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Interactions between non-native parrot species their recipient environments

Hernández Brito, Dailos · 2020

Abstract

Biological invasions are amongst critical causes of biodiversity loss around the world, although interactions between invasive species, native biota and recipient environment are mostly unknown. Their identification may show how certain invasive species are able to establish successfully in novel environments and which potential ecological impacts threaten native biota. However, both interactions and their potential impacts are usually overlooked, especially in first stages of invasion, because they may arise between unexpected interact species as well as their detection depends of long-term studies. Parrots (Order Psittaciformes) are one of the groups most affected by international wildlife trade due to their appraisal as pets, consequently, 16% of parrot species present populations established out of their native range. Between non-native parrots, rose-ringed (Psittacula krameri) and monk (Myiopsitta monachus) parakeets are the most successful to establish and invade novel environments, even though numerous introductions have occurred in more than forty countries worldwide, their biotic interactions and impacts are poorly known. In this thesis, we combine different observations (behavioral, spatial distribution, environmental and population) to assess the role of different biotic interactions and their resultant impacts that arise between a total of X non-native parrot species (mainly rose-ringed and monk parakeets), recipient environment and biota. Our results show that non-native parrot species develop a wide range of biotic interactions (e.g. interspecific competition for nesting sites, facilitation of limiting resources and animal-plant mutualisms) that may threaten or benefit several coexisting species, both native and non-native. Thus, the emergence of adaptive behaviors (e.g. nesting association and innovation) in non-native parrots allows for overcoming barriers from recipient environment, such as biotic resistance and scarcity of limiting resources, or obstacles during their first stages of establishment (e.g. Allee effects). Overall, the wide analysis and study in long-term of biotic interactions here performed shows their importance as one of key mechanisms for success of establishment and invasion process in non-native parrots species. Moreover, their associated ecological impacts are a risk for native species, especially threaten species, hence, the implementation of efficient management plans is urgently necessary to conserve the biodiversity. Key words: Psittaciformes; biological invasions; interspecific competition; biotic resistance; commensalism; behavioral innovation; facilitation; hybridization; Allee effect; animal-plant mutualism; seed dispersal; zoochory; impact; eradication; management plans; population viability

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Interactions between non-native parrot species their recipient environments

Author: Hernández Brito, Dailos
Year: 2020
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In e ac ions be ween non-na i e pa o species
and hei ecipien en i onmen s
Dailos He nández B i o
PhD Thesis
Se ille, 2020
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RECOMMENDED CITATION:
He nández-B i o, D. (2020). In e ac ions be ween non-na i e pa o species and hei
ecipien en i onmen s. PhD Thesis. Uni e si y o Se ille, Se ille, Spain
Co e and co e back pho os a e au ho ship o Dailos He nández-B i o, who emains
his in ellec ual owne . Any o m o ep oduc ion, dis ibu ion, public communica ion o
ans o ma ion o he same wi hou p io au ho iza ion o he au ho is o bidden.
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In e ac ions
and
In e accione
s
y
los
Supe iso s: D .
José Luis
Tu o :
D . Ja ie Ba
Depa men o
Co
ns be ween non
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na i e pa o spe
d hei ecipien en i onmen s
-
s en e especies de psi ácidos
exó
los nue os medios que ocupan
Thesis o he deg ee o Doc o
Uni e si y o Se ille
Se ille, 2020
Dailos He nández B i o
uis Tella Escobedo
D a.
Ma ina
Balbon ín A enas
Conse a ion
Biology
pecies
xó icos
na Ca e e
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Que no puede se sino sal aje y desmedido
Jean-Claude Nicolas Fo es ie
A qui ec o paisajis a
(Du an e el diseño del Pa que de Ma ía Luisa)
A mis pad es
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CONTENT
SUMMARY / RESÚMEN ...............................................................................................1
THESIS INTRODUCTION .............................................................................................7
Biological in asions: an issue o global biodi e si y ..........................................9
No el in e ac ions be ween non-na i e species and ecipien communi ies .......11
Pa o s, unin en ional cha ac e s in he scena io o biological in asion .............12
S udy species: he mos success ul in ade s
and o he non-na i e pa o species ....................................................................15
Objec i es, hypo heses and hesis s uc u e ........................................................18
Re e ences ...........................................................................................................21
SECTION 1: BIOTIC INTERACTIONS IN CAVITY NESTER COMMUNITIES.....32
Chap e I: C owding in he ci y: losing and winning compe i o s
o an in asi e bi d ...............................................................................................34
Re e ences ...........................................................................................................61
Chap e II: Alien ose- inged pa akee s (Psi acula k ame i) a ack
black a s (Ra us a us) some imes esul ing in dea h ......................................68
Re e ences ...........................................................................................................75
Chap e III: Nes -si e compe i ion and killing by in asi e pa akee s
cause he decline o a h ea ened ba popula ion ................................................79
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Resúmen
4
RESÚMEN
Las in asiones biológicas es án en e las causas más impo an es de pé dida de
biodi e sidad en odo el mundo, aunque las in e acciones en e las especies in aso as, la
bio a na i a y el medio ecep o son en su mayo ía desconocidas. Su iden i icación
puede mos a cómo cie as especies in aso as pueden es ablece se exi osamen e en
nue os ambien es y qué impac os ecológicos po enciales amenazan la bio a na i a. Sin
emba go, an o las in e acciones como sus impac os po enciales gene almen e se pasan
po al o, especialmen e en las p ime as e apas de la in asión, po que és os pueden su gi
en e especies que in e ac úan inespe adamen e, po lo que su de ección depende de
es udios a la go plazo. Los lo os (O den Psi aci o mes) son uno de los g upos más
a ec ados po el come cio in e nacional de ida sil es e debido a su alo como
masco as, po lo que el 16% de las especies de lo os p esen an poblaciones es ablecidas
ue a de su á ea de dis ibución na i a. En e los lo os no na i os, las co o as de
K ame (Psi acula k ame i) y a gen ina (Myiopsi a monachus) son las más exi osas
pa a es ablece se e in adi en nue os ambien es, aunque se han p oducido nume osas
in oducciones en más de cua en a países de odo el mundo, sus in e acciones e
impac os bió icos son poco conocidos. En es a esis, combinamos di e en es
obse aciones (conduc uales, de dis ibución espacial, ambien ales y poblacionales)
pa a e alua el papel de las di e en es in e acciones bió icas y sus impac os esul an es
que su gen en e di e en es especies de lo os no na i os (p incipalmen e co o as de
K ame y a gen ina), ambien e y bio a ecep o es. Nues os esul ados mues an que las
especies de lo os no na i os desa ollan una amplia gama de in e acciones bió icas (po
ejemplo, compe encia in e especí ica po si ios de anidación, acili ación de ecu sos
limi an es y mu ualismos animal-plan a) que pueden amenaza o bene icia a a ias
especies coexis en es, an o na i as como no na i as. Po lo an o, la apa ición de
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Resúmen
5
compo amien os adap a i os (po ejemplo, asociación e inno ación en la anidación) en
lo os no na i os pe mi e supe a las ba e as p o enien es del ambien e ecep o , como
la esis encia bió ica y la escasez de ecu sos limi an es, u o os obs áculos du an e sus
p ime as e apas de es ablecimien o (po ejemplo, e ec os Allee). En gene al, el amplio
análisis y es udio a la go plazo de las in e acciones bió icas aquí ealizado mues an su
impo ancia como uno de los mecanismos cla e pa a el éxi o del p oceso de
es ablecimien o e in asión en especies de lo os no na i os. Además, sus impac os
ecológicos esul an es son un iesgo pa a las especies na i as, especialmen e las
especies amenazadas, po lo que la implemen ación de planes de manejo e icien es es
u gen e pa a conse a la biodi e sidad.
Palab as cla e: Psi aci o mes; in asiones biológicas; compe encia in e especí ica;
esis encia bió ica; comensalismo; inno ación compo amen al; acili ación;
hib idación; E ec o Allee; mu ualismo plan a-animal; dispe sión de semillas; zooco ia;
impac o; e adicación; planes de manejo; iabilidad poblacional
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7
THESIS INTRODUCTION
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Thesis In oduc ion
9
Biological in asions: an issue o global biodi e si y
Biodi e si y is esul an o na u al his o y p ocesses ha ope a e in di e en space- ime
scales and in luence on key ac o s om gene ic, h ough species and en i onmen al o
ecologic by which assemble he biological he e ogenei y (Walke 1992). Likewise,
biodi e si y in luences on ecosys em unc ions and p o ides good and se ices o
humani y (Ca dinale e al. 2012). Howe e , he dis up ion o human ac i i y has d i en
deep al e a ions on hese p ocesses and consequen ly endange he local and global
biodi e si y (Vi ousek e al. 1997; Ca dinale e al. 2012). In his sense, he inc easing
biodi e si y loss and ex inc ion a es du ing he las cen u y a e unp eceden ed in
p e ious geological ages, esembling hus massi e ex inc ions om ossil eco d
e idences (Di zo e al. 2014; Ceballos e al. 2015; Wilson 2016). Such human impac
accele a ion on biodi e si y comp omises i s unde s anding and de ec ion o i s causes
and consequences, which in i s u n spoil e o s o p ese e global biodi e si y
(Dawson e al. 2011). Thus, di e en dis up i e human ac ions a e he mos impo an
d i e s o biodi e si y loss such as habi a des uc ion, o e exploi a ion, clima e change
and biological in asions (Sala e al. 2000; Williams e al. 2008), as well syne ge ic
in e ac ions be ween hese d i e s may inc ease u he hei impac s (Man yka-p ingle
e al. 2012).
T ansloca ion o animal and plan s by humans ha e occu ed du ing di e en
human mig a ions wo ldwide. Pimen el (2001) es ima es ha abou 400,000 species
ha e been emo ed om hei na u al geog aphical a eas o ano he by human agency
du ing he pas 10,000 yea s. Al hough he e is a conside able inc easing in
ansloca ion a es since 18 h cen u y due o echnological ad ances ha ha e assis ed
he long-dis ance anspo (Meye son and Mooney 2007; Hulme 2009). By delibe a ely
o acciden ally anspo ing, a la ge numbe o species ha e been in oduced in no el
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Thesis In oduc ion
10
en i onmen s, whe eby hese species ha e o e come geog aphical ba ie s and
es ablished non-na i e iable popula ions ou o hei na i e anges (Blackbu n e al.
2011; Seebens e al. 2017). The es ablishmen and sp ead o hese non-na i e species
(i.e. biological in asion p ocess) may in ol e impac s on na i e species, communi y
dynamics and ecosys em unc ions (Cla e o and Ga cía-Be hou 2005; Vilà e al. 2011),
in addi ion o economic damages and heal h issues (Vi ousek e al. 1996; Simbe lo e
al. 2013). Na i e species may be a ec ed h ough nega i e impac s such as p eda ion,
compe i ion, sp ead o diseases and hyb idiza ion (Mack e al. 2000; Money and
Cleland 2001), hence, hei consequences h ea iabili y o na i e popula ions, e en
hei ex inc ion (Cla e o e al . 2009; Bella d e al. 2016).
Ne e heless, he ole o ac o s ha d i e p ocesses in which non-na i e species
become in asi e, es ablishing and sp eading success ully in new geog aphical a eas, i
has been deba able among nume ous hypo heses (Kola and Lodge 2001; Ca o d e al.
2009; Blackbu n e al. 2011). O e all, hese key ac o s a e mainly ca ego ized in h ee
g oups ha may show syne gy be ween hem and change hei ele ance ega ding a
speci ic s age o in asion p ocess (i.e anspo , in oduc ion, es ablishmen and sp ead)
(Kola and Lodge 2001; Duncan e al. 2003; Blackbu n e al. 2011; Abellán e al. 2017).
Fi s , e en -le el ac o s ega ding he numbe o in oduc ions and indi iduals eleased
ac oss he in asion his o y (i.e p opagule p essu e) (Lockwood e al. 2005). Second,
loca ion-le el ac o s a ibu ed o ecipien en i onmen ai s such as coexis ing na i e
species simila o non-na i e species and clima e ma ching ega ding hei na i e anges
(Fo sy h e al. 2004; Ca dado e al. 2016). Thi d, species-le el ac o s o ad an ageous
ai s ha show non-na i e species such as high abundances ange-wide o hei na u al
dis ibu ion, as demog aphic g ow h a es and beha io al lexibili y (Sol e al. 2005;
Blackbu n e al. 2009; an Kleunen e al. 2010).
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Thesis In oduc ion
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Whe eas se e al bio ic ai s a e conside ed in he unde s anding o biological
in asion success, mechanisms o bio ic in e ac ions a e elega ed o impac s udies on
na i e communi ies, despi e o hei impo ance in shaping species dis ibu ions (Wisz
e al. 2013). Bo h di ec and indi ec bio ic in e ac ions be ween non-na i e species and
ecipien communi ies a e expec ed du ing di e en s ages o in asion p ocess, al hough
i is widely unexplo ed hei in luence on success o es ablishmen and sp ead o non-
na i e species (Ca o d e al. 2009), especially in e eb a es (Blackbu n e al. 2011).
No el in e ac ions be ween non-na i e species and ecipien communi ies
A e hei in oduc ion in no el en i onmen s, non-na i e species may gain o lose
bio ic in e ac ions in ela ion o in e ac ions ha a e p e iously p esen in hei na u al
dis ibu ion anges (Ca o d e al. 2009). These in e ac ions may in luence on non-na i e
and na i e popula ions, so hey a e also keys de e mina ing he success o ailu e o
es ablishmen and in asion p ocesses, bo h h ough he loss (e.g. Enemy Release and
Missed Mu ualisms Hypo heses; (El on 1958; Alpe 2006)) and gain o in e ac ions
(e.g. In asional Mel down and Bio ic Resis ance Hypo heses; (El on 1958; Simbe lo
and Von Holle 1999)). Mo eo e , non-na i e species a e mos ly gene alis (Wol e al.
1996; Cassey e al. 2004; Sol e al. 2005), hence, he eme gence o no el in e ac ions o
eplace gene alis in e ac ions may occu mo e apidly han hose which a e es ic i e
such as specialis (T a ese and Richa dson 2014). As examples o how no el
in e ac ions wo ks bo h ways, on he one hand, ecipien communi y may display bio ic
ba ie s o in asi e species such as a high di e si y o na u al enemies (e.g. p eda o s
and compe i o s) ha p oduce bio ic esis ance agains in asion p ocess (Vilà and
Weine 2004; de Ri e a e al. 2005; Foelke e al. 2018). On he o he hand, coexis ing
in asi e species may de elop bene icial in e ac ions be ween hem, acili a ing hus
hei in asion p ocess as well as hei impac s on na i e communi y h ough in asional
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Thesis In oduc ion
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mel down p ocesses (Adams e al. 2003; Johnson e al. 2009). All hese in e ac ions
usually ca ego ize ega ding hei e ec on each in e ac ing species, namely nega i e,
posi i e and neu al in e ac ions. (Simbe lo and Von Holle 1999). Mos s udies abou
bio ic in e ac ions and associa ed ecological impac s on na i e species ocus on nega i e
ace s such as p eda ion, compe i ion and pa asi ism (Blackbu n e al. 2009; Cla e o e
al. 2009; Ruscoe e al. 2011). Howe e , posi i e in e ac ions, such as acili a ion, may
also de elop be ween na i e and in asi e species and bene i bo h species, al hough ew
s udies ha e es ed hei popula ion e ec s and in asion success (Richa dson e al.
2000; Al ie i 2010; W igh e al. 2016) and only eco ded in plan s and in e eb a es.
Bio ic in e ac ions play a ole be ween di e en ecological hypo heses aimed a
explaining biological in asions al hough a co ec assessmen o impac s on na i e
communi ies may be challenging, chie ly o e looked impac s and long- e m ecological
e ec s (S aye e al. 2006; Simbe lo e al. 2013; Blackbu n e al. 2014). Thus, i is
essen ial he implemen a ion o a holis ic app oach ha displays he complexi y o
ecological in e ac ion ne wo ks and ha includes bio ic in e ac ions om in asi e
species and as well as li e-his o y ai s o in ade s and ecipien communi ies (Ca o d
e al. 2009). Besides, he ole o bio ic in e ac ions may hide due o o he key ac o s i
he ocus is w ong depending on in which s age o in assion p ocess is wo king
(Abellán e al. 2017). The e o e, his holis ic app oach may be undamen al o p edic
impac s on na i e bio a and en i onmen , de eloping hus e ec i e managemen plans
agains biological in asions.
Pa o s, unin en ional cha ac e s in he scena io o biological in asion
A wide a ie y o animal g oups a e emo ed om hei na i e geog aphic anges due o
he in e na ional wildli e ade (Beissinge 2001; Old ield 2003; Young e al. 2016), a
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Thesis In oduc ion
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in e ac ions showed by se e al non-na i e pa o species o in e how limi a ions om
ecipien en i onmen may igge beha io al inno a ions and s a egies o a oid hem,
a o ing hus he in asion p ocess (Sec ion 2). We ocus on animal-plan mu ualism in
which non-na i e pa o s species in e ac wi h ecipien ege al communi y and hei
po en ial e ec s on na i e lo a (Sec ion 3). Finally, we assess he easibili y and
e iciency o a hypo he ical managemen plan ocused on an in asi e pa o species
(Sec ion 4).
