Vol.:(0123456789)
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Jou nal o Pes Science
h ps://doi.o g/10.1007/s10340-022-01489-1
ORIGINAL PAPER
Landscape simpli ica ion inc eases Bac oce a oleae abundance
inoli e g o es: adul popula ion dynamics indi e en land uses
DanielPa edes1 · JoãoF ede icoAl es1· Sa aMendes1 · JoséMiguelCos a1 · JoanaAl es1 ·
An ónioAl esdaSil a1 · JoséPauloSousa1
Recei ed: 10 No embe 2021 / Re ised: 27 Janua y 2022 / Accep ed: 25 Feb ua y 2022
© The Au ho (s) 2022
Abs ac
Bac oce a oleae is he main pes in oli e g o es, and i s managemen equi es a sus ainable pe spec i e o educe he use o
chemical p oduc s. Landscape con ex is being conside ed as an impo an d i e o pes educ ion, bu esul s on B. oleae
show inconsis ency o da e. Mos o landscape-pes con ol s udies ocus on he dynamics o he pes s wi hin he ocal c op,
igno ing hese dynamics in o he land uses. He e we p esen a s udy in which we analyze he seasonal popula ion dynamics
o he oli e pes B. oleae in he mos impo an land uses o a ypical oli e landscape in Po ugal. We ound ha B. oleae is
p esen in all he land uses and he dynamics a e e y simila o hose in he oli e g o es. Howe e , he p esence o hese
land uses in he landscape did no display any inc ease in B. oleae abundance wi hin he oli e g o es. In con as , a landscape
mainly composed by oli e g o es inc eased he abundance o his pes . Impo an ly, mo e di e se landscapes su ounding
oli e g o es educe he abundance o he oli e ly. Based on hese indings, we can conclude ha B. oleae is p esen in all
he land uses o he s udied landscape bu ha his p esence does no imply an inc ease o B. oleae in oli e g o es. Indeed,
o he land uses can p omo e landscape di e si ica ion which is a d i e o he educ ion o B. oleae popula ions in oli e
g o es. We hus encou age oli e s akeholde s o inc ease landscape di e si ica ion a ound hei a ms by p omo ing/ es o -
ing o he c ops/habi a s.
Keywo ds Pes managemen · Land uses· Landscape di e si ica ion· Oli e ly
Key message
• Pes popula ion dynamics ou o he ocal c op has been
poo ly s udied.
• Bac oce a oleae was ound in all he land uses ha com-
pound a ypical oli e landscape in Po ugal.
• The p esence in o he land uses did no lead o a ise o
pes popula ion in he oli e g o es.
• Landscape domina ed by oli e g o es signi ican ly
inc eases he abundance o his pes .
• We encou age oli e g owe s o p omo e landscape di e -
si ica ion o educe he abundance o his pes .
In oduc ion
In he las decades, ag icul u e has passed h ough an in en-
si ica ion p ocess by d as ically inc easing he land de o ed
o a ming o ood p oduc ion (Foley e al. 2005). This p o-
cess has esul ed in he loss o na u al habi a s which in u n
has p o oked a loss o biodi e si y and a educ ion o he
ecosys em se ices associa ed wi h i (Di zo e al. 2014).
In addi ion, his landscape simpli ica ion p ocess inc eases
he likelihood o mo e damaging and ca as ophic pes ou -
b eaks (Pa edes e al. 2021), which has in ensi ied a me s’
dependence on insec icides. Howe e , he esis ance o pes s
o insec icides is augmen ing, e en exceeding he capaci y
o gene a e new p oduc s o comba hese pes s (Gould e al.
2018). Because o his, new policies a e p omo ing he use
o sus ainable solu ions o pes managemen ha do no
Communica ed by Ma io Balzan.
