na i especies(occu inginmo e han19plo s)a e
Ly h um salica ia (25), Pe sica ia lapa hi olia (25), P. maculosa (25),
Calys egia sepium(21),U ica dioica(20)andRumex ob usi olius(20).
Themos  equen alienspecies(occu inginmo e hannineplo s)
a e Paspalum dis ichum (25), Cype us e ag os is (23), Echinochloa
c us-galli (15), Bidens ondosus (12), Cype us longus (10), Digi a ia
sanguinalis(10)andE ige on lo ibundus(10).A o alo 131species
a e ounda leas onceinbo hhabi a s,which ep esen 57%o 
he i e ba da ase and40%o  he o es da ase .O  hese131
sha edspecies,59o  hemoccu in<5%o  o es plo s,includinga
TABLE 1 Meanplo  aluesands anda dde ia iono plan 
g oupsa di e en  axonomicle elsin ipa ian o es s(n =60)and
i e ba communi ies(n =38)o  heno he nIbe ianPeninsula
Fo es s Ri e ba s
Mean SD Mean SD
Angiospe mspecies 41.2 12.2 25.8 16.4
Gymnospe mspecies 0.1 0.2 0 0
P e idophy especies 4.2 20.5 0.6
Plan  amilies 30.8 6.3 14.4 7.5
Plan gene a 41.7 10.7 22.8 13.9
Exclusi elyalien
amilies
0.8 10.3 0.5
Exclusi elyaliengene a 1.9 1.5 3.2 2.5
16541103, 2021, 1, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1111/j s.12933 by Readcube (Lab i a Inc.), Wiley Online Lib a y on [13/06/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
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numbe o annualand/o ni ophilousspeciesmo e ypicalo  i e 
ba s ha canoccasionallybe oundin ipa ian o es s(Appendix
S1).On heo he hand,61o  hesha edspeciesoccu in<5%o 
i e ba plo s,manyo  hembeinghe baceouso woodyspecies
cha ac e is ic o  ipa ian o es s ha  a e occasionally p esen  in
i e ba s(AppendixS1).Nosigni ican  ela ionshipbe weennum-
be so na i eandalienspeciesexis sin ipa ian o es s( P = 0.023;
d = 36; p =0.86),whils  he eisaposi i eandsigni ican  ela ion-
shipbe ween henumbe o na i eandalienspeciesin i e ba s
( P =0.35;d =58;p =0.028).
The mean numbe  o  angiospe m and p e idophy e species,
plan  amilies,andplan gene aishighe in o es plo s(Table1).
The mean numbe o  alien amiliesis also sligh ly highe in o -
es  plo s ep esen ing, on a e age, 2.6% o  plan  amilies com-
pa ed o2.1%o  amilies in i e  ba plo s. On he o he hand,
hemeannumbe o exclusi elyaliengene aishighe in i e ba 
plo s ep esen ing,ona e age,14.0%o plan gene acompa ed
o 4.6% in o es  plo s. When da a a e pooled wi hin habi a s,
o es s and i e  ba s show a simila  pe cen age o  alien ami-
lies (12.5% and 13.6%, espec i ely), whils  a  he genus le el
i e  ba s ha bou  subs an ially mo e alien gene a han o es s
(23.4% and 16.2%, espec i ely). Exclusi ely alien amilies in i-
pa ian o es s include Commelinaceae, My aceae, Mo aceae,
Balsaminaceae, Hyd angeaceae, Juglandaceae (wi h wo gene a,
Juglans and P e oca ya), Pla anaceae, Ga yaceae, Cannaceae
andBignoniaceae.Exclusi elyalien amiliesin i e ba sinclude
Ac inidiaceae,Mo aceae,Pla anaceaeandJuglandaceae.
