16
UNCORRECTED PROOF UNCORRECTED PROOF RESEARCH ARTICLE 1 2 The Pan European Ecological Network: PEEN 3 Rob H. G. Jongman Irene M. Bouwma 4 Arjan Griffioen Lawrence Jones-Walters 5 Anne M. Van Doorn 6 Received: 22 March 2010 / Accepted: 23 December 2010 7 Ó Springer Science+Business Media B.V. 2010 8 Abstract The pan European biological and land- 9 scape diversity strategy (PEBDLS) was developed 10 under the auspices of the Council of Europe in order 11 to achieve the effective implementation of the 12 convention of biological diversity (CBD) at the 13 European level. A key element of PEBLDS has been 14 the development of the Pan European Ecological 15 Network (PEEN) as a guiding vision for coherence in 16 biodiversity conservation. PEEN has been developed 17 in three subprojects: Central and Eastern Europe, 18 completed in 2002; South-eastern Europe, completed 19 in 2006; and Western Europe, also completed in 20 2006. The methodology of the development of the 21 three maps has been broadly comparable but data 22 availability, differences in national databases, tech- 23 nical developments and geographical differences 24 caused variations in the detailed approach. One of 25 the challenges was to find common denominators for 26 the habitat data in Europe; this was solved differently 27 for the subprojects. The project has resulted in three 28 maps that together constitute the PEEN. They differ 29 in terms of ecological coherence and the need for 30 ecological corridors; for example, in Central and 31 Western Europe corridors are essential to provide 32 connectivity, while in Northern, Eastern and South- 33 eastern Europe larger, coherent natural areas still 34 exist. The future steps in developing PEEN should 35 include the implementation of national ecological 36 networks and, in particular, the pursuit of interna- 37 tional coherence through the development of trans- 38 European ecological corridors. The big challenge is 39 to develop a common approach among the over 100 40 European-wide agencies that are responsible for 41 biodiversity conservation. 42 Keywords Ecological network Á Europe Á 43 Ecological corridors Á Indicator species Á Habitat 44 suitability Á Ecological data 45 46 47 Introduction 48 The concept of ecological networks is not new. 49 Ecological networks have been developed over a 50 number of decades in several European countries; 51 beginning in Estonia and former Czechoslovakia in 52 the 1980s and in The Netherlands in 1989 (Jongman 53 et al. 2004). In these countries a strong land use 54 planning tradition created the institutional environ- 55 ment for allocating functions, including the recogni- 56 tion of nature as a designation with its own value. A1 R. H. G. Jongman (&) Á I. M. Bouwma Á A2 A. Griffioen Á A. M. Van Doorn A3 Alterra, Wageningen UR, PO Box 47, 6700 AA A4 Wageningen, The Netherlands A5 e-mail: [email protected] A6 L. Jones-Walters A7 European Centre for Nature Conservation (ECNC), A8 PO Box 90154, 5000 LG Tilburg, The Netherlands 123 Journal : Medium 10980 Dispatch : 31-12-2010 Pages : 16 Article No. : 9567 h LE h TYPESET MS Code : LAND-10-2347 h CP h DISK 4 4 Landscape Ecol DOI 10.1007/s10980-010-9567-x Author Proof

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RESEARCH ARTICLE1

2 The Pan European Ecological Network: PEEN

3 Rob H. G. Jongman • Irene M. Bouwma •

4 Arjan Griffioen • Lawrence Jones-Walters •

5 Anne M. Van Doorn

6 Received: 22 March 2010 / Accepted: 23 December 20107 � Springer Science+Business Media B.V. 2010

8 Abstract The pan European biological and land-

9 scape diversity strategy (PEBDLS) was developed

10 under the auspices of the Council of Europe in order

11 to achieve the effective implementation of the

12 convention of biological diversity (CBD) at the

13 European level. A key element of PEBLDS has been

14 the development of the Pan European Ecological

15 Network (PEEN) as a guiding vision for coherence in

16 biodiversity conservation. PEEN has been developed

17 in three subprojects: Central and Eastern Europe,

18 completed in 2002; South-eastern Europe, completed

19 in 2006; and Western Europe, also completed in

20 2006. The methodology of the development of the

21 three maps has been broadly comparable but data

22 availability, differences in national databases, tech-

23 nical developments and geographical differences

24 caused variations in the detailed approach. One of

25 the challenges was to find common denominators for

26 the habitat data in Europe; this was solved differently

27 for the subprojects. The project has resulted in three

28 maps that together constitute the PEEN. They differ

29 in terms of ecological coherence and the need for

30ecological corridors; for example, in Central and

31Western Europe corridors are essential to provide

32connectivity, while in Northern, Eastern and South-

33eastern Europe larger, coherent natural areas still

34exist. The future steps in developing PEEN should

35include the implementation of national ecological

36networks and, in particular, the pursuit of interna-

37tional coherence through the development of trans-

38European ecological corridors. The big challenge is

39to develop a common approach among the over 100

40European-wide agencies that are responsible for

41biodiversity conservation.

42Keywords Ecological network � Europe �

43Ecological corridors � Indicator species � Habitat

44suitability � Ecological data

45

46

47Introduction

48The concept of ecological networks is not new.

49Ecological networks have been developed over a

50number of decades in several European countries;

51beginning in Estonia and former Czechoslovakia in

52the 1980s and in The Netherlands in 1989 (Jongman

53et al. 2004). In these countries a strong land use

54planning tradition created the institutional environ-

55ment for allocating functions, including the recogni-

56tion of nature as a designation with its own value.

