Using fi e o enhance ewilding when ag icul u al policies ail
João C. Campos
a,
⁎, Julia Be nha d
b
, Nú ia Aquilué
c,d
,LluísB o ons
d,e,
, Jesús Domínguez
g
,ÂngelaLomba
a,h
,
B uno Ma cos
a,h
,Fe nandoMa ínez-F ei ía
a
, F ancisco Mo ei a
a,i
, Sil ana Pais
a,j
,
João P. Hon ado
a,h
,Ad iánRegos
a,g
a
InBIO/CIBIO - Cen o de In es igação em Biodi e sidade e Recu sos Gené icos, Campus Ag á io de Vai ão, Rua Pad e A mando Quin as, n° 7, 4485-661 Vai ão, Po ugal
b
Uni e si ä Leipzig, Ri e s aße 30, 36 04109 Leipzig, Ge many
c
Cen e d'É ude de la Fo ê , Uni e si é du Québec à Mon éal, Mon éal, Canada
d
InFo es J u (CTFC-CREAF), C a. An iga S Llo enç de Mo unys km. 2, 25280 Solsona, Spain
e
CREAF, 08193 Ce danyola del Vallès, Spain
CSIC, 08193 Ce danyola del Vallès, Spain
g
Depa amen o de Zooloxía, Xené ica e An opoloxía Física, Uni e sidade de San iago de Compos ela, 15782 San iago de Compos ela, Spain
h
Depa amen o de Biologia, Faculdade de Ciências da Uni e sidade do Po o, Rua do Campo Aleg e, S/N, Edi ício FC4, 4169-007 Po o, Po ugal
i
InBIO/CIBIO - Cen o de In es igação em Biodi e sidade e Recu sos Gené icos, Labo a ó io Associado, Ins i u o Supe io de Ag onomia, Uni e sidade de Lisboa, Tapada da Ajuda,1349-017 Lisboa,
Po ugal
j
p oMe heus, Resea ch Uni in Ma e ials, Ene gy and En i onmen o Sus ainabili y, Ins i u o Poli écnico o Viana do Cas elo, 4990-706 Pon e de Lima, Po ugal
HIGHLIGHTS
•HNV is a long- e m oppo uni y o fi e
supp ession and biodi e si y conse a-
ion.
•HNV managemen would benefi he
majo i y o species (mo e han 60%).
•Rewilding benefi s up o 20% o species,
including endange ed/endemic axa.
•Up o 33% o species p ofi om open
habi a s c ea ed by fi e.
•Fi e-media ed ewilding is a na u e-
based solu ion when ag icul u al
policies ail.
GRAPHICAL ABSTRACT
abs ac a icle in o
A icle his o y:
Recei ed 22 July 2020
Recei ed in e ised o m 13 Sep embe 2020
Accep ed 2 Oc obe 2020
A ailable online 14 Oc obe 2020
Edi o : Paulo Pe ei a
Keywo ds:
Biodi e si y conse a ion
High Na u e Value a mlands
Land abandonmen
Rewilding has been p oposed as an oppo uni y o biodi e si y conse a ion in abandoned landscapes. How-
e e , ewilding is challenged by he inc easing fi e isk associa ed wi h mo e flammable landscapes, and he
loss o open-habi a specialis species. Con as ingly, suppo ing High Na u e Value a mlands (HNV ) has been
also highligh ed as a aluable op ion, bu he e ec i e implemen a ion o ag icul u al policies o en ails leading
o unce ain scena ios whe ein he e ec s o wildfi e managemen emain la gely unexplo ed. He ein, we simu-
la ed fi e-landscape dynamics o e alua e how fi e supp ession scena ios a ec fi e egime and biodi e si y (102
species o e eb a es) unde ewilding and HNV policies in he u u e (2050), in a ansna ional biosphe e e-
se e (Ge ês-Xu és Moun ains, Po ugal-Spain). Rewilding and HNV scena ios we e modula ed by h ee di e -
en le els o fi e supp ession e ec i eness. Then, we quan ified scena io e ec s on fi e egime (bu ned and
supp essed a eas) and biodi e si y (habi a sui abili y change o 2050). Simula ions confi m HNV as a long-
e m oppo uni y o fi e supp ession (up o 30,000 ha o addi ional supp essed a eas be ween 2031 and 2050
Science o he To al En i onmen 755 (2021) 142897
⁎Co esponding au ho .
E-mail add esses: [email protected] (J.C. Campos), nu ia.aquilue@c c.ca (N. Aquilué), lluis.b o ons@c c.ca (L. B o ons), [email protected] (J. Domínguez),
angelalomba@ c.up.p (Â. Lomba), b uno.ma [email protected] (B. Ma cos), ma inez- ei [email protected] (F. Ma ínez-F ei ía), mo ei [email protected] (F. Mo ei a), jhon ado@ c.up.p
(J.P. Hon ado), ad ian. [email protected] ,ad ian. [email protected] (A. Regos).
h ps://doi.o g/10.1016/j.sci o en .2020.142897
0048-9697/© 2020 The Au ho (s). Published by Else ie B.V. This is an open access a icle unde he CC BY license (h p://c ea i ecommons.o g/licenses/by/4.0/).
Con en s lis s a ailable a ScienceDi ec
Science o he To al En i onmen
jou nal homepage: www.else ie .com/loca e/sci o en
Land-use
Sou he n Eu ope
Wildfi es
in compa ison o ewilding scena io) and o conse a ion (benefi ing a ound 60% o species). Rewilding benefi s
some species (20%), including c i ically endange ed, ulne able and endemic axa, while se e al species (33%)
also p ofi om open habi a s c ea ed by fi e. Al hough HNV emains he bes scena io, ewilding ein o ced
by low fi e supp ession managemen may p o ide a na u e-based solu ion when socie al suppo h ough
ag icul u al policies ails.