This doc o al hesis is cons i u ed by ou sec ions ha desc ibe he amewo k,
jus i ica ion, speci ic hypo heses and p edic ions o main goals. Di e en analy ical
app oaches ha e been used in each sec ion, ma e ial and me hods in his hesis a e
desc ibed in de ail wi hin each sec ion.
In he Sec ion 1, we moni o ed an in asi e popula ion o ose- inged pa akee in
an u ban pa k loca ed in Se ille, Spain, o es po en ial in e speci ic compe i ion o
nes ing si es ( ee ca i ies) be ween he ca i y nes e communi y. Besides, we censuded
se e al monk pa akee popula ions in di e en in aded a eas o assess i s po en ial
acili a ion o nes ing si es o coexis ing ca i y nes e species. We add ess pa icula
ques ions such as (i) whe he ose- inged pa akee exploi s nes ing esou ces
unexploi ed by na i es (oppo unism hypo hesis) o i may sha e same p e e ences o
exploi esou ces ou compe ing agg essi ely wi h he es o ca i y nes e s (compe i ion
hypo hesis) (Sol e al. 2012), (ii) how he dis ibu ion o ose- inged pa akee nes s and
hei in e speci ic agg essions in luence on spa ial nes ing dis ibu ion o es o ca i y
nes e s, (iii) whe he a coexis ing h ea ened ba species, he g ea e noc ule (Alcalde e
al. 2016), is mo e sensi i e o compe i ion p essu e o bo h species do no in e ac ion as
well as wha impac s eme ge on i s popula ion du ing a ime window o 14 yea s
ega ding he g owing ose- inged pa akee popula ion, (i ) whe he monk pa akee also
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Thesis In oduc ion
20
acili a es al e na i e nes ing si es wi h own buil nes s o se e al species in i s in asi e
ange, and ( ) how enan species and hos monk pa akee s bene i o esul an
in e speci ic colonianism. Pa o his wo k ha e been published in he nex jou nals:
PloS One (2014); Hys ix (2014), Royal Socie y Open Science (2018).
In he Sec ion 2, we eco ded se e al adap i e s a egies om se e al
popula ions o h ee di e en non-na i e pa o species (monk pa akee , ose- inged
pa akee , and o ange-winged amazon) o assess hei ole as acili a o s o es ablishmen
and in asion p ocesses o hese pa o species agains unsui able en i onmen al
condi ions. We add ess pa icula ques ions such as (i) whe he he eme gence o
nes ing associa ion be ween monk pa akee s and a na i e species, he whi e s o k,
depends o habi a ea u es ( u al and u ban) and wha bene i s p o ide nes ing
associa ion o bo h species, (ii) how p eda ion p essu e om na i e ap o s
de e mina es he success o es ablishmen and sp ead o monk pa akee (Ca o d e al.
2009), (iii) whe he he sca ci y o a ailable nes ing si es is a ba ie o sp ead o ose-
inged pa akee , (i ) how no el nes ing beha io s om some indi iduals sp ead o he
es popula ion and may assis he in asion p ocess o ose- inged pa akee (Le eb e e
al. 2004), ( ) how an incipien non-na i e popula ion o o ange-winged amazon
o e comes he ma e sho age and o he Allee e ec s (Cou champ e al. 1999), and ( i)
whe he hyb idiza ion be ween non-na i e pa o species may escue popula ions in
ea ly s ages o es ablishmen (Luque e al. 2011). Pa o his wo k has been published
in F on ie s in Zoology (2020).
In he Sec ion 3, we in e he ole o non-na i e pa o species in animal-plan
mu ualisms, in which, pa o s may play a ole as seed dispe se s by di e en
mechanisms. We add ess pa icula ques ions such as (i) which seed dispe sal
mechanisms a e widely p esen be ween di e en pa o species (i.e. epizoocho y and
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Thesis In oduc ion
21
es oma ocho y) (Tella e al. 2015), (ii) whe he pa o s a e p omo e s o seconda y
dispe sal h ough ood was ing when o he animal species bene i o ood acili a ion,
and (iii) how non-na i e pa o species may dis up animal-plan in e ac ions es ablished
be ween na i e bio a and assis non-na i e plan s in in asional mel down p ocesses
(T a ese and Richa dson 2014). Pa o his wo k has been published in Scien i ic
Repo s (2019).
In he Sec ion 4, we assess h ough popula ion iabili y analysis he e iciency
o di e en managemen ac ions acco ding o a hypo he ical e adica ion plan ocused on
an in asi e popula ion o ose- inged pa akee in Se ille, Spain. Gi en ecological
impac s showed in Sec ion 1 and e hic disc epancies ega ding managemen ac ions, i
is u gen ly necessa y a consensus based in scien i ic- echnical e ms. We add ess
pa icula ques ions such as (i) wha demog aphic impac s show di e en managemen
ac ions (i.e. b eeding ailu e o emo ing indi iduals) on ose- inged pa akee
popula ion(ii) whe he popula ion size o in ade de e mina es he e ec i eness o
al e na i e emo ing indi iduals me hods (i.e. apping and shoo ing), and (iii) whe he
e adica ion plans a e mo e easible han con ol plans as well as mo e e ec i e a long-
e m app oach bo h conse a ion and cos -bene i pe spec i es (Simbe lo 2003).
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Thesis In oduc ion
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Sec ion 1: Chap e I
35
ABSTRACT
In asi e species can ake ad an age o esou ces unexploi ed by na i es (oppo unism
hypo hesis) o hey can exploi he same esou ces bu mo e agg essi ely o e icien ly
(compe i ion hypo hesis), hus impac ing na i e species. Howe e , in asi e species end
o exploi an h opogenic habi a s ha a e ine icien ly used by na i es such as u ban
en i onmen s. Focusing on he ing-necked pa akee (Psi acula k ame i), one o he
mos in asi e bi ds wo ldwide, we combined obse a ions o in e speci ic agg essions,
species-speci ic ca i y-nes p e e ences and he spa ial dis ibu ion o he na i e ca i y-
nes ing e eb a e communi y o de e mine he in asion p ocess as well as i s po en ial
impac s on na i e species in a Medi e anean ci y. Ou esul s suppo he compe i ion
hypo hesis, sugges ing ha ing-necked pa akee s a e ou compe ing na i e species
sha ing nes -si e p e e ences. Pa akee s ini ia ed and won mos in e speci ic
agg essions, which we e di ec ed owa ds compe i o s bu also owa ds p eda o s. This
beha iou could explain he spa ial a angemen o na i es, wi h mos bi d species
b eeding close o pa akee s possibly o ake ad an age o hei e ec i e an ip eda o y
beha iou . Howe e , empo al and spa ial pa e ns o seg ega ion sugges ha a
h ea ened ba species is nega i ely a ec ed by pa akee s. This demons a es ha
common species gain bene i s and h ea ened ones (in his s udy, a ba and possibly a
alcon) lose nes si es due o in ade s. The e o e, he conse a ion s a us o he na i e
species ha pay he cos s o compe i ion wi h in ade s should be conside ed. This
scena io o winne s and lose s may, howe e , shi owa ds mo e lose s i he ing-
necked pa akee popula ion con inues o g ow, hus equi ing close moni o ing and
con ol/e adica ion p og ams o a oid u he impac s.
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Sec ion 1: Chap e I
36
INTRODUCTION
Biological in asions a e conside ed a majo h ea o global biodi e si y, since in asi e
species may cause nega i e impac s on na i es h ough inc eased p eda ion isk,
compe i ion, hyb idiza ion o he sp ead o disease (Mack e al. 2000). A smalle
scales, howe e , he ela ionship be ween in asi e species and biodi e si y measu es is
less clea (Rosenzweig 2001), as in oduced species can con ibu e o species gain by i s
es ablishmen (Sax e al. 2002), can educe species ichness h ough ex inc ion
p ocesses (Cla e i e al. 2009) o can ha e no de ec able e ec s on na i e bio a
(Simbe lo 1981). These di e en pa e ns may be explained by he na u e o he
in ade (Sax and Gaines 2008) bu also by he cha ac e is ics o he ecipien
communi y (Ca o d e al. 2009). In an oppo unis ic scena io ( o mally called he
emp y niche, he in asion window o he oppo uni y window hypo heses; Ca o d e al.
2009), in asi e species a e unc ionally di e en om species al eady p esen in he
communi y and hus hei en ance in o a new en i onmen can occu wi hou he
displacemen o ex inc ion o na i es. Con e sely, when exo ic and na i e species
exploi simila esou ces, he ecipien communi y could esis an in asion as a esul o
compe i ion ha s ems om high local di e si y and low niche acancy (MacA hu
1970; Hie o e al. 2005). Howe e , i exo ic species a e able o ou -compe e na i es by
exploi ing esou ces mo e e icien ly o h ough agg essi e beha iou s, hey can
success ully in ade he new a ea causing he displacemen o he na i e compe i o ( an
Wilgenbu g e al. 2010).
U ban en i onmen s ep esen a challenge o biodi e si y, as no all na i e
species inhabi ing he su ounding u al habi a s a e able o colonize hese a eas (Ca e e
and Tella 2011). Di e en s udies ha e ound a educ ion in ichness and di e si y o
na i e species along u ban g adien s, o en in pa allel wi h inc emen s in exo ic in asi e
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Sec ion 1: Chap e I
37
ones (McKinney 2006). Thus, as ci ies expand ac oss he globe, biological
homogeniza ion inc eases as a consequence o he widesp ead inc emen o u ban-
adap able, o en in asi e species a he expense o na i e, o en endemic ones
(McKinney and Lockwood 1999). This pa e n sugges s ha many exo ic and na i e
species may no compe e in na u e (Blackbu n e al. 2009), as he o me end o be
pa icula ly abundan in habi a s ha a e ine icien ly used by na i es, such as in u ban
en i onmen s (Sol e al. 2012). Howe e , ci ies s ill se e as e uges and conse a ion
a eas o some endange ed na i es (Mille and Hobbs 2002), which migh come in o
con lic wi h in asi e species using highly simila esou ces.
Ring-necked pa akee s (Psi acula k ame i) a e na i e o Asia and A ica and
ha e es ablished non-na i e u ban popula ions in a leas 35 coun ies on i e con inen s
(Bu le 2003). Al hough i is conside ed amongs he 100 wo s alien species in Eu ope
(h p://www.eu ope-aliens.o g/speciesTheWo s .do), i s impac on na i e species
emains unclea . The ing-necked pa akee equi es medium-size (4–8 cm en ance size,
S ubbe and Ma hysen 2009) na u al ca i ies o hose exca a ed in ees by o he
species o b eeding. Gi en he usual sho age o ee ca i ies (New on 1998), especially
in u ban en i onmen s whe e decaying ee limbs a e pe iodically emo ed in he
in e es s o public sa e y (Da is e al. 2013), pa akee s could ou compe e na i e ca i y-
nes ing species in agg essi e in e ac ions and hus sp ead a he cos o he numbe s
and/o dis ibu ion o na i es (compe i ion hypo hesis). Al e na i ely, i he na i e
communi y is poo in seconda y ca i y nes e s and/o he esou ce is no limi ed, he
es ablishmen o his in asi e species could be acili a ed by a high a ailabili y o
nes ing si es (oppo unism hypo hesis). P e ious wo k has shown ha pa akee s can
ou compe e only one o he coexis ing na i e ca i y-nes ing bi d species in a cen al-
Eu opean ci y (S ubbe and Ma hysen 2007, 2009), bu la ge -scale s udies compa ing
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Sec ion 1: Chap e I
38
a eas occupied and no occupied by his in ade sugges li le o no impac on
popula ions o na i e bi ds (S ubbe e al. 2010; Newson e al. 2011).
The ou come o in asi e-na i e compe i ion could be con ex -dependen , being
in luenced by he a ailabili y o esou ces and he composi ion o he na i e communi y
(Ca o d e al. 2009). Thus, answe ing simila ques ions bu using di e en sys ems can
help o make gene aliza ions abou p ocesses om local pa e ns. He e, we combined
obse a ions o in e speci ic agg essi e in e ac ions, species-speci ic ca i y-nes
p e e ences and he spa ial dis ibu ion o ca i ies a ailable and used by each species o
in e he p ocess as well as he consequences o ing-necked pa akee in asions in a
Medi e anean ci y. Resul s show a complex scena io whe e, al hough ing-necked
pa akee s ou compe e na i e species in agg essi e encoun e s, mos na i es seem o
bene i om he e ec i e an i-p eda o beha iou o pa akee s. Con e sely, some
h ea ened na i e species can be displaced by pa akee s, esul ing in a dynamic p ocess
o winne s and lose s linked o he popula ion g ow h o he in ade .
MATERIAL AND METHODS
E hics S a emen
Field wo k conduc ed he e was no in asi e and did no equi e he manipula ion o li e
animals. The e o e, his wo k did no equi e speci ic pe mi s by he ele an Spanish
au ho i ies.
S udy Sys em
The ing-necked pa akee was a commonly aded wild species o he Spanish cage-
bi d ma ke (Ca e e and Tella 2008a) and a numbe o u ban popula ions a ose la gely
om acciden al escapes om cages (Ma í and del Mo al 2003). This s udy ocuses on
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Sec ion 1: Chap e I
39
he ci y o Se ille (sou he n Spain), whe e he i s eco ds o he species da e back o
he ea ly 1990’s and he ini ially small popula ion sha ply inc eased (Au ho s’
unpublished da a), eaching ca. 1,000 indi iduals in 2011 (P. Edelaa com. pe s.). We
conduc ed he i s b eeding census o he species in he ci y o Se ille om Ma ch o
July 2013. We i s loca ed po en ial b eeding a eas whe e he species was p esen
aking ad an age o i s conspicuous beha iou . Then, we moni o ed he a ailable
ca i ies o assess whe he o no hey we e occupied by pa akee s based on he
obse a ion o adul s en e ing a minimum o 10 imes on di e en days, he
ocaliza ions o chicks inside he nes , and/o he obse a ion o ju eniles a he
en ance. We loca ed 216 ac i e nes s, 159 (73.6%) in an u ban pa k ( he Ma ía Luisa
Pa k; 37° 22′ 31.57" N, 5° 59′ 19.59" W) and he es o ming smalle b eeding nuclei in
sca e ed g oups o ees o , mo e a ely, in buildings h oughou he ci y. Ma ía Luisa is
he la ges pa k loca ed in he co e o he ci y, comp ising a 40 ha wooded a ea wi h a
a ie y o ee species, mos o hem exo ics such as Pla anus sp., Eucalyp us sp. o
Gledi sia iacan hos. The pa k is comple ely su ounded by s ee s wi h mode a e o
high a ic in ensi y.
A ailabili y and Occupancy o T ee Ca i ies
The assessmen o he a ailabili y o ee ca i ies and hei occupancy by pa akee s and
na i e species was es ic ed o Ma ía Luisa Pa k o a oid po en ial biases when
analysing in e speci ic compe i ion (e.g., small g oups o ees ou side o he pa k
occupied by pa akee s could no be occupied by some na i e species because hey did
no o e su icien o aging habi a ). We GPS loca ed (±3 m) all ee ca i ies ha we
we e able o isually inspec in ees loca ed wi hin he pa k by using 10×50 binocula s.
In each case, we iden i ied he ee species and es ima ed he heigh o he ca i y abo e
g ound (in m) and he wid h o i s en ance (in cm). The en ances o ca i ies we e
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Sec ion 1: Chap e I
40
ca ego ized as small, medium o la ge (<4 cm, 4–8 cm, and >8 cm, espec i ely)
acco ding o p e ious s udies, which showed he p e e ence o pa akee s o ca i ies
wi h en ances be ween 4 and 8 cm wid h (S ubbe and Ma hysen 2007; Czajka e al.
2011). Ca i ies loca ed a <2.5 m abo e g ound we e no conside ed o analyses since
hei accessibili y o humans would p eclude i s use (none we e occupied by na i e o
exo ic species), hus biasing esul s. F om Janua y o Augus (co e ing he en i e
b eeding season o na i e and exo ic species), we epea edly isi ed and obse ed a a
dis ance ( o a minimum o 10 min) each ca i y on a leas 10 di e en days du ing
dayligh hou s o assess whe he o no i was occupied and by wha species, de o ing
48 days (202.5 hou s) o ield wo k. The close p oximi y o many ees wi h ca i ies
o en allowed us o moni o se e al ees simul aneously. A ca i y was conside ed as
occupied by a gi en bi d species when we obse ed adul s en e ing a minimum o 10
imes on di e en days, hea d chicks inside, o obse ed ju eniles a he en ance. In
addi ion, Ma ía Luisa Pa k is also inhabi ed by he g ea e noc ule (Nyc alus
lasiop e us). This ca i y-b eeding o es species is he la ges Eu opean ba (a e aging
48 g; Ibáñez e al. 2001), and he whole popula ion li ing in and a ound Se ille ( oughly
es ima ed a ca. 500 indi iduals in 2003–2004) ga he s o b eed and oos communally
in he ee ca i ies o his pa k (Popa-Lisseanu e al. 2008; Ibáñez e al. 2009). To
iden i y he ca i ies used by g ea e noc ules, we de ec ed hei p esence using an
ul asound de ec o (Pe e sson D 230) and obse ing ba s lea ing ee ca i ies a sunse .
