Changes of isopod assemblages along an urban-suburban-rural gradient in Hungary

UNCORRECTED PROOF O iginal a icle Changes o isopod assemblages along an u bane subu bane u al g adien in Hunga y Elisabe h Ho nung a, *,Be ´la To ´ hme ´ e ´sz b , Tibo Magu a c , Fe enc Vilisics a a Depa men o Ecology, SzIU, FVS, Ins i u e o Zoology, H-1400 Budapes , PO Box 2, Hunga y b Ecological Ins i u e, Uni e si y o Deb ecen, Hunga y c Ho oba´gy Na ional Pa k Di ec o a e, Deb ecen, Hunga y Recei ed 13 Decembe 2005; accep ed 17 Janua y 2007 Abs ac Responses o isopod assemblages o u banisa ion we e s udied along an u banesubu bane u al g adien ep esen ing a dec ease in he in ensi y o human dis u bance. Pi all apping collec ed six species (A madillidium ulga e,Po cellio scabe ,Po cellium collicola,T achelipus a zebu gii,Cylis icus con exus, and T achelipus a hkii). A. ulga e occu ed abundan ly in all si es e lec - ing he b oad ole ance and in asi e na u e o his species. Indica o species analysis demons a ed ha P. scabe and T. a hkii we e signi ican quan i a i e cha ac e species o he u ban si e, while T. a zebu gii was cha ac e is ic o he na u al habi a s (subu ban and u al si es). CANOCO e ealed ha g ound and ai empe a u e show posi i e co ela ion wi h he dis ibu ion o P. scabe and T. a hkii, and nega i e co ela ion wi h T. a zebu gii. Nes ed ANOVA on ap le el showed ha he e we e no signi ican di e - ences be ween he numbe o isopod species and indi iduals, and he di e si y o isopod assemblages in he h ee s udied a eas. Signi ican di e ences we e obse ed a si e le el. The esul s did no suppo he hypo hesis ha di e si y should dec ease in e- sponse o habi a dis u bance. They also con adic ed he in e media e dis u bance hypo hesis; species ichness was no he highes in he mode a ely dis u bed subu ban a ea. Mul i a ia e me hods de ec ed ha he isopod assemblages o he u al and subu ban a eas we e ela i ely simila , while ha o he u ban a ea was ela i ely sepa a ed. Ó2007 Else ie Masson SAS. All igh s ese ed. Keywo ds: Oniscidea; Globene ; Inc eased dis u bance hypo hesis; In e media e dis u bance hypo hesis; Species ichness; Di e si y; U banisa ion 1. In oduc ion The e ec s o u banisa ion can be explo ed h ough in es iga ions o bio ic and abio ic changes along u ban- o- u al g adien s [27,29]. Such g adien s, om densely buil inne ci ies o inc easingly u al su ound- ings, e lec diminishing in ensi ies o human in luence. U ban o es s a e exposed o unique ea u es compa ed o subu ban and u al o es s, including highe ai pollu- ion and dis u bance in ensi y, he hea island phenom- enon and he p esence o g ea e abundance o exo ic species [34,41]. The lo is ic ichness o many u ban habi a s equen ly exceeds ha o less u banised a eas [45], e lec ing he di e se, and mosaic na u e o u ban habi a s and he p esence o in oduced plan s. The ci ye o es eco one also plays an impo an ole in main aining his di e si y [5]. Recen ly, a mul i-na ional esea ch amewo k has been ini ia ed o assess and compa e he in luence o * Co esponding au ho . Tel.: þ36 1 478 42 33; ax: þ36 1 478 42 32. E-mail add ess: [email p o ec ed] (E. Ho nung). 