Mul i axon biodi e si y and unc ional eco e y in es o ed na i e o es s in
a biosphe e ese e
Unai O ega-Ba ue a
1,*
, Unai Se u xa
2
, Ibone Ame zaga-A egi
3
,
Ma kel Mon a˜
no-Lekue
4
, Lo ena Pe˜
na
5
Depa men o Plan Biology and Ecology, Uni e si y o he Basque Coun y (UPV/EHU), Apdo 644, Bilbao 48080, Spain
ARTICLE INFO
Key wo ds:
Exo ic plan a ions
Eucalyp us sp.
Pinus adia a
Plan s
Bi ds
Bac e ia
Fungi
ABSTRACT
Fo es landscapes in Eu ope ha e his o ically unde gone ex ensi e an h opogenic ans o ma ion, leading o he
widesp ead eplacemen o na i e o es s wi h pas u es, exo ic plan a ions, and u ban de elopmen s, ul ima ely
educing biodi e si y. Despi e ecen na u al o es expansion and es o a ion e o s, his has no always been
accompanied by ecological eco e y. In he Basque Coun y (no he n Ibe ian Peninsula), Pinus adia a and
Eucalyp us plan a ions ha e become dominan , aising ecological conce ns due o hei low species di e si y and
impac s on soil p ocesses. In esponse, he EU Na u e Res o a ion Law (2024) has es ablished binding a ge s o
eco e deg aded ecosys ems, p omo ing ac i e es o a ion s a egies ha inco po a e ai -based and mul i axon
app oaches. This s udy e alua es whe he na i e o es es o a ions in he U daibai Biosphe e Rese e suppo
highe biodi e si y and ecosys em unc ioning compa ed o he mos abundan o es sys ems in he a ea. Fou
o es ypes ( es o ed na i e o es s, Pinus adia a plan a ions, Eucalyp us plan a ions, and e e ence na i e o es )
we e compa ed ac oss 36 plo s. We assessed ascula plan s, bi ds, bac e ia, and ungi using ield su eys, soil
DNA sequencing, and unc ional ai assignmen s. Di e si y me ics and bo h axonomic and unc ional
composi ion we e compa ed among sys ems. Res o ed o es s showed signi ican ly highe plan and bi d di-
e si y han exo ic plan a ions, wi h Eucalyp us consis en ly exhibi ing lowes alues ac oss mos indica o s.
Species and unc ional composi ion analyses e ealed ha es o ed o es s closely esembled na i e sys ems,
while Eucalyp us was associa ed wi h ea ly-successional and s ess- ole an ai s. Pine plan a ions exhibi ed
in e media e pa e ns, pa icula ly in mic obial communi ies. O e all, ac i e es o a ion wi h na i e species
p o ed e ec i e in eco e ing biodi e si y and ecosys em unc ion, hough ull con e gence wi h na i e o es s
may equi e longe imescales.
1. In oduc ion
Eu opean o es landscapes ha e unde gone cen u ies o an h opo-
genic ans o ma ion. The eplacemen o agmen a ion o na i e o -
es s by ag icul u al land, pas u es, o es y monocul u e sys ems o
u ban de elopmen s (Kaplan e al., 2009) has led o a decline in biodi-
e si y (Maes e al., 2023). Na u al expansion o na i e o es s, la gely
d i en by u al abandonmen , has con ibu ed o inc eased o es co e
in ecen imes. Howe e , la ge-scale a o es a ion and e o es a ion
p og ammes implemen ed du ing he wen ie h cen u y o es o e o es
co e has mos ly been aimed a supplying imbe ma ke s (Ma he ,
2001; Chazdon, 2008; Vadell e al., 2016). Fu he mo e, in ecen yea s,
his end has in ensi ied unde he umb ella o clima e policy (CO
2
emo al), incen i ising he expansion o as -g owing exo ic o es
plan a ions. This phenomenon has been pa icula ly e iden in he
Ibe ian Peninsula and Scandina ia, e lec ing a g owing p essu e on
* Co esponding au ho .
E-mail add esses: [email p o ec ed] (U. O ega-Ba ue a), [email p o ec ed] (U. Se u xa), [email p o ec ed] (I. Ame zaga-A egi),
[email p o ec ed] (M. Mon a˜
no-Lekue), [email p o ec ed] (L. Pe˜
na).
1
ORCID: 0000–0001-7773–7973
2
ORCID: 0000–0003-1961–5649
3
ORCID: 0000–0003-0683–1190
4
ORCID: 0009–0008-5203–505X
5
ORCID: 0000–0001-8329–9869
Con en s lis s a ailable a ScienceDi ec
Fo es Ecology and Managemen
jou nal homepage: www.else ie .com/loca e/ o eco
h ps://doi.o g/10.1016/j. o eco.2025.123057
Recei ed 2 June 2025; Recei ed in e ised o m 29 July 2025; Accep ed 31 July 2025
Fo es Ecology and Managemen 596 (2025) 123057
A ailable online 14 Augus 2025
0378-1127/© 2025 The Au ho s. Published by Else ie B.V. This is an open access a icle unde he CC BY-NC-ND license ( h p://c ea i ecommons.o g/licenses/by-
nc-nd/4.0/ ).
o es sys ems linked o in ensi y managemen p ac ices (Cecche ini
e al., 2020). Howe e , maximizing sho - e m ca bon seques a ion
h ough monocul u e plan a ions may lead o ade-o s wi h biodi e -
si y, especially in empe a e o es s (Saba ini e al., 2019, Smi h e al.,
2024).
The no he n Ibe ian Peninsula p o ides a clea example o he
ecological impac o hese policies, whe e in ensi e plan a ion o es y
has long shaped he landscape (Vadell e al., 2016). Conc e ely in he
Basque Coun y, o e he pas cen u y, Pinus adia a monocul u es ha e
sp ead apidly ac oss he landscape due o hei apid g ow h and imbe
yield (Michel Rod íguez, 2006). Howe e , in ecen decades, he decline
in pine p o i abili y, d i en by ma ke ins abili y and widesp ead ou -
b eaks o ed and b own needle bligh (Mon ei o e al., 2022; O u˜
no
e al., 2022), has led o an inc easing subs i u ion wi h Eucalyp us sp.
plan a ions (Gobie no Vasco, 2024; Vadell e al., 2022). This change in
land use aises ecological conce ns, as eucalyp us plan a ions exhibi
low species di e si y and al e soil condi ions, he eby a ec ing key
ecosys em p ocesses such as nu ien cycling and li e decomposi ion
(Cal i˜
no-Cancela e al., 2012; Goded e al., 2019; Mallen-Coope e al.,
2022; Se u xa e al., 2025a). These impac s a e associa ed no only wi h
hei in ensi e managemen p ac ices, such as hei sho o a ion cy-
cles, compa ed o pine plan a ions, bu also wi h in insic cha ac e is ics
o eucalyp us, such as i s slow-decomposing li e , p oduc ion o alle-
lopa hic compounds and a high wa e use (Amazonas e al., 2018; Elo-
segi e al., 2020).
