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ARTICLE IN PRESSG ModelNC-25367; No. of Pages 8

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Contents lists available at ScienceDirect

Journal for Nature Conservation

j o ur nal homepage: www.elsev ier .de / jnc

oor effectiveness of Natura 2000 beech forests in protectingorest-dwelling bats

obias Zehetmaira,∗, Jörg Müllerb,c, Volker Runkeld, Peter Stahlschmidte,usanne Winter f, Alexey Zharovg, Axel Gruppea

Institute of Animal Ecology, Department of Ecology and Ecosystem Management, Technische Universität München, Hans-Carl-von-Carlowitz-Platz 2,reising D-85354, GermanyBavarian Forest National Park, Freyunger Str. 2, Grafenau D-94481, GermanyChair for Terrestrial Ecology, Department of Ecology and Ecosystem Management, Technische Universität München, Hans-Carl-von-Carlowitz-Platz 2,reising D-85354, GermanyMondstrasse 213, Münster D-48155, GermanyInstitute for Environmental Sciences, University of Koblenz-Landau, Fortstr. 7, Landau D-76829, GermanyChair of Land Improvement and Nature Conservation, Technische Universität Dresden, Pienner Straße 7, Tharandt D-01737, GermanyDepartment of Ecology and Ecosystem Management, Technische Universität München, Hans-Carl-von-Carlowitz-Platz 2, Freising D-85354, Germany

r t i c l e i n f o

rticle history:eceived 7 April 2014eceived in revised form 2 June 2014ccepted 25 July 2014

eywords:at activityat diversityatcorderagus sylvaticaabitat qualityoosts

a b s t r a c t

With the establishment of the Natura 2000 (N2000) network, the European Union intends to developstrategies to conserve Europe’s threatened habitats and species, including bats. Forest-dwelling bats arehighly reliant on forest structures, such as snags and hollow trees, which the bats need as roosts. Thedecrease in such forest microhabitats significantly affects the habitat use and, therefore, the activity inforests. To determine whether N2000 beech forests under active timber production offer better habitatsfor bats compared to commercially used non-N2000 forests, we measured the bat activity and assessedthe potential roosts in trees and snags in eleven pairs of stands. All survey stands represented mesotrophicbeech forests (Fagus sylvatica L.) of the N2000 habitat type 9130 (Asperulo-Fagetum) in three EuropeanBiogeographic Regions. The activity of all bat species, the activity of priority N2000 species, the speciesnumber, the number of trees with roosts and the snag volume did not differ significantly between theN2000 and non-N2000 stands. We conclude that the current management of the N2000 beech forests is

almost identical to that of non-N2000 commercial forests, and thus, the N2000 status has not led to anincrease of bat-relevant habitat variables yet. Consequently, additional efforts beyond the administrativeassignment of N2000 areas are required to build and ensure an ecologically effective and sustainablenetwork of beech forests in Europe, including increasing important forest requirements for bats, such asroosts and snags.

© 2014 Elsevier GmbH. All rights reserved.

ntroduction

Biodiversity is largely threatened by anthropogenic activities, fact that is well-recognised by international policy. At least 20egional or global conservation agreements have been establishedn recent decades (Boere & Rubec 2002), with the intention of

easurably improving nature (Male & Bean 2005). In addition

Please cite this article in press as: Zehetmair, T., et al. Poor effectivenesJournal for Nature Conservation (2014), http://dx.doi.org/10.1016/j.jnc

o the formulation and implementation of political interventions,t is essential to evaluate their effects on biodiversity (Donaldt al. 2007), but the efforts towards monitoring these effects are

∗ Corresponding author. Tel.: +49 0 8161 714588; fax: +49 0 8161 714598.E-mail address: zehetmair@wzw.tum.de (T. Zehetmair).

ttp://dx.doi.org/10.1016/j.jnc.2014.07.003617-1381/© 2014 Elsevier GmbH. All rights reserved.

