Fo es s2021,12,1730.h ps://doi.o g/10.3390/ 12121730www.mdpi.com/jou nal/ o es s
A icle
E ec so A o es a iononPlan Di e si yandSoilQuali yin
Semia idSESpain
Ca menSegu a
1
,Ma íaN.Jiménez
2
,EmiliaFe nández‐Ondoño
3
andF anciscoB.Na a o
4,
*
1
Depa men o Sus ainableAg icul u eSciences,Ro hams edResea ch,No hWyke,
Okehamp on,De onEX202SB,UK;ca men.segu a‐qui an e@ o hams ed.ac.uk
2
Depa men o Bo any,Facul yo Pha macy,CampusdeCa uja,Uni e si yo G anada,
18071G anada,Spain;mnoelia@ug .es
3
Depa men o SoilScienceandAg icul u alChemis y,Facul yo Science,Uni e si yo G anada,
C/Se e oOchoa,s/n,18071G anada,Spain;[email p o ec ed]
4
A eao Ag icul u eandEn i onmen ,Ins i u eo Ag icul u alResea chandT ainingo Andalusia
(IFAPA),Go e nmen o Andalusia,CaminodePu chils/n,18004G anada,Spain
*Co espondence: b uno.na a o@jun adeandalucia.es
Abs ac :Fa mlanda o es a ionhasbeenp omo edin ecen decadesandisoneo hemains a ‐
egiesincludedin heUNDecadeonEcosys emRes o a ion o eco e deg adeda eas.Howe e ,
heimpac so a o es a iononplan di e si yandsoilquali yindica o sa es illno well‐unde ‐
s oodinsemia iden i onmen s.In hiss udy,weassessed he ela ionshipsbe weenplan di e si y
indica o s(abundance, o al ichness, ichnessby unc ionalg oups,andShannondi e si y)anda
la genumbe o a iablesin48a o es a ionsi esinsou heas Spain.Weconside edassocia eden‐
i onmen al ac o s,suchasgeog aphical,clima ico edaphic a iables,age,andland‐usehis o y.
Wecompa edplan di e si yandsoilp ope ies ollowingland‐usechange omce ealc opping
oa o es a ion,whichisoneo hemos commonland‐usechangesinMedi e aneana eas.Plan
di e si yina o es edsi eswas ound obedependen onp e iouslanduse, hep oximi yo na ‐
u al ege a ion,se e alsoilp ope ies( ex u e,pH,and o alni ogen),andplan a ionage.A o ‐
es edsoilsshowedhighe plan di e si yandanimp o emen inedaphicpa ame e s ela ed o
mul i unc ionali yinsemia idecosys ems(i.e.,soilo ganicca bon,ni ogen,andpo assium) han
a ablec oppedsoils.
Keywo ds:ac i e es o a ion; unc ionalg oups;plan ichness;soilp ope ies;land‐usechange;
Medi e anean egion;UNDecadeonEcosys emRes o a ion
1.In oduc ion
In ecen decades, a iousglobaland egionalini ia i esha ep omo edac i een i‐
onmen al es o a ionasoneo hemainwo ldwidena u e‐basedsolu ions op e en
soile osion, eco e deg adedlands,andlimi hee ec so clima echange[1].In his
sense, hea ea ha hasbeen e o es ed(plan ing eesinp e iously o es edlandscapes)
anda o es ed(plan ing eesinlandscapes ha we eno p e iously o es ed)hasin‐
c easedbyo e 123millionhasince1990,andcu en ly ep esen 7%o hewo ld’s o es
a ea[2].Scan e o sha ebeende o ed omoni o ing hesuccesso eeplan ingin
achie ingobjec i esandassessing hesuccesso es o a ion[3].TheUni edNa ionsGen‐
e alAssemblydecla ed2021–2030as heDecadeonEcosys emRes o a ion,wi hanaim
o plan ingbillionso ees h ough es o a ionini ia i esa ound hewo ld[3].Thiss a ‐
egyisde i ed om heSus ainableDe elopmen GoalsAgenda(i.e.,Sus ainableDe el‐
opmen Goal15.3),add essingbo hclima echangemi iga ionandbiodi e si yandeco‐
sys emse ices[4].
