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Pee Communi y Jou nal
Sec ion: Ecology
RESEARCH ARTICLE
Published
2021-11-22
Ci e as
Ainhoa Mag ach, F ancisco P.
Molina and Ignasi Ba omeus
(2021) Niche complemen a i y
among pollina o s inc eases
communi y-le el plan
ep oduc i e success, Pee
Communi y Jou nal, 1: e1.
Co espondence
ainhoa.mag ach@bc3 esea ch.o g
Pee - e iew
Pee e iewed and
ecommended by
PCI Ecology,
h ps://doi.o g/10.24072/pci.
ecology.100037
This a icle is licensed
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A ibu ion 4.0 License.
Niche complemen a i y among
pollina o s inc eases
communi y-le el plan
ep oduc i e success
Ainhoa Mag ach1,2, F ancisco P. Molina3, and
Ignasi Ba omeus ,3
Volume 1(2021), a icle e1
h ps://doi.o g/10.24072/pcjou nal.1
Abs ac
Ou unde s anding o how he s uc u e o species in e ac ions shapes na u al com-
muni ies has inc eased, pa icula ly ega ding plan -pollina o in e ac ions. Howe e ,
esea ch linking pollina o di e si y o ep oduc i e success has ocused on pai wise
plan -pollina o in e ac ions, la gely o e looking communi y-le el dynamics. He e, we
p esen one o he fi s empi ical s udies linking pollina o isi a ion o plan ep oduc-
ion om a communi y-wide pe spec i e. We use a well- eplica ed da ase encompass-
ing 16 plan -pollina o ne wo ks and da a on ep oduc i e success o 19 plan species
om Medi e anean sh ub ecosys ems. We find ha s a is ical models including simple
isi a ion me ics a e sufficien o explain he a iabili y obse ed. Howe e , a mecha-
nis ic unde s anding o how pollina o di e si y affec s ep oduc i e success equi es
addi ional in o ma ion on ne wo k s uc u e. Specifically, we find posi i e effec s o in-
c easing complemen a i y in he plan species isi ed by diffe en pollina o s on plan
ep oduc i e success. Hence, main aining communi ies wi h a di e si y o species bu
also o unc ions is pa amoun o p ese ing plan di e si y.
1Basque Cen e o Clima e Change-BC3, Edi . Sede 1, 1o, Pa que Cien ífico UPV-EHU, Ba io Sa iena s/n,
48940, Leioa, Spain, 2IKERBASQUE, Basque Founda ion o Science, Ma ía Díaz de Ha o 3, 48013, Bilbao,
Spain, 3Es ación Biológica de Doñana (EBD-CSIC), A da. Amé ico Vespucio 26, Isla de la Ca uja, 41092,
Se illa, Spain
This documen is he Accep ed Manusc ip e sion o a Published Wo k ha appea ed in inal o m in:
Ainhoa Mag ach, F ancisco P. Molina, Ignasi Ba omeus. 2019. Niche complemen a i y among
pollina o s inc eases communi y-le el plan ep oduc i e success. PEER COMMUNITY IN ECOLOGY.
35. DOI (10.1101/629931).
©Pee Communi y Jou nal
This manusc ip e sion is made a ailable unde he CC-BY-NC-ND 3.0 license h p://c ea i ecommons.o g/
licenses/by-nc-nd/3.0/
In oduc ion
Pollina o s p o ide key se ices o plan s by acili a ing pollen low (Ga ibaldi e al. 2013). Declining
ends o some pollina o species in some egions (Po s e al. 2010; Ba omeus e al. 2018) ha e led
esea che s o ocus on he unc ional impac s o hese changes in pollina o di e si y, especially o plan
ep oduc i e success (Biesmeije e al. 2006).
Many s udies ha e e alua ed ep oduc i e success on indi idual plan species (Alb ech e al. 2012;
Thomson 2018), and used ela i ely simple isi a ion me ics (e.g., he numbe o pollina o species isi ing
a plan o he numbe o isi s hey pe o m) o explain he di e ences obse ed (e.g., Bomma co e al.
2012). Con as ingly, communi y-le el analyses emain sca ce (Benne e al. 2018). Ye plan s and
pollina o s do no in e ac in isola ion bu a e embedded wi hin la ge ne wo ks o in e ac ions
encompassing o he plan and pollina o species (Memmo e al. 2004). We a e hus missing an impo an
pa o he pic u e, including di ec in e ac ions be ween he whole ensemble o plan s and pollina o s,
bu also indi ec ones be ween species wi hin one guild (e.g., plan s) h ough hei sha ed esou ces (Pauw
2013; Láza o e al. 2014; Ca alhei o e al. 2014; May ield, S ou e 2017; Johnson, B ons ein 2019).
Unde s anding how changes in pollina o di e si y and communi y s uc u e a ec ecosys em unc ioning
is hus a majo challenge ha equi es a en ion.
The ew s udies ha ha e analyzed he e ec s o pollina o di e si y on ep oduc i e success a he
communi y le el ha e mainly used expe imen al se ups. As an example, a s udy ha expe imen ally
ec ea ed a plan communi y wi h 9 plan species and di e ing le els o pollina o di e si y, ound a
posi i e e ec o pollina o species di e si y on seed se , bu also an impo an e ec o niche
complemen a i y be ween pollina o s, a measu e o communi y s uc u e (F ünd e al. 2013). These
indings show ha no only he di e si y o species p esen , bu also he di e si y o oles hey play and
hus he way in which a communi y is s uc u ed a e de e minan ac o s o ecosys em unc ions.
Indeed, heo e ical esea ch has long sugges ed ha he s uc u e o mul i ophic communi ies has an
e ec o ecosys em unc ioning ( e iewed in (Thompson e al. 2012)). This line o esea ch, oo ed in niche
heo y and e amped by ood-web s udies (Maca hu , Le ins 1967; May, A hu 1972; Tilman 1982; Godoy
e al. 2018), has g ea ly ad anced heo y, bu he ela ionship be ween s uc u e and unc ion has seldom
been es ed using empi ical da a (bu see (Poiso e al. 2013; Kaise -Bunbu y e al. 2017; Láza o e al.
2020)). Speci ically, a majo knowledge gap esides in unde s anding which aspec s o s uc u e de e mine
which aspec s o unc ion (Thompson e al. 2012). This is because al hough a ne wo k pe spec i e has
p omised o encapsula e complex ecological mechanisms occu ing a he communi y le el – such as
indi ec in e ac ions (Hol 1977; Ab ams e al. 1998) o niche o e lap (Woodwa d, Hild ew 2002)- less
a en ion has been gi en o he ways in which hese mechanisms ela e o obse ed ecosys em p ocesses
(Blü hgen 2010). We a e now a a poin whe e we unde s and some o he eme gen pa e ns
cha ac e izing mu ualis ic in e ac ion ne wo ks a he communi y le el, especially in he case o pollina ion
(Bascomp e, Jo dano 2007). Amongs hem is he p e alence o nes ed s uc u es, i.e., a angemen s
whe e specialis species in e ac wi h a subse o he species ha gene alis s in e ac wi h (Bascomp e e
al. 2003). Fu he , plan -pollina o in e ac ion ne wo ks seem o exhibi a ela i ely high ex en o
complemen a y specializa ion a he communi y scale, which may be di ec ly ela ed o key ecosys em
unc ions (Blü hgen, Klein 2011). Howe e , he mechanisms by which hese a ibu es a ec plan
ep oduc ion emain o be unde s ood (Win ee 2013). The ime is hus ipe o explo e he ela ionship
be ween communi y s uc u e and ecosys em unc ioning empi ically, wi h special emphasis on he
unde lying ecological mechanisms ha d i e hese ela ionships.
He e, we p esen an empi ical s udy linking pollina o isi a ion and plan ep oduc i e success a he
communi y le el. We use a well- eplica ed da ase encompassing plan -pollina o in e ac ion ne wo ks
collec ed a 16 si es coupled wi h da a on he ep oduc i e success o 19 plan species eco ded in
Medi e anean sh ub ecosys ems. Ou s udy ocuses on unde s anding whe he adding in o ma ion on
selec ed in e ac ion ne wo k s uc u e indices o p e iously used simple isi a ion me ics (e.g., he
numbe and di e si y o pollina o species isi ing a plan species) aids in be e explaining he di e ences
obse ed in communi y-wide ep oduc i e success. In doing so, we conduc ed ou analyses ocusing on
ep oduc i e success a wo di e en le els: (i) a he species le el by conside ing he associa ion be ween
he posi ion o a ocal species wi hin he la ge ne wo k and i s link o indi idual ep oduc i e success, and
2 Ainhoa Mag ach e al.
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
(ii) a he si e le el, by e alua ing how a ibu es ha desc ibe he whole si e migh a ec a e age alues
o ep oduc i e success o all species measu ed wi hin one pa icula si e. Speci ically, ou s udy ocuses
on how he in e play be ween he complemen a i y in plan species isi ed by di e en pollina o s, and
he edundancy in his unc ion ela e o ep oduc i e success. Plan ep oduc i e success equi es o he
deli e y o conspeci ic pollen and hus o a ce ain deg ee o niche complemen a i y (Blü hgen, Klein 2011).
Ye , g ea e alues o edundancy in species unc ions (e.g., ha p o ided by nes ed s uc u es), a e
hough o p omo e species di e si y (Bas olla e al. 2009) and s abili y (Thébaul , Fon aine 2010) wi hin
plan -pollina o ne wo ks. A p esen , we do no know how ei he o hese ne wo k cha ac e is ics a ec s
he unc ions pe o med by pollina o s.
Ou esul s sugges ha models including in o ma ion on simple isi a ion me ics alone a e able o
explain di e ences in ep oduc i e success. Howe e , a mechanis ic unde s anding equi es addi ional
in o ma ion on ne wo k s uc u e, no ably in o ma ion on he complemen a i y be ween he niches
occupied by di e en pollina o species. Speci ically, we ind a posi i e e ec o inc easing niche
complemen a i y be ween pollina o s on plan ep oduc i e success.