G ea e noc ules, like o he o es ba s, o m ission- usion socie ies ha swi ch oos s
e e y ew days, so each ba colony can con ol a la ge numbe o oos s o which only a
ew a e occupied a a speci ic ime (Popa-Lisseanu e al. 2008). Thus, using p e ious
in o ma ion on adio- acked indi iduals (Popa-Lisseanu e al. 2008), we conside ed
ha a ee was no used by g ea e noc ules du ing sp ing-summe 2013 i we did no
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Sec ion 1: Chap e I
41
obse e ac i i y du ing any o ou 10 spaced isi s. Using in o ma ion on ees used by
noc ules du ing 2003–2004, we also es ed o changes in hei use in ela ion o he
cu en nes ing spa ial dis ibu ion o ing-necked pa akee s. These ees we e loca ed
a e moni o ing 27 noc ules h ough adio- acking o s udy he spa ial pa e n o ee
use by he species, inding ha ca i ies loca ed in 75 ees we e al e na i ely used as
oos si es en yea s ago (Popa-Lisseanu e al. 2008). I is wo h no ing ha he di e en
me hodologies used o iden i y occupied ees could p oduce alse cases o inoccupa ion
in bo h 2003–2004 (a la ge pe iod o ime moni o ing 27 indi iduals) and 2013 (a
sho e pe iod o ime moni o ing all ee ca i ies). Howe e , i may also jus p oduce
s a is ical noise making ou es ima es conse a i e.
Spa ial Dis ibu ion o Occupied Ca i ies
The occupancy o a pa icula ee ca i y by a gi en species could be in luenced by he
spa ial dis ibu ion o ca i ies occupied by he same and/o o he species, d i en no
only by compe i ion bu also by conspeci ic and he e ospeci ic a ac ion p ocesses. We
hus ob ained he dis ance om each occupied ca i y o he nea es ca i y occupied by
conspeci ics and he e ospeci ics (nea es neighbou dis ance) as well as he
co esponding nes agg ega ion indexes. These agg ega ion indexes we e ob ained as
he ela i e posi ion o each occupied ca i y wi hin he whole dis ibu ion o all ca i ies
occupied by conspeci ics o he e ospeci ics in he pa k using ∑ exp (-dij), wi h (i≠j)
whe e dij is he linea dis ance be ween occupied ca i ies i and j, j ep esen ing all
occupied ca i ies (Moilanen and Hanski 1998). These a iables we e complemen a y
measu es depic ing he social en i onmen a ound each nes ca i y a a landscape scale
as well as he exis ence o close compe i o s.
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Sec ion 1: Chap e I
42
The spa ial dis ibu ion o occupied ca i ies could also be in luenced by habi a
he e ogenei y in he pa k. We conside ed he wo main sou ces o habi a he e ogenei y
in ou s udy a ea, i.e. he p oximi y o su ounding s ee s and o es co e . Noise om
ca a ic could al e song pe o mance, ep oduc i e success and e en he spa ial
dis ibu ion o bi ds (Slabbekoo n and Pee 2003; Hal we k e al. 2011). We he e o e
measu ed he linea dis ance om each ca i y o he closes s ee using GIS ools (see
below). On he o he hand, species could di e in hei p e e ences o o es co e age
a ound ca i ies. We ob ained o es co e by measu ing i in a adius o 30 m a ound
each GPS loca ed ca i y on a Google Map image aken in 2013 (Imagens ©2013
Cnes/Spo Image, Digi alGlobe, Ins i u o de Ca og a ía de Andalucía, map da a ©2013
Google, based on BCN IGN Spain), using OpenLaye s Plugin (1.1.0) applica ions in Q-
GIS 1.8.0 (2008 F ee So wa e Founda ion, Inc). Fo es co e was hen sco ed in o ou
main ca ego ies, namely 0–25%, 25–50%, 50–75%, and 75–100%.
In e speci ic In e ac ions
We assessed in e speci ic in e ac ions by andomly sampling he beha iou o di e en
nes ing ing-necked pa akee s du ing a 15-minu e pe iod. We conse a i ely eco ded
he bi d species p esen wi hin a adius o 15 m a ound he ocal pa akee , whe he o
no he e was an agg essi e in e ac ion, wha species s a ed he a ack, and which was
he winne . To inc ease sample size wi hou esampling he same indi iduals o he
numbe o po en ial in e ac ing species, obse a ions we e conduc ed in Ma ía Luisa
Pa k as well as in o he u ban a eas o Se ille occupied by he species (see abo e),
o aling 88 days (351.5 hou s) o ield wo k. These a eas included he main pa ks o he
ci y as well as a chu ch (Di ino Sal ado ) whe e ing-necked pa akee s occupied
ca i ies in walls o b eeding, po en ially compe ing he e wi h lesse kes els (Falco
naumanni), a colonial alcon ha usually b eeds in u ban buildings (Tella e al. 1996).
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Sec ion 1: Chap e I
43
S a is ical Analysis
We employed Gene alized Linea Models (GLM) implemen ed h ough he GENMOD
p ocedu e in SAS 9.2 (SAS Ins i u e 2008) o asce ain which a iables de e mined hole
occupa ion, using he binomial e o dis ibu ion (ca i y occupied o no occupied by a
gi en species) and he logis ic link unc ion. In a i s se o models, we aimed o
de e mine whe he occupied and acan ca i ies di e ed in hei s uc u al
cha ac e is ics. Thus, we modelled he p obabili y o occupancy as a unc ion o he
heigh o he ca i y abo e g ound (in i s linea and quad a ic o ms), he en ance size,
and he ee species. The esul ing species-speci ic pa e ns o ca i y p e e ences (see
esul s) made i di icul o iden i y simila i ies (and hus oppo uni ies o compe i ion)
be ween species. The e o e, we pe o med a ca ego ical p incipal componen analysis
(CATPCA) on en ance size (since i is a ca ego ical a iable) and heigh abo e g ound
o he ca i ies occupied and ook he sco es o he ob ained i s dimension as a single
compiling desc ip o o he ca i ies used by each species. An ANOVA on hese sco es
allowed us o iden i y di e ences in ca i y p e e ences among species, and pos -hoc
Sche e es s pe mi ed us o es ablish homogeneous g oups (i.e., species no di e ing in
hei p e e ences o pa icula ca i y ai s). In a second se o models, we assessed he
spa ial a angemen o each species ega ding ca i y ai s, he dis ibu ion o bo h
conspeci ics and he e ospeci ics, and main habi a ea u es (dis ance o he nea es s ee
and o es co e ) a ound each occupied and a ailable (i.e., unoccupied) ca i y, also
using GLMs wi h a binomial e o dis ibu ion and logis ic link unc ion.
Exac binomial es s we e used o assess whe he he p opo ion o in e speci ic
encoun e s ending in agg essions, he p opo ion o agg essions ini ia ed by ing-necked
pa akee s, and he p opo ion o agg essions won by his species di e ed signi ican ly
om pa i y. To ob ain in e speci ic pa e ns in he equency o agg essions and hei
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Sec ion 1: Chap e I
44
ou pu s, we also used GLMs wi h a binomial e o dis ibu ion and a logis ic link
unc ion, i ing as explana o y a iables he a e age body mass o he species
in e ac ing wi h ing-necked pa akee s (ob ained om Dunning 1993), hei o e lap in
nes -si e p e e ences (as a ac o wi h le els anging om 0– he in e ac ing species was
no a ca i y-nes e - o 3– maximum o e lap in nes -si e ai s), and whe he he
in e ac ing species was a po en ial p eda o o eggs, nes lings o adul s. We expec ed
ha ing-necked pa akee s would be mo e p one o a ack hose species wi h
o e lapping nes -si e p e e ences and po en ial p eda o s, and less p one o a ack la ge -
bodied species.
A backwa d p ocedu e was pe o med o GLM modelling, emo ing om ull
models hose a iables ha we e non-signi ican ly associa ed wi h he esponse a iable
(p>0.05) o ob ain minimum adequa e models (MAM) (Sol e al. 2012). The esul ing
models did no show da a o e dispe sion. We calcula ed he pe cen age o de iance
explained as a measu e o he a iance explained by each MAM.
RESULTS
Occupancy o T ee Ca i ies
We eco ded 1,086 ca i ies in 435 ees loca ed wi hin Ma ía Luisa Pa k du ing he
2013 b eeding season. Ca i ies we e loca ed a an a e age heigh abo e g ound o 13.09
(SD 5.51) m, and he commones ca i y en ances (47%) we e o in e media e size (4–8
cm). Mos ca i ies (62.2%) we e loca ed in London plane ees (Pla anus × ace i olia)
p obably because i is he mos abundan species wi hin he pa k bu is also he species
wi h highes numbe o a ailable ca i ies.
A o al o 10 species we e ound occupying 525 ca i ies (Table 1), including 9
bi d and one ba species. Two bi d species we e exo ics, i.e. he ing-necked pa akee
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Sec ion 1: Chap e I
51
encoun e s (Table 5), he p obabili y ha an encoun e ended in agg ession inc eased
wi h he in e speci ic o e lap in nes ype p e e ences and he body mass o he
in e ac ing species, and dec eased wi h he numbe o indi iduals o he in e ac ing
species in ol ed. The p obabili y ha an agg ession was ini ia ed by ing-necked
pa akee s dec eased wi h he body mass o he in e ac ing species. Finally, he
p obabili y ha a igh was won by ing-necked pa akee s was g ea e i hey ini ia ed
he a ack bu dec eased when he in e ac ing species was a po en ial a ian p eda o .
None heless, ing-necked pa akee s won 25–100% o he agg essions di ec ed owa ds
di e en p eda o species (Figu e 2).
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Sec ion 1: Chap e I
52
Figu e 2. Pe cen age o encoun e s wi h ing-necked pa akee s ha ended in
agg essions (whi e ba s), and pe cen age o agg essions ini ia ed (g ey ba s) and won by
ing-necked pa akee s (black ba s). The numbe o eco ded encoun e s is shown in
b acke s.
Table 5. GLMs explaining he p obabili y ha an in e speci ic encoun e ended in
agg ession (Agg ession), whe he he agg ession was ini ia ed by ing-necked pa akee s
(Figh ini ia ion) and was won by ing-necked pa akee s (Win igh ).
Nes ing o e lap N Body mass A ack ini ia ion P eda o % de iance
Agg ession 78.83 (+)*** 17.13 (-)*** 27.24 (-)*** 19.32
Figh ini ia ion 21.89 (-)*** 9.58
Win igh 24.7 (+)*** 27.89 (-)*** 37.02
The e ained explana o y a iables we e he in e speci ic o e lap in nes ypes (Nes ing o e lap), he
numbe o indi iduals o he in e ac ing species p esen in he encoun e s (N), he a e age body mass o
he in e ac ing species (Body mass), whe he o no ing-necked pa akee s ini ia ed he agg ession (A ack
ini ia ion), and whe he o no he in e ac ing species is a po en ial p eda o o bi ds. χ
2
alues a e gi en
o each a iable. Signs be ween b acke s indica e posi i e o nega i e e ec s o he explana o y
a iables. ***: p<0.001; % de : pe cen age o de iance explained.
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Sec ion 1: Chap e I
53
DISCUSSION
Oppo unism o Compe i ion?
The success ul es ablishmen o exo ic species in no el habi a s cons i u es a poo ly
unde s ood pa adox (Sax and B own 2000). Recen ly, Sol e al. (2012) examined he
in asion pa adox by s udying he use o ood esou ces by in asi e and na i e bi d
species in an Aus alian ci y, concluding ha he success o in ade s is explained by
hei capaci y o exploi ecological oppo uni ies ha mos na i e species a ely use.
Howe e , as he au ho s poin ed ou , compe i ion o e o he esou ces, no ably nes ing
si es, mus be conside ed in u he s udies (Sol e al. 2012).
We in es iga ed wo key aspec s behind he es ablishmen success o ing-
necked pa akee s on a ela i ely newly in aded u ban a ea, namely: he way hey sha ed
nes -si e esou ces wi h he ecipien communi y and he agg essi e in e ac ions hey
expe ienced wi h o he species. This app oach allowed us o show ha his species may
in ade new a eas e en when esou ces a e no o e abundan , hus no suppo ing he
hypo heses p oposing ha sa u a ed communi ies can hal biological in asions h ough
compe i i e p ocesses bu a he suppo ing he compe i ion hypo hesis (ins ead o he
oppo unism hypo hesis) o success ul in asions (Ca o d e al. 2009). Al hough he
a ailabili y o ee ca i ies was ela i ely high in he s udy a ea compa ed o o he ci ies
(S ubbe and Ma hysen 2009; O chan e al. 2013), he la ge popula ions sizes o
di e en ca i y-nes e species oge he wi h he ac ha he cha ac e is ics o
unoccupied ca i ies di e ed om hose o occupied ones sugges a sho age o sui able
b eeding si es o he na i e ca i y-nes ing communi y, coinciden wi h he gene al
pa e n o compe i ion ound ac oss ca i y-nes ing communi ies especially in u ban
en i onmen s (New on 1994; S ubbe and Ma hysen 2009; O chan e al. 2013). Mos o
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Sec ion 1: Chap e I
54
he inabili y o he na i e communi y o esis he pa akee in asion may be due o he
in ade ’s highly agg essi e beha iou ha allows i o ou -compe e na i es, hus
success ully occupying a eas e en when he e is no supe abundan o unde exploi ed
esou ces. In e es ingly, we we e able o sepa a e species in o h ee main unc ional
g oups based on species-speci ic nes si e equi emen s, showing ha pa akee s i in o
one o hese g oups. Thus, e en when hey may be in e ac ing wi h many na i e species,
hey sha e impo an esou ces o popula ion p ospec s (i.e., ep oduc ion) wi h only
some o hem. Howe e , ing-necked pa akee s we e agg essi e (and won mos
agg essi e encoun e s) no only owa ds hose species sha ing nes -si e p e e ences
(including wo o he exo ic pa o species) bu also owa ds o he s, e en non ca i y-
nes ing species and a ian p eda o s. The e o e, he ing-necked pa akee has he
po en ial o modi y he numbe s and spa ial dis ibu ion o coexis ing b eeding species
h ough beha iou -media ed compe i i e exclusion.
Mechanisms behind he Spa ial A angemen o Species
Habi a selec ion models, and species dis ibu ion models in hei b oade sense, a e
ma hema ical desc ip ions o biological pa e ns ha a e a ec ed by en i onmen al
condi ions and a mul i ude o di ec and indi ec in e ac ions (Diamond 1975), hus
in e ing ha causal links om obse a ional da a should be made wi h cau ion. Two
species may co-occu i hey sha e hei habi a equi emen s, bu also i hey acili a e
each o he di ec ly o indi ec ly. Con e sely, species may appea o a oid each o he i
hey show compe i i e exclusion bu also i hey ha e dissimila habi a equi emen s.
Al hough compe i ion o ca i ies can igge in aspeci ic nega i e in e ac ions among
indi iduals, we de ec ed a gene al endency among species o b eed ollowing a pa e n
o conspeci ic agg ega ion. This seems no o esul om he e ogenei ies in habi a and
nes si e a ailabili y, aspec s which we e con olled o in s a is ical analyses, and hus
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Sec ion 1: Chap e I
55
may a he be ela ed o conspeci ic a ac ion p ocesses as p e iously obse ed in many
o he colonial bu also e i o ial species (e.g. S amps 1988; Se ano e al. 2004).
Di e en s udies ha e shown ha b eeding in close p oximi y o conspeci ics bene i s
b eede s om ea lie de ec ion o p eda o s, g oup de ence, and dilu ion o p eda ion
(B own and B own 2001; K ause and Rux on 2002; Se ano e al. 2005).
Rega ding he e ec s o he in asi e species, we ound ha he spa ial
dis ibu ion o nes ing ing-necked pa akee s was impo an o explain he dis ibu ion
pa e ns o all ee-ca i y nes e species o he ecipien communi y while con olling o
he main habi a ea u es. Howe e , he unde lying pu a i e mechanisms (a ac ion o
seg ega ion) we e di e en among species. All bi d species inc eased hei likelihood o
occupying ca i ies loca ed close o pa akee nes s and/o o high densi ies o pa akee s.
Posi i e co-occu ence pa e ns a e indica i e o he e ospeci ic a ac ion (Sebas ián-
González e al. 2010), hus signaling he p esence o di ec o indi ec species
in e ac ions. In ou s udy sys em, a possible explana ion o his associa ion pa e n
could be ound in he high agg essi eness o pa akee s agains a ian p eda o s. In ac ,
ing-necked pa akee s may e en communally a ack p eda o s, as we obse ed a lock
o 60 pa akee s mobbing a boo ed eagle (Aquila penna a) in Ma ía Luisa Pa k in 2008.
