Responses of carabid beetles to urbanization in Transylvania (Romania)

Ou e e ence: LAND 2010 P-au ho que y- 9 AUTHOR QUERY FORM Jou nal: LAND Please e-mail o ax you esponses and any co ec ions o: E-mail: co ec ions.esil@else ie . homsondigi al.com A icle Numbe : 2010 Fax: +353 6170 9272 Dea Au ho , Please check you p oo ca e ully and ma k all co ec ions a he app op ia e place in he p oo (e.g., by using on-sc een anno a ion in he PDF ile) o compile hem in a sepa a e lis . To ensu e as publica ion o you pape please e u n you co ec ions wi hin 48 hou s. Fo co ec ion o e ision o any a wo k, please consul h p://www.else ie .com/a wo kins uc ions. Any que ies o ema ks ha ha e a isen du ing he p ocessing o you manusc ip a e lis ed below and highligh ed by lags in he p oo . Click on he ‘Q’ link o go o he loca ion in he p oo . Loca ion in Que y / Rema k: click on he Q link o go a icle Please inse you eply o co ec ion a he co esponding line in he p oo Re e ence(s) gi en he e we e no ed in he e e ence lis bu a e missing om he ex – please posi ion each e e ence in he ex o dele e i om he lis . Q1 Unci ed e e ence: This sec ion comp ises e e ences ha occu in he e e ence lis bu no in he body o he ex . Please ci e each e e ence in he ex o , al e na i ely, dele e i . Any e e ence no deal wi h will be e ained in his sec ion. Q2 Please check he inse ion o pa label ‘B’ in he cap ions o Figs. 3–5. Thank you o you assis ance. Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 Landscape and U ban Planning xxx (2011) xxx–xxx 1 Con en s lis s a ailable a ScienceDi ec Landscape and U ban Planning jou nal homepage: www.else ie .com/loca e/landu bplan Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania)1 Béla Tó hmé észa,∗, Is án Má héb, Enik˝ o Balázsc, Tibo Magu ad 2 aDepa men o Ecology, Uni e si y o Deb ecen, POB 71, H-4010 Deb ecen, Hunga y3 bDepa men o Technology and Li e Sciences, Sapien ia Uni e si y, RO-530104 Mie cu ea Ciuc, Pia ¸a Libe ˘ a ¸ii 1, Romania4 cRO-537250 Mie cu ea Ciuc, Pia ¸a Majlá h Gusz á Ká oly, 4A/24, Romania5 dHo obágy Na ional Pa k Di ec o a e, POB 216, H-4002 Deb ecen, Hunga y6 7 a icle in o8 9 A icle his o y:10 Recei ed 29 Oc obe 200911 Recei ed in e ised o m 30 Sep embe 2010 12 13 Accep ed 7 Oc obe 201014 A ailable online xxx 15 Keywo ds:16 Globene 17 U ban18 A h opods19 G adien 20 Species ichness21 Fo es specialis species22 abs ac To in es iga e he impac o u baniza ion on ca abid bee les samples we e aken o e wo yea s using pi all aps along a u al–u ban o es g adien ep esen ing inc easinghuman dis u bance in andnea by he ci y o S ân u Gheo ghe (Romania). We p edic ed ha o al numbe o species should dec ease, whe eas numbe o oppo unis ic and ma ix species should inc ease owa ds he u ban end o he g a- dien . Bo h he o e all species ichness and he numbe o indi iduals we e significan ly he highes in he subu ban a ea ollowed by he u al a ea and he lowes in he u ban a ea. These findings con a- dic ed he inc easing dis u bance hypo hesis; he numbe o species did no dec ease by he inc easing dis u bance. The p opo ion o he o es specialis indi iduals and species significan ly dec eased om he u al owa ds he u ban a ea, suppo ing he habi a specialis hypo hesis. An opposi e pa e n was obse ed in species ichness o he gene alis ca abids, suppo ing he oppo unis ic species hypo he- sis. Bo h he p opo ion o ma ix species and hei densi y we e significan ly highe in he u ban a ea, suppo ing he ma ix species hypo hesis. Ou findings also highligh ed ha o e all di e si y is no an app op ia e indica o ; species wi h di e en habi a a fini ies should be analysed sepa a ely o e alua e he eal e ec o u baniza ion. © 2011 Else ie B.V. All igh s ese ed. 1. In oduc ion23 U baniza ion is a con e sion o lands o u ban o o he buil -up24 a eas (Picke , Cadenasso, & G o e, 2001; Xu e al., 2007). These25 a eas accoun only o a ew pe cen age o he ea h’s land su - 26 ace. Howe e , hei influence on he unc ioning and se ices o 27 ecosys ems a e a he la ge (Albe i, 2005; Be ling-Wol & Wu,28 2004; G imm, G o e, Picke , & Redman, 2000). U baniza ion is an29 inc easinglyimpo an o ceshaping helandscape iahabi a ag- 30 men a ion and loss (Gibb & Hochuli, 2002; Miyashi a, Shinkai, &31 Chida, 1998)and he al e a ion o habi a s uc u e (An op, 2000;32 Fe nandez-Ju icic, 2004; Shocha , S e ano , & Whi ehouse, 2004). 33 All hese modifica ions a ec species ichness and communi y 34 s uc u e in u ban a eas. They c ea e oppo uni ies o gene alis 35 species a ou ingu banen i onmen s,and acili a e hein asiono 36 alien and/o in asi e species (Gode oid & Koedam, 2007; Honnay, 37 Piessens, Van Landuy , He my, & Gulinck, 2003). Unde s anding38 he ela ionship be ween u baniza ion and ecological p ocesses is a39 majo objec i e o u ban ecology (B eus e, Feldmann, & Uhlmann, 40 ∗Co esponding au ho . Tel.: +36 52 512900; ax: +36 52 431148. E-mail add esses: [email p o ec ed] (B. Tó hmé ész), [email p o ec ed] (I. Má hé), [email p o ec ed] (E. Balázs), [email p o ec ed] (T. Magu a). 1998; Wu & Da id, 2002), and in i sel is a key esea ch opic in 41 landscape ecology (Wu & Hobbs, 2002). 