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Both local and landscape-level factors are important drivers in shaping ground-dwelling assemblages of sandy grasslands

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1 2 Local and landscape-level factors are equally important drivers in shaping 3 ground-dwelling spider assemblages of sandy grasslands 4 5 Roland Horvátha,, Tibor Maguraa, Béla Tóthmérészc*, János Eichardtb, and Csaba 6 Szinetárb 7 8 a Department of Ecology, University of Debrecen, Egyetem tér 1, Debrecen, H-4032 9 Hungary

10 b Department of Zoology, University of West Hungary, Károlyi Gáspár tér 4, 11 Szombathely, H-9700, Hungary

12 c MTA-DE Biodiversity and Ecosystem Services Research Group, Egyetem tér 1, 13 Debrecen, H-4032 Hungary

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19 Abstract 20 21 Due to agricultural production the area of grasslands decreased dramatically and they 22 also become fragmented. Moreover, the proportion of landscape elements surrounding 23 the grasslands also changed considerably. Earlier studies emphasized that landscape- 24 level factors were essential in shaping assemblages of semi-natural 25 grasslands. Recently several studies demonstrated that local factors were more 26 important than landscape-level factors. Studying ground-dwelling spider assemblages in 27 sandy grassland fragments we found that out of the four investigated landscape-level 28 factors (isolation, total area of croplands in the landscape, total area of forests in the 29 landscape, and landscape diversity), only the area of the sandy grasslands around the 30 fragments was a significant predictor controlling spider diversity. The total number of 31 species, the number of generalist species and the number of hunting species increased 32 significantly with the isolation of fragments. Out of the three studied local factors 33 (fragment size, the shape of grassland fragment, and grazing intensity), only the size of 34 the fragments was a significant predictor of the diversity of specialist , since the 35 number of these species increased significantly as the fragments size increased. Our 36 results suggest that both local and landscape-level factors are equally important drivers 37 in maintaining spider diversity in sandy grasslands; therefore, local and landscape-level 38 factors should be considered simultaneously during the restoration and/or management 39 of grasslands. 40 41 Keywords: fragmentation; generalist species; grazing; habitat specialist species; 42 hunting strategy; isolation

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44 Introduction 45 46 Grasslands are one of the most extensive ecosystems in the world; they cover 40% 47 of the terrestrial area (Sala 2001). In the last century, due to human activities large 48 grassland areas disappeared or decreased considerably, causing serious fragmentation of 49 the remaining grassland habitats (Dengler et al. 2014). Habitat alteration, fragmentation 50 and land use changes are the main anthropogenic processes influencing grasslands at 51 local and global level; these changes caused considerable decrease in populations of 52 certain species by as much as 20-50% throughout Europe as well as change the 53 extinction and colonization rates of grassland inhabiting (Batáry et al. 2012; 54 Horváth et al. 2013). 55 Traditional management practices like low intensity, wildlife-friendly grazing and 56 mowing resulted in diverse semi-natural grasslands (Batáry et al. 2008). During the 57 second half of the 20th century the more efficient large scale farming techniques, such 58 as better drainage, chemical inputs and abandonment of traditional management 59 practices resulted a considerable reduction, alteration and fragmentation of natural and 60 semi-natural grasslands (Buchholz 2010; Habel et al. 2013; Horváth et al. 2009). Most 61 grasslands were cultivated for crops, or left ungrazed and increasingly taken over by 62 shrubbery and forest. Thus, conventionally managed, species-rich grasslands have 63 declined drastically worldwide. These processes have led to the disappearance of many 64 characteristic plant and species in these habitats, contributing to a remarkable 65 loss of biodiversity (Hooftman and Bullock 2012; Krauss et al. 2010; Tscharntke et al. 66 2005). Therefore, it is important that farmers and conservationists across the world re- 67 establish a network of natural and semi-natural grasslands (Ödman and Olsson 2014; 68 Poschold and WallisDeVries 2002). 69 For the maintenance of grasslands and their biodiversity at local scale, appropriate 70 management practices like grazing and/or mowing are necessary, since the lack of 71 managements or very low intensity treatments result in grasslands becoming dominated 72 by perennial rank or tussock grasses and shrubs (Albert et al. 2014; Levin, 2013). At 73 landscape scale, agricultural lands were simplified due to the decrease of landscape 74 diversity caused by the dominance of few crop species and the disappearance of natural 75 and semi-natural grasslands (Concepción et al. 2012). Besides landscape diversity, 76 fragmentation and isolation of natural grasslands have vital influence on local species 77 richness and composition (Horváth et al. 2013, 2015). All these processes have caused 78 habitat degradation, which led to a reduction in the suitability of grasslands for biotas 79 (Benton et al. 2003; Duelli and Obrist 2003). Therefore, to the increase of species 80 diversity in these habitats there is an urgent need to restore natural and semi-natural 81 grasslands and reduce their fragmentation. 82 Sandy grasslands are valuable and the most threatened habitats in Central Europe 83 (Dengler et al. 2014). Since the mid 20th century agricultural management has been 84 causing the fragmentation or disappearance of these grasslands. Thus, rare and sand 85 specialist plant and animal species are highly endangered (Faust et al. 2011; Horváth et