The e o e, na i e species may choose b eeding si es a enough om ing-necked
pa akee s (>15 m) o a oid agg essions bu close enough o be ewa ded by hei
e ec i e an i-p eda o esponse, esul ing in an ac i e b eeding associa ion, which
bene i s he associa ed species (Blanco and Tella 1997). Con e sely, he mu ual spa ial
seg ega ion be ween pa akee s and noc ules, no explained by habi a ea u es, could be
indica i e o di ec compe i ion since hey sha e hei p e e ences o he same kind o
ca i ies. The noc u nal beha iou o ba s p ecluded he sys ema ic obse a ion o
encoun e s wi h ing-necked pa akee s, which would ha e been es ic ed o ins ances
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Sec ion 1: Chap e I
56
when pa akee s would en e ba ca i ies and inspec o po en ial nes si es. Al hough
g ea e noc ules ae ially hun small passe ines when mig a ing a nigh (Ibáñez e al.
2001), hey a e no able o kill bi ds inside hei nes s (J. Jus e com. pe s.) and e en less
so a much la ge species such as he ing-necked pa akee whose body mass (116 g) is
mo e han wice ha o he noc ule (50 g). Gi en ha pa akee s won mos agg essions
when encoun e ing la ge -bodied compe i o s such as e al pigeons and e en powe ul
jackdaws (Figu e 2), hey would be expec ed o also win mos agg essi e in e ac ions
wi h his much smalle ba species. Al hough li le is known abou he e ec s o
agg essi e species like pa akee s on mammals ha shel e and ep oduce in hollows like
ba s, se e al au ho s sugges ed ha hey can e ic hem (Gebha d 1996; S a 1998) and
he e is conce n ha ing-necked pa akee s could cause he loss o sui able ca i ies o
he noc ule ba (Nyc alus noc ula) in The Ne he lands (Haa sma and an de G aa
2009). In ou s udy a ea, a g ea e noc ule was o ui ously obse ed being agg essi ely
expulsed om i s ca i y by a ing-necked pa akee in Ma ía Luisa Pa k in 2005 (E.
Re illa com. pe s.), and i could be expec ed ha he s ong beak o pa akee s could
se iously inju e noc ules o he poin o killing o impeding hei ligh by i e e sibly
damaging hei sensible pa agium (J. Jus e com pe s.). Mo eo e , he e is a published
obse a ion o a simila body-sized exo ic pa akee (Supe b Pa o , Poly elis swainsonii)
killing he much la ge ed squi el (Sciu us ulga is, 295 g; Palomo e al. 2007) in I aly
(Mo i e al. 2013) and e idence o simila cases ha migh had been caused by ing-
necked pa akee s in F ance (Japio 2005; Cle geau and Ve gnes 2009). Du ing his
s udy we obse ed 11 ins ances o ing-necked pa akee s (in ol ing up o 10
indi iduals) a acking and mobbing black a s (Ra us a us 180 g; Palomo e al. 2007),
o cing hem om he p oximi y o hei nes s. Since bo h a s and squi els a e
p eda o s o bi d nes s, including hose o pa akee s (Mo i e al. 2013), hese
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Sec ion 1: Chap e I
57
obse a ions also ein o ce he po en ial bene i s o o he bi d species o b eeding close
o pa akee s.
Impac : Winne s and Lose s in a Con empo a y In asion P ocess
I is di icul o ully asce ain he ecological impac s o in ade s, gi en he a ie y o
po en ial impac s o be assessed, hei sub le bu pe asi e e ec s, and he long ime
gaps be ween he in oduc ion o an exo ic species and i s achie emen o in asi eness
and de ec able impac s (S aye e al. 2006; Simbe lo e al. 2013). Al hough he e a e
well- ecognized cases o nega i e impac s o bi d in asions in island en i onmen s
(Le e 2005; F eed and Cann 2009), hei impac on mainland en i onmen s ha e been
less s udied and much deba ed (Kumschick and Nen wig 2010; Baue and Woog 2011;
S ubbe e al. 2011), o he poin o sugges ing ha in oduced bi d species should be
managed be o e hei nega i e impac s a e p o en (Edelaa and Tella 2012).
Al hough ing-necked pa akee s ha e been shown o ou compe e a small ca i y-
nes ing na i e bi d species in B ussels (S ubbe and Ma hysen 2009), he e is li le
e idence o i s impac on na i e communi ies when compa ing a eas occupied o
unoccupied by his in ade (S ubbe e al. 2010; Newson e al. 2011). Ou di e en
app oach, by eco ding he ou pu o in e -speci ic agg essions and he spa ial
dis ibu ion o species in a Medi e anean ci y, sugges s, howe e , ha ing-necked
pa akee s may igge s ong e ec s on he na i e ecipien communi ies, wi h bo h
posi i e and nega i e esponses depending on he na i e species conside ed.
The spa ial seg ega ion o g ea e noc ules and ing-necked pa akee s oge he
wi h he spa ial pa e ns o ees abandoned du ing he las decade by noc ules, i s
appa en popula ion dec ease and he pa allel inc ease in he ing-necked pa akee
popula ion (Au ho s unpubl. da a) sugges an ac i e displacemen exe ed by he
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Sec ion 1: Chap e I
58
in asi e species. This is a ma e o conce n o his ba species, which shows a
sca e ed dis ibu ion h oughou Eu ope and is classi ied as Vulne able in Spain, wi h
Ma ía Luisa Pa k suppo ing i s la ges known colony (Ibáñez e al. 2004). P e ious
adio- acking s udies showed ha g ea e noc ules o age o e la ge ex ensions o
na u al habi a s (up o 40 km om he u ban pa k (Popa-Lisseanu e al. 2009), including
Doñana Na ional Pa k and su ounding ma shlands, bu hey e u n daily o oos ing o
Ma ia Luisa Pa k and no al e na i e e uges a e known o his popula ion (Popa-
Lisseanu e al. 2008; 2009). This la ge popula ion o g ea e noc ules is he e o e highly
sensi i e o any educ ion in he a ailabili y o ee ca i ies caused by ing-necked
pa akee s. Gi en he sca ci y o ma u e o es s wi h la ge numbe s o adequa e ca i ies
o he species, he o he –al hough smalle - colony o g ea e noc ules known in Sou h
Spain is also loca ed in an u ban pa k (in Je ez de la F on e a, 77 km a om Se ille)
(Ibáñez e al. 2001). Al hough he p esence o ing-necked pa akee s is s ill anecdo ic in
his ci y, i s popula ion expansion migh also pose h ea s o his ba popula ion in he
nea u u e. Fu he s udies a e needed o deep on he popula ion ecology and ends o
g ea e noc ule popula ions, and o o he ba species (Haa sma and an de G aa 2009),
ela ed o hei coexis ence wi h in asi e pa akee s.
Ano he cause o conce n is he ac ha pa akee s began o use wall ca i ies in
2011, b eeding in h ee buildings in 2013, one o hem loca ed in he co e o he ci y
whe e he e is also a colony o lesse kes els. This colonial alcon su e ed a d as ic
decline in Eu ope due o land-use changes ha did no e e un il ecen yea s hanks o
widesp ead conse a ion ac ions, including he p o isioning o nes ca i ies (Ca y e al.
2013). Lesse kes els b eeding in Se ille ha e o o age a om hei b eeding colony
(Tella e al. 1998) bu gain bene i s by he educed p eda ion isk in he ci y (Tella e al.
1996). Al hough hei b eeding success was linked o he quali y o wall ca i ies (Neg o
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Sec ion 1: Chap e I
59
and Hi aldo 1993), he species was no cons ained by nes -si e a ailabili y o
compe i ion wi h e al pigeons and jackdaws in ecen decades (Fo e o e al. 1996).
Howe e , he newly es ablished ing-necked pa akee ough mo e han expec ed wi h
lesse kes els and won mo e han hal o he agg essi e encoun e s (Figu e 2), while
occupying only six wall ca i ies wi hin he lesse kes el colony. I he pa akee
popula ion con inues o g ow exponen ially, i may pose a se ious p oblem o u ban
lesse kes els as well. In con as o noc ules and lesse kes els, which a e o ced o
o age a om he ci y, he abundance o ood esou ces o ing-necked pa akee s in
he u ban pa ks could educe he ene gy hey expend, allowing an inc ease in hei
b eeding success and popula ion g ow h (S ubbe and Ma hysen 2011), hus ein o cing
hei compe i i e supe io i y.
Bo h winne and lose species may esul om an h opogenic-d i en expansions
o species (Ca e e e al. 2010), some in ade s e en a o ing whole communi ies o
na i es (Tablado e al. 2010). Ou esul s sugges ha he p esence o nes ing ing-
necked pa akee s may bene i se e al non- h ea ened na i e bi d species, which may
incu b eeding ad an ages by exploi ing hei e ec i e an i-p eda o y beha io .
Howe e , his si ua ion could change in he nea u u e i he ing-necked pa akee
popula ion con inues o g ow. This is al eady he mos abundan species b eeding in he
pa k and is he only one able o enla ge ee ca i ies up o eaching i s p e e ed size (4–
8 cm; S ubbe and Ma hysen 2009; Czajka e al. 2011; his s udy), as has been shown
in o he u ban pa ks (O chan e al. 2013). In ac , 7 ou o he 28 small-sized ca i ies
(en ance <4 cm) we e enla ged and occupied by pa akee s du ing his s udy. The e o e,
nes si es may become limi ed e en o species using small-sized ca i ies such as i s
and house spa ows. The la e species is a widesp ead commensal whose Eu opean
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Sec ion 1: Chap e I
60
popula ions a e now dec easing, hus d awing a en ion o i s long- e m conse a ion
s a us (Robinson e al. 2005; Shaw e al. 2008).
Conse a ion Implica ions
We ha e shown po en ially se ious impac s o an in asi e bi d a ge ing species ha a e
no easily moni o ed o ha a e no expec ed o in e ac wi h hem, such as a o es ba
and a colonial alcon nes ing in buildings, hus highligh ing he di icul ies in assessing
he en i e se o impac s posed by in ade s (Simbe lo e al. 2013). The po en ial
impac o ing-necked pa akee s (S ubbe and Ma hysen 2009), as well as o o he
pa akee species (Da is e al. 2013) h i ing in u ban habi a s, has been o en discoun ed
since u ban bi d communi ies a e usually composed by ew, gene alis and non-
h ea ened species (McKinney and Lockwood 1999; Sol e al. 2012). Howe e , ou case
s udy sugges s ha u ban ing-necked pa akee s may be nega i ely a ec ing wo
h ea ened species, wi h some common species p obably also a ec ed in he nea u u e
i he pa akee popula ion con inues o g ow. The e o e, bo h he conse a ion s a us o
he na i e species wi h which he in ade in e ac s as well as he popula ion ends o
he in ade should be conside ed. Mo eo e , he posi i e popula ion ends o ing-
necked pa akee s in Spain (au ho s' unpubl. da a) sugges s, as o monk pa akee s
(Muñoz and Real 2006), ha he species could sp ead and in ade u al habi a s, as is
al eady he case in cen al Spain (au ho s' unpubl. da a). In such cases, pa akee s would
in e ac wi h a wide communi y o non-u ban species and new impac s could a ise, hus
equi ing a close moni o ing o in e -speci ic in e ac ions.
As ecommended o o he in asi e o ganisms (Simbe lo e al. 2013),
managemen o a ian in asions should be unde aken be o e popula ions sp ead and
ac ions become cos ly and e en una o dable (Edelaa and Tella 2012). In his ega d,
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Sec ion 1: Chap e II
68
Chap e II
Alien ose- inged pa akee s (Psi acula k ame i) a ack
black a s (Ra us a us) some imes esul ing in dea h
Dailos He nández-B i o, Ál a o Luna, Ma ina Ca e e, José L. Tella
Hys ix I . J. Mamm,. 2014, 25(2):121–123
h ps://doi.o g/10.4404/hys ix-25.2-10992
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Sec ion 1: Chap e II
69
ABSTRACT
The ose- ing pa akee (Psi acula k ame i) is one o he mos success ul in asi e bi ds
in i s es ablishmen wo ldwide. S udies add essing i s po en ial impac on na i e bio a
mos ly ocus on bi ds and li le is known abou how hese and o he pa akee species
in e ac wi h na i e mammals. He e, we epo 21 agg essions o ose- inged pa akee s
owa ds black a s (Ra us a us) in u ban pa ks in Se ille (Sou he n Spain) and
Tene i e (Cana y Islands). Ei he soli a y pa akee s o , mo e o en, g oups o up o 18
a acked a s when hey climbed ees close o pa akee nes s. Mos a acks ended when
he a s descended o he g ound. Howe e , in wo ins ances (9.5% o he agg essions)
he a acks esul ed in he dea h o he a s as a esul o alling o he pa emen . These
obse a ions add u he complexi y o a biological in asion, whe e in oduced
pa akee s ha e nega i e impac s on a p eda o and hus, some na i e bi d species may
bene i om hei an ip eda o beha io . Mo e a en ion should be paid o he
in e ac ions be ween na i e mammals and he non-na i e pa akee s in oduced
wo ldwide.
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Sec ion 1: Chap e II
70
MAIN TEXT
Biological in asions a e conside ed among he main causes o biodi e si y loss
(Cla e o and Ga cía-Be hou, 2005). Since he mid wen ie h cen u y, he in oduc ion
o exo ic species has accele a ed a an ala ming a e (Hulme, 2009), mainly as a
consequence o in e na ional ade (Meye son and Mooney, 2007). Wild bi ds a e
among he mos commonly aded e eb a e axa (Ca e e and Tella, 2008a), wi h a
numbe o species ha ing been acciden ally in oduced in ecen imes (Ca e e and
Tella, 2008b; Blackbu n e al., 2010). Among hese, pa o s a e one o he mos hea ily
aded g oups, mainly because o hei a ac i eness as pe s (Tella and Hi aldo, 2014).
Thi y-eigh pe cen o bi ds o e ed o sale in Spanish pe shops be ween 2004 and
2005 we e pa o s belonging o 72 di e en species (Ca e e and Tella, 2008b). No
su p isingly, se e al pa o species ha e es ablished non-na i e popula ions wo ldwide
(Le e , 2005; Menche i and Mo i, 2014).
In oduced pa o s may ha e a a ie y o ecological and economic impac s
(Menche i and Mo i, 2014). Rega ding ecological impac s, mos esea ch ocuses on
how in oduced pa o s a ec na i e a i auna (e.g., S ubbe and Ma hysen, 2007, 2009;
He nández-B i o e al., 2014). Howe e , e y li le in o ma ion is a ailable on how hey
in e ac wi h na i e mammals. Mo i e al. (2013) obse ed ed squi els p eying on
chicks a wo ose- inged pa akee (Psi acula k ame i) nes s, while an adul
Ba aband’s pa akee (Poly elis swainsonii) was esponsible o he dea h o an adul ed
squi el (Sciu us ulga is) in I aly. Ano he epo documen s ose- inged pa akee s
killing an adul ed squi el in F ance (Cle geau e al., 2009). Mo e ecen ly, Menche i
e al. (2014) eco ded a a al a ack by a ose- inged pa a
kee o a Leisle ’s ba
(Nyc alus leisle i) in I aly. He e, we epo se e al cases o ose- inged pa akee s
a acking and e en causing he dea h o black a s (Ra us a us) in Spain.
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Sec ion 1: Chap e II
71
Obse a ions o agg essions by non-na i e ose- inged pa akee s owa ds black
a s we e made in 2011, 2012, 2013 and 2014 in u ban pa ks o Se ille (Sou he n
Spain), and San a C uz de Tene i e (Cana y Islands). The wo pa ks in Se ille in which
agg essions we e eco ded we e Royal Alcáza (37°23'1.8''N / 5°59'29.6''W) and Ma ía
Luisa Pa k (37°22'31.57''N / 5°59'19.59''W). Rose- inged pa akee s b eed in bo h pa ks,
wi h Ma ía Luisa Pa k holding he la ges b eeding nucleus in Se ille (159 ac i e nes s
in 2013; He nández-B i o e al., 2014). Obse a ions in San a C uz de Tene i e we e
made a Ga cía Sanab ia Pa k (28°28'19''N / 16°15'13''W), whe e he b eeding
popula ion o ose- inged pa akee s eached 12 pai s in 2014 (D. He nández-B i o,
unpubl. da a).
We obse ed 21 agg essions and a acks o ose- inged pa akee s on black a s,
16 in Se ille and 5 in Tene i e. In e ac ions occu ed when a s climbed ees whe e
ose- inged pa akee s we e nes ing o ees in he icini y o nes s. When a a was
de ec ed (Fig.1), ose- inged pa akee s i s emi ed loud ala m calls and hen chased
and a acked he a using physical agg essions, some imes bi ing i and p oducing
isible wounds. All obse ed a acks we e o soli a y a s. A acks we e ca ied ou by
soli a y pa akee s o , mo e equen ly, by g oups o up o 18 indi iduals ec ui ed om
he su ounding b eeding pai s (Fig.2A). In mos cases, pa akee s s opped he a ack
when a s escaped and descended o ell o he g ound. A acks caused he dea h o wo
a s in Ma ía Luisa Pa k. In he i s e en , a a ell o he g ound om a heigh o 15
me e s when i was a acked by i e ose- inged pa akee s in he ee ops o a London
plane. The a ell di ec ly on o he pa emen , dying upon impac . The second a died
unde simila ci cums ances; he a was on a London plane wi h a ose- inged pa akee
nes and was a acked by six pa akee s, ailed o hold on o he b anches and ell o he
pa emen om a heigh o 12 me e s.