mostly not considered in political discussions and often lag greatlybehind other policy fields (Ferraro & Pattanayak 2006). At the Euro-pean scale, the coherent Natura 2000 (N2000) network is a highlypromising approach to safeguard Europe’s biodiversity (Maioranoet al. 2007). The legislation is based on the Birds Directive (EuropeanCouncil 1979) and the Habitats Directive (European Council 1992).Regarding habitats (Annex I) and species (Annex II), 27 MemberStates of the European Union (EU) established Special Areas ofConservation (SAC) where specifically designed management plansshould be developed (Article 6(1), European Commission 2007).

s of Natura 2000 beech forests in protecting forest-dwelling bats..2014.07.003

In addition to protected areas where utilisation is normally pro-hibited or restricted, such as National Parks or nature reserves,the major part of the network consists of still commercially usedsites. This centrepiece of the EU nature conservation strategy covers

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ARTICLENC-25367; No. of Pages 8

T. Zehetmair et al. / Journal for N

ore than 20% of the terrestrial and marine EU territory (Europeanommission 2012a), currently with 26,406 sites (Tsiafouli et al.013), underlining the idea of thinking and acting on an interna-ional scale, according to the demand for effective nature protectionOpermanis et al. 2012).

The overall objective of N2000 is to maintain and restore aFavourable Conservation Status” (FCS) (Articles 1e, 1i) for all habi-ats and species that are listed in Annexes I and II (European Council992). The maintenance of FCS mostly targets the previously pro-ected areas that already provide good habitat quality. In contrast,estoring FCS is a challenge for the commercially managed siteshere habitat quality is often lower and therefore needs to be

mproved. FCS means that the habitats and the species populationsre in good condition with regards to both the quality and the extentnd have a positive future perspective (Evans & Arvela, 2011). Theonservation status must be evaluated and reported every six yearsy all of the Member States (Article 17, Habitats Directive). In therst report, covering 2001–2006, only 17% of both the habitatsnd species were classified as having a “favourable” conservationtatus, whereas for the main proportion, an “unfavourable/bad”r even “unknown” conservation status was reported (Europeanommission 2009).

Forests are an indispensable key for biodiversity. They coverore than 50% of the terrestrial N2000 surface area and repre-

ent the most important habitat type for many endangered speciesEuropean Communities 2003). Many of these species are depend-ng directly or indirectly on particular tree species, such as theuropean beech (Fagus sylvatica L.), which is the main deciduousree species in Europe (Bolte et al. 2007; Packham et al. 2012). Theimitation of beech distribution to Europe bestows great responsi-ility on the EU for the protection and conservation of the species

tself and all habitat types characterised by beech. Forest struc-ural composition plays a decisive role in biodiversity (Cale et al.013; Ishii et al. 2004; Remm & Lohmus 2011; Tews et al. 2004)nd greatly affects the habitat quality for forest-dwelling animalsuch as birds (Pinotti et al. 2012), arthropods (Gossner et al. 2013;usso et al. 2011) and bats (Lacki et al. 2007). With the N2000etwork, the EU and their Member States have vowed to desig-ate SACs for the 13 bat species that are listed in Annex II. Batstrongly depend on the quality of their habitats, especially on thevailability of foraging and breeding structures (Perry et al. 2007;uczynski et al. 2010). Therefore, forest-dwelling bats can be useds good indicators of forest habitat quality (Jones et al. 2009), butlso to examine the habitat – suitability of forest stands under dif-erent management conditions and intensities (Russo et al. 2010).hus they are suitable for evaluating the implementations of con-ervation measures in N2000 forests. In general, the populationsf European bat species have declined because of habitat degrada-ion, fragmentation and land-use changes (Frey-Ehrenbold et al.013; Vié et al. 2008), but little is currently known about batsnd their activity within forests under active timber productionMiller et al. 2003; Russo et al. 2010), even though the large major-ty (97%) of European forests are commercially used (FAO 2011).hus, managed forests play a key role in supporting bats in Europe,ncluding most N2000 forests where active timber production istill common practice (Tsiafouli et al. 2013). With this study, weimed to estimate how bats are affected by N2000 in commer-ially used beech forests at the moment. Therefore, we comparedhe bat activity and the bat diversity in pairs of beech stands usedor timber production inside and outside the N2000 network. Wexpected higher values inside stands protected by N2000 causedy a better management here. Because bats are very dependent

