Carabid assemblages in fragmented sandy grasslands

UNCORRECTED PROOF AGEE 2912 1–5 Ca abid assemblages in agmen ed sandy g asslands Tibo Magu a *, Vik o Ko ¨do ¨bo ¨cz Ho oba ´gy Na ional Pa k Di ec o a e, PO Box 216, H-4002 Deb ecen, Hunga y Recei ed 9 Feb ua y 2006; ecei ed in e ised o m 18 Augus 2006; accep ed 22 Augus 2006 Abs ac E ec s o agmen a ion on ca abid assemblages we e s udied in sandy g assland pa ches in eas e n Hunga y. Rela ionship be ween he habi a cha ac e is ics (a ea, isola ion and shape) and he species ichness o ca abid assemblages was examined by o wa d s epwise mul iple linea eg ession analysis. The o al numbe o he collec ed ca abid species co ela ed nega i ely wi h g assland a ea. O e all ca abid species ichness inc eased as he isola ion o pa ches inc eased. The impo ance o he habi a speci ic ca abid species in he assemblages (exp essed by he a io o he numbe o open-habi a species associa ed wi h sandy soils o he o al numbe o species) inc eased wi h pa ch size. These pa e ns could be explained by he in lux o species en e ing om he su ounding ma ix and edge habi a s causing ele a ed o e all species ichness in habi a pa ches wi h limi ed size o highe deg ee o isola ion. #2006 Published by Else ie B.V. Keywo ds: A ea; F agmen a ion; Habi a island; Isola ion; Specialis species; Species ichness 1. In oduc ion In Hunga y, g asslands co e ed almos 33% o he land su ace in he 19 h cen u y, while hei ecen a ea has d opped o 11% (Anonymous, 2005). A many locali ies, his decline c ea ed a ne wo k o isola ed and agmen ed g assland habi a s. In Hunga y, his a ec ed mos ly he sandy g asslands. Such sandy g asslands, o me ly used as pas u es and/o meadows, we e ploughed and used as c opland, a o es ed wi h non-na i e ees o buil o e . F agmen a ion o na u al and semi-na u al habi a s has wo componen s. Fi s , he o al a ea o he habi a sus aining popula ions dec eases. Secondly, hese habi a s end o be mo e isola ed (Saunde s e al., 1991). Less mobile a h opod species a e especially sensi i e o habi a loss and isola ion (Made , 1980; Samways, 2005). G ound-dwelling ca abid bee les (Coleop e a: Ca abidae) a e excep ionally use ul s udy o ganisms o examining he e ec s o landscape changes, such as agmen a ion because hey a e di e se and abundan , hei ecology and sys ema ics a e ela i ely well known (Lo ¨ ei and Sunde land, 1996), and hey seem o be highly sensi i e o agmen a ion (Magu a e al., 2001; Niemela ¨, 2001; Koi ula and Ve meulen, 2005; Lo ¨ ei e al., 2006). The classical heo y o island biogeog aphy (MacA hu and Wilson, 1967) p edic s ha he numbe o species suppo ed by an island inc eases wi h he a ea o he island, and i dec eases wi h he inc easing deg ee o island isola ion. Fu he mo e, i is emphasised ha o he ac o s, such as he shape o he habi a agmen could ha e an e ec on he numbe o species (Lau ence and Yensen, 1991). Recen ly, se e al pape s e ined he o e simpli ied o iginal assump ions o he island biogeog aphy heo y emphasising he e ec s o su ounding habi a s (edges and ma ix) on he species ichness (Kup e e al., 2006; Lo ¨ ei e al., 2006). In he p esen s udy, ca abid assemblages in agmen ed sandy g asslands we e e alua ed in eas e n Hunga y o examine he ela ionships be ween he habi a cha ac e is ics and he species ichness o ca abid assemblages. We hypo hesised ha he ela ionship be ween species ichness and a ea would be s onge i only he habi a speci ic species we e conside ed. Mo eo e , we expec ed ha he mo e isola ed and elonga ed g assland pa ches would ha e www.else ie .com/loca e/agee Ag icul u e, Ecosys ems and En i onmen xxx (2006) xxx–xxx1 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 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 * Co esponding au ho . Tel.: +36 30 2056372; ax: +36 52 529940. E-mail add ess: magu [email protected] (T. Magu a). 