© 2020 The Au ho (s). Published by Else ie B.V. This is an open access a icle unde he CC BY license (h p://
c ea i ecommons.o g/licenses/by/4.0/).
1. In oduc ion
Land abandonmen in u al landscapes is one o he mos impo -
an d i e s o egional land-use change (Es oque e al., 2019), and
has been sugges ed as an oppo uni y o biodi e si y conse a ion
and he eins a emen o na u al ecological p ocesses (e.g. Quei oz
e al., 2014;Me ckx and Pe ei a, 2015). Rewilding app oaches ely
on his assump ion as hey in end o ees ablish na u al sel -
sus ained ecosys ems wi h educed human in e en ion (Gillson
e al., 2011). In Eu ope, ewilding ini ia i es (e.g. Rewilding Eu ope
Ne wo k; h ps:// ewildingeu ope.com) migh benefi biodi e si y
by p o iding enewed sui able habi a s and dispe sal co ido s, alle-
ia ing o es habi a agmen a ion (Pe ino e al., 2019). This man-
agemen s a egy migh be pa icula ly beneficial o a di e si y o
o es -adap ed species, including emblema ic axa o conse a ion
conce nsuchaswol sandbea s(Na a o and Pe ei a, 2015). Addi-
ionally, ewilded landscapes migh po en ia e a ious ecosys em
unc ions and se ices (Pe ino e al., 2019), such as he suppo o
soil, nu ien s and wa e cycles, he p o isioning o biomass and cli-
ma e egula ion (Na a o and Pe ei a, 2015).
Howe e , ewilding holds some cons ain s ha may limi i s suc-
cess ul implemen a ion. The inhe en homogeniza ion o ewilded
landscapes leads o loss and agmen a ion o open habi a s, mainly
due o sh ub enc oachmen and o es expansion (Mo ei a e al.,
2011). S udies also indica ed ha he g adual cessa ion o adi ional
a ming a eas, many o which known o suppo “High Na u e Value
a mlands”(he ea e HNV ), is a majo cause o local biodi e si y
losses, accele a ing popula ion declines o species adap ed o we g ass-
lands, pas u es and o he ex ensi e ag icul u al a eas (F anks e al.,
2018;Lomba e al., 2020). Fu he mo e, sh ub enc oachmen caused
by land abandonmen con ibu es o uel accumula ions and highe
connec i i y o flammable ege a ion, inc easing he po en ial inci-
dence, se e i y and impac s o la ge wildfi es (see Mo ei a e al., 2011
and e e ences he ein). These changes in he fi e- ege a ion dynamics
could cause se e e impac s on local ecosys em p ocesses and associa ed
biodi e si y, pa icula ly in egions whe e landscapes and bio a ha e
endu ed a long- e m ela ion wi h human ac i i ies and fi e egimes.
The Medi e anean egions o sou he n Eu ope a e pa adigma ic
cases o human- and fi e-media ed landscapes (Loep e e al., 2010;
Fe nandes e al., 2011;Viedma e al., 2015). The ma ked clima e season-
ali y o Medi e anean egions, cha ac e ized by wa m-d y summe s
and cold-humid win e s, p o ides op imal condi ions o equen and
in ense fi e e en s, mos ly concen a ed in he summe season. The
numbe o ex eme fi e e en s has escala ed in sou he n Eu ope du ing
he las decades (San-Miguel-Ayanz e al., 2013;Tedim e al., 2013;
Williams, 2013), mainly due o in ensified e ec s o clima e change
(wa me and d ie clima ic condi ions, and mo e equen hea h
wa es) and inc eased uel accumula ion and connec i i y caused by
u al land abandonmen (Fe nandes e al., 2016a;Tu co e al., 2019).
Fo hese easons, sou he n Eu opean coun ies ha e in es ed as e-
sou ces in fi e supp ession (Fe nandes e al., 2016b). Howe e , exhaus-
i e fi e supp ession migh be coun e -p oduc i e in fi e-p one
ecosys ems, since ege a ion will become dense and p o ide mo e
uel o la ge , mo e in ense and mo ese e e fi es —a p ocess desc ibed
as he “fi efigh ing ap”(Mo ei a e al., 2020). Consequen ly, al e na-
i e managemen s a egies ha e been ecommended, highligh ing
he u gency o a wide- anging policy shi owa ds fi e mi iga ion
and adap a ion (Fe nandes, 2013;Mo i z e al., 2014;Mo ei a e al.,
2020).
The in ica e links be ween land abandonmen and fi e egimes
complexi y he selec ion o app op ia e al e na i e scena ios, and sub-
sequen ly decision-making in fi e managemen and planning (Mo ei a
e al., 2011;McLauchlan e al., 2020). Conside ing he di e en p one-
ness o land co e (LC) ypes o bu n, ag icul u al policies implemen ed
unde he Eu opean Union's Common Ag icul u al Policy ha e been
claimed as an oppo uni y o educe fi e isk by p omo ing HNV in de -
imen o mo e flammable LC ypes, such as dense sc ublands (Mo ei a
and Pe'e , 2018). Al e na i e fi e managemen s a egies ha e been
e alua ed in he las decades o inc ease fi e supp ession e ec i eness
unde he ongoing land abandonmen ends. Fo ins ance, since wild-
fi es migh educe uel load and connec i i y be ween flammable ege-
a ion, se e al s udies ha e sugges ed ha allowing unplanned fi es o
bu n (unde sa e condi ions) would help supp essing u u e la ge o es
fi es unde ad e se clima ic condi ions(e.g. Hou man e al., 2013;Regos
e al., 2014;Duane e al., 2019).
