Breakdown of the species-area relationship in exotic but not in native forest patches

UNCORRECTED PROOF Original article Breakdown of the species-area relationship in exotic but not in native forest patches Tibor Magura a , Andra ´sBa ´ldi b, *,Ro ´bert Horva ´th a a Directorate of the Hortoba ´gy National Park, Sumen u. 2, Debrecen H-4024, Hungary b Animal Ecology Research Group of the Hungarian Academy of Sciences and the Hungarian Natural History Museum, Ludovika te ´r 2, Budapest H-1083, Hungary article info Article history: Received 14 September 2007 Accepted 23 November 2007 Published online - Keywords: Birds Forest specialist species Forestry practice Hungary Generalist species Habitat structure abstract We studied the pattern of bird species richness in native and exotic forest patches in Hungary. We hypothetized that species-area relationship will depend on forest naturalness, and on the habitat specialization of bird species. Therefore, we expected strong species-area relationship in native forest patches and forest bird species, and weaker relationship in exotic forest patches containing generalist species. We censused breeding passerine bird communities three times in 13 forest patches with only native tree species, and 14 with only exotic trees in Eastern Hungary in 2003. Although most bird species (92%) of the total of 41 species occurred in both exotic and native forests, the species-area relationship was significant for forest specialist, but not for generalist species in the native forests. No relationship between bird species and area was found for either species group in the forest with exotic tree species. The comparison of native versus exotic forest patches of similar sizes revealed that only large (>100 ha) native forests harbor higher bird species richness than exotic forests for the forest specialist bird species. There is no difference between small and medium forest patches and in richness of generalist species. Thus, the species-area relationship may diminish in archipelago of exotic habitat patches and/or for habitat generalist species; this result supports the warning that the extension of exotic habitats have been significantly contributing to the decline of natural community patterns. ª2008 Published by Elsevier Masson SAS. 1. Introduction Forests are invaluable for human beings. They are important for recreation and human well being, and provide numerous ecosystem services, and are vital for the maintenance of the majority of biological diversity on Earth (Lacaze, 2000; Dirzo and Raven, 2003; Ozanne et al., 2003; Lewis, 2006). These roles need to be considered when we try to manage forests. For example, the use of wood for heating and timber in construction or paper industry needs fast growing trees, planted in a regular pattern for easy management, without other tree or scrub species. These plantations often are of non-native species and thus are inferior for recreation, and inappropriate for the maintenance of native biodiversity (Koch and Skovsgaard, 1999; Carnus et al., 2006; Gentry et al., 2006). For example, in Hungary 20% of forest cover is the black locust (Robinia pseudoacacia), originally from North America, and 15% is black pine (Pinus nigra), a European species, but not native to * Corresponding author at: Tel.: þ36 1 210 1075; fax: þ36 1 334 2785. E-mail addresses: [email protected],[email protected] (A. Ba ´ldi). available at www.sciencedirect.com journal homepage: www.elsevier.com/locate/actoec ARTICLE IN PRESS 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 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 1146-609X/$ – see front matter ª2008 Published by Elsevier Masson SAS. doi:10.1016/j.actao.2007.11.007 acta oecologica xxx (2008) 1–8 ACTOEC2410_proof  25 February 2008  1/8 Please cite this article in press as: Magura, T. et al., Breakdown of the species-area relationship in exotic but not in native forest patches, Acta Oecolo. (2008), doi:10.1016/j.actao.2007.11.007 UNCORRECTED PROOF Hungary (Ma ´tya ´s, 1997). There are similar patterns for other countries (e.g., EEA, 2006). What are the effects of exotic (i.e. introduced or nonnative) trees and forests on native wildlife? Bird species usually prefer native habitats over exotic patches (Ramos, 1996; Ortega et al., 2006). Native bushes are superior foraging sites for birds (French et al., 2005). Nesting on exotic bushes in urban parks resulted in higher rate of nest failure than nesting on native bushes (Schmidt and Whelan, 1999; Borgmann and Rodewald, 2004). Exotic forest plantations can lure settling birds into such suboptimal habitat, where nest failure is higher than in native forests (Remes, 2003). Knowledge of the effects of exotic tree