Ma e ial and Me hod
Plan pollina o in e ac ions
Ou s udy was conduc ed in SW Spain wi hin he a ea o in luence o Doñana Na ional Pa k (Fig. S1).
Si es we e loca ed wi hin simila ele a ions ( anging om 50 o 150 m a.s.l.), and simila habi a and soil
ypes, educing po en ial con ounding ac o s. Simila i y in plan composi ion be ween si es was 0.41 (plan
mean Sø ensen be a-di e si y). We su eyed 16 Medi e anean woodland pa ches wi h an a e age
dis ance o 7 km be ween hem (min= 3 km, max= 46.5 km). Each si e was su eyed e e y wo weeks o a
o al o 7 imes du ing he lowe ing season o 2015 ( om Feb ua y o May) ollowing a 100-m x 2 m
ansec o 30 mins. Along each ansec , we iden i ied all plan species and eco ded all he lo al isi o s
ha landed on hei lowe s. Only lo al isi o s ( om now on e e ed o as pollina o s) ha could no be
iden i ied in he ield we e cap u ed, s o ed and iden i ied in he labo a o y by FPM and ano he expe
en omologis (see acknowledgemen s). All su eys we e done unde simila wea he condi ions, a oiding
windy o ainy days, du ing mo nings and a e noons wi h he sampling o de being es ablished andomly.
Wi hin each ansec e e y 10 m we su eyed a 2x2 m quad an whe e he numbe o lowe s pe species
we e coun ed, i.e., 10 quad a s pe ansec which makes 40m2 o a ea su eyed o e all.
Plan ep oduc i e success
Wi hin each si e, we ma ked 3-12 indi iduals (mean ± SD: 6.49 ± 2.37) belonging o 1-6 plan species
(mean ± SD:4.06 ± 1.69, Table S2). Fo each indi idual, a he end o he season, we eco ded ui se
(i.e. he p opo ion o lowe s ha se ui ), he a e age numbe o seeds pe ui and he a e age ui
and seed weigh pe ui (1-36 ui s subsampled, mean ± SD: 11.17 ± 6.85, Table S3). These las wo
a iables show a s ong co ela ion (Pea son co ela ion= 0.89), and hus we only p esen esul s on ui
weigh . Ou su ey included a o al o 19 di e en o ally o pa ially sel -incompa ible plan species ha
depend on pollina o s o maximize hei ep oduc ion (Table S4) ac oss ou 16 si es. All plan species we e
common and widesp ead sh ubs. Indi iduals we e selec ed depending on he p esence o lowe s du ing
he sampling e en s. We also calcula ed he a e age ep oduc i e success a he si e le el by a e aging
alues o ep oduc i e success ob ained o each species.
Da a analyses
To e alua e he sampling comple eness, we es ima ed he asymp o ic numbe o species o plan s,
pollina o s and in e ac ions p esen (Chao e al. 2009), a non-pa ame ic es ima o o species ichness o
abundance da a. This es ima o includes non-de ec ed species and allowed us o calcula e he p opo ion
de ec ed wi h ou o iginal da a. We used Chao 1 asymp o ic species ichness es ima o s (Chao e al. 2009)
and es ima ed he ichness o pollina o s, plan s and plan –pollina o links accumula ed as sampling e o
inc eased up o 100% sampling co e age using package iNEXT (Hsieh e al. 2016) wi hin he R en i onmen
(R De elopmen Co e Team 2011). We hen ex ac ed he alues co e ed by ou sampling.
To e alua e di e ences in ne wo k s uc u e be ween communi ies, we cons uc ed plan -pollina o
in e ac ion ne wo ks by pooling he da a o he 7 ounds o sampling. We hus ob ained one in e ac ion
Ainhoa Mag ach e al. 3
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
ne wo k pe si e, ep esen ing he numbe o indi iduals o di e en pollina o species eco ded isi ing
each di e en plan species. Fo each ne wo k, we ex ac ed a se ies o ele an ne wo k me ics a he
species and si e le els.
Addi ionally, we checked o spa ial au oco ela ion in ou da a using Man el co elog ams.
Au oco ela ion alues we e non-signi ican o all a iables, excep o pollina o ichness whe e we ha e
a small bu signi ican e ec a small spa ial scales (Fig. S2). Hence, we ea each si e as independen in
ou analysis.
Species-le el ne wo k analysis
A he species le el, we ocused on a ibu es de ining he posi ion o a ocal plan species wi hin he
la ge communi y. As such, we conside ed wo me ics p o iding complemen a y non- edundan
in o ma ion: (i) a e age niche o e lap in e ms o pollina o s be ween a ocal plan species and each o he
o he plan species in he communi y, and (ii) he con ibu ion o nes edness o each indi idual plan
species. Niche o e lap es ima es he po en ial indi ec in e ac ions be ween plan species h ough sha ed
esou ces (in his case pollina o s) and he po en ial o inc eased he e ospeci ic pollen deposi ion (A ceo-
Gómez e al. 2019). We calcula ed i as he a e age o e lap in pollina o species isi ing a ocal plan and
each o he o he plan s in he communi y using he Mo isi a o e lap index, a measu e o simila i y
be ween wo se s o da a (Zhang 2016). A plan species’ con ibu ion o nes edness is calcula ed by
compa ing he nes edness obse ed in a gi en communi y o ha gene a ed by andomizing he
in e ac ions in which a ocal species is in ol ed. Species ha show impo an con ibu ions o o e all
nes edness will ha e alues >0, while species ha do no con ibu e o o e all nes edness wil show alues
<0 (Saa ed a e al. 2011).
Si e-le el ne wo k analysis
A he si e le el, we ollowed he same logic as he one p esen ed a he species le el. We also
calcula ed wo ne wo k me ics p o iding complemen a y non- edundan in o ma ion. In his case, we
ocused on nes edness, a measu e o he edundancy in he plan s isi ed by di e en pollina o s, and
pollina o niche complemen a i y, a measu e o he complemen a i y in plan species isi ed by di e en
pollina o species.
Nes edness is he p ope y by which specialis s in e ac wi h a subse o he species ha gene alis s
in e ac wi h (Bascomp e e al. 2003). Al hough he e is an ongoing deba e in he li e a u e (e.g., (James e
al. 2012)), some heo e ical s udies ha e ound ha nes ed ne wo ks a e mo e s able and esilien o
pe u ba ions because nes edness p omo es a g ea e di e si y by minimizing compe i ion among species
in a communi y (Bas olla e al. 2009). Howe e , many ne wo k a ibu es a y wi h ne wo k size and
complexi y (Blü hgen e al. 2006). In he case o nes edness, we know i can be a ec ed by ne wo k size
and connec ance (Song e al. 2017). An app oach ha is o en used o co ec o his a e null models,
compa ing null-model co ec ed nes edness alues ac oss di e en ne wo ks. Howe e , his app oach
p esen s he same issues, as z-sco es also change wi h ne wo k size and connec ance (Song e al. 2017).
We hus used a no malized alue o he widely used nes edness me ic NODF based on bina y ma ices
(Almeida-Ne o, Ul ich 2011), 𝑁𝑂𝐷𝐹𝑐 (Song e al. 2017). This no malized alue is calcula ed as 𝑁𝑂𝐷𝐹𝑐=
𝑁𝑂𝐷𝐹𝑛/(𝐶 ∗ 𝑙𝑜𝑔(𝑆)), whe e C is connec ance and S is ne wo k size, calcula ed as 𝑆 = √(𝑛𝑐𝑜𝑙(𝑤𝑒𝑏) ∗
𝑛𝑟𝑜𝑤(𝑤𝑒𝑏)). 𝑁𝑂𝐷𝐹𝑛 is calcula ed as 𝑁𝑂𝐷𝐹/𝑚𝑎𝑥(𝑁𝑂𝐷𝐹), which is independen o ne wo k size and hus
compa able ac oss di e en ne wo ks (Song e al. 2017). To calcula e max(NODF) we used a co ec ed
e sion o he algo i hm (Simmons e al. 2019) whene e possible. Resul s did no change quali a i ely
when using he unco ec ed e sion o he algo i hm o all si es as bo h a e highly co ela ed (Spea man
co ela ion = 0.94).
To calcula e niche complemen a i y, we used a communi y-le el measu e de ined as he o al b anch
leng h o a dend og am based on quali a i e di e ences in isi o assemblages be ween plan s (De o o e
al. 2012; pe chey200?). All ne wo k me ics we e calcula ed using package bipa i e (Do mann e al. 2009).
S a is ical analyses
To e alua e whe he adding in o ma ion on ne wo k s uc u e imp o es ou abili y o explain
di e ences in ep oduc i e success - bo h a he species and he si e le el - we used gene alized linea
(GLMs) and gene alized linea mixed models (GLMMs) espec i ely. In bo h cases we i h ee ypes o
4 Ainhoa Mag ach e al.
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
models: (i) model 0, a null model wi h no explana o y a iables,(ii) model 1, ha only included simple
isi a ion me ics and (iii) model 2 ha addi ionally included in o ma ion on ne wo k s uc u e. These
models a e mean o be addi i e, so ha he ne wo k me ics included a e in ended o complemen a he
han subs i u e he simple me ics adi ionally used.