Please cite this article in press as: Zehetmair, T., et al. Poor effectivenesJournal for Nature Conservation (2014), http://dx.doi.org/10.1016/j.jnc

n foraging and breeding structures (Perry et al. 2007; Ruczynskit al. 2010) the positive influence of N2000 on bats would thus beeflected by more diverse bat-relevant forest microhabitats, whichere estimated in all stands, too. Thus, it would be possible to

PRESSonservation xxx (2014) xxx–xxx

evaluate the current state of the implementation of the N2000network and the N2000 management plans. Studying Asperulo-Fagetum beech stands, we focused on one of the major temperateforest types in Europe that covers more than 13,000 km2 (EuropeanTopic Centre on Biological Diversity, 2009) and therefore repre-sents one of the most important habitats for forest-dwelling batsin Europe.

Methods

Study areas

The survey was conducted in eleven pairs of utilised foreststands in Europe (Fig. 1) representing Asperulo-Fagetum beechforests (N2000 habitat type code 9130) in three BiogeographicRegions (Atlantic: 2 pairs, Continental: 7 pairs, Mediterranean: 2pairs; Fig. 1). Each pair consisted of one stand located within aN2000 site and of one stand within a reference site outside N2000(Table 1). All stands were selected using the following criteria: (i)dominated by European beech (>70%), (ii) managed by a high forestmanagement strategy, (iii) the trees are 80–120 years old and (iv)each stand has a size of at least 10 hectare (ha). Within each stand,eight study plots without conifers in the overstorey (0.1 ha each,ntotal = 176) were randomly selected within the created grids (edgelength = 50 m × 50 m).

Bat survey

The bat surveys were performed in 2011 (seven pairs of stands)and 2012 (four pairs of stands). Bat calls were recorded simulta-neously for four of our eight survey plots each stand on one night inend of May and for the other four plots in the beginning of July from6.00 pm to 6.00 am the next day; this period of time was chosento minimise the confounding factors of seasonal migration (Fordet al. 2005); we standardised the night surveys with regard to thetemperature and wind speed; nights with very high relative humid-ity and rainfall were avoided. Thus, we generated eight recordingdata sets, respectively eight survey nights in each of the 22 stands.In sum, we performed 176 all-night surveys (22 stands × 8 plots)with 96 h recorded in each stand (8 plots × 12 h). Each night, weused eight Batcorder 2.0 instruments (ecoObs GmbH 2010) simul-taneously on the plots, where they were installed in the centre ona metal pole approximately 2.5 m above the forest ground (see Fig.S1). The omnidirectional microphones were orientated upwards ata 10◦ angle and were not directed into dense vegetation to min-imise the effect of the forest structures on the ultrasonic soundtransmission in the understorey (Marten & Marler 1977; Patriquinet al. 2003). After recalibration before each survey season, we usedthe same batcorder-settings (quality: 20, threshold: −27 dB, post-trigger: 60 ms, critical frequency: 16 kHz) and “Auto + Timer” mode(18.00–06.00) for each survey.

Bat-relevant habitat variables

We assessed all trees with roosting resources (cavities, bark-pockets) according to the microhabitat types M12–M15 and M18in Winter and Möller (2008) in each study plot. The identificationand evaluation were performed by visual inspection. The volumeof the standing deadwood [m3] was calculated from the height [m]and the diameter [cm] at breast height (1.3 m; dbh), including allsnags with at least 7 cm (dbh) with bark and 6 cm (dbh) withoutbark (for details, see also Winter & Möller 2008). For the statistical

s of Natura 2000 beech forests in protecting forest-dwelling bats..2014.07.003

calculations, all data on plot level (1000 m2) were standardised tovalues per ha, from which the mean + sd per ha of each stand wascalculated (for details see Table S2). To estimate the characteristicsof the habitat variables, we furthermore compared [a] trees with

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T. Zehetmair et al. / Journal for Nature Conservation xxx (2014) xxx–xxx 3