Ri e ba communi iesha esigni ican ly highe alien ichness,
asRACandRAR, han do ipa ian o es s,bu  hey dono di e 
signi ican lyinalienco e (Table2).Fo es plo sshowposi i emean
MPDSES and MNTDSES alues, indica i e o  a endency owa ds
phylogene ico e dispe sion,while i e ba plo sexhibi nega i e
mean alues o bo hindices,indica i eo a endency ophyloge-
ne icclus e ing(Figu e2).Howe e , heses anda disedindicesa 
he plo  le el e eal ha  he majo i y o  plo s show no e idence
o ei he clus e ingo o e dispe sion(AppendixS3).None heless,
somedi e encesexis be ween hesehabi a s.Bo hMPDSES and
MNTDSESindica e ha  hena i ecomponen o  o es plo sissig-
ni ican lymo eo e dispe sed han ha o  i e ba plo s(Table3,
Figu e2).In ac ,30%o  o es plo sshowMPDSES alues ha in-
dica esigni ican phylogene ico e dispe sion,wi hasligh lylowe 
pe cen age o MNTDSES(AppendixS3).No o es plo isphyloge-
ne icallyclus e ed.On heo he hand,8%o  i e ba plo sp es-
en  MPDSES alues ha  indica e signi ican  clus e ing and his
pe cen ageinc eases o29% o MNTDSES(AppendixS3).Nei he 
indexindica esphylogene ico e dispe sion o any i e ba plo .
Whenp e idophy esa eexcluded om heanalyses,no o es plo 
is signi ican ly o e dispe sed wi h espec  o MPDSES, whils  he
pe cen ageo o e dispe sedplo s ega dingMNTDSES dec eases o
15%(AppendixS3).As o  i e ba plo s, he eisaninc easein he
pe cen ageo signi ican lyclus e edplo s o bo hMPDSES( om8%
o36%)andMNTDSES( om29% o52%).
3.2 | Le el o in asion and communi y
phylogene ic di e si y
Phylogene icdi e si yo  hena i ecomponen o  i e ba plo s,in
e mso bo h heMPDSESandMNTDSESindices,isno signi ican ly
ela ed o hele elo plan in asion(AppendixS4).To hecon a y,
aweaklynega i ebu signi ican  ela ionshipeme gesin o es plo s
be weenMNTDSESand hele elo plan in asion,ei he measu ed
asAR,RARo RAC(AppendixS4),indica ingdec easingle elso 
plan in asionin o es plo swi hhighe MNTDSES alues. In i e
ba s,despi esome eg essionsyieldingsigni ican s a is icale ec s
o someo  hespa ialeigen ec o s,nei he MPDSESno MNTDSES
we esigni ican ly ela ed oany a iable ep esen ing hele elo 
plan in asion(allp >0.05;AppendixS4).
3.3 | E ec o alien species on phylogene ic
ela edness
The eisasigni ican o e allinc easeinphylogene ico e dispe sion
in o es  plo sas indica edbyMPDSES when alien species a e in-
cludedin heanalysis(Figu e2,Table3).This ansla esin oanaddi-
ional10%inc ease( om30% o40%)in he o es plo sexhibi ing
signi ican o e dispe sion(AppendixS3).On hecon a y,nosuch
signi ican changeisobse edin heanalysiso MTNDSES.Simila 
TABLE 2 Mean plo alues and SDo  hele elo plan in asion
and he lo is icdi e si yindicesin ipa ian o es sand i e ba 
communi ieso  heno he nIbe ianPeninsula
Fo es s Ri e ba s
Fo es s s.
i e ba s
Mean SD Mean SD ( es )
Alien
co e (%)
20.64 28.02 30.33 28.10 −1.66(n.s.)
Na i e
co e (%)
245.16 77.15 103.7 46.63 11.32*
To al
co e (%)
265.80 72.05 134.03 54.91 10.23*
RAC(%) 8.21 11.05 21.34 17.42 −4.15*
Alien
ichness
(AR)
2.97 2.15 6.55 5.25 −3.99*
Na i e
ichness
43.10 12.36 19.79 13.98 8.41*
To al
ichness
46.07 12.59 26.34 16.59 6.27*
RAR(%) 6.84 5.11 26.97 14.23 −8.38*
Bon e onico ec ion o mul iplecompa isonswasapplied,
(αBon =0.00625).
Abb e ia ions:n.s.:non-signi ican di e ences;RAC, ela i ealien
co e ;RAR, ela i ealien ichness.
*p < 0.001.
16541103, 2021, 1, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1111/j s.12933 by Readcube (Lab i a Inc.), Wiley Online Lib a y on [13/06/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License

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esul swe eob ainedwhenp e idophy esa eexcluded(Appendix
S3).Thepe cen ageo  i e ba plo swi hsigni ican clus e ingin
MPDSESdec easesbysixpe cen wi h heinclusiono alienspe-
cies(AppendixS3)al hough hisisno signi ican (Figu e2,Table3).