1164-5563/$ - see on ma e Ó2007 Else ie Masson SAS. All igh s ese ed. doi:10.1016/j.ejsobi.2007.01.001 ARTICLE IN PRESS Eu opean Jou nal o Soil Biology xx (2007) 1e8 h p://www.else ie .com/loca e/ejsobi 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 EJSOBI2173_p oo  7 Ma ch 2007  1/8 Please ci e his a icle in p ess as: E. Ho nung e al., Changes o isopod assemblages along an u banesubu bane u al g adien in Hunga y, Eu . J. Soil Biol. (2007), doi:10.1016/j.ejsobi.2007.01.001 +MODEL UNCORRECTED PROOF u banisa ion using in e eb a es and s anda dised ield me hods (Globene ) [30]. Ca abids we e selec ed as he ocal axon, bu o he axa (an s, and spide s) ha e also been s udied ecen ly wi hin he Globene ame- wo k [1]. Acco ding o he Globene p o ocol, we in- ol ed in ou esea ch h ee kinds o o es ed habi a s (u ban pa k, subu ban o es ed a ea, and u al o es ), ep esen ing di e en le els o human dis u bance [31]. The a ge axon was he e es ial, su ace ac i e woodlouse assemblage. Isopods a e ega ded as a use ul and eliable moni o ing g oup. They a e widesp ead, easily iden i ied and in mos cases dominan compo- nen s o he mac odecompose guild in empe a e egions [32]. Isopods belong o he sap ophagous componen s o he soil mac o auna. These o ganisms p ocess he majo i y o dead o ganic ma e ial and so in luence he a e o nu ien elease, an impo an eco- sys em unc ion. The u ban auna can be e y di e se, o en due o in oduced, exo ic species [18,20]. In his pape , we es ed he ollowing p edic ions o isopods in u ban en i onmen s: (1) di e si y should be highes in he subu ban a ea (IDH ein e media e dis u bance hypo hesis); and (2) di e si y should dec ease om a high alue in he u al a ea o a low one in he u ban a ea (inc easing dis u bance hypo hesis). We also in es- iga ed he changes in he isopod assemblages along he u banisa ion g adien , iden i ied he cha ac e is ic and/ o key species ac oss his g adien , and co ela ed ce - ain en i onmen al a iables wi h he obse ed pa e n o isopod abundance and species ichness. 2. Ma e ial and me hods 2.1. S udy a ea The s udy a eas we e si ua ed in and a ound he ci y o Deb ecen (Eas e n Hunga y), he second la ges ci y o he coun y. Th ee o es ed sampling a eas eeach co e ing an a ea o a leas 6 ha ewe e selec ed along an u banisa ion g adien [16]; his ep esen ed u ban, subu ban and u al a eas, acco ding o he Globene p o- ocol [30]. All sampling si es we e in o es s ands dom- ina ed by English oak (Que cus obu ); dis ance among he s udied a eas was a leas 1 km [24]. In he u ban pa k a ea, he e we e se e al asphal -co e ed pa hs and he sh ub laye was s ongly hinned, while in he subu ban a ea he allen ees we e emo ed, while in he u al a ea o es managemen was only occasional a a low-in ensi y le el. The u bane u al g adien ex ended o e a dis ance o app oxima ely 6 km, om he ci y cen e (47320N21 380E), h ough he sub- u bs o he neighbou ing Nagye dTFo es Rese e (47350N21 370E). The c i e ia o dis inguishing u ban, subu ban and u al a ea we e he a io o he buil -up a ea o he na u al habi a s. Dis u bance was es ima ed based on a 1 1 km uni a ound he cen e o he in es iga ed a ea. In he u ban a ea he buil -up a ea exceeded 60%, in he subu ban a ea i was app ox- ima ely 30%, while in he u al a ea he buil -up a ea was 0%. The a ea o he buil -up en i onmen and he na u al habi a s was measu ed by he A cView GIS p og am using an ae ial pho og aph. 2.2. Sampling design Fou si es, a leas 50 m apa , we e selec ed wi hin each sampling a ea (u ban, subu ban, and u al). Iso- pods we e collec ed a each si e using pi all aps, an- domly placing 10 aps a leas 10 m apa om each o he a each si e. This esul ed in a o al o 120 aps sca e ed along he u bane u al g adien (3 a ea 4 si es 10 aps). The pi all aps we e unbai ed, con- sis ing o plas ic cups (diame e 65 mm, olume 