Marine Pollution Bulletin 53 (2006) 2029 www.elsevier.

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Review

The concept of biotope in marine ecology and coastal management

Sergej Olenin
b

a,*

, Jean-Paul Ducrotoy

a Coastal Research and Planning Institute, Klaipeda University, H. Manto 84, 92294, Klaipeda, Lithuania Groupe dEtude des Milieux Estuariens et Littoraux, 115 quai Jeanne dArc, 80230 Saint Valery sur Somme, France

Abstract The term biotope was introduced by a German scientist, Dahl in 1908 as an addition to the concept of biocenosis earlier formulated by Mo bius (1877). Initially it determined the physicalchemical conditions of existence of a biocenosis (the biotope of a biocenosis). Further, both biotope and biocenosis were respectively considered as abiotic and biotic parts of an ecosystem. This notion (ecosystem = biotope + biocenosis) became accepted in German, French, Russian and other European continental ecological literature. The new interpretation of the term (biotope = habitat + community) appeared in the United Kingdom in the early 1990s while classifying marine habitats of the coastal zone. Since then, this meaning was also used in international European environmental documents. This paper examines the evolution of the biotope notion. It is concluded that the contemporary concept is robust and may be used not only for the classication and mapping but also for functional marine ecology and coastal zone management. 2006 Elsevier Ltd. All rights reserved.
Keywords: Biotope; Biocenosis; Ecosystem; Underwater landscape; Ecological terminology; Coastal management

1. Introduction Scientic terms have a life of themselves as they appear, develop, and change content according to emerging scientic paradigms. Sometimes, an old term is re-found again and is given a new meaning. That was the case with the term biotope, which recently entered into the lexicon of national environmental planning literature (i.e., Riecke et al., 1994; Connor, 1995) and international environmental documents (cf. HELCOM, 1998; EUNIS, 2005). Hierarchical levels of biological organization (including the biotope level) are widely used by scientists but also by decision-makers and managers. A recent denition of a biotope was used in the framework of the European programme Biomar-Life: it combines the concepts of habitat and community for dening geographical units (Connor, 1995; Connor et al., 2004). However, the word habitat may be used in various ways: the place where an organism is found (e.g., a sub-tidal sandbank); the area where a

species occurs, or the type of environment where a species could potentially establish itself. The notion of community also varies greatly depending on the authors: from a biological entity consisting of interdependent organisms to a statistically dened assemblage of occasionally cooccurring species (for comprehensive discussion see, e.g., Thorson, 1957a,b; Mills, 1969). Ambiguity has therefore arisen in the use of the biotope concept, in particular in the theoretical eld. This paper aims to outline the possible uses of the word biotope and related ecological concepts. Summarizing the present knowledge on marine biotopes should enable ecologists to agree on a proper use of the concept, remove ambiguity and attribute the same meaning to the word amongst the various paradigms used in functional ecology and coastal management. 2. Origin and evolution of the concept 2.1. Biocenosis, biotope and ecosystem

Corresponding author. Fax: +370 46 398845. E-mail address: serg@gmf.ku.lt (S. Olenin).

In 1877, Mo bius (in Keller and Golley, 2000) was requested by sheries managers to examine an oyster bank,