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86 al. 2013). Spiders are diverse and abundant generalist arthropod predators and play an 87 important role in functioning of natural and semi-natural habitats and agricultural 88 ecosystems and in shaping arthropod communities as natural pest enemies (Horváth et 89 al. 2009, 2013; Marc et al. 1999; Picchi et al. 2016). Moreover, their and 90 ecology are well known, they comprise specialist and web-building species which are 91 sensitive to habitat alteration caused by management regime (burning, mowing, and 92 grazing) and fragmentation (Cattin et al. 2003; Horváth et al. 2009; Valkó et al. 2016). 93 In contrast, generalist and hunting species are more adaptive, and less sensitive than 94 specialist and web-building species (Horváth et al. 2009, 2013, 2015). To understand 95 the effects of land use changes on the structure and composition of specialist, generalist, 96 web-building and hunting spider assemblages in natural and semi-natural habitats, it is 97 necessary to study these influences at both local and landscape-level scales (Batáry et al. 98 2008; Horváth et al. 2015). 99 Earlier studies demonstrated that fragment size (Horváth et al. 2013, 2015; Jonsson 100 et al. 2009; Toft and Schoener 1983) and management like grazing (Eldridge and 101 Whitford 2009; Kovac and Mackay 2009; Batáry et al. 2008; Horváth et al. 2009) were 102 the most important local factors in shaping spider assemblages. However, there are 103 several other local factors which also influence species richness and density of spiders 104 such as the shape of habitat (Barbaro et al. 2005), management types (Batáry et al. 105 2008; Picchi et al. 2016), prey density (Horváth et al. 2005). Habitat isolation proved to 106 be the most influential landscape-level factors on spiders (Horváth et al. 2013; Jonsson 107 et al. 2009), but further factors such as extent of grasslands, croplands and forests in the 108 landscape, and landscape diversity were also important (Clough et al. 2005; Schmidt et 109 al. 2008; Horváth et al. 2015; Picchi et al. 2016). There are relatively few studies that 110 investigate the effects of both local and landscape-level factors on spider assemblages 111 (Horváth et al. 2009, 2015; Torma et al. 2014; Zulka et al. 2014). Therefore, the aim of 112 this study was to examine the effects of local (fragment size, shape of grassland, and 113 grazing intensity) and landscape-level factors (isolation, area of croplands in the 114 landscape, area of forests in the landscape, and landscape diversity) on ground-dwelling 115 spiders in sandy grassland fragments. To get an ecologically relevant picture of the 116 effects of these factors we investigated not only overall species richness but the richness 117 of spiders with different habitat affinity and hunting strategy, because studying only the 118 total number of species could mask the real effects of local and landscape-level factors 119 (Batáry et al. 2008; Horváth et al. 2015). Based on the above we hypothesized: (i) 120 spider species richness should increase with increasing fragment size; (ii) the increasing 121 grazing intensity (measured by vegetation height) decreases the species richness; (iii) 122 more isolated fragments embedded in an impoverished landscape (with a higher extent 123 of croplands and forests) should have fewer species. 124 125 Material and methods 126 127 Study area