A he species le el, esponse a iables included ui se analyzed using a binomial dis ibu ion and he
a e age numbe o seeds pe ui , and he a e age ui weigh i ed using no mal dis ibu ions. The
numbe o seeds pe ui was cen e ed and scaled (i.e., we sub ac ed column means and di ided by
s anda d de ia ion) o allow meaning ul compa isons ac oss species wi h con as ing li e his o ies. As
explana o y a iables, model 1 included he numbe o pollina o species obse ed, and he isi a ion a e
ecei ed by each plan species. Visi a ion a e was calcula ed as he o al numbe o isi s ecei ed by a
plan species di ided by he a e age numbe o lowe s o ha species ound in he 10 2x2 m quad a s pe
ansec . In u n, model 2 added he wo ne wo k a ibu es calcula ed a he species le el: a e age plan
niche o e lap and con ibu ion o nes edness. Fo bo h models, we included plan species iden i y nes ed
wi hin si e and si e as andom e ec s o accoun o mul iple indi iduals o he same plan species
measu ed a each si e.
A he si e le el, esponse a iables we e he a e age ep oduc i e success o all plan s su eyed wi hin
a si e (i.e., a e age ui se analyzed using a binomial dis ibu ion, a e age numbe o seeds pe ui and
a e age ui weigh using a no mal dis ibu ion). We hus had a single alue pe si e and no andom e ec s
a e needed. He e, model 1 included o al pollina o ichness and o al pollina o abundance (i.e. numbe
o isi s ecei ed by all plan s wi hin he communi y) as explana o y a iables. Model 2, in u n, added
in o ma ion on ne wo k s uc u e by including nes edness and pollina o niche complemen a i y.
A e age alues o ep oduc i e success a he si e le el can be d i en by a single plan species. Ye ,
wha will de e mine he pe sis ence o a di e se plan communi y, is he p esence o some so o “equi y”
o e enness in ep oduc i e success ac oss he whole communi y. We he e o e calcula ed he p opo ion
o species wi h no malized (be ween 0 and 1) a e age ui se alues ha we e abo e he 50^ h pe cen ile
as a measu e o equi y. As any selec ed h eshold is a bi a y, we epea ed his using he 25^ h and 75^ h
pe cen ile h esholds (By nes e al. 2014). We hen used he same amewo k as ha used o species and
si e-le el analyses and i he same models 0, 1 and 2 using equi y in ep oduc i e success as esponse
a iable and i ing a binomial dis ibu ion.
In all cases, we used a iance in la ion ac o s o check o collinea i y be ween explana o y a iables.
Addi ionally, we an esidual diagnos ics o check i model assump ions we e me and used he Akaike
In o ma ion C i e ion (AIC) o compa e model pe o mance and complexi y. Whene e he di e ence
be ween he AIC o he models was < 2 (𝛥𝐴𝐼𝐶 < 2), we conside ed all models equally good (Bu nham e
al. 2010). In he case o mixed models, o compa ison, models we e i ed by maximum likelihood and
hen he bes model was e i ed using es ic ed maximum likelihood. All p edic o a iables we e
s anda dized p io o analysis. Fo e e y model we also calcula e he R2 alue using he app oxima ion
sugges ed o GLMMs when necessa y (Nakagawa e al. 2017).
Finally, we es ed whe he he impo ance o ne wo k s uc u e in explaining di e ences in equi y in
ep oduc i e success inc eases wi h he numbe o plan species being conside ed. We expec ha when
only one plan species is conside ed he impo ance o ne wo k s uc u e will be negligible, while we expec
i o inc ease as mo e plan species a e conside ed (up o a maximum numbe o 6 species which is he
maximum we ha e measu ed in ou s udy a a pa icula si e).
To es his, we an a simple simula ion in which he numbe o species conside ed inc eased a each
s ep and o each s ep we e-calcula ed equi y in ep oduc i e success. Ins ead o d awing plan species
andomly o each s ep, we es ed all possible combina ions o each plan numbe le el and ne wo k, as
he numbe o combina ions is small (e.g. o n = 3 plan s selec ed ou o 6 he e a e only 20 possible
combina ions). Then, we es ed i he ela ionship be ween equi y in ep oduc i e success and niche
complemen a i y (gi en i s impo ance in de e mining di e ences in ep oduc i e success, see Resul s
sec ion) changes as a unc ion o he numbe o plan s conside ed wi hin ou simula ed communi ies. To
his end, o each le el o species numbe conside ed, we andomly selec ed one o he gene a ed equi y
alues ac oss each o he 16 communi ies and eg essed hese 16 alues agains ou ne wo k le el
p edic o and ex ac ed he model slope es ima es. We epea ed his p ocess 1,000 imes and a e aged all
slope es ima es. We expec ha he mo e plan s conside ed, he la ge he esul ing a e age es ima es will
Ainhoa Mag ach e al. 5
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
be. No e ha we only in e p e he mean e ec s, as he a iance among di e en plan numbe o species
conside ed depends on he ini ial numbe o possible combina ions.
Resul s
Wi hin ou sampling we eco ded 655 plan -pollina o in e ac ions in ol ing 277 pollina o species and
57 plan species (Table S1). Wi hin he pollina o communi y he dis ibu ion o indi iduals in di e en
o de s was: 92.18% Hymenop e a, 5.69% Dip e a, 1.29% Coleop e a and 0.63% Lepidop e a.
Ou sampling comple eness analyses e ealed ha ou su ey was able o cap u e 17-54% o pollina o
species (a e age = 35%), 43-100% o plan species (a e age = 80%) and 9-32% o plan -pollina o links
(a e age = 20%), in line wi h ha ob ained wi h o he s udies (e.g., (Chaco e al. 2011), Fig. S3). Ou alues
o sampling comple eness we e sligh ly smalle in he case o pollina o s, p obably as a consequence o he
g ea di e si y ound in he Medi e anean egion and wi hin ou s udy a ea in pa icula , a ho spo o
insec di e si y (Eu opean Commission. Di ec o a e Gene al o he En i onmen ., IUCN (In e na ional
Union o Conse a ion o Na u e). 2014).
Species-le el analyses
A he species le el, in he case o ui se , ou esul s showed ha model 2 had he bes i o ou da a
(lowes AIC alue), and ixed e ec s explained 9% o he a iabili y obse ed (condi ional R^2=17%). We
ound a posi i e ela ionship be ween ui se , pollina o species ichness, and a ne wo k s uc u e me ic,
he con ibu ion o nes edness o a ocal plan wi hin he o e all ne wo k (Table 1, Fig. 1, Fig. S4).
Fo he a e age numbe o seeds pe ui a he species le el as well as o ui weigh , ou esul s
showed ha none o he models i ed we e be e han he null model explaining di e ences ac oss plan
species.
Table 1. Resul s o GLMM showing he associa ion be ween simple isi a ion and ne wo k s uc u e
me ics and species-le el ui . Bold le e s indica e a iables wi h la ge e ec s (see Figu e S4 o
es ima e con idence in e als).
F ui se
Es ima e
S d.E o
z. alue
(In e cep )
1.79
0.21
8.38
Pollina o ichness
0.51
0.25
2.04
Rela i e numbe o isi s
-0.16
0.25
-0.64
Plan niche o e lap
0.20
0.23
0.85
Con ibu ion o nes edness
0.47
0.26
1.81
6 Ainhoa Mag ach e al.
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Figu e 1. Pa ial esidual plo s showing he e ec o A) pollina o species ichness and B) he
con ibu ion o nes edness o each plan species on ui se . Do s ep esen each o he indi iduals
sampled o each species wi hin each si e.
Si e-le el analyses
A he si e le el, in he case o ui se and he numbe o seeds pe ui , we ound ha bo h model 1
and 2 we e equally good when penalizing o model complexi y (i.e.,𝛥𝐴𝐼𝐶 < 2; (Bu nham, Ande son
2004)). This sugges s model 2 was a good model despi e i s added complexi y, and ac ually shows a
subs an ially be e p edic i e abili y han model 1 (R^2 = 0.46 o model 2 e sus 0.27 o model 1 in he
case o ui se and R^2 = 0.49 o model 2 e sus 0.35 o model 1 in he case o he numbe o seeds pe
ui ) and he e o e we will commen esul s o his model only. Speci ically, we ound ha bo h ui se
and he numbe o seeds pe ui we e posi i ely ela ed o niche complemen a i y be ween pollina o s
(Tables 2, Fig. 2, Fig. S5). Addi ionally, we ound a nega i e associa ion be ween si e-le el pollina o
ichness and a e age ui se (Table 2A, Fig. 2, Fig. S5).
In he case o ui weigh , we ound ha bo h he null model and model 1 we e equally good
(i.e.,𝛥𝐴𝐼𝐶 < 2; (Bu nham, Ande son 2004)). Model 1, i.e., ha only including simple isi a ion me ics,
showed an R^2 o 0.23. In his case, we ound a posi i e link wi h si e-le el pollina o ichness (Table S5A,
Figs. S5-S6). This associa ion was main ained e en a e emo ing a si e ha has a pa icula ly la ge
pollina o ichness alue (Table S5B, Fig. S7, Fig. S5).
Ainhoa Mag ach e al. 7
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Table 2. Resul s o GLM showing associa ions be ween simple isi a ion and ne wo k s uc u e
me ics and A) si e-le el a e age ui se and B) si e-le el a e age numbe o seeds pe ui based on
bes model selec ed. Bold le e s indica e a iables wi h la ge e ec s.
A) F ui se
Es ima e
S d. E o
z alue
(In e cep )
1.20
0.15
7.79
Pollina o ichness
-0.77
0.26
-2.91
Rela i e numbe o isi s
-0.12
0.19
-0.66
Nes edness
0.02
0.16
0.12
Pollina o niche complemen a i y
0.40
0.26
1.58
B) Seeds pe ui
Es ima e
S d. E o
alue
(In e cep )
45.37
8.84
5.13
Pollina o ichness
1.56
15.80
0.10
Rela i e numbe o isi s
4.37
10.78
0.41
Nes edness
3.94
9.80
0.40
Pollina o niche complemen a i y
26.44
15.49
1.71
Figu e 2. Pa ial esidual plo s showing he e ec o he single p edic o which bes explains he
a iabili y in si e-le el ep oduc i e success. A) Shows he e ec o pollina o ichness, and B) o niche
complemen a i y among pollina o species on si e-le el a e age ui se . C) Shows he e ec o niche
complemen a i y among pollina o species on he a e age numbe o seeds pe ui a he si e le el.