In esponse o inc easing en i onmen al p essu es, in 2024 he EU
Na u e Res o a ion Law was es ablished in he Eu opean Union as pa o
he Eu opean G een Deal and he EU Biodi e si y S a egy o 2030. The
Regula ion es ablishes legally binding es o a ion a ge s o a wide
ange o ecosys ems, wi h he aim o es o ing 20 % o he EU’s deg aded
ecosys ems by 2030 and all ecosys ems in need o es o a ion by 2050
(Regula ion 2024/1991). Indeed, ecological es o a ion has eme ged as
a pi o al s a egy o e e sing biodi e si y loss, wi h a ious o gani-
sa ions employing land s ewa dships o ca y ou es o a ion ac i i ies in
di e en ecosys ems. The goal o es o a ion ecology is o inc ease
ege a ion co e and es o e ecological p ocesses, species in e ac ions,
and communi y s uc u es ha esemble hose obse ed in e e ence
ecosys ems (Chazdon, 2008; Ga ica-Saa ed a e al., 2017).
In his con ex , ac i e es o a ion measu es, encompassing he
plan ing o na i e species, he e adica ion o in asi e alien species, and
he p o ec ion o egene a ing ege a ion, a e equen ly impe a i e in
highly deg aded o agmen ed landscapes (C ouzeilles e al., 2017; Meli
e al., 2017). Ne e heless, assessing he e icacy o es o a ion en-
dea ou s emains a challenge, as con en ional me ics, such as species
ichness and canopy co e , only o e a limi ed pe spec i e on ecological
eco e y. Consequen ly, es o a ion success is inc easingly being e al-
ua ed using biodi e si y me ics ha include axonomic ichness and
composi ion, as well as unc ional di e si y, de ined as he ange o ai s
in luencing ecosys em unc ioning (Cado e e al., 2011;
Ga ica-Saa ed a e al., 2017). The inco po a ion o unc ional ai s
enables a deepe unde s anding o how es o ed communi ies esemble
o e e ence sys ems and how hey egula e key ecological p ocesses
o e ime. The applica ion o unc ional ai s has been demons a ed o
acili a e he design o mo e esilien and adap i e es o a ion s a egies.
This is achie ed by aligning species selec ion and managemen s a e-
gies wi h speci ic en i onmen al conce ns and desi ed ecosys em unc-
ions (Lou ei o e al., 2023).
Fu he mo e, he e is a g owing consensus ha a mul i axon
app oach is c ucial o accu a ely cap u ing es o a ion ou comes ac oss
a ious ophic le els (Seibold e al., 2018). This is pa icula ly ele an
gi en ha di e en axonomic g oups espond asynch onously o en i-
onmen al change and es o a ion in e en ions, in e ms o in ensi y
and di ec ion o change. Fo his eason, me hodological app oaches
based on a single axon ha e he po en ial o yield misleading conclu-
sions (C ouzeilles e al., 2016; Doe le e al., 2020). The inclusion o
bo h abo eg ound and belowg ound o ganisms is he e o e essen ial in
o de o achie e a comp ehensi e e alua ion o he es o a ion ou -
comes, gi en he sensi i i y o ascula plan s, bi ds, bac e ia, and ungi
o al e a ions esul ing om o es managemen p ac ices (Ba dge and
an de Pu en, 2014). Indeed, ascula plan s and bi ds a e equen ly
u ilised as bioindica o s due o hei sensi i i y o habi a s uc u e,
ege a ion complexi y, and landscape composi ion. Fu he mo e, hey
ul il pi o al unc ions in ecosys em p ocesses, including p ima y p o-
duc ion, seed dispe sal and ophic egula ion (Massi e al., 2022; Jung
e al., 2024). Howe e , he majo i y o hese assessmen s demons a e a
pauci y o conside a ion wi h ega d o mic obial di e si y and unc-
ioning. Soil mic obial communi ies, comp ising bac e ia and ungi, a e
undamen al o nu ien cycling and decomposi ion. Thei p onounced
esponse o al e a ions in li e quali y and soil chemis y ende s hem
inc easingly aluable as indica o s o ecological eco e y (Sun e al.,
2017; Sansupa e al., 2021; Huang e al., 2023). The in eg a ion o soil
mic obial communi ies wi h abo eg ound axa has been demons a ed
o enhance he abili y o de ec es o a ion ou comes and assess
ecosys em esilience in a mo e comp ehensi e manne (Hou e al.,
2024). Ne e heless, empi ical assessmen s applying his mul i axon,
unc ional pe spec i e emain sca ce, pa icula ly in biosphe e ese es
cha ac e ised by deg aded o agmen ed landscapes, such as he
U daibai Biosphe e Rese e (UBR). These a eas ha e been designa ed
o he p omo ion o sus ainable de elopmen , wi h a iew o enhancing
he well-being o he local popula ion (Ishwa an e al., 2008) and
achie ing conse a ion objec i es. Fo his eason, in ecen yea s, in
UBR es o a ion e o s led by o ganisa ions such as Fundaci´
on Lu gaia
ha e ocused on ac i ely ans o ming monocul u es in o na i e o es s
and es o ing deg aded and agmen ed na i e o es s. These es o a-
ions ha e in ol ed plan ing o na i e species o imp o e biodi e si y
and landscape connec i i y (O ega e al., 2023).
The aim o his s udy is o e alua e whe he he es o a ion o na i e
o es s in he UBR is mo e e ec i e a suppo ing biodi e si y and key
ecosys em unc ioning compa ed o he mos abundan o es sys ems. In
o de o achie e his objec i e, we conduc ed a compa a i e analysis o
he species and unc ional di e si y and composi ion o ou o es sys-
ems ac oss mul iple axonomic g oups ( ascula plan s, bi ds, bac e ia
and ungi). Conc e ely, es o ed na i e o es s we e compa ed o Pinus
adia a and Eucalyp us sp. plan a ions o simila age, as well as o na i e
o es s ha se e as e e ence ecosys ems in he egion. This mul i axon
compa ison allowed us o examine whe he es o ed o es s exhibi ed
signs o con e gence wi h e e ence o es o wi h o es plan a ions. In
addi ion, we iden i ied he main ecological unc ions associa ed wi h
each o es ype in o de o assess whe he es o ed o es s a e unc-
ionally close o na i e o es s o o exo ic plan a ions.