* Daniel Pa edes
[email p o ec ed]; a [email p o ec ed]
1 Cen e o Func ional Ecology, Associa ed Labo a o y
TERRA, Depa men o Li e Sciences, Uni e si y
o Coimb a, Calçada Ma im de F ei as, 3000-456Coimb a,
Po ugal
Jou nal o Pes Science
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imply he use o his kind o p oduc s (Eu opean Comission
2020). Al hough mos o he echniques ha ha e been es ed
a e o ien a ed owa d i s implemen a ion a he local le el,
ecologis s, ag onomis s and a me s a e inc easingly ecog-
nizing he c i ical ole ha su ounding landscapes can play
in de e mining pes damage (Thies and Tscha n ke 1999;
Bianchi e al. 2006).
Simpli ied landscapes (i.e., expansi e monocul u es) can
inc ease pes ou b eaks because hey p omo e specialized
pes popula ions o de elop and sp ead mo e easily, whe eas
complex landscapes (i.e., mosaics o di e en land uses)
can be a ba ie ha hampe his expansion (Roo 1973;
O’Rou ke and Pe e sen 2017). On he o he hand, di e si ied
landscapes can p omo e pes na u al enemies (p eda o s and
pa asi oids) by p o iding esou ces such as shel e , nec a ,
pollen and/o al e na i e p ey o hei op imal de elopmen ,
hus imp o ing hei ac ion agains pes s (Landis e al. 2000;
Chaplin-K ame e al. 2011). The e o e, plan ing mul iple
c ops o e aining non-c op ege a ion may lowe pes densi-
ies and educe insec icide applica ions (Dainese e al. 2019;
Pa edes e al. 2021). Howe e , pes s can also use o he land
uses in he landscape o hei de elopmen , hus inc eas-
ing hei p esence in o he c op (Tscha n ke e al. 2016).
Fo example, in A ica, co n s embo e s appea ed o be
p omo ed by g asslands (Midega e al. 2014), which aligns
wi h ano he s udy in Aus alia ha also showed na u al
g asslands as a sou ce o pes (Pa y e al. 2015). Pe haps,
he mos c i ical case is he in asi e pes species D osoph-
ila suzukii ha seems o be posi i ely a ec ed by na u al
habi a s a he landscape scale, by using al e na i e hos s
o h i e (San oiemma e al. 2018) and also because hese
habi a s p o ide be e p o ec ion agains ad e se wea he
condi ions du ing win e and summe pe iods (San oiemma
e al. 2019). Thus, he inconsis ency o pes esponses o
landscape complexi y p e en s esea che s o emi clea con-
clusions abou he ole o landscape on pes con ol (Ka p
e al. 2018). I also highligh s he impo ance o pes species’
ai s o elucida e i s esponses o landscape composi ion
(Tambu ini e al. 2020) as well as on how hey in e ac wi h
he di e en elemen s in he landscape.
Bac oce a oleae (Rossi) is he majo oli e pes ha
induces economic losses wo ldwide o aling app oxima ely
15% pe yea (Mon iel-Bueno and Jones 2001). The e ec
o landscape con ex on his pes is no well de ined. Fo
example, a s udy pe o med in sou he n Spain showed a
educ ion o his pes in a eas wi h mo e na u al habi a edge
densi y, landscape di e si y and highe numbe o pa ches
o na u al ege a ion (O ega and Pascual 2014). Howe e ,
ano he s udy ca ied ou in I aly did no ind any e ec o
na u al habi a s on B. oleae abundance (Picchi e al. 2016),
whe eas ano he wo k de eloped in sou he n Spain showed
a posi i e e ec o na u al habi a s on B. oleae abundance
(Manjón-Cabeza e al. 2017). The s ochas ici y ha pes
popula ions ha e may be behind hese con as ing esul s
(Chaplin-K ame e al. 2013; Pa edes e al. 2021). Howe e ,
nei he o hese s udies no mos o he o he s ega ding he
e ec o landscape on o he pes s (bu see San oiemma e al.
2019) ha e e e explo ed whe he he popula ion dynamics
o his pes in o he land uses is di e en o simila om ha
in he oli e g o e.