Ci a ion:Segu a,C.;Jiménez,M.N.;
Fe nández‐Ondoño,E.;Na a o,
F.B.E ec so A o es a iononPlan
Di e si yandSoilQuali yin
Semia idSESpain.Fo es s2021,12,
1730.h ps://doi.o g/10.3390/
12121730
AcademicEdi o :Ba oszAdamczyk
Recei ed:13Oc obe 2021
Accep ed:6Decembe 2021
Published:8Decembe 2021
Publishe ’sNo e:MDPIs aysneu‐
alwi h ega d oju isdic ional
claimsinpublishedmapsandins i u‐
ionala ilia ions.
Copy igh :©2021by heau ho s.Li‐
censeeMDPI,Basel,Swi ze land.
Thisa icleisanopenaccessa icle
dis ibu edunde he e msandcon‐
di ionso heC ea i eCommonsA ‐
ibu ion(CCBY)license(h ps://c e‐
a i ecommons.o g/licenses/by/4.0/).
Fo es s2021,12,17302o 17
Thisnewemphasison eeplan ingisoccu ingwi hinacon ex o in ensedeba e
ega ding hesui abili yo adop ingpassi eo ac i e es o a ions a egies.The ela i e
e ec i enesso ac i e es o a ions a egies ela i e opassi e eco e yo o me ag icul‐
u alland o ecosys em unc ionsisno well‐unde s ood[5].I hasbeenp oposed ha
heac i ea o es a iono deg aded ieldso ag icul u ala easshouldonlybeadop ed
whenecosys em es o a ionmaybeimpossiblepassi ely[6–8].Fo someau ho s,a o ‐
es a ioniso en heonly iableapp oach op e en soile osionand es o eecosys em
unc ions,ando he sp omo eac i ein e en ionasanoppo uni y o clima echange
mi iga ion ha couldha eposi i ee ec sonbiodi e si y,pa icula lywhenna i espe‐
ciesa eplan ed[9–11].
Thein luenceo a o es a iononplan di e si yandsoilquali yisdependen on
land‐usehis o y,whe he eespeciesa eplan ed, he imeelapsedsince es o a ionbe‐
gan,andlandscapecon ex ,amongs o he sen i onmen al a iables[6,12–15].S udies
conduc edunde d ycondi ions ha inco po a ea o es a ionsuccessconside ingbo h
plan di e si yandsoilquali y eco e ya esca ce,e en houghsemia idandsubhumid
a easaccoun o 25%o he e es ialEa h,andahugeag icul u ala ea(cul i a edo
abandoned)hasbeena o es edin ecen decades.These egionsa econside edex emely
ulne able olanddeg ada ionanddese i ica ion[2,10,16].P e iouslanduseisakey
d i e o plan di e si yandsoilquali ychanges ollowinga o es a ion[5,17].Howe e ,
somes udiesconduc edinsemia id ha includedbo hplan di e si yandsoilha e o‐
cusedong asslandsandna u al ege a ionins eado a mlanda o es a ion[18,19].
O he s udies ha epo edposi i elinksbe weenplan di e si yandedaphicp ope ies,
suchassoilo ganicca bon(SOC), o alni ogen(TN),phospho us(P),andpo assium(K),
ha ebeenconduc edinseconda y o es s,abandonedlands,anda oma icspeciescul i‐
a ion[20–22].
S udies ha epo soilquali yimp o emen ollowinga o es a iono a ableland
didno conside plan di e si y[23,24].Abandonmen o ag icul u allandp esen sop‐
po uni y o he eco e yo soilp ope ies ela ed o e ili y,andaninc easeinbiodi‐
e si ywi hli leneed o ac i e es o a ionanda lessexpense[25].Unde s anding
whichedaphicando he en i onmen al a iablesexplainplan di e si yunde semia id
condi ions,pa icula lyina o es a ionp ojec s,couldsuppo anydecision‐makingp o‐
cesses.Howe e ,despi esoilbeing ecognizedasamaind i e o plan di e si y, he
ex en owhichedaphicpa ame e scon olplan di e si yiss illno well‐known[26].
