The significance of habitats as indicators of biodiversity and their links to species

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ARTICLE IN PRESSG ModelCOIND-1273 No of Pages 7

Ecological Indicators xxx (2012) xxxndashxxx

Contents lists available at SciVerse ScienceDirect

Ecological Indicators

jo ur nal homep age wwwelsev ier com locate eco l ind

he significance of habitats as indicators of biodiversity and their links to species

GH Bunceah MMB Bogersa D Evansb L Haladac RHG Jongmanalowast CA Muchera

Bauchd G de Bluste TW Parr f L Olsvig-Whittakerg

Alterra Wageningen UR PO Box 47 6700AA Wageningen The NetherlandsEuropean Topic Centre on Biological Diversity Paris FranceInstitute of Landscape Ecology Slovak Academy of Science Branch Nitra Akademicka 2 POB 22 SK-949 10 Nitra SlovakiaUFZ ndash Helmholtz Centre for Environmental Research Department of Conservation Biology Permoserstr 15 04318 Leipzig GermanyInstitute of Nature Conservation Kliniekstraat 25 B-1070 Brussels BelgiumCentre for Ecology and Hydrology ndash Lancaster Environment Centre Library Avenue Bailrigg Lancaster LA1 4AP United KingdomIsrael Nature and Parks Authority 3 Am VersquoOlamo St Jerusalem 95463 IsraelUniversity of Life Sciences Kreutzwaldi 1 Tartu 51014 Estonia

r t i c l e i n f o

eywordsabitat definitionolicypecieshabitat associationtandardised mappingpatial models

a b s t r a c t

The first section of the paper discusses the background to the use of habitats as indicators for biodiversityincluding a discussion of the range of definitions that have been used Habitats can now be recordedconsistently across Europe at different time intervals in order to estimate stock and change as an indicatorof biodiversity that is efficient and relatively easy to record Habitats are considered to be the third levelin a hierarchy with biomes and landscapes as higher categories and vegetation species and genetic

diversity as lower levels An advantage of using habitats is that many other taxa are associated with themand examples are given from the literature Examples are also given of the association between habitatsand species in European Environmental Zones using expert judgement Statistical analysis using a rangeof procedures can also be used to assess the association between species and habitats Reliable data onthe extent status and changes in European habitats is essential for policy makers across the EuropeanUnion and would also be important for promoting species conservation

Introduction

1 Background

The concept of biodiversity which includes variation at the levelf genes species and ecosystems has been evolving over the past5 years after being proposed by Wilson (1988) The present paperrst emphasises that habitats are important as indicators of biodi-ersity in their own right but then shows that they are also linkedo species and assemblages both of plants and other taxa in a vari-ty of ways This is axiomatic for plant species as habitats are oftenefined by phytosociological syntaxa which are determined by veg-tation composition Plant species can also be used as indicatorsor a habitat as described in the Interpretation Manual for Annex

habitats (European Commission 2007) These species can thene used in conjunction with ancillary data such as soils to predict

Please cite this article in press as Bunce RGH et al The significanceEcol Indicat (2012) httpdxdoiorg101016jecolind201207014

he distribution of the habitats themselves across Europe as showny Muumlcher et al (2004 2009) The landscape ecological literaturelso provides many studies where habitats have been used as a

lowast Corresponding author Tel +31 317 481824 fax +31 317 419000E-mail address robjongmanwurnl (RHG Jongman)

470-160X$ ndash see front matter copy 2012 Published by Elsevier Ltdttpdxdoiorg101016jecolind201207014

copy 2012 Published by Elsevier Ltd

framework for examining the behaviour of species Thus Harveyet al (2006) related tree cover to birds butterflies and dung bee-tles while Dover et al (2010) examined how the structure evenwithin a single habitat can effect butterfly abundance and Hinsleyand Bellamy (2000) show how bird assemblages even within onehabitat a hedge are related to management

Although widely used the term lsquohabitatrsquo remains diverseambiguous and difficult to apply consistently because it is has beendeveloped in different contexts with contrasting meanings