250 ml) con aining 75% e hylene glycol as a killing- p ese ing solu ion. The aps we e co e ed wi h ba k pieces o p o ec hem om li e and ain. T apped iso- pods we e collec ed om he end o Ma ch o he end o No embe 2001. Fo he pu pose o analysis, we pooled samples om he whole yea . G ound empe a u e a 2 cm dep h, and ai empe a- u e and ela i e humidi y a he soil su ace we e mea- su ed adjacen o each ap mon hly on he mo ning o a ypical sunny day. The s a is ical analyses we e based on a e ages. We also es ima ed he pe cen age co e o lea li e , decaying wood ma e ial, he bs, sh ubs and he canopy wi hin a adius o 1 m a ound each ap (see Table 1). Table 1 A e age alues (SE) o he s udied en i onmen al ac o s in he s udy a eas U ban Subu ban Ru al G ound empe a u e 24.9 0.249 a 22.3 0.100 b 21.6 0.249 c Ai empe a u e 31.2 0.146 a 27.3 0.076 b 27.8 0.220 c Rela i e humidi y 60.4 0.744 a 76.6 0.495 b 58.9 0.503 c Co e o lea li e 21.1 4.152 a 57.1 4.181 b 21.1 3.207 a Co e o decaying wood ma e ial 3.8 0.495 a 4.2 0.557 a 11.0 1.442 b Co e o he bs 46.5 5.243 a 29.1 4.108 b 68.6 3.348 c Co e o sh ubs 25.7 3.570 a 55.1 3.602 b 11.6 2.183 c Canopy co e 55.7 3.577 a 49.2 3.035 a 52.5 3.331 a Di e en le e s indica e signi ican (p<0.05) di e ences by ANOVA using he Tukey’s pos hoc es . ARTICLE IN PRESS 2E. Ho nung e al. / Eu opean Jou nal o Soil Biology xx (2007) 1e8 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 EJSOBI2173_p oo  7 Ma ch 2007  2/8 Please ci e his a icle in p ess as: E. Ho nung e al., Changes o isopod assemblages along an u banesubu bane u al g adien in Hunga y, Eu . J. Soil Biol. (2007), doi:10.1016/j.ejsobi.2007.01.001 UNCORRECTED PROOF 2.3. Da a analyses To es di e ences in he o e all isopod abundance, species ichness, and di e si y (Shannon, Simpson, Be ge ePa ke ; see [40]) among he h ee sampling a eas (u ban, subu ban and u al), and among he 12 si es, nes ed analyses o a iance (ANOVA) we e pe - o med using da a om he indi idual aps (si es nes ed wi hin he sampling a eas). The dis ibu ion o da a used in he ANOVA models was no mal ( es ed by he Kolmogo o eSmi no es [39]). The composi ion o he isopod assemblages along he u bane u al g adien was compa ed a ap le el by mul idimensional scaling (MDS) based on he num- be o indi iduals using he B ayeCu is index o dis- simila i y [19]. Cha ac e is ic species o he u ban, subu ban and u al a eas we e iden i ied using he IndVal (indica o alue) p ocedu e [3]. This me hod iden i ies quan i a- i ely he cha ac e is ic species o he s udied habi a ypes, and gene a es a signi icance alue (p- alue) o he s eng h o associa ion using a andomised compu- e ised esampling echnique. The cha ac e is ic alue (IndVal) o a species is exp essed as a p oduc o he speci ici y and ideli y measu es. I ecei es i s maxi- mum (100) when all indi iduals o a species a e ound in a single habi a ype (high speci ici y) and when he species occu s a all si es o ha ype (high ideli y) [3]. The cha ac e is ic species is de ined as he mos cha ac e is ic species o each habi a ype, ound mos ly in ha habi a and p esen in he majo i y o si es be- longing o ha habi a . This p o ed o be a use ul me hod o iden i y he cha ac e is ic in e eb a e spe- cies in se e al habi a s [4,22,23]. The ela ionships be ween he en i onmen al ac o s and he abundance and species ichness o isopods we e examined using he CANOCO package [43,44]. The CANOCO algo i hms pe o m o dina ions o aps and species da a, and a ange he o dina ion acco ding o he en i onmen al a iables ha accompany he ap da a [19]. 