Theunce ain ies ela ed o hee ec so a o es a iononplan di e si yandsoil
ha eimpo an implica ions o Eu opeanUnionCommonAg icul u alPolicysubsidies
[27–29].Eu opeana o es a ione o sha ebeenincludedin heCommonAg icul u al
Policysince1992,mainlyinMedi e aneancoun iessuchasSpain,Po ugal,andI aly
[30].Unde his amewo k, heFa mlandA o es a ionSchemeaimed o educep oduc
su pluses,p omo eea ly e i emen oppo uni ies o a me s,inc ease o es esou ces,
andp o idesoilandbiodi e si yp o ec ionbyo e ing a me sannualincen i epay‐
men s o hecon e siono a mland oa o es a ion[31].Mo e han730,000hao a m‐
landhasbeena o es edinSpain,app oxima ely50%o whichwasp e iouslyde o ed
oa ablec ops[32].Thelacko planningandmeasu ableobjec i eshas esul edina o ‐
es edsi essp eadhaphaza dly,no ollowingany echnicalo e i o ialc i e ia.Thishas
complica ed hewide‐scalemoni o ingo a o es a ionimpac sonbiodi e si yandsoil
quali y,especiallywhenbaselineassessmen hasno beenes ablished[33,34].
Wi hanan icipa edexpansiono a o es edlandby heendo he21s cen u y,i is
impo an oensu eno only heaccu a emanagemen o exis inga o es a ion,bu also
heca e ulplanningo newlyplan ed o es s omaximize hebene i s o biodi e si yand
sus ainabili y[1,3,35].Anaccu a eassessmen o soilchange ollowinga o es a ion
shouldalsobeincludedasap incipalcomponen o es o a ionmoni o ing[36,37],e en
houghmany ac o sa ein ol edand ela ionshipsbe weenplan di e si yandsoilmay
Fo es s2021,12,17303o 17
no bee iden [26,38,39].Iden i yingen i onmen al ac o s ha indica ea o es a ionsuc‐
cessaswellasquan i yingchangesinplan di e si yandsoilquali yindica o s ollowing
plan ingwillhelpimp o e es o a ionsuccess.
In hiss udy,weaimed o(i)elucida ewhichen i onmen al a iablescanexplain
plan di e si y,and(ii)compa eplan di e si yandedaphicp ope iesinpai edce eal
c opsanda o es a ionsi es.We ocusedonplan species ichness,plan di e si y,and
soilquali yindica o sina ypicalsemia idMedi e aneana ea.Species ichnessanddi‐
e si ya eamong hep incipalme icschosen oassess hesuccesso es o a ionini ia‐
i es omabiodi e si ypoin o iew: heya ed i e sandp edic o so ecosys emmul‐
i unc ionali y(unde s oodas heabili yo ecosys ems op o ideandmain ainmul iple
unc ionsandse icessimul aneously), o ins ance,bybu e ing hee ec so clima e
changeanddese i ica ioninsemia idclima es,andbyin luencingnu ien cyclingand
ca bons o age[40,41].Some ecommenda ions o e alua ingplan di e si y es o a ion
ha eencou aged heassessmen o unc ionalg oups[36,38].Iden i ica iono endemism
le elsandconse a ions a usisc ucial o hee alua ingo heac i e es o a ione ec s
onbiodi e si y[12],especiallyinbiodi e si yho spo ssuchas heMedi e anean egion.
Wehypo hesized ha (1)p e iouslanduse,loca ion,clima e,andedaphicp ope ‐
ies,suchas ex u e,SOC,mac onu ien s,andpa ame e s ela ed osoilwa e con en ,
canexplainplan di e si yinMedi e aneansemia ida o es a ion;and(2)a o es a ion
imp o esplan di e si yandsoilquali yindica o s,compa ed oce ealc oppinginsem‐
ia iden i onmen s.Weexplo eden i onmen al a iables ha couldexplainplan di e ‐
si yina o es ed a mlandsusinggene alizedlinea models,quan i yingandcompa ing
plan di e si yandsoilquali yindica o sinpai eda o es a ionandce ealc opsi es.To
sol e helacko baselinesamplingbe o ea o es ing,weselec edce ealc opsascon ol
si esbecausehal o hea o es eda eain hes udy egionwasa ableland[6,27,32].
2.Ma e ialsandMe hods
2.1.S udyA ea
Thes udya ealiesin heno heas o heP o inceo G anada(sou heas Ibe ian
Peninsula)wi hin hecon ineso heBae icmoun ainsys em(Figu e1).I ex endso e
5220km2andincludes headminis a i e egionso Guadix,Baza,andHuésca .Wi hin
hisa ea,a o alo 241 a mswe ea o es edbe ween1993and2006.Thesizeo he a ms
a iedbe ween1.15and97.73ha(a e ageo 26.85ha±1.36SE).Themos common ee
specieswe ePinushalepensisMill.plan edinsingleo mixeds andswi hQue cusilexL.
subsp.ballo a(Des .)Samp.Thesewe eplan eda lowdensi ies(300–500s emha−1)using
helinea subsoiling echnique.Nosil icul u al ea men so g azingwe eadop edin
hea o es ed a mlandso e ime.Weselec eda o alo 48o hesea o es ed a ms o
hiss udyacco ding oc i e iao ep esen a i enesso he h eeadminis a i e egions,
accessibili y,andeconomiccos ela i e osamplinge o andsoilanalysis(Figu e1).