42 A way o es ima e he e ec s o u baniza ion on na u e is 43 o s udy he s uc u e and unc ion o ecological sys ems along 44 u al–u ban g adien s (McDonnell & Picke , 1990; Niemelä, Ko ze, 45 & Ashwo h, 2000). Along hese g adien s, he o iginal, na i e habi- 46 a ( u ala eaand/o wildland)isfi s b okenupbynon-con inuous 47 de elopmen and habi a ion wi h mode a e dis u bance (subu - 48 ban a ea). The emaining habi a agmen s in u ban a eas a e 49 influenced by he densely popula ed, buil -up and o en highly 50 dis u bed ci y cen es and hey a e mo e a ec ed, managed, and 51 agmen ed han hei subu ban and u al complemen s. In 1998, 52 an in e na ional esea ch p ojec called Globene (Global Ne - 53 wo k o Moni o ing Landscape Change) was ini ia ed o assess 54 and compa e he impac o u baniza ion on biodi e si y (Niemelä 55 e al., 2000). This p ojec applies he u al–subu ban–u ban g a- 56 dien app oach (Picke e al., 2001) in o es ed habi a s using a 57 common, s anda dized me hodology (pi all apping) and e al- 58 ua ing he esponses o common in e eb a es o u baniza ion. 59 Un il now, he majo i y o he published pape s in he ame o 60 he Globene p ojec in es iga ed ca abid bee les (Elek & Lö ei, 61 2007; Ishi ani, Ko ze, & Niemelä, 2003; Magu a, Tó hmé ész, & 62 Molná , 2004; Magu a, Lö ei, & Tó hmé ész, 2008; Niemelä e al., 63 2002; Sadle , Small, Fiszpan, Tel e , & Niemelä, 2006; Venn, Ko ze, 64 0169-2046/$ – see on ma e © 2011 Else ie B.V. All igh s ese ed. doi:10.1016/j.landu bplan.2011.02.038 Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 2B. Tó hmé ész e al. / Landscape and U ban Planning xxx (2011) xxx–xxx & Niemelä, 2003). S udies analysing o he a ge in e eb a es a e65 a he limi ed ( o spide s: Ala uikka,Ko ze, Ma einen, & Niemelä,66 2002;Magu a, Tó hmé ész, Ho nung, &Ho á h, 2008; o isopods:67 Ho nung, Tó hmé ész, Magu a, & Vilisics, 2007; Magu a, Ho nung,68 & Tó hmé ész, 2008; Vilisics, Elek, & Lö ei, 2007).69 The aim o he p esen s udy was o in es iga e he e ec s o 70 u baniza ion on ca abid bee les along a u al–u ban g adien ep-71 esen ing inc easing human dis u bance. Se e al hypo heses we e72 o mula ed o explain he e ec s o dis u bance on bio ic com-73 muni ies. We es ed he ollowing hypo heses: (i) Acco ding o74 he inc easing dis u bance hypo hesis o mula ed by G ay (1989),75 an inc ease in dis u bance would mono onously dec ease di e -76 si y. Thus, di e si y should dec ease om a high alue in u al77 a ea o a low one in he hea ily dis u bed u ban a ea. (ii) F equen 78 and/o se e e dis u bance would a ec sensi i e species; i p ima -79 ily a ec s he habi a specialis (he e he o es specialis ) species.80 Thus, he habi a specialis hypo hesis p edic s ha di e si y o o -81 es specialis species should dec ease om he less dis u bed u al82 o es owa ds he mo e dis u bed u ban a ea (Magu a e al., 2004).83 (iii)Species ha a e able ocope wi h dis u bance may benefi om84 he dis u bance caused by u baniza ion, and hey should gain dom- 85 inance in he dis u bed subu ban and hea ily dis u bed u ban a ea;86 oppo unis ic species hypo hesis (G ay,1989).(i )Thes udied o es s87 a e su ounded by a ma ix (open habi a s). U baniza ion changes88 conside able he s uc u e o o es ed habi a s, and i makes hem89 ulne able o he in asion o he ma ix species. Species pene a -90 ing om he su ounding ma ix (he e he open-habi a species)91 may benefi om he habi a al e a ion. We a e men ioning his92 new hypo hesis as ma ix species hypo hesis.93 2. Ma e ials and me hods94 2.1. S udy a ea95 The s udy a eas we e in and a ound he ci y o S ân u Gheo-96 ghe (Sepsiszen gyö gy, Wes e n-T ansyl ania, Romania; 45◦51N;97 25◦47E). The dis ance be ween sampling a eas ( u al, subu ban,98 u ban) was 3–10km and all s udied si es co e ed an a ea o 99 g ea e han 10ha. I has been s essed ecen ly ha a o es pa ch 100 needs o ha e a minimum size o main ain an in ac , habi a -101 specific ca abid assemblage; i is es ima ed o be a leas ens o 102 hec a es (Niemelä, 2001). The e o e, ou si e selec ion ulfilled his103 c i e ion.104 Ru al si es we e in a 90-yea -old oak-ho nbeam-beech o es 105 on no h-wes e n slope a 630–719m ele a ion. Pe cen age co e 106 o he canopy laye was 70–80%; equen species in he canopy:107 Fagus syl a ica,Que cus pe aea,and Ca pinus be ulus. The e we e108 densesh ublaye (co e was 20%) and a ela i ely spa sehe b laye 109 (co e was 5–10%). Subu ban si es we e selec ed in a 60-yea -old110 oak-ho nbeam-beech o es on wes e n slope a 600–700m ele-111 a ion. The same species we e equen in he canopy as in he112 u al o es . Pe cen age co e o he canopy laye was 80–90% 113 wi h mode a e sh ub laye (pe cen age co e was 10%). Co e o 114 he b laye was 10–15%. These subu ban si es we e popula o 115 ec ea ion by he local popula ion. The e we e nume ous pa h-116 ways and ampling in ensi y was high. Dead ees we e ha es ed, 117 and allen ees we e also emo ed. U ban si es we e in a cas le118 pa k wi h mode a ely closed canopy (70–80% pe cen age co e )119 wi h spa se sh ub laye (pe cen age co e was 5%) and dense he b 120 laye (pe cen age co e was 30–40%). Besides he na i e species (C. 121 be ulus,F axinus exelsio ,Que cus obu ,Ace campes e,F. syl a ica, 122 Picea abies,Abies alba,Pinus nig a,Pinus s obus,Tilia co da a,Tilia123 pla yphyllos,Aesculus hippocas anum), se e al non-na i e, exo ic124 species we e also p esen : Li iodend on ulipi e a,Magnolia acumi- 125 na a,Tsuga canadensis,Ca agana a bo escens,F. syl a ica subsp.126 a opu pu ea,andThuja plica a.In hepa k allen eesandb anches 127 we e emo ed. Sh ub laye was s ongly hinned. G ass was eg- 128 ula ly mo ed, and he mowed g ass and lea li