0167-8809/$ – see on ma e #2006 Published by Else ie B.V. doi:10.1016/j.agee.2006.08.014 Please ci e his a icle as: Tibo Magu a, Vik o Ko ¨do ¨bo ¨cz, Ca abid assemblages in agmen ed sandy g asslands, Ag icul u e, Ecosys ems and En i onmen (2006), doi:10.1016/j.agee.2006.08.014 UNCORRECTED PROOF ele a ed species ichness because o species en e ing om he su ounding ma ix and edge habi a s. 2. Ma e ial and me hods Ca abid assemblages o eigh sandy g asslands (see Table 1) loca ed in he Nyı ´ se ´g egion, pa o he G ea Hunga ian Plain (eas e n Hunga y) we e s udied. This egion was co e ed by na u al habi a s (ma shes, en meadows, mi es, sandy g asslands and sandy oak woods) in he 19 h cen u y. A e he in ensi ica ion o landscape managemen du ing he 20 h cen u y, hese habi a s we e elimina ed o became highly agmen ed. Today, hese agmen ed sandy g asslands a e su ounded by a able lands and non-na i e ee plan a ions. In he s udied habi a islands, he p e alen ege a ion associa ion was Po en illo a ena iae–Fes uce um pseudo inae. The s udied pa ches a e emnan s o o me ly con agious g asslands, and ecen ly hey ha e been ligh ly g azed wi h cows and sheep (ca le densi y was <0.25 heads/ha). All o he in es iga ed g asslands we e su ounded bo h by non-na i e deciduous ee plan a ions (black locus and ennobled popla species) and c oplands (maize and co n). The e o e, he ma ix habi a s we e simila o all he s udied pa ches. The dis ance be ween he in es iga ed g assland pa ches was a leas 2km. Du ing he 3-yea apping pe iod (2001–2003) ca abid bee les we e collec ed using unbai ed pi all aps, consis ing o plas ic cups (diame e , 100 mm and olume, 500 ml) wi h 70% e hylene glycol as a killing and p ese ing solu ion. The e we e 10 aps, sca e ed andomly wi hin he indi idual pa ches (a leas 100 m om he g assland edges), and we e checked o nigh ly om he end o Ma ch o he end o Oc obe . Because o he di e ence in he size o he pa ches, i would be assumed ha appinge o wasno hesame o all he s udied g assland pa ches, and he e o e, la ge pa ches could ha e been unde es ima ed ega ding species ich- ness. Howe e , using species ichness es ima ing unc ions i is e iden ha in he la ge pa ches he same p opo ion o he es ima ed species pool wassampledasin hesmalle pa ches ( esul s no shown). Ca abid bee les we e iden i ied o species using he keys o Hu ˚ ka (1996). The a ea o he sandy g assland pa ches was measu ed using he A cView GIS p og am package on a digi ized 1:25,000 map. Isola ion o a habi a island is mos o en measu ed as he dis ance o he nea es pa ch. Howe e , isola ion o a habi a pa ch depends no only on he dis ance o he nea es pa ch, bu also on he a ea o he nea es pa ch. A la ge habi a pa ch is mo e likely o ha e a g ea e numbe o species ha can colonise he neighbou ing pa ches. Isola ion was measu ed by he in e se isola ion measu e (Vos and S umpel, 1995), de ined as he o al sandy g assland a ea wi hin a adius o 1000 m a ound he s udied g assland pa ch. This measu e was used as an in e se o he isola ion, because i s alue