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which failed to produce as much as expected. Writing that the oyster bank was a Bioco nose, or a Social Community, he founded the basis of ecology. He dened the biocenosis as a complex superorganism where animals and plants live together in an interdependent biological community. Even if later it was shown that the biocenosis concept could not be inferred from the Mo bius oyster research (Reise, 1990), it is of note that, in this seminal piece of work, applied ecology was the answer to management and that managers were targeted. Two decades later, Dahl (1908), a colleague of Mo bius, coined the new term (biotope) to dene a complex of factors, which determines physical conditions of existence of a biocenosis. The biotope was related to biocenosis as the biotope of a biocenosis. Later on, Tansley (1935) (in Keller and Golley, 2000) produced the rst denition of ecosystem. Followers introduced complementary notions describing the physical conditions and groups of plants and animals living there and nally suggested that the ecosystem was made up from the biotope (the abiotic environment) and the biocenosis (the biotic communities): Biotope + Biocenosis = Ecosystem (cf. Ramade, 1978; Voronov et al., 2002). 2.2. Semantic eld of related concepts Evolution of the biotope concept can be better dened in the context of relative or accompanying ideas. The important issue is the spatial scale, since by denition, the ecosystem could be of any size (Pickett and Cadenasso, 2002). Consequently, both biocenosis and biotope are also scale independent in the context of the ecosystem concept. A major step towards the synthesis of ecological and geographical approaches was the formulation of the biogeocenosis concept by V. Sukachev in 1942 (Novikov, 1980), as an evolutionary natural phenomenon, located on a relatively limited area, homogeneous inside itself, functionally correlating living organisms and their environment, with a specic type of interactions of its components and a certain type of matter and energy exchange between themselves and with other natural phenomena (Reimers, 1990). An ecosystem may be scale independent, while a biogeocenosis is always attached to a certain space, i.e., it always has a physical location. Since the 1950s, the biogeocenotical (and biotopical as a part of it) approach became popular both in marine and land-based research (Da Silva, 1979; Duvigneaud, 1984; Margalef, 1986). The principles of marine biogeocenology were established (Turpaeva, 1954; Sokolova, 1960; Beklemishev, 1961, 1973; Vinogradov, 1977), and the biotopical methodology was applied to the study of marine biological resources (Moiseev, 1986). Other authors suggested approaching the investigation of marine coastal ecosystems through a structuralfunctional analysis based on biotopes or comparable spatial units (Reise, 1985; Kuznetsov, 1980; Burkovsky and Stoliarov, 2002).

Dierent derivatives of the word landscape (seascape, benthoscape, benthic landscape, sea oor landscape) are often used in underwater studies (Zajac, 1999). However, underwater landscape researchers argue that biotope is a biocentric notion and its usage is appropriate mostly in biological and ecological studies (Arzamascev and Preobrazhensky, 1990). Preobrazhensky et al. (2000) made a comparative analysis of terrestrial and underwater landscape terminology and emphasized that the direct transfer of well-developed terrestrial concepts into the marine realm is not well grounded in most cases due to fundamental dierences in the nature of terrestrial landscapes and underwater complexes. They also compared and evaluated the suitability of other terms for underwater landscape research, such as landscape, facies, biocenosis, biogeocenosis and ecosystem (Arzamascev and Preobrazhensky, 1990). Concluding that none is fully appropriate for underwater classication, these authors proposed a new term, benthem, as a derivate of benthal and system. marec, 1997; OleIn many papers (cf. Reise, 1985; Gle nin, 1997; Ducrotoy, 1999), terms such as substrate or substratum referred to the purely physical bed characteristics of a particular ecological unit, whereas community designated the associated organisms living there. This close linkage between benthic life and the bottom environment, expressed as conformity between types of geographic underwater complexes and characteristic benthic biotic groups was widely used in underwater landscape research (Petrov, 1999; Arzamascev and Preobrazhensky, 1990; Preobrazhensky et al., 2000). To determine the regularly repeating underwater landscape complexes (Petrov, 1999) the term facies (or facium) was used. Originally, this term was suggested by Gressley in 1836 (Arzamascev and Preobrazhensky, 1990) and initially was used by paleo-sedimentologists, who recognized that sediments with certain characteristics would contain typical fossils. These strata would be organized into a sequence reecting changes in climatic and oceanic conditions through time and space. The rst mention of facies in a marine science paper dates from 1913, by Zernov. His denition of the facies related to an area of the sea oor, homogeneous by its natural conditions and occupied by a characteristic community of marine organisms. Presently, there are more than 200 various denitions of the notion of facia (Nesis, 1980). In 1989, Ducrotoy et al., working on megatidal estuaries, referred to the bio-sedimentary facies (or bio-facies in short) as an entity combining characteristics of the sediment (the substratum) and of the biota, leading to a valid typological classication. 2.3. Contemporary meaning and synthesis Up to the early 1990s, the notion biotope in the English literature was not applied widely. The term was rediscovered when the UK Joint Nature Conservation