E en mo e, he oli e ly has a pe iod du ing he yea in
which i disappea s om he oli e g o es. This is called he
“whi e pe iod” ha akes place in he mon hs o May and
June when he abundance o oli e ly in he ield suddenly
d ops, and he e a e no oli e ui s o a ack (Michelakis and
Neuenschwande 1981). Be o e ha pe iod, he e is an en i e
gene a ion o lies (Ma ch–May) (Ma chini e al. 2017) ha
can only lay eggs on he incipien oli e ui s (Boccacio and
Pe achi 2009). By he end o he summe , a e he “whi e
pe iod”,a new gene a ion begins o ise un il he win e
(Ma chini e al. 2017). Du ing he “whi e pe iod,” he oli e
ly can be ound wi hin he oli e ui as la ae, bu he
oli e ly in headul s agecan also mo e in he landscape
looking o o he esou ces ha ul ima ely can ein o ce i s
popula ion when i s a s ising in Au umn. The e o e, o
unde s and he popula ion dynamics o B. oleae in o he land
uses is c i ical o unde s and he impo ance o a landscape
pe spec i e in he managemen o his pes and how can we
be e p omo e na u al pes con ol mechanisms based on
landscape managemen (Daane and Johnson 2010).
He e we pu pose a s udy in which we in es iga e he
dynamics o B. oleae in he di e en land uses ha com-
pound a ypical oli e landscape in Po ugal. We aim a
answe ing he ollowing ques ions: (1) To wha ex en is
Bac oce a oleae p esen in he di e en land uses, includ-
ing oli e g o e? (2) Which a e he dynamics o B. oleae
in hese land uses? and (3) How di e en land uses a he
landscape scale a ec he abundance o Bac oce a oleae in
oli e g o es?
Ma e ial andme hods
S udy si es andinsec sampling
To achie e he objec i es o his s udy, a o al o 79 sam-
pling poin s dis ibu ed among he mos common land uses
de ec ed in he sampling a ea loca ed in he Bei a In e io
egion o Po ugal wi hin he municipali ies o Cas elo
B anco and Idanha-a-No a we e selec ed (Fig.1). The
land uses de ec ed in his a ea we e sh ublands, eucalyp us
o es s, pine o es s, “mon ado” (Ibe ian oaks sa annahs),
g asslands, ineya ds and oli e g o es. As i is he main land
use o he s udy and he place whe e he ac i i y o B. oleae
akes place, o he o al o 79 sampling poin s, 25 poin s
we e loca ed in oli e g o es. The wen y- i e oli e g o es
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we e mos ly cen ena y o he same a ie y (Galega), non-
i iga ed, wi h low g oundco e ege a ion due o li es ock
p esence and we ecul i a ed a he momen o sampling.
Du ing his s udy, oli e g owe s did no apply pes icides and
did no use land plouwing me hods. The es o sampling
poin s we e dis ibu ed among he o he land uses a he a e
o nine o each one (Fig.1).
To de e mine B. oleae abundance, a McPhail ap was
placed a each one o he poin s selec ed in Feb ua y 2019
(Fig.1). Each ap was illed wi h a liquid con en consis -
ing o a 250mL aqueous solu ion wi h 5% diammonium
phospha e and 2% bo ax, which is e y e ec i e in a ac ing
B. oleae adul s (Po cel e al. 2009). Samples we e mon hly
collec ed o one yea s a ing in Feb ua y 2019 and ending
in Janua y 2020. A e collec ion in he ield, samples we e
anspo ed o he labo a o y whe e indi iduals o B. oleae
we e coun ed.
Landscape analysis
A each o he 25 sampling poin s loca ed in oli e g o es,
a geospa ial analysis o he su ounding bu e a ea wi h a
adius o 500m was pe o med using QGIS so wa e (Open-
Sou ce Geospa ial Founda ion. Bea e on. OR. USA), a
Geog aphic In o ma ion Sys ems (GIS) pla o m. Based on
ae ial pho og aphs, we delinea ed polygons by ep esen ing
he di e en pa ches o he di e en land uses ound in he
s udy a ea al eady men ioned abo e. The minimal digi ized
polygon size was 0.8ha. associa ed wi h a ineya d land
use. To alida e misalignmen o he landscape elemen s,
in addi ion o adding da a o he elemen s ha canno be
iden i ied om he ae ial pho og aphs, i was necessa y o
pe o m a isual ield alida ion o con i m he land use o
each selec ed poin . All o his geospa ial in o ma ion was
con e ed o as e images and analyzed in o F ags a s so -
wa e (Uni e si y o Massachuse s. Amhe s , MA. USA).