dec eased as he isola ion o he g assland inc eased. The adius was chosen 1000 m, because e en ligh less ca abids can co e his dis ance (Thiele, 1977). Lau ence and Yensen (1991) s essed ha shape could also in luence he numbe o species inhabi ing habi a island. The shape o g asslands was cha ac e ized by he shape index (Pa on, 1975) de ined as P=200 ffiffiffiffiffiffi pA p, whe e Pwas he pe ime e o he sandy g assland pa ch (m), and A was pa ch a ea (ha). I s alue was 1 o a ound-shaped g assland agmen , while alues g ea e han 1 ep esen ed de ia ion om ci cula i y (Lau ence and Yensen, 1991). The ela ionships be ween he s udied habi a cha ac e - is ics and he species ichness o ca abid assemblages we e examined by o wa d and backwa ds s epwise mul iple linea eg ession analyses. Fo wa d s epwise mul iple linea eg ession analysis p o ided a be e i , hus his me hod was used as sugges ed by Ku ne e al. (1996). We analysed bo h he o al numbe o ca abid species collec ed in he agmen , as well as he habi a specialis s. To ensu e a mo e comple e species in en o y in he s udied g asslands, ca abid ca ches om he h ee apping yea s we e pooled. The dis ibu ion o da a used in he mul iple linea eg ession analyses was no mal ( es ed by he Kolmogo o –Smi no es , Sokal and Rohl , 1995). 3. Resul s Du ing he 3-yea apping pe iod, 8620 ca abids belonging o 67 species we e collec ed om he eigh sandy g asslands. Thi y-one species (7469 indi iduals) we e iden i ied as open-habi a species associa ed wi h sandy soils. The esul o he o wa d s epwise mul iple linea eg ession analysis showed ha ou o he s udied habi a cha ac e is ics, only he a eaand heisola iono he g asslandshadanin luenceon he o al numbe o species. The e was a signi ican nega i e ela ionship be ween he o al numbe o species and he a ea o he sandy g assland (R=0.6983, p= 0.0052, Fig. 1A). The in e se isola ion index and he numbe o all cap u ed ca abid species showed a signi ican nega i e ela ionship (R=0.4211, p= 0.0475): an inc easing isola ion esul ed in ele a ed species numbe s. T. Magu a, V. Ko ¨do ¨bo ¨cz / Ag icul u e, Ecosys ems and En i onmen xxx (2006) xxx–xxx2 AGEE 2912 1–5 Please ci e his a icle as: Tibo Magu a, Vik o Ko ¨do ¨bo ¨cz, Ca abid assemblages in agmen ed sandy g asslands, Ag icul u e, Ecosys ems and En i onmen (2006), doi:10.1016/j.agee.2006.08.014 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 99 100 101 102 103 104 105 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 Table 1 The habi a cha ac e is ics o he eigh s udied sandy g asslands Si e A ea (ha) Isola ion-index (ha) Shape-index 1 99.0 121.5 1.4 2 249.7 122.1 1.3 3 250.6 58.3 1.4 4 353.5 137.6 2.3 5 188.7 130.5 1.2 6 29.1 1.6 1.5 7 51.8 7.3 1.3 8 2.3 137.3 1.2 UNCORRECTED PROOF A signi ican posi i e ela ionship was ound be ween he a io o open-habi a species associa ed wi h sandy soils o he o al numbe o species and he g assland a ea (R=0.7599, p= 0.0287, Fig. 1B; equa ion: a io = 0.4923 + 0.0006 a ea), indica ing he impo ance o habi a speci ic species wi h inc easing pa ch size. Bo h he o e all species ichness and he a io o habi a speci ic species we e un ela ed o he shape index. 