Do s ep esen a e age alues o ui se a he le el o he communi y o all plan species conside ed
(N=16 si es).
Equi y in ui se
When e alua ing he ela ionship be ween communi y composi ion and ne wo k s uc u e on equi y
in ep oduc i e success ac oss he di e en species wi hin a communi y, we ound ha using he 50^ h
pe cen ile all models we e equally good (i.e.,𝛥𝐴𝐼𝐶 < 2; (Bu nham, Ande son 2004)), bu none o he
8 Ainhoa Mag ach e al.
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a iables conside ed showed any s ong associa ions (Table S6). In he case o he o he wo h esholds
conside ed (25^ h and 75^ h pe cen iles) model 0, he null model, was he bes model.
Wi hin ou simula ion e alua ing he ela ionship be ween niche complemen a i y and equi y in
ep oduc i e success a inc easing numbe o plan species conside ed, we ound ha he link o
complemen a i y became mo e impo an as mo e species we e conside ed (Fig. 3). This impo ance
seemed o each a pla eau a 6 species. Howe e , his should be u he e alua ed, as his was he
maximum numbe o species simul aneously obse ed in a communi y o ou s udy, which p ecludes us
om simula ing u he numbe s o species.
Figu e 3. Resul s o simula ion e alua ing he impo ance o niche complemen a i y in de e mining
di e ences in equi y in ep oduc i e ac oss communi ies ha bo ing om one o six species. Poin s
ep esen a e age alues ac oss 1,000 simula ed combina ions.
Discussion
The exis ence o ela ionships be ween in e ac ion ne wo k s uc u e and ecosys em unc ion ha e
been long hypo hesized, ye , he speci ic mechanisms unde lying his ela ionship emain elusi e
(Thompson e al. 2012). Ou esul s sugges ha di e en aspec s o ne wo k s uc u e a ec di e en
dimensions o ecosys em unc ioning. Speci ically, we ind ha he con ibu ion o nes edness o a plan
species wi hin a communi y has a posi i e associa ion wi h i s ui se . F om a plan ’s pe spec i e, his
indica es ha being connec ed o o he plan species ia sha ed pollina o s has a posi i e ou come (e.g. by
ensu ing a s able pollina o supply h ough ime) a he han a nega i e one (e.g. ia he e ospeci ic pollen
anspo ). A he si e le el, we ind ha g ea e alues o niche complemen a i y be ween pollina o s
esul in la ge a e age alues o ep oduc i e success.
Mos o ou analyses e eal ha model 1 and 2 we e equally good, which sugges s ha he added
complexi y o measu ing he ull ne wo k o in e ac ions may no pay o o apid assessmen s. Hence,
simple isi a ion me ics, such as pollina o ichness, migh be enough o desc ibe gene al pa e ns
(Ga ibaldi e al. 2013; Ga ibaldi e al. 2014). Ye , adding ne wo k le el in o ma ion may in o m us o he
po en ial ecological mechanisms unde lying he p ocesses d i ing hese obse ed pa e ns. Fu he ,
al hough we sampled each si e se en imes in a andomized o de in an a emp o be e ep esen
in e ac ions h ough ime, ou su eys we e able o cap u e 20% o in e ac ions gi en he g ea di e si y
o ou s udy sys em. This could be explaining pa o he low e ec sizes we ind a he species le el, whe e
a s onge con ibu ion o pollina o isi s is expec ed gi en hei obliga e dependence. In addi ion, plan
ep oduc i e success is a ec ed by o he a iables which we do no a emp o measu e in his s udy and
ha could explain a la ge po ion o he a iabili y obse ed.
Consis en wi h p e ious expe imen al (Fon aine e al. 2005; F ünd e al. 2013), heo e ical (Pauw
2013), and empi ical s udies (Poiso e al. 2013; Valdo inos e al. 2016), we ind ha niche complemen a i y
Ainhoa Mag ach e al. 9
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Figu e S6. Pa ial esidual plo s showing he e ec o pollina o ichness on si e-le el a e age ui
weigh . Do s ep esen alues o each si e (N=16 si es).
Figu e S7. Pa ial esidual plo s showing he e ec o pollina o ichness on si e-le el a e age ui
weigh . He e, a si e wi h a pa icula ly la ge pollina o ichness alue is emo ed o es whe he i migh
be d i ing he signi ican ela ionship. Do s ep esen alues o each si e (N=15 si es).
16 Ainhoa Mag ach e al.
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Table S1A. Lis o all plan species p esen a each o he si es and included in ne wo k analyses.
Si e
Plan species
Aznalcaza
Asphodelus is ulosus
Aznalcaza
Cis us c ispus
Aznalcaza
Cis us ladani e
Aznalcaza
Cis us monspeliensis
Aznalcaza
Cis us sal i olius
Aznalcaza
Echium plan agineum
Aznalcaza
La andula peduncula a
Aznalcaza
La andula s oechas
Aznalcaza
La a e a c e ica
Aznalcaza
Rosma inus o icinalis
Aznalcaza
Teuc ium u icans
Bona es
And yala in eg i olia
Bona es
Cis us c ispus
Bona es
Cis us ladani e
Bona es
Cis us sal i olius
Bona es
Halimium commu a um
Bona es
La andula peduncula a
Bona es
La andula s oechas
Bona es
Thapsia illosa
Bona es
Thymus mas ichina
Con en odelaLuz
Cis us c ispus
Con en odelaLuz
Cis us ladani e
Con en odelaLuz
Cis us sal i olius
Con en odelaLuz
Halimium halimi olium
Con en odelaLuz
La andula s oechas
Con en odelaLuz
Re ama sp.
Con en odelaLuz
Rosma inus o icinalis
Con en odelaLuz
Spa ium junceum
Con en odelaLuz
Teuc ium u icans
Co i odeSan aTe esa
As agalus lusi anicus
Co i odeSan aTe esa
Cis us c ispus
Co i odeSan aTe esa
Cis us sal i olius
Co i odeSan aTe esa
La andula peduncula a
Co i odeSan aTe esa
La andula s oechas
Co i odeSan aTe esa
Rosma inus o icinalis
Co i odeSan aTe esa
Thapsia illosa
Elpina
Cis us albidus
Elpina
Cis us sal i olius
Elpina
Con ol ulus a ensis
Elpina
Halimium commu a um
Elpina
La andula s oechas
Elpina
Rosma inus o icinalis
Elpozo
Cis us ladani e
Elpozo
Cis us sal i olius
Elpozo
E ica scopa ia
Elpozo
E ica umbella a
Elpozo
Rosma inus o icinalis
Espa agal
A me ia elu ina
Espa agal
Chamaemelum usca um
Espa agal
Cis us libano is
Espa agal
Cis us sal i olius
Ainhoa Mag ach e al. 17
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Si e
Plan species
Espa agal
Halimium commu a um
Espa agal
La andula peduncula a
Espa agal
La andula s oechas
Espa agal
Scabiosa a opu pu ea
LaCunya
And yala in eg i olia
LaCunya
Ce in he gymnand a
LaCunya
Cis us sal i olius
LaCunya
Echium plan agineum
LaCunya
E ica cilia is
LaCunya
Halimium commu a um
LaCunya
La andula peduncula a
LaCunya
Leon odon longi os is
LaCunya
Rosma inus o icinalis
LaCunya
Tube a ia gu a a
LaCunya
Ulex aus alis
LaRocina
Anchusa azu ea
LaRocina
And yala in eg i olia
LaRocina
Cis us sal i olius
LaRocina
Diplo axis i ga a
LaRocina
Halimium commu a um
LaRocina
Halimium halimi olium
LaRocina
La andula peduncula a
LaRocina
La andula s oechas
LaRocina
Lina ia iscosa
LaRocina
Rosma inus o icinalis
LaRocina
Spa ium junceum
Lasmulas
Cis us c ispus
Lasmulas
Cis us ladani e
Lasmulas
Cis us monspeliensis
Lasmulas
Cis us sal i olius
Lasmulas
Echium plan agineum
Lasmulas
La andula s oechas
Lasmulas
Ranunculus sp.
Lasmulas
Rosma inus o icinalis
Lasmulas
Thapsia illosa
Niebla
And yala in eg i olia
Niebla
A c o heca calendula
Niebla
Asphodelus is ulosus
Niebla
As agalus lusi anicus
Niebla
Calendula a ensis
Niebla
Ca duus sp.
Niebla
Cis us c ispus
Niebla
Cis us ladani e
Niebla
Cis us monspeliensis
Niebla
Con ol ulus a ensis
Niebla
La andula peduncula a
Niebla
La andula s oechas
Niebla
Leon odon sp.
Niebla
Lina ia iscosa
Niebla
Linum bienne
Niebla
Lupinus angus i olius
Niebla
Phlomis pu pu ea
Niebla
Ta axacum ulga e
18 Ainhoa Mag ach e al.
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Si e
Plan species
Niebla
Thapsia illosa
Pina esdeHinojos
And yala in eg i olia
Pina esdeHinojos
Cis us c ispus
Pina esdeHinojos
Cis us libano is
Pina esdeHinojos
Cis us sal i olius
Pina esdeHinojos
Diplo axis i ga a
Pina esdeHinojos
Rosma inus o icinalis
Pina esdeHinojos
Spa ium junceum
Pina esdeHinojos
Ulex aus alis
Pinodelcue o
Asphodelus is ulosus
Pinodelcue o
Chamaemelum usca um
Pinodelcue o
Cis us c ispus
Pinodelcue o
Cis us ladani e
Pinodelcue o
Cis us sal i olius
Pinodelcue o
Halimium commu a um
Pinodelcue o
La andula peduncula a
Pinodelcue o
La andula s oechas
Pinodelcue o
Ranunculus sp.