Allowing low-in ensi y wildfi es o bu n migh also con ibu e o
biodi e si y conse a ion, an idea ad oca ed by he “py odi e si y
hypo hesis”, a p ominen concep in fi e ecology (Kelly and B o ons,
2017;Kelly e al., 2017). This hypo hesis elies on he assump ion ha
a high le el o landscape he e ogenei y, ela ed o spa ial- empo al a -
ia ions o esou ces and condi ions media ed by he in ensi y and se e -
i y o fi e egimes, benefi s biodi e si y (see Kellyand B o ons, 2017 and
e e ence he ein). This is s ill a con o e sial hypo hesis since s udies
ha e ound bo h posi i e (e.g. Ponisio e al., 2016;Tingley e al., 2016)
and nega i e (e.g. Taylo e al., 2012;Be y e al., 2015) e ec s o
py odi e si y on di e en biological communi ies. The incong uen
ou comes o py odi e si y unde line he complex ela ions be ween
biodi e si y, fi e egime, and landscape s uc u e and composi ion
(McLauchlan e al., 2020). In o ma i e assessmen s should hus con-
side how land-use change and fi e can a ec bo h he di e si y, a ail-
abili y and connec i i y o sui able habi a s o mul iple species (Kelly
and B o ons, 2017), p e e ably om di e en axonomic g oups.
He e, we assess he impac s o al e na i e landscape ajec o ies and
fi e supp ession managemen s a egies on u u e fi e egimes and on
biodi e si y conse a ion. We ocused on changes in bu ned and
supp essed a eas and habi a sui abili y o 102 e eb a e species
om h ee axonomic g oups (amphibians, bi ds, and ep iles) in a
ansbounda y p o ec ed moun ain egion be ween Po ugal and
Spain, ep esen a i e o he ypical sou he n Eu opean moun ain
landscapes unde going land abandonmen o a leas hal a cen u y
(Regos e al., 2015;Lasan a e al., 2017). We aimed o answe he ol-
lowingques ions: 1)How would di e en land-use and fi esupp ession
managemen scena ios con ibu e o u u e fi e mi iga ion (i.e. ewe
a eas bu ned associa ed wi h highe supp ession e ficiency)?; 2) How
would hose scena ios a ec biodi e si y?; 3) Which is he bes man-
agemen scena io o p omo ing fi e mi iga ion and biodi e si y conse -
a ion?; and 4) Wi h ag icul u al policies ailing o cope wi h u al
abandonmen , could a ewilding ajec o y in eg a ed wi h fi e supp es-
sion policies con ibu e o enhance biodi e si y conse a ion? We fi s
modelled he fi e- ege a ion dynamics om 2010 o 2050, unde wo
con as ing land-use managemen scena ios ( ewilding e sus HNV )
coupled wi h h ee fi e managemen s a egies di e ing in fi e
J.C. Campos, J. Be nha d , N. Aquilué e al. Science o he To al En i onmen 755 (2021) 142897
2
supp ession le els. Then, based on u u e p ojec ions o fi e- ege a ion
and on species dis ibu ion modelling, we quan ified he po en ial im-
pac s o hese con as ingpolicy op ions on fi e egime, landscape s uc-
u e, and hei subsequen e ec s on biodi e si y.
2. Me hods
2.1. S udy a ea
The s udy a ea is he T ansbounda y Biosphe e Rese e Ge ês-Xu és
(BR-GX; c.a. 2670 km
2
) in he NW Ibe ian Peninsula (Fig. 1). The ese e
encompasses ou Na u a 2000 si es and wo na ional p o ec ed a eas:
he Peneda-Ge ês Na ional Pa k (PGNP; Po ugal) and he Baixa Limia-
Se a do Xu és Na u al Pa k (BLXNP; Spain). This moun ain egion is o-
pog aphically complex, wi h al i udes a ying be ween 15 and 1545 m
and is composed by a di e sified o ien a ion elie , deep alleys, plains
and s eep slopes. The clima e is p edominan ly A lan ic, cha ac e ized
by high p ecipi a ion le els (a ound 1000 o 3000 mm/yea ), mos ly
concen a ed in au umn and win e seasons (Oc obe –Ma ch), al-
hough he a ied elie and wide- anging al i udes a o d di e se mi-
c oclima ic condi ions. The ese e is loca ed be ween he
Medi e anean and Eu osibe ian biogeog aphical egions, holding high
biodi e si y le els and se ing as e uge o se e al emblema ic species
o conse a ion conce n. The landscape is domina ed by sc ublands and
agmen ed o es s o deciduous ees (mainly oaks) and coni e s
(mainly pines). The ese e is expe iencing a long- e m u al exodus
(ha ing a cu en low popula ion o 29.4 inhabi an s/km
2
), which has
p og essi ely con ibu ed o sc ub and woodland expansion a he ex-
pense o c opland and pas u eland (Regos e al., 2015). The e ec s o
land abandonmen a e pa icula ly in ensified in he moun ainous and
ha dly accessible a eas o he ese e, ollowing con empo a y
endencies obse ed ac oss o he Medi e anean egions (e.g., Ga cía-
Ruiz and Lana-Renaul , 2011; an Leeuwen e al., 2019). S ill, he a ea
is subjec ed o a high equency o human-induced wildfi es (e.g. adi-
ional use o fi e; Chas-Amil e al., 2015), leading o a fi e egime cha ac-
e ized by ecu en fi e e en s and conside able o al bu ned a eas
(195,000 ha bu ned be ween 1983 and 2010) wi h a fi e e u n in e al
o 37 yea s. Despi e inc eased in es men s in fi e supp ession, hese e -
o s ha e been p o en unsuccess ul o alle ia e he local fi e egime
and he impac s o la ge wildfi es, a pa e n ha has been obse ed
ac oss o he moun ain u al a eas in sou he n Eu ope, such as in
Po ugal (e.g. Fe nandes e al., 2016b) and Spain (e.g. Loep e e al.,
2010;Pausas and Fe nández-Muñoz, 2012;Viedma e al., 2015), o
F ance (San Roman Sanz e al., 2013), I aly (U sino and Romano,
2014;Bajocco e al., 2019) and G eece (Xys akis e al., 2017).