2. Me hods
2.1. S udy a ea
The UBR, loca ed in he Basque Coun y in no he n Ibe ian Penin-
sula (Fig. 1), encompasses app oxima ely 22,000 ha and was designa ed
a UNESCO Biosphe e Rese e in 1984 and subsequen ly in eg a ed in o
he Na u a 2000 ne wo k. This designa ion has p omo ed he sus ain-
able use o na u al esou ces while os e ing ecosys em p o ec ion,
gi ing ise o a socio-ecological sys em cha ac e ised by he coexis ence
be ween conse a ion goals and economic ac i i ies (Onaindia e al.,
2013b, Cas illo-Eguski za e al., 2017). The UBR landscape is cha ac-
e ised by a he e ogeneous mosaic shaped by his o ical land ans-
o ma ions and cu en an h opogenic p essu es. The egion has a
humid A lan ic clima e, wi h mean annual empe a u es exceeding 13◦C
and annual p ecipi a ion anging om 1200 o 1700 mm, dis ibu ed
ela i ely e enly h oughou he yea . The al i udinal g adien anges
om sea le el o 776 m, hough mos o he ese e lies below 400 m.
The li hology o he a ea is composed o c e aceous o ma ions, p i-
ma ily lysch, ma l and sands one, ha con ibu e o he de elopmen o
acidic, oligo ophic soils (pH 3.5–6.0) ha suppo acidophilous o es
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
2
communi ies (Loidi e al., 2011; O an ia e al., 2019).
The Can ab ian g een oak o es is conside ed one o he mos highly
alued na u al o es s o he ese e; howe e , mixed-oak o es s
domina ed by Que cus obu L. a e he dominan po en ial ege a ion in
he UBR (Onaindia e al., 2013a). Mos o hese mixed-oak o es s a e
cons i u ed o small s ands, ypically be ween 35 and 70 yea s, wi h
minimal o i ually no sil icul u al in e en ion. Cu en ly, hey
occupy less han 9 % o he ese e, while monocul u es o as -g owing
exo ic species o sil icul u al pu poses, including Pinus adia a and
Eucalyp us sp., co e o e 50 % o he landscape (Gobie no Vasco,
2024). Conc e ely, Pinus adia a, also known as Mon e ey pine, is na i e
o a es ic ed coas al ange in Cali o nia and no he n Baja Cali o nia
(Michel Rod íguez, 2006), whe e i plays a ele an ole in main aining
ecosys em dynamics h ough apid g ow h and high dis u bance esil-
ience. Rega ding eucalyp us plan a ions, he main species plan ed in he
egion a e Eucalyp us globulus Labill. and Eucalyp us ni ens H. Deane &
Maiden, which a e na i e o sou heas e n Aus alia and cons i u e a
undamen al elemen o empe a e scle ophyll o es s in ha egion.
The es ablishmen o pine plan a ions in he Basque Coun y, which
a e p edominan ly si ua ed on p i a ely owned land ha p e iously
unc ioned as pas u es, ma ginal ag icul u al c oplands and na i e o -
es s, can be a ibu ed o he mid-20 h cen u y a o es a ion policies.
Howe e , a ea o eucalyp us plan a ions has doubled om 2005 o 2024
due o subs i u ion o pine wi h eucalyp us due o he b own and ed
band disease ha ha e ecen ly a ec ed pine plan a ions in he egion.
These changes in he landscape a e a esponse o socio-economic
changes and declining p o i abili y o adi ional ag icul u al p ac ices
and o es y p ac ices (Se u xa e al., 2025a).
In he Basque Coun y, he p ocess o es ablishing o es plan a ions
is cha ac e ised by a sequence o s eps, beginning wi h soil p epa a ion
ollowed by ee plan ing. In he case o pine plan a ions, subsequen ly
wo o h ee hinning ope a ions a e ypically conduc ed du ing he
s and’s de elopmen , wi h o a ion cycles o 30–40 yea s. In he case o
eucalyp us plan a ions, o a ion cycles a e o 10–15 yea s and subse-
quen o ha es ing, he s umps a e pe mi ed o esp ou . These plan-
a ions a e o en si ua ed in a eas wi h challenging opog aphy, leading
o ecological agmen a ion and loss o unc ional connec i i y. Despi e
hese challenges, he egion ha bou s habi a s o high conse a ion
alue, including ipa ian o es s, Can ab ian holm oak g o es, and
es ua ine ma shes. These habi a s con ibu e o he a ea’s ele a ed
lo is ic and aunal di e si y (Cas illo-Eguski za e al., 2017).
2.2. S and selec ion
Fi s ly, 6 sampling poin s we e es ablished andomly in he wo
oldes o es es o a ion a eas ha a e managed by Fundaci´
on Lu gaia, 3
sampling poin s in each one wi h a minimum dis ance o 200 m (Ba low
e al., 2007). The numbe o sampling poin s pe si e was de e mined
based on he size o he es o a ion a eas (5 and 7 ha) and he minimum
dis ance equi ed be ween sampling poin s o educe spa ial au oco -
ela ion. These a eas ha e unde gone ecological es o a ion o
app oxima ely 15–20 yea s and a e e e ed o he ein as “Res o a ion”.
The es o a ion managemen s a egy encompassed he plan ing o
na i e ee species, including na i e ees such as Que cus obu , Co ylus
a ellana, and F axinus excelsio . P o ec i e ba ie s we e placed a ound
each ee, and he a ea was subjec ed o moni o ing o p e en he
p oli e a ion o in asi e species. Subsequen ly, 10 Eucalyp us sp. plan-
a ions s ands (he ea e e e ed o as “Eucalyp us”) and 10 Pinus
adia a plan a ions s ands (he ea e e e ed o as “Pine”) we e selec ed
andomly; bo h we e app oxima ely 20 yea s old. Finally, 10 mixed oak
o es s s ands olde han 35 yea s (he ea e e e ed o as “Na i e”)
we e selec ed andomly (Fig. 1). O hopho os we e used o e i y he age
Fig. 1. Map showing he geog aphic loca ion o he sampling poin s o Na i e o es s, Res o a ion si es, Pine plan a ions, and Eucalyp us plan a ions in he UBR
loca ed in he Basque Coun y.
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
3
o he o es sys ems. In each s and, a sampling poin was es ablished a
he cen e o a oid edge e ec s. A summa y able wi h key si e de-
sc ip o s (ele a ion, slope, o es s uc u e me ics, soil pH, and li hol-
ogy) o each sampling poin is p o ided in Appendix B (Table B.1).