12 The legal and policy context

Habitats are specifically referred to in the European Union (EU)Habitats Directive which includes a list (Annex I) of habitats tobe protected (European Commission 1992) These habitats aredescribed in various levels of detail in the Interpretation Man-ual (European Commission 2007) which has been revised severaltimes to include additional habitats and revised definitions as the

of habitats as indicators of biodiversity and their links to species

EU has expanded In the Manual there are some diagnostic featuresas well as lists of associated species Criteria to identify a particularhabitat or to distinguish between them are not included in the legaltext of the Directive

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ARTICLECOIND-1273 No of Pages 7

RGH Bunce et al Ecologi

These habitats together with selected species listed in AnnexI have been used to construct the Special Areas of ConservationSAC) which together with Special Protection Areas (SPA) desig-ated under the 1979 EU Birds Directive and Marine Protectedreas (MPA) form the Natura 2000 network which is the primary

ramework for site based nature protection in the EU EU Membertates are also obliged to report on the conservation status of theseabitats every six years under Article 17 of the Habitats Directivend Article 12 of the Birds Directive

In addition the RAMSAR Convention (httpwwwramsarorg)elates to one habitat group wetlands Habitats are also referred ton the Berne Convention with an agreed list of habitats to be usedforelection of sites for its Emerald Network (Council of Europe 2010)abitat conservation is also one of the 2020 targets (Aichi targets) of

he Convention on Biological Diversity and its status is an importantategory to report on (GEO BON 2011)

Habitats are therefore a central pillar of European nature conser-ation policy The categorisation of European habitats started withhe earlier CORINE biotopes classification (Devillers et al 1991)rom which the original list of habitats for Annex I was selectednd then the Palearctic classification (Devillers and Devillers-erschuren 1996) Annex I has been progressively modified withhe accession of new member states with the most recent versioneing 2007 and recent changes have been based on the Palearcticlassification Evans (2006) has discussed the background to thennex I habitats Although Natura 2000 also protects species listed

n Annex II of the Habitats Directive together with selected birdpecies listed in the Birds Directive it is often considered that theaintenance of a series of habitats in good condition is one of the

est ways to conserve species

3 The scientific ecological research context

The term habitat has often been defined as the spatial extent of resource for a particular species Habitat in this sense is explicitlyinked to a species or species group that share the same ecolog-cal requirements Dictionary definitions are also not specific andsually refer generally to communities The terms lsquohabitat patchrsquo

micro-habitatsrsquo and lsquotemporary habitatrsquo are also often used in thisespect Other terms such as biotope and ecosystem are also usedn similar contexts in the literature but are rarely defined In recentears the latter term has been increasingly used in the concept ofcosystem services but as described by Fisher et al (2009) it is usu-lly applied at a range of different scales from the specific eg arop field to the general such as a riparian zone Ecosystem servicesave to be considered as a policy category additional to traditionalature conservation (Haslett et al 2010) The increasing literaturen this subject uses ecosystem as a general term and leaves thehrase open to a range of interpretations although Haines-Youngnd Potschin (2007) specifically mention the use of habitats as anption to assess

The scientific use of the term habitat shows an evolution ineaning from the vague and broad to the narrow and precise as

hown following examples of definitions

ldquoPlace living space where an organism livesrdquo (Odum 1963) ldquoPlace where a species normally lives often described in termsof physical factors such as topography and soil moisture and byassociated dominant forms (eg intertidal rock pools or mesquitewoodland)rdquo (Calow 1999)

Please cite this article in press as Bunce RGH et al The significanceEcol Indicat (2012) httpdxdoiorg101016jecolind201207014

ldquoHabitat is a zone (area) comprising a set of resources con-sumables and utilities for the maintenance of an organism Theresources occur in union andor intersect and may also be equiva-lent links between resource outlets are established by individual

PRESSicators xxx (2012) xxxndashxxx

searching movements of the organismrdquo (Dennis and Shreeve1996)

Reviews of the application of the term have been made by egHall et al (1997) and Dennis et al (2003)

