Accep ed Manusc ip
Ti le: Solidago canadensis impac s on na i e plan and
pollina o communi ies in di e en -aged old ields
Au ho : Annam´
a ia Fenesi Csongo I. V´
ag´
asi Monica
Beldean Ri a F¨
oldesi Le en e-P´
e e Kolcs´
a Julie Te esa
Shapi o Edina T¨
o ¨
ok Anik´
oKo
´
acs-Hos y´
anszki
PII: S1439-1791(15)00042-0
DOI: h p://dx.doi.o g/doi:10.1016/j.baae.2015.03.003
Re e ence: BAAE 50871
To appea in:
Recei ed da e: 27-8-2014
Re ised da e: 26-2-2015
Accep ed da e: 12-3-2015
Please ci e his a icle as: Fenesi, A., V´
ag´
asi, C. I., Beldean, M., F¨
oldesi, R., Kolcs´
a , L.-
P., Shapi o, J. T., T¨
o ¨
ok, E., and Ko ´
acs-Hos y´
anszki, A.,Solidago canadensis impac s
on na i e plan and pollina o communi ies in di e en -aged old ields, Basic and
Applied Ecology (2015), h p://dx.doi.o g/10.1016/j.baae.2015.03.003
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Solidago canadensis impac s on na i e plan and pollina o communi ies in di e en -1
aged old ields 2
3
Annamá ia FENESIa,b,*, Csongo I. VÁGÁSIc,d, Monica BELDEANb, Ri a FÖLDESIe, 4
Le en e-Pé e KOLCSÁRb, Julie Te esa SHAPIROe, , Edina TÖRÖKb, Anikó KOVÁCS-5
HOSTYÁNSZKIe 6
7
aDepa men o Ecology, Uni e si y o Deb ecen, Egye em é 1, H–4032 Deb ecen, Hunga y
8
bHunga ian Depa men o Biology and Ecology, Babeş-Bolyai Uni e si y, Republicii s ee 9
42, RO–400015 Cluj-Napoca, Romania 10
cE olu iona y Ecology G oup, Hunga ian Depa men o Biology and Ecology, Babeş-Bolyai 11
Uni e si y, Clinicilo s ee 5–7, RO–400006 Cluj-Napoca, Romania
12
dMTA-DE “Lendüle ” Beha iou al Ecology Resea ch G oup, Depa men o E olu iona y 13
Zoology, Uni e si y o Deb ecen, Egye em é 1, H–4032 Deb ecen, Hunga y 14
eMTA ÖK, Lendüle Ecosys em Se ices Resea ch G oup, Alko mány ú 2–4, H–2163 15
Vác á ó , Hunga y 16
School o Na u al Resou ces and En i onmen and Depa men o Wildli e Ecology and 17
Conse a ion, Uni e si y o Flo ida, Gaines ille, FL 32611, USA 18
19
*Co esponding au ho . Tel.: +40 744916952; ax: +40 264431858. 20
E-mail add ess: [email p o ec ed] 21
22
*Manusc ip
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Abs ac 23
24
Seconda y succession in o me a able ields (i.e. old ields) migh be al e ed by he 25
coloniza ion o in asi e alien species, wi h possible communi y-wide impac s, hinde ing he 26
abili y old ields o become species- ich communi ies. Howe e , he e ec s o in asi e 27
species on local communi ies ha e a ely been add essed in he ligh o seconda y succession. 28
The e o e, we s udied he impac o he highly in asi e Solidago canadensis on plan and 29
pollina o communi ies along a g adien o in asion se e i y in old ields wi h di e en ages 30
(1–20 yea s since las ploughing) in Sou he n T ansyl ania, Romania. We asked whe he he 31
in asion o S. canadensis causes shi s in (1) he composi ion and di e si y o plan 32
communi ies, and (2) pollina o communi ies along he successional g adien . Fu he , we 33
asked (3) o wha ex en he p esence o S. canadensis a ec ed lowe isi a ion o na i e 34
plan species by pollina o s. Acco ding o ou esul s, he in asion educed he na i e plan 35
species ichness h oughou succession, al hough he mos p o ound nega i e e ec on plan 36
di e si y and ege a ion na u alness was exe ed in olde successional communi ies. The 37
in asion o S. canadensis had a nega i e e ec on he abundance o bees i espec i e o he 38
old ield age; howe e , he e was no simila nega i e e ec on ho e lies. Na i e lowe s 39
expe ienced educed isi a ion by wild bees, honey bees and ho e lies due o he augmen ed 40
p esence o S. canadensis. The e o e, he in asion o his pe ennial plan species di e s he 41
ajec o y o ege a ion succession, al e s he mu ualis ic links be ween he na i e elemen s o 42