The egionhasaMedi e aneanmac oclima ewi hinmeso‐,sup a‐,ando omedi e ‐
anean he mo ypesandsemia id,d y,andsubhumidomb o ypes[42].Themos e‐
quen soilsa eLep osols(Li hicLep osolsandRendzicLep osols),Calcisols(Pe icCal‐
cisolsandHaplicCalcisols),Cambisols(Eu icCambisolsandCalca icCambisols),and
Regosols(Lep icRegosols,Calca icRegosolsandEu icRegosols).The ea ealsosmall
a easo Calca icPhaeozems[43].
Fo es s2021,12,17304o 17
Figu e1.S udya eaandsi esincluded.Pu pleci clesindica e heloca iono a o es edsi essampledin2006(N=43).
G eenpinsindica e heloca iono pai edsi essampledin2006–2007 ope o m hedi e si ycompa isonbe weena o ‐
es a ion(N=6)andce ealc ops(N=6).Blackc ossesindica e heloca iono pai edsi es,a o es a ion(N=18),andce eal
c ops(N=18)whe esoilp ope ieswe emeasu edin2011.
2.2.Da aCollec ion
F om he48 a mlandsa o es a ion,weselec ed43si es o hes udyo explana o y
en i onmen al a iables o plan di e si y(Table1andFigu e1;Supplemen a yma e ial
TableS1).Wi hineachsi e,asampleplo (20×20m)wasiden i ied,a leas 50maway
om heedge oa oidanyedgee ec s.Fo eachplo , i e20mlonglinea ansec swe e
selec ed,sepa a ed omeacho he by4m,andalignedpe pendicula o u owsmade
du ingplan ing.
Theda ao ascula plan s(includinga o es edspecies)we e eco dedwhe e he
ipo apoin e ouched he ege a ionpe pendicula lye e y100cmalong ansec s,ina
manne simila o hepoin quad a echnique(c .[44]).Wees ima edspeciesabundance
(numbe o indi idualso eachspeciespe plo )and ela i eco e (%)o heae ialpa s
o eachspecies.Ba esoil(%)wasalsoes ima ed.Allspecieswe eiden i iedandclassi ied
in o unc ionalg oups:annuals(includingannualo biennialplan s),pe ennial o bs,pe ‐
ennialg asses,andwoodyspecies(includingdwa sc ubsandsh ubs).Conse a ions a‐
uswaschecked oquan i yendemismand oiden i yendange edspecies,acco ding o
Cabezudoe al.[45].
Plan di e si ywasconside edin e mso species ichnessandspeciesdi e si y.Spe‐
cies ichnessis epo edas he o alnumbe o speciespe plo (To al_R)andas ichness
by unc ionalg oup(Annuals_R,Fo bs_R,G asses_R,andWoody_R).Speciesdi e si y
wascalcula edas heShannon–Wiene index(H’):
H’=−Σp
i
lnp
i
,(1)
whe ep
i
is heabundanceo iexp essedasap opo iono he o alspeciesabundancein
eacha o es edplo .
Di e encesinplan di e si ybo hincludingandexcludinga o es edspecieswe e
examined(To al_HandH_noa , espec i ely).
Fo es s2021,12,17305o 17
Table1.En i onmen alda acollec ed o each a mlanda o es a ion(N=43).*Nominal a iables.