The above definitions are primarily theoretical and are notdesigned for field mapping with the objective to determine theextent of habitats and their spatial distribution However recentlyBunce et al (2008) have adapted the principles originally devel-oped in the Great Britain Countryside Survey for mapping Europeanhabitats and rules are provided for assignment of a given patch toa habitat class at a defined scale Bunce et al (2008) define habitatas ldquoan element of the land surface that can be consistently definedspatially in the field in order to define the principal environmentsin which organisms liverdquo

Habitat classification can be related to the UN Forest and Agri-culture Organisation (FAO) Land Cover Classification System (LCCS)described by Di Gregorio and Jansen (2000) which is widely usedas a land cover classification system in Africa In addition to theirrecognition in their own right habitats also have the following prac-tical advantages

1 Aerial photographs especially infra-red can be used to estimatehabitat extent and its change over time eg Staringhl et al (2011)

2 Remote sensing data from satellites can be linked to in situ mapsof habitats to larger units eg Muumlcher (2009) Vanden Borre et al(2011) provide an overview of the use of satellite imagery in thisrespect

3 Relationships between habitats and species assemblage compo-sition or particular taxa important to biodiversity can be usedto link habitat records to other biodiversity indicators such asspecies eg Petit and Usher (1998)

4 Habitat records can be linked to changes over time at the land-scape level and to vegetation assemblages as described byHaines-Young and Potschin (2007)

Protocols are now available and can be used to link extanthabitat data across Europe for five national major monitoring pro-grammes (Bunce et al 2012) and could also be developed for othersurveys

14 Application of habitats

The advantages listed above are the principal reasons behind theadoption of habitats as the linking framework in the European Bio-diversity Network (EBONE) an EU 7th Framework project In thisproject the definition of habitat was that described by Bunce et al(2008) in which a restricted list General Habitat Categories (GHCs)were developed for habitat mapping and for linking extant dataThe project extended the use of GHCs outside Europe (Bunce et al2011) and also carried out a quality control and assurance exercisethroughout Europe The main limitation of using habitats is thatthe links with assemblages and individual species are not alwaysexplicit and need interpretation further survey or data analysis asdescribed in Section 5 Habitats have also been used as a frameworkfor collecting other biodiversity data eg in the Great Britain (GB)Countryside Survey (Firbank et al 2003) data on plants vegeta-tion birds and aquatic species have been collected and linked to 19Broad Habitats equivalent to the GHCs These analyses were car-ried out using the following hierarchical structure based on Bunce

of habitats as indicators of biodiversity and their links to species

(1999) with the addition of biome at the highest level

1 Biomes are traditionally defined by a combination of life formsand climate with the balance between then being rarely explicit

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3

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5

6

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2

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ARTICLECOIND-1273 No of Pages 7

RGH Bunce et al Ecologi

Biomes are widely used to express biodiversity resources at theintercontinental scale

Landscapes are comprised of complexes of habitats but theircomplexity is often ignored in traditional ecology but in recentyears in landscape ecology it has been accepted

Habitats may also be termed biotopes or ecosystems and aremainly made up of mixtures of vegetation types although asdiscussed below in some cases they may equate with a singlevegetation class

Vegetation consists of complexes or assemblages of plants Thesemay be described as syntaxa in phytosociology or by samplingobjectively within defined domains In Annex 1 vegetation typesare often synonymous with habitats Assemblages of other taxasuch as birds and butterflies are also used

Species assemblages of plants make vegetation Species may beimportant to conservation in their own right or because of theirimportance in vegetation structure or links with other speciesas described in the present paper

Even within a given species there are often different pools ofgenetic diversity in different parts of the geographical distribu-tion especially in wide ranging species such as Pinus sylvestrisThe use in forestry of provenances for planting appropriate treesin specific regions is a specific example

Other major national survey and monitoring schemes have used comparable structure built around habitat records eg Northernreland (Cooper and McCann 2000) Sweden (Staringhl et al 2011) andustria (Wrbka et al 2004) In the EBONE project a regionalisationf habitats has been applied according to the European Environ-ental Stratification (Metzger et al 2005 Jongman et al 2006)abitats have also been linked with recording of vegetation plotsnd in Israel links have been made between habitat in sites withssemblages of reptiles and insects (Olsvig-Whittaker et al 2012)