plantations on birds at the community level is limited. Several studies aimed to compare communities in native and exotic forests, using simple parameters. Species richness and abundance is less variable than species composition in native versus exotic forests (Hausner et al., 2002; Johnson and Freedman, 2002; Steverding and Leuschner, 2002; Bakker and Higgins, 2003), although some studies have failed to detect any differences (Donald et al., 1998; Fleishman et al., 2003; Wilson et al., 2006). The underlying mechanism of community differences between native and exotic forests may be the predation pressure on avian broods (Barber et al., 2001; Carignan and Villard, 2002), or the selective habitat preferences of species (Lerner and Stauffer, 1998). However, we have been unable to find any comparison of the species-area relationship (SAR) in exotic versus native forests. This is surprising, because SAR is a basic rule of ecology, stating that species richness increases with increasing sample area (Rosenzweig, 1995; Ba ´ldi and McCollin, 2003; Drakare et al., 2006). Thus, the important question is whether a fundamental ecological relationship is altered by one of the most peculiar and pervasive human activities of modern times – to introduce plant species to areas outside their native ranges. In this study we investigated if exotic forests harbour fewer species than native patches. We compared the species-area relationship of bird assemblages in native versus exotic forest patches in Eastern Hungary. We hypothetized that SAR will depend on forest naturalness, and on the habitat specialization of species. Naturalness probably acts via heterogeneity, which is higher in native than in exotic forests (Thompson et al., 2003; Bartha et al., 2006). Forest patches are islands for forest specialist species, but less so for generalist species, which may occur in the surrounding landscape. This may mask the general species-area relationship (e.g. Magura et al., 2001). Therefore, we expect significant SAR in native forest patches and forest specialist bird species, and weak, if any in exotic forest patches with generalist species. Further, we expect similar species richness values in small native and exotic forest patches, where no real interior habitat is available. However, different species richnesses are expected in large patches due to difference in the species-area relationships. Such finding may have important nature conservation consequences for the maintenance of biodiversity. 2. Study area and methods The study area is located in Eastern Hungary on the Szatma ´rBereg plain (Fig. 1)(N48 0505500,E22 3002200). The plain is covered by pastures and agricultural land with scattered forest patches. The natural vegetation of Central-European plains is forest, but the millennia of human activities (e.g., cattle grazing) modified it into primarily grassland areas with small forest patches (Standova ´r and Primack, 2001). Roughly 80% of the region is farmland. The surroundings was similar for all studied patches, grassland and/or arable fields. Native and exotic tree species were present in the patches. We chose 13 forest patches with only deciduous native tree species (Quercus robur,Carpinus betulus,Quercus petraea,Populus canescens,Populus alba, with a few Acer campestre,Salix alba and Fraxinus angustifolia), and 14 with only deciduous exotic tree species (Robinia pseudoacacia,Populus canadensis, with few individuals of Quercus rubra,Acer negundo,Salix matsudana,Amorpha fruticosa and Fraxinus pennsylvanica). The age of studied forest patches was 35–70 years. Three area categories of forest fragments were established: small (<10 ha), medium (10 <100 ha), and large (>100 ha) (Table 1). Within size class fragments with natural tree composition and those with exotic trees were distinguished. There was no significant difference between the mean area of these two groups within area category (t 7 ¼0.289, p¼0.781 for small forests, t 7 ¼ 0.812, p¼0.443 for medium forests, and t 7 ¼2.853, p¼ 0.064 for large forests). Forest patch heterogeneity was estimated by eye as percent cover of shrubs and number of nest holes within the censused areas. Breeding bird communities were censused in the forest patches in the breeding season of 2003. Three censuses were carried out during the season (April, May and early June), only under good weather conditions (no wind or rain), from sun rise to 9 a.m. (Moska ´t, 1987). We applied a standard point census technique (100 m radius, 5 min census time); all birds seen or heard were recorded. Sampling effort was standardized for all patches – since many patches were only a few hectares large, one point per patch at such patches was possible to census. We