The eisasigni ican o e allinc easeinphylogene icclus e ing,as
indica edbyMNTDSES,whenalienspeciesa eincluded(Figu e2,
Table3),which ansla esin oaninc easein heincidenceo clus-
e ingin i e ba plo sby5%(AppendixS3).Whenp e idophy es
a eexcluded, hepe cen ageo plo swi hsigni ican clus e ingin
MPDSESdec easesby25%wi h heinclusiono alienspecies,whils 
o MNTDSES he eisadec easein heincidenceo clus e ingby2%
(AppendixS3).
Simila ly,plo ing heinc emen ale ec o inclusiono aliens
agains  heMPDSESandMNTDSESo  hena i ecomponen illus-
a es hee ec o alieninclusiononPD(“aliene ec ”;Figu e3).
In o es  plo s, posi i e alien e ec s inc ease wi h inc easing
MPDSESo  hena i ecomponen (i.e.,wi hinc easingo e dis-
pe sion);mo e-o e dispe sedplo sa ee enmo eo e dispe sed
wi h headdi iono aliens(Figu e3a).Addi ionally, heMPDSES is
signi ican ly ela ed o healiene ec (ß1 =0.148;F1,58 =17.57;
p <0.001;AppendixS4).Theanalogouse ec o alieninclusion
o  i e ba da a,inwhichMPDSES alues a e p edominan ly neg-
a i e,isnon-signi ican .Whenp e idophy esa eexcluded om
hephylogene ic ee hee ec o alieninclusionin o es sbe-
comesnon-signi ican ,bu  a s ong inc ease o  healien e ec 
wi hdec easingMPDSESisde ec edin i e ba s,i.e., hemo e
clus e ed hena i ecommuni yis, hes onge  heposi i ee -
ec o alienspeciesisin hephylogene ics uc u e(AppendixS5).
On heo he hand, hee ec o  heinclusiono aliensin o es 
and i e ba plo sleads oli lechangeinMNTDSES when his
index o na i especiesisnega i e(i.e.,inphylogene icallyclus-
e edna i ecommuni ies).In i e ba plo s,howe e ,i becomes
inc easinglynega i ewhenMNTDSESo na i ecommuni iesin-
c eases owa dsze o(i.e.,inless-s onglyclus e edcommuni ies;
Figu e3b).Addi ionally, heMNTDSESissigni ican ly ela ed o he
aliene ec (ß1 =−0.192;F1,36 =6.24;p =0.018;AppendixS4).
Whenp e idophy esa eexcluded om heanalyses, he eg es-
siono aliene ec onMNTDSESisalsosigni ican in i e ba plo s
(AppendixS5).
FIGURE 2 Boxplo sshowing hemedianand hedispe siono  hes anda disedmeanphylogene icdis ance(MPDSES,le )and he
s anda disedmeannea es  axondis ance(MNTDSES, igh )in ipa ian o es sand i e ba communi ies.N:na i ecomponen ;T:whole
communi y(na i e+alienspecies).Le e sindica elacko signi ican di e encesa e applying heBon e onico ec ion o mul iple
compa isons(αBon =0.00625;seeTable3)
TABLE 3 T es sbe ween hena i eand o al(na i e+alien)
componen so  ipa ian o es sand i e ba swi h espec  o
hes anda disedmeanphylogene icdis ance(MPDSES)and he
s anda disedmeannea es  axondis ance(MNTDSES)
MPDSES MNTDSES
Fo es
To al s. na i e 6.46*−0.03(n.s.)
Ri e ba
To al s. na i e 2.37(n.s.) −4.45*
To al
Fo es  s. i e ba 11.60*12.10*
Na i e
Fo es  s. i e ba 11.78*9.88*
Bon e onico ec ion o mul iplecompa isonswasapplied
(αBon =0.00625).
Abb e ia ion:n.s.,non-signi ican di e ences.
*p < 0.001.
