Solidago canadensis impacts on native plant and pollinator communities in different-aged old-fields

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 This is a PDF ile o an unedi ed manusc ip ha has been accep ed o publica ion. As a se ice o ou cus ome s we a e p o iding his ea ly e sion o he manusc ip . The manusc ip will unde go copyedi ing, ypese ing, and e iew o he esul ing p oo be o e i is published in i s inal o m. Please no e ha du ing he p oduc ion p ocess e o s may be disco e ed which could a ec he con en , and all legal disclaime s ha apply o he jou nal pe ain. Page 1 o 34 Accep ed Manusc ip 1 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 Page 2 o 34 Accep ed Manusc ip 2 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 Page 3 o 34 Accep ed Manusc ip 3 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 Page 4 o 34 Accep ed Manusc ip 4 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 Page 5 o 34 Accep ed Manusc ip 5 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 Page 6 o 34 Accep ed Manusc ip 6 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 Page 7 o 34 Accep ed Manusc ip 7 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 Page 8 o 34 Accep ed Manusc ip 8 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 Page 15 o 34 Accep ed Manusc ip 15 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 Page 16 o 34 Accep ed Manusc ip 16 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 Page 17 o 34 Accep ed Manusc ip 17 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 in asi e Solidago canadensis s.l. es ain he na i e Eu opean lo a? J. Ecol., 96, 993–418 1001. 419 Page 18 o 34 Accep ed Manusc ip 18 Aizen, M.A., Mo ales, C.L., Mo ales, J.M. (2008). In asi e mu ualis s e ode na i e 420 pollina ion webs. PLoS Biology, 6, e31. 421 Ba ha, S., Szen es, S., Ho á h, A., Házi, J., Zimme mann, Z., Molná , C., ... , Molná , Z. 422 (2014). Impac o mid‐successional dominan species on he di e si y and p og ess o 423 succession in egene a ing empe a e g asslands. 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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 Page 24 o 34 Accep ed Manusc ip 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 Page 31 o 34 Accep ed Manusc ip Figu e Page 32 o 34 Accep ed Manusc ip Figu e Page 33 o 34 Accep ed Manusc ip 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 Page 34 o 34 Accep ed Manusc ip 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.