UNCORRECTED PROOF
O iginal a icle
B eakdown o he species-a ea ela ionship in
exo ic bu no in na i e o es pa ches
Tibo Magu a
a
, And a
´sBa
´ldi
b,
*,Ro
´be Ho a
´ h
a
a
Di ec o a e o he Ho oba
´gy Na ional Pa k, Sumen u. 2, Deb ecen H-4024, Hunga y
b
Animal Ecology Resea ch G oup o he Hunga ian Academy o Sciences and he Hunga ian Na u al His o y Museum,
Ludo ika e
´ 2, Budapes H-1083, Hunga y
a icle in o
A icle his o y:
Recei ed 14 Sep embe 2007
Accep ed 23 No embe 2007
Published online -
Keywo ds:
Bi ds
Fo es specialis species
Fo es y p ac ice
Hunga y
Gene alis species
Habi a s uc u e
abs ac
We s udied he pa e n o bi d species ichness in na i e and exo ic o es pa ches in
Hunga y. We hypo he ized ha species-a ea ela ionship will depend on o es na u al-
ness, and on he habi a specializa ion o bi d species. The e o e, we expec ed s ong
species-a ea ela ionship in na i e o es pa ches and o es bi d species, and weake ela-
ionship in exo ic o es pa ches con aining gene alis species. We censused b eeding
passe ine bi d communi ies h ee imes in 13 o es pa ches wi h only na i e ee species,
and 14 wi h only exo ic ees in Eas e n Hunga y in 2003. Al hough mos bi d species (92%)
o he o al o 41 species occu ed in bo h exo ic and na i e o es s, he species-a ea ela-
ionship was signi ican o o es specialis , bu no o gene alis species in he na i e
o es s. No ela ionship be ween bi d species and a ea was ound o ei he species g oup
in he o es wi h exo ic ee species. The compa ison o na i e e sus exo ic o es pa ches
o simila sizes e ealed ha only la ge (>100 ha) na i e o es s ha bo highe bi d species
ichness han exo ic o es s o he o es specialis bi d species. The e is no di e ence be-
ween small and medium o es pa ches and in ichness o gene alis species. Thus, he
species-a ea ela ionship may diminish in a chipelago o exo ic habi a pa ches and/o
o habi a gene alis species; his esul suppo s he wa ning ha he ex ension o exo ic
habi a s ha e been signi ican ly con ibu ing o he decline o na u al communi y pa e ns.
ª2008 Published by Else ie Masson SAS.
1. In oduc ion
Fo es s a e in aluable o human beings. They a e impo an
o ec ea ion and human well being, and p o ide nume ous
ecosys em se ices, and a e i al o he main enance o he
majo i y o biological di e si y on Ea h (Lacaze, 2000; Di zo
and Ra en, 2003; Ozanne e al., 2003; Lewis, 2006). These oles
need o be conside ed when we y o manage o es s. Fo ex-
ample, he use o wood o hea ing and imbe in cons uc ion
o pape indus y needs as g owing ees, plan ed in a egula
pa e n o easy managemen , wi hou o he ee o sc ub
species. These plan a ions o en a e o non-na i e species
and hus a e in e io o ec ea ion, and inapp op ia e o
he main enance o na i e biodi e si y (Koch and Sko sgaa d,
1999; Ca nus e al., 2006; Gen y e al., 2006). Fo example, in
Hunga y 20% o o es co e is he black locus (Robinia
pseudoacacia), o iginally om No h Ame ica, and 15% is black
pine (Pinus nig a), a Eu opean species, bu no na i e o
* Co esponding au ho a : Tel.: þ36 1 210 1075; ax: þ36 1 334 2785.
E-mail add esses: [email protected],[email p o ec ed] (A. Ba
´ldi).
a ailable a www.sciencedi ec .com
jou nal homepage: www.else ie .com/loca e/ac oec
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1146-609X/$ – see on ma e ª2008 Published by Else ie Masson SAS.
doi:10.1016/j.ac ao.2007.11.007
ac a oecologica xxx (2008) 1–8
ACTOEC2410_p oo  25 Feb ua y 2008  1/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007
UNCORRECTED PROOF
Hunga y (Ma
´ ya
´s, 1997). The e a e simila pa e ns o o he
coun ies (e.g., EEA, 2006).