Va iableTypeVa iableUni Mean(SD)MinimumMaximum
Geog aphicEle a ionabo eseale el(Ele a ion)m1237.0(304.2)881.01939.0
Dis ance oc ops(D_c ops)m188.5(308.0)0.01488.0
Dis ance ona u al ege a ion(D_n_ eg)m61.9(133.7)0.0531.5
Dis ance osh ubpa ches(D_sh ub)m1241.0(1022.0)90.54675.0
Dis ance oQue cuswoodland(D_Qil)m1727.0(2219.0)0.09834.0
Clima icAnnualP ecipi a ion(Pp)mm394.0(65.2)311.0547.6
MeanTempe a u e(T)°C12.68(0.98)10.9014.10
EdaphicA ailablewa e (AW)%6.82(2.90)2.2212.74
Soilmois u ea ieldcapaci y(33kPa)
(SM33)%16.24(5.78)5.0928.58
Soilmois u eo hewil ingpoin (1500
kPa)(SM1500)%9.42(3.81)2.8719.02
Wa e sa u a ion(WS)%38.39(7.59)27.6356.42
G a el(>2mm)%37.24(18.02)0.0073.42
Sand(2–0.05mm)%45.28(16.75)3.2375.65
Coa sesil (0.05–0.02mm)(CSil )%10.27(4.35)3.0724.67
Finesil (0.02–0.002mm)(FSil )%22.15(9.66)8.3757.25
Clay(<0.002mm)%22.30(8.99)6.1238.47
Ca ionexchangecapaci y(CEC)Cmol(+)
kg−111.28(6.13)3.2725.39
Elec icalconduc i i y(EC)mS
cm−10.89(0.40)0.512.21
Exchangeablesodium(Na)Cmol(+)
kg−10.06(0.07)00.29
Exchangeablemagnesium(Mg)Cmol(+)
kg−11.86(1.28)0.305.98
Exchangeablepo assium(K)Cmol(+)
kg−10.36(0.27)0.061.28
Solublecalcium(s_Ca)mgL−13.71(2.41)0.679.81
Solublemagnesium(s_Mg)mgL−10.65(0.47)0.162.59
Solublepo assium(s_K)mgL−10.40(0.51)0.023.05
Solublesodium(s_Na)mgL−11.61(0.98)0.134.78
CaCO3(CaCO3)%24.36(23.24)0.1073.78
pH7.90(0.55)6.728.83
Soilo ganicca bon(SOC)%1.24(0.82)0.303.19
To alni ogen(TN)%0.11(0.05)0.050.24
NO2−(NO2)mgL−13.29(8.65)0.0041.60
NO3−(NO3)mgL−111.10(44.41)0.00233.40
SO42−(SO4)mgL−146.60(64.58)6.81301.50
Cl−(Cl)mgL−130.25(22.01)10.53117.2
GeologicalLi hology*
LanduseAgeo a o es a ion yea s8.4(2.1)5.012.0
P e iouslanduse(ce ealc opo old‐
ield)*
Plan co e (Co e )%67.49(13.15)40.0091.00
Co e o Pinushalepensis(Co Pha)%5.93(6.44)0.0032.00
Co e o Que cusilexsubsp.ballo a
(Co Qil)%0.26(0.62)0.003.00
Fo es s2021,12,17306o 17
Theedaphiccha ac e is icsandgeog aphic,clima ic,geological,andlanduse a ia‐
bles o eacha o es edsi ea esumma izedinTable1.Composi esoilsampleswe ecol‐
lec ed om hecen e andco ne so eachquad a oadep ho 0–10cmandmixed o
ob ainone ep esen a i esamplepe plo .Soilsampleswe eai ‐d iedandsie ed(<2
mm).G a els(>2mm)we eweighedands o edsepa a ely.Soil ex u ewasanalyzedby
heRobinsonpipe eme hod[46].A ailablewa e con en wascalcula edby hedi e ‐
encebe weenmois u econ en a ieldcapaci yex ac edinap essu epla ea −33kPa
andmois u ea plan wil ingpoin ,measu eda −1500kPa[47].Exchangeablebases
(Mg2+,K+,andNa+)we eex ac edwi h1NNH4OAc,andca ion‐exchangecapaci ywas
de e minedbysa u a ioninsodiumbywashing hesoilsampleswi halcoholandex ac ‐
ing hesodiumadso bedwi h1NNH4OAc[46].pHwasmeasu edinasoilsuspension
indis illedwa e (1:2.5).Soilo ganicca bonwasde e minedusing heWalkleyandBlack
me hod[48],modi iedbyTyu in[49].TheKjeldahlme hodwasused ocalcula e o alN
[50].CaCO3equi alen swe ede e minedusing hemanome icme hod[51].Asa u a ed
ex ac wasp epa ed omeachsample ode e minei selec icalconduc i i y[52].Solu‐
blecalcium,magnesium,po assium,andsodiumin hesa u a edex ac swe ede e ‐
minedbya omic‐abso p ionspec ome y.A ailablephospho us(onlymeasu edin2011)
wasde e minedusingOlsen’sme hod[53].NO2−,NO3−,SO42−,andCl‐in hesa u a ed
ex ac swe ede e minedbyhigh‐p ecisionliquidch oma og aphy(HPLC).