3. Resul s 3.1. Isopod di e si y along he u bane u al g adien 3.1.1. Abundance The o al isopod sample consis ed o 9115 indi iduals ep esen ing six species; 3548 indi iduals belonging o six species we e cap u ed in he u ban a ea [A ma- dillidium ulga e (La eille, 1804), Po cellio scabe La eille, 1804, Po cellium collicola (Ve hoe , 1907), T achelipus a hkii (B and , 1833), T achelipus a ze- bu gii (B and , 1833), Cylis icus con exus (De Gee , 1778)], i e species and 2720 indi iduals in he subu ban a ea (A. ulga e,P. collicola,T. a hkii,T. a zebu gii, C. con exus), and ou species and 2847 indi iduals in he u al a ea (A. ulga e,P. collicola,T. a hkii, T. a zebu gii). Analysing he ap-le el da a (numbe o indi iduals) by ANOVA we ound no signi ican di e ence in o e all isopod abundance (Table 2; Fig. 1A) a he g adien le el, bu he e we e signi ican di e ences a si e le el. 3.1.2. Numbe o species and di e si y indices The gene al pa e n o changes in species ichness along he g adien is simila o he changes in abun- dance (Table 2;Fig. 1B). We ound no signi ican di e - ence in o e all isopod species ichness. The e was no signi ican di e ence in he Shannon-, Simpson- and Be ge ePa ke di e si y along he g adien (Table 2; Fig. 2). Signi ican di e ences we e obse ed a si e le el o bo h he Shannon and Simpson di e si y indices. 3.2. Isopod assemblage composi ion along he u bane u al g adien The e was no ma ked sepa a ion among he si es along he u bane u al g adien . The MDS o dina ion based on he abundance da a e ealed ha he u ban assemblage was sepa a ed om he subu ban and u al Table 2 Nes ed ANOVA showing di e ences in isopod di e si y, abundance and species ichness along he u banesubu bane u al g adien and among he 12 si es Sou ce o a ia ion d MS Fp Shannon di e si y G adien 2 0.0678 1.3230 ns Si es 9 0.0512 3.5755 <0.01 E o 108 0.0143 Simpson di e si y G adien 2 1.3796 1.5376 ns Si es 9 0.8972 2.2529 <0.05 E o 108 0.3983 Be ge ePa ke di e si y G adien 2 0.6152 1.7406 ns Si es 9 0.3534 1.6312 ns E o 108 0.2167 All species, numbe o species G adien 2 0.0089 0.6306 ns Si es 9 0.0141 0.6205 ns E o 108 0.0228 All species, numbe o indi iduals G adien 2 0.2479 0.4858 ns Si es 9 0.5102 2.2880 <0.05 E o 108 0.2230 ARTICLE IN PRESS 3E. Ho nung e al. / Eu opean Jou nal o Soil Biology xx (2007) 1e8 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 288 289 290 291 292 293 294 295 296 297 298 299 300 301 302 303 304 305 306 307 308 309 310 311 312 EJSOBI2173_p oo  7 Ma ch 2007  3/8 Please ci e his a icle in p ess as: E. Ho nung e al., Changes o isopod assemblages along an u banesubu bane u al g adien in Hunga y, Eu . J. Soil Biol. (2007), doi:10.1016/j.ejsobi.2007.01.001 UNCORRECTED PROOF assemblage and he assemblage o he subu ban and u ban a eas was simila o each o he (Fig. 3). We iden i ied he quan i a i e cha ac e species o he s udied habi a ypes by he IndVal p ocedu e (Table 3): (1) habi a gene alis s, nume ous in all a eas (A. ulga e, P. collicola); (2) synan opic species (p e e ing he u ban a ea) o species wi h b oad ole ance, ei he eco ded exclusi ely (P. scabe ) o mos abundan in he u ban a ea (T. a hkii); (3) species cha ac e is ic o he subu ban a ea (C. con exus); and (4) species cha ac- e is ic o he subu ban and u al a eas (T. a zebu gii). 3.3. En i onmen al ac o s and isopods A Canonical Co espondence Analysis (CANOCO) e ealed ha g ound and ai empe a u es showed pos- i i e co ela ions wi h he abundance dis ibu ions o P. scabe and T. a hkii, and co ela ed nega i ely wi h T. a zebu gii (Fig. 4). 