16541103, 2021, 1, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1111/j s.12933 by Readcube (Lab i a Inc.), Wiley Online Lib a y on [13/06/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
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4 | DISCUSSION
Ou esul s a e consis en wi h p e ious esea ch showing ha
i e ba communi iesa eno ablymo ein aded han ipa ian o -
es s,wi h alueso in asionle elsimila  o hoseob ained o bo h
habi a salongA lan ic i e s(TabacchiandPlan y-Tabacchi,2005;
Campose al.,2013).Thehighe le elo in asionobse edin i e 
ba communi iesisp obably he esul o  hes onge hyd ological
dis u bance heyexpe ience ela i e o ipa ian o es (Da ise al.,
2000).Ri e ba seme geseasonallyinla esp inganda ecolonised
p ima ilybyannualspecies h oughseeddispe sal(Assini,2001).By
la eau umn oea lywin e , his ege a ionisalmos  o ally emo ed
bywa e  lowand,consequen ly,a a ie yo speciesmaycolonise
in a low-compe i ion en i onmen  (Hobbs and Huenneke, 1992).
Addi ionally, i e ba sin hes udya eaa econcen a eda lowel-
e a ionsnea humanse lemen s(Liendoe al.,2016).Asa esul ,
hey ecei eplen i ulnu ien s,especiallyni ogen,allowing o  he
es ablishmen o oppo unis icni ophilousalien he ophy es,such
as Ama an husspp.,Xan hium s uma ium and B. ondosus(Campos
e al.,2013;BolpagniandPio i,2015).
4.1 | Habi a ype and phylogene ic s uc u e
We oundsigni ican di e encesinMPDSESandMNTDSES be ween
ipa ian o es sand i e ba communi ies,apa e n ha isconsis -
en wi hp e ious indingsina a ie yo habi a so Cen alEu ope
andSou hA ica(P ocheşe al.,2015;Lososo áe al.,2015a).These
di e encesa eexp essedino e allnega i e alues o bo hPDin-
dicesin i e ba plo sandbyo e allposi i e aluesin o es plo s
(Figu e2).Thisindica es ha ,asexpec ed, hena i ecomponen 
is p edominan ly clus e ed in i e ba plo s and o e dispe sed in
o es plo s.Phylogene icallyclus e edpa e nsa ealso epo ed
omna u alhabi a s,suchascoas aldunes(B unbje ge al.,2014)
and oman h opogenichabi a ssuchasu banplan communi ies
(Čeplo á e  al., 2015). Phylogene ic clus e ing is consis en  wi h
en i onmen al il e ingd i enbydis u bance(B unbje ge al.,2012;
B unbje ge al.,2014;Lososo áe al.,2015a),whichwouldselec 
closely ela edspecies om ewlineages ha sha esimila ecologi-
cal equi emen s(Čeplo áe al.,2015),assuming ai conse a ism
ac oss hephylogene ic ee(bu seeGe holde al.,2015).Incon-
as , equen o e dispe sedphylogene ics uc u ein o es plo s
is consis en wi h less in ense hyd ological dis u bance and g ea e
habi a s abili y han oundon i e ba s.In hese o es s, hismay
p omo ecoexis enceo species omdis an lineages ianichedi -
e ences(Ki agawae al.,2015;Lososo áe al.,2015a).Addi ionally,
habi a  age has also been linked wi h he phylogene ic s uc u e
o plan communi ies(Lososo áe al.,2015b).In his ega d,o e -
dispe sed phylogene ic s uc u e may p e ail in his o ically olde
communi ies in which bio ic in e ac ions may be s onge han in
ecen lydis u bedcommuni ies.Fu he , o es s,andespecially i-
pa ian o es sinou s udya ea,ha bou se e alspeciesbelonging
ophylogene icallyoldanddis inc lineages,suchasp e idophy es,
which in luences phylogene ic pa e ns o  hese communi ies by
con ibu ing oPD(Lososo áe al.,2015b;Šipoše al.,2020).
Despi e i e ba communi ies being mo e in aded and phylo-
gene icallylessdi e se han ipa ian o es s,we indli leasso-
cia iono  hePDo  hena i ecomponen o ei he habi a wi h
le elso plan in asion.Wehadexpec edanega i e ela ionship
be weenPDandle elo speciesin asion(Lososo áe al.,2015a).