2.3. Sampling design
Field su eys we e pe o med be ween Ap il and June in 2023. In
o de o sample he ascula plan species, om each sample poin , wo
pe pendicula ansec s o 50 m we e es ablished, o ming a c oss. A
o al o i e subplo s, each measu ing 10 m² (2 m ×5 m), we e es ab-
lished a egula in e als along he ansec s. The cen al subplo was
sha ed by bo h ansec s, esul ing in a o al sampled a ea o 90 m² pe
plo . In each subplo , he pe cen age o co e o each plan species was
isually es ima ed. The mean alues o each species pe plo we e ob-
ained by a e aging he co e es ima es ac oss subplo s (B owe and Za ,
1978; Onaindia e al., 2004). Plan species and hei occu ences a e in
Table B.3.
Bi d sampling was ca ied ou by poin coun me hods a he same
cen al subplo es ablished o ege a ion sampling. Sampling was
conduc ed du ing he sp ing season, which coincides wi h he pe iod o
maximum b eeding ac i i y, om dawn un il wo hou s a e sun ise and
wi hou ad e se wea he condi ions. Samplings we e ca ied ou by he
same pe son. One plo o each o es sys em was sampled each day,
changing he o de e e y day o a oid e ec s o ime o day (Goded
e al., 2019). A each sampling poin , a 15-minu e obse a ion pe iod
was conduc ed, du ing which all bi d species obse ed o hea d we e
eco ded (P oença e al., 2010). Bi d species and hei occu ences a e in
Table B.4.
Soil sampling o mic obial communi y analysis was conduc ed a
h ee andomly selec ed subplo wi hin each sampling poin in o de o
cap u e spa ial he e ogenei y. A each cen oid, composi e soil samples
we e collec ed by combining h ee soil co es aken om a dep h o
20 cm. The sampling dep h was se a 20 cm, as his le el encompasses
he biologically ac i e su ace laye p io o he ansi ion zone in soil
mic obial communi ies (Guo e al., 2022). Soil samples we e i s ho-
mogenized and sie ed o emo e coa se deb is and oo s. F om each
homogenized sample, 250 mg o soil we e used o o al DNA ex ac ion
using he DNeasy Powe Soil P o Ki (Qiagen), in acco dance wi h he
manu ac u e ’s p o ocol, o ensu e he p ocu emen o high-quali y
nucleic acid o downs eam applica ions in sequencing. Ex ac ed
DNA was subsequen ly dispa ched o No ogene (Camb idge, UK) o
amplicon-based sequencing o bac e ial and ungal communi ies. The
16S RNA gene (V3 - V4 egion) was a ge ed o bac e ial iden i ica ion,
while he ITS2 egion was ampli ied o ungal iden i ica ion (Bu´
ee e al.,
2009; Ch is ensen e al., 2023). The PCR ampli ica ion was pe o med
using 15 µL o Phusion® High-Fideli y PCR Mas e Mix (New England
Biolabs), wi h a concen a ion o 0.2 µM o each o wa d and e e se
p ime and app oxima ely 10 ng o empla e DNA. The sequencing was
conduc ed on a pai ed-end Illumina pla o m, gene a ing 250 bp
pai ed-end eads. The aw sequences we e hen subjec ed o s ingen
quali y con ol measu es, which included p ime imming and quali y
il e ing. The denoising and chime a emo al p ocesses we e conduc ed
using he DADA2 pipeline, he eby yielding in high- esolu ion amplicon
sequence a ian s (ASVs). Finally, an a e age alue o ASV abundance
o each plo was calcula ed conside ing he h ee cen oids.
2.4. Func ional ai s assignmen
The unc ional ai s o plan s, bi ds, bac e ia, and ungal species
we e cha ac e ised o e alua e he unc ional di e si y and composi ion
o communi y unc ions. The selec ion o hese ai s was based on hei
ecological ele ance and hei po en ial o in luence ecosys em unc-
ioning. The desc ip ion o he unc ional ai s o each axon appea in
Appendix A.
The assignmen o plan unc ional ai s was conduc ed a he
species le el using he Flo aVeg.eu da abase. The ollowing ai s we e
assigned: li e o m ( ee, sh ub, he baceous, and e n); dispe sal mode
(endozoocho y, my mecocho y, anemocho y, dyszoocho y, and local
dispe sal); dispe sal dis ance (ca ego ised in o six dis ance classes,
anging om sho (1) o long dis ances (6)); Speci ic Lea A ea (SLA)
(low, medium, and high); shade ole ance (low, medium, and high); and
nu ien equi emen (oligo ophic, meso ophic, eu ophic). The ca e-
go isa ion o SLA and shade ole ance was conduc ed h ough he
implemen a ion o he Jenks na u al b eaks classi ica ion me hod. In he
case o shade ole ance, he da a we e de i ed om he Ellenbe g- ype
ligh indica o alue.
The unc ional ai s exhibi ed by bi d species we e assigned based
on ecological beha iou and mo phological cha ac e is ics. The da a
om Pedley e al. (2019) and Seobi dli e (h ps://seo.o g/) we e used o
assign he ollowing ai s: body mass (small and medium); o aging si e
(g ound, ege a ion, and mixed); nes ing si e (g ound, ca i ies, ee,
sh ub, and mixed); habi a ype ( o es species and non- o es species);
and die (insec i o ous and mixed-die ). The mean body mass o each
species was ca ego ised in o he wo g oups using he Jenks na u al
b eaks classi ica ion me hod.
In he case o bac e ia, i s ly a axonomic assignmen was conduc ed
using he SILVA 138.1 da abase o bac e ial 16S sequences, employing
a 97 % iden i y h eshold (Quas e al., 2013). Subsequen ly, hei
unc ional cha ac e is ics we e de e mined, by means o he FAPROTAX
da abase (Louca e al., 2016), a sys em ha enables he alloca ion o
unc ional oles based on axonomic classi ica ion. In ou da ase , only
21.4 % o bac e ial ASVs could be assigned o a leas one unc ional
g oup, while 78.6 % emained unassigned. This is a common limi a ion
in bac e ial unc ional anno a ion, as many soil axa emain poo ly
cha ac e ized, which may lead o unde es ima ion o ce ain unc ional
g oups. Func ions ha we e p esen in a minimum o i e s ands we e
selec ed. The ollowing unc ional g oups we e selec ed: ca bon me a-
bolism (chemohe e o ophy, pho ohe e o ophy, cellulolysis, a oma ic
hyd oca bon deg ada ion, e men a ion, and me hylo ophy); ni ogen
me abolism (ni ogen ixa ion, ni i ica ion, deni i ica ion, ni a e
espi a ion, and u eolysis); i on espi a ion; manganese oxida ion;
human-associa ed bac e ia; and animal pa asi es o symbion s.