e we e aken 129 away. The e we e se e al pa ed and asphal -co e ed pa hs in he 130 pa k. 131 2.2. Sampling design 132 Sampling design ollowed he Globene p o ocol (Niemelä e al., 133 2002). Fo es ed sampling a eas we e selec ed along a u al–u ban 134 g adien wi hin he ci y, and in he su ounding o es , as equi ed 135 by he Globene p o ocol. Fou si es, a leas 100m apa we e 136 selec ed wi hin each sampling a ea. Ca abid bee les we e collec ed 137 by andomly placing en pi all aps a leas 10m apa a each si e. 138 This esul ed in a o al o 120 aps along he u al–u ban g adien 139 (3 a eas×4 si es×10 aps). Pi all aps consis ed o plas ic cups 140 (diame e 65mm, olume 250ml) con aining 75% e hylene glycol 141 as a killing-p ese ing solu ion. The aps we e co e ed wi h ba k 142 pieces o p o ec hem om li e and ain. T apped bee les we e 143 collec ed o nigh ly om he end o Ap il o he end o Sep embe 144 in bo h 2004 and 2005. T aps we e placed a he same loca ions in 145 bo h yea s. Ca abids we e iden ified o species using keys in H˚ u ka 146 (1996).147 2.3. Da a analyses 148 The ca abid assemblages along he u al–u ban g adien was 149 displayed by mul idimensional scaling (MDS) using he Man- 150 ha an dis ance o he ela i e abundance o ca abid species 151 (Legend e & Legend e, 1998). Nes ed analyses o a iance wi h 152 epea ed measu es (using Gene al Linea Models) we e pe o med 153 o es di e ences in he o e all ca abid densi y, species ichness, 154 s anda dized species ichness, he a io o o es , gene alis and 155 open-habi a species in he assemblages among he h ee sampling 156 a eas ( u al, subu ban, u ban), among he 12 si es, and be ween he 157 wo yea s (2004 and 2005). Da a om he indi idual aps we e 158 used. Si es we e nes ed wi hin he sampling a eas and yea s we e 159 conce nedas epea ing(Sokal&Rohl ,1995).Toelimina e hee ec 160 o sample size, species ichness was s anda dized o e e y ap 161 using species a e ac ion o expec ed species ichness (Heck, an 162 Belle, & Simbe lo , 1975; Niemelä & Ko ze, 2009). The minimum 163 a iance, unbiased es ima es o he expec ed numbe o species 164 was used (Smi h & G assle, 1977): 165 ES(m)=ST − ST  i=1 N−ni m N n,166 whe e ES(m) is he expec ed numbe o species in a subsample con- 167 aining mindi iduals; ST is he o al numbe o species, niis he 168 abundance o he i h species and Nis he o al numbe o indi- 169 iduals. We choose m=10 indi iduals ( he lowes ca ch in a ap). 170 Calcula ions we e pe o med by he Di O d package (Tó hmé ész, 171 1993). 172 Ca abid bee les we e ca ego ised in o o es , gene alis and 173 open-habi a species acco ding o he in o ma ion in H˚ u ka (1996).174 The dis ibu ion o da a used in he ANOVA model was no mal 175 ( es ed by he Kolmogo o –Smi no es , Sokal & Rohl , 1995). 176 When ANOVA e ealed a significan di e ence be ween he means, 177 a Tukey es was pe o med o mul iple compa isons among 178 means. 179 Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 B. Tó hmé ész e al. / Landscape and U ban Planning xxx (2011) xxx–xxx 3 3. Resul s 180 3.1. Ca abid assemblages along he g adien 181 The o al ca abid ca ch consis ed o 6971 indi iduals ep e-182 sen ing 50 species (3651 indi iduals o 39 species in 2004, and183 3320 indi iduals o 41 species in 2005; Table 1). In he u al a ea184 20 species and 2076 indi iduals we e caugh (999 indi iduals 19185 species in 2004, 1077 indi iduals 15 species in 2005); 26 species186 and 4194 indi iduals we e cap u ed in he subu ban a ea (2352187 indi iduals22speciesin2004,1842indi iduals21speciesin 2005),188 and 701 indi iduals belonging o 36 species we e cap u ed in he189 u ban a ea (300 indi iduals 26 species in 2004, 401 indi iduals190 29 species in 2005). The mos nume ous species was P e os ichus191 oblongopunc a us in bo h yea s, and in o al, made up 26.7% o he192 o al ca ch. Howe e , i occu ed a ely in he u ban a ea. In he193 u al o es , P. oblongopunc a us,Ca abus glab a us,Abax pa allelus 194 andMolops piceus we e hemos abundan in bo h yea s.In he sub- 195 u bana ea,P. oblongopunc a us,P e os ichus hunga icus,C. glab a us 196 and Ca abus iolaceus we e he mos nume ous. In he u ban a ea A. 197 pa allelus,Pseudoophonus ufipes,Abax ca ina us and Ha palus la us 198 we e he mos common (Table 1). 199 U ban ca abid assemblages di e ed om subu ban and u al 200 assemblages; MDS o dina ion e ealed a clea sepa a ion be ween 201 hem (Fig. 1). The assemblages o subu ban and u al a eas we e 202 e y simila o each o he . The ca abid assemblages in he u ban 203 si es we e sepa a ed om he o he s along he fi s axis. The size 204 o he con ex hull on he o dina ion sca e plo was he highes in 205 he case o u ban a ea, indica ing a high he e ogenei y, ha is he 206 composi ion o he apped ca abids changed conside ably om 207 ap o ap (Fig. 1). 