A1 R. H. G. Jongman (&) � I. M. Bouwma �

A2 A. Griffioen � A. M. Van Doorn

A3 Alterra, Wageningen UR, PO Box 47, 6700 AA

A4 Wageningen, The Netherlands

A5 e-mail: [email protected]

A6 L. Jones-Walters

A7 European Centre for Nature Conservation (ECNC),

A8 PO Box 90154, 5000 LG Tilburg, The Netherlands

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57 One of the main drivers for the development of

58 ecological networks is that biodiversity in Europe

59 was continuing to decline due to a decrease of habitat

60 quality, habitat extent and increase in fragmentation.

61 The ecological consequences of these processes are

62 the destabilisation of landscape level processes,

63 diminishing ecosystem functions, declining popula-

64 tions of natural species and a threat to their collective

65 sustainability (Rodoman 1974; Jongman 2002; Man-

66 der et al. 2003). There was therefore a realisation that

67 for many natural species the existing area and

68 distribution of nature reserves and national parks

69 was too small and fragmented (Somma et al. 2004;

70 Hepcan et al. 2009).

71 The concept of ecological connectivity is implicit

72 in several international conventions such as the

73 Ramsar convention and the Bern convention, Euro-

74 pean agreements (habitats and species directive) and

75 related EU policy implementation (Natura 2000). It

76 has become operational in national strategies

77 (national ecological networks) and at European level

78 through the pan European biological and landscape

79 diversity strategy (PEBLDS), Council of Europe

80 1996). The PEBLDS was Europe’s response to the

81 need for effective implementation of the convention

82 of biological diversity (CBD) at the European level;

83 the Pan European Ecological Network—PEEN

84 formed one of its central delivery mechanisms. The

85 indicative map of the pan-European ecological net-

86 work was developed in the context of the following

87 aim (Council of Europe 1996): ‘The pan-European

88 ecological network addresses the development of an

89 ecological network at a European level. It will consist

90 of core areas, corridors and buffer zones. Restoration

91 areas will be identified where they are considered

92 necessary. The pan-European ecological network

93 aims to conserve the full range of ecosystems,

94 habitats, species and landscapes of European impor-

95 tance and to counteract the main causes for decline

96 by creating the right spatial and environmental

97 conditions’.

98 The aim of establishing ecological networks is

99 therefore the protection of nature and biodiversity.

100 Their development has been stimulated by a combi-

101 nation of science and nature management practice.

102 In the USA and some European countries (e.g.

103 Portugal), the ecological network is termed ‘Green-

104 way’ (Ahern 2004) and it attempts to integrate wider

105 societal interests with biodiversity conservation such

106as outdoor recreation and builds on the tradition of

107greenbelt planning and parkway planning (Jongman

108and Pungetti 2004). Bennet and Wit (2001) and

109Bennett (2004) define an ecological network as,

110‘‘A coherent system of natural and/or semi-natural

111landscape elements that is configured and managed

112with the objective of maintaining or restoring

113ecological functions as a means to conserve biodi-

114versity while also providing appropriate opportunities

115for the sustainable use of natural resources’’. The

116aims of the pan-European ecological network were

117further elaborated to ensure that (Rientjes and

118Roumelioti 2003):

119– a full range of good quality ecosystems, habitats,

120species and landscapes of European importance is

121conserved;

122– habitats are large enough to guarantee key species

123a favourable conservation status;

124– sufficient opportunities exist for dispersal and

125migration of species;

126– damaged parts of the key environmental systems

127are restored; and

128– the key environmental systems are buffered from

129threats.

130The Pan European Ecological Network presents a

131vision for the development of coherency in ecological

132networks throughout Europe and its implementation

133by national and regional governments. A method was

134therefore developed to take into consideration both a

135European perspective and national ecological net-

136works and, when applicable, to include regional

137networks developed within countries.

138The Pan European Ecological Network project

139was developed in three stages. The first PEEN

140subproject was carried out for Central and Eastern

141Europe (Bouwma et al. 2002) and had to deal with

142restricted data availability. This was also the case for

143the second subproject in South-eastern Europe that

144covers the Balkan, Greece and Turkey (Biro and

145Grobelnik 2006). Although there was not a perfect set

146of data for the third subproject in Western Europe,

147this subproject was able to make use of developments

148in data availability and new ancillary data (Jongman

149et al. 2006a).

150The indicative map of the pan-European ecolog-

151ical network in Central and Eastern Europe (PEEN-

152CEE) was presented at the fifth ‘Environment for

153Europe’ ministerial conference in Kyiv, in May 2003,

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154 where the results were received as an important

155 contribution to the establishment of PEEN and the

156 promotion of ecological networks in general. At the

157 Belgrade ministerial conference in 2007 the whole

158 process and all three maps were reported (Bonnin

159 et al. 2007).

160 This paper presents the common approach and

161 regional differences in relation to the PEEN project

162 for the development of the PEEN vision and maps for

163 Central and Eastern Europe (PEEN-CEE), South-

164 eastern Europe (PEEN-SEE) and Western Europe

165 (PEEN-WE). It presents the results and discusses the

166 differences.

167 Data and methods

168 All three of the pan European networks maps were

169 developed at a scale of 1:3,000,000. For the design of

170 the map data were used that were consistently

171 available for Europe or the project area. Species

172 with a widespread occurrence in Europe, but for

173 which data were not available for all Europe or at

174 least for the project area, were not used. They would

175 have been confused with species or habitats with a

176 regionally specific distribution. In the development of

177 the maps two lines of analysis were followed: (1) the

178 analysis of habitats; and (2) the analysis of species

179 requirements in relation to habitat size, quality and

180 extent. Both lines were integrated in the final stage

181 (Fig. 1). The core of the methodology was the same

182 for all three maps. Adaptations were made in the

183 methodology to take the differences between regions

184 into account. For PEEN-CEE the land cover data that

185 were used for the determination of core areas of the

186 network were only available at different levels of

187 accuracy; this caused differences in overall accuracy

188 of determining habitats (Table 1). Three land cover

189 databases were used: CORINE 2000 (http://www.eea.