Pinodelcue o
Rosma inus o icinalis
Pinodelcue o
Thapsia illosa
Pinodelcue o
Ulex aus alis
U banizaciones
Calendula a ensis
U banizaciones
Cis us c ispus
U banizaciones
Cis us sal i olius
U banizaciones
Halimium commu a um
U banizaciones
La andula peduncula a
U banizaciones
La andula s oechas
U banizaciones
Rosma inus o icinalis
U banizaciones
Tube a ia gu a a
U banizaciones
Ulex aus alis
Villaman iquees e
Cis us c ispus
Villaman iquees e
Cis us ladani e
Villaman iquees e
Cis us sal i olius
Villaman iquees e
Genis a hi su a
Villaman iquees e
Rosma inus o icinalis
Villaman iquees e
Spa ium junceum
Villaman iquesu
And yala in eg i olia
Villaman iquesu
A me ia elu ina
Villaman iquesu
Cis us c ispus
Villaman iquesu
Cis us sal i olius
Villaman iquesu
Con ol ulus a ensis
Villaman iquesu
Genis a hi su a
Villaman iquesu
Halimium halimi olium
Villaman iquesu
La andula s oechas
Villaman iquesu
Rosma inus o icinalis
Table S1B. Lis o all pollina o species p esen a each o he si es and included in ne wo k analyses.
Si e
Pollina o species
Aznalcaza
And ena la ipes
Aznalcaza
And ena nig oaenaea
Ainhoa Mag ach e al. 19
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Si e
Pollina o species
Aznalcaza
And ena ni idiuscula
Aznalcaza
And ena sp.
Aznalcaza
And ena enuis ia a
Aznalcaza
An hopho a dispa
Aznalcaza
An hopho a sp.
Aznalcaza
Apis melli e a
Aznalcaza
Bombus e es is
Aznalcaza
Callipho a sp.
Aznalcaza
Ce ce is sabulosa
Aznalcaza
Dasypoda a gen a a
Aznalcaza
Dasypoda cingula a
Aznalcaza
Dasypoda c assico nis
Aznalcaza
Empis mo pho1
Aznalcaza
E is alis a bus o um
Aznalcaza
Euce a al e nans
Aznalcaza
Euce a codinai
Aznalcaza
Euce a colla is
Aznalcaza
Euce a elonga ula
Aznalcaza
Euce a hispaliensis
Aznalcaza
Euce a sp.
Aznalcaza
Fla ipanu gus enus us
Aznalcaza
Helio au us u icollis
Aznalcaza
Hopli is adunca
Aznalcaza
Lasioglossum mo pho1
Aznalcaza
Mac oglossum s ella a um
Aznalcaza
Me odon sp.
Aznalcaza
Osmia leaiana
Aznalcaza
Panu gus calca a us
Aznalcaza
Pseudoan hidium li u a um
Aznalcaza
Rhyncomyia cup ea
Aznalcaza
Sy phidae sp.
Aznalcaza
Tabanus mo pho1
Aznalcaza
Tabanus mo pho2
Aznalcaza
Volucella elegans
Aznalcaza
Xylocopa can ab i a
Bona es
Ammophila heydeni
Bona es
Ancis oce us biphale a us
Bona es
And ena hispania
Bona es
And ena nig oaenaea
Bona es
And ena o a ula
Bona es
And ena hyssono a
Bona es
And ena ulpecula
Bona es
An haxia mo pho1
Bona es
An hidium sep emspinosum
Bona es
Apis melli e a
Bona es
Bombus e es is
Bona es
Bombylius sp.
Bona es
Ce a ina cucu bi ina
Bona es
Colle es acu us
Bona es
Colle es liga us
Bona es
Dasypoda hi ipes
Bona es
Dasypogon mo pho1
Bona es
Empis mo pho1
20 Ainhoa Mag ach e al.
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Si e
Pollina o species
Bona es
Empis sp.
Bona es
E is alis sp.
Bona es
Euce a colla is
Bona es
Euce a elonga ula
Bona es
Euce a sp.
Bona es
G aphosoma linea um
Bona es
Halic us scabiosae
Bona es
Hopli is papa e is
Bona es
Lasioglossum sp.
Bona es
Megachile sp.
Bona es
Musca sp.
Bona es
Pla ynochae us se osus
Bona es
T ypoxylon mo pho1
Con en odelaLuz
Ammophila heydeni
Con en odelaLuz
An hopho a e usa
Con en odelaLuz
Apis melli e a
Con en odelaLuz
Bombus e es is
Con en odelaLuz
Chasma op e us illosulus
Con en odelaLuz
Euce a al e nans
Con en odelaLuz
Exosoma lusi anicum
Con en odelaLuz
Ichneumonidae mo pho1
Con en odelaLuz
Oxy hy ea unes a
Con en odelaLuz
Pla ynochae us se osus
Con en odelaLuz
Sy phidae sp.
Con en odelaLuz
T opino a squalida
Con en odelaLuz
Vespula ge manica
Con en odelaLuz
Xylocopa can ab i a
Co i odeSan aTe esa
And ena hyssono a
Co i odeSan aTe esa
An hopho a aes i alis
Co i odeSan aTe esa
An hopho a hispanica
Co i odeSan aTe esa
Apis melli e a
Co i odeSan aTe esa
Bombus e es is
Co i odeSan aTe esa
Dasypoda cingula a
Co i odeSan aTe esa
Dasypoda c assico nis
Co i odeSan aTe esa
Euce a ch ysopyga
Co i odeSan aTe esa
Euce a codinai
Co i odeSan aTe esa
Euce a sp.
Co i odeSan aTe esa
He iades c enula us
Co i odeSan aTe esa
Lasioglossum albocinc um
Co i odeSan aTe esa
Lasioglossum malachu um
Co i odeSan aTe esa
Lasioglossum sp.
Co i odeSan aTe esa
Les ica clypea a
Co i odeSan aTe esa
Mac oglossum s ella a um
Co i odeSan aTe esa
Me odon sp.
Co i odeSan aTe esa
Musca mo pho1
Co i odeSan aTe esa
Nemo elus mo pho1
Co i odeSan aTe esa
Nomada ag es is
Co i odeSan aTe esa
Nomada sp.
Co i odeSan aTe esa
Pla ynochae us se osus
Co i odeSan aTe esa
T ypoxylon mo pho1
Co i odeSan aTe esa
Xylocopa can ab i a
Elpina
And ena e ugineic us
Elpina
And ena nig oaenaea
Ainhoa Mag ach e al. 21
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Si e
Pollina o species
Elpina
Apis melli e a
Elpina
Bombus e es is
Elpina
Ce a ina cucu bi ina
Elpina
Empis essella a
Elpina
Euce a al e nans
Elpina
Euce a sp.
Elpina
Les ica clypea a
Elpina
Pla ynochae us se osus
Elpina
Psilo h ix i idicoe ulea
Elpina
Xylocopa can ab i a
Elpozo
And ena hispania
Elpozo
And ena sp.
Elpozo
Apis melli e a
Elpozo
Bombus e es is
Elpozo
Bombylius mo pho1
Elpozo
Bombylius sp.
Elpozo
Bombylius o qua us
Elpozo
Colle es nig icans
Elpozo
Dasypoda c assico nis
Elpozo
Helio au us u icollis
Elpozo
Lasioglossum bimacula us
Elpozo
Lasioglossum imminu us
Elpozo
Me odon sp.
Elpozo
Musca sp.
Elpozo
Panu gus sp.
Elpozo
Psilo h ix i idicoe ulea
Elpozo
Xylocopa iolacea
Espa agal
And ena sp.
Espa agal
An hopho a a oalba
Espa agal
Apidae sp.
Espa agal
Apis melli e a
Espa agal
Ce ce is mo pho1
Espa agal
Chasma op e us illige i
Espa agal
Dasypoda sp.
Espa agal
Episy phus bal ea us
Espa agal
Euce a colla is
Espa agal
Halic us idi isus
Espa agal
Lasioglossum bimacula us
Espa agal
Lasioglossum leucozonium
Espa agal
Lasioglossum malachu um
Espa agal
Lasioglossum mo pho1
Espa agal
Osmia ul i en is
Espa agal
Pie is apae
Espa agal
Ten h edo sp.
Espa agal
Usia mo pho1
LaCunya
And ena hyssono a
LaCunya
An hopho a dispa
LaCunya
An hopho a e usa
LaCunya
Apis melli e a
LaCunya
Bombus e es is
LaCunya
Ce a ina cucu bi ina
LaCunya
Dasypoda cingula a
LaCunya
Empis mo pho1
22 Ainhoa Mag ach e al.
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Si e
Pollina o species
LaCunya
Helio au us u icollis
LaCunya
Lasioglossum albocinc um
LaCunya
Lasioglossum imminu us
LaCunya
Lasioglossum malachu um
LaCunya
Lasioglossum sp.
LaCunya
Lasioglossum idi isus
LaCunya
Loma ia mo pho1
LaCunya
Panu gus banksianus
LaCunya
Pie is b assicae
LaCunya
Pseudoan hidium melano um
LaRocina
And ena sp.
LaRocina
An hopho a bimacula a
LaRocina
An hopho a e usa
LaRocina
Apis melli e a
LaRocina
A achnospila mo pho1
LaRocina
Bombus e es is
LaRocina
Ce a ina sp.
LaRocina
Colle es acu us
LaRocina
Colle es sp.
LaRocina
Dasypoda cingula a
LaRocina
Dasypoda c assico nis
LaRocina
Dasypoda sp.
LaRocina
Dischis us mo pho1
LaRocina
Dischis us senex
LaRocina
Episy phus bal ea us
LaRocina
E is alis enax
LaRocina
Helophilus i i a us
LaRocina
He iades c enula us
LaRocina
He iades unco um
LaRocina
Hopli is iden a a
LaRocina
Lasioglossum imminu us
LaRocina
Lasioglossum malachu um
LaRocina
Lasioglossum sp.