History of habitat mapping

The concept of habitat developed initially from the biomes wasescribed by the classical bio-geographers of the 19th century egon Humboldt and Bonpland (1807) Their maps defined the mainiomes across the world eg desert and tropical rainforest and wereased on a combination of observed vegetation and climate Somef these biomes are synonymous with modern concepts of habitatsg deciduous forest but most are on a larger scale Biomes haveontinued to be used at a global scale for modelling impacts oncosystems across the world eg of climate change (Woodward987)

Early in the 20th Century Raunkiaer (1904) formalised vegeta-ion structure by using plant life form spectra in order to defineegions according to their actual vegetation rather than by alsonvolving climate This has been elaborated subsequently for veg-tation mapping (Kuchler 1967 Kuumlchler and Zonneveld 1988ignatti 1982) Recently Bunce et al (2008) based the definition ofabitats on plant life forms in order to transcend species and localerms and to utilise the underlying regression relationship withnvironment In the early 20th century the discipline of vegeta-ion science developed as scientists recognised that plants formedecognisable assemblages leading in due course to the science ofhytosociology (Braun-Blanquet 1932) Mapping has been an inte-ral part of phytosociology at a variety of scales (Pedrotti 2004) butecause of the need to apply expert judgement it has not widelyeen used for monitoring Later in the field of landscape ecology

Please cite this article in press as Bunce RGH et al The significanceEcol Indicat (2012) httpdxdoiorg101016jecolind201207014

he relationships of land use land cover and ecology has been elab-rated by applying earth observation for air and space into landlassifications (Zonneveld 1995) and these are now integrated inabitat interpretation

PRESSicators xxx (2012) xxxndashxxx 3

Areas have been protected for many reasons but the use ofnature reserves to conserve areas for vegetation and indirectlyhabitats did not start until the 1940s eg with the 1947 NationalParks and Access to the Countryside Act in the UK Progressivelyhabitats have been recognised as an essential basis for the selec-tion of representative series of sites for conservation objectives forexample in both the EU Natura 2000 and the Council of EuropersquosEmerald networks Approximately 60 of the habitats of Annex 1of the Habitats Directive are based on plant communities (Evans2010) emphasising the link between vegetation classification andhabitat definition There has also been work providing cross walksbetween the different classifications eg Rodwell et al (2002)related the EUNIS habitats classification (Davies and Moss 2004) tovegetation syntaxa Other national surveys eg the Phase One habi-tats in England described by JNCC (1992 with subsequent revisions)have been undertaken to record habitats as a basis for policy formu-lation There are also a range of national habitat descriptions oftenproduced in support of Natura 2000 eg Czech Republic (Chytryet al 2010) and Estonia (Paal 2007)

3 Incorporation of species data in spatial models ofhabitats

In recent years several models have been produced that dependon knowledge of the correlation between species and habitats Anexample is LARCH (from Landscape Assessment using Rules forConfiguration of Habitat) (Verboom and Pouwels 2004) LARCHdetermines ecological networks for specific species in a patchylandscape and assesses the sustainability of these networks LARCHcomputes potential sustainability of networks based on habitatrequirements and traits of selected species Therefore results candiffer from actual distribution data of species A number of recentLARCH analyses use eco-profiles instead of individual speciesEco-profiles form a classification of animal species that have param-eters within certain limits regarding dispersal capacities and arearequirements In this way results become more general and lessspecific for a single species

Another approach is the BEETLE model described by Watts et al(2007) As in LARCH the spatial arrangement of habitat patches inlandscapes is used and the model assumes that species respond tohabitat fragmentation Eycott et al (2007) also describe the use offocal species eg surrogates and indicators as a means of evaluatingbiodiversity at the landscape scale