distinguished between forest specialist and habitat generalist species based on literature data (Snow and Perrins, 1998), considering local conditions. The species-area relationship was established with the most frequently used log-log transformed model (Rosenzweig, 1995), using individual patch areas (not the categories) in the calculation. The number of forest specialist and generalist bird species between the two forest types (native and exotic) with similar size were examined by repeated measure analysis of variance (ANOVA). Forest type (native or exotic) was considered as factor and the time of the counting (April, May and June) were used as repeated measures. The data were normalized by log(xþ1) transformation. When the results of the ANOVA showed that there was difference in the species richness among the forest types, this was tested by a Tukey-type multiple comparison (Sokal and Rohlf, 1981). The analyses were carried out using the SPSS-PC program (SPSS, 1999). 3. Results Shrub cover did not differ significantly between native and exotic forest patches (t 7 ¼0.324, p¼0.755 for small forests, t 7 ¼0.040, p¼0.969 for medium forests, and t 7 ¼0.828, acta oecologica xxx (2008) 1–82 ARTICLE IN PRESS 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 223 224 225 226 227 228 ACTOEC2410_proof  25 February 2008  2/8 Please cite this article in press as: Magura, T. et al., Breakdown of the species-area relationship in exotic but not in native forest patches, Acta Oecolo. (2008), doi:10.1016/j.actao.2007.11.007 UNCORRECTED PROOF p¼0.435 for large forests). There were more nest holes in native forest patches compared to the exotic patches in small and large forests, but not in the medium size category (t 7 ¼3.653, p¼0.008 for small forests, t 7 ¼1.005, p¼0.348 for medium forests, and t 7 ¼2.543, p¼0.038 for large forests). Altogether 41 bird species were observed during the censuses. Most species (92%) were observed in both the native and the exotic forest patches (Appendix). The species-area relationship was significantly positive in the native forest patches for forest specialist species, and positive but not significant for generalist species (Table 2). No species-area relationship were found for bird assemblages in exotic forest patches (Table 2). Comparing bird species richness between native and exotic forest patches of similar size classes revealed that the small and medium patches were not significantly different for either forest specialist or generalist species. In large forests, however, bird species richness was significantly higher in native than in exotic forests for forest specialist species, but not for generalist species (Table 3). This relationship between the native and exotic forest patches seems to be stable, at least within season, because the interaction term of time * naturalness was on no occasion significant (Table 3). The time effect alone indicated significant decline in species number from April to June in the small forest patches (for both forest and generalist species), and for forest specialist species in medium forest patches. The number of generalist species did not change within season in medium-sized forest patches, and none of the groups declined over the season in the large forest patches (Table 3). Fig. 1 – Location of the study area in Eastern Hungary. Triangles represent forest patches of exotic tree species and circles native tree species. The size of the mark represents three area categories of the patches (<10 ha, 10–100 ha, >100 ha). Table 1 – Characteristics of fragments and the number of observed bird species in three size groups of native and exotic forest patches in Eastern Hungary Small Medium Large Native Exotic Native Exotic Native Exotic Number of fragments 5 4 4 5 4 5 Area (ha) s.d. 7.1 1.86 6.6 2.92 41.0 28.94 58.7 34.99 425.0 206.16 130.0 18.71 Number of nestholes s.d. 697 344 55 49 465 447 204 336 725 284 172 351 Cover of shrubs (%) s.d. 44 27.9 38 32.3 33.75 31.5 33 24.4 46 35.0 28 31.2 Number of observed forest specialist species 12.2 1.10 10.5 1.73 13.8 4.99 14.0 5.61 16.5 1.26 12.4 2.19 Number of observed generalist species 3.2 1.64 3.3 2.06 1.8 1.50 3.0 1.58 1.8 0.96 4.0 1.58 acta oecologica xxx (2008) 1–8 3 ARTICLE IN PRESS 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 288 289 290 291 292 293 294 295 296 297 298 299 300 301 302 303 304 305 306 307 308 309 310 311 312 313 314 315 316 317 318 319 320 321 322 323 324 325 326 327 328 329 330 331 332 333 334 335 336 337 338 339 340 341 342 ACTOEC2410_proof  25 February 2008  3/8 Please cite this article in press as: Magura, T. et al., Breakdown of the species-area relationship in exotic but not in native forest patches, Acta