LaRocina
Malachius mo pho1
LaRocina
Me odon sp.
LaRocina
Nomada ag es is
LaRocina
Nomada uca a
LaRocina
Nomada mela ho acica
LaRocina
Osmia cae ulescens
LaRocina
Panu gus banksianus
LaRocina
Panu gus sp.
LaRocina
Rhyncomyia cup ea
LaRocina
Sphecodes sp.
LaRocina
Sy phidae sp.
LaRocina
Xylocopa can ab i a
Lasmulas
And ena la ipes
Lasmulas
And ena nig oaenaea
Lasmulas
And ena hyssono a
Lasmulas
An hopho a dispa
Lasmulas
An hopho a hispanica
Lasmulas
Apis melli e a
Lasmulas
Bombylius sp.
Lasmulas
Bombylius o qua us
Ainhoa Mag ach e al. 23
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Si e
Pollina o species
Lasmulas
Conopidae sp.
Lasmulas
Dasypoda albimana
Lasmulas
Empis sp.
Lasmulas
Empis es acea
Lasmulas
E is alis similis
Lasmulas
Euce a ch ysopyga
Lasmulas
Euce a sp.
Lasmulas
Fla ipanu gus enus us
Lasmulas
Helio au us u icollis
Lasmulas
Lasioglossum imminu us
Lasmulas
Lasioglossum malachu um
Lasmulas
Myc e us cu culioides
Lasmulas
Panu gus calca a us
Lasmulas
Panu gus da gius
Lasmulas
Xylocopa can ab i a
Niebla
And ena la ipes
Niebla
And ena labialis
Niebla
And ena o a ula
Niebla
And ena hyssono a
Niebla
And ena enuis ia a
Niebla
An hidium sep emspinosum
Niebla
An hopho a dispa
Niebla
An hopho a hispanica
Niebla
An hopho a sp.
Niebla
Apis melli e a
Niebla
Bombus e es is
Niebla
Bombylius imb ia us
Niebla
Bombylius sp.
Niebla
Ce a ina callosa
Niebla
Colle es sp.
Niebla
Episy phus bal ea us
Niebla
Euce a colla is
Niebla
Euce a no a a
Niebla
Exosoma lusi anicum
Niebla
Halic us scabiosae
Niebla
Helio au us u icollis
Niebla
He iades c enula us
Niebla
Lasioglossum malachu um
Niebla
Lasioglossum sp.
Niebla
Mac ophya mon ana
Niebla
Me odon sp.
Niebla
Osmia bico nis
Niebla
Osmia submicans
Niebla
Panu gus banksianus
Niebla
Panu gus da gius
Niebla
Pla ynochae us se osus
Niebla
Po osia cup ea
Niebla
Rhodan hidium s ic icum
Niebla
Sphae opho ia sc ip a
Niebla
Sys opha planidens
Niebla
Usia mo pho1
Niebla
Usia mo pho2
Niebla
Usia sp.
24 Ainhoa Mag ach e al.
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Si e
Pollina o species
Niebla
Vespula ge manica
Niebla
Xylocopa iolacea
Pina esdeHinojos
And ena hispania
Pina esdeHinojos
Apis melli e a
Pina esdeHinojos
Bombus e es is
Pina esdeHinojos
Ch ysu a e ulgens
Pina esdeHinojos
Colle es acu us
Pina esdeHinojos
Colle es nig icans
Pina esdeHinojos
Colle es sp.
Pina esdeHinojos
Dasypoda c assico nis
Pina esdeHinojos
Lasioglossum bimacula us
Pina esdeHinojos
Lasioglossum malachu um
Pina esdeHinojos
Lasioglossum sp.
Pina esdeHinojos
Nomada mela ho acica
Pina esdeHinojos
Panu gus da gius
Pina esdeHinojos
Psilo h ix i idicoe ulea
Pina esdeHinojos
Ten h edo co yne es
Pina esdeHinojos
Xylocopa can ab i a
Pinodelcue o
Ancis oce us gazella
Pinodelcue o
Ancis oce us econdi us
Pinodelcue o
And ena hispania
Pinodelcue o
And ena sp.
Pinodelcue o
Apis melli e a
Pinodelcue o
Bombylella a a
Pinodelcue o
Bombylius sp.
Pinodelcue o
Ce a ina cucu bi ina
Pinodelcue o
Ce ce is mo pho1
Pinodelcue o
Dasypoda cingula a
Pinodelcue o
Fla ipanu gus enus us
Pinodelcue o
Lasioglossum sexno a um
Pinodelcue o
Loma ia mo pho1
Pinodelcue o
Megascolia macula a
Pinodelcue o
Me odon sp.
Pinodelcue o
Musca sp.
Pinodelcue o
Nomada mela ho acica
Pinodelcue o
Nomada me ce i
Pinodelcue o
Nomada sp.
Pinodelcue o
Panu gus cephalo es
Pinodelcue o
Pelecoce a icinc a
Pinodelcue o
Sys oechus mo pho1
Pinodelcue o
Usia sp.
Pinodelcue o
Xylocopa can ab i a
U banizaciones
And ena sp.
U banizaciones
And ena ulpecula
U banizaciones
Apis melli e a
U banizaciones
Bombus e es is
U banizaciones
Bombylidae mo pho1
U banizaciones
Bombylius sp.
U banizaciones
Ce a ina sp.
U banizaciones
Colle es nig icans
U banizaciones
Dasypoda cingula a
U banizaciones
Dasypoda sp.
U banizaciones
Dischis us senex
Ainhoa Mag ach e al. 25
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Table S4. Lis o plan species su eyed and hei ma ing sys em.
Plan _ amily
Plan _genus
Plan _species
ep oduc i e_sys em
Cis aceae
Cis us
monspeliensis
sel -incompa ible
Cis aceae
Cis us
c ispus
sel -incompa ible
Cis aceae
Cis us
ladani e
sel -incompa ible
Cis aceae
Cis us
sal ii olius
sel -incompa ible
Cis aceae
Cis us
albidus
sel -incompa ible
Cis aceae
Cis us
libano is
sel -incompa ible
Cis aceae
Halimium
commu a um
sel -incompa ible
Cis aceae
Halimium
halimi olium
sel -incompa ible
Lamiaceae
La andula
peduncula a
pa ially sel -compa ible
Lamiaceae
La andula
s oechas
pa ially sel -compa ible
Lamiaceae
Teuc ium
u icans
pa ially sel -compa ible
Lamiaceae
Rosma inus
o icinalis
pa ially sel -compa ible
Lamiaceae
Phlomis
pu pu ea
sel -incompa ible
Xan ho hoeaceae
Asphodelus
is ulosus
pa ially sel -compa ible
Fabaceae
Ulex
aus alis
sel -incompa ible
Fabaceae
Spa ium
junceum
sel -incompa ible
Fabaceae
As agalus
lusi anicus
pa ially sel -compa ible
Fabaceae
Re ama
sphae oca pa
pa ially sel -compa ible
Bo aginaceae
Anchusa
azu ea
sel -incompa ible
32 Ainhoa Mag ach e al.
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Table S5. Resul s o GLM showing e ec o simple isi a ion me ics on A) si e-le el a e age ui
weigh based on bes model selec ed and B) he same analysis emo ing one si e ha has a pa icula ly
la ge pollina o ichness alue o es whe he his poin migh be d i ing he ela ionship.
A)
Es ima e
S d.
E o
alue
(In e cep )
0.08
0.01
8.56
Pollina o ichness
0.02
0.01
2.11
Rela i e numbe o
isi s
0.01
0.01
0.78
B)
Es ima e
S d.
E o
alue
(In e cep )
0.08
0.01
8.04
Pollina o ichness
0.02
0.01
1.97
Rela i e numbe o
isi s
0.01
0.01
0.76
Ainhoa Mag ach e al. 33
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Table S6. Resul s o GLM showing e ec o simple isi a ion me ics on equi y in ep oduc i e success
ac oss plan species wi hin a si e based on bes model selec ed (0.50 h eshold).
Es ima e
S d. E o
z
alue
(In e cep )
0.40
0.63
0.64
Pollina o ichness
-0.41
1.18
-0.35
Rela i e numbe o
isi s
-0.28
0.72
-0.39
Nes edness
0.87
1.02
0.86
Pollina o niche
complemen a i y
-0.51
1.32
-0.38
Re e ences
Ab ams PA, Hol RD, Ro h JD (1998) APPARENT COMPETITION OR APPARENT MUTUALISM? SHARED
PREDATION WHEN POPULATIONS CYCLE. Ecology, 79, 201–212. h ps://doi.o g/10.1890/0012-
9658(1998)079[0201:acoams]2.0.co;2
Alb ech M, Schmid B, Hau ie Y, Mülle CB (2012) Di e se pollina o communi ies enhance plan
ep oduc i e success. P oceedings o he Royal Socie y B: Biological Sciences, 279, 4845–4852.
h ps://doi.o g/10.1098/ spb.2012.1621
Almeida-Ne o M, Ul ich W (2011) A s aigh o wa d compu a ional app oach o measu ing nes edness
using quan i a i e ma ices. En i onmen al Modelling & So wa e, 26, 173–178.
h ps://doi.o g/10.1016/j.en so .2010.08.003
A ceo-Gómez G, Sch oede A, Albo C, Ashman T-L, Knigh TM, Benne JM, Sua ez B, Pa a-Tabla V (2019)
Global geog aphic pa e ns o he e ospeci ic pollen eceip help unco e po en ial ecological and
e olu iona y impac s ac oss plan communi ies wo ldwide. Scien i ic Repo s, 9.
h ps://doi.o g/10.1038/s41598-019-44626-0
Bagchi R, Swin ield T, Galle y RE, Lewis OT, G ipenbe g S, Na ayan L, F eckle on RP (2010) Tes ing he
Janzen-Connell mechanism: pa hogens cause o e compensa ing densi y dependence in a opical ee.
