conservation policy and the eu habitats directive: favourable conservation status as a measure of...

Copyright © 2007 John Wiley & Sons, Ltd and ERP Environment

European EnvironmentEur. Env. 17, 363–375 (2007)

Published online 22 October 2007 in Wiley InterScience

(www.interscience.wiley.com) DOI: 10.1002/eet.458

Conservation Policy and the EUHabitats Directive: Favourable

Conservation Status as a Measure ofConservation Success

Johanna Mehtälä* and Timo VuorisaloDepartment of Biology, University of Turku, Turku, Finland

ABSTRACTThe Habitats Directive is the key instrument for biodiversity conservation in the Euro-pean Union. In the Habitats Directive, maintenance or restoration of natural habitatsand populations of wild species of Community interest at a favourable conservationstatus (FCS) is defined as an overall objective of conservation measures. We arguethat the concept could serve as a realistic and useful measure of conservationsuccess, as for as practical problems related to its application can be solved. Theproblems with FCS stem primarily from difficulties in organising comprehensivespecies and habitat monitoring networks at both the national and European scales,lack of data on historical changes in habitat and species distributions, habitat identi-fication problems, difficulties in the study of habitat-specific structures and functions,and scaling problems. We present conceptual clarifications to the concept, and rec-ommend inclusion of within-species genetic variability and functional characteristicsof ecosystems into assessment of FCS. We also present recommendations on theestablishment of nationally representative species and habitat monitoring networks.Copyright © 2007 John Wiley & Sons, Ltd and ERP Environment.

Received 23 November 2006; revised 6 July 2007; accepted 10 July 2007

Keywords: favourable conservation status; precautionary principle; Habitats Directive; Bonn Convention; ecology; species

monitoring; habitat monitoring; sustainability

Introduction

THE CONCEPT OF SUSTAINABLE DEVELOPMENT HAS BECOME INCREASINGLY IMPORTANT SINCE 1980when the first World Conservation Strategy was published (IUCN, UNEP and WWF, 1980). Espe-

cially since the Brundtland Commission’s report Our Common Future (WCED, 1987), it has

achieved prominence in international law and environmental policy, as exemplified by the Rio

*Correspondence to: Johanna Mehtälä, Department of Biology, University of Turku, FI-20014 Turku, Finland. E-mail: [email protected]

364 J. Mehtälä and T. Vuorisalo

Earth Summit in 1992 and the Johannesburg Summit on Sustainable Development in 2002. The

general idea of sustainable development is reconciliation of environmental protection and economic

growth. An important component of sustainability is maintenance of ecological stability and resilience

(Howarth, 1997). It is understood that life on earth may be our most critical natural resource, which

however is now threatened by a multitude of interrelated environmental problems ranging from loss of

fertile soils to climate change. Consequently, at the Johannesburg Summit a rather ambitious global

target was set by the governments, i.e. to significantly reduce the current rate of biodiversity loss by 2010

(United Nations, 2002).

In the European Union, the Habitats Directive is the key instrument for biodiversity conservation

(Council Directive 92/43/EEC). It takes into account not only biodiversity but also economic, social, cul-

tural and regional requirements. Thus the Habitats Directive also contributes to the general objective of

sustainable development in the territory of the European Union (Palerm, 2006). Its practical objective

is to create a coherent ecological network of special areas of conservation (SACs) called ‘Natura 2000’,

covering the types of natural habitat and wild animal and plant species that are considered to have special

importance to the Community. In addition to the SACs, the Natura 2000 network includes special

protection areas for birds (SPAs) defined by the Birds Directive (Barberán et al., 2005). As stated in the

preamble of the Habitats Directive, the overall aim of this network is ‘to ensure the restoration or main-

tenance of natural habitats and species of Community interest at a favourable status’. It is thus clear

that the Habitats Directive will have a central role in attempts to halt biodiversity decline in the Union’s

territory by 2010 (Göteborg European Council, 2001).

In this paper we focus on the concept of favourable conservation status (FCS hereafter), which serves

as an overall target of conservation measures in the European Union, and thus promotes sustainable

development. In addition to the Habitats Directive, the concept is used in the Bonn Convention, or the

Convention on the Conservation of Migratory Species of Wild Animals. Thus, it has potential influence

upon conservation policies and practices not only in the 25 member states of the European Union, but

in all the 101 states all over the world that by the 1 April 2007 had become parties of the Bonn Con-

vention (Convention on Migratory Species, 2007). Considering the importance of the concept, it is sur-

prising that FCS is not even mentioned in major textbooks of ecology or conservation biology (e.g. Meffe

and Carroll, 1997; Begon et al., 2006; Frankham et al., 2004; Primack, 2002). Also, only a few papers

have been published that critically analyse its contents from an ecological perspective (but see Syrjänen,

2001; Halahan and May, 2003).