hese old ields, and causes a non-desi ed al e na i e s able s a es o be ins alled. 43
44
Keywo ds: Canada golden od; plan in asion; plan –pollina o in e ac ion; allow; se -aside; 45
Apis; Romania; Na u a 2000; biodi e si y conse a ion 46
47
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In oduc ion 48
49
O e he las ew decades, a ious social and economic changes ha e d i en he abandonmen 50
o ex ensi e a eas o o me ly a able lands in se e al pa s o he wo ld. I hese a eas a e le 51
alone and p opagule sou ces a e a ailable, seconda y succession akes place and di e se semi-52
na u al ege a ion can be o med in sho ime (S oa e e al. 2009). Fo me a able lands 53
(he ea e old ields) may suppo a mland biodi e si y and p o ide aluable ecosys em 54
se ices (e.g. pollina ion) a he beginning o seconda y succession (Tscha n ke, Ba á y & 55
Do mann 2011) and hei conse a ion alue may inc ease in he la e s ages (Co be 1995). 56
Howe e , eaching a s age o di e se seconda y ege a ion is no ee om challenges, as old 57
ields a e highly suscep ible o in asion by alien plan species (Ca o d e al. 2012). Non-58
na i e in asi e species a e likely o colonize he eshly abandoned c oplands because o 59
inc eased soil nu ien a ailabili y associa ed wi h p e ious e ilize applica ions, and limi ed 60
compe i ion o esou ces (S andish, C ame & Hobbs 2008). Longe -li ed alien species can 61
pe sis un il he la e s ages o succession, he e o e pe u bing na u al succession pa hways 62
and al e ing he pa e n o ege a ion eco e y (Meine s, Picke & Cadenasso 2002). By 63
p e en ing he es ablishmen o la e-successional species, alien plan in asion may cause he 64
in aded habi a s o emain in he ea ly successional s ages o a long ime (C ame , Hobbs & 65
S andish 2008). 66
The es ablishmen o in asi e plan s may in luence no only he na i e lo a and he 67
ajec o y o i s seconda y succession, bu can also ha e an impac on he ela ed in e eb a e 68
communi ies, such as pollina o insec s ( an Hengs um e al. 2014). Fo me s udies p esen ed 69
highly con o e sial esul s, epo ing ei he posi i e (Ba omeus, Vilà & San ama ía 2008) o 70
nega i e e ec s (de G oo , Kleijn & Jogan 2007; Mo oń e al. 2009) on pollina o abundance 71
and di e si y. In asi e plan s a e epo ed o in luence he abundance and di e si y o smalle , 72
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mos ly soli a y wild bees (Apoidea) o limi ed dispe sal abili y (Ga hmann & Tscha n ke 73
2002), as well as ho e lies (Sy phidae) and bumblebees, which gene ally ha e la ge 74
o aging anges (G eenlea e al. 2007). The e o e, in asi e species migh h ea en 75
biodi e si y no only a he local le el, bu a he landscape scale as well (Hejda, Pyšek & 76
Ja ošík 2009). Mo eo e , in asi e plan species can also ac indi ec ly on ecological 77
communi ies by weakening he mu ualis ic links be ween na i e plan s and hei pollina o s 78
(Aizen, Mo ales & Mo ales 2008) by lu ing pollina o s away om na i e plan s, ei he 79
pa ially (Baske , Eme y & Rudge s 2011) o comple ely (Ba omeus, Vilà & San ama ía 80
2008). This expansion o plan p e e ence o pollina ion can esul in ewe pollina o isi s, 81
pe asi e in e speci ic pollen ans e (Baske , Eme y & Rudge s 2011), and consequen ly, 82
dec eased ep oduc ion success and di e si y o he na i e lo a. Despi e hese possible 83
de imen al e ec s, he di ec e ec o in asi e plan species on seconda y succession o old 84
ields is ela i ely unde s udied (Flo y and Clay 2010), while he e ec s o in asi e species 85
on he pollina o communi ies along he successional ajec o y has so a ne e been s udied. 86
He e, we ocus on a la ge-scale biological in asion phenomenon in sou he n 87
T ansyl ania, Romania. La ge-scale abandonmen o a able lands has occu ed o e he las 88