Ecology Le e s, 13, 1262–1269. h ps://doi.o g/10.1111/j.1461-0248.2010.01520.x
Ba omeus I, S a e JR, Wa d D, Aguado O (2018) His o ical collec ions as a ool o assessing he global
pollina ion c isis. Philosophical T ansac ions o he Royal Socie y B: Biological Sciences, 374, 20170389.
h ps://doi.o g/10.1098/ s b.2017.0389
Bascomp e J, Jo dano P (2007) Plan -Animal Mu ualis ic Ne wo ks: The A chi ec u e o Biodi e si y. Annual
Re iew o Ecology, E olu ion, and Sys ema ics, 38, 567–593.
h ps://doi.o g/10.1146/annu e .ecolsys.38.091206.095818
Bascomp e J, Jo dano P, Melian CJ, Olesen JM (2003) The nes ed assembly o plan -animal mu ualis ic
ne wo ks. P oceedings o he Na ional Academy o Sciences, 100, 9383–9387.
h ps://doi.o g/10.1073/pnas.1633576100
Bas olla U, Fo una MA, Pascual-Ga cía A, Fe e a A, Luque B, Bascomp e J (2009) The a chi ec u e o
mu ualis ic ne wo ks minimizes compe i ion and inc eases biodi e si y. Na u e, 458, 1018–1020.
h ps://doi.o g/10.1038/na u e07950
Benadi G, Blü hgen N, Ho es ad T, Poe hke H-J (2013) When Can Plan -Pollina o In e ac ions P omo e
Plan Di e si y? The Ame ican Na u alis , 182, 131–146. h ps://doi.o g/10.1086/670942
Benadi G, Pauw A (2018) F equency dependence o pollina o isi a ion a es sugges s ha pollina ion
niches can allow plan species coexis ence (I Ba omeus, Ed,). Jou nal o Ecology, 106, 1892–1901.
h ps://doi.o g/10.1111/1365-2745.13025
34 Ainhoa Mag ach e al.
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Benne JM, Thompson A, Goia I, Feldmann R, Ş e an V, Bogdan A, Rakosy D, Beloiu M, Bi o I-B, Bluemel S,
Filip M, Madaj A-M, Ma in A, Passonneau S, Kalisch DP, Sche e G, Knigh TM (2018) A e iew o
Eu opean s udies on pollina ion ne wo ks and pollen limi a ion, and a case s udy designed o ill in a
gap. AoB PLANTS, 10. h ps://doi.o g/10.1093/aobpla/ply068
Biesmeije JC, Robe s SPM, Reeme M, Ohlemulle R, Edwa ds M, Pee e s T, Scha e s AP, Po s SG,
Kleuke s R, Thomas CD, Se ele J, Kunin WE (2006) Pa allel Declines in Pollina o s and Insec -Pollina ed
Plan s in B i ain and he Ne he lands. Science, 313, 351–354. h ps://doi.o g/10.1126/science.1127863
Blü hgen N (2010) Why ne wo k analysis is o en disconnec ed om communi y ecology: A c i ique and an
ecologis ’s guide. Basic and Applied Ecology, 11, 185–195. h ps://doi.o g/10.1016/j.baae.2010.01.001
Blü hgen N, Klein A-M (2011) Func ional complemen a i y and specialisa ion: The ole o biodi e si y in
plan pollina o in e ac ions. Basic and Applied Ecology, 12, 282–291.
h ps://doi.o g/10.1016/j.baae.2010.11.001
Blü hgen N, Menzel F, Blü hgen N (2006) BMC Ecology, 6, 9. h ps://doi.o g/10.1186/1472-6785-6-9
Bomma co R, Ma ini L, Vaissiè e BE (2012) Insec pollina ion enhances seed yield, quali y, and ma ke alue
in oilseed ape. Oecologia, 169, 1025–1032. h ps://doi.o g/10.1007/s00442-012-2271-6
B uno JF, S achowicz JJ, Be ness MD (2003) Inclusion o acili a ion in o ecological heo y. T ends in
Ecology & E olu ion, 18, 119–125. h ps://doi.o g/10.1016/s0169-5347(02)00045-9
Bu nham KP, Ande son DR (Eds.) (2004) Model selec ion and mul imodel in e ence. Sp inge New Yo k.
h ps://doi.o g/10.1007/b97636
Bu nham KP, Ande son DR, Huy ae KP (2010) AIC model selec ion and mul imodel in e ence in beha io al
ecology: some backg ound, obse a ions, and compa isons. Beha io al Ecology and Sociobiology, 65,
23–35. h ps://doi.o g/10.1007/s00265-010-1029-6
By nes JEK, Gam eld L, Isbell F, Le check JS, G i in JN, Hec o A, Ca dinale BJ, Hoope DU, Dee LE, Emme
Du y J (2014) In es iga ing he ela ionship be ween biodi e si y and ecosys em mul i unc ionali y:
challenges and solu ions (R F eckle on, Ed,). Me hods in Ecology and E olu ion, 5, 111–124.
h ps://doi.o g/10.1111/2041-210x.12143
Ca alhei o LG, Biesmeije JC, Benadi G, F ünd J, S ang M, Ba omeus I, Kaise -Bunbu y CN, Baude M,
Gomes SIF, Me ckx V, Baldock KCR, Benne ATD, Boada R, Bomma co R, Ca a R, Chaco N, Dänha d
J, Dicks LV, Do mann CF, Ek oos J, Henson KSE, Holzschuh A, Junke RR, Lopeza aiza-Mikel M, Memmo
J, Mon e o-Cas año A, Nelson IL, Pe anidou T, Powe EF, Rundlö M, Smi h HG, S ou JC, Temi ope K,
Tscha n ke T, Tscheulin T, Vilà M, Kunin WE (2014) The po en ial o indi ec e ec s be ween co-
lowe ing plan s ia sha ed pollina o s depends on esou ce abundance, accessibili y and ela edness
(F Jo dan, Ed,). Ecology Le e s, 17, 1389–1399. h ps://doi.o g/10.1111/ele.12342
Chaco NP, Vázquez DP, Lomáscolo SB, S e ani EL, Do ado J, Pad ón B (2011) E alua ing sampling
comple eness in a dese plan -pollina o ne wo k. Jou nal o Animal Ecology, 81, 190–200.
h ps://doi.o g/10.1111/j.1365-2656.2011.01883.x
Chao A, Colwell RK, Lin C-W, Go elli NJ (2009) Su icien sampling o asymp o ic minimum species ichness
es ima o s. Ecology, 90, 1125–1133. h ps://doi.o g/10.1890/07-2147.1
Delmas E, Besson M, B ice M-H, Bu kle LA, Dalla Ri a GV, Fo in M-J, G a el D, Guima ães PR, Hemb y DH,
Newman EA, Olesen JM, Pi es MM, Yeakel JD, Poiso T (2018) Analysing ecological ne wo ks o species
in e ac ions. Biological Re iews, 94, 16–36. h ps://doi.o g/10.1111/b .12433
De o o M, Bailey S, C aze P, Memmo J (2012) Unde s anding and planning ecological es o a ion o plan -
pollina o ne wo ks. Ecology Le e s, 15, 319–328. h ps://doi.o g/10.1111/j.1461-0248.2012.01740.x
Do mann CF, F und J, Blu hgen N, G ube B (2009) Indices, G aphs and Null Models: Analyzing Bipa i e
Ecological Ne wo ks. The Open Ecology Jou nal, 2, 7–24.
h ps://doi.o g/10.2174/1874213000902010007
Eu opean Commission. Di ec o a e Gene al o he En i onmen ., IUCN (In e na ional Union o
Conse a ion o Na u e). (2014) Eu opean ed lis o bees. Publica ions O ice, LU.
h ps://doi.o g/10.2779/77003
Flanagan RJ, Mi chell RJ, Knu owski D, Ka on JD (2009) In e speci ic pollina o mo emen s educe pollen
deposi ion and seed p oduc ion in Mimulus ingens (Ph ymaceae). Ame ican Jou nal o Bo any, 96,
809–815. h ps://doi.o g/10.3732/ajb.0800317
Ainhoa Mag ach e al. 35
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Fon aine C, Dajoz I, Me igue J, Lo eau M (2005) Func ional Di e si y o Plan Pollina o In e ac ion Webs
Enhances he Pe sis ence o Plan Communi ies (N Wase , Ed,). PLoS Biology, 4, e1.
h ps://doi.o g/10.1371/jou nal.pbio.0040001
F ünd J, Do mann CF, Holzschuh A, Tscha n ke T (2013) Bee di e si y e ec s on pollina ion depend on
unc ional complemen a i y and niche shi s. Ecology, 94, 2042–2054. h ps://doi.o g/10.1890/12-
1620.1
Ga ibaldi LA, Ca alhei o LG, Leonha d SD, Aizen MA, Blaauw BR, Isaacs R, Kuhlmann M, Kleijn D, Klein
AM, K emen C, Mo andin L, Schepe J, Win ee R (2014) F om esea ch o ac ion: enhancing c op yield
h ough wild pollina o s. F on ie s in Ecology and he En i onmen , 12, 439–447.
h ps://doi.o g/10.1890/130330
Ga ibaldi LA, S e an-Dewen e I, Win ee R, Aizen MA, Bomma co R, Cunningham SA, K emen C,
Ca alhei o LG, Ha de LD, A ik O, Ba omeus I, Benjamin F, Bo eux V, Ca i eau D, Chaco NP,
Dudenho e JH, F ei as BM, Ghazoul J, G eenlea S, Hipoli o J, Holzschuh A, Howle B, Isaacs R, Ja o ek
SK, Kennedy CM, K ewenka KM, K ishnan S, Mandelik Y, May ield MM, Mo zke I, Munyuli T, Naul BA,
O ieno M, Pe e sen J, Pisan y G, Po s SG, Rade R, Ricke s TH, Rundlo M, Seymou CL, Schuepp C,
Szen gyo gyi H, Taki H, Tscha n ke T, Ve ga a CH, Viana BF, Wange TC, Wes phal C, Williams N, Klein
AM (2013) Wild Pollina o s Enhance F ui Se o C ops Rega dless o Honey Bee Abundance. Science,
339, 1608–1611. h ps://doi.o g/10.1126/science.1230200
Godoy O, Ba omeus I, Roh RP, Saa ed a S (2018) Towa ds he In eg a ion o Niche and Ne wo k Theo ies.