Wha a e he e ec s o exo ic (i.e. in oduced o non-
na i e) ees and o es s on na i e wildli e? Bi d species
usually p e e na i e habi a s o e exo ic pa ches (Ramos,
1996; O ega e al., 2006). Na i e bushes a e supe io o aging
si es o bi ds (F ench e al., 2005). Nes ing on exo ic bushes
in u ban pa ks esul ed in highe a e o nes ailu e han
nes ing on na i e bushes (Schmid and Whelan, 1999;
Bo gmann and Rodewald, 2004). Exo ic o es plan a ions
can lu e se ling bi ds in o such subop imal habi a , whe e
nes ailu e is highe han in na i e o es s (Remes, 2003).
Knowledge o he e ec s o exo ic ee plan a ions on bi ds
a he communi y le el is limi ed. Se e al s udies aimed o
compa e communi ies in na i e and exo ic o es s, using
simple pa ame e s. Species ichness and abundance is less
a iable han species composi ion in na i e e sus exo ic
o es s (Hausne e al., 2002; Johnson and F eedman, 2002;
S e e ding and Leuschne , 2002; Bakke and Higgins, 2003),
al hough some s udies ha e ailed o de ec any di e ences
(Donald e al., 1998; Fleishman e al., 2003; Wilson e al.,
2006). The unde lying mechanism o communi y di e ences
be ween na i e and exo ic o es s may be he p eda ion p es-
su e on a ian b oods (Ba be e al., 2001; Ca ignan and Villa d,
2002), o he selec i e habi a p e e ences o species (Le ne
and S au e , 1998). Howe e , we ha e been unable o ind
any compa ison o he species-a ea ela ionship (SAR) in ex-
o ic e sus na i e o es s. This is su p ising, because SAR is
a basic ule o ecology, s a ing ha species ichness inc eases
wi h inc easing sample a ea (Rosenzweig, 1995; Ba
´ldi and
McCollin, 2003; D aka e e al., 2006). Thus, he impo an
ques ion is whe he a undamen al ecological ela ionship is
al e ed by one o he mos peculia and pe asi e human
ac i i ies o mode n imes – o in oduce plan species o a eas
ou side hei na i e anges. In his s udy we in es iga ed i
exo ic o es s ha bou ewe species han na i e pa ches. We
compa ed he species-a ea ela ionship o bi d assemblages
in na i e e sus exo ic o es pa ches in Eas e n Hunga y. We
hypo he ized ha SAR will depend on o es na u alness,
and on he habi a specializa ion o species. Na u alness p ob-
ably ac s ia he e ogenei y, which is highe in na i e han in
exo ic o es s (Thompson e al., 2003; Ba ha e al., 2006). Fo -
es pa ches a e islands o o es specialis species, bu less so
o gene alis species, which may occu in he su ounding
landscape. This may mask he gene al species-a ea ela ion-
ship (e.g. Magu a e al., 2001). The e o e, we expec signi ican
SAR in na i e o es pa ches and o es specialis bi d species,
and weak, i any in exo ic o es pa ches wi h gene alis spe-
cies. Fu he , we expec simila species ichness alues in
small na i e and exo ic o es pa ches, whe e no eal in e io
habi a is a ailable. Howe e , di e en species ichnesses a e
expec ed in la ge pa ches due o di e ence in he species-a ea
ela ionships. Such inding may ha e impo an na u e con-
se a ion consequences o he main enance o biodi e si y.