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128 129 We selected eight dry sandy grassland fragments in the Nyírség region (Great 130 Hungarian Plain, Eastern Hungary) (Table 1). In the 19th century the typical natural 131 habitat types were dry habitats (sandy grasslands, and sandy oak woods) and wetlands 132 (marshes, mires and fen meadows) in this region. Due to intensive landscape 133 management during the 20th century, these habitats disappeared or became highly 134 fragmented. Nowadays, these altered and fragmented dry sandy grasslands are 135 encompassed by croplands and non-native tree plantations. In each dry sandy grassland 136 fragment, the prevalent grassland association was Cynodonti-Festucetum pseudovinae 137 (Török et al. 2008). These studied fragments have been lightly, moderately or heavily 138 grazed by cattle and/or sheep. All of the investigated grassland fragments were 139 surrounded by both croplands (maize and corn) and non-native deciduous tree 140 plantations (black locust and ennobled poplar species). Therefore, the neighbouring 141 habitats were similar to the studied fragments. The shortest distance between the 142 investigated grassland fragments was 2 km, while the furthest distance was 75 km. 143 144 # Table 1 approximately here # 145 146 Sampling design 147 148 We collected spiders using pitfall traps during a three-year studying period (2001- 149 2003). The traps consisted of 100 mm diameter plastic cups (volume of 500 ml) and 150 contained about 200 ml 70% ethylene glycol as killing and preserving solution. The 151 traps were covered by fibreboard to be protected from litter and rain. We placed ten 152 traps randomly in each studied grassland fragment. All traps were placed at least 30m 153 from each other and from the grassland edges to avoid edge effect (Lafage and Pétillon 154 2014). We emptied the traps fortnightly from the end of March to the end of October. 155 We identified spider specimens to species level using standard keys (Nentwig et al. 156 2017). We followed the World Spider Catalog’s nomenclature (World Spider Catalog 157 2017). 158 159 Data analysis 160 161 We investigated the effects of local and landscape-level factors on spider 162 assemblages (Table 1). Fragment size, shape of grassland fragment, and grazing 163 intensity were regarded as local factors. We measured the size of the studied dry sandy 164 grassland fragments on digitized 1:25000 maps and aerial photographs using the 165 ArcGIS program. Laurence and Yensen (1991) emphasized that the patch shape could 166 also affect the number of species within a habitat island. We characterized the shape of 167 the grassland fragments by the shape index (Magura et al. 2001) defined as 168 P/(200√(πA)), where P was the perimeter of the sandy grassland fragment (m), and A 169 was fragment size (ha); a fragment is round shaped (isodiametric), when the value of the

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170 shape index is 1, while a value greater than 1 suggest deviation from circularity 171 (Laurence and Yensen 1991). We characterized grazing intensity by the average height 172 of the vegetation. We measured vegetation height monthly in each year within a 1 × 1 m 173 plot next to all traps. Grazing is a traditional management practice in this region; cattle 174 and sheep stay all summer in the grassland fragments. Vegetation height was a reliable 175 proxy for grazing intensity, as there was a significant relationship between grazing 176 intensity (density of grazing ) and vegetation height (r = 0.8472, p < 0.001). 177 Former papers reported that the height of vegetation is more important than the type of 178 grazing animals (e.g. Horváth et al. 2009). For the statistical analyses we used the 179 average value of vegetation height from the three years. Landscape-level factors were 180 calculated at 5 scales with varied radii around each studied grassland fragment (100m, 181 500m, 1000m, 2000m and 3000m; Fig. 1). Four landscape-level factors were analyzed: 182 (i) isolation, (ii) total area of croplands in the landscape, (iii) total area of forests in the 183 landscape, and (iv) landscape diversity (Table 1). Isolation of a fragment also depends 184 on the size of the nearest fragment; therefore, we measured the isolation of the grassland 185 fragments by the inverse isolation index (Vos and Stumpel 1995), defined as the total 186 dry sandy grassland area within a given radius around the investigated grassland 187 fragment. This value decreases as the isolation of the grassland fragment increases 188 (Magura et al. 2001). We identified the habitat patches based on aerial photographs 189 within the buffer around the investigated grassland fragments. We expressed the 190 landscape diversity with the Shannon diversity based on the area of eight habitat types: 191 grasslands, croplands, forests, orchards, artificial areas, anthropogenic areas, marshes, 192 fens and water bodies. 193 The effects of local and landscape-level factors on species richness of spiders were 194 analysed separately for each scale with radii between 100m and 3000m. To determine 195 the spatial scale at which the surrounding landscape was relevant, first we fitted the full 196 model including each (local and landscape-level) factor by generalized linear models 197 using the multiple regression design based on Akaike’s Information Criterion (GLMs, 198 Bolker et al. 2009). Model with the lowest AIC was accepted as the final model. 199 Secondly, we plotted the goodness of fit of the final model against the 5 scales for 200 which landscape-level factors were calculated. That spatial scale, at which the final 201 model had the highest value of goodness of fit, was regarded as relevant spatial scale. 202 We used quasi-Poisson log link function to account for overdispersion in the data 203 (Bolker et al. 2009). We pooled the spider catches from the three years before the 204 statistical analyses to ensure a complete species inventory. Spider species with different 205 habitat affinity (specialist and generalist) and hunting strategy (hunting and web- 206 building species) respond variously to local factors (fragment size: Cronin et al. 2004; 207 Horváth et al. 2013; shape of grassland: Barbaro et al. 2005; grazing intensity: Horváth 208 et al. 2009; Kovac and Mackay 2009) and landscape-level factors (isolation: Horváth et 209 al. 2013; Jonsson et al. 2009; area of forests in the landscape: Caprio et al. 2015; Gallé 210 et al. 2011). Thus, it is essential to investigate species with different habitat affinity and 211 hunting strategy separately. Besides the total number of species, we also analyzed the