The purpose of this paper is to consider, from both biological and environmental policy perspectives,

how well the concept of FCS succeeds in serving as a target of biodiversity conservation efforts. This

largely depends on how well FCS can be applied to more specific and ecologically relevant questions,

and how crucial these questions are from the perspective of practical conservation biology and policy

objectives. For practical reasons we focus on applications of the concept within the European Union.

FCS as a Concept: History, Definition and Links to Related Concepts

The history of FCS can be traced back to the early 1970s, when preparations for the Bonn Convention

started as a consequence of Recommendation 32 of the Action Plan accepted at the United Nations Con-

ference on the Human Environment, held in Stockholm in 1972 (Lyster, 1994). The Bonn Convention

was signed on 23 June 1979 (Convention on Migratory Species, 2007). At the same time preparations

of two other multilateral commitments were under way in Europe. These were the EC Birds Directive

(Council Directive 79/409/EEC of 2 April 1979) and the Convention on the Conservation of European

Wildlife and Natural Habitats (i.e. the Berne Convention). Although these two documents did not

Copyright © 2007 John Wiley & Sons, Ltd and ERP Environment Eur. Env. 17, 363–375 (2007)DOI: 10.1002/eet

Conservation Policy and the EU Habitats Directive 365

explicitly use the concept of FCS, both of them referred to a targeted population level that should cor-

respond ‘in particular to ecological, scientific and cultural requirements, while taking account of eco-

nomic and recreational requirements’ (Berne Convention, 1979; Council Directive 79/409/EEC). All

three conventions shared a strong ecological emphasis (Lyster, 1994), and influenced the contents of

the EC Habitats Directive, which came into force on 21 May 1992. In particular, the Berne Convention

can be regarded as a precursor to the Habitats Directive (Bennett and Ligthart, 2001).

The concept of favourable conservation status in the Habitats Directive was derived from the Bonn

Convention. In the Bonn Convention the concept is applied only for endangered migratory species

included in Appendices I or II, while in the Habitats Directive it is used for both habitat types listed in

Annex I and species listed in Annexes II, IV and V. Not surprisingly, the two definitions of FCS for

species are practically identical, as well as criteria 1–3 for species (Table 1). Although the Bonn

Convention does not define FCS for habitats, its Criterion 4 has implications for the management of

habitats.

Summarized from Table 1, the key criteria for FCS of a habitat are (1) stable or increasing natural

range, (2) long-term survival of its specific structure and functions and (3) favourable conservation status

of its ‘typical’ component species. The criteria for FCS of a species are (1) the population dynamics data

must indicate good chances for long-term survival in its natural habitats, (2) the natural range of the

species must be stable or increasing, (3) there has to be a sufficient amount of suitable habitat and (4)

the distribution and abundance of the species approach historic coverage and levels. Criterion 4 is not


EC Habitats Directive Bonn Convention

Specified objectives Specified objectivesTo promote the maintenance of biodiversity, Conservation of migratory species and their habitats,

taking account of economic, social, cultural ‘paying special attention to migratory species theand regional requirements (preamble) conservation status of which is unfavourable’

(Article II)

Use of FCS/criteria Use of FCS/criteriaFCS of a habitat: (1) its natural range and areas Criteria of FCS: (1) population dynamics data

it covers within that range are stable or increasing, indicate that the species is maintaining itself on a(2) the specific structure and functions that long-term basis as a viable component of itsare necessary for its long-term maintenance exist ecosystems, (2) the range of the species is neitherand are likely to continue to exist for the foreseeable currently being reduced, nor is likely to be reducedfuture and (3) the conservation status of its typical on a long-term basis, (3) there is, and will be in thespecies is favourable (Article 1(e)) foreseeable future, sufficient habitat to maintain the

population on a long-term basis and (4) the distributionFCS of a species: (1) population dynamics data on and abundance of the species approach historic

the species concerned indicate that it is maintaining coverage and levels to the extent that potentiallyitself on a long-term basis as a viable component of suitable ecosystems exist to the extent consistentits natural habitats, (2) the natural range of the with wise wildlife management (Article 1c)species is neither being reduced nor likely to bereduced for the foreseeable future and (3) there is,and will probably continue to be, a sufficiently largehabitat to maintain its populations on a long-termbasis (Article 1(i))

Table 1. Definitions of the conservation status and criteria of the FCS in the EC Habitats Directive and in the Bonn Convention(Council Directive 92/43/EEC; Convention on Migratory Species, 2007). These criteria apply at the national or European Unionlevel. In Annex III of the Habitats Directive criteria are listed also for individual site assessment. For instance, in site assessmentfor a given natural habitat type also the degree of representativity of natural habitat type on the site should be evaluated


explicitly included in the Habitats Directive, but is often mentioned as a possible target of conservation

efforts.