190 europa.eu) for the EU (then candidate) countries;

191 PELCOM (Mucher et al. 2001) for Belarus, Ukraine

192 and the western part of Russia; and the IGBP-DIS for

193 the eastern part of European Russia (Loveland and

194 Belward 1997). The IGBP-DIS was used as common

195 denominator as it is the most general map for anal-

196 ysis. In PEEN-WE the Norwegian and Swiss national

197 land cover maps had to be integrated with the CO-

198 RINE land cover map to develop a composite habitat

199 map. General habitat categories (GHC’s), Bunce

200et al. 2008) have been used to develop a common

201legend. In all phases of the projects intensive inter-

202action was maintained with stakeholders from policy

203and science in order to check the validity of the

204scenarios and ecological content.

205The PEEN is based on habitat maps showing

206existing non-fragmented natural and semi-natural

207areas that are considered large enough to sustain

208viable populations of large species and species of

209European importance requiring large areas. The

210subproject PEEN-WE includes Europe from the Polar

211Circle in Norway through to Crete, Sicily and

212Gibraltar. Madeira and the Canary islands have

213not been included as they are a separate biome

214(Macaronesian) and are situated at a large distance

215from the European mainland. However, for nature

216conservation they are of utmost importance because

217of their tertiary island endemics. The area needed for

218the survival of species differs largely (from one to

219another) in particular due to climatic and geograph-

220ical conditions. Habitat maps had to be developed

Land cover Europe

EnZ

Land Cover per EnZ

DEM

Land cover per AEnZ

Species list Europe

EnZ

Species list per EnZ

Habitatconditions

Habitat conditions per species and AEnZ

Habitat map per zone

Merging into PEEN-WE habitat map

Fig. 1 Procedure for identifying core areas for the PEEN-WE

region

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221 based on land cover data and ancillary data, such as

222 digital elevation models, protected areas databases

223 and important bird areas; this was because land cover

224 maps do not deliver sufficient ecological information

225 as most data they provide are too general and do not

226 include species dynamics and geographical and

227 biogeographical differences (Table 1). For PEEN-

228 WE (Jongman et al. 2006a) use was made of the

229 environmental stratification of Europe (Metzger et al.

230 2005) so that habitats in different biogeographical

231regions could be identified. As mountain ranges play

232an important role use was also made of a digital

233elevation model (DEM) of Europe in order to

234distinguish between high mountains and lowlands.

235In the PEEN-CEE and the PEEN-SEE subprojects

236shortcomings were observed by experts reviewing the

237results. These were mainly that the method applied

238might underestimate the value of fragmented land-

239scapes and small scale (hedgerow) landscapes; only

240large unfragmented landscapes are identified in land

Table 1 Data used in the PEEN projects

Dataset Source Used for Coverage

Topographic information

Digital elevation model USGS Data centre Habitat types All

Hydrology (rivers, lakes) ESRI, WDBII, Bartholomew Corridors All

National borders ESRI, WDBII Background All

Towns and cities GEOnet Names Server Background All

Roads ESRI Counterchecking corridors All

Rivers Bartholomew River corridors All

Internationally protected areas

Ramsar sites UNEP-WCMC Protected areas All

World heritage sites UNEP-WCMC Protected areas All

Man and biosphere reserve sites UNEP-WCMC Protected areas All

Natura 2000 European Environmental Agency Protected areas EU15

Protected areas norway UNEP On line GIS and map database Protected areas Norway

Internationally acknowledged areas

Important bird areas BirdLife International Protected areas All

Prime butterfly areas Butterfly foundation Protected areas All

Landcover information

PELCOM Alterra Habitat types Russia (not entirely to

Ural)

IGBP-DIS global land cover IGBP Habitat types Eastern Russia

Corine land cover (CLC2000) European environment agency Habitat types EU15

LC Database Norway Nijos Habitat types Norway

LC database Switzerland WSL Habitat Types Switzerland

Peatlands Wetland International Habitat types CEE-region

Coastal systems of Europe EUCC Habitat types West and Central Europe

Boreal forests Taiga rescue network Habitat types Russia

Other

Biogeographical regions European environment agency Habitat types All

Potential natural vegetation Alterra Habitat types All

Soil map FAO-Unesco Soil Database Habitat types All

European environmental

stratification

Wageningen UR Habitat types All

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241 cover maps. In the PEEN-WE subproject, where this

242 is even more important, small scale areas have been

243 analysed separately for their connectivity function.

244 Here regional expert knowledge was an important

245 source of information as land cover data deliver

246 insufficient information; it was used specifically for a

247 second round of analysis to check for connectivity

248 especially in urbanised and small scale agricultural

249 landscapes (Denmark, Germany, The Netherlands).