2. S udy a ea and me hods
The s udy a ea is loca ed in Eas e n Hunga y on he Sza ma
´ -
Be eg plain (Fig. 1)(N48
0505500,E22
3002200). The plain is
co e ed by pas u es and ag icul u al land wi h sca e ed
o es pa ches. The na u al ege a ion o Cen al-Eu opean
plains is o es , bu he millennia o human ac i i ies (e.g., ca -
le g azing) modi ied i in o p ima ily g assland a eas wi h
small o es pa ches (S ando a
´ and P imack, 2001). Roughly
80% o he egion is a mland. The su oundings was simila
o all s udied pa ches, g assland and/o a able ields. Na i e
and exo ic ee species we e p esen in he pa ches. We chose
13 o es pa ches wi h only deciduous na i e ee species
(Que cus obu ,Ca pinus be ulus,Que cus pe aea,Populus canes-
cens,Populus alba, wi h a ew Ace campes e,Salix alba and F ax-
inus angus i olia), and 14 wi h only deciduous exo ic ee
species (Robinia pseudoacacia,Populus canadensis, wi h ew indi-
iduals o Que cus ub a,Ace negundo,Salix ma sudana,Amo -
pha u icosa and F axinus pennsyl anica). The age o s udied
o es pa ches was 35–70 yea s. Th ee a ea ca ego ies o o es
agmen s we e es ablished: small (<10 ha), medium
(10 <100 ha), and la ge (>100 ha) (Table 1). Wi hin size class
agmen s wi h na u al ee composi ion and hose wi h
exo ic ees we e dis inguished. The e was no signi ican
di e ence be ween he mean a ea o hese wo g oups wi hin
a ea ca ego y (
7
¼0.289, p¼0.781 o small o es s,
7
¼
0.812, p¼0.443 o medium o es s, and
7
¼2.853, p¼
0.064 o la ge o es s). Fo es pa ch he e ogenei y was
es ima ed by eye as pe cen co e o sh ubs and numbe o
nes holes wi hin he censused a eas.
B eeding bi d communi ies we e censused in he o es
pa ches in he b eeding season o 2003. Th ee censuses we e
ca ied ou du ing he season (Ap il, May and ea ly June),
only unde good wea he condi ions (no wind o ain), om
sun ise o 9 a.m. (Moska
´ , 1987). We applied a s anda d poin
census echnique (100 m adius, 5 min census ime); all bi ds
seen o hea d we e eco ded. Sampling e o was s anda d-
ized o all pa ches – since many pa ches we e only a ew
hec a es la ge, one poin pe pa ch a such pa ches was
possible o census. We dis inguished be ween o es specialis
and habi a gene alis species based on li e a u e da a (Snow
and Pe ins, 1998), conside ing local condi ions.
The species-a ea ela ionship was es ablished wi h he
mos equen ly used log-log ans o med model (Rose-
nzweig, 1995), using indi idual pa ch a eas (no he ca e-
go ies) in he calcula ion. The numbe o o es specialis
and gene alis bi d species be ween he wo o es ypes
(na i e and exo ic) wi h simila size we e examined by e-
pea ed measu e analysis o a iance (ANOVA). Fo es ype
(na i e o exo ic) was conside ed as ac o and he ime o
he coun ing (Ap il, May and June) we e used as epea ed
measu es. The da a we e no malized by log(xþ1) ans o -
ma ion. When he esul s o he ANOVA showed ha he e
was di e ence in he species ichness among he o es
ypes, his was es ed by a Tukey- ype mul iple compa ison
(Sokal and Rohl , 1981). The analyses we e ca ied ou using
he SPSS-PC p og am (SPSS, 1999).
3. Resul s
Sh ub co e did no di e signi ican ly be ween na i e and
exo ic o es pa ches (
7
¼0.324, p¼0.755 o small o es s,
7
¼0.040, p¼0.969 o medium o es s, and
7
¼0.828,
ac a oecologica xxx (2008) 1–82
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ACTOEC2410_p oo  25 Feb ua y 2008  2/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007
UNCORRECTED PROOF
p¼0.435 o la ge o es s). The e we e mo e nes holes in
na i e o es pa ches compa ed o he exo ic pa ches in small
and la ge o es s, bu no in he medium size ca ego y
(
7
¼3.653, p¼0.008 o small o es s,
7
¼1.005, p¼0.348 o
medium o es s, and
7
¼2.543, p¼0.038 o la ge o es s).
Al oge he 41 bi d species we e obse ed du ing he cen-
suses. Mos species (92%) we e obse ed in bo h he na i e
and he exo ic o es pa ches (Appendix). The species-a ea
ela ionship was signi ican ly posi i e in he na i e o es
pa ches o o es specialis species, and posi i e bu no
signi ican o gene alis species (Table 2). No species-a ea
ela ionship we e ound o bi d assemblages in exo ic o es
pa ches (Table 2).