In addition to these national or Community-level criteria, additional site-specific criteria are given in

Annex III of the Habitats Directive. These will be discussed below. In Table 2 we have listed ecological

questions that can be derived from the criteria for FCS in the Bonn Convention and in the Habitats

Directive. Some of them, and practical guidelines for integrating ecology into political decision-making

related to FCS, will also be discussed below.


Conservation criteria Specific ecological questions Suggested methods

Species:1. Long-term viability of species What is the minimum viable Population viability analysis

populations population size for each species? (PVA); metapopulationHow big a proportion of local studies; landscape analyses;

populations needs to be viable in genetic studies of localorder to make conservation status populations.nationally favourable?

What are the population structuresof protected species?

Are the conservation statuses ofgenetically distinct local populationsfavourable?

2. Stable or increasing natural How to define natural range? Organizing comprehensiverange national population monitoring

networks.

3. Long-term persistence of What is the minimum dynamic area Organizing comprehensivesufficiently large areas of for each species? national habitat monitoringsuitable habitat Is habitat restoration required? networks.

4. The range and levels of Are historical data available? Collection of archaeologicaldistribution and abundance Is suitable habitat available outside and historical data; habitatapproach historic coverage current range? monitoring.

Is habitat restoration required?

Habitats:1. Stable or increasing natural range How to identify habitat types? Organizing comprehensive

national habitat monitoringnetworks.

2. Long-term persistence of specific What are specific structures and Comparative ecosystem studies;structure and functions functions? organizing comprehensive

How much diversity is required monitoring networks forto maintain specific structures and selected habitat structures andfunctions? functions.

Do species substitutions affectspecific structures and functions?

3. Favourable conservation status of What are ‘typical’ species? Organizing comprehensivetypical species of the habitat How large a habitat area is required national monitoring networks for

to maintain a characteristic species keystone or ‘charismatic’ species.pool?

Table 2. Ecological interpretation of the criteria of FCS listed in Table 1


FCS belongs to a family of environmental concepts closely related to the precautionary principle, and

as highlighted above to sustainable development, which is clearly an important general objective behind

the concept of the FCS. The precautionary principle is a general rule of public policy action to be used

in situations of potentially serious or irreversible threats to health or the environment, where there is a

need to act to reduce potential hazards before there is strong proof of harm, taking into account the

expected costs and benefits of action and inaction (Harremoës et al., 2001). The idea behind this defin-

ition is that beyond some threshold level the risk of serious or irreversible changes is thought to markedly

increase, although the scientific evidence for such an increase may not yet be conclusive. Setting a target

conservation level, such as the FCS of a habitat type or a species, is thus inevitably a precautionary act,

as it is assumed that failing to maintain or achieve the targeted level may result in a potentially irre-

versible loss, i.e. extinction. It is also very typical that scientific consensus as to what constitutes the

optimal target level in conservation is lacking (Berrens, 2001).

The safe minimum standard (SMS) approach of natural resource economics, first proposed by Ciriacy-

Wantrup (1952), also parallels the use of the FCS in many respects. The SMS approach requires, in

accordance with the precautionary principle, that some safe minimum level of a renewable natural

resource be protected unless the social costs of doing so are somehow intolerable (Berrens et al., 1998;

Berrens, 2001). For instance, the United States Endangered Species Act of 1973 is generally considered

to be consistent with an SMS-type approach.

As applied to biodiversity conservation, an important initial element of the SMS approach involves

identifying a critical biological threshold that should be respected if extinctions are to be avoided. Berrens

(2001) emphasized that the SMS approach can be applied not only to individual species, but also to habi-

tats, number of hectares preserved or some ‘emerging’ indices of biodiversity. The main difference

between the SMS approach, as applied in the US Endangered Species Act, and the FCS approach in the

Habitats Directive, is that economic considerations have greater importance in the former. The SMS

approach by definition requires that the costs of conservation efforts should not be ‘excessive’, ‘intoler-

able’ or ‘unacceptably large’ (Berrens et al., 1998; Berrens, 2001). According to the Endangered Species