250 In order to identify existing non-fragmented nat-

251 ural and semi-natural areas considered large enough

252 to sustain viable populations of species of European

253 importance the following steps were conducted

254 (Fig. 1):

255 – development of a combined land cover map for

256 the entire region (CORINE, Switzerland and

257 Norway for PEEN-WE; CORINE, PELCOM

258 and IGBP for PEEN-CEE);

259 – development of a simplified but ecologically

260 interpreted habitat classification map based on

261 land cover data and ancillary data;

262 – identification of habitats of sufficient size accord-

263 ing to the working scale of the map, in order to

264 identify size classes based on indicator species;

265 – identification of Emerald Network sites (the

266 Council of Europe in 1996 launched the Emerald

267 Network, which is based on the same principles

268 as the legally binding Natura 2000 network within

269 the European Union (Council of Europe 2009),

270 Zapovedniks (nationally protected sites in Russia,

271 Ukraine and Belarus (Sobolev et al. 1995) and

272 NATURA 2000 sites [protected under the EU

273 Species and Habitats Directive, Commission of

274 the European Communities (2003)];

275 – identification of Ramsar sites, important bird

276 areas (IBAs) and prime butterfly areas;

277 – identification of area requirements of indicator

278 species;

279 – linking indicator species to the identified habitats;

280 and

281 – estimation of the area required in order to

282 maintain sustainable populations of the indicator

283 species.

284 In all three subprojects the first step was to develop

285 a combined land cover map for the region based

286 on the existing land cover data. In Western Europe

287 the integration was made by using general habitat

288 categories (GHC’s) that allowed the three maps to be

289linked (Bunce et al. 2008). Based on the compiled

290land cover map of the region, habitats were identified

291by overlaying various land cover classes with addi-

292tional information about the environmental zones of

293Europe (EnZ), (Fig. 2), altitude and soil information

294(wet soils, calcareous soils, etc.) using:

295– size of the habitat area;

296– naturalness for the biogeographic region/environ-

297mental zone; which in practice meant that most

298coniferous forests in the Continental and Atlantic

299zones were not included as their function is

300predominantly for wood production, while the

301coniferous forests in the high Alpine region were

302included because they are natural; and

303– importance for natural species occurring in the

304region.

305The next step in the identification of large non-

306fragmented areas was the selection of indicator

307species and then to link them to the identified

308habitats (Fig. 1). This was done by analysis of

309existing priority species data and the habitat require-

310ments of their populations. Experts in various coun-

311tries were consulted about certain species and about

312the basic lists. Given the scale of the map, the grain of

313the land cover information and the available species

314data, it was decided to consider only larger mammal

315and bird species as possible indicator species for

316PEEN-WE and PEEN-SEE. Migrating fish were

317included for PEEN-CEE through an assessment of

318the accessibility of rivers and the presence of existing

319barriers. In PEEN-SEE they have not been included;

320in PEEN-WE they were substituted by the occurrence

321of natural, non-regulated larger rivers that are con-

322sidered as a proxy for the existence of migrating fish

323(in both subprojects these measures were taken

324because of a lack of harmonised European data).

325For PEEN-WE it was possible to use the European

326environmental stratification (Fig. 2) to select differ-

327ent habitat sizes for species in different parts of

328Europe; for example, the potential density of wolf

329and brown bear is lower in boreal Scandinavia than in

330the Iberian Peninsula and the Balkan (Swenson

3312000). Moreover habitats do differ between different

332altitudinal bands. This means that within the envi-

333ronmental stratification for identification of habitats

334substrata have been identified (altitudinal environ-

335mental zones, AEnZ) based on the DEM for Europe

336(Jongman et al. 2006b). Grasslands at higher altitudes

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337 are identified as alpine grasslands, but these altitudes

338 are different for different latitudes. For Alpine North

339 (ALN, Scandinavia) the border between low and high

340 has been identified at 400 m altitude, while in Alpine

341 South (ALS, eg. Alps, Pyrenees, Carpathian) three

342 zones have been identified below 700, from 700 to

343 1200 m and above 1200 m. In the Mediterranean

344 mountains zone (MDM) the difference between low

345 and high is identified at 1000 m. Various types of

346 grasslands such as calcareous grasslands have been

347 identified by interpreting the CORINE land cover

348 map with help of the ancillary data such as the

349 national soil maps and geological data.

350 Mammal and bird species have been selected on

351 their existing international protection status: species

352occurring in the annexes of the EU Habitats and Birds

353Directives (Commission of the European Communi-

354ties 2003), cross checked with the Bern Conven-

355tion—Annex II and the vulnerable-extinct status of

356species according to EBBC or IUCN. For Western

357Europe in total 84 bird species and 14 mammal

358species were selected and then differentiated for

359EnZ’s. For South-eastern Europe 90 bird species and

36020 mammal species (Biro and Grobelnik 2006) and

361for Central and Eastern Europe 115 bird species and

36219 mammal species were selected (Bouwma et al.

3632002).

364The importance of species for PEEN was identi-

365fied in relation to their occurrence in European

366habitats. For PEEN-WE the occurrence of habitats

Fig. 2 Environmental

stratification of Europe

(Metzger et al. 2005). The

zones are ALN Alpine

North, ALS Alpine South,

ATN Atlantic North, ATC

Atlantic Central, LUS

Lusitanian, BOR Boreal,

NEM Nemoral, CON

Continental, PAN

Pannonian, MDN

Mediterranean North, MDM

Mediterranean Mountains,

MDS Mediterranean South

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367 was specified for EnZ’s (Table 2). Within these zones

368 species were linked to habitat types including the

369 threshold estimated for survival of a population per

370 region (Table 3). All the threshold areas were iden-

371 tified for the environmental zones and later merged

372 into the joint legend for European habitats as there

373 are differences in carrying capacity between boreal

374 and mediterranean regions.

375 The size thresholds for different habitat areas were

376 determined to support sustainable populations accord-

377 ing to a number of steps. First species were linked to

378 the identified habitat type and habitat types were

379 identified for each EnZ. Then standards were assigned

380 to each species based on the minimum population size

381 considered large enough to be sustainable in the long-

382 term. This allowed the minimum size of the areas

383 needed to support viable populations of all selected

384 species, per habitat type, to be estimated (Table 3).