4. Discussion 4.1. Di e si y U banisa ion causes se e al o ms o dis u bance, such as al e a ion, agmen a ion and isola ion o indig- enous habi a s, changes in empe a u e, mois u e and edaphic condi ions, and pollu ion [29]. G ay [6] hypo h- esised ha in habi a s in luenced by dis u bance, o e all di e si y should dec ease. Ou esul s did no suppo his hypo hesis. The o e all species ichness and di e - si y o isopods we e almos as high in he u ban a ea as in he u al one. O e all di e si y changes along he dis- u bance g adien (u bane u al g adien ) can be com- plex, because in a g oup o axa, species ichness may U ban Subu ban Ru al 0.0 1.5 3.0 a a a Numbe o species U ban Subu ban Ru al 0 20 40 60 80 100 B A a a a Numbe o indi iduals Fig. 1. Mean alues (SE) o o e all isopod abundance (A) and o e all isopod species ichness (B) pe ap along he u bane subu bane u al g adien . Di e en le e s indica e signi ican (p<0.05) di e ences based on he LSD (leas signi ican di e ence) mul iple compa ison. U ban Subu ban Ru al 0.0 0.5 1.0 1.5 a a a Be ge -Pa ke di e si y U ban Subu ban Ru al 0.0 0.5 1.0 1.5 2.0 a a a Simpson di e si y U ban Subu ban Ru al 0.0 0.1 0.2 0.3 a a a Shannon di e si y Fig. 2. Mean alues (SE) o o e all isopod di e si y pe ap along he u banesubu bane u al g adien . Di e en le e s indica e signi - ican (p<0.05) di e ences based on he LSD (leas signi ican di - e ence) mul iple compa ison. ARTICLE IN PRESS 4E. Ho nung e al. / Eu opean Jou nal o Soil Biology xx (2007) 1e8 313 314 315 316 317 318 319 320 321 322 323 324 325 326 327 328 329 330 331 332 333 334 335 336 337 338 339 340 341 342 343 344 345 346 347 348 349 350 351 352 353 354 355 356 357 358 359 360 361 362 363 364 365 366 367 368 369 370 371 372 373 374 375 376 377 378 379 380 381 382 383 384 385 386 387 388 389 390 391 392 393 394 395 396 397 398 399 400 401 402 403 404 405 406 407 408 409 410 411 412 413 414 415 416 EJSOBI2173_p oo  7 Ma ch 2007  4/8 Please ci e his a icle in p ess as: E. Ho nung e al., Changes o isopod assemblages along an u banesubu bane u al g adien in Hunga y, Eu . J. Soil Biol. (2007), doi:10.1016/j.ejsobi.2007.01.001 UNCORRECTED PROOF inc ease o dec ease wi h dis u bance depending on hei habi a p e e ence. Ou indings also con adic he IDH [2]. Species ichness and/o di e si y we e no highes in he mode a ely dis u bed subu ban a ea as he IDH p edic s. This may be because basal species in ood webs p obably con o m o his hypo hesis, bu isopods, conside ed as decompose s do no [49]. Mo eo e , u ban a eas do suppo impo an pools o biodi e si y. Wi hin many u ban a eas s udied in he Uni ed S a es and Eu ope, he e a e places wi h le els o biodi e si y ha a e compa able o he su ounding na i e habi a s (e.g. [28,8,37]). The a e age isopod spe- cies ichness in Hunga ian deciduous o es s is a ound 5e6[21]. I is a gued ha di e ences in he landscapes (he e: u ban, subu ban, and u al a ea) migh a ec spe- cies abundance[13]. Howe e , di e ences in o e all Oniscidea abundance be ween si es we e no obse ed. 4.2. Isopod assemblage composi ion along he u bane u al g adien Few s udies ha e been published on he ecological cha ac e is ics o u ban isopod assemblages (commu- ni y s uc u e, di e si y and abundance ela ions) [18,20]. Simila u bane u al o es su eys, ocusing on he soil mac o auna (ea hwo ms, isopods, and mil- lipedes) a e