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212 number of habitat specialist and generalist species, in addition the number of hunting 213 and web-building species. We considered those species as generalists which can be 214 found in several open habitat types. We regarded the following species as habitat 215 specialists: (i) species which occur solely in open, sandy habitats, (ii) species which 216 occur in severe open habitat types, but in lowlands they live only in open, sandy habitats 217 (Buchar and Ruzicka 2002). We considered those spiders as hunting species which 218 move throughout their habitats and find their prey or attack their prey from ambush 219 (Uetz et al. 1999). We regarded those spiders as web-building species which use 220 different types of webs (orb webs, space webs and sheet webs) to catch their prey (Uetz 221 et al. 1999). For every response variable, the goodness of fit of the final model was the 222 highest at the spatial scale with radius 1000m; thus, we used this spatial scale during the 223 statistical analyses (Fig. 2). We did not find ecologically relevant correlations between 224 studied local and landscape-level factors at spatial scales with radius 1000 m (using 225 Spearman’s rank correlations; results are not shown), thus we did not study interactions 226 between factors when analyzing their effects on the species richness. Neither the local 227 factors (except the vegetation height) nor the landscape-level factors did change during 228 the three-year study period; therefore, we used data from the last study year (2003) for 229 the statistical analyses. 230 231 # Figure 1 and Figure 2 approximately here # 232 233 Results 234 235 During the three-year study, we sampled 4988 individuals of 80 species, including 236 17 habitat specialist species with 793 individuals (Electronic Supplementary Material 237 (ESM) Table 1). The total number of species, the number of habitat generalist species, 238 the number of hunting species and the number of web-building species did not show 239 significant relationship with the fragment size (Table 2), while the number of habitat 240 specialist species increased significantly with the increase of the size of dry sandy 241 grassland fragments (Fig. 3a and Table 2). The other two local factors (shape of 242 grassland fragment and grazing intensity) had no effect on the species richness (Table 243 2). There was no relationship between the number of habitat specialist species and the 244 number of web-building species and the isolation (Table 2). We found a significant 245 positive relationship between the total number of species, the number of generalist 246 species and the number of hunting species and the isolation (Fig. 3b-d and Table 2). The 247 species richness did not depend on the area of croplands, and the area of forests in the 248 landscape, and the landscape diversity (Table 2). 249 250 # Table 2 and Figure 3 approximately here # 251 252 Discussion 253