In he case o ungi, he axonomic assignmen was conduc ed using
and he UNITE da abase o ungal ITS2 sequences, employing a 97 %
iden i y h eshold (Aba enko e al., 2023). Subsequen ly, he assign-
men o ungal unc ional ai s was conduc ed using he FUNGuild
da abase (Nguyen e al., 2016), which associa es ungal axa wi h
ecological guilds. In ou da ase , 34 % o ungal ASVs we e success ully
assigned o ecological guilds, whe eas 65 % could no be linked o
speci ic unc ional ca ego ies. The low assignmen a e e lec s known
gaps in ungal guild da abases, po en ially limi ing he de ec ion o
less-s udied ecological unc ions. I is impo an o no e ha he s udy
only encompassed guild assignmen s cha ac e ised by a high le el o
con idence (high p obable and p obable) and guilds ha we e p esen in
a minimum o i e s ands. The ollowing guilds we e conside ed: sap-
o ophs (wood, plan , dung, and unde ined); myco hizal ungi (ec o-
myco hizal and e icoid myco hizal); pa hogens (plan and animal);
pa asi es (plan and animal); and endophy es.
I is impo an o no e ha unc ional anno a ions de i ed om bo h
FAPROTAX and FUNGuild e lec po en ial me abolic o ecological
unc ions in e ed om axonomic iden i ies, a he han di ec ly
measu ed ac i i ies. This incomple e unc ional co e age mus be
conside ed when in e p e ing he ecological oles o mic obial com-
muni ies in ou s udy.
2.5. Species di e si y and composi ion
Plan and bi d species ichness and Shannon di e si y index
(Shannnon and Wea e , 1949) we e calcula ed o each s and. In he
case o plan s, he pe cen age o co e o each species was used,
whe eas o bi ds, he bi d abundance was conside ed. Concu en ly,
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
4
species composi ion was e alua ed using he B ay-Cu is dissimila i y
index.
In he case o bac e ia and ungi, alpha di e si y me ics we e
calcula ed a e a e ac ion o s anda d sequencing dep h. Speci ically,
he numbe o obse ed ea u es (ASV ichness) and he Shannon di-
e si y index we e compu ed. Fo composi ion, a phylogene ic ee was
cons uc ed using Fas T ee based on mul iple sequence alignmen s o
ASV ep esen a i e sequences. This ee, oge he wi h ASV abundance
da a, was used o compu e pai wise communi y dissimila i ies
employing he Weigh ed UniF ac me ic o each s and. This me ic in-
co po a es bo h phylogene ic ela edness and ela i e abundances o
axa.
2.6. Func ional di e si y and composi ion
The calcula ion o unc ional di e si y o he ou axonomic g oups
was conduc ed using Rao’s quad a ic en opy, a me ic ha accoun s o
bo h species abundances and unc ional ai dissimila i ies wi hin a
communi y. The unc ional dis ances be ween species we e compu ed
using Gowe ’s simila i y coe icien , and o imp o e he accu acy o he
esul s a squa e oo co ec ion was applied (Mammola e al., 2021).
Al hough we did no explici ly es o mul icollinea i y among unc-
ional ai s, he calcula ion o Rao’s Q inhe en ly accoun s o ai
edundancy h ough he use o a dis ance ma ix, which educes he
in luence o o e lapping ai in o ma ion.
Func ional composi ion was assessed using he p opo ional ep e-
sen a ion o each unc ional ai wi hin he communi y. These ela i e
abundances we e used o calcula e B ay-Cu is dissimila i y ma ices
be ween s ands. Collinea i y be ween ai s was no add essed p io o
NMDS o PERMANOVA, as hese a e explo a o y, non-pa ame ic ana-
lyses. T ai co ela ions we e expec ed due o he ca ego ical and
mu ually exclusi e na u e o ce ain ai s (e.g., SLA o shade ole ance),
bu all ai s we e e ained o p ese e ecological in e p e abili y.
2.7. S a is ical analysis
To compa e species di e si y ( ichness and Shannon index) and
unc ional di e si y among he ou o es sys ems, we i s assessed he
no mali y o he da a using he Shapi o-Wilk es . Homogenei y o a -
iances was hen e alua ed wi h Le ene’s es . Based on he ou comes o
hese p elimina y es s, di e en s a is ical app oaches we e pe o med:
when da a me bo h no mali y and homoscedas ici y assump ions, we
used one-way ANOVA ollowed by Tukey’s Hones Signi ican Di e -
ence es o pos -hoc pai wise compa isons. I he da a we e no mally
dis ibu ed bu showed unequal a iances, Welch’s ANOVA was pe -
o med, wi h pai wise di e ences e alua ed using he Games-Howell
es . In cases o non-no mal bu homoscedas ic da a, he K uskal-
Wallis es was used, ollowed by Dunn’s es wi h app op ia e co ec-
ion o mul iple compa isons. Finally, when nei he assump ions we e
me , a boo s ap-based p ocedu e was employed o assess pai wise di -
e ences obus ly.
In he case o species and unc ional composi ion, we compu ed a
P incipal Coo dina es Analysis (PCoA) and a Non-Me ic Mul idimen-
sional Scaling (NMDS) analysis, espec i ely, o isualize he di e ence
pa e ns o species composi ion. Subsequen ly, o bo h cases, a
Pe mu a ional Mul i a ia e Analysis o Va iance (PERMANOVA) was
conduc ed o examine he s a is ic di e ences among o es sys ems.
Finally, in o de o iden i y he unc ional ai s ha we e mos indica-
i e o each o es sys em, we conduc ed an Indica o Species Analysis a
unc ional le el. This acili a ed he iden i ica ion o speci ic unc ional
a ibu es ha cha ac e ised each sys em. All s a is ical es s we e pe -
o med using R so wa e ( 4.2.2; R Co e Team, 2024) in R S udio
pla o m (RS udio Team, 2023).
3. Resul s
3.1. Species and unc ional di e si y
Plan s species ichness and Shannon di e si y index we e ound o be
signi ican ly highe in Na i e and Res o a ion han in o es plan a ions.
The Eucalyp us exhibi ed he lowes alues o bo h di e si y me ics. In
he con ex o plan unc ional di e si y, Eucalyp us exhibi ed signi i-
can ly lowe alues in compa ison o he o he o es sys ems, wi h he
excep ion o Pine. Fu he mo e, he ichness o bi d species was ound o
be signi ican ly highe in Na i e han in o es plan a ions, and signi i-
can ly highe in Res o a ion han in Eucalyp us. In he case o Shannon
di e si y, Na i e and Res o a ion demons a ed signi ican ly highe
alues han Eucalyp us. Howe e , bi d unc ional di e si y did no
demons a e signi ican di e ences among any o es sys ems (Fig. 2;
Table B.2).