in p og ess in he amewo k o he Bal i- mo e Ecosys em S udies Long Te m Ecological P ojec , in he Uni ed S a es [7,15,33]. In he Bal imo e me opoli an a ea, he abundance o isopods in he u al o es was ex emely low. Mos o he isopod species and specimens we e collec ed in ci y pa ks. Two isopod species (C. con exus and T. a hkii) domina ed hese samples. C. con exus in Hunga y occu ed always in low numbe s in human in luenced habi a s. I was also in oduced o No h Ame ica, whe e i became one o he mos common oniscid species especially in he u ban o es s o he No h-eas e n Uni ed S a es [14]. A su p isingly la ge ac ion o he egional auna may be ound in ci ies [18,20]. In a p e ious s udy in Budapes we ound al oge he 18 isopod species, o -0.6 -0.6 -0.4 -0.4 -0.2 -0.2 0.0 0.0 0.2 0.4 u ban subu ban u al 1s MDS axis 2nd MDS axis 0.2 Fig. 3. O dina ion o he isopod assemblages along he s udied u bane u al g adien using he B ayeCu is index o dissimila i y and MDS o dina ion. Table 3 Two-way indica o able showing he species indica o powe o he habi a clus e ing hie a chy IndVal U ban Subu ban Ru al All habi a A madillidium ulga e 95.83 ns 2088/37 2280/39 2218/39 Po cellium collicola 80.83 ns 272/31 245/35 226/31 Subu ban and u al T achelipus a zebu gii 83.62* 16/3 183/36 339/35 U ban T achelipus a hkii 75.07* 1143/31 10/3 64/19 Po cellio scabe 35.00* 28/14 0/0 0/0 Subu ban Cylis icus con exus 3.33 ns 1/1 2/2 0/0 The IndVal column indica es he species indica o alue o he co e- sponding clus e ing le el, which is he maximum indica o alue ob- se ed in all he clus e ing hie a chy. ns, no signi ican ; *p<0.05. In he columns o each species, he i s alue indica es he numbe o specimens p esen and he second alue co esponds o he numbe o aps whe e he species is p esen , in his sample g oup. Fig. 4. Canonical Co espondence Analysis o he abundance da a o isopods and he measu ed en i onmen al a iables. Cylis icus con- exus was omi ed because i was ep esen ed by only h ee indi iduals. ARTICLE IN PRESS 5E. Ho nung e al. / Eu opean Jou nal o Soil Biology xx (2007) 1e8 417 418 419 420 421 422 423 424 425 426 427 428 429 430 431 432 433 434 435 436 437 438 439 440 441 442 443 444 445 446 447 448 449 450 451 452 453 454 455 456 457 458 459 460 461 462 463 464 465 466 467 468 469 470 471 472 473 474 475 476 477 478 479 480 481 482 483 484 485 486 487 488 489 490 491 492 493 494 495 496 497 498 499 500 501 502 503 504 505 506 507 508 509 510 511 512 513 514 515 516 517 518 519 520 EJSOBI2173_p oo  7 Ma ch 2007  5/8 Please ci e his a icle in p ess as: E. Ho nung e al., Changes o isopod assemblages along an u banesubu bane u al g adien in Hunga y, Eu . J. Soil Biol. (2007), doi:10.1016/j.ejsobi.2007.01.001 UNCORRECTED PROOF which i e (27%) we e in oduced. Th ee species ep e- sen ed u al, o es cha ac e is ic ones [O home opon planum (Budde-Lund, 1885), T. a zebu gii and P o acheoniscus poli us (C. Koch, 1841)]. A. ulga e and P. collicola a e gene alis and expansi e in Hun- ga y. They we e p esen in plan ed, mode a ely dis- u bed o es s. The species dis ibu ion among locali ies e lec ed he deg ee o an h opogenic impac . C. con exus and Po cellionides p uinosus (B and , 1833) indica ed s ong human in luence [20]. U ban habi a s a e o en conside ed o be ho spo s o species in oduc ions [26,42]. Exo ic species can easily colo- nize hese hea ily dis u bed a eas. Once es ablished hey can g ow in numbe and begin o expand hei ange, occasionally becoming in asi e by he gene