Geog aphic a iableswe eob ained om heLandUseandVege a ionCo e Map
o Andalusia(scale1:25,000)[54].Clima eda awe ede i ed omin o ma ionp o ided
by heNa ionalMe eo ologicalIns i u eo Spainusing hein e pola ionme hodp o‐
posedbySánchez‐Paloma ese al.[55].Li hologicalda awe eob ained omgeological
mapspublishedby heSpanishIns i u e o GeologyandMining[56].Li hologywasdi‐
idedin o he ollowingca ego ies:(1)ca bona e ocks,includinglimes ones,dolomi es,
ma bles,limes onec us s,andconglome a es;(2)siliceous ocks,includingmicaschis s,
phylli es,andqua zi es;(3)sil swi hclay;and(4)sands.
Da a ela ing oplan a ionage,land‐useandmanagemen we ep o idedby he
Minis yo Ag icul u eandFishe ieso Andalusia.Rela i eco e so P.halepensisandQ.
ilexsubsp.ballo ameasu edwi hin hesampleda o es eda easwe eincludedasland‐
use a iables.
Apai edsi ecompa isonapp oachwas ollowed ocompa eplan di e si yandsoil
cha ac e is icsbe weena o es edandce ealc opsi esbecausebaselinesamplingbe o e
a o es ingwasuna ailable.A o alo sixpai edsi eswi hsimila ecologicalcha ac e is‐
icswe echosen ocompa eplan di e si y(Figu e1).Addi ionalsoilsamplingwasca ‐
iedou in18pai edsi es ocompa esoilpa ame e sa wosoildep hs(0–5and5–10cm)
be weenbo hlanduses(Figu e1).Adjacen ce ealc opsconside edin hesepai edcom‐
pa isonswe esampled hesameasa o es edplo s.
2.3.S a is icalAnalyses
Ap elimina yexplo a iono di e encesbe ween hep e iouslanduses(a o es a‐
ionplan edinc oplands e susold‐ ields,N=43)waspe o medby wo‐sample ‐ es s
and wo‐sampleWilcoxon es (non‐pa ame ic es (W)) o all hes udy a iables.
Gene alizedlinea models(GLMs)we eused oanalyzeplan di e si yindica o sin
hea o es edsi es(N=43)includinggeog aphic,clima ic,edaphic,geological,andland‐
useda aasexplana o y a iableso di e si y.Tos a ,co ela ionanalysiswaspe o med
onquan i a i eexplana o y a iables,andhighlyco ela ed a iables( >|0.75|)we e
excluded omsubsequen modellings eps.APoissone o dis ibu ionwasused o
modelAbundance,To al_R,and ichnessby unc ionalg oup(Annuals_R,Fo bs_R,
G asses_R,andWoody_R).AGaussiane o dis ibu ionwasused omodelTo al_H.
Collinea i yando e dispe sionwe echecked o model alida ion.Va iancein la ion
ac o s(VIFs)we eused oiden i ycollinea i ybe ween a iables.Colinea a iables
we e emo ed om hemodels.Homoscedas ici yandno mali ywe ee alua edbyplo ‐
ing esidualsandp edic ed alues.Thebes modelswe eselec edacco ding o helowe
Fo es s2021,12,17307o 17
Akaikein o ma ionc i e ion(AIC).Thep opo iono a ianceexplainedby hemodel
wascalcula edasapseudoR2=[(nullde iance— esidualde iance)/nullde iance]×100.
Ini ialmodelselec ionwaspe o med ollowingas epwisep ocedu e.Ino de o i he
mos pa simoniousmodel,anynonsigni ican ac o swe e emo ed om he a iables
includedin heini ialmodelwhen heAICwascompa able.Finally, hee ec o each
signi ican explana o y a iableon he esponse a iablewasexplo edbyexamining ac‐
o coe icien sandplo ingadjus edp edic ionso he esponse a iableusing hemos
pa simonious i edmodel.Compa isonsbe weena o es edandadjacen ce ealplo s
we econduc edby wo‐sample ‐ es s.Alls a is icalanalyseswe epe o medusingR
so wa e,andggplo 2,co plo ,ca ,MASS,andgge ec spackages[57–62].