Wi h ega d o bac e ial species ichness and Shannon di e si y, no
signi ican di e ences we e obse ed among o es sys ems; howe e ,
unc ional di e si y was highe in Na i e and Res o a ion compa ed o
Eucalyp us. Ul ima ely, he ichness o ungal species did no demon-
s a e a signi ican di e ence among o es sys ems. In con as , Shan-
non di e si y was signi ican ly lowe in Eucalyp us han in Na i e and
Pine, while, unc ional di e si y was only lowe in Eucalyp us han in
Na i e (Fig. 2; Table B.2).
3.2. Species composi ion
In he con ex o plan species, Na i e was dis inguished om
Eucalyp us and Pine, as e idenced by he esul s o he PCoA (Fig. 3).
This esul was con i med by signi ican di e ences be ween Na i e and
Pine, and Na i e and Eucalyp us in he PERMANOVA esul s. In a simila
manne , Res o a ion di e ed signi ican ly om Pine and Eucalyp us, ye
no om Na i e, indica ing a eco e y o plan communi y composi ion
in es o ed si es. Finally, Pine and Eucalyp us also exhibi ed signi ican
di e ences in species composi ion be ween bo h (Table 1).
In he case o bi ds, he o dina ion exhibi ed a less clea ly de ined
sepa a ion among o es sys ems in compa ison o ha obse ed in
plan s (Fig. 3). Howe e , signi ican di e ences we e de ec ed be ween
he Na i e and Pine, and Na i e and Eucalyp us. In he case o Res o-
a ion, no signi ican di e ences we e obse ed wi h Na i e, Pine o
Eucalyp us (Table 1).
Fo bac e ial communi ies, he PCoA demons a ed conside able
o e lap among all o es sys ems, hough some di e en ia ion was
e iden be ween Eucalyp us and Res o a ion (Fig. 3). Indeed, Eucalyp us
showed signi ican di e ences wi h Na i e, Pine, and Res o a ion in
PERMANOVA analysis. Mo eo e , a signi ican di e ence was obse ed
be ween he Pine and Res o a ion, in con as o he non-signi ican
di e ences be ween he Pine and Na i e, and Na i e and Res o a ion
(Table 1).
In he case o ungal communi ies, a mode a e sepa a ion was
exhibi ed be ween Eucalyp us and he o he o es sys ems (Fig. 3). This
inding was co obo a ed by he PERMANOVA esul s, which e ealed
signi ican di e ences be ween Eucalyp us and he emaining o es
sys ems (Table 1).
3.3. Func ional composi ion
In plan s, Eucalyp us and Pine exhibi ed a g ea e di e gence om
bo h Na i e and Res o a ion, as e idenced by he NMDS analysis
(Fig. 4). Howe e , he PERMANOVA analysis e ealed signi ican di -
e ences among all o es sys ems (Table 2). Fu he mo e, he ollowing
ai s we e ound o be mo e associa ed wi h Na i e and Res o a ion: li e
o ms ( ees, and he bs), animal dispe sal (endozoocho ous and dys-
zoocho ous), dispe sal dis ance (low), SLA (high), shade ole an (high),
and nu ien equi emen (eu ophic and meso ophic). In con as , Pine
and Eucalyp us we e dis ibu ed in p oximi y o he ollowing ai s: li e
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
5
o ms ( e ns), dispe sal mode (anemocho ous and my mecocho ous),
dispe sal dis ance (high), shade ole ance (low and medium), and
nu ien equi emen (oligo ophic).
In bi ds, he NMDS analysis indica ed ha Eucalyp us we e sligh ly
sepa a ed om he o he o es sys ems (Fig. 4). Indeed, i s unc ional
composi ion di e ed signi ican ly wi h bo h Na i e and Pine, ye did no
di e om Res o a ion (Table 2). T ai s’ dis ibu ion is no clea ly
associa ed wi h o es sys ems bu Eucalyp us seemed o be mo e asso-
cia ed wi h he ollowing ai s: o aging si e ( ege a ion), habi a ype
(non- o es species), and die (insec i o ous). In con as , Na i e, Pine
and Res o a ion we e ela i ely mo e closely linked o he ollowing
ai s: body mass (medium), o aging si e (g ound and mixed), nes ing
si e ( ee and mixed), habi a ype ( o es species), and die (mixed-die ).
In bac e ia, he NMDS exhibi ed a dis inc sepa a ion be ween
Fig. 2. Boxplo s o species ichness, Shannon di e si y, and unc ional di e si y o plan s, bi ds, bac e ia, and ungi ac oss he ou o es sys ems: Na i e, Pine,
Eucalyp us, and Res o a ion. The cen al line o each box ep esen s he median, while he lowe and uppe edges co espond o he 25 h and 75 h pe cen iles,
espec i ely. Whiske s ex end o 1.5 imes he in e qua ile ange, and do s ep esen ou lie s. S a is ical di e ences among o es sys ems we e es ed using di e en
me hods depending on da a no mali y and homoscedas ici y (Table B.2). Di e en le e s indica e signi ican di e ences be ween g oups (p <0.05).
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
6
Eucalyp us and Res o a ion (Fig. 4). Indeed, he esul s o PERMANOVA
indica ed signi ican di e ences be ween hese o es sys ems, as well as
be ween Eucalyp us and Na i e, and Res o a ion and Pine (Table 2). The
dis ibu ion o unc ions ela ed o ni ogen me abolism and manganese
oxida ion we e dis ibu ed nea Res o a ion, Na i e and Pine plo s,
while hose unc ions associa ed wi h ca bon me abolism, i on espi a-
ion and human-associa ed bac e ia we e dis ibu ed in he p oximi y o
Eucalyp us.
In ungi, Eucalyp us exhibi ed dis inc i e unc ional composi ion
ha se i sligh ly apa om o he o es sys ems (Fig. 4). A de ailed
analysis e ealed ha i s unc ional composi ion di e ed signi ican ly
om ha o he o he sys ems (Table 2). Speci ically, he e icoid
myco hizal guild was iden i ied associa ed wi h Eucalyp us sys ems.
Howe e , i is e iden ha Na i e, Res o a ion and Pine we e mo e
Fig. 3. PCoA plo s ep esen ing communi y composi ion o plan s, bi ds, bac e ia and ungi ac oss he ou o es sys ems (Na i e, Res o a ion, Pine, and Euca-
lyp us). Fo plan s and bi ds, B ay-Cu is dis ances we e used, while o bac e ia and ungi Weigh ed UniF ac dis ances we e applied. Axes ep esen he i s wo
p incipal coo dina es, which explain he highes p opo ion o a iance in communi y dissimila i y. Small poin s indica e he s ands and he iangles indica e he
cen oid o each o es sys em.