Oecolo. (2008), doi:10.1016/j.actao.2007.11.007 UNCORRECTED PROOF 4. Discussion We studied bird communities in a landscape with patches of natural and exotic tree species. The majority of the landscape was farmland (arable fields and grasslands), with scattered forest patches, including the studied patches. Although it is known that different landscape matrix influences species diversity even in similar forest patches (Lindenmayer et al., 2002; Watson et al., 2005), here we had the same landscape type as matrix. Therefore, we supposed that the matrix effect was similar for all patches, thus, did not bias the results. There was no difference in bird species composition of natural versus exotic forest patches. In a study of birds in tree plantations versus native shrub patches in the Negev, Israel, Shochat et al. (2001) found only a small overlap of bird species. Probably the difference between exotic and native patches in our study was not large enough to exclude bird species, only to influence the frequency of occurrence. The communities showed different responses to area; the species-area relationship explained the number of bird species (i.e. significant positive species-area relationship) in native forests, but not in exotic forest patches. A similar pattern was found by Shochat et al. (2001) for native scrub fragments versus planted forests; bird species richness depended on area in the former, but not in the latter. Santos et al. ( Q1 2006) compared native oak and mature pine plantations in Spain, and they found similar bird species richness in native and plantation archipelagoes, and different species-area relationship, just in this study. However, all species-area regression models were significant in their study (Santos et al., 2006). The difference between native and exotic forests was more pronounced, if species were classified according to their specificity to the forest habitat. Species richness of generalists was not related to forest area at all. The species – area relationship of forest specialist bird species was not significant in exotic forest patches, but was significant (positive) in native patches. The clear difference between the response of specialist and generalist (including exotic) species to fragmentation was described for several taxa, including birds (McCollin, 1993; Germaine et al., 1998), plants (Abbott, 1992; Bakker and Higgins, 2003) and invertebrates (Magura et al., 2001; Ostergard and Ehrlen, 2005; Ouin et al., 2006). More generally, species traits are known to confound the SAR (Ewers and Didham, 2006). There may be two potential mechanisms to explain the presence of SARin native, but notin exotic forest patches.First, there is usually a basic difference between exotic and native forests in spatial heterogeneity; native forestsare more heterogeneous (Ma ´tya ´s, 1997; Thompson et al., 2003). We also found some indications of such a trend (more nest holes in native forests), although others were in our case not statistically significant (shrub cover). Habitat heterogeneity and species richness have a positive correlation (Tews et al., 2004), thus, the more heterogeneous and complex habitat structure of native forests may promote forest specialist bird species rather than generalist species (MacNally et al., 2000). Second, native forests are more island-like patches, because their complex structure is more different from the surrounding landscape, than the exotic forests with a simple structure. Therefore, forest specialist species are probably restricted to the native forest ‘‘isolates’’, with the subsequent SAR, while exotic forests might not function as isolates. This landscape effect may also be responsible for the absence of a SAR in some cases (Estades and Temple, 1999; Wethered and Lawes, 2003; Lo ¨vei et al., 2006). Increased habitat fragmentation results in the relative increase of edge habitats due to the increase of edge/core ratio. Edges are favored by generalist and/or early successional species (Harris and Silva-Lopez, 1992; Imbeau et al., 2003; Lo ¨vei et al., 2006), therefore we expected a reverse relationship for generalist species with forest area: as forest area increases the proportion of edges, and the generalists species, is decreasing. Although these were significant neither for native (R¼0.1) nor for exotic (R¼0.1) forests, the trends of the regression coefficients (Table 2) support this hypothesis. The SAR is a fundamental rule of ecology, but it still suffers from several biases (Ba ´ldi and McCollin, 2003). Here we demonstrated the key role of native/exotic species composition, that is the quality of habitats and the specialist/generalists character of target species on the SAR (Lo ¨vei et al., 2006). An