Only heMNTDSES alueso  o es plo sa eweakly, houghsigni i-
can ly,nega i ely ela ed o hele elo plan in asion(Appendix
S4).Thismeans ha ,in ipa ian o es s, hoseplo swhicha ephy-
logene icallymo edi e sea  he ipo  hephylogene ic ee(i.e.,
wi hspeciesbelonging odi e en gene a)a elesscolonisedby
alienplan s.Thus,ou  esul sa eonlypa lyconsis en wi hp e i-
ous s udies in which plan communi ies wi h g ea e phylogene ic
clus e ingexpe ienceinc easedin asionbyalienspecies(Ge hold
e al.,2011;Lososo áe al.,2015a;Nge al.,2018).Theses udies
epo  he ela ionshipbe weenPDandin asionac ossa a ie y
o spa ialex en sand esolu ions.Howe e ,ou s udywascon-
duc ed wi h small ege a ion plo s a a limi ed spa ial ex en and
FIGURE 3 Rela ionshipbe ween healiene ec (SES o –SESna )and heSES alueso  hemeanphylogene icdis ance(MPD;a)and
meannea es  axondis ance(MNTD;b)calcula ed o  hena i ecomponen in ipa ian o es sand i e ba communi ies.SES,s anda dised
e ec sizes.Fo pa ame e so  hesigni ican  eg essions,seeAppendixS4
16541103, 2021, 1, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1111/j s.12933 by Readcube (Lab i a Inc.), Wiley Online Lib a y on [13/06/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
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wi hinjus  wocommuni y ypes.Thescopeo habi a  a ia ionin
as udycanin luenceobse ed ela ionshipsbe weenin asionin-
ensi yandphylogene iccommuni ys uc u e(Loiolae al.,2018),
and hismaybe hecauseo ou mixed esul s,wi haninc easein
in asionwi hinc easingphylogene icclus e ingonlyde ec ed o 
MNTDSESo  ipa ian o es plo s.
4.2 | Alien in asion, phylogene ic s uc u e and
scope o habi a a ia ion
We ound ha  hele elo in asionbyalienspeciesisassocia ed
wi hphylogene ics uc u eo  ipa ian o es sand i e ba com-
muni ies in con as ing ways.
Fi s ,inclusiono alienplan sinanalysiso  o es plo da asig-
ni ican lyinc easedo e allo e dispe sionasindica edbyMPDSES,
which ansla edin oa en-pe cen inc easeo  o es plo sexhib-
i ing signi ican  o e dispe sion, while no signi ican  change was
de ec edinMNTDSES(Figu e2,AppendixS3).This esul sugges s
ha  alien plan s in ading no he n Ibe ian ipa ian o es s ep-
esen  phylogene ically dis an  amilies no  p esen  in he na i e
lo a(Table1).These amiliesincludespeciessuchasT adescan ia
luminensis(Commelinaceae),P. hispanica(Pla anaceae)o Impa iens
bal ou ii(Balsaminaceae) ha inc ease hemeanphylogene icdis-
anceamongallspeciesin he o es plo s,inc easingphylogene ic
o e dispe sioninin adedplo s(Figu es2aand3a).Plo swi h el-
a i elys ongphylogene ico e dispe siono na i esmayp o ide
en i onmen al o bio ic condi ions ha p omo e in asion p ima ily
bynon- ela edand unc ionallydis inc  axa,pe hapsdue oexis -
ingle elso compe i iono niche illing, u he inc easing hele el
o o e dispe sion.Elsewhe einEu opealienspecieses ablishmen 
caninc easeo e dispe sionin o es communi ies(Lososo áe al.,
2015a). Despi e his gene al pa e n, Lososo á e  al. (2015a) also
showed ha allu ial o es sp esen con as ingpa e ns,wi hno
signi ican e ec due o heinclusiono alienspeciesonMPDSES,
bu inc easedclus e ingsignalledby endsinMNTDSES. These con-
as ing esul smayindica e ha  hein luenceo alienplan in asion
onphylogene ics uc u eisno uni o mac ossscaleso  ege a ion
assessmen .S udiesemployingsmallplo s ha co espondclosely
o hescalea whichbio icin e ac ionsoccu (Ca bonie al.,2013)
andwhe ealienspeciesmigh a ec na i especiesmos nega i ely
(S ohlg ene al.,1999;Ge holde al.,2011)mayp esen di e en 
phylogene ic pa e ns han s udies wi h la ge plo s ha con ain
subs an ial en i onmen al he e ogenei y. Al e na i ely, communi-
iesmay a yin hein luenceo pa icula  ai s ha a eac edon
byen i onmen al il e s,andin luen ial ai smayp esen di e ing
le elso phylogene icconse a ismindi e ing lo asandcommu-
ni ies.Thissugges s heneed o addi ional esea ch,including ield
expe imen s, ode e minekey unc ional ai s ha in luencecom-
muni y composi ion and he deg ee o which hose ai s demon-
s a e phylogene ic conse a ism.