Act of 1973, as amended, setting aside of critical habitats for species preservation can only be done after

an economic analysis of the resulting impacts. In practice this has sometimes resulted in a substantial

reduction of the proposed critical habitat (Berrens et al., 1998). The situation is quite the reverse in the

Habitats Directive, which explicitly requires that site choice for the Natura 2000 network will be done

entirely on ecological grounds (see Annex III of the Habitats Directive). This does not, however, mean

that all forms of economic land use are prohibited within or near areas belonging to the Natura

2000 network. Article 6(3) of the Habitats Directive establishes a procedure of environmental

impact assessment in which the potential negative impacts of proposed projects on a Natura 2000 site

are examined. Article 6(4) also allows in some cases natural capital to be traded off for economic and

social capital.

Problems and Challenges

Missing Historical Reference Data

The word ‘favourable’ implicitly refers to some ideal or target conservation level of a species or a habitat

type, with which the ‘favourability’ of the prevailing situation is compared. There are two problems with

this. First, the ‘ideal’ conservation status needs to be defined. Second, we need to decide how large devi-

ations from the ‘ideal’ long-term mean of population size or distribution area, for instance, are permit-

ted without losing the presumed ‘favourability’ of the situation.



One possibility is to set the historical range of a species or habitat type, or some historical population

size, as targets of conservation efforts. This makes sense if the historical range or population size was

larger than, or as large as, the present one. Interestingly, also in assessment of the recently introduced

Biodiversity Intactness Index the optimal comparison is considered to be the population size that

prevailed before landscape alteration by modern industrial society (Scholes and Biggs, 2005).

Nothing prevents adopting this policy if historical ranges or population sizes are known, suitable

vacant habitats still exist and estimated costs are not intolerable. This may be the case if some other

factor than habitat loss, for instance active persecution, has been the primary cause of original popula-

tion decline or shrinking distribution. For example, the ecologically tolerant but historically severely per-

secuted gray wolf (Canis lupus) has been successfully reintroduced to many areas within its former range

(Carroll et al., 2006).

The most compelling counterarguments against use of historical ranges or population sizes are that

in most cases they are unknown, or if known it is not possible to restore them due to irreversible habitat

changes. Reintroductions present well known examples of such difficulties. Before the large-scale

deforestation in Central and Western Europe (including Britain), the Eurasian lynx (Lynx lynx) was a

widespread feline species (Kurtén, 1968; Yalden, 1999). There have been attempts to reintroduce lynxes

to some areas within their former range in Central Europe. Kramer-Schadt et al. (2005) studied the

recovery and reintroduction of lynxes in the fragmented forest landscapes of Germany. They argued that

the suitable habitats for lynx are so patchily distributed that they may in the long run fail to sustain

viable reintroduced populations. The conclusion is, unfortunately, that due to habitat loss the historical

range of this species may be unfit to serve as target of conservation efforts.

As suggested by Syrjänen (2001) and Scholes and Biggs (2005), it is generally more realistic to con-

centrate on maintaining at least the current ranges or population sizes. As regards population sizes, the

target population sizes should be determined on the basis of long-term means or trends to account for

natural population fluctuations. Population viability analyses (PVAs) can be applied to assess the chances

for long-term survival of local populations. The problem, however, is how to extrapolate from these local

studies to national or regional levels in the hierarchical assessment of FCS (see below).

Habitat Identification Problems

According to the Habitats Directive, the habitat of a species means an environment defined by specific

abiotic and biotic factors, in which the species lives at any stage of its biological cycle (Article 1(f)). In

practice it may not be easy to say whether a certain habitat type occurs in an area or not. This is because

the distinctiveness of habitat types is much lower than that of separate species. While different species

are, by definition, reproductively isolated from one another, the different habitat types are often poorly

defined in terms of their ‘characteristic structure and functions’, and habitats with intermediate char-

acteristics are common. Especially in the marginal areas of their natural ranges the habitats often mix

with other habitat types, and are therefore difficult to discern.

The reason for this is the dynamic nature of the constituent communities of habitat types. The bio-

logical communities typical of certain habitat types change continuously through several processes,

including community ‘microevolution’ (i.e. evolutionary changes in genetic constitution of populations

inhabiting the habitat) and succession, in which replacement of community types takes place in one

locality (Aarssen and Turkington, 1983). The natural range of a habitat type is especially difficult to study,

because the species composition and hence apparently also ‘the specific structure and functions’ of the

habitat type usually vary geographically. Defining any ‘natural ranges’ is also becoming increasingly dif-

ficult due to climate change, that is predicted to cause major shifts in ranges of species, especially in

the higher latitudes.