F om his spa ial pa e n analysis, we ob ained class-le el
landscape me ics, in which he o al a ea o each land use
wi hin each o he landscape bu e zones was quan i ied
as a pe cen age. F om hese alues, we hen ob ained he
Shannon’s di e si y index (SHDI) ha we used as a p oxy
o landscape di e si y. To es ima e landscape simpli ica ion,
we used he p opo ion o he oli e land use su ounding he
sampling poin .
Modeling
To de e mine he p esence o B. oleae in he di e en land
uses p esen a he s udy a ea, we i s added all he indi idu-
als coun ed a each sampling da e o ge a single alue pe
sampling poin . We hen s ablisheds a is ically signi ican
di e ences be ween se s o pai s o land uses by using a non-
pa ame ic Wilcoxon ank sum es wi h p- alues adjus ed
by he Benjamini & Hochbe g me hod (Benjamini and
Hochbe g 1995). Then, o be e ep esen a ion in a box-
plo we log- ans o med he da a used o pe o m his es .
G asslands
Legend
Ecualyp us o es s
Sh ublands
O he landuses(e.g.,u ban a eas)
Oli eg o es
Mon ado
Pine o es s
Vineya ds
ab
0Km5Km10 Km
Fig. 1 Loca ion o he s udy a ea in Po ugal (a) and dis ibu ion o sampling poin s ac oss he di e en land uses o he s udy a ea (b)
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To accoun o he dynamics o B. oleae in he di e -
en habi a s du ing one yea , we used Gene alized Addi-
i e Mixed Models (GAMMs) wi h a p edic o in he o m
o an in e ac ion be ween he Julian day in which he aps
we e sampled and he land use in which hey we e loca ed.
Numbe o kno s we e se a i e o allow o he ep esen a-
ions o he luc ua ions o he popula ion a each land use.
Finally, we added he loca ion o he ap (sampling poin ) as
a andom ac o o con ol o he pseudo eplica ion o e he
ime as se e al samples we e collec ed in he same poin a
di e en momen s o he yea . We i s modeled his combi-
na ion o ac o s wi h a Poisson e o dis ibu ion o being
he na u e o he esponse a iable a coun bu a e es ing
o o e dispe sion, we inally decided o model i wi h a
nega i e binomial e o dis ibu ion wi h a log link unc ion
o accoun o o e dispe sion. To accoun o model s abili y,
we pe o med he popula ion dynamic models o each one
o he land uses sepa a ely and ob ained e y simila esul
in e ms o signi ican dynamics (p- alue < 0.05).
To accoun o he e ec o he di e en land uses on B.
oleae abundance in oli e g o es, we also used he pooled
da ase applied in he i s analysis ha con ains a single
alue o each one o he 25 oli e g o es sampled. We hus
decided o pe o m a model selec ion app oach based on
he Akaike in o ma ion c i e ia co ec ed o small sample
size (AICc) o choose he bes model. Models wi h he low-
es AICc and hose wi h a di e ence o less han wo AICc
uni s om he lowes we e chosen o u he explana ion.
Thus, we c ea ed a se o gene alized linea models each one
including as p edic o he pe cen age o a single land use
ha we e: oli e, sh ublands, “mon ado,” g asslands, euca-
lyp us o es , pine o es s, ineya ds and Shannon di e si y
index. We comple e his se o models wi h a null model
o accoun o non-e ec s. Simila ly o popula ion dynamic
models, we i s op ed o a Poisson e o dis ibu ion bu
a e es ing o o e dispe sion we decided o model he da a
wi h a nega i e binomial e o dis ibu ion and log link unc-
ion. All analyses we e pe o med in R using he packages
“mgc ” o GAMMs (Wood 2011), “lme4” o GLMs (Ba es
e al. 2015), “DHARMa” o o e dispe sion analysis (Ha ig
e al. 2018) and “ggplo 2” o isualized esul s (Wickham
2016).