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Sec ion 1: Chap e II
72
Figu e 1 – Ins ance o a black a pe secu ed and a acked by a ose- inged pa akee in
Ma ia Luisa Pa k, Se ille, May 25 2013. Da kness p e en ed a be e -quali y image
(Pho o: Dailos He nández-B i o).
The numbe o obse a ions as well as he ime o he day a which hey we e
eco ded sugges s ha a acks by ose- inged pa akee s owa ds a s a e no anecdo al,
bu a he a di icul o obse e beha io . Ou obse a ions we e mos ly ca ied ou
du ing scheduled ield-wo k aimed o assess in e ac ions be ween ose- inged pa akee s
and na i e a i auna (He nández-B i o e al., 2014), so mos a acks we e eco ded in he
mo ning o ea ly a e noon (Fig.2B). In he wiligh hou s, ose- inged pa akee s ga he
in la ge locks and lea e o he oos (Pi hon and Dy ham, 1999; He nández-B i o and
Luna, unpubl. obse .). Only b eede emales s ay in hei nes o e nigh when hei
pai s e u n o he communal oos in he b eeding season (Bu le , 2003). The e o e, we
expec mo e a acks would ha e been eco ded i ou obse a ions included he
c epuscula pe iod be o e pa akee s e u n o he oos , gi en he noc u nal beha io o
a s (Hooke and Innes, 1995; Cox e al., 2000).
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Sec ion 1: Chap e II
73
Figu e 2 – Pe cen age o cases showing A) he numbe o ose- inged pa akee s
in ol ed in a acks on a s; and B) he ime o day when a acks we e eco ded.
Ra s (Ra us sp.) a e o en in asi e p eda o s, and hei nega i e e ec s on
na i e and endemic a i auna in oceanic islands a e well known A kinson (1985);
T a ese e al. (2009). On he o he hand, u baniza ion changes he abundance and
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Sec ion 1: Chap e II
74
ichness o p eda o communi ies Chace andWalsh (2006), and hus a s can each high
popula ion densi ies in he ci ies McKinney (2008). They can wipe ou small
popula ions o exo ic pa o s, as Sco ecci (1953) eco ded in he icini y o a zoo in
I aly, whe e a colony o monk pa akee s (Myiopsi a monachus), he i s es ablished in
his coun y (Spanò and T u i, 1986), ceased o exis as a esul o a p eda ion on
pa akee eggs and chicks. A his poin , and al hough we do no ha e di ec e idence in
he wild, black a s a e po en ial p eda o s o ose- inged pa akee nes s and e en o
adul s (while incuba ing o sleeping, as has been eco ded in cap i i y; J.L. Tella and M.
Ca e e, unpubl. obse .), so hese agg essions may se e he same pu pose as a acks
on ed squi els, also known p eda o s o pa akee nes s (Mo i e al., 2013). This
agg essi e beha io , along wi h he agg essi eness o ose- inged pa akee s owa ds
a ian p eda o s, could explain why he nes s o some bi d species a e agg ega ed in he
p oximi y o nes ing ose- inged pa akee s in Se ille (He nández-B i o e al., 2014).
Al hough ose- inged pa akee s may displace (He nández-B i o e al., 2014) and e en
kill coexis ing ba s, which compe e o nes ing ca i ies (Menche i e al., 2014), o he
bi d species may be a o ed by he an ip eda o y beha io o pa akee s, hus inc easing
hei b eeding success. Th ee bi d species (common blackbi d Tu dus me ula, spo less
s a ling S u nus unicolo and colla ed do e S ep opelia decaoc o) b eeding in he
p oximi y o ose- inged pa akee s joined he pa akee s in chasing he a s in ou cases.
Toge he , hese obse a ions sugges ha species bene i om o he species’
agg essi eness owa d p eda o s (e.g., Blanco and Tella, 1997), and indica e ha he
impac o ose- inged pa akee s on na i e auna may be bo h nega i e and posi i e
depending on he species conside ed and i s conse a ion s a us (He nández-B i o e al.,
2014).
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Sec ion 1: Chap e II
75
Despi e he ac ha non-na i e popula ions o ose- inged pa akee s ha e sp ead
h oughou a leas 35 coun ies (Bu le , 2003), esea ch aimed o assess hei impac on
na i e auna has ocused mos ly on bi ds and has cen e ed on only a ew Eu opean ci ies
(S ubbe and Ma hysen, 2007, 2009; O chan e al., 2013; He nández-B i o e al., 2014).
Mo e a en ion should be de o ed o hei in e ac ions wi h mo e elusi e mammals, such
as a s and ba s, wi h a widesp ead occu ence in ci ies wo ldwide.
ACKNOWLEDGEMENTS
We hank he s a o Ga cía Sanab ia Pa k, Ma ía Luisa Pa k and Royal Alcáza o
Se ille o allowing us access o hese enclosu es. Thanks o Emiliano Mo i o
commen s ha imp o ed he manusc ip . Field wo k was suppo ed by Fundación
Repsol and he Spanish Minis y o Economy and Compe i i eness (p ojec AIC-A-
2011-0706). The au ho s would like o acknowledge he suppo p o ided by he
Pa o Ne COST – Eu opean Coope a ion in Science and Technology- Ac ion, allowing
ui ul discussions on da a p o ided in his pape .
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expe imen al s udy. Aus al Ecology, 25(4), 375-385.
− He nández-B i o, D., Ca e e, M., Popa-Lisseanu, A. G., Ibáñez, C., & Tella, J.
L. (2014). C owding in he ci y: losing and winning compe i o s o an in asi e
bi d. PloS one, 9(6), e100593.
− Hooke , S., & Innes, J. (1995). Ranging beha iou o o es ‐dwelling ship a s,
Ra us a us, and e ec s o poisoning wi h b odi acoum. New Zealand jou nal
o zoology, 22(3), 291-304.
− Hulme, P. E. (2009). T ade, anspo and ouble: managing in asi e species
pa hways in an e a o globaliza ion. Jou nal o applied ecology, 46(1), 10-18.
− Le e , C. (2005). Na u alized Bi ds o he Wo ld. Poyse T. & Poyse , A.D,
London.
− McKinney M.L., 2008. E ec s o u baniza ion on species ichness: a e iew o
plan s and animals. U ban Ecosys. 11: 161-176.
− McKinney, M. L. (2008). E ec s o u baniza ion on species ichness: a e iew
o plan s and animals. U ban ecosys ems, 11(2), 161-176.
− Menche i M., Scale a R., Mo i E., 2014. Fi s eco d o a possibly o e looked
impac by alien pa o s on a ba (Nyc alus leisle i). Hys ix 25: 61-62.
− Mo i, E., Ancillo o, L., Menche i, M., Romeo, C., & Fe a i, N. (2013). I alian
ed squi els and in oduced pa akee s: ic ims o pe pe a o s?.
− Meye son, L. A., & Mooney, H. A. (2007). In asi e alien species in an e a o
globaliza ion. F on ie s in Ecology and he En i onmen , 5(4), 199-208.
− O chan, Y., Chi on, F., Shwa z, A., & Ka k, S. (2013). The complex in e ac ion
ne wo k among mul iple in asi e bi d species in a ca i y-nes ing communi y.
Biological In asions, 15(2), 429-445.
− Pi hon, J. A., & Dy ham, C. A. L. V. I. N. (1999). Census o he B i ish Ring-
necked Pa akee Psi acula k ame i popula ion by simul aneous coun s o oos s.
Bi d S udy, 46(1), 112-115.
− Sco ecci, G. (1953). Monaco (Myiopsi a monachus). In: Animali: come sono,
do e i ono, come i ono. Milano, I aly: Labo Edi o s.
− Spanò, S., & T u i, G. (1986). Il Pa ocche o dal colla e, Psi acula k ame i,
allo s a o libe o in Eu opa, con pa icola e i e imen o alle p esenze in I alia, e
p imi da i sul Pappagallo monaco, Myiopsi a monachus. Ri is a i aliana di
O ni ologia, 56, 231-239.
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Sec ion 1: Chap e III
83
nes ing species. The e a e no p ima y ca i y-nes ing species in his pa k (e.g.
woodpecke s) ha could exca a e ca i ies, so all a ailable ca i ies s em om ee
decay. In addi ion, he land su ounding Se ille is highly de o es ed and mos ly de o ed
o ag icul u e, hus o e ing li le habi a o o es -dwelling species.
The g ea e noc ule has a e y sca e ed dis ibu ion h oughou cen al and
sou he n Eu ope, and Spain cons i u es he main co e a ea o he species (Alcalde e al.
2016). This species shows spa ial sexual seg ega ion (Ibáñez e al. 2009) and, du ing he
b eeding season, emales ga he in small ma e ni y colonies (Alcalde e al. 2016). The
Ma ía Luisa Pa k was hough o hold he la ges local popula ion o his species, as
ea lie s udies conduc ed in jus a sec o o he pa k oughly es ima ed he p esence o
500 adul s (mos ly emales) (Ibáñez e al. 2009). Females o m ission– usion socie ies,
esul ing in se e al di e en ia ed ma e ni y colonies ha occupy se e al ca i ies in
di e en ees, wi h an a e age o 27 emales simul aneously sha ing a ee ca i y
(Popa-Lisseanu e al. 2008; Fo una e al. 2009). Females may use di e en ees ac oss
he ep oduc i e season and hus each ma e ni y colony may occupy a minimum o 30
ees wi h adequa e ca i ies, wi h he numbe o emales using a pa icula ca i y
changing o e ime (Popa-Lisseanu e al. 2008). None heless, emales show long- e m
ideli y o he same g oups o ees (some ees had been used o a leas 14 yea s;
Popa-Lisseanu e al. 2008). Females usually a i e a he pa k in Ma ch, gi ing bi h o
one o wo pups in May–June. Al hough mos o hem lea e he pa k om Augus o
No embe , o he s s ay yea ound (Popa-Lisseanu e al. 2008; Ibáñez e al. 2009). They
ha e la ge o aging anges, usually mo ing 15–40 km om he pa k in a single nigh o
hun insec s and mig a ing songbi ds (Popa-Lisseanu e al. 2009). Thei annual su i al
a es we e simila o hose o o he long-dis ance ae ial-hawking ba species (Papada ou
e al. 2012). Gene ically, his popula ion is closely ela ed o he wo closes colonies o
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84
he species ound in Je ez de la F on e a and Doñana Na ional Pa k (San os e al. 2016),
loca ed 60 km and 75 km om Ma ía Luisa Pa k, espec i ely.
The ose- inged pa akee is a success ul in ade in Eu ope. Thei success is a
esul o he la ge numbe s o indi iduals impo ed as cage bi ds, equen acciden al
escapes o delibe a ed eleases and niche simila i y be ween a eas o in oduc ion and
hei na i e Asian habi a s (Ca dado e al. 2016; Abellán e al. 2017). The i s
in oduc ion in Se ille mos p obably occu ed in 1992 wi h a e y small g oup o
indi iduals delibe a ely eleased in Ma ía Luisa Pa k. This is a highly sociable species
ha nes s in ca i ies (mos ly in ees bu also in buildings; He nández-B i o e al. 2014a)
and p oduces one o ou o sp ing pe yea . In Ma ía Luisa Pa k, mos pa akee s selec
ee ca i ies o nes ing be ween la e Decembe and Feb ua y and lay eggs mos ly in
Ma ch. The e is, howe e , la ge a ia ion in b eeding phenology among pai s, wi h
ledglings lea ing he nes s be ween May and ea ly July. Mo eo e , newly o med
imma u e pai s and g oups o non-b eeding indi iduals inspec and use ee ca i ies
h oughou he b eeding season.
Ano he eigh ca i y-nes ing bi d species a e b eeding in Ma ía Luisa Pa k
(He nández-B i o e al. 2014a). Howe e , we ha e p e iously shown ha he only
species compe ing wi h noc ules o ee ca i ies was he ose- inged pa akee
(He nández-B i o e al. 2014a). Thus, he es o he species we e no conside ed in his
s udy.
Popula ion moni o ing
Al hough ou i s s udies on noc ules in Ma ía Luisa Pa k da e back o 1992 (Popa-
Lisseanu e al. 2008), i was no un il 2003 ha we geoloca ed all ees used by he
species in a single yea . Un o una ely, we we e no able o ob ain a census o he whole
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Sec ion 1: Chap e III
85
adul popula ion o noc ules. Ob aining accu a e popula ion sizes o o es ba s o ming
ission– usion socie ies, such as he g ea e noc ule, is ex emely di icul (Hayes e al.
2009; Kunz e al. 2009). In ac , he e is no published in o ma ion on popula ion sizes
o his o simila ba species. The e o e, we elied on empo al changes in he numbe
o ees used by he species as a su oga e o changes in popula ion size. We did no
know he b eeding popula ion size o pa akee s in 2003 as he species was s ill e y
sca ce and li le a en ion was paid o i because no impac s we e expec ed a ha ime.
We can, howe e , ob ain a eliable es ima e o he numbe o pa akee nes s in 2003
om a demog aphic model buil wi h he yea o he i s in oduc ion, de ailed
popula ion coun s conduc ed since 2011 and b eeding pa ame e s ob ained om he
same popula ion (Ca e e e al. in p ep.).
In 2013 and 2016–2017, we simul aneously ook a census o he numbe o nes s
o pa akee s and he numbe o ees used by noc ules in he pa k. Ou moni o ing
p og amme ex ended om Janua y o ea ly Augus (co e ing he b eeding season o
bo h species), and obse a ions we e conduc ed in he mo ning (08.00–12.00 h) and
a e noon (17.00–21.00 h). We GPS-loca ed (±3 m) each yea all ee ca i ies ha we
we e able o isually inspec using 10 × 50 binocula s. We isually es ima ed he heigh
o he ca i y abo e g ound (in m) and he wid h o i s en ance (in cm), which was
sco ed as small, medium o la ge (less han 4 cm, 4–8 cm and g ea e han 8 cm,
espec i ely) (He nández-B i o e al. 2014a). We did no conside he o ien a ion o he
ca i ies (He nández-B i o e al. 2014a), as in o he s udies on nes -si e compe i ion
be ween pa akee s and na i e species (O chan e al. 2013; Doda o e al. 2014; Cha e e
al. 2016; Yose e al. 2016; Mo i e al. 2017), because o ien a ion did no a ec he
selec ion o ca i ies in a p e ious s udy (S ubbe and Ma hysen 2009). Ou p e ious
analyses showed a p e e ence o bo h pa akee s and noc ules o ca i ies well abo e
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Sec ion 1: Chap e III
86
g ound (app ox. 15 m) and wi h medium-sized en ances (He nández-B i o e al.
2014a). The ea e , we epea edly isi ed and obse ed a a dis ance ( o a minimum o
10 min) each ca i y on a leas 10 di e en days e enly spaced h oughou he b eeding
season o he wo species. Gi en ha pa akee s can en e ca i ies ha a e no used as
nes s (see abo e), we conse a i ely conside ed as ac i e pa akee nes s hose ca i ies
whe e we obse ed adul s en e ing a minimum o 10 imes on di e en days, hea d
chicks inside o obse ed ju eniles a he en ance. Rega ding he iden i ica ion o ees
used by noc ules, we complemen ed he p e ious me hodology o moni o ing ee
ca i ies wi h he use o an ul asound ba de ec o (Pe e sson D230) and sys ema ic
obse a ions a sunse o obse e noc ules lea ing ee ca i ies (He nández-B i o e al.
2014a).
Agg essi e in e ac ions
In e ac ions be ween pa akee s and noc ules we e obse ed du ing he moni o ing
ac i i ies desc ibed abo e. When we obse ed ha assmen and a acks o pa akee s
owa ds noc ules p esen inside o in he en ance o hei ee ca i ies, we eco ded he
du a ion o he agg ession (in minu es), whe he he noc ule was expulsed and whe he
he pa akee la e en e ed he ca i y. Dead and inju ed noc ules we e also ound, always
unde ees occupied by noc ules. A e unexpec edly encoun e ing he i s case o a
dead noc ule, we p oceeded o eco d hese e en s mo e sys ema ically. We a e
con inced ha many cases we e o e looked because dead and inju ed noc ules could be
hidden by g ound ege a ion, sca enged by ca s and a s o , as we la e lea ned,
eco e ed by people wo king in he pa k o isi o s. The co pses we ound we e
anspo ed o he labo a o y o a de ailed isual examina ion o damage o he skin and
bones.
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Sec ion 1: Chap e III
87
Spa ial a angemen o noc ules
Each yea (2013, 2016 and 2017) we geoloca ed he ca i ies used by pa akee s and
noc ules as well as hose ha emained unoccupied. The occupancy o a pa icula ee
ca i y by noc ules could be in luenced by he a ailabili y o unoccupied ca i ies as well
as by he spa ial dis ibu ion o ca i ies occupied by he same and/o o he species,
d i en no only by compe i ion bu also by conspeci ic a ac ion p ocesses (He nández-
B i o e al. 2014a). We hus measu ed he Euclidean dis ance om each ca i y o he
nea es ca i y occupied by noc ules and pa akee s (nea es -neighbou dis ance) as well
as he co esponding agg ega ion indexes. Agg ega ion indexes we e ob ained as he
ela i e posi ion o each ca i y wi hin he whole dis ibu ion o all ca i ies occupied by
conspeci ics o he e ospeci ics using Σexp(−d
ij
), wi h (i ≠ j), whe e d
ij
is he linea
dis ance be ween ca i ies i and j, and j ep esen s all occupied ca i ies (Moilanen and
Hanski 1998). Nea es -neighbou dis ances and agg ega ion indexes a e complemen a y
and depic he social en i onmen a ound each ca i y a a landscape scale as well as he
exis ence o close conspeci ics and compe i o s in i s p oximi y (He nández-B i o e al.