3.Resul s
3.1.En i onmen alCha ac e is icsandPlan SpeciesinA o es edSi es
Thi y‐nineen i onmen al a iableswe e eco ded o eacho he43a o es edsi es
(Table1;TableS1).A o alo 252plan specieswe eiden i ied(TableS2, axonomyac‐
co ding oBlancae al.[63]).Onlyeigh o hesehadbeena i iciallyin oduced:Pinus
halepensis,Que cusilexsubsp.ballo a,PinuspineaL.,P unusmahalebL.,Junipe usphoenicea
L.,Pinusnig aJ.F.A nold,P unusa iumL.,andQue cus agineaLam.The emainingna i e
specieswe edi idedas53.57%annuals,14.29% o bs,6.35%pe ennialg asses,and
25.79%woodyspecies.O hesespecies,a o alo 52(20.63%)belonged oa es ic eda ea
o dis ibu ion(TableS2):28we eendemic o heIbe ianpeninsulaandNo hA ica
(11.11%),15we eendemic o heIbe ianpeninsula(5.95%),and13we eendemic o he
sou handsou heas egionso heIbe ianpeninsula(5.15%).Weiden i iedonlyoneen‐
dange edspecies(0.40%) ha appea sin heAndalucianRedLis o h ea enedspecies
(Cen au eapul ina a(Blanca)Blanca).Themeandi e si yes ima edpe samplewhenin‐
oduced eespecieswe eincluded(To al_H)was2.54±0.40(SD),and2.49±0.44when
heywe edis ega ded(H_noa ).
3.2.E ec o heP e iousLandUseonPlan Di e si yandEn i onmen alVa iables
Annual_R,Co Phal,D_n_ eg,Mg,andAgewe esigni ican lyhighe ina o es ed
si es haninpai edc oppedsi es( =4.2305,d =41,p‐ alue<0.001;Wilcoxon es ,W=
319.5,p‐ alue<0.05;W=305.5,p‐ alue<0.05;W=320,p‐ alue<00.05;andW=351,p‐
alue<0.01, espec i ely).G asses_R,Woody_R,andD_c opswe ehighe ina o es ed
old‐ ields(W=74,p‐ alue<0.001;W=33,p‐ alue<0.0001;andW=120,p‐ alue<0.01,
espec i ely).Nosigni ican di e enceswe e oundbe weenp e iouslanduse o Abun‐
dance,To al_R,To al_H,andFo bs_R,and he emainingen i onmen al a iables(Sup‐
plemen a yMa e ial,Figu esS1–S4).
3.3.E ec so En i onmen alVa iablesandSoilP ope iesonPlan Di e si y
Thequan i a i epa ame e sincludedasexplana o y a iablesinGLMswe ese‐
lec ed akingin oaccoun aco ela ioncoe icien lowe han|0.75|(Figu eS5).Addi‐
ionally,p e iouslanduseandli hologywe eincludedin hemodelsasca ego icalex‐
plana o y a iables.Allmodel esul sandp edic ed esponsesg aphsa ea ailablein
Supplemen a yMa e ial.
Fo Abundance, hemos pa simoniousmodel e ealedasigni ican associa iononly
wi hCo PhaandCo e (TableS3andFigu eS6).
Theselec edmodel o To al_Rshowedsigni ican nega i e ela ionshipswi h
D_n_ eg,D_sh ub,s_K,CSil ,SOC,Age,andOld‐ ield ac o (Table2;Figu e2;TableS4).
Themodel e ealede idenceo posi i ee ec so s_Mg,TN,pH,andCo e onTo al_R
(Table2andFigu e2).
Fo es s2021,12,17308o 17
Table2.Resul s o To al_Rbes andmos pa simoniousmodel(pseudoR2=75.92%;AIC=258.3).
Es ima eSEzValueP (>|z|)
In e cep 1.390000.734101.8940.058
D_n_ eg−0.000960.00037−2.5660.010
D_sh ub−0.000080.00004−2.0910.037
s_Mg0.263800.092702.8460.004
s_K−0.306200.09801−3.1240.002
CSil −0.029480.00944−3.1210.002
SOC−0.212000.07265−2.9190.004
pH0.227200.086422.6290.009
TN3.186001.179002.7030.007
Age−0.066330.02660−2.4940.013
P e iouslanduseOld‐ ield−0.351900.09792−3.593<0.001
Co e 0.015080.002895.219<0.001
D_n_ eg=dis ance ona u al ege a ion(m);D_sh ub=dis ance osh ubpa ches(m);s_Mg=
solubleMg(mgL−1);s_K=solubleK(mgL−1);CSil =coa sesil ac ion(0.05–0.02mm,%);SOC=
soilo ganicca bon(%);TN=soil o alni ogen(%);Age=a o es a ionage(yea s);Co e =plan
co e (%);P e iouslanduse=landusep io oa o es a ion(old‐ ieldo ce ealc ops).