385 For determining the viability of species different

386 thresholds in area size per habitat type were set,

387 related to estimated population size. ‘Very large

388 areas’ are defined as: ‘‘greater than five times the

389 minimum area required to make the long-term

390 survival of all populations of the selected species

391 quite probable’’. ‘Large areas’ (defined as: ‘‘two times

392the minimum area required to make the long-term

393survival of all populations of the selected species quite

394probable’’) make it possible for the present population

395of the selected species to occur in that location.

396However, when isolated, these areas may suffer

397the loss of some species and immigration might be

398required; connection to other areas or area enlarge-

399ment is therefore recommended. Areas with a sub-

400optimal size were estimated to maintain a maximum

401of 70% of the selected species populations; the most

402demanding species can only be maintained or restored

403by either enlarging habitat size or by making con-

404nections with comparable habitats by corridors,

405or both.

406The required minimum area sizes (considered as

407sub-optimal for the various habitats) range from:

40850 km2 for wetlands, peat lands and grasslands; to

409300 km2 for different types of forests. Habitat area

410size has been calculated in three classes. The spatial

411patterns of habitat types that exceed each of the

412thresholds were assessed in a GIS analysis.

413The identification of corridors for the pan-Euro-

414pean ecological network in PEEN-CEE was based on

415the location of forested landscapes, known bird

416migration routes, mountain ranges and river related

Table 2 Simplified habitat classification for forest habitats of Western Europe based on CORINE 2000, Norwegian land cover and

the Swiss land cover

Forest Land cover database Habitat Environmental Zone

Scandinavia CORINE Broad-leaved forest ALN, BOR, ATN

CORINE Coniferous forest ALN, ATN

CORINE Mixed forest ALN, NEM, BOR, ATN

Norway Non-productive forest. A lot

of mountain forest

ALN, BOR, NEM

CORINE Transitional woodland-shrub ALN, BOR, NEM

Others CORINE Broad-leaved forest ATC, ATN, LUS, MDM, MDN, MDS, PAN,

CON, ALS

CORINE Coniferous forest ATC, ATN, MDM, MDN, MDS, PAN,

ALS

CORINE Mixed forest ATC, ATN, LUS,MDM, MDN, MDS, PAN,

ALS, CON

CORINE Burnt areas MDM, MDN, MDS, ALS, LUS

Switzerland Forests ALS,MDM, CON

CORINE Transitional woodland-shrub ATC, ATN, LUS,MDM, MDN, MDS, PAN, ALS

Switzerland Scrub and/or herbaceous vegetation

associations

CON

The codes of the environmental zones (EnZ) are given in Fig. 2 (Metzger et al. 2005)

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417 wetlands. All corridors were discussed with regional

418 experts. For PEEN-WE corridors were based on:

419 – bird migration routes;

420 – the need for connectivity between core areas,

421 based on their size required to support the

422 connectivity needs of mobile species that had

423 been identified as related to the habitat;

424 – relative forest density per km2 in areas between

425 large habitat blocks and smaller areas and

426 between small habitat blocks based on forest

427 presence in the land cover map;

428 – location of hedgerow landscapes (small scale farm-

429 ing areas) based on expert information and situated

430 mainly in CORINE land cover classes 20, 21, 22—

431 that include small scale agricultural landscapes

432 potentially functioning as corridors between core

433 areas;

434 – location of mountain ranges (part of habitat

435 classes);

436 – location of natural or semi-natural rivers;

437– expert judgment (project members and consulted

438experts); and

439– comparison with existing national networks (core

440areas and corridors).

441Results

442PEEN project has resulted in three maps: the map for

443Central and Eastern Europe (PEEN-CEE), (Fig. 3);

444the map of South-eastern Europe (PEEN-SEE),

445(Fig. 4); and the map of Western Europe (PEEN-

446WE), (Fig. 5). All three maps have a comparable

447legend showing the three categories for habitat size

448needed for population survival (marginal, 100%,

449greater than five times population size). In all three

450maps existing protected areas have been included

451such as: Zapovedniks in Russia, Belarus and Ukraine;

452Natura 2000 (European Union); and Emerald sites

453(Berne Convention outside European Union); official

Table 3 Mammal species, their occurrence in environmental zones and the estimated claim on habitat

EnZs ALN ALS BOR NEM CON ATN ATC LUS PAN MDN MDM MDS

1 2 3 4 5 6 7 8 9 10 11 12

Castor fiber Beaver 4 4 4 4 4 4 4

Alopex lagopus Polar fox 7

Lutra lutra Otter 4 4 4 4 4 4 4 4 4 4 4

Mustela lutreolaa European mink 5 5

Canis lupusa Wolf 7 6 7 6 6 6

Gulo guloa Forest dormouse 8

Lynx lynx Lynx 8 8 8

Lynx pardinaa Pardel lynx 8 8 8 8

Rupicapra rupicapra Gems/chamois 5

Capra hircus (aegagrus) Ibex 5 5

Ursus arctos Brown bear 8 8 8 8

Cervus elaphus

corsicanus

Red deer 6

Alces alces Moose 7 7

Ramgofer taramdis

femmicus

Reindeer 8

The bold values are excluded in Annex 1 of the European Habitats and Species Directive and therefore not included herea European priority species. Habitat size classes for a sustainable population: 1:\10 km2, 2: 10–50 km2, 3: 50–150 km2, 4:

150–300 km2, 5: 300–600 km2, 6:600–1000 km2, 7: 1000–2000 km2, 8:[2000 km2. Mustela lutreola: Normandy, Brittany and

Luneburger Heide; Canis lupus: all EU 15 except Finland; Lynx lynx: all Europe except Finland and Baltic states; Lynx pardina: only

Iberian Peninsula; Cervus elaphus corsicanus: only Corsica; Rangifer tarandus fenicus: only Finnish forest Reindeer. The following

mammal species have not been considered as they are marine mammals: Monachus monachus, Phoca hispida (Botnica and

saimensis), Halichoerus grypus, Phoca vitulina. For the legend of the EnZ see Fig. 2

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454 Ramsar convention sites. Sites known to be important

455 but not protected, such as important bird areas

456 (IBA’s) and prime butterfly areas, were not included

457 but were used for checking the accuracy of the maps.