Second,inclusiono alienspeciesinanalysiso  i e ba plo s
didno ha e asigni ican e ec  on heo e all alueso MPDSES,
al hough hepe cen ageo plo swi hsigni ican lyclus e eds uc-
u e dec eased by 6% (Figu e 2, Appendix S3). On he con a y,
inclusiono alienspeciesp oducedaweak,signi ican dec easein
MNTDSES,wi ha5%inc easein heincidenceo clus e inga  he
plo le el.The esponseo MPDSESsugges s ha onlya ewalien
species colonising i e  ba s come om majo  lineages wi h li le
ep esen a ion in he na i e communi y (Qian and Sandel, 2017),
asMPDSESmeasu es heo e allPDamongallspeciesin hecom-
muni y.In ac ,less hanoneplan  amilyis,ona e age,exclusi ely
aliena  heplo le el(Table1).Incon as , heinc easein hedeg ee
o phylogene icclus e ing egis e edwi hMNTDSES sugges s ha
alien plan species es ablishing in i e ba communi ies ha e some
close(congene ico con amilial) ela i esin hena i especiesas-
semblages,as hisindex ocuseson he ipso  hephylogene ic ee
(Swenson,2014).Thisispa lycon i medby he ac  ha in i e 
ba s he eisaconside ablenumbe o exclusi elyalienplan gene a
pe plo ,apa e nno  ounda  amilyle el(Table1).This a ia ion
couldmean ha aliengene abelong o amiliesal eadyp esen in
hena i e lo a(especiallyAs e aceaeandPoaceae),so ha  hee -
ec o alienspecieson hephylogene ics uc u eo  i e ba sis
close o he ipso  hephylogene ic ee, owhichMNTDSES is mos
sensi i e.Asimila pa e nhasbeen epo ed o plan communi-
ies along clima e g adien s in Cali o nia (Qian and Sandel, 2017)
andsugges s hep eadap a ionhypo hesis,whichs a es ha alien
specieswi haclosena i e ela i eha ebe e chanceso es ab-
lishmen (Schlaep e e al.,2010;Cado ee al.,2018).Inou case,
gi en he in ense hyd ological dis u bance o which i e ba s a e
subjec ed, hesealienspecieswouldincludespecieswi had an a-
geous ai s(annual, as -g owing,seed-dispe sedspecies;Campos,
2010)andecological equi emen ssimila  o hoseo na i especies,
aslikelyoccu samongAma an husspp.,E ag os isspp.,E ige on spp.
and Paspalum spp.
4.3 | E ec s o p e idophy e inclusion/exclusion
Finally,wehighligh  he implica ionso  p e idophy e inclusion on
obse edpa e ns.When heseplan sa eexcluded om heanaly-
ses,PDin o es plo sis educedandalienplan sha enosigni i-
can e ec onphylogene ics uc u e,measu edei he asMPDSES
o  MNTDSES (Appendix S5). The signi ican  posi i e ela ionship
be weenaliene ec andinc easingMPDSES(i.e.,inc easingo e d-
ispe sion,PD)o  o es plo sonlyappea swhenp e idophy esa e
included.Aliensdono inc easePDwhenp e idophy esa eomi ed,
which sugges s ha alien species a e dis ibu ed on he phylogene ic
eesimila ly ona i enon-p e idophy es.Theinclusiono p e ido-
phy esallowsde ec iono asigni ican e ec o alieninclusionon
MPDSES,pe hapsbecauseo  heg ea e PDin hemo einclusi e
da ase .Sincenoaliensa ep e idophy es, hise ec onMPDSES is
ullya ibu able oalienangiospe ms.