Compa ing bi d species ichness be ween na i e and exo ic
o es pa ches o simila size classes e ealed ha he small
and medium pa ches we e no signi ican ly di e en o ei he
o es specialis o gene alis species. In la ge o es s, how-
e e , bi d species ichness was signi ican ly highe in na i e
han in exo ic o es s o o es specialis species, bu no o
gene alis species (Table 3). This ela ionship be ween he
na i e and exo ic o es pa ches seems o be s able, a leas
wi hin season, because he in e ac ion e m o ime * na u al-
ness was on no occasion signi ican (Table 3). The ime e ec
alone indica ed signi ican decline in species numbe om
Ap il o June in he small o es pa ches ( o bo h o es and
gene alis species), and o o es specialis species in medium
o es pa ches. The numbe o gene alis species did no
change wi hin season in medium-sized o es pa ches, and
none o he g oups declined o e he season in he la ge o es
pa ches (Table 3).
Fig. 1 – Loca ion o he s udy a ea in Eas e n Hunga y. T iangles ep esen o es pa ches o exo ic ee species and ci cles
na i e ee species. The size o he ma k ep esen s h ee a ea ca ego ies o he pa ches (<10 ha, 10–100 ha, >100 ha).
Table 1 – Cha ac e is ics o agmen s and he numbe o obse ed bi d species in h ee size g oups o na i e and exo ic
o es pa ches in Eas e n Hunga y
Small Medium La ge
Na i e Exo ic Na i e Exo ic Na i e Exo ic
Numbe o agmen s 5 4 4 5 4 5
A ea (ha) s.d. 7.1 1.86 6.6 2.92 41.0 28.94 58.7 34.99 425.0 206.16 130.0 18.71
Numbe o nes holes s.d. 697 344 55 49 465 447 204 336 725 284 172 351
Co e o sh ubs (%) s.d. 44 27.9 38 32.3 33.75 31.5 33 24.4 46 35.0 28 31.2
Numbe o obse ed o es
specialis species
12.2 1.10 10.5 1.73 13.8 4.99 14.0 5.61 16.5 1.26 12.4 2.19
Numbe o obse ed
gene alis species
3.2 1.64 3.3 2.06 1.8 1.50 3.0 1.58 1.8 0.96 4.0 1.58
ac a oecologica xxx (2008) 1–8 3
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ACTOEC2410_p oo  25 Feb ua y 2008  3/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007
UNCORRECTED PROOF
4. Discussion
We s udied bi d communi ies in a landscape wi h pa ches o
na u al and exo ic ee species. The majo i y o he landscape
was a mland (a able ields and g asslands), wi h sca e ed
o es pa ches, including he s udied pa ches. Al hough i is
known ha di e en landscape ma ix in luences species di-
e si y e en in simila o es pa ches (Lindenmaye e al.,
2002; Wa son e al., 2005), he e we had he same landscape
ype as ma ix. The e o e, we supposed ha he ma ix e ec
was simila o all pa ches, hus, did no bias he esul s.
The e was no di e ence in bi d species composi ion o
na u al e sus exo ic o es pa ches. In a s udy o bi ds in
ee plan a ions e sus na i e sh ub pa ches in he Nege ,
Is ael, Shocha e al. (2001) ound only a small o e lap o
bi d species. P obably he di e ence be ween exo ic and
na i e pa ches in ou s udy was no la ge enough o exclude
bi d species, only o in luence he equency o occu ence.
The communi ies showed di e en esponses o a ea; he spe-
cies-a ea ela ionship explained he numbe o bi d species
(i.e. signi ican posi i e species-a ea ela ionship) in na i e
o es s, bu no in exo ic o es pa ches. A simila pa e n
was ound by Shocha e al. (2001) o na i e sc ub agmen s
e sus plan ed o es s; bi d species ichness depended on a ea
in he o me , bu no in he la e . San os e al. (
Q1 2006)
compa ed na i e oak and ma u e pine plan a ions in Spain,
and hey ound simila bi d species ichness in na i e and
plan a ion a chipelagoes, and di e en species-a ea ela ion-
ship, jus in his s udy. Howe e , all species-a ea eg ession
models we e signi ican in hei s udy (San os e al., 2006).