458 When needed the map was corrected and the

459 correction was checked with local experts.

460 The PEEN-CEE map (Fig. 3) shows ecological

461 coherence in the Russian taiga forest and tundra that

462still can act as a coherent ecological system. The

463wetlands of Pripjat and Polesia also function as amajor

464core area as do the steppe grasslands in south-east

465European Russia around the delta of the Volga. Alpine

466habitats only occur in the Carpathian and Ural moun-

467tains. In general, towards the south and the west frag-

468mentation has occurred and for ecological coherence

469ecological corridors are needed. They have been

Fig. 3 The map of PEEN for Central and Eastern Europe (Bouwma et al. 2002)

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470 indicated on the map by lines. Two types of corridors

471 are indicated: forested corridors; and river and wetland

472 corridors.

473 In PEEN-SEE (Fig. 4) the major habitat types that

474 form core areas are forests, saline marshes and other

475 marshlands, various types of grasslands and mountain

476 habitats. Alpine habitats are restricted to the eastern

477 part of Turkey, the Rodopi and Balkan Mountains

478 and the Dalmatian Alps. Forest and grassland habitats

479 are dominant. Fragmentation is regional and partly

480 due to differences in natural habitats. All terrestrial

481 corridors are forest corridors as this is the major

482 habitat type. These are indicated by arrows. The

483 major species corridors, however, are the long

484 distance migration corridors for birds from Central

485 and Eastern Europe over the Bosporus and the

486 Caucasus to Africa and western Asia.

487 The PEEN-WE (Fig. 5) map includes major core

488 areas of rivers bogs and wetlands, forest, heathlands

489 and Mediterranean scrub, several grassland types and

490 alpine habitats. Fragmentation is much stronger than

491in the PEEN-CEE and PEEN-SEE. The major

492coherent nature areas are found in Scandinavia, the

493Alps and in the Iberian mountain ranges. Western

494Europe is highly urbanised and therefore nature is

495highly fragmented. The structure of the network

496contains many more ecological corridors than in the

497other two subprojects. However, this process of

498fragmentation is already clearly visible in the western

499part of PEEN-CEE (Fig. 6).

500In all three subprojects ecological corridors have

501been identified. In PEEN-CEE and PEEN-SEE these

502have been mainly indicated on the basis of habitat

503pattern and following consultation with regional

504experts. In PEEN-WE experts were consulted but

505use was also made of existing national ecological

506network (NEN) plans that, in that period, were partly

507available and included ecological corridors. That

508means that the ecological corridors in PEEN-WE

509have different origins. For Denmark ecological

510corridors were proposed based on habitat informa-

511tion; these were then discussed and decided upon in

Fig. 4 The map of PEEN for South-eastern Europe (Biro et al. 2006)

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512 consultation with landscape planning experts. For

513 The Netherlands use was made of the national

514 ecological network and its robust corridors which

515 were included in the rural development plan (Ministry

516 of ANF 2006). For Germany the parallel process of

517 developing the ecological networks for the regional

518 units represented by the Bundeslander was utilised

519(see Ssymank et al. 2005). Use has also been made of

520NGO projects such as the Alpine corridor project of

521WWF. As the maps are ‘live’ documents not all

522information, particularly about ecological corridors,

523could be integrated. For Great Britain the information

524developed by Natural England became available too

525late to be included (Catchpole 2007).

Fig. 5 The map of PEEN

for Western Europe

(Jongman et al. 2006a)

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526 Discussion

527 The PEEN project was initiated during a period of

528 great political and related policy change in Europe;

529 the data that could be used and its quality was

530 therefore very much dependent on the sources and the

531 moment in time that it was abstracted. The land cover

532 data, especially for the PEEN-CEE map, had differ-

533 ent sources with different pixel sizes and legends and

534 direct habitat interpretation was not possible. The

535 available land cover categories had to be utilised, but

536 resulted in coarse habitat categories. It was therefore

537 only possible to use the lowest common denominator

538 in order to develop a common legend. In this case it

539 was the IGBP-DIS, used for eastern part of European

540 Russia (Loveland and Belward 1997).

541 The land cover data used for PEEN-WE also had

542 three different sources: CORINE Land cover; the

543 Swiss land cover map; and the Norwegian land cover

544 map, each with a different legend. This required the

545 development of a series of interpretation steps that

546 would lead to harmonised habitat maps. Other dat-

547 abases, such as soil maps, river basin maps, geomor-

548 phological databases and geological maps could not be

549 used at the European level due to differences in

550 approach betweenEuropean countries; however, it was

551 possible to use certain specific information, such as for

552 calcareous soils.