For instance, the species composition of vascular plants in the forest types of Finland changes latitu-

dinally (Kalliola, 1973). In all forest types dominated by coniferous trees the relative abundance of aspen

(Populus tremula) decreases from south to north, probably for climatic reasons. Some characteristic

species of particular forest types may even be entirely absent from certain areas. A good example is the

typical northern forest herb, the dwarf cornel Cornus suecica, which in the south only occurs in coastal

areas. Opposite examples are the may lily Maianthemum bifolium and the stone bramble Rubus saxatilis,which are missing from the high latitudes (Kalliola, 1973). Still, all these species are considered as

‘typical’ for certain forest types in Finland.

This presents practical difficulties for any attempts to classify habitat types or measure their conser-

vation statuses. Habitat types simply are not spatio-temporally restricted objects with clear boundaries

that can be observed in the field. As Looijen and van Andel (2002) have put it, ‘There are two ways of

describing changes in species composition (species turnover) in some area in terms of community

dynamics. The one is to say that the community that used to exist in the area still exists, but that, because

of the change in species composition, it has “internally” changed . . . The other way is to say that, because

of the change in species composition, the community that used to exist in the area no longer exists but

has turned into another community of another type’.

Such problems can be tackled only by explicitly including geographical variability of habitat types in

the regional monitoring programmes of biodiversity. This presents challenges also for listing of ‘typical’

species of habitats, as also these vary geographically.

Problems with Identification of Habitat-Specific Structures and Functions

In spite of the practical difficulties listed above it is clear that, based on physical conditions and a mul-

titude of biotic interactions, habitats can be, and have for a long time been, classified for both practical

and scientific purposes. For instance, in Finland the so-called forest type theory of Cajander (1909), pri-

marily based on indicator species of ground-layer vegetation, has for nearly a century dominated both

forestry practices and forest ecology. A more widespread habitat categorization is included in Annex I

of the Habitats Directive. The list includes the natural (or semi-natural) habitat types of Community

interest, and provides the basis for their protection.

The favourability of conservation status of a particular habitat type requires that the specific struc-

tures and functions that are necessary for its long-term maintenance continue to exist for the foresee-

able future. Obviously, we first need to identify these important habitat characteristics (Table 2). A good

candidate for a distinctive habitat characteristic is species or functional group diversity, especially for

some highly productive habitat types. Darwin argued for ‘the principle that the greatest amount of life

can be supported by great diversification of structure’, implying that high diversity often results in high

productivity. The mechanism was niche differentiation (although he did not use the term): ‘So in the

general economy of any land, the more widely and perfectly the animals and plants are diversified for

different habits of life, so will a greater number of individuals be capable of there supporting them-

selves’ (Darwin, 1859).

There is now, nearly 150 years after publication of Darwin’s book, strong evidence that biodiversity

may indeed influence ecosystem processes such as productivity and nutrient cycles. Although there is

some uncertainty on whether it is diversity per se, or the presence of a few keystone species, represent-

ing the most important functional groups, that determines the level of functioning of the ecosystem,

the general pattern seems to be that of increasing productivity up to a certain limit (Kinzig et al., 2002).

High productivity can undoubtedly be considered a specific characteristic of ecosystems such as tropi-

cal rainforests and estuarine ecosystems.



In most cases, however, the specificity of ecosystem structure and functions will most likely be found

in some complex combination of structural and functional characteristics of the ecosystem. Besides the

level of productivity, potentially diagnostic habitat characteristics include species or functional group

richness, species composition (including the presence of some indicator species), the number of trophic

links and their distribution among species, the distribution of species interaction strengths and also

physical conditions (e.g. annual rainfall) that set limits to the range of particular habitat types (Ebenman

and Jonsson, 2005). The so-called ‘response diversity’, defined as the diversity of responses to environ-

mental change among species contributing to the same ecosystem function, may be of special impor-

tance to ecosystem resilience (Nyström, 2006).

Considering the practical need to assess the conservation status of habitat types, two questions are

here of particular interest. First, how large a habitat area is required to maintain a characteristic local

species pool? Second, if some species loss will happen anyway, to what extent can habitats tolerate species

loss without losing their specific structure and functions?