4 Habitats as policy indicators

The development of biodiversity indicators in Europe has beenheavily influenced by the requirements of the Convention of Bio-logical Diversity (CBD) The CBD set a target ldquoto achieve by 2010a significant reduction of the current rate of biodiversity loss atthe global regional and national level as a contribution to povertyalleviation and to the benefit of all on Earthrdquo whilst in Europe 51countries in the United Nations Economic Commission for Europe(UNECE) region adopted the Kiev Resolution in 2003 to ldquoreinforceour objective to halt the loss of biological diversity at all levels bythe year 2010rdquo These targets could not be reached due to adversesocietal processes but there were also difficulties in monitoringthem due to the lack of (harmonised) data (Braat and ten Brink2007) New targets for 2020 were adopted by the CBD in 2010 ndashlsquothe Aichi targetsrsquo (see httpwwwcbdintsptargets) At its Con-ference of the Parties to the Convention in Nagoya in 2010 the

of habitats as indicators of biodiversity and their links to species

CBD agreed to the new strategic plan for biodiversity including thetwenty lsquoAichi targetsrsquo for the period 2011ndash2020 and asked GEOBON to help advice on how the datasets can be assembled The ini-tial step in this process was an Adequacy Report (GEO BON 2011)

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ARTICLECOIND-1273 No of Pages 7

RGH Bunce et al Ecologi

ichi target 5 is to ldquoBy 2020 the rate of loss of all natural habi-ats including forests is at least halved and where feasible broughtlose to zero and degradation and fragmentation is significantlyeducedrdquo Within the EU the Commission has published a biodi-ersity strategy for 2020 which also includes a series of targetsEuropean Commission 2011)

In the Streamlining European 2010 Biodiversity Indicators (SEBI010) project a set of indicators was developed to meet the CBDequirements The 26 SEBI ldquoheadlinerdquo indicators are clusteredithin the 7 CBD focal areas and were selected according to defined

riteria The set is not designed to be comprehensive but to pro-ide the best coverage on the basis of available information andesources The technical report containing specifications of the 26ndicators selected was published in EEA (2007) One of these indi-ators related to the conservation status of habitats of Europeannterest as listed in Annex I of the Habitats Directive

The extent and condition of habitats is an important and use-ul measure of biodiversity It is also a legal requirement in theU to monitor and regularly report on the Conservation Status ofhe habitats of Annex 1 The Countryside Survey of the UK (Haines-oung et al 2000) has shown how habitats can be used to integrateata from the landscape vegetation and species scales The EBONEroject has recently shown how the same principles can be usedhroughout Europe and beyond (Roche and Geijzendorffer 2012)he condition of habitats is considered to be related to the distri-ution and abundance of many other species and populations ofalue and there has been much discussion of whether biodiversityan best be conserved by focusing on habitats rather than indi-idual species Habitats can also provide the basis for assessmentsf ecosystem services as suggested by Haines-Young and Potschin2007) and in the Countryside Survey (2000) and GHCs could bedapted for the same objectives

The current policy measures do not however cover many habi-ats typical of the wider countryside that are often highly managednd disturbed In practice the majority of European agriculturalandscapes have no habitats protected under current EU legisla-ion but yet much biodiversity is present within them albeit oftenestricted to habitat fragments Such landscapes and habitats there-ore need to be included in any overall assessment of biodiversityesources in the EU

Links between habitats and species

Relationships between habitats and species can be identified inhree ways

Species or assemblages of species can be expected to be foundwithin a certain habitat within a given location in an Environ-mental Zone as described by Metzger et al (2012)

Species occurring in a certain habitat are characterising thathabitat by their traits such as Ellenberg values

The occurrence of a given habitat within an Environmental Zonean be predicted from the presence of a certain species known toe present within it Information about such links can be carriedut using the following sources of information

Distribution maps of species eg the Atlas Flora Europea can belinked to known distribution of habitats

Databases on Ellenberg values can be exploited such as

Please cite this article in press as Bunce RGH et al The significanceEcol Indicat (2012) httpdxdoiorg101016jecolind201207014

httpstatedvbokuacatzeigerwerte httpwwwcehacukproductssoftwareCEHSoftware-MAVIShtm and literature andrelated databases can be searched (Pignatti et al 2005 Fanelliet al 2007)