553It is generally assumed that to provide for the

554adequate protection of biodiversity, a specified area

555of land in any country should be designated; thus, in

556the convention on biological diversity (CBD) 10% is

557mentioned as a minimum area of the land that should

558be in protected areas per country. This figure is,

559however, a political construct and its effectiveness for

560biodiversity conservation has not been proven. As

561figures tend to live a life of their own in a policy

562environment, it is now set as a target in the plans for

563ecological networks of the German Bundeslander

564(Von Haaren and Reich 2006).

565Land use change, infrastructure planning and

566climate change are important challenges for the

567development of effective ecological networks world-

568wide. International (cross-border) cooperation and

569renewal of national and regional strategies for

570biodiversity, as well as evidence of their effective-

571ness, is required. This presents a challenge for policy

572makers, land managers and planners, as well as for

573the research community that supports them. This is

574clearly the key challenge for the implementation of

575ecological networks as, at present, within the Euro-

576pean Union, Norway and Switzerland around 120

577regional and national governments and agencies share

578the responsibility for biodiversity conservation. Real

579ecological networks are being developed at the

580country or regional level, but at the European and

581global level there are mainly visions. In general the

582size of network components serves as a criterion of

583the network hierarchy (Mander et al. 2003). Thus,

584mega-scale ecological networks can be considered at

585global level such as the Atlantic flyways (Boere et al.

5862006); and the macro-scale of ecological networks is

587represented by macro-regional-level plans such as

588PEEN (Bouwma et al. 2002; Jongman et al. 2006),

589the Wildlands project (Noss 1992) or national level

590projects within larger countries such as Russia

591(Sobolev et al. 1995). Most ecological networks have

592been planned at the mesoscale, and these include

593examples such as the Cheshire ECONET (http://

594www.cheshire.gov.uk/SREP/NHE_Econet_index.htm)

595and the Dutch Ecological Network (Jongman et al.

5962006b). The most detailed analyses and implemen-

597tation schemes have been undertaken at a micro-

598scale, for example increasing the accessibility of

599streams for Salmonid fish (see http://www.tweed

600foundation.org.uk) or adapting forest management to

601prevent fragmentation (Harris 1984).

Fig. 6 Main ecological corridors in Poland for mammals such

as wolfs (Jedrzejewski et al. 2005)

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602 PEEN does fulfil a role in developing a vision for

603 coherence in natural habitats for Europe, The Con-

604 vention on Migratory Species (CMS) aims to con-

605 serve migratory species throughout their range. One

606 of the agreements under the aegis of the CMS focuses

607 on ‘African–Eurasian Migrant Waterbirds’. The fly-

608 ways of birds as a whole must be considered as

609 ecological networks linking Siberian, European and

610 African wetlands. Flyways are defined as: the entire

611 range of a migratory bird species (or groups of related

612 species or distinct populations of a single species)

613 through which it moves on an annual basis from the

614 breeding grounds to non-breeding areas, including

615 intermediate resting and feeding places as well as the

616 area within which the birds migrate (Boere et al.

617 2006). This links Siberian and Greenland habitats to

618 those in Western Europe and Africa and, in this way,

619 emphasizes the importance of coherency in European

620 nature. PEEN gives that coherency a territorial basis.

621 At the pan-European level, the convention on the

622 conservation of European wildlife and natural habi-

623 tats (Bern convention, adopted in 1979) binds con-

624 tracting parties to the protection of habitats and

625 species of European concern and promotes cooper-

626 ation between countries for the protection of migra-

627 tory species. The recommendation, adopted within

628 the convention, on the conservation of natural areas

629 outside protected areas specifically addresses these

630 issues (Recommendation 25, adopted in 1991). It

631 encourages the conservation and, where necessary,

632 the restoration of ecological corridors, habitat types

633 and landscape features that are important for wildlife

634 conservation.

635 Connectivity can have several forms. It can be

636 flyways as for migratory birds including stepping

637 stones, but also terrestrial or river corridors. The most

638 obvious terrestrial corridors are forest corridors, but

639 they can also exist as wetland/river related corridors

640 or mountain corridors.

641 The Species and Habitats Directive and the Birds

642 Directive form the main legal framework for pro-

643 tecting nature and biodiversity in the EU; together

644 they implement some of the international require-

645 ments outlined above, including the Bern convention

646 (Commission of the European Communities 2007).

647 Article 3 of the Habitats Directive states that ‘‘where

648 they consider it necessary, member states shall

649 endeavour to improve the ecological coherence of

650 Natura 2000 by maintaining, and where appropriate

651developing, features of the landscape which are of

652major importance for wild fauna and flora, as referred

653to in Article 10’’.

654Connectivity is considered a task of national or

655regional competent authorities. It is recognised that

656the implementation of connectivity measures may be

657constrained by the lack of detailed knowledge of the

658ecological requirements of species and habitats.

659Article 18 of the Habitats Directive therefore calls

660for research and exchange of information. It specif-

661ically states that: ‘‘Particular attention shall be paid to

662scientific work necessary for the implementation of

663Articles 3 and 10, and transboundary co-operative

664research between member states shall be encour-

665aged’’. Conservation actions under other EU legisla-

666tion may also help to deliver connectivity measures

667required under the Birds and Habitats Directives. In

668particular, the Water Framework Directive (WFD)

669includes measures, such as the development of river

670basin management plans that will help to maintain and

671restore connectivity in the wider environment (Com-

672mission of the European Communities 2007). How-

673ever, this also means that no authority has the

674responsibility for continental European corridors.

675Their development depends solely on the willingness

676of countries and regions to cooperate. This is not an

677effective approach as countries have an inside focus

678and there is no incentive for European cooperation.

679National and regional ecological networks are always

680defined and constructed within a complex interaction

681of specific political, social, economic, and natural

682conditions resulting in specific cultural traditions.