T ends in Ecology & E olu ion, 33, 287–300. h ps://doi.o g/10.1016/j. ee.2018.01.007
Godoy O, K a NJB, Le ine JM (2014) Phylogene ic ela edness and he de e minan s o compe i i e
ou comes (J Cha e, Ed,). Ecology Le e s, 17, 836–844. h ps://doi.o g/10.1111/ele.12289
Goldbe g DE, Ba on AM (1992) Pa e ns and Consequences o In e speci ic Compe i ion in Na u al
Communi ies: A Re iew o Field Expe imen s wi h Plan s. The Ame ican Na u alis , 139, 771–801.
h ps://doi.o g/10.1086/285357
Hol RD (1977) P eda ion, appa en compe i ion, and he s uc u e o p ey communi ies. Theo e ical
Popula ion Biology, 12, 197–229. h ps://doi.o g/10.1016/0040-5809(77)90042-9
Hsieh TC, Ma KH, Chao A (2016) iNEXT: an R package o a e ac ion and ex apola ion o species di e si y
( H ill numbe s) (G McIne ny, Ed,). Me hods in Ecology and E olu ion, 7, 1451–1456.
h ps://doi.o g/10.1111/2041-210x.12613
James A, Pi ch o d JW, Plank MJ (2012) Disen angling nes edness om models o ecological complexi y.
Na u e, 487, 227–230. h ps://doi.o g/10.1038/na u e11214
Johnson CA, B ons ein JL (2019) Coexis ence and compe i i e exclusion in mu ualism. Ecology, e02708.
h ps://doi.o g/10.1002/ecy.2708
Kaise -Bunbu y CN, Mougal J, Whi ing on AE, Valen in T, Gab iel R, Olesen JM, Blü hgen N (2017)
Ecosys em es o a ion s eng hens pollina ion ne wo k esilience and unc ion. Na u e, 542, 223–227.
h ps://doi.o g/10.1038/na u e21071
Lanuza JB, Ba omeus I, Godoy O (2018) Opposing e ec s o lo al isi o s and soil condi ions on he
de e minan s o compe i i e ou comes main ain species di e si y in he e ogeneous landscapes (JM
Gómez, Ed,). Ecology Le e s, 21, 865–874. h ps://doi.o g/10.1111/ele.12954
Láza o A, Gómez-Ma ínez C, Aloma D, González-Es é ez MA, T a ese A (2020) Linking species-le el
ne wo k me ics o lowe ai s and plan i ness (N Ra e y, Ed,). Jou nal o Ecology, 108, 1287–1298.
h ps://doi.o g/10.1111/1365-2745.13334
Láza o A, Lundg en R, To land Ø (2014) Expe imen al educ ion o pollina o isi a ion modi ies plan -plan
in e ac ions o pollina ion. Oikos, 123, 1037–1048. h ps://doi.o g/10.1111/oik.01268
Maca hu R, Le ins R (1967) The Limi ing Simila i y, Con e gence, and Di e gence o Coexis ing Species.
The Ame ican Na u alis , 101, 377–385. h ps://doi.o g/10.1086/282505
May RM, A hu RHM (1972) Niche O e lap as a Func ion o En i onmen al Va iabili y. P oceedings o he
Na ional Academy o Sciences, 69, 1109–1113. h ps://doi.o g/10.1073/pnas.69.5.1109
May ield MM, S ou e DB (2017) Highe -o de in e ac ions cap u e unexplained complexi y in di e se
communi ies. Na u e Ecology & E olu ion, 1. h ps://doi.o g/10.1038/s41559-016-0062
Memmo J, Wase NM, P ice MV (2004) Tole ance o pollina ion ne wo ks o species ex inc ions.
P oceedings o he Royal Socie y o London. Se ies B: Biological Sciences, 271, 2605–2611.
h ps://doi.o g/10.1098/ spb.2004.2909
36 Ainhoa Mag ach e al.
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1
Mo ales CL, T a ese A (2008) In e speci ic Pollen T ans e : Magni ude, P e alence and Consequences o
Plan Fi ness. C i ical Re iews in Plan Sciences, 27, 221–238.
h ps://doi.o g/10.1080/07352680802205631
Nakagawa S, Johnson PCD, Schielze h H (2017) The coe icien o de e mina ion R 2 and in a-class
co ela ion coe icien om gene alized linea mixed-e ec s models e isi ed and expanded. Jou nal o
The Royal Socie y In e ace, 14, 20170213. h ps://doi.o g/10.1098/ si .2017.0213
Pauw A (2013) Can pollina ion niches acili a e plan coexis ence? T ends in Ecology & E olu ion, 28, 30–
37. h ps://doi.o g/10.1016/j. ee.2012.07.019
Poiso T, Mouque N, G a el D (2013) T ophic complemen a i y d i es he biodi e si y-ecosys em
unc ioning ela ionship in ood webs (F Adle , Ed,). Ecology Le e s, 16, 853–861.
h ps://doi.o g/10.1111/ele.12118
Po s SG, Biesmeije JC, K emen C, Neumann P, Schweige O, Kunin WE (2010) Global pollina o declines:
ends, impac s and d i e s. T ends in Ecology & E olu ion, 25, 345–353.
h ps://doi.o g/10.1016/j. ee.2010.01.007
R De elopmen Co e Team R (2011) R: A language and en i onmen o s a is ical compu ing (RDC Team,
Ed,). h ps://doi.o g/10.1007/978-3-540-74686-7
Saa ed a S, S ou e DB, Uzzi B, Bascomp e J (2011) S ong con ibu o s o ne wo k pe sis ence a e he
mos ulne able o ex inc ion. Na u e, 478, 233–235. h ps://doi.o g/10.1038/na u e10433
Simmons BI, Hoeppke C, Su he land WJ (2019) Bewa e g eedy algo i hms (A Eklö , Ed,). Jou nal o Animal
Ecology, 88, 804–807. h ps://doi.o g/10.1111/1365-2656.12963
Song C, Roh RP, Saa ed a S (2017) Why a e some plan pollina o ne wo ks mo e nes ed han o he s? (A
Eklö , Ed,). Jou nal o Animal Ecology, 86, 1417–1424. h ps://doi.o g/10.1111/1365-2656.12749
S a e JR, Ba omeus I, Beggs JR, Gaske AC, Pa emo e DE (2019) Plan species dominance inc eases
pollina ion complemen a i y and plan ep oduc i e unc ion. Ecology, 100.
h ps://doi.o g/10.1002/ecy.2749
Thébaul E, Fon aine C (2010) S abili y o Ecological Communi ies and he A chi ec u e o Mu ualis ic and
T ophic Ne wo ks. Science, 329, 853–856. h ps://doi.o g/10.1126/science.1188321
Thompson RM, B ose U, Dunne JA, Hall RO, Hladyz S, Ki ching RL, Ma inez ND, Ran ala H, Romanuk TN,
S ou e DB, Tylianakis JM (2012) Food webs: econciling he s uc u e and unc ion o biodi e si y.
T ends in Ecology & E olu ion, 27, 689–697. h ps://doi.o g/10.1016/j. ee.2012.08.005
Thomson DM (2018) E ec s o long- e m a ia ion in pollina o abundance and di e si y on ep oduc ion
o a gene alis plan (I Ba omeus, Ed,). Jou nal o Ecology, 107, 491–502.
h ps://doi.o g/10.1111/1365-2745.13055
Tilman D (1982) Resou ce compe i ion and communi y s uc u e. (MPB-17), olume 17. P ince on Uni e si y
P ess. h ps://doi.o g/10.1515/9780691209654
Valdo inos FS, B osi BJ, B iggs HM, Moisse de Espanés P, Ramos-Jilibe o R, Ma inez ND (2016) Niche
pa i ioning due o adap i e o aging e e ses e ec s o nes edness and connec ance on pollina ion
ne wo k s abili y (J Chase, Ed,). Ecology Le e s, 19, 1277–1286. h ps://doi.o g/10.1111/ele.12664
Vanbe gen AJ, Woodcock BA, G ay A, G an F, Tel o d A, Lambdon P, Chapman DS, Pywell RF, Hea d MS,
Ca e s S (2013) G azing al e s insec isi a ion ne wo ks and plan ma ing sys ems (A B ody, Ed,).
Func ional Ecology, 28, 178–189. h ps://doi.o g/10.1111/1365-2435.12191
Win ee R (2013) Global change, biodi e si y, and ecosys em se ices: Wha can we lea n om s udies o
pollina ion? Basic and Applied Ecology, 14, 453–460. h ps://doi.o g/10.1016/j.baae.2013.07.004
Woodwa d G, Hild ew AG (2002) Body-size de e minan s o niche o e lap and in aguild p eda ion wi hin
a complex ood web. Jou nal o Animal Ecology, 71, 1063–1074. h ps://doi.o g/10.1046/j.1365-
2656.2002.00669.x
Zhang J (2016) Spaa: SPecies associa ion analysis. h ps://gi hub.com/helixcn/spaa
Ainhoa Mag ach e al. 37
Pee Communi y Jou nal, Vol. 1 (2021), a icle e1 h ps://doi.o g/10.24072/pcjou nal.1