Table 1
Resul s o PERMANOVA analysis o species composi ion o plan s, bi ds, bac e ia and ungi among he pai s o he ou o es sys ems (Na i e, Res o a ion, Pine, and
Eucalyp us). Signi icance: ***: p ≤0.001; **: p ≤0.01; *: p ≤0.05.
Abo eg ound axon Compa ison be ween o es sys ems p- alue Belowg ound axon Compa ison be ween o es sys ems p- alue
Plan s Na i e - Pine 0.002 ** Bac e ia Na i e - Pine 0.21
Na i e - Eucalyp us 0.001 *** Na i e - Eucalyp us 0.017 *
Na i e - Res o a ion 0.072 Na i e - Res o a ion 0.097
Pine - Eucalyp us 0.001 *** Pine - Eucalyp us 0.025 *
Pine - Res o a ion 0.001 *** Pine - Res o a ion 0.016 *
Eucalyp us - Res o a ion 0.001 *** Eucalyp us - Res o a ion 0.001 ***
Bi ds Na i e - Pine 0.019 * Fungi Na i e - Pine 0.48
Na i e - Eucalyp us 0.011 * Na i e - Eucalyp us 0.002 **
Na i e - Res o a ion 0.603 Na i e - Res o a ion 0.345
Pine - Eucalyp us 0.067 Pine - Eucalyp us 0.034 *
Pine - Res o a ion 0.306 Pine - Res o a ion 0.096
Eucalyp us - Res o a ion 0.052 Eucalyp us - Res o a ion 0.001 ***
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
7
Fig. 4. NMDS plo s ep esen ing he unc ional composi ion o plan , bi d, bac e ial, and ungal communi ies ac oss he ou o es sys ems (Na i e, Res o a ion,
Pine, and Eucalyp us). B ay-Cu is dis ances based on ela i e abundances o unc ional g oups we e used o compu e dissimila i ies be ween s ands. See he meaning
o plan and bi d ai s ac onyms in Table A.1 and A.2.
Table 2
Resul s o PERMANOVA analysis o unc ional composi ion o plan s, bi ds, bac e ia and ungi among he pai s o he ou o es sys ems (Na i e, Res o a ion, Pine,
and Eucalyp us). Signi icance: ***: p ≤0.001; **: p ≤0.01; *: p ≤0.05.
Abo eg ound axon Compa ison be ween o es sys ems p- alue Belowg ound axon Compa ison be ween o es sys ems p- alue
Plan s Na i e - Pine 0.001 *** Bac e ia Na i e - Pine 0.363
Na i e - Eucalyp us 0.002 *** Na i e - Eucalyp us 0.021 *
Na i e - Res o a ion 0.016 * Na i e - Res o a ion 0.226
Pine - Eucalyp us 0.033 * Pine - Eucalyp us 0.061
Pine - Res o a ion 0.001 *** Pine - Res o a ion 0.017 *
Eucalyp us - Res o a ion 0.001 *** Eucalyp us - Res o a ion 0.001 ***
Bi ds Na i e - Pine 0.161 Fungi Na i e - Pine 0.274
Na i e - Eucalyp us 0.011 * Na i e - Eucalyp us 0.001 ***
Na i e - Res o a ion 0.717 Na i e - Res o a ion 0.366
Pine - Eucalyp us 0.016 * Pine - Eucalyp us 0.02 *
Pine - Res o a ion 0.912 Pine - Res o a ion 0.211
Eucalyp us - Res o a ion 0.268 Eucalyp us - Res o a ion 0.002 **
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
8
closely associa ed wi h he ollowing guilds: sap o ophs (wood, plan
and dung) and pa hogens (plan and animal).
3.4. Func ional indica o s
In plan s, he ai s indica i e o Na i e and Res o a ion we e as
ollows: dispe sal mode (dyszoocho us) (p =0.010), dispe sal dis ance
(class 4) (p =0.001), and shade ole ance (high) (p =0.008). Mean-
while, he ollowing ai s we e indica i e o Na i e, Res o a ion and
Pine: SLA (high), and dispe sal mode (local dispe sal) and dis ance (class
2) (p =0.001 o all o hem). Concu en ly, he ai o bi ds nes ing si e
( ee) (p =0.009) and bac e ial g oups ela ed o ni ogen me abolism
(deni i ica ion (p =0.019) and ni a e espi a ion (p =0.031)) we e
iden i ied as indica o s o hem as well. In he case o ungi, no signi i-
can indica o s we e iden i ied.
4. Discussion
4.1. Ecological impac s o o es plan a ions
Despi e he long-s anding p esence and spa ial dominance o
monocul u e plan a ions in he UBR, hei biodi e si y emains ma k-
edly in e io o ha o na i e o es s. Indeed, he esul s ob ained
demons a e ha plan a ions o Pinus adia a and Eucalyp us sp. suppo
signi ican ly lowe species and unc ional di e si y, and a dis inc
composi ion o abo e- and belowg ound o ganisms, compa ed o bo h
na i e and es o ed o es s. Conc e ely, plan species and unc ional
di e si y in monocul u e plan a ions we e ound o be signi ican ly
lowe han in na i e o es s, as hese plan a ions end o simpli y o es
s uc u e, educe niche a ailabili y, and cons ain ecological p ocesses
(Liu e al., 2018; Wang e al., 2022; Vu Ho e al., 2023). Species and
unc ion composi ional analyses u he e ealed ha bo h species and
unc ional assemblages in plan a ions we e signi ican ly dis inc om
hose in na i e o es s. Indeed, ai s associa ed wi h ea ly-successional
o s ess- ole an s a egies, such as low shade ole ance, long-dis ance
dispe sal, oligo ophic a ini y, and p e alence o anemocho ous syn-
d ome o my mecocho ous synd ome, ha e been linked o pine and
eucalyp us s ands. These ai p o iles a e ypical o communi ies ha
a e es ablished in simpli ied o es s uc u es and unde esou ce-limi ed
condi ions, which a e cha ac e is ic o in ensi ely managed o es s
(B ocke ho e al., 2003; Mee s e al., 2010; Cal i˜
no-Cancela e al.,
2012). In con as , na i e and es o ed o es s we e associa ed wi h
zoocho ous dispe sal, high SLA, eu ophic condi ions, and highe shade
ole ance, e lec ing mo e ad anced successional s ages and inc eased
ecological complexi y (Suganuma e al., 2014; Blandino e al., 2021).