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254 We examined the effects of local and landscape-level factors on spider assemblages 255 in dry sandy grassland fragments by pitfall trapping. Previous studies (Samu et al. 1996; 256 Topping and Sunderland 1992) emphasised that pitfall trapping under-represented the 257 vegetation-dwelling spiders. In our investigation only 8 of the 80 sampled species were 258 vegetation-dwelling. Therefore, a combination of several sampling methods is needed to 259 have a comprehensive picture of the spider assemblages in the studied grassland 260 fragments. However, the aim of our study was to examine the effects of local and 261 landscape-level factors on ground-dwelling spiders, thus pitfall trapping could be 262 regarded as a reliable sampling method (Gallé et al. 2011; Zulka et al. 2014). 263 We found that both local (fragment size) and landscape-level factors (isolation) 264 played important role in shaping spider assemblages. The fragment size influenced only 265 the number of habitat specialist species, as it is increased with the fragment size. The 266 total number of spiders, the species richness of generalist species and the richness of 267 hunting species increased with the increasing isolation. 268 A significant positive relationship was demonstrated between the total number of 269 animal species and the size of habitat islands in a number of earlier papers (spiders: Toft 270 and Schoener 1983; butterflies: Tscharntke et al. 2002; birds: Watson et al. 2005). 271 Similarly to previous results (Baldissera et al. 2013; Gallé 2008; Horváth et al. 2015; 272 Torma et al. 2014; Webb and Hopkins 1984; Zulka et al. 2014), we also found that 273 overall spider species richness was unrelated to the fragment size. Contrary to the 274 prediction of the classical theory of island biogeography, several authors reported a 275 significant negative correlation between the total number of animal species and the area 276 of habitat islands (spiders: Horváth et al. 2013; Jonsson et al. 2009; beetles: Lövei et al. 277 2006; Magura et al. 2001; Webb and Hopkins 1984). However, similarly to our result, 278 several studies demonstrated significant positive relationship between fragment size and 279 species richness of habitat specialist species (spiders: Bonte et al. 2002; Horváth et al. 280 2013; Scott et al. 2006; Webb and Hopkins 1984; specialist herbivores: Sanchez and 281 Parmenter 2002; Zabel and Tscharntke 1998; birds: Mohd-Azlan and Lawes 2011). The 282 reason for the contradiction is that the original theory of island biogeography has been 283 regarded real islands, which differ considerably from terrestrial habitat islands. Around 284 real islands the neighbouring matrix are usually unsuitable for organisms living on the 285 islands; therefore, the chance of the penetration of species from the neighbouring matrix 286 is very low on these islands. Matrix habitats surrounded terrestrial habitat islands, 287 however, are usually less inhospitable; therefore, terrestrial habitat islands could be 288 inhabited by species from the bordering matrix habitats (Tscharntke et al. 2012). 289 Therefore, it is not surprising that the observed species-area relationship may be 290 positive, negative or neutral, depending on the ratio of habitat specialist and non-habitat 291 specialist species and/or hunting and web-building species. These findings lead to the 292 call for the necessity of refinement of the classical hypothesis (Cook et al. 2002). We 293 have to separate specialist and non-specialist species, as well as hunting and web- 294 building species, when we study the force of this prediction in terrestrial habitat islands. 295 Specialist species truly perceive the habitat fragments as islands and they occur in large

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296 numbers only in these fragments, while non-habitat specialist species, hunting and web- 297 building species can survive and reproduce both in habitat fragments and surrounding 298 matrix (Bonte et al. 2002; Cook et al. 2002; Lövei et al. 2006; Magura et al. 2001; 299 Mohd-Azlan and Lawes 2011). 300 We found no effect of grazing intensity on spider species richness. Contrarily, 301 several earlier studies reported that intensive grazing influenced negatively the diversity 302 of ground-dwelling spiders (Bell et al. 2001; Pétillon et al. 2007). Similarly to our 303 results, other papers found that species richness of ground-dwelling spiders was 304 unaffected by grazing (European grasslands: Batáry et al. 2008; Dennis et al. 2001; Ford 305 et al. 2013; Szinetár and Samu 2012; Australian grasslands: Harris et al. 2003). In the 306 present study none of the examined spider diversity measures decreased significantly by 307 grazing intensity and related trampling, even though both processes change 308 considerably soil and vegetation conditions (Eldridge and Whitford 2009). Trampling 309 reduces soil roughness and causes soil compaction, while grazing decreases the cover, 310 complexity and height of vegetation (Eldridge and Whitford 2009; Horváth et al. 2009). 311 Based on our results, it seems that grazing and trampling change exclusively the species 312 composition. The majority of disturbance-sensitive species disappear from heavily 313 grazed grassland fragments, but several less sensitive species preferring short vegetation 314 and more open and warmer patches can spread from neighbouring grasslands and 315 agricultural areas to these fragments. 316 The isolation is also an important factor in shaping the diversity of habitat specialist 317 and non-habitat specialist species, as well as hunting and web-building species (Horváth 318 et al. 2013; Jonsson et al. 2009; Torma et al. 2014). Nevertheless, several papers did not 319 show relationship between isolation and species richness (Horváth et al. 2013, 2015; 320 Jonsson et al. 2009; Torma et al. 2014; Webb and Hopkins 1984), while other studies 321 demonstrated a significant negative relationship (Miyashita et al. 1998; Toft and 322 Schoener 1983; Usher et al. 1993). There was no positive relationship reported for 323 spiders. Our results showed that the total number of species, the species richness of 324 generalist spiders and the richness of hunting species increased as the isolation of 325 grassland fragments increased. This could be explained by the higher ratio of other 326 habitat types (crops and forests) around the isolated grassland fragments, which may 327 serve as source habitats of the generalist and hunting spiders allowing them to colonize 328 the more isolated grassland fragments. In several studies the correlation between 329 diversity of specialists or web-building species and isolation was significantly negative 330 (Horváth et al. 2013; Jonsson et al. 2009; Zulka et al. 2014), but other papers 331 demonstrated neutral relationship (Cardoso et al. 2010; Horváth et al. 2015; Sanchez 332 and Parmenter 2002). We found no significant relationship between the richness of 333 specialist spiders or the richness of web-building spiders and the isolation. However, 334 isolation has considerable influence on generalist and hunting spiders, which occurred 335 with high species number only in the more isolated fragments. The reason for the lack 336 of isolation effect on specialist and web-building species is that these species have good 337 dispersal ability. Due to their spreading, mainly with ballooning, they can penetrate in a