wo decades due o a signi ican class o absen ee landowne s and he dec easing p o i abili y 89
o adi ional subsis ence ag icul u e (Fische , Ha el & Kuemme le 2012). Solidago 90
canadensis L. (Canada golden od) is he mos success ul in asi e colonize o hese old 91
ields, o en becoming dominan e en a he landscape scale. I s impac on he species 92
ichness o ascula plan s and pollina o s has al eady been s udied in o he coun ies (de 93
G oo , Kleijn & Jogan 2007; Mo oń e al. 2009). Howe e , hese s udies analysed only he 94
inal s age o in asion, when he in asi e species had al eady es ablished dominance in he 95
communi y, and compa ed his s age wi h non-in aded, semi-na u al communi ies, 96
ep esen ing only he wo con as ing ex emes o he in asion g adien . 97

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To ill hese gaps in ou knowledge ega ding he impac o in asion on bo h plan and 98
pollina o communi ies o e he cou se o seconda y succession on old ields along a 99
con inuous g adien o in asion, we ini ia ed a la ge-scale ield s udy. We aimed o answe he 100
ollowing ques ions: (1) Does he in asion o S. canadensis cause shi s in he composi ion 101
and di e si y o plan communi ies o e he cou se o seconda y succession o old ields? (2) 102
Does abundance and species ichness o pollina o s change due o he in asion o he old 103
ields? (3) To wha ex en does he p esence o S. canadensis a ec lowe isi a ion o na i e 104
plan species by pollina o s? To answe hese ques ions, we s udied he e ec o S. canadensis 105
densi y (0–80% ela i e co e ) in 36 old ields wi h a ious ages (1–20 yea s since he las 106
ploughing). 107
108
Ma e ials and me hods 109
The in asi e species 110
S. canadensis is a hizoma ous, pa ch- o ming pe ennial he b o he As e aceae. I has become 111
an excep ionally success ul in asi e species since i s in oduc ion o Eu ope in he 18 h 112
cen u y and is now widesp ead h oughou he con inen (Webe 1998). In addi ion o i s 113
p oli ic ege a i e p opaga ion (Meye & Schmid 1999), S. canadensis eleases chemicals 114
ha inhibi he g ow h, ge mina ion and su i al o na i e plan s (Abhilasha e al. 2008), and 115
change he soil composi ion by di e ing nu ien s and mine als (Zhang e al. 2009). 116
117
Loca ion and s udy si es 118
Ou s udy a ea was loca ed in he middle sec ion o he Tâ na a Ma e Valley in Sou he n 119
T ansyl ania, Romania (Fig. 1A). Clima e is classi ied as mode a e con inen al. Mean annual 120
ain all is app oxima ely 650 mm, while mean annual empe a u e is 8 °C (D ăgulescu 2003). 121
The egion has been designa ed as Na u a 2000 Si e o Communi y In e es (Sighişoa a-122
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Tâ na a Ma e Na u a 2000 si e, ROSCI0227) because 18 habi a s lis ed in he EU Habi a s 123
Di ec i e Annex I can be ound in his egion. 124
We examined he succession o ege a ion and pollina o communi ies in his si e 125
using a ch ono-sequence o old ields ep esen ing di e en s ages o succession (space o 126
ime subs i u ion, sensu Picke 1989). We chose old ields wi h a known abandonmen da e in 127
he icini y o ou illages (Bie an, Laslea, Malânc a , Richiş; Fig. 1A). Abandonmen he e 128
can be de ined as he cessa ion o ploughing, hough o he uses and ac i i ies such as mowing 129
o g azing may be p esen in he ields. These old ields may also be in equen ly bu ned by 130
locals. The 36 ields examined anged in age om 1 o 20 yea s since abandonmen . These 131
si es we e also chosen o ep esen he whole ange o S. canadensis abundance om highly o 132
less in aded a eas ( ela i e co e 0–80%). Mos si es we e qui e small (mean a ea: 1.3 ha, 133