2.2. Scena io design
Two land-use scena ios we e es ablished (see Fig. 2 and Table 1 o
de ails conce ning annual con e sion a es): (1) a ewilding scena io,
ep esen ing he ongoing end o land abandonmen epo ed in ou
s udy a ea (see Regos e al., 2015), as well as in o he moun ain egions
ac oss Sou he n Eu ope unde he cu en EU Common Ag icul u al Pol-
icy (CAP) (see Lasan a e al., 2017 and e e ences he ein); and (2) an
HNV scena io, eflec ing he adop ion o policies ha e ec i ely main-
ain and expand HNV ag icul u al a eas (i.e. a g eene pa h o he CAP;
Pe'e e al., 2014, 2020). Fo each land-use scena io, h ee fi e manage-
men scena ios di e ing in supp ession policies we e compa ed (Fig. 2):
(1) a scena io o le ing fi es bu n in o es and sc ubland a eas while
fi efigh ing esou ces a e p io i ized owa ds human asse s p o ec ion
(i.e. fi efigh ing e o s mos ly ocused on ag icul u al mosaics, in
which human asse s a e mos ly loca ed; low fi e supp ession-LS; i.e.
Fig. 1. Loca ion o he T ansbounda y Biosphe e Rese e Ge ês-Xu és in heIbe ian Peninsula ( hick black con ou s) and o hemain p o ec ed a eas o Peneda-Ge êsNa ional Pa k and he
Baixa Limia-Se a do Xu és Na u al Pa k (yellow con ou s).
J.C. Campos, J. Be nha d , N. Aquilué e al. Science o he To al En i onmen 755 (2021) 142897
3
using fi e o p omo e biodi e si y; see e iew in Kelly and B o ons,
2017); (2) a scena io ep esen a i e o cu en fi e supp ession ap-
p oaches and capabili ies (cu en fi e supp ession-CS; see model pa-
ame iza ion in Supplemen a y ma e ial in Pais e al., 2020); and
(3) a scena io wi h highe in es men in fi e supp ession (i.e. inc eased
fi efigh ing capaci y; high fi e supp ession-HS).
2.3. Fi e-landscape modelling and e ec s o managemen scena ios on fi e
egime
Fi e-media ed landscape dynamics we e add essed using a landscape
dynamic modelling app oach based on he REMAINS model (see Pais
e al., 2020 conce ning REMAINS model). B iefly, he model inco po a es
landscape change p ocesses d i ing fi e- ege a ion dynamics, gene a ing
dis inc scena ios o landscape dynamics based on p e iously designed
condi ions. The model simula es fi e igni ion, sp ead and ex inc ion, cali-
b a ed unde business-as-usual scena ios wi h he cu en fi efigh ing e -
ec i eness specifically de e mined o each adminis a i e egion o he
s udy a ea (please see Supplemen a y ma e ial in Pais e al., 2020), na u-
alsuccessionandpos -fi e egene a ion. Annual bu n a eas a e quan i-
fied o define a bu n a ea dis ibu ion and o consequen ly inco po a e
his o ical fi e egime in o he model. In addi ion, he fi e-succession
model is coupled wi h a land-use model o implemen land-use policies
a he landscape le el. Pa icula ly, he model used (1) dynamic a iables
in o ming abou he main LC ype, ime since las change o LC ype and
which ansi ions has ecen ly aken place; and (2) s a ic a iables, such
as oad densi y, ele a ion, slope and aspec . The a iables we e imple-
men ed in he model a 30-m esolu ion o he ese e. Then, he
model es ima ed he fi e igni ion p obabili y and simula ed he fi e
sp ead based on h ee majo d i e s. Fi s ly, he land-use changes (e.g.
u al abandonmen , ag icul u al in ensifica ion and o es con e sions)
a e simula ed by using a demand-alloca ion app oach whe e he quan i y
o change by ime s ep is based on his o ical land-use ends, which al-
lows emula ing landscape dynamics unde cu en land-use policies. Sec-
ondly, wildfi e and fi e supp ession, in which he fi e egime was
calib a ed by p e iously speci ying a a ge annual bu ned a ea based
on fi e s a is ical da a be ween 1983 and 2010 a he municipal le el o
he Spanish Minis y o Ag icul u e and Fishe ies, Food and En i onmen
(MAPAMA, 2018) o Spain, and on fi e pe ime e s o he pe iod
1990–2010 o he Ins i u e o Na u e Conse a ion and Fo es s (h ps://
www.icn .p /) o Po ugal. Thi dly, he ege a ion dynamics, including
na u al succession and pos -fi e egene a ion, which was compu ed by
iden i ying pos -fi e egene a ion cu es o each pixel o he s udy a ea
Fig. 2. Schema ic ep esen a ion o he s udy app oach. The land-use and fi e supp ession managemen scena ios implemen ed in his s udy a e p esen ed, as well as he esul ing
al e na i e scena ios es ed o imp o e fi e mi iga ion and biodi e si y conse a ion in he s udy a ea.
Table 1
Land-use and fi e supp ession scena ios implemen ed in he modelling simula ions. The
land-use scena io, he annual ege a ion con e sion a es unde each land-use policy
and he fi e supp ession le els a e p esen ed o each scena io.