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338 given fragment not only from adjacent grassland patches, but also from further ones. 339 Thus unfavourable habitats like croplands and non-native tree plantations do not act as 340 barriers for these species. 341 Dry grasslands are the most threatened habitat types in Europe. These grasslands 342 have a high priority in European Union. Thus, protection of this habitat type is 343 important to ensure the survival of their plant and animal communities. Our results 344 suggest that both local and landscape-level factors play important role in maintaining 345 and enhancing spider diversity in sandy grasslands; therefore, during the restoration 346 and/or management of these grassland fragments these local and landscape-level factors 347 must be considered simultaneously. Furthermore, it is recommended to increase the size 348 of the remnant grassland fragments, converting adjacent croplands and non-native tree 349 plantations to grasslands, because it may contribute to maintaining and enhancing the 350 diversity of habitat specialist species. 351 352 Acknowledgements 353 354 The field study was part of the National Biodiversity Monitoring System in Hungary 355 funded by the Ministry of Rural Development. The authors were supported by OTKA K 356 116639, and KH 126477 projects. Trapping and identification is the work of JE and 357 CsSz. The concept of the paper, statistical evaluation and paper writing are the work of 358 RH, TM, BT. 359 360 References 361 362 Albert ÁJ, Kelemen A, Valkó O, Miglécz T, Csecserits A, Rédei T, Deák B, 363 Tóthmérész B, Török P (2014) Secondary succession in sandy old fields: a 364 promising example of spontaneous grassland recovery. Appl Veg Sci 17:214–224 365 Baldissera R, Rodrigues ENL, Hartz SM (2013) Assessment of the probability of 366 colonization of local spider communities in an experimental landscape. J Arachnol 367 41:160167. 368 Barbaro L, Pontcharraud L, Vetillard F, Guyon D, Jactel H (2005) Comparative 369 responses of bird, carabid, and spider assemblages to stand and landscape diversity 370 in maritime pine plantation forests. Ecoscience 12:110–121 371 Batáry P, Báldi A, Samu F, Szűts T, Erdős S (2008) Are spiders reacting to local or 372 landscape scale effects in Hungarian pastures? Biol Conserv 141:2062–2070 373 Batáry P, Holzschuh A, Orci KM, Samu F, Tscharntke T (2012) Responses of plant, 374 insect and spider biodiversity to local and landscape scale management intensity in 375 cereal crops and grasslands. Agric Ecosyst Environ 146:130–136 376 Bell J, Wheater C, Cullen W (2001) The implications of grassland and heathland 377 management for the conservation of spider communities: a review. J Zool 255:377– 378 387

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546 Zabel J, Tscharntke T (1998) Does fragmentation of Urtica habitats affect phytophagous 547 and predatory insects differentially? Oecologia 116:419–425 548 Zulka KP, Abensperg-Traun M, Milasowszky N, Bieringer G, Gereben-Krenn BA, 549 Holzinger W, Hölzler G, Rabitsch W, Reischütz A, Querner P, Sauberer N, 550 Schmitzberger I, Willner W, Wrbka T, Zechmeister H (2014) Species richness in 551 dry grassland patches of eastern Austria: a multi-taxon study on the role of local, 552 landscape and habitat quality variables. Agric Ecosyst Environ 182:25–36 553

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554 Figure captions 555 556 Fig. 1 Digital map of Rohod sampling site, with a high percentage of croplands (54%), 557 moderate forest (24.2%), but little grassland (4.55%) and other habitat types. The circles 558 show the 5 spatial scales of 100, 500, 1000, 2000 and 3000 m radius around the study 559 grassland at which landscape composition was measured. 560 561 Fig. 2 The scale dependence of the strength of relationship between total number of 562 species and studied variables. The value of goodness of fit are plotted against the spatial 563 scale (in m) at which landscape composition was measured. 564 565 Fig. 3 Relationship between the size of dry sandy grassland fragments and the number 566 of sandy grassland specialist species (a), and relationship between the inverse isolation 567 index and the total number of spider species (b), the number of generalist species (c) 568 and the number of hunting species (d). Dashed lines represent the confidence bands (95 569 %). 570