208 Table 1 The numbe s and habi a p e e ence o ca abid bee le species cap u ed in pi all aps in and a ound he ci y o S ân u Gheo ghe, T ansyl ania (Romania), in 2004 and 2005. Species sequence is acco ding o he biannual o al (mos common fi s ). F= o es specialis species, G=habi a gene alis species, O =open-habi a species. Species Habi a a fini y 2004 2005 To al Ru al Sub-u ban U ban Ru al Sub-u ban U ban P e os ichus oblongopunc a us F 361 797 6 281 412 2 1859 Ca abus glab a us F 266 396 0 227 369 0 1258 Abax pa allelus F 117 112 119 219 144 255 966 P e os ichus hunga icus G 9 492 0 2 305 0 808 Ca abus iolaceus G 30 236 6 49 313 9 643 Molops piceus F 94 78 0 91 43 0 306 Ca abus co iaceus F 28 80 1 16 34 1 160 P e os ichus nige G 26 8 0 77 41 0 152 Abax pa allelepipedus F 2 41 0 20 81 0 144 Cych us semig anosus F 23 35 0 19 15 0 92 Pseudoophonus ufipes G 0 10 45 0 1 25 81 Pla yde us u us G 0 41 6 0 31 1 79 Ca abus au oni ens F18402610058 Abax ca ina us G 0025622154 Abax schueppeli F 4 8 0 23 18 0 53 Leis us u oma gina us F 528814643 Ha palus la us G 0013001932 Ca abus in ica us F 110 0130 024 Leis us piceus G101100921 Ha palus quad ipunc a us F00800917 Licinus dep essus O001200416 Laemos enus e icola G10900515 Ha palus p og ediens O00600612 T echus quad is ia us G10300711 No iophilus ufipes G02402210 Ca abus a ensis G1500208 Badis e bulla us O0020057 Panagaeus bipus ula us G0050016 Ama a con exio G0020013 No iophilus bigu a us G0100023 Pla ynus assimilis G0120003 Synuchus i alis G0000213 Ama a amilia is G0020002 Ca abus con exus G1100002 Cymindis hume alis O0000202 Leis us e ugineus G0000022 Poecilus cup eus O0110002 S omis pumica us G0000022 Ama a mon i aga O0010001 Ama a simila a O0010001 Anchomenus do salis O0000011 Anysodac ylus bino a us O0000011 Cala hus melanocephalus O0000011 Ha palus dis inguendus O0100001 Lo ice a pilico nis G0000101 No iophilus palus is G0000011 Ophonus a finis O0000011 Ophonus co da us O0000011 P e os ichus mace O0010001 P e os ichus melana ius G0010001 Numbe o indi iduals 999 2352 300 1077 1842 401 6971 Numbe o species 19 22 26 15 21 29 50 Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 4B. Tó hmé ész e al. / Landscape and U ban Planning xxx (2011) xxx–xxx Fig. 1. O dina ion (non-me ic mul idimensional scaling using he Manha an dis- ance o he ela i e equency o he species) o he ca abid assemblages along he s udied Romanian u baniza ion g adien based on he ca ches o pi all aps in 2004 and 2005. S ess o he wo-dimensional configu a ion was 22.86%. Open symbols deno e da a om 2004, while filled ones da a om 2005. 3.2. Ca abid di e si y along he g adien 209 The o al numbe o indi iduals was significan ly he highes in210 he subu ban a ea ollowed by he u al a ea and i was he low-211 es in he u ban a ea (Fig. 2aand Table 2). The o al numbe o 212 ca abid species was also significan ly he highes in he subu ban213 a ea ollowed by he u al a ea and was he lowes in he u ban214 a ea (Fig. 2b and Table 2). A e s anda dizing he sample size by 215 species a e ac ion, he species ichness was significan ly highe 216 20 40 60 d d c b a Numbe o indi iduals 2004 2005 a 4 6 8 c c b b a Numbe o species a Ru al Subu ban U ban 3 4 5 C B b b a a a Es ima ed numbe o species a A Fig. 2. Mean (±SE) alues o he o al numbe o ca abid indi iduals (A), he o al numbe o ca abidspecies(B) and hees ima ednumbe o species o 10 indi iduals (C) along he s udied u baniza ion g adien calcula ed o he pi all aps. Di e en le e s indica e significan di e ences by Tukey es . in he u al and subu ban a eas han in he u ban one. The e was 217 no s a is ically significan di e ence be ween he u al and sub- 218 u ban a eas (Fig. 2c and Table 2). These findings con adic ed he 219 inc easing dis u bance hypo hesis. 220 Bo h he a io o o es specialis ca abid species and he a io 221 o hei abundance dec eased significan ly om he u al a ea 222 owa ds he u ban one (Fig. 3 and Table 2) suppo ing he habi a 223 specialis hypo hesis. An opposi e endency was obse ed o gen- 224 e alis s. The sha e o bo h he gene alis species and indi iduals 225 inc eased significan ly om u al o u ban a ea, albei di e ence in 226 he a io o gene alis indi iduals we e no s a is ically significan 227 be ween hesubu banandu bana eas(Fig.4andTable2).Ou find- 228 ings pa ially suppo ed he oppo unis ic species hypo hesis. Bo h 229 he a io o he open-habi a indi iduals and species we e signifi- 230 can ly highe in he u ban a ea compa ed o he u al o subu ban 231 ones (Fig. 5 and Table 2), suppo ing he ma ix species hypo hesis. 232 4. Discussion 233 The dis u bance g adien om u al o u ban is a g adien o 234 a numbe o dis u bance e en s, such as ampling, managemen , 235 and pe haps pollu ion. We ound ha bo h he species ichness and 236 he numbe o indi iduals we e he highes in he subu ban a ea 237 ollowed by he u al a ea and he lowes in he u ban a ea, con- 238 adic ing he inc easing dis u bance hypo hesis. P opo ion o he 239 o es specialis s dec eased om he u al owa ds he u ban a ea, 240 suppo ing he habi a specialis hypo hesis. Gene alis ca abids 241 showed he opposi e pa e n, suppo ing he oppo unis ic species 242 hypo hesis. Bo h he p opo ion o ma ix species and hei densi y 243 we e significan ly highe in he u ban a ea, suppo ing he ma ix 244 species hypo hesis. 245 4.1. Ra ios s. o als 246 Analysing o al numbe o indi iduals and species ichness as an 247 indica o o he impac s o u baniza ion on in e eb a es was no 248 an en i ely sui able pa ame e because gi en g oups o species may 249 su e (e.g. habi a specialis s), while o he g oups may benefi (e.g. 250 gene alis s and/o ma ix species) om he dis u bance and habi- 251 a al e a ion caused by u baniza ion. Species wi hdi e en habi a 252 a fini ies ( o es specialis s, gene alis s, ma ix species) should be 253 conside ed sepa a ely o de ec accu a ely he di e si y pa e n 254 along he u baniza ion g adien (McIn y e, 2000; Magu a e al., 255 2004; Magu a, Tó hmé ész, & Molná , 2008). The o e all impac 256 o u baniza ion is di e en on di e en species, so a mo e a icu- 257 la ed in e p e a ions is no possible using he summa y di e si y 258 desc ip o s. These limi a ions could be esol ed by conside ing he 259 a ios ( s. o al numbe s) o species wi h di e en habi a a fini ies 260 in an assemblage. 