Fig. 1. Location of the 11 pairs of forest stands used in the study. Forest sites of the Continental region: [1] WUE (D-Würzburg), [2] EB (D-Ebrach), [3] FH (D-Forchheim),[ eusiedL mpshe RGEN

rtaswd

TD−

4] KH (D-Kelheim), [5] LL (D-Landsberg/Lech), [6] SZ (AT-Salzach), [7] NSS (AT-N (FR-Montagne de Lure); forest sites of the Atlantic region: [10] EHH (UK-East Haxplained in Table 1. Hachures indicate occurrence of beech stands in Europe (EUFO

oosts, [b] trees with cavities and [c] trees with bark pockets withhe published data from Winter and Möller (2008) referring to themount of these structures within managed and protected beech

Please cite this article in press as: Zehetmair, T., et al. Poor effectivenesJournal for Nature Conservation (2014), http://dx.doi.org/10.1016/j.jnc

tands. Regarding the [d] snag volume we compared our valuesith the data from Christensen et al. (2005) from beech stands withifferent protection histories.

able 1etails of beech stands included in the study. Coordinates = centre of stand. N2000 site-cod

= non-Natura 2000 site. BR = Biogeographic Region (C: Continental, M: Mediterranean a

Country Site-number Site-code Coordinates

Germany 1 WUE a N49.69316 E9.86465

1 WUE b N49.68693 E9.84414

2 EB a N49.81963 E10.51930

2 EB b N49.79550 E10.61388

3 FH a N49.89099 E11.06900

3 FH b N49.88195 E11.07602

4 KH a N48.91093 E11.83536

4 KH b N48.93317 E11.75266

5 LL a N47.93401 E11.08484

5 LL b N48.05427 E11.00376

Austria 6 SZ a N48.10881 E12.75224

6 SZ b N48.10637 E12.75685

7 NSS a N47.98129 E16.73221

7 NSS b N47.98010 E16.68826

France 8 S a N44.64747 E5.17914

8 S b N44.64546 E5.21374

9 L a N44.09582 E5.78427

9 L b N44.09406 E5.79385

Great Britain 10 EHH a N51.04975 W0.96389

10 EHH b N50.99633 W1.05619

11 WVW a N51.82563 W2.65891

11 WVW b N51.72506 W2.69089

ler See); forest sites of the Mediterranean Region: [8] S (FR-Massiv de Saou), [9]ire Hangers), [11] WVW (UK-Wye Valley Woodlands). The numbers refer to sites

2013).

Data analysis

The software batIdent 1.03 (www.ecoobs.de) was used to auto-

s of Natura 2000 beech forests in protecting forest-dwelling bats..2014.07.003

matically determine the bat species by their specific calls and toexclude the non-bat recordings. For the manual identification ofnonspecific calls, we used bcAnalyze 2.0 (www.ecoobs.de) and

e = each Natura 2000 site is recognised by a unique code which forms the key-item.nd A: Atlantic).

Size of (ha) Altitude Year N2000 site-code BR

17.0 334 m a.s.l. 2012 DE6225372 C15.0 359 m a.s.l. 2012 – C12.0 412 m a.s.l. 2012 DE6029371 C10.0 418 m a.s.l. 2012 – C12.0 570 m a.s.l. 2011 DE6032371 C15.0 540 m a.s.l. 2011 – C15.0 450 m a.s.l. 2011 DE7036372 C15.0 470 m a.s.l. 2011 – C20.0 630 m a.s.l. 2011 DE8032372 C10.0 630 m a.s.l. 2011 – C

10.3 412 m a.s.l. 2012 AT3110000 C10.9 437 m a.s.l. 2012 – C10.3 263 m a.s.l. 2012 AT1124823 C11.2 257 m a.s.l. 2012 – C

14.5 1150 m a.s.l. 2011 FR8201686 M10.3 950 m a.s.l. 2011 – M11.5 1340 m a.s.l. 2011 FR9301537 M10.5 1230 m a.s.l. 2011 – M

10.2 210 m a.s.l. 2011 UK0012723 A11.0 180 m a.s.l. 2011 – A10.3 130 m a.s.l. 2011 UK0012727 A10.1 200 m a.s.l. 2011 – A

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4 T. Zehetmair et al. / Journal for Nature Conservation xxx (2014) xxx–xxx

Table 2Recorded calls of species/species groups and their calculated activity as min-counts (mc) in N2000 and non-N2000 stands. The species listed in Annex II of the Directive areshown in bold. p-level = significance as obtained from the Wilcoxon signed-rank test for differences in min-counts on species level between N2000 stands and non-N2000stands.