Some insight into the first question can be derived from the well known species–area relationships

(Tilman, 1999). According to the classic formulation the number of species in a locality (S) depends on

the area of the sites (A) raised to the power of z (MacArthur and Wilson, 1967). If A1 is the area of the

local protected ecosystem, and A2 is the size of the larger region within which the protected ecosystem

is embedded, to maintain the local species diversity S1 the larger area needs to include total species rich-

ness of S2, derived as S2 = S1(A2/A1)z (Kinzig et al., 2002). For instance, to guarantee that the local ecosys-

tem of 10 hectares can in the long run maintain its species diversity of 100 species, S2 of an area of 100

km2 needs to be 282 species (with z = 0.15). This result emphasizes the importance of maintaining a

large enough regional species pool for maintenance of local ecosystem diversity and functions. The FCS

of particular habitat types obviously cannot be maintained in the long run without ensuring sufficient

connectivity between habitat patches of similar type. The importance of these ecological ‘stepping

stones’, especially for actively dispersing taxa such as birds and mammals, was apparently first empha-

sized by MacArthur and Wilson (1967; for insects see Wilson, 1961). Establishment of stepping stones

is mentioned in Article 10 of the Habitats Directive as a possible method of improving the ecological

coherence of the Natura 2000 network.

Potential answers to the second question may be provided by community viability analysis, a method

recently developed for identification of fragile habitat types and keystone species (Ebenman and Jonsson,

2005). The objective of community viability analysis is to predict the response of ecological communi-

ties to species loss, or more specifically to address the risk of secondary extinctions. A so-called dynamic

community viability analysis can be performed if we can find ways to estimate population growth rates

of species and the strength of interactions among species. The problem is that collection of such data

is laborious, especially as part of standard monitoring programmes. A possibility for overcoming this

problem is to estimate interaction strength parameters from the body sizes of species using allometric

scaling relationships (Ebenman and Jonsson, 2005).

Strict Requirements for Species Monitoring

In Table 2 we have listed some ecological questions related to the FCS of species. Investigating these

questions usually requires rather detailed population biological data. For instance, population viability

analyses require data on population demography and sometimes also management options (Beissinger

and McCullough, 2002). Such data is seldom available without comprehensive monitoring efforts, which

clearly form the basis of any ecologically based conservation programme. This makes selection of species

into monitoring programmes especially important. As monitoring programmes may be costly, moni-

toring efforts should obviously focus on species that are at a risk to lose the favourability of their



conservation status. This objective is partially fulfilled by listing in the annexes of the Habitats Direc-

tive the species that need particular protection efforts.

However, the concept of FCS is not only applied to the (mostly) endangered animal and plant species

listed in annexes II, IV and V. The Habitats Directive also requires that the conservation statuses of

‘typical species’ of natural habitat types of Community interest should be kept favourable (Table 1).

Although it is not made clear in the Directive, this must refer either to so-called keystone or to charis-

matic species, or both. Keystone species are those having disproportionately large effects on community

structure (Holt and Loreau, 2002), or whose removal would have strong effects on community diversity

and composition (Ebenman and Jonsson, 2005). Keystone species usually belong to some of the most

important functional groups in the community, such as pollinators, nitrogen fixers or herbivores

(Nyström, 2006). Charismatic species are well known and highly valued species such as the elephants,

large carnivores or apes, whose conservation is nowadays commonly considered important by the general

public.

Identification of keystone species, especially rare ones, may be difficult. Due to this, the overall con-

clusion thus seems to be that not only endangered species need to be monitored, but also the entire

communities represented by natural habitat types of Community interest. Without such a comprehen-

sive monitoring system some keystone species would probably remain undiscovered.

Monitoring efforts are also required below the species level. Inspired by the concept of distinct pop-

ulation segments (DPSs), introduced into the US Endangered Species Act in 1978 (Clark, 2002) and

based on extensive surveys of genetic differentiation of local populations (Cohn, 1990; Frankham et al.,2004), various concepts and methods have been developed for assessment of conservation value of units

below the species level. These include the evolutionarily significant units (ESUs: DeWeerdt, 2002) and

management units based on the exchangeability method of Crandall et al. (2000). The ultimate objec-

tive is to allocate conservation efforts to subpopulations considered as particularly important for the evo-

lutionary potential of the species. The Convention on Biological Diversity (UNEP, 1992) also requires

that diversity at the genetic level has to be included in conservation targets and monitoring programmes.

Although genetically distinct populations within species are not specifically mentioned in the Habitats

Directive, site assessment criteria listed in Annex III (Stage 1 B; Stage 2, 2b, 2e) assume that the degree

of isolation of Annex II species present in the proposed site is known. Isolated populations may often be

genetically distinct for instance due to genetic drift or inbreeding. It is not mentioned whether a high

degree of isolation should be considered a positive or negative factor from the conservation perspective.