4. Discussion Se e al pape s in es iga ing he ela ionship be ween he a ea o habi a island and he numbe o all sampled animal species epo ed signi ican posi i e co ela ion (Made , 1980; Pein inge e al., 2003; Wa son e al., 2005). Some s udies ound ha o e all animal species ichness was un ela ed o he habi a a ea (Hopkins and Webb, 1984; Juliao e al., 2004). Mo eo e , se e al o he pape s, simila ly o ou esul s, and con a y o he p edic ion o he classical heo y o island biogeog aphy, desc ibed a signi ican nega i e ela ionship be ween he a ea o habi a island and he o al numbe o animal species (Baue , 1989; Magu a e al., 2001; Lo ¨ eie al.,2006). The abo e con adic ion could a ise om he ac ha he o iginal heo y conside ed eal islands. Howe e , he e is a majo di e ence be ween eal and habi a islands. In eal islands, he su ounding habi a is usually inhospi able o o ganisms occu ing on islands, while in he case o habi a islands, he bo de ing habi a ( he ma ix) is usually less hos ile. Due o he abo e di e ence, habi a islands could be inhabi ed by colonis s om he ma ix: ‘‘species can colonize he islands om he sea’’ (Cook e al., 2002). Consequen ly, he species–a ea ela ionship obse ed may be posi i e, nega i e o nei he , depending on he balance o con ibu- ion by habi a speci ic and non-habi a speci ic species. Such esul s lead o call o u he e inemen s o he pa adigm (Cook e al., 2002). The e o e, when s udying p edic ions o island biogeog aphy heo y on habi a islands, a dis inc ion mus be d awn be ween specialis species ha uly pe cei e he habi a pa ches as islands and a e unable o su i e in he su ounding ma ix and hose species ha occu in bo h he habi a pa ch and he ma ix (Baue , 1989; Magu a e al., 2001; Cook e al., 2002; Lo ¨ ei e al., 2006). In he p esen s udy, he open-habi a ca abid species associa ed wi h sandy soils could su i e and ep oduce in la ge numbe s only in he sandy g assland pa ches and we e unable o su i e in he adjacen non-na i e deciduous plan a ions. Howe e , he o he ca abid species could su i e in he su ounding ma ix as well. Acco dingly, emo ing he non-habi a speci ic species and analysing he impo ance o only habi a speci ic open-habi a ca abid species associa ed wi h sandy soils, he signi ican nega i e ela ionship became signi ican posi i e as p edic ed by he heo y o island biogeog aphy. This duali y in he species– a ea ela ionship conce ning ca abid bee les is no a special case, as se e al s udies epo ed simila esul s in di e en habi a s (Baue , 1989; Ushe e al., 1993; Halme and Niemela ¨, 1993; De V ies, 1994; Magu a e al., 2001; Lo ¨ ei e al., 2006). Ou indings ha o e all species ichness inc eased as g assland a ea dec eased and g assland isola ion inc eased, as well as ha he impo ance o habi a specialis species in he assemblage inc eased wi h inc easing g assland a ea could be explained by he in luence o species en e ing om he su ounding ma ix and by he need o habi a speci ic species o a minimum a ea o hei habi a . Se e al p e ious s udies on ca abids showed ha species ichness pa e ns in habi a pa ches wi h limi ed size and/o high deg ee o isola ion could be in luenced by species om he neighbou ing ma ix and om he edge, causing inc eased o e all species ichness (Baue , 1989; Halme and Niemela ¨, 1993; Desende e al., 1999; Magu a e al., 2001; Lo ¨ ei e al., 2006). Fu he mo e, highly isola ed pa ches may ‘‘sample’’ mo e gene alis species wi h good dispe sal abili y han less isola ed ones. O he in es iga- ions ocusing on ca abids in g asslands also emphasised ha landscape ea u es (p oximi y o o he habi a s, T. Magu a, V. Ko ¨do ¨bo ¨cz / Ag icul u e, Ecosys ems and En i onmen xxx (2006) xxx–xxx 3 AGEE 2912 1–5 Please ci e his a icle as: Tibo Magu a, Vik o Ko ¨do ¨bo ¨cz, Ca abid assemblages in agmen ed sandy g asslands, Ag icul u e, Ecosys ems and En i onmen (2006), doi:10.1016/j.agee.2006.08.014 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 Fig. 1. Rela ionship be ween he sandy g assland pa ch a ea and he o al numbe o ca abid species collec ed (A) and he a io o open-habi a ca abid species associa ed wi h sandy soils o he o al numbe o ca abid species (B). Bo h eg essions a e signi ican ( p<0.05). Do ed lines ep esen he con idence bands (95%). UNCORRECTED PROOF p opo ion o na u al, semi-na u al habi a s, e c.) we e especially impo an , as species incoming om he adjacen habi a s may conside ably al e he local species composi ion and di e si y (Jeanne e e al., 2003; Pu au e al., 2005). Fu he mo e, a habi a pa ch needs o be o a minimum size o ensu e habi a he e ogenei y and special en i onmen al condi ions equi edbyspecialis species (Niemela ¨e al., 1987; De V ies and Den Boe , 1990). Mo eo e , acco ding o Den Boe (1985) he su i al ime and he ex inc ion a e o a ca abid popula ion a e g ea ly in luenced by he a ea o he habi a . In a sui ably la ge habi a pa ch a popula ion could consis o se e al, asynch onously luc ua ing subpopula ions con ibu ing o he enhancemen o su i al and he educ ion o ex inc ion. The abo e discussed minimum equi ed a ea o ca abids can a y, depending on geog aphic loca ion, habi a s uc u e o he age o he agmen bu i is es ima ed o be ens o hec a es (Niemela ¨, 2001). Fac o s like land use o managemen in ensi y (Daube e al., 2005), mowing egime (Haysom e al., 2004; G andchamp e al., 2005), g azing p essu e (G andchamp e al., 2005) and e iliza ion (K omp, 1999) could also ha e signi ican impac s on bo h o e all species ichness and di e si y o habi a speci ic ca abids. Howe e , in ou si ua ion he s udied g asslands we e managed (g azed) in a simila way. Because o he adhesion o he common Eu opean ag icul u al policy he s udy a ea is bound o be subjec o e en u he landscape scale modi ica ion in he u u e. The e o e, o he conse a ion o he habi a speci ic open-habi a species associa ed wi h sandy soils in he s udied egion he ollowing measu es a e ecommended du ing he sandy g assland managemen . All o he g assland pa ches should be p ese ed because habi a speci ic species a e impo an e en in he assemblages o he smalles pa ches. Mo eo e , o enla ge small pa ches, con e ing adjacen a able lands o g asslands is ecom- mended. Recen pape s (Niemela ¨e al., 2002; Magu a e al., 2004) emphasised ha in habi a pa ches in aded by gene alis species om he su ounding habi a s he numbe o habi a speci ic species may dec ease indica ing in e speci ic compe i i e in e ac ions. In ou s udy, howe e , i was no he case, because o e all ca abid species ichness and he numbe o habi a speci ic open- habi a species associa ed wi h sandy soils showed a signi ican posi i e co ela ion. Acknowledgemen s This s udy was pa o he Na ional Biodi e si y Moni o ing Sys em in Hunga y unded by he Minis y o En i onmen and Wa e . We a e especially g a e ul o Ga ´bo Lo ¨ ei and wo anonymous e iewe s o commen s on ea lie d a s o he manusc ip . We hank Zol a ´n Elek and Ti ada Molna ´ o ield assis ance. Re e ences Anonymous, 2005. Hunga ian online da abase. A h p://www.ksh.hu. Baue , L.J., 1989. Moo land bee le communi ies on limes one ‘habi a islands’. I. Isola ion, in asion and local species di e si y in ca abids and s aphylinids. J. Anim. Ecol. 58, 1077–1098. Cook, W.M., Lane, K.T., Fos e , B.L., Hol , R.D., 2002. Island heo y, ma ix e ec s and species ichness pa e ns in habi a agmen s. Ecol. Le . 