Species Calls N2000 (mc) Non-N2000 (mc) Total (mc) p-Level

Barbastella barbastellus (Schreber, 1774) 389 50 19 69 nsEptesicus nilssonii (Keyserling & Blasius, 1839) 2 – 1 1 nsEptesicus serotinus (Schreber, 1774) 205 14 2 16 nsHypsugo savii (Bonaparte, 1837) 19 2 – 2 nsMiniopterus schreibersii (Kuhl, 1817) 83 2 4 6 nsMyotis bechsteinii (Kuhl, 1817) 327 7 2 9 nsMyotis brandtii/mystacinus (Eversmann, 1845/Kuhl, 1817) 9075 124 245 369 nsMyotis dasycneme (Boie, 1825) 38 – 3 3 nsMyotis daubentonii (Kuhl, 1817) 1811 51 35 86 nsMyotis emarginatus (É. Geoffroy, 1806) 25 – 2 2 nsMyotis myotis (Borkhausen, 1797) 563 28 28 56 nsMyotis nattereri (Kuhl, 1817) 798 41 18 59 nsNyctalus leisleri (Kuhl, 1817) 105 5 1 6 nsNyctalus noctula (Schreber, 1774) 1212 141 18 159 *

Pipistrellus kuhlii (Kuhl, 1817) 46 2 – 2 nsPipistrellus nathusii (Keyserling & Blasius, 1839) 8209 141 356 497 nsPipistrellus pipistrellus (Schreber, 1774) 30,443 341 1129 1470 nsPipistrellus pygmaeus (Leach, 1825) 6207 268 129 397 nsPlecotus auritus/austriacus (Linnaeus, 1758/Fischer, 1829) 149 5 3 8 nsRhinolophus hipposideros (Bechstein, 1800) 2 1 – 1 nsSum 59,708 1223 1995 3218

*

ciuriccteDtIADa(

pvupnafiAwtawiTIRtaarlR

p < 0.05.

ompared the sonograms and oscillograms of the bat calls withmages from Hammer and Zahn (2009) and (Skiba 2009). The doc-mentation was organised with bcAdmin 2.0 (www.ecoobs.de). Batecords were excluded from the analysis in cases of doubtful speciesdentification. The activity of each bat species was calculated byounting the number of recorded 1-min intervals (min-counts) ofalls per night (for details, see Müller et al. 2012). Additionally,he bats being found and their activity were separated to the cat-gories of all species and those which are listed in Annex II of theirective. Thus, we could use the following bat-variables for fur-

her analyses: number of [a] all bat species; number of [b] AnnexI bat species; [c] total bat activity (all species included); and [d]nnex II bat activity (only species listed in Annex II of the Directive).ifferences in min-counts on species level between N2000 standsnd non-N2000 stands were tested by Wilcoxon signed-rank testTable 2).

The statistical analyses were performed with R-3.0.2 (www.r-roject.org) by using the packages lme4 (Bates et al. 2013) andegan (Oksanen et al. 2013). First, Mann–Whitney U tests weresed to check the structural variables (roosts, cavities, bark-ockets and snag volume) for differences between the N2000 andon-N2000 stands. Then, to evaluate how bats in beech forestsre affected by N2000, we used linear mixed effect models (LMM)t by REML (lmer). All values of variables – [a] all bat species; [b]nnex II bat species; [c] total bat activity; [d] Annex II bat activityere log10(x + 1) transformed to obtain normal distributions. In

he models, the N2000 status (yes/no) was set as a fixed factor,nd the stand, the Biogeographic Region, the year and the seasonere set as random factors with an observation-specific random

ntercept to account for possible overdispersion (Ye et al. 2013).o assess the overall effect of N2000 we used the following model: lmer (log10(x + 1) ∼ N2000 status + (1|stand) + (1|Biogeographicegion) + (1|year) + (1|season) + (1|Obs), data = dataset1). In addi-ion, we included the interaction between the N2000 statusnd the corresponding site to assess the effects of N2000

Please cite this article in press as: Zehetmair, T., et al. Poor effectivenesJournal for Nature Conservation (2014), http://dx.doi.org/10.1016/j.jnc

t the regional scale (site level). The second model withegard to the site level is similar to model I above (model II):mer (log10(x + 1) ∼ N2000 status:site + (1|stand) + (1|Biogeographicegion) + (1|year) + (1|season), data = dataset2).