Scena io
ac onym
Land-use
scena io
C opland o
sc ubland (ha)
Sc ubland o
c opland (ha)
Fi e supp ession
le el
RWild_LS RWild 400 0 Low
RWild_CS RWild 400 0 Cu en
RWild_HS RWild 400 0 High
HNV _LS HNV 0 400 Low
HNV _CS HNV 0 400 Cu en
HNV _HS HNV 0 400 High
J.C. Campos, J. Be nha d , N. Aquilué e al. Science o he To al En i onmen 755 (2021) 142897
4
by using a No malized Di e ence Vege a ion Index ime se ies acco ding
o Gu man and Igna o (1998).Finalfi e pe ime e s esul ed om he
spa io- empo al in e ac ions be ween he landscape i sel and he imple-
men a ion o fi efigh ing s a egies.
2.4. Species dis ibu ion modelling and e ec s o managemen scena ios on
biodi e si y
2.4.1. Species da a and en i onmen al a iables
In o de o assess he impac s o al e na i e managemen scena -
ios on biodi e si y, we buil species dis ibu ion models (SDMs) o
102 e eb a es (79 bi ds, 15 ep iles and 8 amphibians; Appendix
A) using a se o en i onmen al a iables. To do so, we ga he ed p es-
ence/absence da a om he PGNP (Pimen a and San a ém, 1996;
Soa es e al., 2005)and heBLXNP(Domínguez e al., 2005, 2012)
a lases. Bi ds da a we e a ailable a 2 km esolu ion o he PGNP
( o 1990–1995) and a 1 km ( o 1998–2000) and 2 km esolu ion
( o 2010) o he BLXNP. The da a o amphibians and ep iles we e
a ailable a 1 km esolu ion o bo h PGPN ( o 1998–2003) and
BLXNP ( o 2010). SDMs we e hen conduc ed on o a 1-km g id co -
e ing he en i e s udy a ea, using a minimum o 30 p esences pe
species (Appendix A).
En i onmen al a iables we e selec ed in acco dance wi h he
empo al pe iods o he a lases da a. Al i ude, slope and aspec
we e acqui ed a 30 m esolu ion om a Global Digi al Ele a ion
Model o he Ad anced Spacebo ne The mal Emission and Reflec ion
Radiome e (h ps://as e web.jpl.nasa.go /gdem.asp). LC a iables,
conc e ely pe cen age o e e g een o es s, deciduous o es s, ag i-
cul u al a eas, ocky a eas wi h spa se ege a ion and sh ublands,
we e ob ained om Landsa -de i ed da a o 2010, and om
REMAINS simula ions o he pe iod be ween 2020 and 2050,
unde he p e iously defined six managemen scena ios (RWild_LS,
RWild_CS, RWild_HS, HNV _LS, HNV _CS and HNV _HS). All en i on-
men al a iables we e consequen ly esampled in o 1 km o spa ial
esolu ion.
2.4.2. Model fi ing and e alua ion
We buil ou SDMs using six modelling echniques (Gene alized
Linea Models, Gene alized Addi i e Models, Random Fo es s, A ifi-
cial Neu al Ne wo ks, Gene alized Models o Boos ed Reg ession,
and Mul i a ia e Adap i e Reg ession Splines) a ailable in he R
package ‘biomod2’(h p:// - o ge. -p ojec .o g/p ojec s/biomod/).
The models we e fi using 80% o he da a, while he emaining 20%
we e used o model e alua ion h ough he a ea unde he cu e
(AUC) o a ecei e ope a ing cha ac e is ics (ROC; Fielding and
Bell, 1997). Subsequen ly, we buil consensus p edic ions based on
weigh ed a e ages o single-model p ojec ions o each species
(Ma mion e al., 2009;A aújo e al., 2011). Ensemble models we e
de i ed using a minimum o 10 model eplica es (see Appendix A
conce ning model eplica es and AUC quali y h esholds o each
species). These models we e hen p ojec ed o 2010 (one p ojec-
ion) and 2050 (using 40 fi e-landscape model uns pe manage-
men scena io, hus o aling 240 p ojec ions pe species). Finally,
consensus p edic ions we e eclassified in o p esence/absence
maps h ough ROC op imized h esholds a ailable in ‘biomod2’
(Thuille e al., 2009).
2.4.3. E ec s o managemen scena ios on biodi e si y
To e alua e he e ec s o land-use and fi e supp ession managemen
scena ios on biodi e si y, we calcula ed he p edic ed habi a sui abili y
change be ween 2010 and 2050 o each species. We fi ed mul iple lin-
ea eg ession models using he species habi a sui abili y changes as
dependen a iables, and fi e supp ession le el (low/cu /high) and
land-use managemen ( ewilding/HNV ) as independen a iables.
The e ec s o land-use and fi esupp ession managemen on speciesdis-
ibu ions we e assessed based on model significance s a is ics (Appen-
dix B). Finally, habi a sui abili y changes (be ween 2010 and 2050)
we e e alua ed acco ding o (i) species g oups significan ly a ec ed
by land-use and fi e supp ession managemen ; and (ii) species' egional
IUCN s a us (Appendix A).
2011−2030 2031−2050
Bu ned A ea Oppo unis ic Fi e Supp esion
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
0
50
100
150
200
250
0
20
40
60
Scena ios
Thousands hec a es
Land−use managemen
HNV
Rewilding
Fig. 3. Es ima ed bu ned and supp essed a eas ( ollowing an oppo unis ic fi e supp ession,i.e. adop ing open mosaics o igina ed om ag icul u al ac i i ies as oppo uni ies o s op fi es)
o wo u u e pe iods (2011–2030 and 2031–2050), acco ding o di e en land-use and fi e managemen scena ios. Poin s and ba s ep esen he mean and he mean s anda d e o ,
espec i ely.