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Committee, working on a classication of the coastal marine environment, produced a new denition of the biotope (Connor, 1995; Hiscock, 1995): Biotope = habitat + community, broader than under its former accepted denition where the biotope was considered as the physical part of the ecosystem. The new biotope concept combines the physical environment (habitat) and its distinctive assemblage of conspicuous species. The habitat was dened according to geographical location, physiographic features and the physical and chemical environment (including salinity, wave exposure, strength of tidal streams, etc.), while the community was described as a group of organisms occurring in a particular environment, presumably interacting with each other and with the environment, and identiable by means of ecological survey from other groups (Hiscock and Tyler-Walters, 2003). The community was interpreted as a biotic element of a biotope. The new meaning of the word biotope should be distinguished from the ecosystem denition, which also includes both the physical environment and community (e.g., Odum, 1975; Ramade, 1978). Connor (1995) refers to the use of wave action and tidal current in his denition of biotope/habitat whereas the earlier denition of ecosystem does take into account this energy aspect. However, strictly speaking (according to its original denition), the new concept of biotope does not take directly into consideration the energy and other ecosystem linkages between its abiotic and biotic components. The community (particularly one of its partsthe complex of the most distinctive, conspicuous species) is mentioned only as one of the distinctive characteristics, which enables one to distinguish and classify the biotopes (Olenin et al., 1996). Thus, the new interpretation of biotope diers essentially from the traditional one because it combines both habitat and community, whereas the original word biotope (sensu Dahl, 1908) indicated only a physical habitat. Moreover, nowadays, for practical reasons of interpretation of terms used in directives, statutes and conventions, in some documents, biotope is sometimes synonymized with habitat (Connor et al., 2004; Hiscock and TylerWalters, 2003). It is argued here that the best word to t the underlying concept would have been bio-facies but, since the 1990s, not only scientists but also policy-makers and managers have named it biotope. The new understanding of biotope now dominates in the international scientic and applied environmental literature (CORINE, 1991; HELCOM, 1998; EUNIS, 2005; Connor et al., 2004). For instance, the Internet search engine for the scientic literature SCIRUS (www.scirus.com) gives more than 1700 links to journal articles, in which the biotope term is used in the elds of biodiversity, benthic research, agro-landscape, agriculture, landscape ecology and others. According to the short descriptions in the articles, it can be concluded that the new interpretation of the biotope is used in most of them.

This is why, further in this paper, the word biotope will be used in its most recent meaning. 3. Nature and nomenclature of the biotopes 3.1. Physical characteristics At a given scale, a habitat encompasses a spatial domain, relatively homogeneous with regard to environmental parameters. The environments physical and chemical characteristics are taken to encompass the substratum and the particular local conditions. The substratum of a benthic habitat would be rock or sediment while that of a pelagic habitat would be a water mass, e.g., a permanent thermocline or just a lens of brackish water drifting out from an estuary to the sea. McCoy and Bell (1991) identied three structural parameters in relation to the ecological signicance of a particular habitat: its heterogeneity, its complexity, and the scale at which the habitat is dened. Heterogeneity refers to the relative abundance (per unit volume or area) of the various structural components and their variability (e.g., SD according to the mean) while complexity deals with the absolute abundance of the various structural components. The scale relates to the unit volume or area used to measure heterogeneity and complexity, including macrofeatures and microfeatures sometimes called microhabitats (Le Hir, 2002; Le Hir et al., 2003). Hence the biotope may be from the size of underneath a boulder or a rockpool, up to the size of a very large mussel bed or sub-tidal sand bank. Consequently, the scale of the biotope will depend on the size of the habitat supporting the dominant biota or on the functional unit (see below). The physical and chemical conditions vary within a range, which is characteristic of the habitat. This means that a habitat is limited in space. Due to a problem of continua occurring with physico-chemical variables, the heterogeneity (or its opposite, homogeneity) of the habitat would always be relative, depending on the aims of a study: coastal zone mapping or meiobenthic community research, evaluation of biological resources or study of benthicpelagic interactions. The biotope integrates the environmental factors which structure the habitat, and is indicated by a limited set of words summarizing the local conditions, i.e., littoral muddy sand or sub-littoral rock platform. Such expressions integrate the various parameters which play a role in the habitat of a particular community. 3.2. Biological features From a biological point of view, the biotope results from a balance between the regional pool of species and the local environmental conditions. The species composition will be dependent on their access to the habitat and on other biological requirements, such as recruitment of young stages, trophic relations and food availability.