683Thus, perceptions of nature and interpretations of the

684same words differ across borders. The key is therefore

685to focus on interactions that might occur between

686regions, levels, hierarchies, organisations, NGOs, and

687departments. A European initiative is needed for the

688implementation ecological connectivity at the Euro-

689pean level as national initiatives will only cross one

690border and do not have a European view.

691A further complication of present conservation

692policy is that, although the EU Habitats Directive

693recognises that priority habitats and species need to

694be conserved, it does not provide any accommodation

695for the changes that can take place to ecosystems over

696time, through succession and inherent ecosystem

697dynamics (Evans 2006), and does not give guidelines

698for the design of ecological corridors. Some research

699has been done on how species respond to the

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700 cumulative effects of land use and climate change

701 (Devictor et al. 2008). Site-based conservation strat-

702 egies, whether local or linked to the Natura 2000

703 network, may not be written to provide a response to

704 such challenges. A shifting climate could result in

705 species extinctions within designated areas or move-

706 ment of species populations towards sites not opti-

707 mally designed for the delivery of conservation

708 objectives or within other countries. Well designed

709 ecological corridors are therefore also required for

710 the ecological network of Europe and the first studies

711 have been carried out (Vos et al. 2008).

712 Ecological corridors can be designed or make use

713 of existing landscape structures. However, proof is

714 needed that these structures might function as such.

715 For many centuries transhumance has existed in

716 Europe as part of the agricultural system (Bunce et al.

717 2004). For Spain this has resulted in a network of

718 north–south drove roads (Canadas), hundreds of

719 kilometres in length, which have been functioning

720 as corridors for species and propagules (Manzano

721 et al. 2005). In the framework of the PEEN-WE

722 project a study has been carried out on the role of the

723 Canadas as north–south corridors, but so far no

724 conclusions could be drawn in relation to their actual

725 function as their condition is unknown (Bunce et al.

726 2006). Part are still functioning, part have changed

727 their function. They have therefore not been included.

728 Corridors are being developed at the national level

729 and terrestrial ecological corridors at the European

730 level only exist at the conceptual level of PEEN.

731 Ecological corridors are being developed for national

732 purposes (Fig. 7). Initiatives however do exist to

733 coordinate within countries (between regions) and

734 between countries; as is evident from the outcomes of

735 the Vilm meeting (Ssymank et al. 2005) and the

736 Bialowieza meeting (Jedrzejewski and Jedrzejewski

737 2008) which convened in order to explore such

738 issues.

739 The different levels of implementation of ecolog-

740 ical networks imply that data requirements are

741 different and that different parties are involved as

742 stakeholders. For ecological networks to be imple-

743 mented the inclusion of the right stakeholders at the

744 right level is therefore essential. For the international,

745 European level agreements equivalent to those for

746 flyways do not yet exist, but they are needed. Success

747 in biodiversity conservation and sustainable develop-

748 ment requires not only that a diverse range of

749planning initiatives should be developed for the

750design and management, in this case specifically of

751ecological networks, but also that a range of stake-

752holders and land use and management interests

753should be involved.

754Conclusion

755The PEEN project was successful in reaching its goal

756to promote the idea of a pan-European vision of

757biodiversity conservation through a European eco-

758logical network. However, there are three important

759challenges for the further development of a coherent

760ecological network. Knowledge transfer is needed as

761well as new knowledge especially in relation to the

762impact of changing environmental and land use

763conditions on species and habitats in the wider

764countryside. Climate and global change will affect

765the patterns of many ecological and other relation-

766ships in the landscape, potentially leading to a level

767of complexity that may prove intractable and difficult

768to resolve. Research on changing population patterns

769in relation to landscape permeability should be

770directed towards the provision of essential knowledge

771needed for the limitation and prevention of irrevers-

772ible damage, adaptation and mitigation measures. In

773the present situation, where countries are pursuing

774national initiatives at the expense of international and

775cross boundary cooperation, the major problem is that

776European ecological corridors are not being devel-

777oped; a situation that is only exacerbated because

778there is presently no responsible institution or coor-

779dination mechanism in place. That means that

780coherence between countries and regions is hard to

781realise in practice.

782Developing connectivity is one of the recommen-

783dations of the CBD Conference of Parties in Nagoya

784(Japan) in October 2010. Ecological networks need to

785be developed at the field and regional scales, and at

786the national and trans-national scales. Monitoring

787habitat diversity, and not just connectivity, is a key

788element in any biodiversity strategy for agricultural

789landscapes. Quantifying the economic benefits of

790ecological networks and making them explicit

791through interdisciplinary research and integrated

792long-term research on the social, economic and

793ecological mechanisms that maintain biodiversity

794and its ecological services is a clear necessity.

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795 Acknowledgments The preparation of an indicative PEEN796 maps started in 1999, was finalised in 2006 and reported to the797 European Ministers of Environment in 2007. We acknowledge798 all those who made the work possible by funding it, the799 Ministry of Agriculture Nature and Food Quality of The800 Netherlands through its Policy Supporting Research801 Instrument, the Dutch BBI-MATRA fund, the Council of802 Europe and the Swiss Government. We also acknowledge the803 Walloon Government for their support as well as the national804 and international data providers that made the technical805 development of the maps possible, the Committee of Experts806 for the development of the Pan-European Ecological Network807 that supported the process from its inception, and the hundreds808 of policy advisors who have been consulted. We are also809 grateful to the stakeholders who made their knowledge and810 data available for this project. Without their willingness to811 support and contribute the project would have been impossible812 in the context of the complex situation in Europe that has813 prevailed throughout this decade.

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