J.C. Campos, J. Be nha d , N. Aquilué e al. Science o he To al En i onmen 755 (2021) 142897
5

3. Resul s
3.1. E ec s o managemen scena ios on fi e egime
Acco ding o ou simula ions, u u e fi es would bu n a ound
175,000 (RWild_CS) and 150,000 (HNV _CS) hec a es be ween 2031
and 2050 unde he cu en fi e supp ession sys em (mo e han 8500
and 7500 ha/yea , espec i ely). Scena ios wi h he highes fi e sup-
p ession le els would la gely educe bu ned a eas (Fig. 3). Howe e ,
fi efigh ing oppo uni ies om open a eas gene a ed by ag icul u al ac-
i i ies would become mos impac ul unde low supp ession scena ios
(see fi e supp ession in Fig. 3), wi h significan di e ences (up o
30,000 ha; i.e. 1500 ha/yea ) be ween ewilding and HNV scena ios
in he long e m (2031–2050) as a mland a eas g adually inc ease
(see “c opland”in Appendix C).
3.2. E ec s o managemen scena ios on biodi e si y
Ou ensemble models p esen ed high p edic i e accu acy (AUC =
0.92 ± 0.06; TSS = 0.72 ± 0.14; Appendix A) and e ealed di e en e-
sponses o biodi e si y o land-use and fi e supp ession managemen
(Fig. 4 and Appendix D).
Land-use managemen significan ly a ec ed mo e species (83%)
han fi e supp ession managemen (48%) (Fig. 4 and Appendix B).
HNV scena ios would inc ease habi a sui abili y o mo e han hal o
he species (N = 64), whe eas ewilding scena ios a e p edic ed o in-
c ease sui able habi a s o 20% o species (N = 21). Howe e , se e al
h ea ened and endemic species a e significan ly benefi ed by ewilding
scena ios (Fig. 5), including he only c i ically endange ed species (hab-
i a gains o mo e han 50%), and mos o he endange ed (3 ou o 5
species; Appendix E) and ulne able species (6 ou o 11 species;
15
34
53
0
20
40
High FS SE Low FS SE No FS e ec
% species
A
21
64
17
0
20
40
60
RWild SE HNV SE No land−use e ec
% species
B
RWild SE HNV SE No land−use e ec
High FS SE Low FS SE No FS e ec
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
−50
−25
0
25
−50
−25
0
25
−50
−25
0
25
Scena ios
% o change om 2010
Land−use managemen
Rewilding
HNV
C
Fig. 4. E ec s o managemen scena ios on biodi e si y. (A) Pe cen age o speciesbenefi ed by fi e supp ession (FS) managemen scena ios (2050), independen ly o he land-use scena io
and (B) pe cen age o species benefi ed by land-use managemen scena ios, independen ly o he fi e managemen scena ios (2050). (C) To al habi a sui abili y change (2010–2050)
acco ding o species g oups significan ly a ec ed by (1) land-use managemen (significan e ec s - SE; species wi h no significan e ec s a e plo ed as “No FS/land-use e ec s”) and
(2) fi e supp ession (FS) managemen scena ios, i.e. species significan ly a ec ed by high (High FS SE) and low (Low FS SE) fi e supp ession endencies, and wi h no significan e ec s
(No FS e ec ). Poin s and ba s a e ep esen ing he mean and he mean s anda d e o , espec i ely.
J.C. Campos, J. Be nha d , N. Aquilué e al. Science o he To al En i onmen 755 (2021) 142897
6
Appendix F). Ibe ian endemics would be pa icula ly benefi ed by HNV
unde low fi e supp ession le els (15% o habi a gain). S ill, se e al en-
demic species also p ofi om ewilding scena ios, conc e ely hose sig-
nifican ly a ec ed by high supp ession s a egies and hose no a ec ed
by fi e supp ession managemen (Fig. 5).
Conce ning he e ec s o fi e supp ession managemen , a high p o-
po ion o species (33%) is p edic ed o benefi om low fi e supp es-
sion (i.e., fi e-media ed landscapes), while only 15% p ofi om high
fi e supp ession, bo h in de imen o he cu en fi e supp ession sys-
em (Fig. 4).
4. Discussion
4.1. E ec s o managemen s a egies on fi e mi iga ion
The cu en endency o land abandonmen would inc ease he im-
pac s o u u e fi es (Fig. 3), which migh be explained by enc oachmen
o ege a ion domina ed by mo e flammable species in a eas p e iously
occupied by a mlands and spa se ege a ion (Appendix D). The imple-
men a ion o an HNV scena io could ep esen a mo e e ficien (and
cos -e ec i e) fi e mi iga ion s a egy in he long- e m (Fig. 3), as also
demons a ed in o he egions o he Ibe ian Peninsula (Aquilué e al.,
2020). Ou fi e- ege a ion simula ions ein o ce hese assump ions, by
demons a ing po en ial ad an ages o le ing unplanned fi es bu n o
inc ease fi e supp ession oppo uni ies de i ed om new open a eas
(Fig. 3). Le ing unplanned fi es bu n unde a o able wea he
condi ions in landscapes wi h highe a ailabili y o semi-open a eas
migh con ibu e o inc ease fi e-supp ession oppo uni ies by educing
uel accumula ion and p omo ing accessibili y o fi efigh ing (Duane
e al., 2019).