5, 619–623. Daube , J., Pu au , T., Allspach, A., F isch, J., Voig lande , K., Wol e s, V., 2005. Local e sus landscape con ols on di e si y: a es using su ace- dwelling soil mac oin e eb a es o di e ing mobili y. Global Ecol. Biogeog . 14, 213–221. Den Boe , P.J., 1985. Fluc ua ions o densi y and su i al o ca abid popula ions. Oecologia 67, 322–330. Desende , K., E ynck, A., Tack, G., 1999. Bee le di e si y and his o ical ecology o woodlands in Flande s. Belg. J. Zool. 129, 139–155. De V ies, H.H., 1994. Size o habi a and p esence o g ound bee le species. In: Desende , K., Du e ˆne, M., Lo eau, M., Lu , M.L., Mael ai , J.-P. (Eds.), Ca abid Bee les: Ecology and E olu ion. Kluwe Academic Publishe s, Do d ech , pp. 253–259. De V ies, H.H., Den Boe , P.J., 1990. Su i al o popula ions o Agonum e ice i Panz. (Col.: Ca abidae) in ela ion o agmen a ion o habi a s. Ne h. J. Zool. 40, 484–498. G andchamp, A.C., Be gamini, A., S o e , S., Niemela ¨, J., Duelli, P., Scheidegge , C., 2005. The in luence o g assland managemen on g ound bee les (Ca abidae: Coleop e a) in Swiss mon ane meadows. Ag ic. Ecosys . En i on. 110, 307–317. Halme, E., Niemela ¨, J., 1993. Ca abid bee les in agmen s o coni e ous o es . Ann. Zool. Fenn. 30, 17–30. Haysom, K.A., McC acken, D.I., Fos e , G.N., So he on, N.W., 2004. De eloping g assland conse a ion headlands: esponse o ca abid assemblage o di e en cu ing egimes in a silage ield edge. Ag ic. Ecosys . En i on. 102, 263–277. Hopkins, P.J., Webb, N.R., 1984. The composi ion o he bee le and spide aunas on agmen ed hea hlands. J. Appl. Ecol. 21, 935–946. Hu ˚ ka, K., 1996. Ca abidae o he Czech and Slo ak Republics. Kabou ek, Zlin, Czech Republic. Jeanne e , P., Schupbach, B., Luka, H., 2003. Quan i ying he impac o landscape and habi a ea u es on biodi e si y in cul i a ed landscapes. Ag ic. Ecosys . En i on. 98, 311–320. Juliao, G.R., Ama al, M.E.C., Fe nandes, G.W., Oli ei a, E.G., 2004. Edge e ec and species–a ea ela ionships in he gall- o ming insec auna o na u al o es pa ches in he B azilian Pan anal. Biodi e s. Conse . 13, 2055–2066. Koi ula, M.J., Ve meulen, H.J.W., 2005. Highways and o es agmen a- ion—e ec s on ca abid bee les (Coleop e a: Ca abidae). Landsc. Ecol. 20, 911–926. K omp, B., 1999. Ca abid bee les in sus ainable ag icul u e: a e iew on pes con ol e icacy, cul i a ion impac s and enhancemen . Ag ic. Ecosys . En i on. 74, 187–228. Kup e , J.A., Malanson, G.P., F anklin, S.B., 2006. No seeing he ocean o he islands: he media ing in luence o ma ix-based p ocesses on o es agmen a ion e ec s. Global Ecol. Biogeog . 15, 8–20. Ku ne , M., Nach schiem, C., Wasse man, W., Ne e , L., 1996. Applied Linea S a is ical Models. McG aw-Hill, Bos on, USA. Lau ence, W.F., Yensen, E., 1991. P edic ing he impac s o edge e ec s in agmen ed habi a s. Biol. Conse . 55, 45–67. Lo ¨ ei, G.L., Sunde land, K.D., 1996. Ecology and beha io o g ound bee les (Coleop e a: Ca abidae). Annu. Re . En omol. 41, 231–256. Lo ¨ ei, G.L., Magu a, T., To ´ hme ´ e ´sz, B., Ko ¨do ¨bo ¨cz, V., 2006. The in luence o ma ix and edges on species ichness pa e ns o g ound bee les (Coleop e a: Ca abidae) in habi a islands. Global Ecol. Biogeog . 