Resul s
A o al o 10,315 B. oleae indi iduals we e collec ed and
coun ed du ing he whole expe imen . They we e p esen in
all he land uses es ed in his s udy (Fig.2). As expec ed,
he highes abundance was ound in oli e g o es wi h a o al
o 8290 indi iduals. In he land use “mon ado,” a o al o
794 indi iduals we e apped ollowed by eucalyp us o es s
in which 634 indi iduals we e collec ed. Fo he o he land
uses, he o al numbe o cap u ed was lowe , wi h 246 indi-
iduals in pine o es , 196 in ineya ds, 114 in sh ublands
and only 34 in g asslands. These di e ences a e e lec ed in
Fig.2 and align wi h he esul s ob ained in he Wilcoxon
es (Table1). Oli e g o es appea s as he land use ha -
ing signi ican di e ences wi h all he es o he land uses.
“Mon ado” showed di e ences wi h g asslands, pine o es
Fig. 2 Boxplo o he loga i hm o he abundance o B. oleae o he
di e en land uses in ol ed in he s udy. The op and bo om o he
box a e i s qua ile (Q1) and hi d qua ile (Q3), and he cen e line
is median (Q2). Whiske s ep esen 1.5 imes he IQR (Q3–Q2) in
ela ion o he uppe and lowe qua iles. Poin s ep esen helog o
he abundance a each sampling poin
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and oli e g o es bu no wi h eucalyp us o es s and ine-
ya ds. Simila esul s we e ob ained o eucalyp us o es s.
Sh ublands only showed signi ican di e ences wi h oli e
g o es and “mon ado.” A simila si ua ion occu ed wi h
pine o es and g asslands, bu hese land uses also showed
signi ican di e ences wi h eucalyp us o es s. Finally,
ineya ds only displayed signi ican di e ences wi h oli e
g o es.
Pes dynamics we e signi ican in i e o he se en land
uses es ed in his s udy (Fig.3). Speci ically, B. oleae
showed signi ican dynamics in oli e g o es, “mon ado,”
eucalyp us o es s, g asslands and ineya ds (Table2). Bac-
oce a oleae popula ion in oli e g o es s a ed aising in
Table 1 S a is ically signi ican
di e ences be ween se s o
pai s o land uses by using a
nonpa ame ic Wilcoxon ank
sum es wi h p- alues adjus ed
by he Benjamini & Hochbe g
me hod
Signi ican di e ences (p < 0.05) highligh ed in bold
Eucalyp us o es s Sh ublands Mon ado Oli e g o es G asslands Pine o es s
Sh ublands 0.053
Mon ado 0.691 0.029
Oli e g o es < 0.001 < 0.001 0.005
G asslands 0.006 0.442 0.009 < 0.001
Pine o es s 0.027 0.376 0.029 < 0.001 0.685
Vineya ds 0.329 0.442 0.156 < 0.001 0.074 0.137
Fig. 3 Bac oce a oleae empo al dynamics in he di e en land
uses ha compound he s udy along wi h he ep esen a ion o all he
dynamics oge he . Blue, pink, g een, b own and ed lines ep esen
g asslands, ineya ds, oli e g o es, “mon ado” and eucalyp us o -
es s, espec i ely. Shadowed a eas ep esen he con iden in e als a
95%
Table 2 Signi icance o alues o oli e ly popula ion dynamics in
he di e en land uses used o he Gene alized Addi i e Model
Va iable p- alue
Popula ion dynamics in Eucalyp us o es s < 0.001
Popula ion dynamics in Sh ublands 0.371
Popula ion dynamics in Mon ado 0.003
Popula ion dynamics in Oli e g o es < 0.001
Popula ion dynamics in G asslands 0.019
Popula ion dynamics in Pine o es s 0.615
Popula ion dynamics in Vineya ds < 0.001
R2 0.204
De iance explained 70.6%

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Ma ch, eaching a peak in May. Then, i sligh ly dec eased
un il mid-July when s a ed no ably spiking o ind he maxi-
mum abundance in mid-Oc obe o hen dec ease du ing
he win e . G asslands and ineya ds showed a e y simi-
la pa e n ha e lec ed a sus ained g ow h om Ma ch o
Oc obe and hen dec ease du ing he win e . In con as , he
dynamics o B. oleae in eucalyp us o es s and “mon ado”
displayed a wo-hump pa e n wi h an in ense g ow h om
Feb ua y o mid-Ap il and hen a dec ease inding he bo -
om a ound July and Augus and hen expe ienced ano he
g ow h ha peaked in he beginning o No embe . These
dynamics we e e y simila o he one in oli e g o es, bu
he in ensi y o he i s inc ease was highe han in oli e
g o es, especially o eucalyp us o es s (Fig.3).