al- ized s a is ical ‘‘ ens ule’’ [48]. The exclusi e appea ance o P. scabe in he u ban pa k is in acco dance wi h his species’ habi a p e e - ence. Con a y o he A lan ic and Medi e anean a eas o Eu ope, in Hunga y his species can be ound only in human se lemen s, and in and a ound houses (i s Hun- ga ian name, ‘‘cella bug’’ e e s o i s mos equen occu ence) whe e hea island e ec succeeds, while i is a species o ubiqui ous, eu o opic na u e in B i ain [10]. Habi a p e e ences o species may depend on he biogeog aphical egion. Fo example he o iginally Eas - and Middle-Eu opean C. con exus inhabi s na u- al coas al and di e en human si es in B i ain [9]. T. a hkii, simila ly o A. ulga e has a wide ange o habi a s in whole Eu ope which a e mainly di e en g asslands and synan opic places. P. collicola is dis- ibu ed om no he n G eece h ough Hunga y o sou h-eas Ge many [38]. In Hunga y i has a b oad ol- e ance li ing bo h in humid g asslands and mode a ely d y o es s, om occasionally inunda ed galle y o es s o u ban pa ks. The ela i ely s onge sepa a ion o he u ban a ea om he subu ban and u al ones in ou s udy was likely caused by he highe abundances o he gene alis species. Fu he mo e, T. a zebu gii was sensi i e o changes in en i onmen al condi ions in he u ban a ea, and i was only abundan in he subu ban and u al si es. 4.3. En i onmen al ac o s and isopods Al hough e es ial isopods a e dependen i s o all on soil mois u e and humidi y, he e we could use he alues o indi ec ac o s measu ed: g ound empe a u e a 2 cm dep h and he ai empe a u e on he su ace we e highe in he u ban a ea, which appea ed o be a ou ed by T. a hkii and P. scabe . These species p e- e ed he u ban habi a s, which is usually cha ac e ised by highe a e age empe a u e (hea islands). The neg- a i e co ela ion o T. a zebu gii o hese ac o s is ex- plained by he ac ha his species p e e s unmanaged habi a s, which a e usually cha ac e ised by lowe g ound and ai empe a u e. Species dis ibu ion, spa ial and empo al pa e n, abundance o isopods may depend on dis u bance, on ood quali y and shel e si e a ailabili y. P opo ion o dico yledonous plan s as good quali y ood may play an impo an egula ing ac o in isopod abundance [35,36]. The quali y o lea li e did no in luence mi- c ohabi a selec ion o A. ulga e al hough i had signi - ican a ec s on i s g ow h and su i al. Soil ex u e and he o e all luc ua ion o yea ly clima e had an in lu- ence on he species’ spa ial dis ibu ion [11]. The abundance o T achelipus nodulosus C.L. Koch and A. ulga e p o ed o be co ela ed wi h soil pa ame e s and ege a ion composi ion, espec i ely [12]. 4.4. Findings o o he Globene p ojec s The e a e published pape s abou he esul s o he Globene p ojec conce ning ca abid bee les in i e coun ies, including Finland, Canada, Bulga ia, Japan and Hunga y. The a e age numbe o indi iduals and species we e compa ed in each p ojec along an e subu bane u al g adien . In Finland bo h he numbe o indi iduals and he numbe o species inc eased signi ican ly om he u ban a ea owa ds he u al a ea [31]. Ano he s udy in Finland epo ed no signi - ican di e ence in he numbe o indi iduals, while he numbe o species inc eased signi ican ly om he u - ban owa ds he u al a ea [47]. In a hi d s udy no sig- ni ican di e ence was ound nei he in he numbe o indi iduals no in he numbe o species [1]. In Canada he numbe o indi iduals was he highes in he subu - ban a ea, while he numbe o species was he highes in he u al a ea [31]. In