2014a). The wo main sou ces o habi a he e ogenei y in ou s udy a ea, i.e. he
p oximi y o su ounding s ee s and o es co e , we e no conside ed because ou
p e ious wo k showed no e ec s on he spa ial a angemen o noc ules and pa akee s
(He nández-B i o e al. 2014a).
We employed gene alized linea models (GLMs) wi h a binomial e o
dis ibu ion and logis ic-link unc ion o asce ain ac o s explaining he p obabili y o
occupa ion o a ca i y by noc ules, i ing as explana o y a iables i s ai s (en ance
size and heigh abo e g ound) and i s nea es dis ance and agg ega ion o bo h noc ules
and pa akee s. As alues o nea es dis ance and agg ega ion o he same species we e
highly co ela ed (all p < 0.001), we al e na i ely included in models only one o hese
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Sec ion 1: Chap e III
88
spa ial desc ip o s. Con inuous a iables we e s anda dized o modelling. Models we e
sepa a ely buil o he mos dis an yea s (2013 and 2017) om which we ga he ed
spa ial in o ma ion om bo h species, using he Akaike in o ma ion c i e ion co ec ed
o small sample sizes (AICc) o model selec ion (Bu nham and Ande son 2002).
Wi hin each se o candida e models, we calcula ed ∆AICci as he di e ence be ween
he AICc o model i and ha o he bes -suppo ed model (i.e. he model wi h he lowes
AICc). Models wi hin 2 AICc uni s o he bes suppo ing model we e conside ed as
al e na i es. We also quan i ied he plausibili y o each model as being he bes
app oxima ion using Akaike weigh s, w (Bu nham and Ande son 2002), and pe o med
model a e aging (MuMIn package) o es ima e he ela i e impo ance o all a iables
h ough he calcula ion o model-a e aged es ima es and con idence in e als (CIs)
using he se o al e na i e models. A gi en e ec ecei ed no, weak o s ong suppo
when he 95% CI o he coe icien es ima e s ongly o e lapped ze o, ba ely
o e lapped ze o o did no o e lap ze o, espec i ely. We calcula ed he pe cen age o
de iance explained by he bes -suppo ed models, ob ained as 100 − (de iance o model
i/de iance null model) * 100), o assessing hei goodness o i . All s a is ical analyses
we e conduc ed in R . 3.1.2 (R Co e Team 2013), and aw da a used o analyses a e
p o ided in he elec onic supplemen a y ma e ial.
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Sec ion 1: Chap e III
89
RESULTS
In e speci ic agg essions
Du ing he b eeding seasons o 2016 and 2017, we eco ded 36 agg essions o pa akee s
owa ds noc ules in ees occupied by bo h species. Mos agg essions (55.6%) we e
eco ded in May and du ing he 3 h be o e sunse (83.3% o agg essions). All o hese
agg essions happened a he en ance o ee ca i ies, and in 16 cases, we obse ed
pa akee s a acking noc ules wi h hei beaks while emi ing loud sounds ( igu e 1).
These agg essions usually las ed om 1 o 25 min, eaching in one case up o 145 min
(median: 13.13 min), and ended wi h he pa akee s en e ing he ee ca i y a e he
a acked noc ule was expulsed and obliged o lee du ing he day ime. Once pa akee s
en e ed he ee ca i y, we we e no able o obse e whe he o no hey a acked o he
noc ules ha may ha e been inside he ca i y. In 20 o he ins ances, pa akee s ceased
hei agg ession and le he ee a sunse wi hou success ully expelling he noc ules
om hei e uge ca i ies.
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Sec ion 1: Chap e III
90
Figu e 1. Main pic u e: an adul emale ose- inged pa akee a acks a g ea e noc ule a
he en ance o a ee ca i y used as a e uge by noc ules in Ma ía Luisa Pa k, Se ille,
on 17 May 2016. One noc ule was ound dead unde he same ee he nex day, wi h
wounds caused by pa akee s. Accompanying pic u es: de ails o di e en ypes o
wounds caused by pa akee s on noc ules ound dead unde hei ee e uges
(pho og aphs: Dailos He nández-B i o).
Du ing he same pe iod, we ound 20 dead and wo inju ed noc ules unde 18
di e en ees, all o hem wi h ac i e nes s o pa akee s. Se en o hese noc ules we e
ound dead unde he same ee whe e we eco ded agg essions by pa akee s he
p e ious day. Dead noc ules included h ee ew-day-old pups, 10 lac a ing young and
se en adul s, as well as one p egnan emale. Eigh o hese co pses we e oo pu e ied
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Sec ion 1: Chap e III
91
o allow us o obse e wounds o o he indica ions o a ack ( empe a u es eached 33–
40°C du ing he s udy pe iod, so co pses decomposed apidly). The o he 12 co pses
we e esh enough o show clea wounds caused by he beaks o pa akee s ( igu e 1).
These wounds we e p esen mainly on he wings, consis ing o holes in he memb anes
(dac ylopa agium and plagiopa agium a eas), cha ing on he skin co e ing he phalanges
and o ea m, b eakages o phalanges, and wounds bo h on he uppe side o he body
and he abdomen. The wo inju ed, bu s ill li ing adul noc ules showed simila wounds
and we e no able o ly o o climb he unks o ees o e u n o hei ca i y e uges.
We suspec ha noc ules we e inju ed and killed when pa akee s en e ed hei ca i ies
a e expelling o he noc ules (see abo e), because pups and lac a ing young do no
lea e he ca i ies. We did no obse e agg essions add essed o noc ules by o he
species, nei he did we ind e idence o o he species p eying on o killing noc ules in
he pa k.
Tempo al ends in he occupa ion o ees
Figu e 2 shows he numbe s o nes s o pa akee s and ees occupied by noc ules in
Ma ía Luisa Pa k om 2003 o 2017. Pa akee s we e sca ce in his a ea in 2003 and,
al hough we did no conduc a de ailed census, ou demog aphic-based es ima ion
sugges s ha he local b eeding popula ion would ha e been as low as 13 nes s a ha
ime. Th ee annual de ailed censuses conduc ed since 2013 showed an inc ease o 311
ac i e nes s in 2017 ( igu e 2). This means a 96% popula ion inc ease om 2013 o
2017, and an es ima ed 2192% popula ion inc eases om 2003 o 2017. Noc ules
showed an opposi e end: ca i ies used as e uges we e ound in 75 ees in 2003, he
numbe dec easing o only 14 ees in 2017 ( igu e 2). This esul ed in a 70% dec ease
be ween 2013 and 2017, and an 81% dec ease be ween 2003 and 2017.
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Figu e 2.
Changes in he n
by g ea e noc ules ac oss y
Spa ial seg ega ion be ween
Figu e 3 shows he spa ial
ac oss yea s in he pa k, ind
con ac ion o noc ule e ug
nes s. In 2013, he i s y
pa akee nes s (n
= 159 nes
p obabili y a ee ca i y wa
heigh abo e g ound and o
nes s ( able 1). In his yea
medium- o la ge-
size en a
pa akee nes ( able 2). In
declined o 14 while he nu
Sec
e numbe o nes s o ose
-
inged pa akee s and
s yea s in Ma ía Luisa Pa k.
een noc ules and pa akee s
ial dis ibu ion o pa akee nes s and noc ule
indica ing no only a educ ion in he numbe b
uges along wi h an inc ease and spa ial expan
yea we eco ded bo h noc ule e uges (n
nes s) ( igu e 3), he bes
-
suppo ed model in
was occupied by noc ules was ela ed o i s en
o he nea es dis ances o o he noc ule e ug
ea , noc ules used ca i ies well abo e g ound
ances, close o o he noc ule e uges and a
In 2017, when he numbe o ees occupied
numbe o pa akee nes s had inc eased o 31
ec ion 1: Chap e III
92
and ees occupied
e e uges eco ded
bu also a spa ial
ansion in
pa akee
= 47 ees) and
indica es ha he
en an
ce size and
uges and pa akee
nd le el and wi h
om he nea es
d by noc ules had
311 ( igu e 3), he
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Sec ion 1: Chap e III
99
p eda ion a es is e y limi ed (Ba clay and Ha de 2003). Mo eo e , he annual
su i al a es o noc ules li ing in Ma ía Luisa Pa k be o e he popula ion g ow h o
pa akee s was low (0.74) compa ed o o he o es ba species ha o age a sho e
dis ances (Papada ou e al. 2012). The e o e, an inc emen in mo ali y a es caused by
pa akee s bo h on lac a ing and adul noc ules would se e ely a ec he popula ion
dynamics o his popula ion and con ibu e o i s d ama ic decline. Un o una ely, we
we e unable o measu e empo al changes in popula ion size owing o he
me hodological di icul ies o accu a ely es ima ing popula ion sizes o o es ba
species ha o m ission– usion socie ies (Hayes e al. 2009; Kunz e al. 2009).
Howe e , i is easonably expec ed ha he popula ion size would ha e declined in
pa allel wi h he 81% educ ion in he numbe o ee ca i ies used. Popula ion size, as
well as social s uc u e, is expec ed o be a ec ed: emales we e o iginally dis ibu ed
ac oss he pa k o ming di e en ia ed, s able ma e ni y colonies (Popa-Lisseanu e al.
2008; Fo una e al. 2009; igu e 3), some hing ce ainly dis up ed gi en he sca ci y
and agg ega ion o he ees cu en ly used ( igu e 3). This social dis up ion may ha e
unexpec ed e ec s on he b eeding biology o he species.
Ac ions o educe he popula ion o pa akee s and o p o ide a i icial e uges o
noc ules a e u gen ly needed. Wi hou ha , his popula ion, which was once he la ges
known o his h ea ened ba species in i s ange (Fo una e al. 2009), could be
comple ely ex inc wi hin a ew yea s. The p o ision o a i icial e uges alone would be
insu icien , gi en ha noc ules lea n o use hem slowly (C. Ibáñez 2003, unpublished
da a) and ha he popula ion size and di ec impac s o pa akee s a e inc easing a a
much as e a e. An e adica ion plan o pa akee s was planned o ea ly 2017 by he
ci y go e nmen o Se ille, bu his was cancelled owing o p essu es by animal wel a e
associa ions, e en hough i was suppo ed by a Spanish law which speci ically deals
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Sec ion 1: Chap e III
100
wi h he managemen o in asi e species (Real Dec e o 630/2013). P og ammes o he
con ol o e adica ion o in asi e species o en ace public opposi ion (Blackbu n e al.
2010), especially when dealing wi h cha isma ic species such as pa akee s (Ca e e and
Tella 2008).
Ou wo k exempli ies he challenges in unde s anding he ue ecological
impac s o in asi e bi ds. While he numbe o in oduced non-na i e bi d species
inc eases wo ldwide (Dye e al. 2017) and in asion isks inc ease in new egions
(Ca dado e al. 2017), he scien i ic communi y is only able o s udy a small ac ion o
hese popula ions and hei mul i ace ed po en ial impac s (Ma in-Alba acin e al.
2015). Mo eo e , some impac s, such as hose epo ed he e, a e unexpec ed and can be
easily o e looked in he absence o long- e m esea ch. In his sense, we specula e
whe he o he impac s on ba s emain hidden. Many Eu opean ba species, some o
hem h ea ened, a e closely linked o buildings and u ban habi a s (Ancillo o e al.
2015; Russo and Ancillo o 2015; Rydell e al. 2017)); meanwhile, pa akee s a e
sp eading ac oss Eu opean ci ies (Ca dado e al. 2016; Abellán e al. 2017). The e is
conce n ha pa akee s could educe he a ailabili y o sui able ee ca i ies o he
noc ule ba (Nyc alus noc ula) in he Ne he lands (Haa sma and an de G aa 2013).
One Leisle 's ba (Nyc alus leisle i) was ound killed by pa akee s in I aly (Menche i e
al. 2014), and pa akee s seem o be compe ing o g ea e noc ule e uges in Je ez de la
F on e a (I. Sánchez and D. He nández-B i o 2017, pe sonal obse a ion). Pa akee s can
also compe e o ca i ies in buildings ha hey use o nes ing, which can be usu ped
om o he bi d (He nández-B i o e al. 2014a) and ba species. One o he au ho s (D.
He nández-B i o) obse ed in Se ille, on 7 July 2017, h ee pa akee s a emp ing o
o ce isabelline se o ine ba s (Ep esicus isabellinus) ou om a wall ca i y in a all
building du ing he day ime. Se en y-eigh ba s la e lew ou om ha ca i y a sunse .
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Sec ion 1: Chap e III
101
All o hese obse a ions can be conside ed as anecdo al and do no necessa ily imply
an impac on ba popula ions. Simila ly, he i s obse a ion o pa akee s ha assing a
noc ule in Ma ía Luisa Pa k in 2005 (He nández-B i o e al. 2014a) would no ha e
indica ed a popula ion impac un il he long- e m esea ch p esen ed he e. As a ma e o
conce n, a ecen me a-analysis shows ha la ge impac s caused by in asi e species can
o en be missed owing o small sample sizes, esul ing in high Type II e o a es and
alse ce ain y o no impac (Da idson and Hewi 2014). The e o e, much mo e
esea ch is needed o p ope ly assess he impac o pa akee s and o he in asi e species
on a a ie y o ba species (Welc and Leppanen 2017).
ACKNOWLEDGEMENTS
We hank he Ma ía Luisa Pa k s a o acili a ing he wo k wi hin he pa k. J.
Nogue as helped wi h da a managemen , and se e al people om he Depa men o
Conse a ion Biology and E olu iona y Ecology o EBD-CSIC, s uden s om Pablo de
Ola ide Uni e si y and o he olun ee s helped o eco d ield obse a ions. E. Mo i
and an anonymous e iewe helped o imp o e his pape . This wo k was unded by he
O ganismo Au ónomo de Pa ques Nacionales (MMA) o Spain (p ojec 021/2002) o
C.I., and Fundación Repsol, he Se e o Ochoa P og am (SVP-2014-068732) and Ac ion
COST ES1304 (Pa o Ne ) o D.H-B., J.L.T. and M.C
SUPPLEMENTARY MATERIAL
Appendix S1. (see Supplemen a y Ma e ial Sec ion 1-Chap e III).
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Sec ion 1: Chap e III
102
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124
SECTION 2:
NOVEL INTERACTIONS
AGAINST BIOTIC BARRIES
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Sec ion 2: Chap e V
131
a e also ole a ed by whi e s o ks and use hei nes s as nes ing subs a es (Indykiewicz
2006). The la ge-bodied whi e s o k has ex emely low nes p eda ion a es, as lea ned
om long- e m b eeding moni o ing p og ams in Spain (e.g.; Baos e al. 2012; Ve ga a
e al. 2006), and p eda ion o adul s is anecdo al (Tobolka 2014). The e o e, smalle
bi d species may also associa e wi h whi e s o ks o educe p eda ion isk (Bocheński
2005; Kosicki e al. 2007), as i has been demons a ed o o he p o ec i e nes ing
associa ions (Quinn and Ue a 2008).
Fig. 1 Pa akee s and s o ks. Nes s o monk pa akee s (yellow a ows) associa ed wi h
whi e s o k nes s. (Pho os: D. He nández-B i o)
He e, we hypo hesised ha monk pa akee s associa e wi h s o ks no because o
s uc u al bene i s bu o ob ain p o ec ion agains p eda ion, hus allowing hei sp ead
in o u al habi a s despi e he exis ence o a la ge p eda o communi y. To es his
hypo hesis, we i s compa ed he p obabili y o pa akee -s o k associa ions be ween
u al and u ban habi a s, p edic ing ha hey should be mo e equen in he o me ,
whe e p eda o s a e mo e abundan han in he la e . We hen assessed whe he hese
associa ions occu a andom, o i hey can be explained by a combina ion o bio ic and
abio ic ac o s such as he p oximi y o he s o k nes o ano he pa akee colony
(conspeci ic densi y), he ype o subs a e (pylons, ees, o oo s), o he densi y o
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Sec ion 2: Chap e V
132
p eda o s in he su oundings (p eda ion isk). We p edic ed ha besides p oximi y o
conspeci ics, pa akee s should b eed p e e en ially in s o k nes s si ed in pylons ( hus
educing p eda ion isk a he nes ) in a eas wi h a low densi y o ae ial p eda o s ( hus
educing p eda ion isk while o aging). The low densi y o p eda o s in he su ounding
o a nes ensu es sa e a eas o pa akee s o pe o m basic ac i i ies such as o aging.