Obviously, this would depend on genetic uniqueness of the local population. Annex III specifically

requires taking into account of the ‘global ecological value of the site for the biogeographical regions

concerned’ in Community-level assessment of national lists of sites. This can be interpreted to support

conservation of local races or subspecies of animal and plant species of Community interest.

To summarize, it seems that assessment of FCS for species requires ambitious national monitoring

programmes that include (genetic) analyses of population structures, and regular collection of popula-

tion biological data of both species listed in Annex II and keystone or otherwise ‘typical’ species of natural

habitats listed in Annex I.

FCS as a Hierarchical Concept

A great methodological challenge is that for legislative and administrative reasons FCS of habitat types

and species will be assessed over entire countries or even larger areas (the EU as a whole included),

while most biological monitoring methods have been developed either for local populations (e.g. popu-

lation viability analysis, PVA) or for individual communities or ecosystems in need of protection (bird

census methods, community viability analysis). An obvious question is how big a proportion of local



populations or ecosystems need to be viable before the conservation statuses of the corresponding

species or habitat types of Community interest can be considered as nationally favourable. Finally, could

the conservation statuses at the Community level be considered as favourable although the criteria of

the FCS would not be met in one or more of the member states?

One solution is to consider FCS as a hierarchical concept. Such an approach is, in fact, already embed-

ded in the process of site selection for the Natura 2000 network described in detail in Article 4 and

Annex III of the Habitats Directive. The first step (or Stage 1) in this hierarchical procedure is prepara-

tion by each member state of its own list of potential sites for the network. The second step (Stage 2) is

assessment of these national lists from the Community perspective. In this latter process the Commu-

nity importance of each site listed by a member state is assessed, taking into account both the relative

value of the site at national level and its global ecological value for the biogeographical region or for the

entire Community. The hierarchical procedure clearly allows for instance the possibility that a site con-

sidered as valuable at the national level may be judged less so at the level of the entire Community.

Theoretically, the conservation status of an individual habitat or population is always either favourable

or not, depending on how we define favourability. In the evolutionary timescale the fate of every

population is extinction (Endler, 1986). However, conservation policies are made within the ecological

timescale, in which extinction probabilities of populations differing in size, age structure and sex ratio

can be compared in different environments. The concept of minimum viable population size (MVP),

which refers to the number of individuals necessary to ensure the long-term survival of the species

(Shaffer, 1981), is one possible measure of favourability of conservation status at the site level. If the

local population size is equal to or exceeds the MVP, its conservation status can be considered favourable.

Analogously, the conservation status of a local habitat can be considered as favourable if the population

sizes of its typical component species exceed or are equal to their MVPs, so that the specific structure

and functions necessary for the long-term maintenance of the local habitat are likely to continue to exist

for the foreseeable future (Table 1). Obviously, this is more difficult to assess than the population sizes

of the habitat’s typical species.

At the national level the conservation status is evidently a statistical outcome of fates of individual

habitats and animal and plant populations at the local scale. Based on Annex III of the Habitats Direc-

tive (e.g. Stage 1 A), it is considered an important prerequisite for favourability of conservation status

that a representative sample of each natural habitat type or species of Community interest will be pre-

served in each member state. This goal is more ambitious than merely ensuring the long-term survival

of the habitat type or species in the country. For the latter goal even a single large enough sample of a

habitat type or a large individual population of a species might be enough. The criterion of representa-

tivity requires taking into account the geographic variability of each natural habitat type and animal and

plant species of Community interest. Due to the requirement of representativity, it is not possible to

present any ‘rule of thumb’ on how large a proportion of the total area of a particular natural habitat

type of Community interest should be protected in a single country. This obviously depends on the geo-

graphic variability of the habitat type in question. If the degree of variability is high, a larger proportion

needs to be protected than in a country where the sites representing this habitat type are rather uniform.

Implications for Current European Union Conservation Policies

In this paper we have listed a number of problems and challenges related to the concept of FCS, pri-

marily from the biological perspective. The problems include those related to the hierarchical structure

of the concept, lack of historical data, the diverse requirements for species monitoring, and problems

of identification of habitat types and their specific structures and functions. In spite of these problems



we believe that FCS has potential to serve as a unifying, heuristic target for international conservation

efforts, especially within the EU, and could serve as a link urgently needed between conservation theory

and practice. The conceptual links to a safe minimum standard approach of natural resource econom-

ics, shared by the US Endangered Species Act and EC Habitats Directive, could also enable sharing of

experience in practical implementation of conservation targets between these areas.