261 4.2. Inc easing dis u bance hypo hesis 262 Inc easing dis u bance hypo hesis p edic s ha inc easing dis- 263 u bancewould mono onously dec ease di e si y(G ay,1989).Ou 264 esul s, howe e , did no suppo his p edic ion as he o al num- 265 be o ca abid species was significan ly he highes in he subu ban 266 a ea ollowed by he u al a ea and was he lowes in he u ban 267 a ea. Some pape s published in he ame o he Globene p ojec 268 also con adic ed his hypo hesis (Ala uikka e al., 2002; esul s 269 om Bulga ia in Elek & Lö ei, 2007; Magu a e al., 2004; Niemelä 270 e al., 2002), whe eas o he s suppo ed i ( esul s om Canada 271 and Finland in Gaublomme, Hend ickx, Dhuy e e , & Desende , 272 2008; Ishi ani e al., 2003; Niemelä e al., 2002; Sadle e al., 2006; 273 Venn e al., 2003). As he e is a significan ela ionship be ween 274 he apped numbe o indi iduals and he collec ed numbe o 275 Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 B. Tó hmé ész e al. / Landscape and U ban Planning xxx (2011) xxx–xxx 5 Table 2 Nes ed ANOVA wi h epea ed measu es showing di e ences in o al numbe o indi iduals and species, in es ima ed species ichness and in p opo ion o o es specialis , gene alis and open-habi a indi iduals and species along he u al–u ban g adien and among he 12 si es. Yea = he e ec o s udy yea (2004 and 2005). Va iable Sou ce d MS F p To al numbe o indi iduals Be ween-subjec s e ec s G adien 2 38703.30 214.66 <0.001 Si es 9 180.30 1.14 ns E o 108 158.70 Wi hin-subjec s e ec s Yea 1 456.50 4.68 <0.05 Yea ×G adien 2 1499.20 15.37 <0.001 Yea ×Si es 9 403.70 4.14 <0.001 E o 108 97.50 To al numbe o species Be ween-subjec s e ec s G adien 2 444.87 85.27 <0.001 Si es 9 5.22 2.35 <0.05 E o 108 2.22 Wi hin-subjec s e ec s Yea 1 0.42 0.17 ns Yea ×G adien 2 6.02 2.48 ns Yea ×Si es 9 4.87 2.01 <0.05 E o 108 2.42 Es ima ed numbe o species ichness Be ween-subjec s e ec s G adien 2 48.37 8.96 <0.01 Si es 9 5.40 6.11 <0.001 E o 108 0.88 Wi hin-subjec s e ec s Yea 1 1.12 0.92 ns Yea ×G adien 2 4.72 3.85 <0.05 Yea ×Si es 9 1.39 1.14 ns E o 108 1.23 P opo ion o o es indi iduals Be ween-subjec s e ec s G adien 2 2.57 12.81 <0.01 Si es 9 0.20 7.16 <0.001 E o 108 0.03 Wi hin-subjec s e ec s Yea 1 0.05 2.74 ns Yea ×G adien 2 0.31 17.10 <0.001 Yea ×Si es 9 0.05 2.65 <0.01 E o 108 0.02 P opo ion o o es species Be ween-subjec s e ec s G adien 2 3.50 45.98 <0.001 Si es 9 0.08 2.97 <0.01 E o 108 0.03 Wi hin-subjec s e ec s Yea 1 0.01 0.60 ns Yea ×G adien 2 0.08 3.52 <0.05 Yea ×Si es 9 0.06 2.78 <0.01 E o 108 0.02 P opo ion o gene alis indi iduals Be ween-subjec s e ec s G adien 2 1.95 15.80 <0.01 Si es 9 0.12 4.31 <0.001 E o 108 0.03 Wi hin-subjec s e ec s Yea 1 0.03 2.17 ns Yea ×G adien 2 0.26 16.74 <0.001 Yea ×Si es 9 0.05 3.36 <0.01 E o 108 0.02 P opo ion o gene alis species Be ween-subjec s e ec s G adien 2 2.06 38.28 <0.001 Si es 9 0.05 2.12 <0.05 E o 108 0.03 Wi hin-subjec s e ec s Yea 1 0.01 0.38 ns Yea ×G adien 2 0.06 2.92 ns Yea ×Si es 9 0.06 2.71 <0.01 E o 108 0.02 P opo ion o open-habi a indi iduals Be ween-subjec s e ec s G adien 2 0.11 6.37 <0.05 Si es 9 0.02 6.37 <0.001 E o 108 0.002 Wi hin-subjec s e ec s Yea 1 0.001 0.51 ns Yea ×G adien 2 0.002 0.67 ns Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 6B. Tó hmé ész e al. / Landscape and U ban Planning xxx (2011) xxx–xxx Table 2 (Con inued) Va iable Sou ce d MS F p Yea ×Si es 9 0.002 0.38 ns E o 108 0.003 P opo ion o open-habi a species Be ween-subjec s e ec s G adien 2 0.22 7.59 <0.05 Si es 9 0.03 4.07 <0.01 E o 108 0.01 Wi hin-subjec s e ec s Yea 1 0.001 0.15 ns Yea ×G adien 2 0.001 0.21 ns Yea ×Si es 9 0.0002 0.06 ns E o 108 0.004 species, a possible eason o he inconsis en esul s is he di e -276 ence in he numbe o ca abid indi iduals cap u ed by pi all aps.277 Using a e ac ion, he p edic ion om he dec easingdi e si ywi h278 inc easing dis u bance was no suppo ed: he ( a ified) numbe 279 o species was significan ly highe in he u al and subu ban a eas280 han in he u ban one. One possible eason o his ailu e is ha he 281 u al–u ban g adien is a complex sys em whe e many en i on-282 men al ac o s ( empe a u e, mois u e, edaphic condi ions, acidi y,283 pollu ion, decomposi ion, e c.) in e ac (Niemelä, 1999). These ac-284 o s a e likely o be di e en in he s udied coun ies, which could285 lead o a ia ion in esponses o ca abids along he g adien s286 (Ishi ani e al., 2003). Mo eo e , in he modified subu ban and/o 287 u ban a eas wi h inc easing edge o edge-like habi a s he species288 pa e n may be s ongly modified (Lö ei, Magu a, Tó hmé ész, &289 Ködöböcz, 2006). A mo e ob ious eason is he di e se esponses o 290 ca abids wi h di e en habi a a fini ies o dis u bance. Fo es spe- 291 cialis s may su e , while gene alis s and species pene a ing om292 hesu oundingma ixmay benefi om he dis u bance and habi-293 a al e a ioncausedbyu baniza ion.Fo ha eason,i islikely ha 294 he o e all di e si y is no he mos app op ia e indica o o dis- 295 u bance. The e o e, species wi h di e en habi a a fini ies should296 be analysed sepa a ely o e alua e he eal e ec o u