Rega ding he e ec o he di e en land uses on he
abundance o B. oleae in oli e g o es, he selec ed mod-
els displayed a posi i e e ec su ounding oli e g o es
(Es ima e = 0.015: p- alue = 0.019) and a nega i e e ec o
he Shannon di e si y index on B. oleae abundance (Es i-
ma e = − 0.734: p- alue = 0.045) (Table3; Fig.4). The abun-
dance o B. oleae inc eased mo e han wo imes om oli es
g o es wi h o he land uses su ounding hem o places wi h
a e y simpli ied landscape > 90% o oli e g o es in he
bu e a ea). In con as , landscape di e si ica ion displayed
a nega i e e ec educing he abundance o B. oleae wo
imes om less di e si ied o mo e di e si ied landscapes.
I is impo an o no ice ha he pa allel e ec ha shows
he ends o landscape simpli ica ion and landscape di e -
si y is due o he high nega i e co ela ion among hese wo
p edic o s.
Discussion
This s udy demons a ed ha B. oleae is p esen in he main
land uses ha comp ise his ypical Po uguese oli e land-
scape. As expec ed, maximum abundance was ound in
oli e g o es, bu also impo an abundances we e ound in
Table 3 Akaike alues
co ec ed o small sample size
(AICc) o he di e en models
gene a ed o es ing he e ec
o he amoun o a land use in
he landscape on he abundance
o he pes Bac oce a oleae
Minimum AICc in bold
Land use AICc
Null 345.45
Mon ado 363.32
Eucalyp us o es s 347.87
G asslands 345.68
Pine o es s 346.20
Sh ublands 347.91
Vineya ds 347.81
Oli e g o es 341.46
Shannon di e si y 342.91
Fig. 4 Es ima ed e ec s o g adien s o landscape di e si y (b own line) and landscape simpli ica ion (g een line) on Bac oce a oleae abun-
dance. Shadowed a eas ep esen he con iden in e als a 95% (N = 25)
Jou nal o Pes Science
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eucalyp us o es s and “mon ado.” Ou expe imen al design
does no allow us o in e he easons o he p esence o B.
oleae in he di e en land uses, bu we can hypo hesize ha
hey a e no h i ing on al e na i e hos s due o B. oleae
cha ac e is ics as a specialis oli e pes (Daane and Johnson
2010; Cla ke 2018). In con as , he ac ha his ly appea s
in all he land uses es ed may be ela ed o i s dispe sal
beha io , being able o ly a ound 400m pe week (Fle che
and Kapa os 1981). When popula ions s a o build, new
adul lies dispe se o o he oli e a eas o de elop he nex
gene a ion (Kouna idis e al. 2008). Du ing his mig a ion,
hey can pass h ough pa ches o o he land uses in which
hey es and sea ch o esou ces o su i e. Consequen ly,
i makes sense ha he dynamics in o he land uses has o be
ela ed o he dynamics in he oli e g o es as hey a e going
o ac as a sou ce o oli e lies. Ne e heless, his pa allelism
is no pe ec in all he land uses. Fo example, he popula-
ions in ineya ds and g assland peak a he same ime as in
oli e g o es, in he all, bu no peak is displayed in sp ing.