In con as  o o es plo s,alien plan sin i e ba plo s e-
duce phylogene ic clus e ing mo e in s ongly clus e ed plo s
(i.e.,s onge posi i e e ec s o  aliensonPD a  mo e-nega i e
16541103, 2021, 1, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1111/j s.12933 by Readcube (Lab i a Inc.), Wiley Online Lib a y on [13/06/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
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LIENDO E aL.
MPDses alues) haninweaklyclus e edones.Thisis e lec ed
in hep edominance o  i e ba  poin sin heuppe -le quad-
an and henega i eslopeo  he eg ession(Figu e3a,Appendix
S5A).Alienslead oamo e- andomphylogene ics uc u einplo s
wi hclus e edspeciesdis ibu ions,asmeasu edbyMPDSES,bu 
dono subs an iallya ec phylogene ics uc u ewhenisal eady
andom.Incon as ,aliensinc ease clus e ingin i e ba plo s
(g ea e nega i ealiene ec )whenMNTDSES aluesa enea ze o
(i.e.,lessclus e ing;Figu e3b),meaningaliensinc easeclus e ing
a  he e minal b anches o  he phylogene ic ee. When p e i-
dophy esa eexcluded, his end emainssigni ican (Appendix
S5B).Thus,ou  esul sdono  a ywi h heexclusiono p e ido-
phy es,demons a ingconsis en suppo  o e ec so alienson
PDin heda a om i e ba s.P e idophy esshouldbeincludedin
analysiso p e idophy e- ichcommuni iesino de  ounde s and
how alienspeciesmayin luence pa e nso  PDand unde lying
ecological p ocesses.
5 | CONCLUDING REMARKS
Thiss udy epo sda a om ege a ionplo sin wohabi a  ypes,
ipa ian o es sand i e ba  communi ies, ha di e inle elso 
na u al dis u bance wi hin he same ecosys em. I  o e s insigh 
on hepo en iale ec so alienplan soncommuni yphylogene ic
s uc u ein ipa ianhabi a s.Thesehabi a sdi e ma kedlyinle el
o plan in asion,whichshowsanuanced ela ionshipwi h hePD
o  hena i ecomponen o  he ege a ion.Alienplan sin luence
indiceso phylogene ics uc u eo bo hhabi a s,albei di e en ly.
Inaddi ion,ou  esul sdi e  omp e ious esea ch ha wasbased
onda a omwidelydi e ingcommuni y ypes,especially ega ding
ipa ian o es s,whe eMPDSES is he phylogene ic index ha sig-
ni ican lychangesdue o heinclusiono alienplan species.Fu he 
esea chshouldin es iga e he ela ionshipbe ween hescopeo 
habi a  a ia ionand hee ec so alienspeciesonobse edindi-
ceso di e si y,and he oleo p e idophy esinalienin asiono 
p e idophy e- ich plan communi ies.
ACKNOWLEDGEMENTS
Wewouldlike o hankMai eSolísandMe cedesHe e a o  hei 
helpwi h he ieldwo kandMa aTo ca o me hodologicalad ice
and aluable commen s on he manusc ip .
AUTHOR CONTRIBUTIONS
DL,IB,IGM,JACandPBPconcei edanddesigned he esea ch;
DLandPBPconduc ed heanalysesandled hew i ing;allau-
ho s discussed he esul s and c i ically commen ed on he
manusc ip .
DATA AVAILABILITY STATEMENT
Ripa ian o es and i e ba  ege a ionplo sused in hep esen 
wo ka eincludedin heVege a ion-Plo Da abaseo  heUni e si y
o  heBasqueCoun y, egis e edin heGlobalIndexo Vege a ion
Da abases(GIVDcode:EU-00-011;Biu une al.,2012).Theya e
a ailableinAppendixS1.Addi ionally,Rcode o  hecalcula iono 
hePDindicesa eincludedinAppendixS2.
ORCID
Diego Liendo h ps://o cid.o g/0000-0002-4816-4093
Idoia Biu un h ps://o cid.o g/0000-0002-1454-0433
Juan An onio Campos h ps://o cid.o g/0000-0001-5992-2753
I zia Ga cía-Mijangos h ps://o cid.o g/0000-0002-6642-7782
Pe e B. Pea man h ps://o cid.o g/0000-0002-0794-101X
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