ange: 0.08–2.7 ha). The yea o las ploughing, and cu en land-use p ac ices we e 134
de e mined by in e iewing landowne s, and based on ha in o ma ion, we ca ego ized he 135
old ields as mown, g azed o wi hou managemen . To de e mine whe he he si es had been 136
ecen ly bu ned, o no , we looked o local signs o i e (bu ned ees and sh ubs, incine a ed 137
li e o g ass ussocks) and asked local people. 138
139
Vege a ion su ey 140
Pe cen co e o ascula plan species wi h a esolu ion o 1% was isually es ima ed wi hin 141
h ee 4 × 4 m plo s pe si e (Fig. 1B). We assigned 0.5% o species wi h a co e smalle han 142
1% (species ep esen ed by one small indi idual o by seedlings only). The le el o in asion 143
was cha ac e ized in each plo by he ela i e co e o S. canadensis and a e aged o e he 144
h ee plo s a si e le el. The su eys we e conduc ed once pe si e in May–June 2012. 145
In o de o accu a ely depic composi ional changes du ing he cou se o succession, 146
we calcula ed he ollowing a iables o each si e: 147
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1. Na i e ascula plan species ichness and di e si y. Species ichness was calcula ed by 148
a e aging he numbe o species in he h ee plo s pe si e. The Shannon di e si y index o 149
each si e was calcula ed based on he p opo ional co e o each species. S. canadensis was 150
no included in he calcula ion. 151
2. Na u alness. All plan species we e assigned o one o he h ee na u alness g oups 152
acco ding o Sanda e al. (1983): (i) species o na u al and semi-na u al habi a s, he ea e 153
called “g assland species”, (ii) species common o bo h na u al and ude al communi ies, 154
he ea e “gene alis species”, and (iii) species o ude al communi ies, he ea e “ ude al 155
species”. The p opo ion o each ca ego y was calcula ed o each si e and used in subsequen 156
analyses. 157
3. Func ional guilds. The p opo ion o he main unc ional g oups was aken in o accoun by 158
dis inguishing (i) g aminoid species belonging o he Poaceae, Cype aceae o Juncaceae, (ii) 159
legume species o he Fabaceae and (iii) o bs, i.e. he baceous plan s belonging o o he 160
amilies. As he p esence o ee and sh ub species ne e exceeded 1% a e age co e a si e 161
le el, we excluded hose species om he calcula ion o his a iable. 162
163
Pollina o sampling 164
We sampled pollina o insec s in 22 si es (ou o 36 si es wi h bo anical su eys). We selec ed 165
hese si es o co e he whole ange o successional age (1 o 20 yea s) and low o high 166
densi ies o S. canadensis co e . The si es we e a leas 250 m om each o he . Pollina o s 167
we e sampled along wo 100 m ansec s pe s udy si e; each ansec assigned was a leas 15 168
m om he ield edge and 15 m apa om each o he (Fig. 1B). We su eyed he pollina o s 169
wice: i s du ing he peak lowe ing season o indigenous ege a ion (21–25 July 2012) and 170
second du ing he peak lowe ing o S. canadensis (19–23 Augus 2012). T ansec coun s 171
we e pe o med be ween 9 a.m. and 6 p.m. unde a ou able wea he condi ions wi h li le 172
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wind and ew clouds a mos . A pai o obse e s walked along each ansec o 20 min and 173
eco ded all insec s ac i ely pollina ing, lying o pe ching on he ege a ion. In he case o 174
ac i e pollina ion, he isi ed plan species was also eco ded. Pollina o s we e hand-ne ed 175
and p ese ed in 96% e hanol o la e iden i ica ion a he species le el. 176
We dis inguished ou pollina o g oups: honey bee (Apis melli e a L.), bumblebees 177
(Bombus spp.), o he wild bees (Apoidea: Colle idae, Meli idae, Halic idae, Megachilidae, 178
And enidae, Apidae excep Bombus spp. and honey bee), and ho e lies (Sy phidae). 179
Bumblebees and o he wild bees we e analysed sepa a ely because hese wo g oups ha e 180