Nes s in pylons, con a y o hose loca ed in ees o on oo s, a e no accessible o
mammalian o ep ilian p eda o s (Bocheński 2005), so he an ip eda o y e ec o
nes ing wi h a s o k is ocused on ae ial p eda o s (i.e., ap o s) and, hus, maximised.
Finally, o disca d he po en ial bene i s de i ed om he nes s uc u e pe se, we
e alua ed he e ec o nes abandonmen by s o ks on he subsequen nes abandonmen
by pa akee s. We p edic ha pa akee s should abandon hei nes s a e s o k
abandonmen due o he disappea ance o i s p o ec i e e ec agains p eda o s;
o he wise, pa akee s could associa e wi h s o ks o simply ake ad an age o hei nes
s uc u es. Complemen a ily, we compa ed he beha iou o pa akee s owa d
app oaching a ian p eda o s when b eeding in associa ion wi h s o ks o no . We
p edic ed ha pa akee s should lush mo e equen ly om ap o s when b eeding alone
han when b eeding wi h s o ks, whe e hey can ake ad an age o he p esence o he
p o ec i e species, which can de e ap o a acks.
MATERIAL AND METHODS
S udy a ea and ieldwo k
The s udy was ca ied ou in an ex ensi e a ea including he ci y o Mad id and i s
su ounding u al habi a s along he Manzana es and Ja ama i e s (Fig. (Fig.2),2), an
a ea mos ly de o ed o i iga ed, in ensi e ag icul u e (mainly maize and ege ables)
and g a el ex ac ion. In his a ea, ap o s nes mos ly in ipa ian o es s, while whi e
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Sec ion 2: Chap e V
133
s o ks nes in he same o es as well as on elec ic pylons and building oo s (Blanco
1994; 1996).
Du ing he b eeding seasons (Ap il–Augus ) o 2014 and 2015, he s udy a ea
was epea edly isi ed o GPS-loca e all nes s o pa akee s and s o ks p esen in he
u ban and u al habi a s (Fig. (Fig.2),2), also eco ding he ype o subs a e in which
he nes s we e loca ed ( ee, pylon o oo ). Mo eo e , we moni o ed he communi y o
medium-sized ap o species p esen in he s udy a ea, including he black ki e Mil us
mig ans, he boo ed eagle Hie aae us penna us, he common buzza d Bu eo bu eo, he
no he n goshawk Accipi e gen ilis, he ed ki e Mil us mil us and he pe eg ine alcon
Falco pe eg inus. P eda ion o monk pa akee s by ap o s has been eco ded in i s
in aded ange (Bu ge and Goch eld 2009; Reed e al. 2014; B iceño e al. 2019), as
well in ou s udy a ea, such as pe eg ine alcon (Sánchez-Ma eos and Sánchez-Alba án
2017) and golden eagle Aquila ch ysae os (E. Na a o pe s. comm.). We eco ded
emains o monk pa akee s in se e al nes s o black ki es, boo ed eagles and pe eg ine
alcons and obse ed hei hun ing a emp s on lying pa akee s. The o he wo ap o
species a e o simila size and beha iou , and also include bi ds in hei die s (Fo sman
1999). Thus, we conside ed he i e ap o species as po en ial p eda o s. We did no
ind e idence o o he bi d species, such as co ids, p eying upon monk pa akee s o
hei nes s.
Fo s o ks and ap o s, each nes co esponds o a single b eeding pai , whe eas
o pa akee s each nes can house om one o se e al b eeding pai s accommoda ed in
di e en chambe s ( ange: 1–35 ac i e chambe s). A pa akee nes was classi ied as
being associa ed wi h a s o k i hey sha ed he same nes ing subs a e, i he pa akee
nes was loca ed on he same s uc u e (i.e., he same ee o elec ici y pylon) as he
s o k nes , o i he pa akee nes was wi hin a adius o 15 m o a s o k nes (Fig.
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Sec ion 2: Chap e V
134
(Fig.1).1). Al hough mammals and snakes can p eda e on pa akee s, hey we e no
conside ed in his s udy as we do no ha e accu a e in o ma ion abou hei dis ibu ion
and abundance. Howe e , hei po en ial impac on pa akee s is discussed based on nes
subs a e (see p e ious).
In e ac ions be ween pa akee s and ap o s
We eco ded he esponses o pa akee s owa ds in uding ap o s (i.e., ap o s lying
wi hin less han 15 m o an ac i e pa akee nes ) ac oss he s udy a ea, ollowing
p e ious wo k conduc ed on a simila species, he ose- inged pa akee (Psi acula
k ame i; He nández-B i o e al. 2014). Pa akee esponses we e classi ied as s ay (i.e.,
when pa akee s s ayed in hei nes s, showing no sign o ea owa d he ap o ),
mobbing (i.e., when pa akee s lew o a ack he app oaching ap o ), o lush (i.e., when
pa akee s lew away om he ap o ).
S a is ical analyses
We used Gene alised Linea Models o es i pa akee -s o k associa ions we e mo e
likely in u al han in u ban a eas (logis ic link unc ions, binomial e o dis ibu ions)
by conside ing all pa akee nes s p esen in he s udy a ea and including he habi a
whe e hey we e loca ed (i.e., u ban o u al) as an explana o y a iable. We hen
e alua ed i pa akee s used u al s o k nes s (p obabili y o pa akee -s o k associa ion
conside ing all s o k nes s p esen in he u al a eas; logis ic link unc ions, binomial
e o dis ibu ions) based on a combina ion o conspeci ic densi y, ype o subs a e and
p eda ion isk. Conspeci ic densi y was ob ained as he ela i e posi ion o each
pa akee nes wi hin he pa akee popula ion. We used he o mula Σexp(−dij)*A, whe e
dij is he linea dis ance be ween each pa akee nes i and all pa akee nes s j, and A is
he numbe o chambe s pe nes j (Moilanen and Hanski 1998). Highe alues o his
index poin o a highe densi y o conspeci ics a ound a selec ed loca ion. P eda ion isk
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Sec ion 2: Chap e V
135
was assessed by using he agg ega ion o ap o s nes s as a p oxy, calcula ed using he
same index explained p e iously (no e ha he e A always equals 1). Models o he
p obabili y o pa akee -s o k associa ions we e sepa a ely un o 2014 and 2015
because o con e gence p oblems when using gene alised linea mixed models and nes
as a andom e m. Finally, we ela ed he p obabili y o nes abandonmen by u al
pa akee s wi h nes abandonmen by he s o k (independen a iable), conside ing i
colonies occupied in 2014 emained occupied in 2015 (logis ic link unc ions, binomial
e o dis ibu ions). All con inuous a iables we e included in hei linea and quad a ic
o ms and s anda dised be o e modelling. Model selec ion was pe o med using he
Akaike In o ma ion C i e ion co ec ed o small sample sizes, AICc; (Bu nham and
Ande son 2002). Wi hin each se o models (which includes he null model), we
calcula ed he ∆AICci (as he di e ence be ween he AICc o model i and ha o he
bes model) and he weigh (w) o each model. Models wi hin 2 AICc uni s o he bes
one we e conside ed as al e na i es and used o pe o m model a e aging (MuMIn
package). We conside ed ha a gi en e ec ecei ed no, weak o s ong s a is ical
suppo when he 95% con idence in e al (CI) s ongly o e lapped wi h ze o, ba ely
o e lapped wi h ze o, o did no o e lap wi h ze o, espec i ely. S a is ical analyses
we e conduc ed in R 3.1.2 (R Co e Team, 2013).
Di e ences in beha iou al esponses o pa akee s owa d app oaching ap o s
we e compa ed among habi a s (u ban o u al) and be ween nes s associa ed o no
associa ed wi h s o ks using gene alised linea models (mul inomial e o dis ibu ion).
We included he numbe o pa akee s p esen in each ap o in usion and he in e ac ing
ap o species as co a ia es o con ol o hei po en ial e ec s.
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Sec ion 2: Chap e V
136
RESULTS
P o ec i e nes associa ions
We eco ded mo e han 900 monk pa akee nes s, mos o hem loca ed in he u ban
habi a , while a ian p eda o s we e much mo e abundan in u al habi a s (Fig. 2; Table
1). Pa akee s nes ing in u al habi a s we e a e and mainly associa ed wi h s o ks
(97.06% o pa akee nes s associa ed wi h s o ks in 2014, n = 34; 73.53% o pa akee
nes s associa ed wi h s o ks in 2015, n = 34), an associa ion ha was nea comple ely
absen among u ban pa akee s (habi a (u ban): es ima e: -2.30; 95% CI: − 2.88 - -1.73).
Impo an ly, pa akee s b ed mainly associa ed wi h s o k nes s loca ed in pylons (70 and
100% o associa ed nes s in 2014 and 2015 we e in pylons, Table Table1).1). Howe e ,
models un o 2014 and 2015 show ha he p obabili y o pa akee -s o k associa ion
was only ela ed o habi a (i.e., mo e likely in u al han in u ban a eas; Table 2).
Fig. 2 S udy a ea. U ban (da k g ey) and su ounding u al (whi e) a eas o Mad id
Me opoli an a ea (40° 21′ 03.1“ N, 3° 30’ 06.1” W). Di e en colou ed poin s show he
loca ion o nes s o ap o s ( ed), s o ks (yellow), and pa akee s associa ed (blue) o no
associa ed (pu ple) wi h s o ks. Black dashed lines a e i e s
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Sec ion 2: Chap e V
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Table 1 Abundance o species in he s udy a ea. Numbe o nes s o Monk pa akee s,
whi e s o ks and a ian p eda o s in u ban and u al a eas. The ype o subs a e (i.e.,
pylon, ee o oo ) is indica ed o Monk pa akee and whi e s o k nes s
Species 2014 2015
U ban Ru al U ban Ru al
Monk pa akee s Myiopsi a monachus 867 34 890 34
Associa ed wi h s o ks
Pylons 0 23 0 25
T ees 9 10 9 0
Roo s 0 0 0 0
No associa ed wi h s o ks
Pylons 2 0 2 0
T ees 856 1 879 9
Roo s 0 0 0 0
Whi e s o ks Ciconia ciconia 41 466 47 440
Pylons 6 233 7 209
T ees 32 217 37 216
Roo s 3 16 3 15
Black ki es Mil us mig ans 0 239 0 244
Common buzza ds Bu eo bu eo 0 15 0 14
Boo ed eagles Hie aae us penna us 0 5 0 4
No he n goshawks Accipi e gen ilis 0 3 0 3
Red ki es Mil us mil us 0 1 0 1
Pe eg ine alcon Falco pe eg inus 6 0 6 0
Wi hin u al habi a s, s o k nes s we e no used a andom, and pa akee s
selec ed, among hose a ailable, s o k nes s loca ed in a eas whe e conspeci ics we e
mo e abundan and p eda ion isk was lowe (Table 3; Fig. 3). I is wo h no ing ha he
co ela ion be ween conspeci ic densi y and p eda ion isk inc eased om 2014 o 2015
(− 0.38 and − 0.48, espec i ely), so p eda ion isk ecei ed weake suppo in models
ob ained o he second yea . Howe e , when conspeci ic densi y was excluded om
models, p eda ion isk was s ongly ela ed o s o k nes s also used in 2015 (es ima e: -
1.34, 95% CI: − 2.60 - -0.47).
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Sec ion 2: Chap e V
138
**
*
97.5%
1852.17
-3.58
0.03
6577.41
2.5%
-1814.86
-5.61
-1.73
-6642.7
Es ima e
18.66
-4.59
-0.85
-32.65
Va iables
subs a e (pylon)
habi a (u ban)
yea (2015)
habi a (u ban)*subs a e (pylon)
Weigh
0.58
0.24
0.13
0.06
0
0
0
0
0
0
0
∆AICc
0
1.81
3.08
4.64
22.49
26.35
26.95
84.12
85.48
399.36
400.26
AICc
234.58
236.4
237.66
239.22
257.07
260.93
261.53
318.7
320.07
633.94
634.84
k
5
4
4
3
4
3
2
3
2
1
2
Model
subs a e*habi a + yea
subs a e*habi a
subs a e + habi a + yea
subs a e + habi a
habi a *yea
habi a + yea
habi a
subs a e + yea
subs a e
null
yea
R
2
=0.70
Table 2 Fac o s a ec ing he p obabili y o associa ion be ween pa akee s and s o ks. Rela i e impo ance o
habi a (u ban and u al) and ype o subs a e (pylon, ee o oo ) on he p obabili y o p o ec i e nes ing
associa ions be ween monk pa akee s Myiopsi a monachus and whi e s o ks Ciconia ciconia. Es ima es and 95%
con idence in e als (2.5 and 97.5%) we e assessed a e model a e aging (∆AIC ≤2). We conside ed ha a
gi en a iable has no, weak o s ong suppo when he 95% con idence in e al s ongly o e lapped ze o, ba ely
o e lapped ze o (*), o did no o e lap ze o (**), espec i ely. k: numbe o pa ame e s. AICc: Akaike
In o ma ion C i e ion co ec ed o small sample sizes. ∆AICc: di e ence be ween he AICc o model i and ha
o he bes -suppo ed model (i.e. he model wi h he lowes AICc); w: Akaike weigh s. R
2
: measu e o how well
he model explains he da a
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Sec ion 2: Chap e V
139
**
**
**
97.50%
-1.94
3.81
2563.8
2564.79
97.50%
2.84
13256.1
13745.84
1.25
2.50%
-5.47
1.56
-2533.35
-2532.36
2.50%
1.19
-
13219.31
-
13742.57
-2.75
Es ima e
-3.7
2.69
15.22
16.21
Es ima e
2.02
18.4
1.63
-0.75
Va iables
p eda ion isk
conspeci ic densi y
subs a e (pylon)
subs a e ( ee)
Va iables
conspeci ic densi y
subs a e (pylon)
subs a e ( ee)
p eda ion isk
weigh
0.67
0.33
0
0
0
0
0
0
weigh
0.36
0.31
0.19
0.15
0
0
0
0
∆AICc
0
1.38
21.19
26.31
66.34
72.52
75.76
95.04
∆AICc
0
0.32
1.26
1.8
48.71
58.04
63.18
92.88
AICc
107.56
108.94
128.75
133.87
173.9
180.08
183.32
202.61
AICc
101.07
101.39
102.34
102.87
149.78
159.11
164.26
193.96
k
3
5
4
2
4
2
3
1
k
4
2
3
5
4
3
2
1
Model 2014
p eda ion isk + conspeci ic densi y
p eda ion isk + conspeci ic densi y + subs a e
conspeci ic densi y + subs a e
conspeci ic densi y
p eda ion isk + subs a e
p eda ion isk
subs a e
null
Model 2015
conspeci ic densi y + subs a e
conspeci ic densi y
p eda ion isk + conspeci ic densi y
p eda ion isk + conspeci ic densi y + subs a e
p eda ion isk + subs a e
subs a e
p eda ion isk
null
2014: R
2
=0.39
2015: R
2
=0.40
Table 3 Fac o s a ec ing he p obabili y o associa ion be ween pa akee s and s o ks in u al a eas. Rela i e impo ance o p eda ion isk,
conspeci ic agg ega ion and subs a e on he p obabili y o p o ec i e nes ing associa ions be ween Monk pa akee s Myiopsi a monachus and
whi e s o ks Ciconia ciconia. Es ima es and 95% con idence in e als (2.5 and 97.5%) we e assessed a e model a e aging (∆AIC ≤2). We
conside ed ha a gi en a iable has no, weak o s ong suppo when he 95% con idence in e al s ongly o e lapped ze o, ba ely o e lapped
ze o (*), o did no o e lap ze o (**), espec i ely. Models we e un sepa a ely o 2014 and 2015. k: numbe o pa ame e s. AICc: Akaike
In o ma ion C i e ion co ec ed o small sample sizes. ∆AICc: di e ence be ween he AICc o model i and ha o he bes -suppo ed model (i.e.
he model wi h he lowes AICc); w: Akaike weigh s. R2: measu e o how well he model explains he da a
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Sec ion 2: Chap e V
140
Fig. 3 P obabili y o pa akee -s o k associa ion in u al habi a s. Pa akee s selec u al
s o k nes s loca ed a he om p eda o s (agg ega ion o p eda o s) and su ounded by
la ge densi ies o conspeci ics (agg ega ion o pa akee s). Es ima es (solid lines),
con idence in e als (dashed lines) and aw da a (black do s) a e shown o 2014 and
2015
P obabili y o nes abandonmen by pa akee s
F om 2014 o 2015, 44% o he 34 u al pa akee nes s we e abandoned, while all u ban
pa akee nes s emained ac i e. This high a e o abandonmen eco ded among u al
nes s was s ongly explained by he abandonmen o he nes by he s o k (Table 4).
Subs a e, conspeci ic densi y and p eda ion isk we e weakly suppo ed, wi h nes s
loca ed in pylons, a om conspeci ics and in a eas wi h a low densi y o p eda o s
being mo e p one o abandonmen when hey we e abandoned by s o ks.
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Sec ion 2: Chap e V
147
inb eeding dep ession o he e olu ion o adap a ions o he new habi a - is an impo an
pa o he colonisa ion p ocess (Sakai e al. 2001; C ooks 2005; Lockwood 2013).