archipelago of non-native habitats, and/or non-native species in the archipelago may lead to the breakdown of the SAR. The age of forest patches may be relevant factor in determining species richness. In this study the age of patches were 35–70 years, that none of them were old growth. Humphrey (2005) and Santos et al. (2006), for example, showed that 100–200 year old plantations already conferring substantial benefits to many species. It is clear that for the studied forest archipelago and landscape only large (>100 ha) forest patches containing native tree species can preserve natural patterns. Such patches support forest specialist bird species during the whole breeding season. We warn, however, against using only species presence in forest patches for patch evaluation: on our study nearly all the observed species (92%) were present in both native and exotic forest patches. Therefore, simple presence-absence survey may be misleading, because it can not identify the superior value of native forests. Other studies also highlighted the subordinate role of exotic trees and bushes for nesting, foraging and community assemblage (Schmidt and Whelan, 1999; Remes, 2003; Borgmann and Rodewald, 2004; French et al., 2005). This evidence support the conclusion that the expansion of exotic trees and bushes via forestry practice and gardening will harm natural patterns and the underlying processes, hence accelerating the decline of birds in isolated forests. Table 2 – Equations of the species-area relationship of forest specialist birds in patches of native and exotic trees in Eastern Hungary. Equations are given for forest specialist and generalist species separately. Asterisks indicate significant relationship Equation Rnp Native forests Number of forest specialist species Y¼0.82 þ0.09X0.66 13 0.02* Number of generalist species Y¼0.46 0.01X0.07 13 0.84 Exotic forests Number of forest specialist species Y¼0.76 þ0.09X0.40 14 0.16 Number of generalist species Y¼0.47 þ0.03X0.10 14 0.73 acta oecologica xxx (2008) 1–84 ARTICLE IN PRESS 343 344 345 346 347 348 349 350 351 352 353 354 355 356 357 358 359 360 361 362 363 364 365 366 367 368 369 370 371 372 373 374 375 376 377 378 379 380 381 382 383 384 385 386 387 388 389 390 391 392 393 394 395 396 397 398 399 400 401 402 403 404 405 406 407 408 409 410 411 412 413 414 415 416 417 418 419 420 421 422 423 424 425 426 427 428 429 430 431 432 433 434 435 436 437 438 439 440 441 442 443 444 445 446 447 448 449 450 451 452 453 454 455 456 ACTOEC2410_proof  25 February 2008  4/8 Please cite this article in press as: Magura, T. et al., Breakdown of the species-area relationship in exotic but not in native forest patches, Acta Oecolo. (2008), doi:10.1016/j.actao.2007.11.007 UNCORRECTED PROOF Acknowledgement We are indebted to Duncan McCollin, Be ´la To ´thme ´re ´sz and two anonymous referees for comments on previous versions of the paper. The research was supported by a grant of the Hungarian Scientific Research Fund (OTKA research grant no. F61651, to T. M.), and the Directorate of the Hortoba ´gy National Park. T. M. and A. B. were Bolyai Research Fellows of the Hungarian Academy of Sciences. Table 3 – Results of the repeated measures ANOVA for the bird species richness in deciduous forests with similar size. Naturalness of the forests (native, exotic) comprised the factor and the time of the counting (April, May and June in 2003) were used as repeated measures. Results of the Tukey test indicate which forest category differs significantly (p<0.05) from the other; for example ‘Native >Exotic’ indicates that the species richness was significantly higher in the native forests than in the exotic patches Variable Source SS df MS FpTukey posteriori test Small forests, Number of forest specialist species Within-Subjects Effects Time 0.130 2 0.065 18.100 0.000 Time Naturalness 0.004 2 0.002 0.558 0.585 Error 0.050 14 0.004 Between-Subjects Effects Naturalness 0.013 1 0.013 0.653 0.446 Error 0.143 7 0.020 Small forests, Number of generalist species Within-Subjects Effects Time 0.150 2 0.075 4.830 0.025 Time Naturalness 0.043 2 0.021 1.382 0.283 Error 0.217 14 0.016 Between-Subjects Effects Naturalness 0.028 1 0.028 0.294 0.604 Error 0.662 7 0.095 Medium forests, Number of forest specialist species Within-Subjects Effects Time 0.080 2 0.040 7.235 0.007 Time Naturalness 0.005 2 0.003 0.474 0.632 Error 0.077 14 0.006 Between-Subjects Effects Naturalness 0.006 1 0.006 0.072 0.797 Error 0.625 7 0.089 Medium forests, Number of generalist species Within-Subjects Effects Time 0.006 2 0.003 0.210 0.813 Time Naturalness 0.084 2 0.042 2.751 0.098 Error 0.213 14 0.015 Between-Subjects Effects Naturalness 0.011 1 0.011 0.255 0.629 Error 0.297 7 0.042 