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571 Figure 1. 572

573

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574 Figure 2. 575

576

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577 Figure 3. 578

579 580

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581 Tables 582 583 Table 1. The studied local and landscape-level factors of the sandy grassland fragments in the Nyírség region. Fragments/variables Fragment size Shape index Average height Inverse Total area of Total area of Landscape (ha) of vegetation isolation index forests in the croplands in diversity (cm) (ha) landscape (ha) the landscape (ha) 1. Bagamér 244.8 2.674 15 28.2 1409.0 236.0 0.213 2. Bátorliget 159.6 3.251 15 234.6 503.4 325.6 0.475 3. Hajdúbagos 226.4 2.312 35 7.3 518.6 412.1 0.469 4. Martinka 332.1 2.782 9 76.0 600.0 789.0 0.406 5. Nyíregyháza 173.9 1.204 3 101.9 74.0 485.6 0.540 6. Nyírtura 23.7 1.277 25 25.8 147.3 339.4 0.380 7. Rohod 52.5 1.341 7 38.9 273.9 259.0 0.526 8. Újtanya 1.9 1.220 14 12.1 96.5 236.0 0.426

584

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585 Table 2. Relationship between the number of spider species and the local and landscape-level 586 factors by generalized linear models (GLMs) using the multiple regression design. Significant 587 negative (–) and significant positive (+) relationships are marked. Total Number of Number of Number of Number of number of specialist generalist hunting web-building species species species species species Fragment size not entered +** not entered not entered not entered Shape index not entered not entered not entered not entered not entered Average height of not entered not entered not entered not entered not entered vegetation Inverse isolation –** not entered –*** –*** not entered index Area of croplands not entered not entered not entered not entered not entered in the landscape Area of forests in not entered not entered not entered not entered ns the landscape Landscape diversity not entered not entered not entered not entered not entered 588 **: p < 0.01 589 ***: p < 0.001 590 not entered: the factors were not entered into the final model based on Akaike’s Information 591 Criterion (AIC)

21 Electronic Supplementary Material

Title: Local and landscape-level factors are equally important drivers in shaping ground- dwelling spider assemblages of sandy grasslands Journal: Biodiversity and Conservation Authors: Roland Horváth, Tibor Magura, Béla Tóthmérész*, János Eichardt, Csaba Szinetár *Corresponding author: MTA-DE Biodiversity and Ecosystem Services Research Group, Egyetem tér 1, Debrecen, H-4032 Hungary; e-mail: [email protected]