PRESSicators xxx (2012) xxxndashxxx

3 Information can be extracted from field guides and formal keysto link species to habitats eg Loxia scotica in Caledonian pineforest (Annex I habitat 91C0)

4 Literature reviews can be carried out of detailed work torelate species to habitats eg Sitta europaea to deciduous forestpatches

5 There are monographs on individual species eg Prunella mod-ularis or overviews of species given in for instance the Floraof the British Isles (see httpwwwbritishecologicalsocietyorgjournals publicationsjournalofecologybiologicalfloraphp)

6 Expert knowledge of individual scientists can be utilised todevelop specific relationships

7 Individual studies can be carried out to determine relationshipswithin given habitats in a given landscape

Some Red Data Book species are related to specific habitats egarable weeds in crops However in many cases the distribution ofrare species is accurately known because of the interest of conser-vation biologists in such species In these cases the relationship isnot with a habitat in general but with a locality eg Cabo da Roca(Portugal) for Armeria pseudarmeria and Omphalodes kuzinskyanae

Distribution maps of some vascular plant species are availablefrom the Flora Europea and in local flora and atlases eg the CzechRepublic and the United Kingdom National Biodiversity ActionPlans for species usually specify the habitat measures required toprotect such species Other taxa can be related to individual plantspecies on which they depend for food eg butterflies and changescan then be modelled eg Schweiger et al (2012)

It is also possible to derive links statistically from vegeta-tion databases for some plant species eg through the use ofSynBioSys (httpwwwsynbiosysalterranlsynbiosyseu) which isderived from the phytosociological literature When detailed infor-mation is available from samples structured at the landscape leveleg Firbank et al (2003) then vegetation assemblages can be relatedstatistically to habitats in order to present results in a format whichis understandable to policy makers They can also be used as a basisfor comparing changes in biodiversity

There are a range of different types of relationships betweenspecies and habitats and some examples are given below

1 Generalist species without relationships to specific habitats egPica pica Streptopelia decaocto Agrostis capillaris

2 Species that are linked to several habitats eg Columba palum-bus Falco columbarius Tamus communis

3 Species which occur in one habitat in different EnvironmentalZones eg Branta leucopsis Briza media Cirsium acaule

4 Species that occur in one habitat in one Environmental Zone egPinus mugo Abies pinsapo Ziziphus lotus

5 Species which are dependent on other species for food and can bepredicted from the occurrence of that species eg many bumblebees or butterflies with certain plants Phengaris alcon is depend-ing on both Gentiana pneumonanthe as a host plant and coloniesof various ant species (Myrmica spp)

6 Species which need a high quality of a given habitat type egsome Sphagnum species are only present in undisturbed bogs

7 Species which have different habitat requirements in differentEnvironmental Zones eg Huperzia selago Silene acaulis Sax-ifraga tridactylites

It is often difficult to attach species to habitat types except forthose available in individual scientific papers This is especially truefor generalist species which can occur in a wide range of habitats

of habitats as indicators of biodiversity and their links to species

As described above these have been coordinated in some modelseg LARCH to enable prediction at the landscape sale Firbank et al(2003) have also demonstrated links between habitats defined bystatistical analysis of vegetation can be linked to species within

ARTICLE IN PRESSG ModelECOIND-1273 No of Pages 7

RGH Bunce et al Ecological Indicators xxx (2012) xxxndashxxx 5

Table 1Relationships between Annex I forest habitat types and environmental zones including characteristic species and compared with non-Annex I forest types (EuropeanCommission 2007 Polunin and Walters1985 Noirfalise 1987 Jongman and Bunce 2000 Bohn et al 2000)

Env ZONE Annex I forest habitats Characteristic species taken from theEU interpretation manual