15, 283–289. MacA hu , R.H., Wilson, E.O., 1967. The Theo y o Island Biogeog aphy. P ince on Uni e si y P ess, P ince on, NJ, USA. T. Magu a, V. Ko ¨do ¨bo ¨cz / Ag icul u e, Ecosys ems and En i onmen xxx (2006) xxx–xxx4 AGEE 2912 1–5 Please ci e his a icle as: Tibo Magu a, Vik o Ko ¨do ¨bo ¨cz, Ca abid assemblages in agmen ed sandy g asslands, Ag icul u e, Ecosys ems and En i onmen (2006), doi:10.1016/j.agee.2006.08.014 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 287 288 289 290 291 292 293 294 295 296 297 298 299 300 301 302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317 318 319 320 321 322 323 324 325 326 327 328 329 330 331 332 333 334 335 336 337 338 339 340 341 342 343 344 345 346 347 348 349 350 351 352 UNCORRECTED PROOF Made , H.-J., 1980. Die Ve inselung de Landscha aus ie o ¨kologische Sich . Na u Landscha 55, 91–96. Magu a, T., Ko ¨do ¨bo ¨cz, V., To ´ hme ´ e ´sz, B., 2001. E ec s o habi a ag- men a ion on ca abids in o es pa ches. J. Biogeog . 28, 129–138. Magu a, T., To ´ hme ´ e ´sz, B., Molna ´ , T., 2004. Changes in ca abid bee le assemblages along an u banisa ion g adien in he ci y o Deb ecen, Hunga y. Landsc. Ecol. 19, 747–759. Niemela ¨, J., 2001. Ca abid bee les (Coleop e a: Ca abidae) and habi a agmen a ion: a e iew. Eu . J. En omol. 98, 127–132. Niemela ¨, J., Haila, Y., Halme, E., Pajunen, T., Pun ila, P., Tukia, T., 1987. Habi a p e e ences and conse a ion s a us o Agonum manne heimii Dej. in Ha ¨me, sou he n Finland. No . En omol. 67, 175–179. Niemela ¨, J., Ko ze, J.D., Venn, S., Pene , L., S oyano , I., Spence, J., Ha ley, D., Mon es de Oca, E., 2002. Ca abid bee le assemblages (Coleop e a: Ca abidae) ac oss u ban– u al g adien s: an in e na ional compa ison. Landsc. Ecol. 17, 387–401. Pa on, D.R., 1975. A di e si y index o quan i ying habi a edge. Wildli e Soc. B 3, 171–173. Pein inge , M., Be gamini, A., Schmid, B., 2003. Species–a ea ela ionships and nes edness o ou axonomic g oups in agmen ed we lands. Basic Appl. Ecol. 4, 385–394. Pu au , T., Roschewi z, L., Daube , J., Thies, C., Tscha n ke, T., Wol e s, V., 2005. Landscape con ex o o ganic and con en ional a ms: in luences on ca abid bee le di e si y. Ag ic. Ecosys . En i on. 108, 165–174. Samways, M.J., 2005. Insec Di e si y Conse a ion. Camb idge Uni e si y P ess, Camb idge, UK. Saunde s, D.A., Hobbs, R.J., Ma gules, C.R., 1991. Biological conse- quences o ecosys em agmen a ion: a e iew. Conse . Biol. 5, 18– 32. Sokal, R.R., Rohl , F.J., 1995. Biome y. F eeman, New Yo k, USA. Thiele, H.U., 1977. Ca abid Bee les in hei En i onmen s. Sp inge , New Yo k, USA. Ushe , M.B., Field, J.P., Bed o d, S.E., 1993. Biogeog aphy and di e si y o g ound-dwelling a h opods in a m woodlands. Biodi . Le . 1, 54–62. Vos, C.C., S umpel, H.P., 1995. Compa ison o habi a isola ion pa ame e s in ela ion o agmen ed dis ibu ion pa e ns in he ee og (Hylea a bo ea). Landsc. Ecol. 11, 203–214. Wa son, J.E.M., Whi ake , R.J., F eudenbe ge , D., 2005. Bi d communi y esponses o habi a agmen a ion: how consis en a e hey ac oss landscapes? J. Biogeog . 32, 1353–1370. T. Magu a, V. Ko ¨do ¨bo ¨cz / Ag icul u e, Ecosys ems and En i onmen xxx (2006) xxx–xxx 5 AGEE 2912 1–5 Please ci e his a icle as: Tibo Magu a, Vik o Ko ¨do ¨bo ¨cz, Ca abid assemblages in agmen ed sandy g asslands, Ag icul u e, Ecosys ems and En i onmen (2006), doi:10.1016/j.agee.2006.08.014 352353 354 355 356 357 358 359 360 361 362 363 364 365 366 367 368 369 370 371 372 373 373 374 375 376 377 378 379 380 381 382 383 384 385 386 387 388 389 390 391 392 393 394394