di e en biological ai s in e ms o lo al equi emen s, lying abili ies and sociali y 181
(Ga hmann & Tscha n ke 2002; G eenlea e al. 2007; Michene 2007), and he e o e 182
di e en esponses o landscape and local scale en i onmen al condi ions we e expec ed. 183
Al hough he e a e some semi-social species and/o gene a among he collec ed bees (e.g. 184
some Halic us spp.), we use ‘soli a y bees’ in he cu en pape o wild bees o he han 185
bumblebees. 186
As he p esence o pollina o s depends on he pollen and nec a supply, we ga he ed 187
addi ional bo anical in o ma ion: lowe ing plan species and he numbe o lowe s a species 188
le el we e eco ded a en 1 × 1 m quad a es placed equidis an ly along he same wo ansec s 189
pe si e (Fig. 1B). We coun ed he numbe o heads in he case o As e aceae species and 190
simple umbels o Apiaceae, bu e e o bo h as lowe s o he sake o simplici y. 191
We calcula ed se e al a iables o de ec po en ial changes in he pollina o 192
communi ies du ing succession: 193
1. Abundance o each pollina o g oup (bumblebees, soli a y bees, honey bee, ho e lies) pe 194
ansec was calcula ed as he numbe o indi iduals pe g oup pe ansec ; 195
2. Species ichness o bumblebees, soli a y bees and ho e lies; species da a om he wo 196
ansec s pe si e we e pooled because o he ela i ely low species numbe s; 197
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S. canadensis is conside ed one o he mos impo an plan s yielding uni lo al honey 346
sou ces in Cen al Eu ope (Fa kas and Zajácz 2007). The e o e, a highly in e es ing inding is 347
ha he inc easing co e o S. canadensis signi ican ly dec eased he abundance o honey 348
bees, e en du ing mass lowe ing. Al hough honey bees a e impo an pollina o s o S. 349
canadensis, hey a e likely o begin isi ing golden od lowe s only when he abundance o 350
o he lowe ing plan species has declined (G oss and We ne 1983). As he adi ionally 351
managed landscape we s udied is a di e se mosaic o c oplands, old ields and managed 352
g asslands (pas u es o hay ields), pollina o s can easily ind o he pollen sou ces in he 353
icini y o he in aded old ields. 354
Al hough plan species ichness and composi ion wen h ough conside able al e a ion 355
along he 20 yea s o succession, nei he soli a y bees no ho e lies showed any di e ence 356
among old ields o di e en age. The e o e, he bimodal species ichness dis ibu ion o 357
pollina o s along seconda y succession p esen ed by S e an-Dewen e and Tscha n ke (2001) 358
could no be eplica ed o bees o o ho e lies in ou s udy. This is because nei he he 359
numbe o lowe ing species, no he numbe o lowe s depended on he age o old ields 360
(da a no shown). One excep ion was he honey bee, which seemed o p e e plan species 361
ypical o he beginning o succession in younge old ields. 362
363
Indi ec e ec s o in asion: lowe isi a ion o na i e species 364
Plan s and hei pollina o s a e igh ly in e wined componen s o ecological communi ies, 365
he e o e he ad e se e ec o S. canadensis migh ha e u he indi ec epe cussions as well. 366
Alien plan species can ha e a s ong e ec on he pollina ion success o na i e species, o en 367
compe ing wi h hem by causing ei he educed pollina o isi a ion a es o inc eased 368
he e ospeci ic pollina ion o na i e lowe s. Howe e , in some cases, bo h a local and 369

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landscape scales, hese o en mass- lowe ing in asi es may acili a e na i e plan pollina ion 370
by inc easing pollina o densi ies (Bje knes e al. 2007). 371
We ound a nega i e co ela ion be ween he co e o S. canadensis and lowe 372
isi a ion o na i e plan s (he ea e lowe s) by bees and ho e lies. On he one hand his 373