4.2. E ec s o managemen s a egies on biodi e si y
Th ee main findings should be highligh ed omou esul s: 1) he e
is a high a iabili y o biodi e si y esponses o di e en combina ions
o land-use and fi e supp ession managemen s a egies; 2) HNV man-
agemen would benefi he majo i y o species (mo e han 60%) in de -
imen o ewilding scena ios (Fig. 4); and 3) a subs an ial numbe o
species (mo e han a hi d) would p ofi om scena ios wi h mo e
fi e in he landscape (i.e., low fi e supp ession le els; Fig. 4). The majo -
i y o he s udied species p ofi ed om HNV scena ios, ein o cing ha
main aining some ag icul u al a eas con ibu es o biodi e si y a he
landscape le el (Halada e al., 2011;Lomba e al., 2020). The e ec i e
implemen a ion o HNV policies could ha e an impac ul epe cussion
on biodi e si y conse a ion in ou s udy a ea, gi en he small ex en
o ag icul u al a eas in compa ison o o he LC ypes such as sh ublands
(c.a. 60%) and o es s (c.a. 20%). Land abandonmen and subsequen
ege a ion enc oachmen leads o habi a losses o a mland specialis s
and o he species adap ed o open and semi-open a eas (Appendices B
and D).Howe e , o he species would benefi om hese condi ions, es-
pecially when associa ed wi h high fi e supp ession s a egies (Fig. 4;
Appendices B and D). These species a e mos ly ep esen ed by o es
High FS SE Low FS SE No FS e ec
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
−50
0
50
Managemen scena ios
% o change om 2010
IUCN s a us
CR
EN
VU
SE − Signi ican e ec
Fi e managemen e ec s (% habi a change) on h ea ened species (IUCN c i e ia)
A
High FS SE Low FS SE No FS e ec
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
RWild_LS
RWild_CS
RWild_HS
HNV _LS
HNV _CS
HNV _HS
−75
−50
−25
0
Managemen scena ios
% o change om 2010
Endemic species
SE − Signi ican e ec
Fi e managemen e ec s (% habi a change) on Ibe ian endemic species
B
Fig. 5. E ec s o managemen scena ios on h ea ened and endemic biodi e si y. (A) Changes (%) o habi a sui abili y o 2050 (in compa ison o 2010) o h ea ened ( egional IUCN
c i e ia) and (B) endemic species om he Ibe ian Peninsula. The species a e plo ed acco ding o he fi e supp ession (FS) managemen scena ios significan e ec s (SE), i.e. species
significan ly a ec ed by high (High FS SE) and low (Low FS SE) fi e supp ession endencies, and wi h no significan e ec s (No FS e ec ). Poin s and ba s ep esen he mean and he
mean s anda d e o , espec i ely.
J.C. Campos, J. Be nha d , N. Aquilué e al. Science o he To al En i onmen 755 (2021) 142897
7
bi d specialis s and amphibians (Appendices B and D), which a e mo e
dependen on we e and coole habi a s wi h dense ege a ion.
The impo ance o fi e lies in he c ea ion and main enance o open
habi a s, pa icula ly in a eas suscep ible o g adual a mland abandon-
men ha would p oduce mo e homogeneous and flammable land-
scapes. In ac , he posi i e esponse o a la ge g oup o species o
lowe fi e supp ession le els (Fig. 4; Appendices B and D) migh be p e-
cisely ela ed o he c ea ion o open spaces(and agmen a ion o wood
and sc ub ma ices) p omo ed by wildfi es. New open habi a s can be
po en ially ecolonized om su ounding habi a pa ches, a pa e n al-
eady acknowledged o ep iles and ea ly successional bi d species
(Regos e al., 2016;Che gui e al., 2019). S ill, p e ious s udies ha e
obse ed dis inc esponses o biodi e si y communi ies o fi e acco d-
ing o di e en pos -fi e eco e y and ecoloniza ion ajec o ies in
Medi e anean bu n a eas (B os e al., 2011;Ma eos e al., 2011;
San os e al., 2016). All hese con as ing pa e ns sugges a con ex -
dependen esponse (i.e. habi a dominance a he landscape le el and
habi a p e e ence o he species) o biodi e si y and bio ic communi-
ies o fi e-landscape dynamics (Pausas and Pa , 2018).
4.3. Implica ions o biodi e si y conse a ion and landscape managemen
4.3.1. Op imal managemen scena io: HNV wi h high fi e supp ession
The implemen a ion o HNV combined wi h high fi e supp ession
s a egies, al hough demanding a ca e ul s a egic planning o a oid ex-
eme fi e haza ds de i ed om fi efigh ing ap e ec s, would ep e-
sen an op imal managemen app oach ha could con ibu e o bo h
fi e mi iga ion and biodi e si y conse a ion (Fig. 4). The his o ical
human in e en ions ha shaped Medi e anean landscapes o e
millennia sus ained high le els o biodi e si y. The p omo ion o HNV
would benefi se e al species adap ed o hese semi-na u al habi a s
and dependen on he con inua ion o specific a ming sys ems (Fig. 4
and Appendix D), including many axa o conse a ion conce n cu -
en ly h ea ened by land abandonmen p ocesses (F anks e al., 2018;
Pais e al., 2020;Fig. 5). The con inuous abandonmen o hese p ac ices
will cause nega i e impac s o se e al species (Fig. 4 and Appendix D),
which highligh s he need o s able u al policies and long- e m socio-
economic p ojec s o sus ain ag icul u al ac i i ies in u al moun ain
a eas.
None heless, he successand con inua ion o HNV demands ailo ed
socio-economic and poli ical s a egies explo ing al e na i e and mo e
e ficien solu ions han economic incen i es (Lomba e al., 2020). In-
c easing capaci y building, socie al coope a ion and aising awa eness
o he alues o biodi e si y and ecosys em se ices in hese egions
a e pa amoun o sus aining he na u al and cul u al he i age associ-
a ed wi h hese adi ional ac i i ies. S ill, gi en he cu en ly domina -
ing endencies o land abandonmen and he di ficul ies o ein o ce he
cu en fi e supp ession sys em (due o he high cos s in ol ed and he
low accessibili y in s eep moun ainous a eas), HNV should be consid-
e ed as complemen a y o o he al e na i e managemen scena ios o
assu e cos -e ec i eness and adequa e implemen a ion.