Figu e2.Rela ionshipsbe weenTo al_Randexplana o yand esponse a iablesiden i iedby hebes i model.Black
linesa e he i edlinescalcula edusing heggp edic unc ioninR,p o iding hep edic ed alueson hescaleo he
esponse.Theshadeda ea ep esen s he95%con idencein e al.Poin s ep esen measu ed alues(N=43).(a)D_n_ eg
=dis ance ona u al ege a ion(m);(b)D_sh ub=dis ance osh ubpa ches(m);(c)s_Mg=solubleMg(mgL−1);(d)s_K
=solubleK(mgL−1);(e)CSil =coa sesil ac ion(0.05–0.02mm,%);( )SOC=soilo ganicca bon(%);(g)TN=soil o al
ni ogen(%);(h)pH;(i)Age=a o es a ionage(yea s);(j)Co e =plan co e (%);(k)landusep io oa o es a ion.
Fo unc ionalg oups,Annuals_Rwassigni ican lynega i elyassocia edwi h
D_Qil,K,CSil ,NO3,SandandSil +clayli hology,Old‐ ieldp e iouslanduse,andAge
(TableS5andFigu eS7).Posi i ee ec swe e ound o Mg,AW,EC,pH,Siliceous ocks,
Co Phal,andCo e .
Fo es s2021,12,17309o 17
Signi ican posi i eassocia ionswe e ound o pHandCo e onFo bs_R;howe e ,
Kseemed oexe anega i ee ec (TableS6andFigu eS8).
Fo G asses_R, heselec edmodelshowedaposi i e ela ionshipwi hAW,CSil ,
andOld‐ ieldp e iouslanduse(TableS7;Figu eS9).E idenceo anega i ee ec o
D_sh ubandCaCO3onG asses_Rwas ound.
Theselec edmodel e ealede idenceo henega i ee ec so G a elandCaCO3on
Woody_R(TableS8;Figu eS10).Ono he hand,aposi i ee ec was oundonWoody_R
byOld‐ ieldas hep e iouslanduse.
Themos pa simoniousmodel o To al_Hincluded12 a iables,no allassocia ed
wi hsigni ican e ec s(Table3;TableS9).D_n_ eg,D_sh ub,s_K,CaCO3,andP e i‐
ous_land_use(Old‐ ield ac o )hadanega i ee ec onTo al_H,whe eass_Mg,pH,
Co Qil,andCo e showedaposi i ee ec (Figu e3).
Table3.In luenceo ac o siden i iedby hemos pa simoniousmodelpe plan ShannonIndex
di e si y(To al_H)(pseudoR2=74.71%;AIC=11.24).
Fac o Es ima eSE ValueP (>| |)
(In e cep )−0.922200.91590−1.0070.322
D_n_ eg−0.000980.00039−2.5310.017
D_sh ub−0.000180.00004−4.285<0.001
D_Qil−0.000050.00003−1.9690.058
s_Mg0.503300.135303.7200.001
s_K−0.260500.11160−2.3350.026
CSil −0.017160.01152−1.4900.147
CaCO3−0.006720.00261−2.5760.015
pH0.476300.115504.123<0.001
Age−0.062440.03068−2.0360.051
Old‐ ieldp e iouslanduse−0.359100.10910−3.2920.003
Co Qil0.136400.064512.1150.043
Co e 0.013120.004053.2410.003
D_n_ eg=dis ance ona u al ege a ion(m);D_sh ub=dis ance osh ubpa ches(m);D_Qil=
dis ance oQue cuswoodland(m);s_Mg=solubleMg(mgL−1);s_K=solubleK(mgL−1);CSil =
coa sesil ac ion(0.05–0.02mm,%);CaCO3(%);pH;Age=a o es a ionage(yea s);Co Qil=
Que cusilexco e (%);Co e =plan co e (%)
Fo es s2021,12,173016o 17
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