The di e ence be ween na i e and exo ic o es s was
mo e p onounced, i species we e classi ied acco ding o
hei speci ici y o he o es habi a . Species ichness o
gene alis s was no ela ed o o es a ea a all. The species
– a ea ela ionship o o es specialis bi d species was no
signi ican in exo ic o es pa ches, bu was signi ican
(posi i e) in na i e pa ches. The clea di e ence be ween
he esponse o specialis and gene alis (including exo ic)
species o agmen a ion was desc ibed o se e al axa, in-
cluding bi ds (McCollin, 1993; Ge maine e al., 1998), plan s
(Abbo , 1992; Bakke and Higgins, 2003) and in e eb a es
(Magu a e al., 2001; Os e ga d and Eh len, 2005; Ouin
e al., 2006). Mo e gene ally, species ai s a e known o
con ound he SAR (Ewe s and Didham, 2006).
The e may be wo po en ial mechanisms o explain he
p esence o SARin na i e, bu no in exo ic o es pa ches.Fi s ,
he e is usually a basic di e ence be ween exo ic and na i e
o es s in spa ial he e ogenei y; na i e o es sa e mo e he e o-
geneous (Ma
´ ya
´s, 1997; Thompson e al., 2003). We also ound
some indica ions o such a end (mo e nes holes in na i e o -
es s), al hough o he s we e in ou case no s a is ically signi i-
can (sh ub co e ). Habi a he e ogenei y and species ichness
ha e a posi i e co ela ion (Tews e al., 2004), hus, he mo e
he e ogeneous and complex habi a s uc u e o na i e o es s
may p omo e o es specialis bi d species a he han gene al-
is species (MacNally e al., 2000). Second, na i e o es s a e
mo e island-like pa ches, because hei complex s uc u e is
mo e di e en om he su ounding landscape, han he ex-
o ic o es s wi h a simple s uc u e. The e o e, o es specialis
species a e p obably es ic ed o he na i e o es ‘‘isola es’’,
wi h he subsequen SAR, while exo ic o es s migh no unc-
ion as isola es. This landscape e ec may also be esponsible
o he absence o a SAR in some cases (Es ades and Temple,
1999; We he ed and Lawes, 2003; Lo
¨ ei e al., 2006).
Inc eased habi a agmen a ion esul s in he ela i e in-
c ease o edge habi a s due o he inc ease o edge/co e a io.
Edges a e a o ed by gene alis and/o ea ly successional spe-
cies (Ha is and Sil a-Lopez, 1992; Imbeau e al., 2003; Lo
¨ ei
e al., 2006), he e o e we expec ed a e e se ela ionship o
gene alis species wi h o es a ea: as o es a ea inc eases
he p opo ion o edges, and he gene alis s species, is de-
c easing. Al hough hese we e signi ican nei he o na i e
(R¼0.1) no o exo ic (R¼0.1) o es s, he ends o he e-
g ession coe icien s (Table 2) suppo his hypo hesis.
The SAR is a undamen al ule o ecology, bu i s ill su e s
om se e al biases (Ba
´ldi and McCollin, 2003). He e we dem-
ons a ed he key ole o na i e/exo ic species composi ion,
ha is he quali y o habi a s and he specialis /gene alis s
cha ac e o a ge species on he SAR (Lo
¨ ei e al., 2006). An
a chipelago o non-na i e habi a s, and/o non-na i e species
in he a chipelago may lead o he b eakdown o he SAR.
The age o o es pa ches may be ele an ac o in de e -
mining species ichness. In his s udy he age o pa ches
we e 35–70 yea s, ha none o hem we e old g ow h.
Humph ey (2005) and San os e al. (2006), o example, showed
ha 100–200 yea old plan a ions al eady con e ing subs an-
ial bene i s o many species. I is clea ha o he s udied o -
es a chipelago and landscape only la ge (>100 ha) o es
pa ches con aining na i e ee species can p ese e na u al
pa e ns. Such pa ches suppo o es specialis bi d species
du ing he whole b eeding season. We wa n, howe e , agains
using only species p esence in o es pa ches o pa ch e alu-
a ion: on ou s udy nea ly all he obse ed species (92%) we e
p esen in bo h na i e and exo ic o es pa ches. The e o e,
simple p esence-absence su ey may be misleading, because
i can no iden i y he supe io alue o na i e o es s. O he
s udies also highligh ed he subo dina e ole o exo ic ees
and bushes o nes ing, o aging and communi y assemblage
(Schmid and Whelan, 1999; Remes, 2003; Bo gmann and
Rodewald, 2004; F ench e al., 2005). This e idence suppo
he conclusion ha he expansion o exo ic ees and bushes
ia o es y p ac ice and ga dening will ha m na u al pa e ns
and he unde lying p ocesses, hence accele a ing he decline
o bi ds in isola ed o es s.