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Emphasizing the relation between physical conditions and living organisms Beklemishev (1961) postulated fundamental principles of biotopes homology (Text Box 1). It is interesting to note that the Beklemishevs biotope homology concept, based on the study of large oceanic pelagic systems, has many common features with the parallel bottom level communities theory of Thorson (1957a,b). The latter was built on comparative analysis of marine benthic macrofauna of the European, North American and Greenland shelves.

4. Biotope approach to marine studies and management 4.1. Coastal typology The management of coasts and natural resources rely on the ease to map geographical units. In the late 19801990s, with many European Directives being promulgated, law has become a driving force for ecology (Ducrotoy and Elliott, 1997; Elliott et al., 1999). Classications were the most widely used aspects of biotopes. The EU CORINE biotope classication was developed in the 1980s and used to derive the habitats, meeting the requirements of the EU Habitats Directive (EU, 1992). Because of signicant shortcomings in its structure, the CORINE classication remains very broad and alternatives were proposed. The marine biotope classication was published by the Joint Nature Conservation Committee (JNCC) in the United Kingdom (Connor et al., 2004). In France, the Zones Nationales re t Scientique, Faunistique et Floristique (ZNIEFF) dInte classication was developed (Dauvin et al., 1996); similar activity took place in Lithuania (Olenin et al., 1996). A regional international classication of coastal biotopes and their complexes was developed for the Baltic Sea (HELCOM, 1998). With the establishment of the European Environment Agency, a rationalised and restructured classication is being proposed: European Nature Information System (EUNIS) used in coastal zone planning and management (EUNIS, 2005). Recently, the notion of biotope was suggested by Olenin and Daunys (2004) for the development of a coastal typology meeting further the requirements from the EU Water Framework Directive (EU, 2000) (Fig. 1). The WFD typology works at landscape level, then at a broader scale than biotopes. In a regional case study, the coastal types were eectively dened as complexes of biotopes (Olenin and Daunys, 2004). It was noted that the biotope notion integrates several, if not all, obligatory and optional factors (the tidal range, salinity, depth, current velocity, wave exposure, turbidity, etc.) listed in the WFD requirements for coastal typology (Guidance document, 2003). Furthermore, the biotope classication procedure requires the analysis of matching between physical and biological features used to characterize the biotopes. The next step, following the creation of the biotope classication system and its use for coastal mapping, includes identication of coastal types as the complexes of interrelated neighboring biotopes. This step gives the coastal typology a robust scientic background and provides it with essential ecological relevance. The major argument for the use of biotopes for the coastal typology is that there are already several national and international biotope classication systems developed for the coastal zones of Europe (see above). 4.2. Monitoring and surveillance The application of an international classication system should oer an opportunity for monitoring changes in

Text Box 1. Beklemishevs (1961, 1973) postulates of biotope homology: (1) Homological biotopes are formed under the inuence of similar physical factors and occupy similar place among other biotopes. (2) Principles used to identify the homology of plant and animal organs are applicable for dening homologies of biotopes. (3) Marine biological structure is a function of its biotopic structure (the ocean physical organization). (4) The concept of biotope homology is applicable to explain and predict species distributions.

In the new accepted use of the term biotope, the community is the second strong element. From the discussion above, it is possible to give a denition of a community: it is a species assemblage occupying a well-dened physical structurethe habitat. However, it would seem dicult to integrate the complete composition of the community into the naming of a biotope. Traditionally the bottom communities have been designated by names of the dominant species, for example, the Macoma community in coastal areas of Denmark (Petersen, 1914; Thorson, 1957a,b). The same approach was proposed for nomenclature of estuarine bio-sedimentary facies (Ducrotoy et al., 1989). In contemporary classications, benthic biotopes are identied by brief descriptions of the physical environment and the Latin name of the conspicuous and/or dominant species (Dauvin et al., 1996; Olenin, 1997; Connor et al., 2004). Not only living organisms themselves can be considered as biological features, but these include also the signs of their presence and activity (empty shells, sandy refuges, borrows, traces, faecal pellets, etc.). These give indications of the physico-chemical qualities of the substratum and how the vital activities of the bottom fauna aect them (cf. McCall and Teversz, 1982; Bromley, 1996). Consequently, the qualities of biotopes themselves depend on correlations between biological and physico-chemical processes. That is why the application of further biotic features in classication of biotopes is not only useful but also necessary from a methodical point of view.