esul migh be he consequence o an indi ec compe i ion o pollina ion be ween he 374
supe io in asi e and supp essed na i e species, compe ing possibly o ligh , space, soil 375
nu ien s. On he o he hand, he posi i e e ec o S. canadensis on bumblebees and 376
ho e lies sugges s also di ec compe i ion o pollina ion be ween in asi e and na i e species 377
in Augus due o he la ge amoun o pollen and nec a p o ided by S. canadensis. The lowe 378
equency o lowe isi a ion induced by in asi es can possibly esul in a pollina ion de ici 379
in na i e species (Chi ka & Schü kens 2001). The educ ion in he pollina o pool by alien 380
plan s migh hus co up he ep oduc i e success o insec -pollina ed na i e plan s (S ou & 381
Mo ales 2009). Al hough lowe s o S. canadensis a e sel -incompa ible and dependen on he 382
p esence o pollina o s o seed p oduc ion (G oss & We ne 1983), i can also igo ously 383
sp ead locally by hizomes, hus i s ep oduc i e gain is less a ec ed by a dec eased 384
pollina o pool. Th ough his wo-way compe i ion by di ec and indi ec ou es, S. canadensis 385
can u he p opaga e i s dominance and, he e o e, con ibu e o he de elopmen o an 386
al e na i e, sel -pe pe ua ing s able s a e in which i domina es. 387
388
Conclusions and ecommenda ions 389
Wi hou managemen in e en ions, deg aded, al e na i e s a es d i en by S. canadensis, 390
ins ead o aluable seconda y g asslands, migh domina e he landscape o decades ( his 391
in asi e species can pe sis in he ields o up o 50–75 yea s; Ha ne and Bazzaz 1983), 392
and could a es he na u al ajec o y o seconda y succession in an ea lie , less di e se 393
phase. The e o e, we canno allow spon aneous p ocesses o e ege a e he old ields in aded 394
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by S. canadensis. Ins ead, ac i e managemen ac ions a e needed o guide ege a ion 395
eco e y. 396
Long- e m expe imen s es ing he e ec s o di e en land-use echniques on 397
con olling S. canadensis a e al eady a ailable om Cen al-Eas e n Eu ope (Ho á h 2012). 398
Acco ding o hese expe imen s, a combina ion o p esc ibed dis u bances, such as egula 399
mowing and ex ensi e au umnal g azing wi h ca le o sheep, could educe he a ge in asi e 400
species’ abundance and enhance he eco e y o na i e ege a ion. 401
402
Acknowledgemen s 403
We g ea ly hank And ás Báldi, Zol án Bo a-Duká and Esz e Rup ech o use ul commen s 404
and sugges ions on he manusc ip . Ede Gábos kindly p oduced he map o he s udy a ea and 405
Zsol Józan iden i ied he bees. A.F. was suppo ed by he Eu opean Union and he S a e o 406
Hunga y, co- inanced by he Eu opean Social Fund in he amewo k o TÁMOP 4.2.4. A/2-407
11-1-2012-0001 ‘Na ional Excellence P og am’. A.K-H. and R.F. we e suppo ed by 408
‘Lendüle ’ p og am o he Hunga ian Academy o Sciences, OTKA p ojec (101940), A.K-H. 409
was a Bolyai Fellow and a MTA Pos doc o al Fellow. 410
411
Appendix A. Supplemen a y da a 412
Supplemen a y da a associa ed wi h his a icle can be ound, in he online e sion, a 413
XXXXX." 414
415
Re e ences 416
Abhilasha, D., Quin ana, N., Vi anco, J., Joshi, J. (2008). Do allelopa hic compounds in 417
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Table 1. The e ec s o Solidago canadensis co e , age o he old ields and land-use (M – mown, W – wi hou managemen ) on ege a ion 525
cha ac e is ics acco ding o he linea mixed-e ec s models. Posi i e e ec s a e indica ed by “+” and nega i e e ec s by “–”. E ec sizes (Pa ial 526
) and hei 95% con idence in e al a e also shown. P- alues o signi ican e ec s a e in bold. Signi ican in e ac ions be ween he co a ia es S. 527
canadensis co e and old ield age ha e no con as s bu a e plo ed on Fig. 2. 528
d F p Con as s E ec
size 95% C.I.