Our analysis indicates that conservation objectives of the Habitats Directive require planning, estab-

lishment and execution of rather ambitious monitoring programmes of natural habitats and species of

Community interest. The need for periodical reviews of the success of the Natura 2000 network is gen-

erally acknowledged in Articles 9 and 11 of the Habitats Directive, but without any specifications. An

important conclusion is that the criteria of FCS listed in Article 1(c) and 1(i) of the Habitats Directive,

and summarized in Table 1, in fact require not only monitoring of mere presence of species or habitat

types of community interest in a certain area, but also regular monitoring of populations, species inter-

actions and other ecological characteristics of the most important ecological communities. As this cer-

tainly requires professional ecological expertise, we consider it of great importance that conservation

biologists actively participate in planning of ecologically based guidelines for assessment of FCS. After

all, the concept will in any case be widely applied in decision making related to conservation measures

at least in the European context. Involvement in the required monitoring programmes for biological

diversity apparently also means many job opportunities for conservation biologists.

Addressing all these challenges will require active involvement of policy makers, conservation admin-

istration and the scientific community. On the basis of this paper, developing a national monitoring

programme should include at least the following steps.

(1) Listing of species and habitat types to be protected. It is important to notice that within the Euro-

pean Union ‘typical species’ of habitat types of Community interest also need to be protected and

their FCSs estimated.

(2) Establishment of geographically representative and methodologically unified monitoring networks

for species and habitat types. Partially separate networks may be necessary, because the ranges of

protected species and habitat types do not necessarily overlap, and because collection of species

monitoring data should not be confined to particular habitat types only.

(3) For each protected species of Community interest, national programmes should be established for

mapping of genetic structure of subpopulations. The objective would be to discover ecologically or

genetically distinct units that most urgently need protection.

(4) Species monitoring networks should collect accurate data on population sizes and demographic

properties, and other data needed for appropriate PVAs, performed periodically.

(5) Habitat monitoring networks should be based on a priori mapping of representative sites, and

include monitoring of specific community structures and functions, and conservation statuses of

keystone or charismatic species. These need to be identified for each habitat type and geographi-

cal region separately.

It is equally important to decide what to do with the collected data. The primary concern will be assess-

ment of conservation statuses of species and habitat types, and development of special conservation pro-

grammes for those species or habitat types whose conservation statuses are not regionally favourable.

Another possibility is to develop or use indices such as the Biodiversity Intactness Index as overall esti-

mates of pursued conservation policies in particular administrative areas. The Biodiversity Intactness

Index (BII) basically combines species and habitat data, and can be compared directly within and across

scales (Scholes and Biggs, 2005). The monitoring data collected according to the Habitats Directive

requirements can be used to calculate BIIs for various geographical scales. Such indices can mainly



serve as overall estimates of success of conservation policies, but for obvious reasons cannot replace

species or habitat specific analyses in conservation planning.

The Habitats Directive in its present form is rather expert oriented, and largely excludes economic

considerations or public hearings from the site selection process to the Natura 2000 network (Annex

III). This can be considered a problem, since it is generally observed that active participation of all stake-

holders in conservation planning results in a more effective policy outcome. In Finland the neglect of

public opinion resulted in widespread criticism of the entire network and the EC conservation policy as

a whole in the late 1990s (Oksanen, 2003). It should also be noted that, in the preamble of the Habi-

tats Directive, taking into account economic, social, cultural and regional requirements in maintenance

of biodiversity is mentioned as an objective. In its present form the directive partially fails to fulfil this

objective, in spite of inclusion of these requirements in Article 6. The hierarchical structure of the con-

servation status assessment process in fact provides possibilities for increasing the role of public hear-

ings. Planning and management necessarily take place primarily at the local or regional level. The

authorities at these levels are probably familiar with natural conditions and conservation issues, but may

fail to observe for instance the small-scale recreational activities that in many cases could be taken into

account in planning and execution of the Natura 2000 network without seriously compromising the

most important conservation targets. We believe that a more active role of public opinion in the plan-

ning procedure would in the long run greatly benefit European conservation policy by creating more

positive attitudes toward conservation issues.

Whatever the exact targets of biodiversity conservation, it is of central importance that the progress

towards these goals is reliably monitored. This increases the potential significance of unifying concepts

such as the FCS. Obviously, the concept of FCS needs to be developed to incorporate discoveries of

modern conservation science. For instance, in its present form it does not explicitly include the genet-

ically distinct subpopulations of endangered species. However, the fairly flexible definition of the FCS

makes inclusion of such novel aspects quite possible.

Acknowledgements

Comments of Michael B. Usher and three anonymous referees were of great help in improving the

manuscript. The study was financially supported by the Maj and Tor Nessling Foundation.

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