Table 1. List of species with number of individuals, habitat affinity and hunting strategy according to Buchar and Ruzicka (2002) and Cardoso et al. (2011) in the eight dry sandy grassland fragments from 2001 to 2003. Notations: S = Sandy grassland specialist species, G = Generalist species, F = Forest species, H = Hunter species, W = Web-builder species. Locations: 1 = Bagamér, 2 = Bátorliget, 3 = Hajdúbagos, 4 = Martinka, 5 = Nyíregyháza, 6 = Nyírtura, 7 = Rohod, 8 = Újtanya. Species Habitat Hunting 1 2 3 4 5 6 7 8 affinity strategy Dysderidae Harpactea rubicunda G H 0 0 0 0 1 0 0 0 Eresidae Eresus kollari S W 0 3 3 0 0 0 0 0 phalerata G W 0 0 0 0 0 0 0 4 Steatoda albomaculata G W 0 0 0 0 3 0 0 0 Linyphiidae Ceratinella brevipes G W 1 0 1 0 0 0 0 0 Improphantes geniculatus S W 0 0 0 0 0 0 1 0 Pelecopsis radicicola G W 0 0 0 1 0 0 0 0 Trichoncus affinis F W 0 0 0 0 0 0 14 0 Trichopterna cito S W 0 0 0 0 0 1 0 0 Tetragnathidae Pachgnytha clercki G W 0 0 0 1 0 1 0 0 Pachygnatha degeeri G H 0 1 0 0 1 10 0 4 Araneidae Neoscona adianta S W 0 2 0 0 0 0 0 0 Lycosidae accentuata G H 22 41 51 6 0 25 24 13 Alopecosa aculeata G H 1 0 1 0 1 0 0 0 Alopecosa cuneata G H 12 0 48 0 0 4 1 7 Alopecosa cursor G H 2 0 1 65 147 2 1 2 Alopecosa mariae G H 8 5 4 0 1 13 60 27 Alopecosa psammophila S H 0 0 1 27 0 0 1 3 Alopecosa pulverulenta G H 10 7 2 3 38 252 3 14 Alopecosa schmidti S H 8 16 21 4 3 0 8 0 Alopecosa sulzeri G H 0 2 0 0 73 9 3 8 Arctosa lutetiana G H 0 1 0 0 0 0 0 0 Arctosa maculata S H 0 0 0 0 1 0 0 0 Hogna radiata G H 0 0 1 0 0 0 0 0 Lycosa sp. H 0 0 1 0 0 0 0 0 Pardosa agrestis G H 0 0 0 0 11 0 1 0 Pardosa alacris F H 1 0 2 0 0 0 0 0 Pardosa bifasciata S H 198 3 247 4 0 3 2 17 Pardosa paludicola G H 0 0 0 0 1 0 0 0 Pardosa palustris G H 2 3 0 2 26 73 62 65 Piratula latitans G H 1 0 0 0 0 0 0 0 robusta G H 1 1 0 0 1 1 1 0 Trochosa ruricola G H 2 0 0 1 0 23 0 4 Trochosa terricola G H 4 3 3 2 0 164 5 10 Xerolycosa miniata S H 0 0 0 5 1 4 8 27 Xerolycosa nemoralis G H 1 29 3 30 61 58 419 393 Hahniidae Hahnia pusilla F W 0 0 0 0 0 1 0 0 Dictynidae Argenna subnigra S W 0 0 0 0 0 1 0 0 Titanoecidae Titanoeca quadriguttata G W 0 0 0 0 0 0 0 2 Miturgidae Cheiracanthium elegans F H 0 0 0 0 0 0 1 0 Zora armillata G H 0 0 0 1 0 0 0 0 Liocranidae Agroeca brunnea F H 1 0 0 0 0 0 0 0 Agroeca cuprea G H 0 0 1 0 0 5 1 0 Clubionidae Clubionidae sp. H 0 0 0 0 0 1 0 0 Corinnidae Phrurolithus festivus G H 0 0 2 0 0 3 0 0 Zodariidae Zodarion germanicum G H 0 0 46 1 0 1 0 0 Gnaphosidae Beralndia cinerea S H 9 0 21 32 0 0 0 0 Callilepis nocturna S H 0 0 0 0 0 1 0 0 Drassodes cupreus G H 0 0 0 0 0 1 1 0 Drassodes lapidosus G H 0 0 0 0 0 0 2 0 Drassodes pubescens G H 2 1 2 0 1 4 0 2 Drassyllus praeficus G H 2 2 14 1 1 58 11 8 Drassyllus pusillus G H 2 0 1 0 2 43 3 4 Gnaphosa mongolica S H 0 0 1 0 0 0 0 0 Haplodrassus dalmatensis G H 0 0 0 0 0 0 1 0 Haplodrassus signifier G H 12 25 23 15 10 5 68 62 Phaeocedus braccatus G H 0 0 0 0 0 1 0 0 Trachyzelotes pedestris G H 1 0 0 0 0 0 0 0 Zelotes aeneus G H 0 0 0 2 0 0 0 0 Zelotes apricorum G H 10 0 3 0 0 3 1 2 Zelotes electus G H 16 10 24 1 8 26 40 13 Zelotes erebeus G H 1 0 1 0 0 0 0 1 Zelotes latreillei G H 1 0 0 0 0 159 4 4 Zelotes longipes G H 95 18 143 42 40 77 142 83 Zelotes subterraneus F H 1 1 0 0 0 0 0 1 Philodromidae Thanatus arenarius G H 12 2 8 0 2 7 9 20 Thomisidae Ozyptila atomaria G H 3 0 1 0 0 3 2 2 Ozyptila scabricula G H 1 1 0 3 39 26 10 10 Xysticus acerbus G H 0 0 3 1 1 0 0 0 Xysticus audax G H 0 1 0 0 0 1 0 0 Xysticus cristatus G H 1 0 2 2 0 0 0 0 Xysticus kochi G H 1 1 5 0 5 0 11 7 Xysticus lineatus G H 0 0 1 0 0 1 1 0 Xysticus mongolicus S H 0 0 0 0 3 0 0 0 Xysticus ninnii S H 16 2 22 1 1 0 0 43 Xystcus sabulosus S H 3 0 0 0 0 0 0 1 Xysticus striatipes G H 0 0 1 1 2 10 4 4 Salticidae Aelurillus v-insignitus S H 1 1 1 3 3 0 0 0 Euophrys frontalis G H 0 0 1 0 0 0 0 0 Phlegra fasciata S H 1 0 0 0 1 1 2 0 Σ 466 182 717 257 489 1082 928 867

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