Sp ead lag phases may ake yea s o decades, can a y among species and popula ions
o he same species subjec o di e en ecological condi ions (Aagaa d 2014) and a e
highly unp edic able (Cou s e al. 2018). The e o e, i is no p uden o assume ha
exo ic species ha ha e been obse ed in u ban a eas o a long pe iod will emain
s ic ly u ban in he u u e. In ac , he dis ibu ion o monk pa akee s in ading Is ael
has inc eased and shi ed om p edominan ly u ban a eas o ag icul u al landscapes in
less han wo decades (Cou s e al. 2018). This may ha e been acili a ed by he long-
e m decline and poo conse a ion s a us o ap o s in ha coun y (Yom-To e al.
2012). Thus, accep ing in asi e species as pa o he u ban ecosys em may some imes
esul in hei sp ead in o adjacen u al landscapes, whe e hey could ha e di e en ,
o en unknown impac s (Bo ham e al. 2009, Mo ei a-A ce e al. 2014), such as c op
damage in he case o monk pa akee s (Sena e al. 2016; Pos igo e al. 2017).
The monk pa akee was i s in oduced in Spain in 1976, inc easing i s
dis ibu ion since hen a a a e o 8.14 g id cells (5 × 5 km) pe yea (Abellán e al.
2017). Howe e , i was no un il ecen ly ha he Spanish popula ion began o g ow
exponen ially, inc easing om c. 6000 indi iduals in 2010 o c. 20,000 in 2015
dis ibu ed ac oss > 130 u ban popula ions (Molina e al. 2016). The e o e, i seems ha
he species has o e come he lag phase, and he p o ec i e nes ing associa ion
acili a ing i s sp ead ou side o ci ies, a he han anecdo al, may be occu ing in o he
a eas. P edic i e models indica e ha he e is s ill plen y o sui able habi a o he
species (Muñoz e al. 2006; Molina e al. 2016). Fu he mo e, whi e s o ks a e widely
dis ibu ed ac oss Spain (wi h > 33,000 nes s in 2004; Molina e al. 2004) and
popula ions a e apidly inc easing hanks o he use o human- ela ed ood subsidies
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Sec ion 2: Chap e V
148
(in asi e Ame ican c ay ish and ubbish dumps (Blanco 1996; Tablado e al. 2010;
Sanz-Aguila e al. 2015). In ac , in ecen yea s, monk pa akee s also sp ead om
ano he Spanish ci y (Za agoza, 270 km dis an o Mad id), by nes ing in s o k nes s
(J.L. Tella obs. Pe s.), bu such a sp ead was hal ed by he esponsible au ho i ies by
shoo ing he whole pa akee popula ion. The e o e, he isk o monk pa akee expansion
om u ban habi a s, hanks o he widesp ead dis ibu ion o whi e s o ks, should be
conside ed when designing managemen s a egies o his highly in asi e species,
which is g owing exponen ially in Medi e anean coun ies (Pos igo e al. 2019),
including Spain, Po ugal, F ance, I aly, G eece, Mo occo, and Is ael, whe e he wo
species coexis (Hagemeije and Blai 1997; Pos igo e al. 2019). Managemen ac ions
could be equi ed in he case u al popula ions o monk pa akee s would cause
signi ican impac s (Pos igo e al. 2019), al hough hese ac ions usually show low
suppo om he socie y (Shackle on e al. 2019), e en lowe when dealing wi h
cha isma ic species such as pa akee s (C owley e al. 2019). Consequen ly, mo e
esea ch and awa eness campaigns a e necessa y no only o know he ac ual magni ude
o he impac s de i ed om in asi e species (Blackbu n e al. 2010) bu also o make
managemen ac ions e ec i e. In ou case, ac ions only ocus on he a oidance o monk
pa akee s nes ing in whi e s o k nes s may no be e icien . On he one hand, an i-
nes ing de ices ins alled in pylons o whi e s o ks do no p e en hei nes ing, e en
a e g ea managemen e o s (Ma ica o e al. 2016). On he o he hand, nes emo al
would be no e icien because bo h species show s ong ideli y o hei nes ing
subs a es and o en ebuild hei nes s e y soon (Sol e al. 1997; Ve ga a e al. 2006;
P ue -Jones e al. 2007; Bu ge and Goch eld 2009). Con a ily, ac ions aimed o
imp o e p eda o popula ions as biological con olle s, should be e ec i e o hal he
sp ead o his species in o u al a eas in he long e m.
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Sec ion 2: Chap e V
149
This s udy assesses he nes ing associa ion be ween an in asi e bi d, he monk
pa akee , and a na i e bi d species, he whi e s o k, showing a commensalism
ela ionship in which pa akee colonies associa ed wi h s o k nes s bene i om he
e ec i e an ip eda o y de ence o s o ks. This associa ion was mo e likely in u al
a eas, whe e p eda ion p essu e is highe han in nea by u ban ones, assis ing hus he
sp ead o monk pa akee s ac oss he u al en i onmen . Mo eo e , he abandonmen o
pa akee colonies a e he p e ious nes abandonmen o associa ed s o ks, as well as
hei di e en beha iou al eac ions agains ap o s when associa ed wi h s o ks, sugges
ha pa akee s ha e a s ong dependence o hei hos s. Howe e , his p o ec i e
associa ion is limi ed as pa akee colonies also a oided high densi ies o b eeding
ap o s in he s udy a ea. Wi hou he acili a ion p o ided by s o ks, he bio ic
esis ance om he ap o communi y p e en s he in asion success o pa akee s. Fu u e
s udies a e needed o assess he complexi y o in e ac ions be ween in asi e species and
he ecipien communi y, which may be undamen al o de elop e ec i e managemen
plans agains biological in asions.
ACKNOWLEDGEMENTS
We hank J. López o helping du ing ieldwo k and he owne s o Case ío del Hena es
and El Piul o allowing us o access hei p i a e lands ( he es o he s udy a ea was
access- ee). Ó. F ías and F. Ma ínez also helped in he ieldwo k and con ibu ed da a
on he loca ion o nes s o ap o s, s o ks and pa akee s. Logis ic and echnical suppo
we e p o ided by Doñana ICTS-RBD. This s udy was suppo ed by he Se e o Ochoa
P og am (SVP-2014-068732) and Ac ion COST ES1304 (Pa o Ne ). Logis ical and
echnical suppo o ieldwo k was p o ided by Doñana ICTS-RBD.
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Sec ion 2: Chap e V
150
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using he ose‐ inged pa akee (Psi acula k ame i) as a case s udy. Di e si y and
Dis ibu ions, 15(2), 310-318.
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Py hu a c uen a a Neo opical 1 0 0
Py hu a on alis Neo opical 0 0 1
Py hu a g iseipec us Neo opical 1 0 0
Py hu a molinae Neo opical 1 0 1
Py hu a pe la a Neo opical 1 0 0
Thec oce cus acu icauda us Neo opical 0 1 1
T ichoglossus haema odus Aus alasia 0 0 1
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Table S2. Lis o plan species whe e pa o s was ed ood. We show he species, amily
and he numbe o imes we de ec ed ood was ing a he species.
Species Family Numbe o obse a ions
Acacia angus issima Fabaceae 16
Acacia a oma Fabaceae 1
Acacia au iculi o mis Fabaceae 12
Acacia ka oo Fabaceae 13
Acacia mea nsii Fabaceae 10
Acacia melanoxylon Fabaceae 11
Acaena splendens Rosaceae 1
Ac ocomia o ai A ecaceae 65
Aesculus hippocas anum Sapindaceae 1
Agonand a b asiliensis Opiliaceae 2
Ailan hus al issima Sima oubaceae 2
Aiphanes minima A ecaceae 2
Aizoon cana iense Aizoaceae 1
Albizia ca bona a Fabaceae 1
Albizia guachapele Fabaceae 2
Albizia julib issin Fabaceae 1
Albizia p oce a Fabaceae 9
Albizia saman Fabaceae 19
Anaca dium excelsum Anaca diaceae 3
Anaca dium occiden ale Anaca diaceae 9
Anadena he a colub ina Fabaceae 2
Anisocappa is speciosa Cappa idaceae 4
Annona mu ica a Annonaceae 2
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A achis hypogaea Fabaceae 36
A auca ia angus i olia A auca iaceae 303
A auca ia a aucana A auca iaceae 34
A auca ia bidwillii A auca iaceae 51
A auca ia cunninghamii A auca iaceae 5
A bu us unedo E icaceae 7
A disia escallonioides P imulaceae 4
Aspa agus a bo escens Aspa agaceae 2
Aspidospe ma queb acho-blanco Apocynaceae 2
As onium u undeu a Anaca diaceae 1
A alea bu y acea A ecaceae 2
A alea phale a a A ecaceae 0
A alea p inceps A ecaceae 3
A alea speciosa A ecaceae 9
A alea o ai A ecaceae 1
A e hoa ca ambola Oxalidaceae 3
Azadi ach a indica Meliaceae 2
Bac is gasipaes A ecaceae 1
Bambusa ulga is Poaceae 4
Be ula pendula Be ulaceae 4
B achychi on populneus Mal aceae 3
B osimum alicas um Mo aceae 1
Bucida buce as Comb e aceae 4
Bu se a sima uba Bu se aceae 3
Bu se a omen osa Bu se aceae 2
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By sonima c assi olia Malpighiaceae 7
By sonima pachyphylla Malpighiaceae 34
By sonima e basci olia Malpighiaceae 5
Cajanus cajan Fabaceae 1
Callica pa ampla Lamiaceae 3
Calophyllum an illa um Calophyllaceae 6
Ca apa guianensis Meliaceae 2
Ca duus hoe me i As e aceae 112
Ca ya illinoinensis Juglandaceae 7
Cassia ja anica Fabaceae 2
Cas anea sa i a Fagaceae 3
Cas illa elas ica Mo aceae 1
Casua ina c is a a Casua inaceae 1
Casua ina equise i olia Casua inaceae 11
Ca alpa bignonioides Bignoniaceae 7
Cec opia pel a a U icaceae 1
Cec opia sch ebe iana U icaceae 9
Ced ela odo a a Meliaceae 15
Ced us a lan ica Pinaceae 1
Ced us libani Pinaceae 1
Ceiba pen and a Mal aceae 28
Ceiba pen hand a Mal aceae 1
Ceiba speciosa Mal aceae 9
Cel is aus alis Ulmaceae 45
Cen au ea sols i ialis As e aceae 46
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Ce as ium glome a um Ca yophyllaceae 3
Ce a onia siliqua Fabaceae 5
Ce cis siliquas um Fabaceae 67
Chenopodium mu ale Chenopodiaceae 4
Cipadessa bacci e a Meliaceae 1
Ci ha exylum donnell-smi hii Ve benaceae 1
Ci ha exylum spinosum Ve benaceae 7
Ci us × au an ium Ru aceae 2
Ci us × sinensis Ru aceae 37
Ci us au an i olia Ru aceae 1
Clusia p a ensis Clusiaceae 1
Cnidoscolus que ci olius Eupho biaceae 1
Coccoloba ca acasana Polygonaceae 1
Coccoloba u i e a Polygonaceae 52
Cocos nuci e a A ecaceae 7
Co ea a abica Rubiaceae 5
Commipho a lep ophloeos Bu se aceae 1
Compsoneu a sp ucei My is icaceae 1
Condalia buxi olia Rhamnaceae 8
Co dia alba Bo aginaceae 1
Co dia collococca Bo aginaceae 5
Co dia cymosa Bo aginaceae 1
Co ymbia cla ksoniana My aceae 7
Co ymbia essalla is My aceae 2
Couepia g andi lo a Ch ysobalanaceae 15
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Couepia polyand a Ch ysobalanaceae 1
Couma mac oca pa Apocynaceae 2
C a aegus monogina Rosaceae 1
C o on magdalenensis Eupho biaceae 1
Cupania ame icana Sapindaceae 3
Cup essus sempe i ens Cup essaceae 87
Cu a ella ame icana Dilleniaceae 16
Cyanophalla e usa Cappa idaceae 2
Cype aceus sp. Cype aceae 1
Cy a ia ha io i Cy a iaceae 2
Delonix egia Fabaceae 15
Dend opanax la ilobus A aliaceae 1
Do yalis ca a Salicaceae 47
D acaena d aco Aspa agaceae 2
D imys win e i Win e aceae 1
Elaeis guianensis A ecaceae 1
Emmo um ni ens Icacinaeae 7
E iobo ya japonica Rosaceae 75
E y h ina ca a Fabaceae 21
E y h ina usca Fabaceae 16
E y h ina poeppigiana Fabaceae 10
E y h oxylum deciduum E y h oxylaceae 6
Eucalyp us camaldulensis My aceae 45
Eucalyp us globulus My aceae 35
Eucalyp us minia a My aceae 5
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Euc yphia co di olia Cunoniaceae 1
Eugenia uni lo a My aceae 38
Ficus benghalensis Mo aceae 19
Ficus ca ica Mo aceae 115
Ficus insipida Mo aceae 2
Ficus ly a a Mo aceae 27
Ficus mac ophylla Mo aceae 57
Ficus maxima Mo aceae 1
Ficus mic oca pa Mo aceae 106
Ficus nymphaei olia Mo aceae 1
Ficus eligiosa Mo aceae 9
Ficus ubiginosa Mo aceae 16
Ficus sp. Mo aceae 1
Ficus s uhlmannii Mo aceae 6
Ficus igona a Mo aceae 2
Flacou ia ine mis Salicaceae 5
F axinus angus igolia Oleaceae 33
Geo oea deco icans Fabaceae 1
Gledi sia iacan hos Fabaceae 18
Gli icidia sepium Fabaceae 9
Gmelina a bo ea Lamiaceae 15
G e illea glauca P o eaceae 3
G e illea obus a P o eaceae 91
Gua ea guidonia Meliaceae 27
Guazuma ulmi olia Mal aceae 1
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Hanco nia speciosa Apocynaceae 3
Hand oan us ch ysan hus Bignoniaceae 2
Helian hus annuus As e aceae 99
Helioca pus sp. Mal aceae 1
Hibiscus osa-sinensis Mal aceae 4
Hi ella ugosa Ch ysobalanaceae 2
Hymenaea cou ba il Fabaceae 1
Hymenaea s igonoca pa Fabaceae 17
Hypochae is adica a As e aceae 2
Inga adenophylla Fabaceae 1
Inga chocoensis Fabaceae 1
Inga cylind ica Fabaceae 3
Inga edulis Fabaceae 1
Inga euilleei Fabaceae 1
Inga punc a a Fabaceae 1
Inga spec abilis Fabaceae 11
Jaca anda mimosi olia Bignoniaceae 39
Ja opha hie onymi Eupho biaceae 11
Ja opha humbold iana Eupho biaceae 7
Ja opha mollisima Eupho biaceae 4
Juglans egia Juglandaceae 3
Junipe us p oce a Cup essaceae 1
Junipe us i giniana Cup essaceae 9
Lage s oemia indica Ly h aceae 2
Lage s oemia speciosa Ly h aceae 9
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Lau eliopsis philippiana A he ospe ma aceae 2
Leucaena leucocephala Fabaceae 2
Ligus um japonicum Oleaceae 38
Ligus um lucidum Oleaceae 13
Li is ona chinensis A ecaceae 165
Luehea seemannii Mal aceae 4
Magonia pubescens Sapindaceae 2
Malus domes ica Rosaceae 17
Mangi e indica Anaca diaceae 96
Mangi e a zeylanica Anaca diaceae 1
Manilka a biden a a Sapo aceae 3
Map ounea guianensis Eupho biaceae 2
Ma an hes panamensis Ch ysobalanaceae 7
Mau i ia lexuosa A ecaceae 46
Mau i iella a ma a A ecaceae 1
May enus boa ia Celas aceae 5
Melaleuca ci ina My aceae 17
Melia azeda ach Meliaceae 311
Melicoccus bijuga us Sapindaceae 10
Miconia impe iola is Melas oma aceae 10
Misodend um linea i olium Misodend aceae 2
Mo us alba Mo aceae 84
Mo us nig a Mo aceae 127
Mou i i pusa Melas oma aceae 2
Musa × pa adisiaca Poaceae 3
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My ceugenia exsucca My aceae 1
My us communis My aceae 16
Neea hei e a Nyc aginaceae 16
Nephelium lappaceum Sapindaceae 1
No ho agus dombeyi No ho agaceae 3
No ho agus obliqua No ho agaceae 10
Och oma py amidale Mal aceae 1
Olea eu opaea Oleaceae 49
Olea eu opaea syl es is Oleaceae 21
Onopo dum acan hium As e aceae 1
O yza sa i a Poaceae 11
Pachi a quina a Mal aceae 10
Pa kia biglandulosa Fabaceae 4
Pa kinsonia aculea a Fabaceae 3
Pa kinsonia p aecox Fabaceae 1
Pa odiodend on ma gini illosum Pic odend aceae 2
Passi lo a edulis Passi lo aceae 2
Pa elli olia pa ella is Ama an haceae 5
Pa elli olia p ocumbens Ama an haceae 1
Pel opho um p e oca pum Fabaceae 1
Pe sea ame icana Lau aceae 11
Phala is cana iensis Poaceae 3
Phoenix cana iensis A ecaceae 260
Phoenix dac yli e a A ecaceae 437
Phy olacca dioica Phy olaccaceae 33
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