baniza ion297 (Magu a e al., 2004; Magu a, Ho nung, e al., 2008).298 4.3. Habi a specialis hypo hesis 299 In acco dance wi h he habi a specialis hypo hesis, bo h he300 p opo ion o indi iduals and he species o o es specialis ca a-301 bids dec eased significan ly om he u al a ea owa ds he u ban302 one. All he Globene pape s, which s udied o es species sep-303 a a ely, demons a ed ha u baniza ion caused a p onounced304 change in he assemblages wi h he s onges e ec upon he o -305 es specialis species (Magu a, Lö ei, & Tó hmé ész, 2010; Niemelä 306 & Ko ze, 2009). Fo es specialis species equi e mic osi es wi h a307 pa icula kind o en i onmen al he e ogenei y, such as a ou able308 mic oclima e, p esence o dead and decaying ees, significan 309 co e o lea li e , sh ubs and he bs, oge he o ming an undis-310 u bed o es habi a (Desende , E ynck, & Tack, 1999). Habi a 311 al e a ion caused by u baniza ion appea s o elimina e a ou able312 mic osi es o o es specialis s and con ibu es o he decline o 313 o es specialis s’ p opo ion in he assemblage. Along he s ud- 314 ied g adien , dis u bance was he highes in he u ban a ea (pa ed315 pa hs, hinned sh ub laye ), i was mode a e in he subu ban a ea316 (dead ees ha es ed, and allen ees and b anches emo ed),317 and lowes in he u al a ea. This dec easing dis u bance was also318 exp essed by he inc eased abundance and species ichness o o -319 es specialis ca abid species. 320 4.4. Oppo unis ic species hypo hesis 321 Oppo unis ic species hypo hesis p edic s ha species ha 322 a e able o cope wi h dis u bance would inc ease hei dom-323 0.5 0.6 0.7 0.8 0.9 b c b b a P opo ion o o es indi iduals 2004 2005 a Ru al Subu ban U ban 0.3 0.4 0.5 0.6 0.7 0.8 Ba a c c b b P opo ion o o es species A Fig. 3. Mean (±SE) p opo ions o he o es specialis indi iduals (A) and he o es Q2 specialis species (B) along he s udied u baniza ion g adien o he pi all aps. Di e en le e s indica e significan di e ences by Tukey es . inance (G ay, 1989). Ou esul s did suppo his hypo hesis, 324 as he p opo ion o bo h he indi iduals and species in gen- 325 e alis s we e significan ly he highes in he hea ily dis u bed 326 u ban a ea compa ed o he o he mode a ely o ligh ly dis- 327 u bed subu ban and u al a eas. Da a om Canada (Niemelä 328 e al., 2002), Denma k (Elek & Lö ei, 2007), Finland (Niemeläe 329 al., 2002; Venn e al., 2003) and Hunga y (Magu a e al., 2004)330 suppo ed his p edic ion, as oppo unis ic species we e domi- 331 nan ; he gene alis species we e equen , o hei p opo ion 332 was he highes in he u ban a eas. The e was no di e ence in 333 he numbe o gene alis indi iduals along he u al–u ban g adi- 334 en in Belgium (Gaublomme e al., 2008) o Japan (Ishi ani e al., 335 2003), and none o he species gained clea dominance in he 336 u ban a ea in Bulga ia (Niemelä e al., 2002). A su p ising pa - 337 e n was ound in Finland whe e mo e gene alis indi iduals we e 338 collec ed om u al a eas han ei he u ban o subu ban ones 339 (Ala uikka e al., 2002). 340 Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 B. Tó hmé ész e al. / Landscape and U ban Planning xxx (2011) xxx–xxx 7 0.1 0.2 0.3 0.4 0.5 B a a bc bc c P opo ion o gene alis indi iduals 2004 2005 b A Ru al Subu ban U ban 0.2 0.3 0.4 0.5 c c b b a P opo ion o gene alis species a Fig. 4. Mean (±SE) p opo ions o he gene alis indi iduals (A) and he gene alis species (B) along he s udied u baniza ion g adien o he pi all aps. Di e en le e s indica e significan di e ences by Tukey es . 0.00 0.03 0.06 0.09 b b a a a P op. o open-habi a indi iduals 2004 2005 a Ru al Subu ban U ban 0.00 0.04 0.08 0.12 B b b a a a P opo ion o open-habi a species a A Fig. 5. Mean (±SE) a ios o he open-habi a indi iduals (A) and he open-habi a species (B) along he s udied u baniza ion g adien o he pi all aps. Di e en le e s indica e significan di e ences by Tukey es . 4.5. Ma ix species hypo hesis 341 Ou esul sdidsuppo hishypo hesis,as hep opo iono bo h 342 he indi iduals and species o open-habi a ca abids we e signifi- 343 can ly he highes in he hea ily dis u bed u ban a ea compa ed o 344 he o he mode a ely o ligh ly dis u bed subu ban and u al a eas. 345 The significan al e a ion o he o iginal habi a s in he u ban a ea 346 was eflec ed by he high numbe o ma ix species in he species 347 pool;s ill hei p opo ionwe elowcompa ed ogene alis and o - 348 es species in caseo aps. In heu bana ea, he o es pa cheswi h 349 closed canopy and mode a e closu e because o he walking pa hs 350 and hinned sh ubs allows he colonisa ion and su i al o open- 351 habi a species. Resul s conce ning he ma ix species a e epo ed 352 om Finland and Hunga y; open-habi a species we e mo e abun- 353 dan in he u ban a ea in Finland (Venn e al., 2003) and in Hunga y 354 (Magu a e al., 2004). P o ound changes in habi a quali y du ing 355 u baniza ion (Gilbe , 1989; Niemelä, 1999) p o ide possibili y o 356 he ma ix species o in ade he al e ed u ban habi a s. Koi ula and 357 Niemelä (2003) also poin ed ou ha ma ix species can in ade dis- 358 u bed o es habi a sbecauseo heal e a iono abio ic ac o sand 359 bio ic in e ac ions. 360 4.6. Summa y and ecommenda ions 361 Themodifica ions caused by u baniza ionchangedconside ably 362 he s uc u e o o es ed habi a s. They a ec ed species ichness 363 andcommuni y s uc u e in u ban a eas. Di e si y o o es special- 364 is species adap ed o he o es habi a