Canopy a chi ec u e could con ibu e o hese ecological con as s, as
he e ical lea o ien a ion o eucalyp us and he gene ally lowe c own
closu e o young pine plan a ions allow highe ligh pene a ion, he eby
p omo ing unde s o ey dominance by heliophilous and gene alis spe-
cies (Alday e al., 2017; Elosegi e al., 2020). Such species include he
e n P e idium aquilinum and he sh ubs like Rubus sp. and Ulex sp. ha
ha e been obse ed o agg essi ely colonise open o es loo s and can
domina e he unde s o ey in coni e ous o eucalyp us plan a ions
con ibu ing o unde s o ey homogeniza ion (Onaindia e al., 2013a;
Se u xa e al., 2024).
The mos ma ked di e gence was obse ed in Eucalyp us plan a ions
o all axonomic g oups. In he case o bi ds, Eucalyp us plan a ions
suppo ed a signi ican ly lowe species di e si y han Res o ed and
Na i e, and a dis inc species composi ion. Fu he mo e, he unc ional
composi ion was signi ican ly di e en om bo h he Na i e and Pine
sys ems, hus, being less associa ed wi h ai s such as o es specialis s.
These indings a e consis en wi h p e ious s udies in no he n Spain,
which ha e shown ha eucalyp us s ands ha bou only a nes ed subse
o o es bi d assemblages, leading o bio ic homogenisa ion and he loss
o o es specialis s (Goded e al., 2019). The quali y o a ian habi a is
cons ained by educed sh ub di e si y, low canopy he e ogenei y, and
he lack o nes ing subs a es (e.g., ca i ies and deadwood). Fu he -
mo e, Eucalyp us li e and ba k chemis y educe epiphy ic and
in e eb a e abundance, he eby diminishing ood esou ces o bi ds
(Cal i˜
no-Cancela.,2013; Ga cía-Fe nandez e al., 2025). This end was
u he con i med by he analysis o soil mic obial communi ies. Despi e
he absence o a signi ican di e ence in bac e ial and ungal species
ichness ac oss o es sys ems, a no able di e gence in unc ional di-
e si y and composi ion was obse ed, wi h Eucalyp us plan a ions
exhibi ing he mos p onounced dispa i ies. In hese plan a ions, bac-
e ial communi ies we e domina ed by unc ional g oups associa ed
wi h ca bon me abolism, i on espi a ion, and human- ela ed axa,
while unc ional g oups associa ed wi h ni ogen me abolism we e less
ep esen ed. This shi in bac e ial communi ies e lec s al e ed edaphic
condi ions, including lowe nu ien a ailabili y, mo e acidic soils, and
educed li e quali y, which a e ecognised as in luen ial conce ning
mic obial me abolism (Qu e al., 2020; Guo e al., 2024). Fungal com-
muni ies we e simila ly a ec ed by eucalyp us plan a ions and showed a
lowe Shannon di e si y o ungi han bo h Na i e o es s and Pine
plan a ions. In ac , eucalyp us plan a ions showed a clea associa ion
wi h e icoid myco hizal ungi ha is p obably ela ed o he a ailabili y
o E icaceae in he unde s o y o hese plan a ions, whe e high ligh
a ailabili y usually a ou s unde s o y de elopmen , in con as o de-
ciduous o es s. These species also end o exhibi my mecocho ous
dispe sal s a egies, which we e mo e p e alen in eucalyp us plan a-
ions, u he con ibu ing o he di e en ia ion o hei unc ional
composi ion (Cal i˜
no-Cancela.,2013). In addi ion, p e ious s udies in
he Ibe ian Peninsula ha e shown ha Eucalyp us plan a ions suppo
lowe ec omyco hizal di e si y compa ed o na i e o es s and o en
ha bou in oduced ungal axa om he ee’s na i e ange (Díez, 2005;
San olamazza-Ca bone e al., 2019). These in oduced axa may ha e
con ibu ed o he dis inc ungal communi y composi ion obse ed in
Eucalyp us plan a ions.
Beyond he pa e ns o axonomic and unc ional di e en ia ion, he
obse ed ai s and mic obial unc ions in Eucalyp us plan a ions indi-
ca e dis up ions in key ecological p ocesses. One o he mos e iden is
he al e a ion o nu ien cycling, pa icula ly he ni ogen cycle. In his
con ex , Eucalyp us plan a ions p oduce nu ien -poo li e ich in
polyphenols and ligni ied compounds, which inhibi mic obial ac i i y
and supp ess ni i ica ion, he eby educing soil ni a e a ailabili y and
impai ing ni ogen cycling (Zancada e al., 2003; Cas o-Díez e al.,
2021; Mallen-Coope e al., 2022). In pa allel, he p e alence o
ea ly-successional plan ai s such as anemocho y, my mecocho y, and
low shade ole ance sugges s a limi ed capaci y o s uc u al de elop-
men and unc ional ma u a ion, po en ially cons aining successional
p og ession and o es egene a ion (Cal i˜
no-Cancela e al., 2012).
Mo eo e , he unde ep esen a ion o o es specialis bi ds and lowe
bi d di e si y and ichness may u he limi bio ic in e ac ions such as
seed dispe sal, which a e essen ial o ecosys em eedbacks and plan
ec ui men (Ga cía-Fe n´
andez e al., 2025; Cal i˜
no-Cancela.,2013).
Impo an ly, such dis up ions a e likely d i en no only by he in insic
biological ai s o Eucalyp us plan a ions, bu also by he in ensi e
sil icul u al p ac ices commonly associa ed wi h i s cul i a ion and
ha es ing. Finally, i is impo an o acknowledge ha Eucalyp us
plan a ions we e es ablished p ima ily o imbe p oduc ion, no con-
se a ion. Howe e , in mul i unc ional landscapes such as biosphe e
ese es, unde s anding hei ecological ole emains ele an . In his
con ex , ou esul s indica e ha hese plan a ions con ibu e li le o
biodi e si y conse a ion, highligh ing he need o ca e ully balance
p oduc ion goals wi h ecological alues in land-use planning.
In con as , Pine plan a ions exhibi ed an in e media e posi ion
ac oss a conside able p opo ion o he ob ained esul s. Despi e a
decline in plan ichness and di e si y, a decline in bi d species ichness
and a di e en species composi ion when compa ed o bo h Na i e and
Res o ed sys ems, pine plan a ions did no di e signi ican ly in bi d o
mic obial di e si y when con as ed wi h na i e and es o ed o es s.
This pa e n may be pa ly explained by he highe p esence o na i e
U. O ega-Ba ue a e al.
Fo es Ecology and Managemen 596 (2025) 123057
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