Results

Bat survey

We recorded a total of 60,563 bat call sequences in 176sampling-nights (2112 h of recording). Of these, 98.6% (59,708)were assigned to 18 species and two species-groups with acous-tically inseparable species (Myotis brandtii/M. mystacinus andPlecotus auritus/P. austriacus). The most frequent species was Pip-istrellus pipistrellus with 30,443 individual calls (51.0%), followedby Myotis brandtii/mystacinus with 9075 calls (15.2%) and Pipistrel-lus nathusii with 8209 calls (13.7%). By comparing the bat activity(min-counts) between N2000 beech stands and non-N2000 standson species level we found significant higher values for Nyctalusnoctula, which was detected more often inside N2000 stands thanin the reference stands outside the network. The summary of allrecorded calls per bat species as well as the calculated min-countsare shown in Table 2.

Impact of Natura 2000

We applied two linear mixed effect models (lmer) to analysethe effects of the protection status N2000 on several bat variables.In general (model I), we found that the bat diversity representedby the number of bat species was not significantly affected for allbats (LMM: z = 1.876, p = 0.061) nor was the number of Annex IIbat species (LMM: z = 0.828, p = 0.408). When considering bat activ-ity in the survey stands by calculating minute-counts, we couldnot detect any significant effect of N2000 status for the generalbat activity (LMM: z = 0.763, p = 0.445) or the activity of Annex IIbats (LMM: z = 0.812, p = 0.417). The values of the bat-variables aresummarised in Table S1. For a more detailed analysis, we nexttested how bats were affected by N2000 status at the regional scale(model II). In doing so, it was possible not only to evaluate the gen-eral effect of N2000 (model I) but also to evaluate the influence at

s of Natura 2000 beech forests in protecting forest-dwelling bats..2014.07.003

the site level to consider regional aspects of the N2000 implemen-tation. Similar to the general approach, we also could not detect adecisive impact of N2000 on the bat variables at the regional scale.Only one significant effect was found for site NSS and general bat

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T. Zehetmair et al. / Journal for Nature Conservation xxx (2014) xxx–xxx 5

Fig. 2. Box–Whisker plots illustrating the amount of habitat variables in beech stands with protection status (N2000) and without protection status (non-N2000).M ng varb ).

acf

bnebmfRis

D

t2ZKrpfsIoaretaaSfitcAatei2aRtpf

ann–Whitney U-tests indicated no significant differences in values for the followiark-pockets (W = 72, p-value = 0.462) and (d) snag volume (W = 52.5, p-value = 0.622

ctivity (LMM: z = 2.185, p = 0.029). The differences at the site levelorresponding to the LMM II are illustrated by Box–Whisker plotsor all five bat variables (Fig. S2).

When evaluating the values of bat-relevant habitat variablesetween beech stands within N2000 areas and stands outside theetwork (reference sites), we did not detect any significant differ-nces here, too (Fig. 2). Comparing our findings with data publishedy Winter and Möller (2008) they were in the same order ofagnitude as within beech stands under active management but

ar below the values of long-term unmanaged stands (Fig. 3a–c).egarding snag volume (Fig. 3d), we found significantly less stand-

ng deadwood in all our survey stands than in protected beechtands calculated by Christensen et al. (2005).