Table 2 – Equa ions o he species-a ea ela ionship o
o es specialis bi ds in pa ches o na i e and exo ic ees
in Eas e n Hunga y. Equa ions a e gi en o o es
specialis and gene alis species sepa a ely. As e isks
indica e signi ican ela ionship
Equa ion Rnp
Na i e o es s
Numbe o o es
specialis species
Y¼0.82 þ0.09X0.66 13 0.02*
Numbe o gene alis species Y¼0.46 0.01X0.07 13 0.84
Exo ic o es s
Numbe o o es
specialis species
Y¼0.76 þ0.09X0.40 14 0.16
Numbe o gene alis species Y¼0.47 þ0.03X0.10 14 0.73
ac a oecologica xxx (2008) 1–84
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ACTOEC2410_p oo  25 Feb ua y 2008  4/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007
UNCORRECTED PROOF
Acknowledgemen
We a e indeb ed o Duncan McCollin, Be
´la To
´ hme
´ e
´sz and
wo anonymous e e ees o commen s on p e ious e sions
o he pape . The esea ch was suppo ed by a g an o he
Hunga ian Scien i ic Resea ch Fund (OTKA esea ch g an
no. F61651, o T. M.), and he Di ec o a e o he Ho oba
´gy Na-
ional Pa k. T. M. and A. B. we e Bolyai Resea ch Fellows o he
Hunga ian Academy o Sciences.
Table 3 – Resul s o he epea ed measu es ANOVA o he bi d species ichness in deciduous o es s wi h simila size.
Na u alness o he o es s (na i e, exo ic) comp ised he ac o and he ime o he coun ing (Ap il, May and June in 2003)
we e used as epea ed measu es. Resul s o he Tukey es indica e which o es ca ego y di e s signi ican ly (p<0.05)
om he o he ; o example ‘Na i e >Exo ic’ indica es ha he species ichness was signi ican ly highe in he na i e
o es s han in he exo ic pa ches
Va iable Sou ce SS d MS FpTukey pos e io i es
Small o es s, Numbe o
o es specialis species
Wi hin-Subjec s E ec s
Time 0.130 2 0.065 18.100 0.000
Time Na u alness 0.004 2 0.002 0.558 0.585
E o 0.050 14 0.004
Be ween-Subjec s E ec s
Na u alness 0.013 1 0.013 0.653 0.446
E o 0.143 7 0.020
Small o es s, Numbe o
gene alis species
Wi hin-Subjec s E ec s
Time 0.150 2 0.075 4.830 0.025
Time Na u alness 0.043 2 0.021 1.382 0.283
E o 0.217 14 0.016
Be ween-Subjec s E ec s
Na u alness 0.028 1 0.028 0.294 0.604
E o 0.662 7 0.095
Medium o es s, Numbe o
o es specialis species
Wi hin-Subjec s E ec s
Time 0.080 2 0.040 7.235 0.007
Time Na u alness 0.005 2 0.003 0.474 0.632
E o 0.077 14 0.006
Be ween-Subjec s E ec s
Na u alness 0.006 1 0.006 0.072 0.797
E o 0.625 7 0.089
Medium o es s, Numbe o
gene alis species
Wi hin-Subjec s E ec s
Time 0.006 2 0.003 0.210 0.813
Time Na u alness 0.084 2 0.042 2.751 0.098
E o 0.213 14 0.015
Be ween-Subjec s E ec s
Na u alness 0.011 1 0.011 0.255 0.629
E o 0.297 7 0.042
La ge o es s, Numbe o
o es specialis species
Wi hin-Subjec s E ec s
Time 0.021 2 0.011 3.024 0.081
Time Na u alness 0.005 2 0.003 0.734 0.498
E o 0.050 14 0.004
Be ween-Subjec s E ec s
Na u alness 0.162 1 0.162 10.228 0.015 Na i e >Exo ic
E o 0.111 7 0.016
La ge o es s, Numbe o
gene alis species
Wi hin-Subjec s E ec s
Time 0.000 2 0.000 0.005 0.995
Time Na u alness 0.031 2 0.016 0.458 0.642
E o 0.478 14 0.034
Be ween-Subjec s E ec s