4.3.2. Al e na i e managemen scena io: using fi e o enhance ewilding
Ou simula ions suppo ewilding as a beneficial scena io o o es -
adap ed species and axonomic g oups conside ed mos ulne able o
en i onmen al and landscape dis u bances (e.g. amphibians). Mo e-
o e , ewilding scena ios would p omo e s abiliza ion o sui able a eas
and conside able inc eases o a ailable habi a s o se e al h ea ened
and endemic species inside he ese e (Fig. 5). The habi a gains a e
pa icula ly no iceable o c i ically endange ed (Appendix E) and ul-
ne able axa (Appendix F), highligh ing he po en ial o ewilding poli-
cies o p o ec ing unique biodi e si y o conse a ion conce n.
Acco ding o hese esul s, and gi en he o e all ine ec i eness o
cu en Eu opean ag icul u al policies (Pe'e e al., 2014, 2019), ou
s udy sugges s an al e na i e and p ac icable scena io cen e ed on
fi e-media ed ewilding, consis en wi h he con inuous end o
depopula ion in he s udy a ea. In such a scena io, he po en ial haza ds
o ewilding would be bu e ed by low fi e supp ession s a egies, bo h
by con olling he e ec s o unplanned fi es and by using s a egic p e-
sc ibed fi es. This scena io would con ibu e o an oppo unis ic fi e
mi iga ion and o po en ial inc eases in py odi e si y o suppo ing
biodi e si y conse a ion (Kelly e al., 2015, 2017).
Al e na i e scena ios in eg a ing bo h ewilding and HNV manage-
men a he landscape le el should also be conside ed (Me ckx and
Pe ei a,2015;Lomba e al., 2020),since hei combinedimplemen a ion
would po en ially con ibu e o achie e di e en conse a ion goals
(e.g. conse a ion o biodi e si y adap ed o ag icul u al and o es ed
a eas) and o secu e a wide ange o ecosys em se ices (Pais e al.,
2020). Fu u e s a egic planning and decision-making in he egion,
howe e , should conside he p ospec i e ade-o s be ween fi e mi i-
ga ion, ecosys em se ices and conse a ion o di e en ace s o
biodi e si y.
5. Conclusions
This s udy con ibu es o he inc easing e idence o ag icul u al
policies as essen ial ools o ensu e biodi e si y while educing fi e
haza d, an aspec ha has been equen ly neglec ed when assessing
he beneficial e ec s o ag icul u al policies. Also, ou s udy sugges s
using fi e o enhance ewilding as an al e na i e managemen s a -
egy in ou s udy a ea —an issue ha decision make s and manage s
should conside when implemen ing ewilding ini ia i es in o he
fi e-p one egions. Addi ionally, ou s udy highligh s he need o
enewed poli ical and socio-economic e o s explo ing di e en so-
lu ions o economic incen i es and/o managemen s a egies
in eg a ing bo h ewilding and HNV . In his con ex , ou s udy dem-
ons a es how an e ec i e implemen a ion o Eu opean ag icul u al
policies could benefi biodi e si y ( h ough he c ea ion o new open
habi a s o endange ed species) while p o iding u he fi e-
supp ession oppo uni ies. Ou s udy also shows how fi e supp es-
sion policies can help he implemen a ion o ewilding ini ia i es in
o he abandoned, fi e-p one moun ain a eas ac oss Sou he n
Eu ope. I also goes beyond he business-as-usual scena ios and p o-
ides plausible u u e pa hways whe ein ewilding modula ed by
fi e supp ession can eme ge as na u e-based solu ion i he new EU
Common Ag icul u al Policy con inues o ail a e e sing u al aban-
donmen ends. F om a me hodological pe spec i e, ou s udy high-
ligh s he need o aking in o accoun he landscape dynamics and
fi e- ege a ion eedbacks when modelling u u e managemen sce-
na ios in moun ain landscapes o Sou he n Eu ope —highly dynamic
a eas his o ically d i en by he spa ial in e ac ion be ween fi e-
ela ed p ocesses (such as fi e and pos -fi e eco e y) and human in-
e en ions (land-use and fi e supp ession policies).
CRediT au ho ship con ibu ion s a emen
Concei ed and designed he expe imen s: JCC, AR. Expe imen s and
da a analyses: JCC, JB, AR. Con ibu ed da a/analysis ools: NA, JD, BM,
FM-F. D a p epa a ion: JCC. Visualiza ion: JCC. W i ing, e iew and
edi ing: all au ho s.
Decla a ion o compe ing in e es
The au ho s decla e ha hey ha e no known compe ing financial
in e es s o pe sonal ela ionships ha could ha e appea ed o influ-
ence he wo k epo ed in his pape .
Acknowledgemen s
The au ho s hank Ad ián Lamosa To es, Xosé Pa da ila and Albe o
Gil o hei help du ing fieldwo k in Xu és; and Ra ael Vázquez o p o-
iding addi ional da a o amphibians and ep iles.
J.C. Campos, J. Be nha d , N. Aquilué e al. Science o he To al En i onmen 755 (2021) 142897
8
Funding
This esea ch was suppo ed by Po uguese na ional unds
h ough FCT - Founda ion o Science and Technology, I.P., unde
he Fi ESma p ojec (PCIF/MOG/0083/2017), and by p ojec
INMODES (CGL2017-89999-C2-2-R) unded by he Spanish Minis y
o Science and Inno a ion.
AL, FM and FM-F we e suppo ed by FCT (DL57/2016/CP1440/
CT0001, IF/01053/2015; and DL57/2016/CP1440/CT0010, espec-
i ely). AR was suppo ed by he Xun a de Galicia (ED481B2016/
084-0) and IACOBUS p og am (INTERREG V-A España–Po ugal,
POCTEP 2014–2020).
Appendix A–F. Supplemen a y da a
Supplemen a y da a o his a icle can be ound online a h ps://doi.
o g/10.1016/j.sci o en .2020.142897.
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