iscussion

Several studies have already analysed bat activity in relationo timber-harvesting strategies (Humes et al. 1999; Owen et al.004), managed and unmanaged stands (Ruczynski et al. 2010;ielinski & Gellman 1999) and different habitat types (Celuch &ropil 2008; Wermundsen & Siivonen 2008). Humes et al. (1999)eported significantly higher bat activity in old-growth stands com-ared with unthinned stands, and significant differences were alsoound between forest types, such as conifer, deciduous and mixedtands (Patriquin & Barclay 2003; Wermundsen & Siivonen 2008).n contrast to these approaches, we focused on the current situationf bats and bat-relevant habitat variables in beech stands underctive management (high forest) inside and outside N2000 withegard to one specific forest type (Asperulo-Fagetum) to evaluate theffect of the N2000. The results of our general approach indicatedhat there are currently no significant differences between N2000nd non-N2000 stands for the four variables describing bat activitynd diversity, nor for the bat relevant habitat variables (Fig. 2, Table2). We could not find an increased use of N2000 stands by batsor neither all of the species nor for those species that are listedn Annex II as target species of the network. Taking into accounthe pairwise comparison of stands, we only detected one signifi-ant effect of N2000, that being on the general bat activity in theustrian stand NSS a. For all other pairs of stands, both positivend negative influences were found (Fig. S2). These influences athe site-level become less important when looking at the overallffect of the N2000 status. The stand structures play a key rolen bat occurrence and their behaviour in forests (Kunz & Fenton003). Roosts, especially, such as tree cavities and bark pockets,re essential for resting and reproduction (Ruczynski et al. 2010;

Please cite this article in press as: Zehetmair, T., et al. Poor effectivenesJournal for Nature Conservation (2014), http://dx.doi.org/10.1016/j.jnc

usso et al. 2004; Webala et al. 2010) as well as for reducinghe risk of predation (Ruczynski & Bogdanowicz 2008) and thuslay a key role in bat conservation concepts. However, even inorests managed under conservation guidelines, such as those in

iables: (a) roosts (W = 63.5, p-value = 0.869), (b) cavities (W = 72, p-value = 0.465), (c)

the N2000 network, the deficit of roosts could be detrimental forbats (Hutson et al. 2001). In our study sites, the values of habitatvariables did not differ between beech stands inside and outsidethe N2000 network (Fig. 2, Table S2), which implies that for theseAsperulo-Fagetum stands, the conservation status did not lead toimprovements. Compared with the published data of Winter andMöller (2008) referring to the amount of structures within man-aged and protected beech stands, the values in our survey standswere of the same order of magnitude as within stands under activemanagement but far below the values of long-term unmanagedstands (Fig. 3a–c). Similarly, significantly less standing deadwoodwas found in our survey stands compared with protected stands asreported in Christensen et al. (2005) where snag volume increasedsignificantly by time without forest utilisation (Fig. 3d). This illus-trates that the period of time under natural development is the mostimportant criterion for the development of higher snag volume inforest stands (Fig. 3d). The fact that N2000 has been implementedfor almost 20 years might explain the low values we found, as itmight take several decades of active conservation strategies imple-mented in forest management to improve habitat quality in anecological sense. This inevitably leads to the question of how thenetwork supports the protection of biodiversity in European beechforests now and how it will do so in future. Our findings of the non-existent effects of N2000 on bat activity and bat diversity may beinterpreted as a result of the characteristics of bat-relevant micro-habitats, where we also found no differences inside and outside thenetwork. This indicates that forest management strategies in N2000sites do not differ significantly compared to commercial forest uti-lisation and thus did not improve habitat quality, so far. Therefore,the current N2000 beech forests do not represent better habitatsfor bats than the managed reference stands outside the networkat the moment. Instead, they only fulfil their role in maintainingthe initial status but do not to play a pioneering role for conserva-tion issues in Europe. Real improvements by N2000 in biodiversitywill most likely be visible after some more decades depending onthe development of specific conservation strategies which shouldbe integrated and implemented in management plans. Here, espe-cially the protection and the enrichment of forest microhabitatslike cavities or snags as key components for biodiversity must beincluded to improve habitat quality. Previous studies on bats (Lisónet al. 2013), vertebrates in general (Maiorano et al. 2007), saprox-ylic beetles (Bosso et al. 2013; D’Amen et al. 2013), bird species(Albuquerque et al. 2013; Pellissier et al. 2013), lichen species(Rubio-Salcedo et al. 2013) and plant diversity (Dimitrakopouloset al. 2004) have already indicated the unfortunately currently low

s of Natura 2000 beech forests in protecting forest-dwelling bats..2014.07.003

effectiveness of the N2000 network. Furthermore, Mazaris et al.(2013) explicitly addressed the need to establish species-specificmanagement plans with flexible conservation strategies to ensurefuture protection under the changing conditions of land use and

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6 T. Zehetmair et al. / Journal for Nature Conservation xxx (2014) xxx–xxx