Non-annex I forest habitats occurringin environmental zone

ALN 9040 Nordic subalpinesubarcticforests with Betula pubescens sspCzerepavonii

Cornus suecica Pinus sylvestris forest

BOR 9010 Western Taiga Maianthemum bifolium Betula pubescens forestNEM 9030 Natural forests of primary

succession stages of land upheavalcoast

Molinia caerulea Populus tremula forest

ATN 91A0 Old sessile oak woods with Ilexand Blechnum in the British Isles

Blechnum spicant Picea sitchensis plantations

ATC 9120 Atlantic acidophilous beechforests with Ilex and sometimes alsoTaxus in the shrub layer (Quercionrobori-petraeae)

Teucrium scorodonia Picea abies plantations

CON 9110 Luzulo-Fagetum beech forests Luzula albida Pinus sylvestris plantationsPAN 91H0 Pannonian woods with Quercus

pubescensCarex humilis Populus spp plantations

LUS 9240 Quercus faginea and Quercuscanariensis Iberian woods

Quercus faginea Pinus radiata plantations

ALS 9420 Alpine Larix decidua andor Pinuscembra forests

Rhododendron ferrugineum Alnus incana forest

MDN 9540 Mediterranean pine forests with Erica arborea Populus spp plantations

nea sativa Pinus sylvestris forestia lentiscus Eucalyptus spp plantations

dht

hwGpbaiZ

sqmht

1

2

3

6b

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Table 2Examples of species in Environmental Zones The species are linked to conifer forests(FPHCON) and deciduous forest (FPHDEC) in these zones (European Commission2007 Polunin and Walters 1985 Noirfalise 1987 Jongman and Bunce 2000)

Env zone GHC Birds Plants

ALN FPHCON Loxia curvirostra Trientalis europeusFPHDEC Carduelis flavirostris Matteuchia struthiopteris

BOR FPHCON Lophophanes cristatus Linnae borealisFPHDEC Dryocopus martius Galium boreale

NEM FPHCON Tetrao urogallus Lycopodium annotinumFPHDEC Parus major Anemone ranunculoides

ATN FPHCON Loxia scotica Goodyera repensFPHDEC Sitta europea Lonicera periclymenum

ATC FPHCONFPHDEC

Dendrocopos major Pinus sylvestris

Certhia familiaris Maianthemum bifoliaCON FPHCON Columba palumbus Pyrola minor

FPHDEC Phylloscopus collybita Luzula albidaPAN FPHCON Tetrao tetrix Stachys pannonica

FPHDEC Cyanistes caeruleus Staphylea officinalisLUS FPHCON Carduelis spinus Lithospermum diffusum

FPHDEC Sylvia undata Narcissus triandraALS FPHCON Lophophanes cristatus Homogyne alpine

FPHDEC Dryocopus martius Alnus incanaMDN FPHCON Milvus milvus Pinus nigra

FPHDEC Ciconia ciconia Quercus fagineaMDM FPHCON Streptopelia risoria Pinus mariacutetima

FPHDEC Pyrrhocorax pyrrhocorax Quercus pubescens

endemic Mesogean pinesMDM 9260 Castanea sativa woods CastaMDS 9320 Olea and Ceratonia forests Pistac

ifferent landscapes The links between plants and invertebratesave also been used to assess changes in associated taxa eg but-erflies and bumble bees

The sparsely vegetated habitats described by Bunce et al (2011)ave been used in the desert in Israel to assess their relationshipsith other taxa as described by Olsvig-Whittaker et al (2012)HCs were mapped in Avdat in the Negev desert and statisticalrocedures used to correlate reptiles flowering plants and inverte-rates with the habitat records Although species richness was notssociated both assemblages and individual species showed signif-cant correlations Further work is required in other Environmentalones and habitats

Several theoretical exercises eg Halada et al (2010) have linkedpecies to existing habitat classifications Such relationships are notuantitative and provide no details of probabilities There are alsoany papers in the landscape ecological literature demonstrating

ow faunal species interact with habitats in the landscape of whichhe following are examples

Birds respond to habitat composition and structure for examplebird identification books specify the habitat in which a givenbird is likely to be found In the scientific literature There arealso papers such as Hinsley and Bellamy (2000) which describethe influence of habitat structure and management on bird pop-ulations