Richness and di e si y
Plan species ichness
S. canadensis co e 1, 29
4.43
0.044
– 0.09 -0.24; 0.43
Old ield age 1, 29
32.88
<0.001
+ 0.72 0.38; 1.06
SCC × OFA 1, 29
3.09
0.088
D
i e si y
S. canadensis
co e
1, 29
20.43
<0.001
–
0.24
-
0.09; 0.58
Old ield
age
1, 29
6.07
0.019
+
0.61
0.27; 0.95
SCC × OFA 1, 29
15.19
<0.001
-0.58 -0.92; -0.24
Na u alness
G assland species
S. canadensis co e 1, 29
0.01
0.956
Old ield age 1, 29
53.51
<0.001
+ 0.80 0.46; 1.14
SCC × OFA 1, 29
5.30
0.028
-0.39 -0.73; -0.05
Gene alis species
S. canadensis
co e
1, 29
0.94
0.339
Old ield
age
1, 29
21.43
<0.001
–
-
0.72
-
1.06;
-
0.38
SCC
×
OFA
1, 29
10.69
0.002
0.51
0.17; 0.86
Rude al species
Old ield age 1, 31
45.09
<0.001
– -0.76 -1.11; -0.42
Func ional guilds
G aminoids
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24
S. canadensis
co e
1, 29
1.56
0.221
Old ield
age
1, 29
9.53
0.004
+
0.21
-
0.12; 0.55
SCC
×
OFA
1, 29
5.78
0.022
0.40
0.06; 0.74
Legumes
S. canadensis co e 1, 27
5.42
0.027
+ 0.52 0.18; 0.86
Old ield age 1, 27
8.10
<0.008
+ 0.68 0.34; 1.02
SCC × OFA 1, 27
8.39
0.007
-0.48 -0.82; -0.14
Land-use 1, 27
4.47
0.021
W>M -0.47 -0.81; -0.13
Fo bs
S. canadensis co e 1, 30
12.30
0.001
– -0.50 -0.84; -0.16
Old ield
age
1, 30
38.31
<0.001
–
-
0.74
-
1.09;
-
0.40
529
530
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Figu e

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Figu e
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Zusammen assung
Die sekundä e Sukzession au ehemaligen Felde n
könn e du ch die Besiedelung mi in asi en emden
A en e ände we den, die mögliche weise die ganze
Gemeinscha beein lussen, indem sie die En wicklung
hin zu a en eichen Gemeinscha en e schwe en.
Indessen wu den die Ein lüsse on in asi en A en au
lokale Gemeinscha en sel en in Hinblick au die
sekundä e Sukzession be ach e . Wi un e such en
deshalb den Ein luss de hoch-in asi en Kanadischen
Gold u e (Solidago canadensis) au P lanzen- und
Bes äube gemeinscha en en lang eines G adien en
un e schiedlich s a ke In asion au al en Felde n
un e schiedlichen Al e s (1 bis 20 Jah e sei dem le z en
P lügen) in Süd-T anssil anien (Rumänien). Wi ag en,
ob In asion du ch die Gold u e Ve ände ungen in de
Zusammense zung und Di e si ä de P lanzen- und
Bes äube gemeinscha en en lang des
Sukzessionsg adien en he o u . Deswei e en ag en
wi , inwiewei die Anwesenhei de Gold u e den
Blü enbesuch an einheimischen P lanzen beein luss .
Die In asion eduzie e den A en eich um de
einheimischen P lanzen übe die gesam e Sukzession
hinweg, abe de s ä ks e nega i e E ek au die
P lanzendi e si ä und die Na ü lichkei de Vege a ion
e olg e in den äl e en Sukzessionsgemeinscha en. Die
Gold u en-In asion ha e unabhängig om Al e de
Felde einen nega i en E ek au die Abundanz de
Bienen, abe es gab keinen gleicha igen E ek au die
Schweb liegen. Einheimische P lanzen e uh en einen
eduzie en Blü enbesuch du ch Wildbienen,
Honigbienen und Schweb liegen, de de e meh en
Anwesenhei de Gold u e geschulde wa . Deshalb
ände die In asion du ch diese ausdaue nde
Abs ac in Ge man
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P lanzena den Ve lau de Vege a ionssukzession,
ände die mu ualis ischen Ve bindungen zwischen den
einheimischen A en de ehemaligen Felde und
e u sach die E ablie ung eines un e wünsch en,
al e na i en s abilen Zus andes.