C i ically, howe e , B. oleae abundance in eucalyp us o es
and in “mon ado” eally encompassed oli e g o e dynamics
wi h wo clea ly de ined peaks: one in sp ing and he o he
in au umn. This ac could be explained by he simila dis-
ibu ional cha ac e is ics o hese habi a s.
The dis ibu ional cha ac e is ic o a habi a can be
e y impo an o pes s o ecognize new po en ial sui -
able e i o ies o dispe se (P opoki and Owens 1983; Jac-
el e al. 2021). This is especially ele an in he bi o ous
diu nal species such as eph i id lies, he g oup o which
B. oleae belongs. Fo hese insec s, isual cues may play
an impo an ole in he loca ion o hos plan s and essen-
ial esou ces, such as ood, ma ing and o iposi ion si es
(P opoki and Owens 1983). Indeed, some eph i id lies ha e
been de ec ed isually eac ing o shapes and silhoue es o
plan popula ions (Moe icke e al. 1975). The e o e, size o
ege a ional pa ch, densi y and dispe sion pa e n o hos s
plan s wi hin a pa ch, mo phological di e ences be ween
hos s and non-hos s, and he o e all aspec o hos habi-
a s (Feeny 1976, Rausche 1981) in space and ime may all
ha e an e ec ha can be eally impo an wi hin an ag i-
cul u al con ex (Bach 1981; Smi h 1976). The e o e, when
dispe sing, B. oleae could po en ially selec eucalyp us o es
and, especially, “mon ados” as sui able habi a s o look o
esou ces du ing he sp ing, as hey show dispe sion pa -
e ns and densi ies e y simila o oli e g o es wi h soli a y
sca e ed ees. Indeed, his could be he eason behind he
ac ha B. oleae popula iondynamic in “mon ados” mimic
hose in oli e g o es. Howe e , eucalyp us o es appea ance
is no simila o oli e g o es and he easons o epea ing
some empo al dynamics could be ela ed o his habi a ac -
ing as a shel e o ex eme wea he condi ions specially
in summe when B. oleae is highly a ec ed by high em-
pe a u es in he adul s age (Gu ie ez e al. 2009; Wang
e al. 2009; Abd El-Salam e al. 2019). Howe e , empo al
dynamics in eucalyp us o es s show a decline in he oli e
ly popula ion du ing summe . This is no consis en wi h
he clima e shel e hypo hesis, and mo e esea ch should be
done o elucida e why B. oleae eels a ac ion o eucalyp us
o es s.
Despi e he ac ha mos o he land uses s udied dis-
played a signi ican popula ion dynamic o he pes , ou
esul s show ha none ac as a ese oi o boos Bac oce a
oleae popula ions. This is co obo a ed wi h he null e ec
ha any habi a had on ly abundance wi hin he oli e g o e.
C i ically, landscape simpli ica ion (la ge a eas jus co e ed
wi h oli e g o es in he landscape) and landscape di e si-
ica ion (di e si y o land uses in he landscape) displayed
he mos no able e ec s. On he one hand, simpli ied land-
scapes allow B. oleae o easily g ow as hey ha e plen y o
esou ces o build hei popula ion and sp ead easily because
hey ha e no ba ie s o impede i s dispe sion (Roo 1973;
O’Rou ke and Pe e sen 2017). Impo an ly, he p esence o
B. oleae indi iduals in o he land uses, as demons a ed in
his s udy, suppo s he idea ha hese landscape elemen s
could ac as a ba ie o he dispe sion o his pes , which
e lec s i s low abundance in di e si ied landscapes. Mo eo-
e , simpli ied landscapes lack na u al enemies ha can exe
a p ope op-down con ol o he pes popula ion, whe eas
di e si ied landscapes can p o ide wi h esou ces o hese
na u al enemies allowing he deli e y o na u al pes con ol
se ices (Chaplin-K ame e al. 2011; Rusch e al. 2016).