s dec eased conside ably 365 by he inc easing u baniza ion. Rega ding he o al numbe o 366 species, his dec ease was compensa ed by he in asion o gen- 367 e alis and open-habi a species. In he u ban a ea he e we e open 368 pa ches p oduced by walking pa hs, hinned sh ubs and lawn; he 369 open pa ches allowed he colonisa ion and su i al o open-habi a 370 species and suppo ed gene alis species. In he modified subu ban 371 and/o u ban a eas he e was an inc easing edge o edge-like habi- 372 a s which also may ha e a con ibu ion o he inc eased species 373 ichness o hese a eas. Fo es specialis species equi e mic osi es 374 wi h a pa icula kind o en i onmen al he e ogenei y. Thus, i 375 is i al o inc ease he pa chiness o he u ban pa ks and c ea e 376 closed-canopy o es pa ches wi h allen ee unks, sh ubs, he bs 377 and hick li e laye . I is also impo an o minimize he open 378 pa ches c ea ed by wide pa hs and/o oads; he asphal -co e ed 379 pa hs/ oads a e ba ie s o he ca abids and many o he compo- 380 nen s o he soil auna, hus hey a e especially ha m ul and pa ed 381 pa hs a e p e e ed. 382 Unci ed e e ence Q1 383 Lö ei and Sunde land (1996).384 Acknowledgemen s 385 We a e hank ul o Gabo Lö ei and Johan Ko ze o he 386 help ul commen s and/o p oposal ega ding ou manusc ip . 387 We a e g a e ul o he Sapien ia Founda ion Ins . Resea ch P o- 388 g ammes (Romania) suppo ing he field wo k ( esea ch g an no. 389 1357/2004) and TÁMOP 4.2. IM is g a e ul o he Domus Hun- 390 ga ica Founda ion (Hunga y) suppo ing his isi in Deb ecen. TM 391 hanks he Bolyai Resea ch Fellowship o he Hunga ian Academy 392 o Sciences o suppo ing his esea ch. 393 Re e ences394 Ala uikka, D., Ko ze, D. J., Ma einen, K., & Niemelä, J. (2002). Ca abid bee le and 395 spide assemblages along a o es ed u ban– u al g adien in Sou he n Finland. 396 Jou nal o Insec Conse a ion,6, 195–206. 397 Please ci e his a icle in p ess as: Tó hmé ész, B., e al. Responses o ca abid bee les o u baniza ion in T ansyl ania (Romania). Landscape U ban Plan. (2011), doi:10.1016/j.landu bplan.2011.02.038 ARTICLE IN PRESS GModel LAND20101–8 8B. Tó hmé ész e al. / Landscape and U ban Planning xxx (2011) xxx–xxx Albe i, M. (2005). The e ec s o u ban pa e ns on ecosys em unc ion. In e na ional 398 Regional Science Re iew,28, 168–192.399 An op, M. (2000). Changing pa e ns in he u banized coun yside o Wes e n400 Eu ope. Landscape Ecology,15, 257–270. 401 Be ling-Wol , S., & Wu, J. (2004). Modeling u ban landscape dynamics: A e iew.402 Ecological Resea ch,19, 119–129.403 B eus e, J., Feldmann, H., & Uhlmann, O. (1998). U ban ecology. Be lin: Sp inge .404 Desende , K., E ynck, A., & Tack, G. (1999). Bee le di e si y and his o ical ecology405 o woodlands in Flande s. Belgian Jou nal o Zoology,129, 139–156.406 Elek, Z., & Lö ei, G. L. (2007). Pa e ns in g ound bee le (Coleop e a: Ca abidae)407 assemblages along an u banisa ion g adien in Denma k. Ac a Oecologica,32,408 104–111.409 Fe nandez-Ju icic, E. (2004). Spa ial and empo al analysis o he dis ibu ion o 410 o es specialis s in an u ban- agmen ed landscape (Mad id, Spain) implica- 411 ions o local and egional bi d conse a ion. Landscape and U ban Planning,69,412 17–32.413 Gaublomme, E., Hend ickx, F., Dhuy e e , H., & Desende , K. (2008). The e ec s o 414 o es pa ch size and ma ix ype on changes in ca abid bee le assemblages in415 an u banized landscape. Biological Conse a ion,141, 2585–2596.416 Gibb, H., & Hochuli, D. F. (2002). Habi a agmen a ion in an u ban en i onmen 417 la ge and small agmen s suppo di e en a h opod assemblages. Biological418 Conse a ion,106, 91–100.419 Gilbe , O. L. (1989). The ecology o u ban habi a s. London UK: Chapman and Hall.420 Gode oid, S., & Koedam, N. (2007). U ban plan species pa e ns a e highly d i en 421 by densi y and unc ion o buil -up a eas. Landscape Ecology,22, 1227–1239.422 G ay, J. S. (1989). E ec s o en i onmen al s ess on species ich assemblages. Bio-423 logical Jou nal o he Linnean Socie y,37, 19–32. 424 G imm, N. B., G o e, J. M., Picke , S. T. A., & Redman, C. L. (2000). In eg a ed425 app oaches o long- e m s udies o u ban ecological sys ems. Bioscience,50,426 571–584.427 Heck,K. L., an Belle,G., &Simbe lo , D.(1975). Explici calcula iono he a e ac ion 428 di e si y measu emen and he de e mina ion o su ficien sample size. Ecology, 429 56, 1459–1461.430 Honnay, O., Piessens, K., Van Landuy , W., He my, M., & Gulinck, H. (2003). Sa elli e431 based land use and landscape complexi y indices as p edic o s o egional plan 432 species di e si y. Landscape and U ban Planning,63, 241–250.433 Ho nung, E., Tó hmé ész, B., Magu a, T., & Vilisics, F. (2007). Changes o isopod 434 assemblages along an u ban–subu ban– u al g adien in Hunga y. Eu opean435 Jou nal o Soil Biology,43, 158–165.436 H˚ u ka, K. (1996). Ca abidae o he Czech and Slo ak Republics. Zlin: Kabou ek.437 Ishi ani, M., Ko ze, D. J., & Niemelä, J. (2003). Changes in ca abid bee le assemblages438 ac oss an u ban– u al g adien in Japan. Ecog aphy,26, 481–489.439 Koi ula, M., & Niemelä, J. (2003). Gap elling as a o es ha es ing me hod in bo eal 440 o es s: Responses o ca abid bee les (Coleop e a, Ca abidae). Ecog aphy,26,441 179–187.442 Legend e, P., & Legend e, L. (1998). Nume ical ecology. Ams e dam: Else ie Science.443 Lö ei,G. L.,Magu a, T.,Tó hmé ész,B.,&Ködöböcz,V.