Fig. 3. Data values of the stand structures (mean + sd) from the study plots compared with the data from the literature: (a) roosts, (b) cavities, (c) bark-pockets, (d) snagv fferenw (data

coHu2t

laptmatdHeTufHtStiaapeiwcFlTibagob

olume. In a–c, our data were compared with the data from beech stands under dias compared with the data from beech stands with a different protection history

limate change, and Louette et al. (2011) indicated the importancef regional conservation strategies under the N2000 framework.owever, to evaluate the effects of strategies, it is also necessary tose simple but effective monitoring programmes (Chiarucci et al.008; Simonson et al. 2013) to develop adapted management planso enhance their effectiveness in terms of habitats and species.

Currently, most forests are facing intensive economic chal-enges and rapid changes, especially regarding their importance as

source of renewable biomass (Mansikkasalo 2012). This growingressure even on the N2000 forests (Tsiafouli et al. 2013) necessi-ates a strong policy at all levels (Jones-Walters & Civic 2013) and

akes the development and implementation of realisable man-gement plans indispensable for ensuring nature conservation inhe future (Jones-Walters & Civic 2013). Thus, politics, stakehol-ers and society have to accept the costs of nature conservation.owever, regarding the commercial N2000 beech forests, moreffort is clearly required to improve habitat quality for bats here.herefore, modifications towards a forestry that is based on nat-ral conditions are needed, especially as forests are key habitatsor the roosting and feeding of European bats (Lacki et al. 2007).owever, many species are currently threatened by the applica-

ion of modern forest management practices (Hutson et al. 2001).pecifically, non-selective and intensive harvesting practices leado the loss of large-sized living trees and deadwood, which resultsn the decline of habitat quality for bats (Erickson & West 2003)nd many other species. Well-adopted and sustainable forest man-gement strategies implicated in concrete N2000 managementlans will be necessary to prevent this decline. This could lead tonhancement of the situation of all forest-dwelling bats by improv-ng their habitats. First of all, bat-relevant structures such as trees

ith existing cavities and bark pockets (habitat trees) must beonsequently excluded from felling operations (Russo et al. 2004).urthermore, large dead trees and an adequate number of matureiving trees (recruitment trees) must also be excluded from felling.hese recruitment trees will develop into habitat trees, providingmportant microhabitats for the following decades. In addition toats, many other forest-dwelling species, such as other mammals,

Please cite this article in press as: Zehetmair, T., et al. Poor effectivenesJournal for Nature Conservation (2014), http://dx.doi.org/10.1016/j.jnc

rthropods and birds, will benefit from such management strate-ies. These strategies will help enhance the success of achieving theverall long-term target of the N2000 network: preserving Europe’siodiversity.

t management conditions (data from Winter & Möller 2008). The snag volume (d)from Christensen et al. 2005).

Acknowledgements

The data collection and analyses were performed as a part ofthe joint project “Beech Forest for the Future” (BeFoFu), whichis being conducted within the BiodivERsA network of the EU 7thFramework Programme for Research. This study was funded by theGerman Federal Ministry of Education and Research (funding code01LC0814B). We give special thanks to the Bayerische StaatsforstenAöR (FB Landsberg a. Lech, FB Kelheim, FB Forchheim, FB Ebrach, FBArnstein), Österreichische Bundesforste AG Forstbetrieb Wiener-wald, Castell-Castellsche Forstverwaltung, HeeresforstverwaltungBruckneudorf, Chargé de mission Forêt de Saoû, Forestry Commis-sion Coleford, Forestry Commission Itton and W. Tyrwhitt-Drakefor the permission to collect the data from their sites. Further-more, we thank O. Chambon, E. Engeßer, A. Friedel, R. Gable, A.Jones, S. Keilholz, G. Kiefer, L. Kiennen, U. Mergner, Dr. B. Mitter-bacher, R. Patrick, G. Rühling, G. Schebeck, N. Stöger, T. Thron, Dr.A. Wieshaider and D. Winterstein for their support and help withthis survey.

Appendix A. Supplementary data

Supplementary data associated with this article can be found, inthe online version, at http://dx.doi.org/10.1016/j.jnc.2014.07.003.

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