Na u alness 0.077 1 0.077 1.012 0.348
E o 0.536 7 0.077
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ACTOEC2410_p oo  25 Feb ua y 2008  5/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007

UNCORRECTED PROOF
Appendix
Lis o obse ed bi d species in axonomic o de om 27 o es pa ches in Eas e n Hunga y. To al numbe o obse a ions
du ing he h ee censuses in 2003 is gi en. Snow and Pe ins (1998) was ollowed o nomencla u e. As e isks indica e
o es specialis species
Small Medium La ge
Na i e Exo ic Na i e Exo ic Na i e Exo ic
Columbidae
Columba palumbus Woodpigeon 062300
S ep opelia u u Tu le Do e 774556
Upupidae
Upupa epops Hoopoe 000002
Picidae
Dend ocopos majo * G ea Spo ed Woodpecke 723689
Dend ocopos medius* Middle Spo ed Woodpecke 000010
Dend ocopos mino * Lesse Spo ed Woodpecke 002100
D yocopus ma ius* Black Woodpecke 101430
Jynx o quilla* W yneck 641339
Picus i idis* G een Woodpecke 001001
Mo acillidae
An hus i ialis T ee Pipi 6 4 3 12 2 2
Tu didae
E i hacus ubecula* Robin 12 4 8 12 8 5
Luscinia mega hynchos* Nigh ingale 10 18 4 13 3 6
Tu dus me ula* Blackbi d 14 9 11 12 20 8
Tu dus philomelos* Song T ush 100230
Syl iidae
Locus ella lu ia ilis* Ri e Wa ble 131209
Phylloscopus collybi a* Chi cha 9389128
Phylloscopus sibila ix* Wood Wa ble 6567121
Phylloscopus ochilus* Willow Wa ble 011020
Syl ia a icapilla* Blackcap 22 13 18 19 18 13
Syl ia cu uca Lesse Whi e h oa 010000
Muscicapidae
Muscicapa s ia a* Spo ed Flyca che 001221
Ficedula sp.* Flyca he sp. 000010
Aegi halidae
Aegi halos cauda us* Long- ailed Ti 001100
Pa idae
Pa us a e * Coal Ti 000030
Pa us cae uleus* Blue Ti 3236114
Pa us majo * G ea Ti 17 8 11 12 14 9
Pa us palus is* Ma sh Ti 000001
Si idae
Si a eu opaea* Nu ha ch 104232
Ce hiidae
Ce hia sp.* T eec eepe 10 848510
O iolidae
O iolus o iolus* Golden O iol 9 9 4 12 8 14
Laniidae
Lanius collu io Red-backed Sh ike 120304
Co idae
Ga ulus glanda ius* Jay 102321
S u nidae
S u nus ulga is S a ling 21 21 17 8 10 48
Passe idae
Passe mon anus T ee Spa ow 000100
F ingillidae
Ca duelis ca duelis Gold inch 020102
Ca duelis chlo is* G een inch 320227
Cocco h aus es cocco h aus es* Haw inch 6 13 6 7 11 4
F ingilla coelebs* Cha inch 29 23 28 24 33 37
Se inus se inus Se in 000001
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ACTOEC2410_p oo  25 Feb ua y 2008  6/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007
UNCORRECTED PROOF
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Appendix (con inued)
Small Medium La ge
Na i e Exo ic Na i e Exo ic Na i e Exo ic
Embe izidae
Embe iza ci inella Yellowhamme 5227210
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ACTOEC2410_p oo  25 Feb ua y 2008  7/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007
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ACTOEC2410_p oo  25 Feb ua y 2008  8/8
Please ci e his a icle in p ess as: Magu a, T. e al., B eakdown o he species-a ea ela ionship in exo ic bu no in na i e o es
pa ches, Ac a Oecolo. (2008), doi:10.1016/j.ac ao.2007.11.007