Bulga ia he e we e no signi ican di e ences nei he in he numbe o indi iduals no he numbe o species [31]. In Japan bo h he numbe o in- di iduals and he numbe o species inc eased om he u ban a ea owa ds he u al a ea [17]. In Hunga y he numbe o indi iduals was signi ican ly highe in he u- al a ea han in he subu ban and u ban a eas, bu he numbe o species was signi ican ly lowe in he subu - ban a ea han in he u ban and u al a eas du ing 2001. The e we e no di e ences in he numbe o species be- ween he u ban and u al a ea [24]. Howe e , he e was no signi ican di e ence in he numbe o species along he u banisa ion g adien in Hunga y du ing 2002 [25]. In spi e o ha he numbe o species was no di e en , he numbe o o es specialis species inc eased ARTICLE IN PRESS 6E. Ho nung e al. / Eu opean Jou nal o Soil Biology xx (2007) 1e8 521 522 523 524 525 526 527 528 529 530 531 532 533 534 535 536 537 538 539 540 541 542 543 544 545 546 547 548 549 550 551 552 553 554 555 556 557 558 559 560 561 562 563 564 565 566 567 568 569 570 571 572 573 574 575 576 577 578 579 580 581 582 583 584 585 586 587 588 589 590 591 592 593 594 595 596 597 598 599 600 601 602 603 604 605 606 607 608 609 610 611 612 613 614 615 616 617 618 619 620 621 622 623 624 EJSOBI2173_p oo  7 Ma ch 2007  6/8 Please ci e his a icle in p ess as: E. Ho nung e al., Changes o isopod assemblages along an u banesubu bane u al g adien in Hunga y, Eu . J. Soil Biol. (2007), doi:10.1016/j.ejsobi.2007.01.001 UNCORRECTED PROOF signi ican ly om he u ban owa ds he u al a ea in bo h yea s [24,25]. In Finland spide species we e also s udied in he Globene p ojec . The e we e no signi ican di e ences nei he in he numbe o indi iduals no in he species ichness [1]. The ou come o hese s udies is a he di e se. I is likely, ha he numbe o indi iduals and he numbe o species a e no he mos app op ia e measu es cha - ac e ising he e ec o u banisa ion. G oups o species wi h di e en ecological cha ac e is ics ( o es special- is s, open habi a species, gene alis s, e c.) in luenced in a s ikingly di e en way by he u banisa ion. These sub le di e ences in he ecological beha iou and/o habi a p e e ence o he species should be conside ed du ing he s udies. The habi a a ini y indices may p o- ide a use ul way o quan i y hese ecological cha ac e - is ics o he assemblages [46]. Acknowledgmen s We wish o exp ess ou hanks o Z. Elek (DE, Deb ecen, Hunga y) o his help in ield wo k, o K. Szla ecz (JHU, Bal imo e, Uni ed S a es), Z. Ko so ´s and Cs. Csu ´zdi (NHM, Budapes , Hunga y) o he many aluable consul a ions. OTKA T 043508 ( o E.H.), OTKA F 61651 ( o T.M.) and Bolyai Resea ch Fellow g an o he Hunga ian Academy o Sciences ( o T.M.) helped ou esea ch. We app ecia e he com- men s and he sugges ions o Johan Ko ze who helped o imp o e he o me d a o he manusc ip . Re e ences [1] D.M. Ala uikka, D.J. Ko ze, K. Ma einen, J. Niemela ¨, Ca abid and spide assemblages along an u ban o u al g adien in Sou he n Finland, J. Insec Conse . 6 (2002) 195e206. [2] J.H. Connell, Di e si y in opical ain o es s and co al ee s, Science 199 (1978) 1302e1310. [3] M. Du e ˆne, P. Legend e, Species assemblages and indica o species: he need o a lexible asymme ical app oach, Ecol. Monog . 67 (1997) 345e366. [4] Z. Elek, T. Magu a, B. 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Ho nung e al., Changes o isopod assemblages along an u banesubu bane u al g adien in Hunga y, Eu . J. Soil Biol. (2007), doi:10.1016/j.ejsobi.2007.01.001