Large forests, Number of forest specialist species Within-Subjects Effects Time 0.021 2 0.011 3.024 0.081 Time Naturalness 0.005 2 0.003 0.734 0.498 Error 0.050 14 0.004 Between-Subjects Effects Naturalness 0.162 1 0.162 10.228 0.015 Native >Exotic Error 0.111 7 0.016 Large forests, Number of generalist species Within-Subjects Effects Time 0.000 2 0.000 0.005 0.995 Time Naturalness 0.031 2 0.016 0.458 0.642 Error 0.478 14 0.034 Between-Subjects Effects Naturalness 0.077 1 0.077 1.012 0.348 Error 0.536 7 0.077 acta oecologica xxx (2008) 1–8 5 ARTICLE IN PRESS 457 458 459 460 461 462 463 464 465 466 467 468 469 470 471 472 473 474 475 476 477 478 479 480 481 482 483 484 485 486 487 488 489 490 491 492 493 494 495 496 497 498 499 500 501 502 503 504 505 506 507 508 509 510 511 512 513 514 515 516 517 518 519 520 521 522 523 524 525 526 527 528 529 530 531 532 533 534 535 536 537 538 539 540 541 542 543 544 545 546 547 548 549 550 551 552 553 554 555 556 557 558 559 560 561 562 563 564 565 566 567 568 569 570 ACTOEC2410_proof  25 February 2008  5/8 Please cite this article in press as: Magura, T. et al., Breakdown of the species-area relationship in exotic but not in native forest patches, Acta Oecolo. (2008), doi:10.1016/j.actao.2007.11.007 UNCORRECTED PROOF Appendix List of observed bird species in taxonomic order from 27 forest patches in Eastern Hungary. Total number of observations during the three censuses in 2003 is given. Snow and Perrins (1998) was followed for nomenclature. Asterisks indicate forest specialist species Small Medium Large Native Exotic Native Exotic Native Exotic Columbidae Columba palumbus Woodpigeon 062300 Streptopelia turtur Turtle Dove 774556 Upupidae Upupa epops Hoopoe 000002 Picidae Dendrocopos major* Great Spotted Woodpecker 723689 Dendrocopos medius* Middle Spotted Woodpecker 000010 Dendrocopos minor* Lesser Spotted Woodpecker 002100 Dryocopus martius* Black Woodpecker 101430 Jynx torquilla* Wryneck 641339 Picus viridis* Green Woodpecker 001001 Motacillidae Anthus trivialis Tree Pipit 6 4 3 12 2 2 Turdidae Erithacus rubecula* Robin 12 4 8 12 8 5 Luscinia megarhynchos* Nightingale 10 18 4 13 3 6 Turdus merula* Blackbird 14 9 11 12 20 8 Turdus philomelos* Song Trush 100230 Sylviidae Locustella fluviatilis* River Warbler 131209 Phylloscopus collybita* Chiffchaff 9389128 Phylloscopus sibilatrix* Wood Warbler 6567121 Phylloscopus trochilus* Willow Warbler 011020 Sylvia atricapilla* Blackcap 22 13 18 19 18 13 Sylvia curruca Lesser Whitethroat 010000 Muscicapidae Muscicapa striata* Spotted Flycatcher 001221 Ficedula sp.* Flycather sp. 000010 Aegithalidae Aegithalos caudatus* Long-tailed Tit 001100 Paridae Parus ater* Coal Tit 000030 Parus caeruleus* Blue Tit 3236114 Parus major* Great Tit 17 8 11 12 14 9 Parus palustris* Marsh Tit 000001 Sittidae Sitta europaea* Nuthatch 104232 Certhiidae Certhia sp.* Treecreeper 10 848510 Oriolidae Oriolus oriolus* Golden Oriol 9 9 4 12 8 14 Laniidae Lanius collurio Red-backed Shrike 120304 Corvidae Garrulus glandarius* Jay 102321 Sturnidae Sturnus vulgaris Starling 21 21 17 8 10 48 Passeridae Passer montanus Tree Sparrow 000100 Fringillidae Carduelis carduelis Goldfinch 020102 Carduelis chloris* Greenfinch 320227 Coccothraustes coccothraustes* Hawfinch 6 13 6 7 11 4 Fringilla coelebs* Chaffinch 29 23 28 24 33 37 Serinus serinus Serin 000001 acta oecologica xxx (2008) 1–86 ARTICLE IN PRESS 571 572 573 574 575 576 577 578 579 580 581 582 583 584 585 586 587 588 589 590 591 592 593 594 595 596 597 598 599 600 601 602 603 604 605 606 607 608 609 610 611 612 613 614 615 616 617 618 619 620 621 622 623 624 625 626 627 628 629 630 631 632 633 634 635 636 637 638 639 640 641 642 643 644 645 646 647 648 649 650 651 652 653 654 655 656 657 658 659 660 661 662 663 664 665 666 667 668 669 670 671 672 673 674 675 676 677 678 679 680 681 682 683 684 ACTOEC2410_proof  25 February 2008  6/8 Please cite this article in press as: Magura, T. et al., Breakdown of the species-area relationship in exotic but not in native forest patches, Acta Oecolo. 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Effects of growth stage and tree species composition on breeding bird assemblages of plantation forests. Bird Study 53, 225–236. acta oecologica xxx (2008) 1–88 ARTICLE IN PRESS 799 800 801 802 803 804 805 806 807 808 809 810 811 812 813 814 815 816 817 818 819 820 821 822 823 824 825 826 827 828 829 830 831 832 833 834 835 836 837 838 839 840 841 842 843 844 845 846 847 848 849 850 851 852 ACTOEC2410_proof  25 February 2008  8/8 Please cite this article in press as: Magura, T. et al., Breakdown of the species-area relationship in exotic but not in native forest patches, Acta Oecolo. (2008), doi:10.1016/j.actao.2007.11.007