(2006). Theinfluence o ma ix444 andedgesonspecies ichness pa e nso g ound bee les(Coleop e a,Ca abidae)445 in habi a islands. Global Ecology and Biogeog aphy,15, 283–289.446 Lö ei, G. L., & Sunde land, K. D. (1996). Ecology and beha io o g ound bee les447 (Coleop e a Ca abidae). Annual Re iew o En omology,41, 231–256. 448 Magu a, T., Ho nung, E., & Tó hmé ész, B. (2008). Abundance pa e ns o e es ial449 isopods along an u baniza ion g adien . Communi y Ecology,9, 115–120.450 Magu a, T., Lö ei, G. L., & Tó hmé ész, B. (2008). Time-consis en ea angemen 451 o ca abid bee le assemblages by an u banisa ion g adien in Hunga y. Ac a 452 Oecologica,34, 233–243. 453 Magu a, T., Tó hmé ész, B., Ho nung, E., & Ho á h, R. (2008). U banisa ion and454 g ound-dwelling in e eb a es. In L. N. Wagne (Ed.), U baniza ion 21s cen u y455 issues and challenges (pp. 213–225). No a Science Publishe s. 456 Magu a, T., Tó hmé ész, B., & Molná , T. (2008). A species-le el compa ison o occu -457 ence pa e ns in ca abids along an u banisa ion g adien . Landscape and U ban458 Planning,86, 134–140.459 Magu a,T., Lö ei,G.L.,&Tó hmé ész, B.(2010). Doesu banisa iondec easedi e si y 460 in g ound bee le (Ca abidae) assemblages? Global Ecology and Biogeog aphy,19,461 16–26. Magu a, T., Tó hmé ész, B., & Molná , T. (2004). Changes in ca abid assem- 462 blages along an u banisa ion g adien . Landscape Ecology,19, 747– 463 759. 464 McDonnell, M. J., & Picke , S. T. A. (1990). Ecosys em s uc u e and unc ion along 465 u ban– u al g adien s an unexploi ed oppo uni y o ecology. Ecology,71,466 1232–1237. 467 McIn y e, N. E. (2000). Ecology o u ban a h opods: A e iew and a call o ac ion. 468 Annals o he En omological Socie y o Ame ica,93, 825–835. 469 Miyashi a, T., Shinkai, A., & Chida, T. (1998). The e ec s o o es agmen a ion 470 on web spide communi ies in u ban a eas. Biological Conse a ion,86, 357– 471 364. 472 Niemelä, J. (1999). Ecology and u ban planning. Biodi e si y and Conse a ion,8,473 119–131. 474 Niemelä, J. (2001). Ca abid bee les (Coleop e a: Ca abidae) and habi a agmen a- 475 ion: A e iew. Eu opean Jou nal o En omology,98, 127–132. 476 Niemelä, J., Ko ze, J., & Ashwo h, A. (2000). The sea ch o common an h opogenic 477 impac s on biodi e si y: A global ne wo k. Jou nal o Insec Conse a ion,4, 3–9. 478 Niemelä, J., & Ko ze, J. D. (2009). Ca abid bee le assemblages along u ban o u al 479 g adien s: A e iew. Landscape and U ban Planning,92, 65–71. 480 Niemelä,J., Ko ze,J. D.,Venn, S.,Pene ,L.,S oyano ,I.,Spence,J.,e al. (2002).Ca abid 481 bee le assemblages (Coleop e a, Ca abidae) ac oss u ban– u al g adien s: An 482 in e na ional compa ison. Landscape Ecology,17, 387–401. 483 Picke , S. T. A., Cadenasso, M. L., & G o e, J. M. (2001). U ban ecological sys- 484 ems: Linking e es ial ecological, physical, and socioeconomic componen s 485 o me opoli an a eas. Annual Re iew o Ecology, E olu ion, and Sys ema ics,32,486 127–157. 487 Sadle , J. P., Small, E. C., Fiszpan, H., Tel e , M. G., & Niemelä, J. (2006). In es iga - 488 ing en i onmen al a ia ion and landscape cha ac e is ics o an u ban– u al 489 g adien using woodland ca abid assemblages. Jou nal o Biogeog aphy,33,490 1126–1138. 491 Shocha , E., S e ano , W. L., & Whi ehouse, M. E. A. (2004). U baniza ion and spide 492 di e si y: Influences o human modifica ion o habi a s uc u e and p oduc i - 493 i y. Ecological Applica ions,14, 268–280. 494 Smi h, W., & G assle, F. J. (1977). Sampling p ope ies o a amily o di e si y mea- 495 su es. Biome ics,33, 283–292. 496 Sokal, R. R., & Rohl , F. J. (1995). Biome y. New Yo k: F eeman. 497 Tó hmé ész, B. (1993). NuCoSA 1.0: Numbe c unche o communi y s udies and 498 o he ecological applica ions. Abs ac a Bo anica,17, 283–287. 499 Venn, S. J., Ko ze, D. J., & Niemelä, J. (2003). U baniza ion e ec s on ca abid di e si y 500 in bo eal o es s. Eu opean Jou nal o En omology,100, 73–80. 501 Vilisics, F., Elek, Z., & Lö ei, G. L. (2007). Composi ion o e es ial isopod assem- 502 blages along an u banisa ion g adien in Denma k. Pedobiologia,51, 45–53. 503 Wu, J., & Da id, J. (2002). A spa ially explici hie a chical app oach o modeling 504 complex ecological sys ems: Theo y and applica ions. Ecological Modelling,153,505 7–26. 506 Wu, J., & Hobbs, R. (2002). Key issues and esea ch p io i ies in landscape ecology: 507 An idiosync a ic syn hesis. Landscape Ecology,17, 355–365. 508 Xu, C., Liu, M., Zhang, C., An, S., Yu, W., & Chen, J. M. (2007). The spa io empo al 509 dynamics o apid u ban g ow h in he Nanjing me opoli an egion o China. 510 Landscape Ecology,22, 925–937. 511 Béla Tó hmé ész is p o esso o ecology a he Ecological Ins i u e, Uni e si y o 512 Deb ecen.His esea ch in e es sinclude he heo yo di e si y,communi yecology, 513 and u baniza ion. 514 Is án Má hé is assis an p o esso o he Depa men o Technical and Na u al Sci- 515 ences a he Sapien ia Hunga ian Uni e si y o T ansyl ania, Romania. His esea ch 516 in e es s include ca abid’s ecology and u baniza ion. 517 Enik˝ o Balázs is a mas e o sciences s uden a he Babes¸-Bolyai Uni e si y, Cluj 518 Napoca, Romania. He esea ch in e es is he e ec o u baniza ion on Ca abids. 519 Tibo Magu a is field biologis a he Ho obágy Na ional Pa k Di ec o a e, and 520 leade o he Ca abidology Resea ch G oup a he Uni e si y o Deb ecen. His main 521 esea ch in e es s include he dis ibu ion, biogeog aphy and ecology o g ound 522 bee les. 523