Beetles have also been widely studied as they also respond tohabitat composition and structure For example Petit and Usher(1998) discuss the ground beetle communities in woody uncul-tivated habitats

Some butterflies depend upon specific plants for food and nectarbut they also depend on habitats For example Dover et al (2010)describe how butterflies depend on management within a singlehabitat

Worked examples of potential links at a European level

Please cite this article in press as Bunce RGH et al The significanceEcol Indicat (2012) httpdxdoiorg101016jecolind201207014

etween species and habitats

Expert opinion linked to Environmental Zones Tables 1 and 2iven below are based on expert judgement and show the type

MDS FPHCON Cyanopica cyana Pinus halepensisFPHDEC Sylvia mystacea Fraxinus ornus

of relationships which are widely known These tables could beimproved by a more detailed literature review of individual studiesdescribing species behaviour in habitats

7 Discussion

The primary policy requirement for habitats in Europe is for esti-mates of their extent and the changes taking place The EBONE

of habitats as indicators of biodiversity and their links to species

project has provided a framework for obtaining such estimatesSeveral individual countries eg Great Britain have already pro-duced figures for stock and change of habitats and these have beenlinked with vegetation samples at both species and assemblage

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ARTICLECOIND-1273 No of Pages 7

RGH Bunce et al Ecologi

evels as presented by Haines-Young et al (2000) However therere still many gaps in the coverage especially in the Mediterraneanones One of the problems is that member states have existingonitoring programmes and the production of international fig-

res is usually not a priority However a programme of screeningxisting data sources could provide a basis for setting up a longerm monitoring system A statistical procedure to develop a sam-ling framework for the EU has been designed by Metzger et althis volume)

Several extant EU databases could also be used to developxplicit links between species and habitats eg phytosociologicalata in SynBioSys (httpwwwsynbiosysalterranlsynbiosyseu)ould provide a statistical framework for plant species Some ofhe European databases on other taxa eg birds and butterflieslso have the potential for statistical analyses Studies could leado better integrated models of habitats and species and provide atimulus for policy support measures Inevitably rare species couldot be included in such analyses although they are often related topecific habitats

Finally many countries eg the Czech Republic and Estonia haveublished their own habitat classifications with links to Annex Ihile others (eg France Italy) have published local interpretations

f Annex I habitat types (Evans 2012) It would be valuable if theseexts could be linked to provide better overall descriptions of Annex

habitats and species

Conclusions

The present paper emphasises the importance of habitats in theevelopment of biodiversity policies in their own right and alsoemonstrates that there are strong links with species These may beormalised through statistical analysis or by expert knowledge butither way habitats are a central pillar of nature protection policyn the EU The eventual production of estimates of the extent statusnd changes in European habitats would provide an important toolor policy makers across the EU

cknowledgements

This research has been carried out in the framework of the ECP7 project EBONE (EC-FP7 Contract ENV-CT-2008-212322) ando-financed by the Dutch Ministry of Economics Agriculture andnnovation Project code KB14-002-007 It is part of the Europeanontribution to GEO BON

eferences

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Staringhl G Allard A Esseen P-E Glimskaumlr A Ringvall A Svensson H SundquistS Christensen P Gallegos Torell A Houmlgstroumlm M Lagerqvist K Marklund LNilsson B Inghe O 2011 National Inventory of Landscapes in Sweden (NILS)ndash scope design and experiences from establishing a multiscale biodiversitymonitoring system Environ Monit Assess 173 579ndash595

Vanden Borre J Paelinckx D Muumlcher CA Kooistra L Haest B De Blust GSchmidt A 2011 Integrating remote sensing in Natura 2000 habitat monitor-ing prospects on the way forward J Nat Conserv 19 (2) 116ndash125

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of habitats as indicators of biodiversity and their links to species

Linking pattern and process in cultural landscapes An empiricalstudy based on spatially explicit indicators Land Use Policy 12 12httpdxdoiorg101016jlandusepol200310012

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