Thus, he implica ion o inc easing landscape di e si y
seems ob ious o his pes and align wi h o he s udies ha
de ec ed landscape di e si ica ion as a majo d i e o B.
oleae abundance educ ion (O ega and Pascual 2014).
Managemen implica ions
As a consequence o hese indings, we encou age a me s,
echnicians and poli icians o p omo e landscape di e si i-
ca ion in oli e g o es. Inc easing a eas wi h di e en land
uses will likely impend he dispe sion o B. oleae, ul ima ely
p e en ing sudden popula ion peaks, which would o he wise
esul in ou b eaks ha would necessi a e insec icide appli-
ca ions. A he indi idual le el, a me s could be e con ol
B. oleae popula ions by plan ing na i e ege a ion a ound
hei a ms. Howe e , he inc ease in land use di e si y will
depend on he coo dina ion be ween g oups o neighbo ing
a me s o edesign he a angemen o he di e en elemen s
o he landscape, aking ad an age o he syne gies p o ided
by he o og aphy and changing land uses in a eas whe e
he composi ion is no balanced. A good way o p omo e
coo dina ion be ween di e en ag icul u al s akeholde s is
he inancial compensa ion p og ams ha go e nmen s can
pu in place o help inc ease he di e si y o he landscape
(Ba á y e al. 2015). This ype o coo dina ion based on he
Jou nal o Pes Science
1 3
in o ma ion we ob ain om he landscape is an impo an
and necessa y s ep o ob aining colla e al bene i s ela ed
o he p omo ion o ecosys em se ices such as biodi e si y
conse a ion and human heal h.
Au ho s’ con ibu ion
DP, SM, JA, AAS and JPS concei ed he p ojec and
designed he me hods; DP, SM, JFA and JMC collec ed he
da a; DP and JPS analyzed he da a; DP w o e he s udy. All
au ho s made c i ical con ibu ions o he d a and ga e inal
app o al o publica ion.
Acknowledgemen s The au ho s a e g a e ul o he Associa ion o
Oli e Oil P oduce s o Bei a In e io (APABI, Po ugal) o all he
e o and ca e in iden i ying and de ec ing he landowne s we equi ed,
as well as o all he oli e g owe s who kindly allowed he aps o
emain on hei p ope ies du ing he ial pe iod. We hank M. Sc.
Danielle Rudley o p oo eading he documen . We would also like
o hank M. Sc. Mikola Rasko, M. Sc. Rúben Mina and M. Sc. Sand a
Simões o hei con ibu ion o he sampling p ocess.
Funding This esea ch was unded by P og ama Ope acional Regional
do Cen o, G an Numbe Cen o-01-0145-FEDER-000007 (ReNA-
TURE), by he Eu opean Union’s Ho izon 2020 Resea ch and Inno-
a ion P og amme, unde G an Ag eemen No. 773554 (EcoS ack),
and by Fundação pa a a Ciência e Tecnologia h ough he p ojec
PTDC/ASP-PLA/30003/2017 (OLIVESIM). JA was inanced by
FCT/MCTES, h ough na ional unds (PIDDAC), o he Cen e o
Func ional Ecology-Science o People and he Plane (CFE), wi h he
e e ence UIDB/04004/2020.
Decla a ions
Con lic o in e es All au ho s decla e ha hey ha e no con lic o in-
e es . The au ho s ha e no ele an inancial o non- inancial in e es s
o disclose. The au ho s ha e no compe ing in e es s o decla e ha a e
ele an o he con en o his a icle. All au ho s ce i y ha hey ha e
no a ilia ions wi h o in ol emen in any o ganiza ion o en i y wi h
any inancial in e es o non- inancial in e es in he subjec ma e o
ma e ials discussed in his manusc ip . The au ho s ha e no inancial
o p op ie a y in e es s in any ma e ial discussed in his a icle.
E hical app o al This a icle does no con ain any s udies wi h human
pa icipan s o animals pe o med by any o he au ho s.
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