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DEVELOPMENT OF A COASTAL TYPOLOGY

Identification of complexes of neighboring interrelated biotopes the coastal types

Identification, mapping and description of biotopes

Development of a biotope classification

Justification of ecological relevance by the analysis of matching between physical and biological features

Inventory of physical factors shaping benthic environment

Salinity Depth Wave exposure Substratum Shape, bottom relief Water temperature Turbidity Others

Inventory of biological features characterizing biotopes

Characteristic species Coverage and dominant forms of macrophytes and/or macrofauna Visible biogenic signs (empty shells, traces of crawling, etc.) Community structure

Fig. 1. Scheme showing the benthic biotope classication procedure and its relevance to the coastal typology (modied from Olenin and Daunys, 2004). Explanation in text.

marine ecosystems. Following the 1992 Rio UNCED Earth Summit, the European Community passed the Habitats Directive (1992/EC192) (Bell, 1997) which places a requirement on member states to designate Special Areas of Conservation (SACs). The establishment of the Natura 2000 network is an integrated approach to the designation of protected habitats to represent Europes environmental diversity, including SACs but also Special Protection Areas (SPAs) designated under the Birds Directive. The philosophy adopted here is that if the habitat is protected, then the health of the biota will also be safeguarded. Once an area is assigned SAC status, the Habitats Directive (Article 17) requires that the member state government reports at regular 6-year intervals on the conservation status of the habitats and of the species for which the site is designated (Ducrotoy, 1999). The information provided includes a broad scale assessment of the complete range of habitats and their associated communities and whereas they meet conservation objectives for the site. The biotope concept could be used in managing marine SACs, but it would then be necessary to further rene existing classications to

ensure they are suciently accurate for monitoring changes in the long term. Operational monitoring and surveillance provide tools for understanding and assessing natural and humaninduced disturbances and produces scientic information for environmental management, as well as protection and conservation. They require the collection of quantitative data over more or less long periods of time (Ducrotoy and Sylvand, 1997). Surveillance implies measurements using validated methods and selected stations, distributed throughout the ecosystem, and are normally repeated through time (Ducrotoy et al., 1989). This demands that large geographical areas are surveyed at a broad scale in a short period of time, so methods for rapid sampling that produce sets of permanent baseline data are required. This is exactly the case when mapping biotopes (Davies and Foster-Smith, 1995). However, biotope classications were not devised as a monitoring tool per se but were designated to promote consistent interpretation of data. Assuming that the biotope classication is ecologically sound, complementary methodologies will need to be introduced at a

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suitable scale if certain designated sites should be surveyed using such classications. 4.3. The use of biotopes in functional coastal ecology Estuarine and coastal ecosystems can be considered as inherently variable. The steady state is a dynamical system which changes and varies in a stochastic way; therefore the knowledge of key abiotic environmental conditions is a

prerequisite for understanding the functional properties of the system under consideration. Many studies (cf. Reise, 1985; Brown and McLachlan, 1990; Bek, 1997; Burkovsky and Stoliarov, 2002) have shown peculiarities of coastal ecosystems which make them fundamentally dierent from terrestrial ecosystems. Since coastal marine ecosystems consist of two interacting subsystems (benthic and pelagic), most of primary production is not used where it is produced, but rather it is transported

Fig. 2. (Above) Scheme showing distribution of benthic biotopes at the exposed coast of the Baltic Sea (based on biotope mapping results in the Seaside Regional Park, Lithuania, unpublished): (1) shallow hollows between the shoreline and the rst sand bar with decomposing algal mats (in summer time); (2) mobile sand in the upper part of the slope with amphipods and mysids; (3) boulders in the swash zone with green algae; (4) stony bottoms with red algae Furcellaria lumbricalis; (5) boulder reefs with dense blue mussel colonies; (6) gravel and pebble bottoms with rare macrofauna and no macroalgae; (7) sandy bottoms with bivalve Macoma balthica and polychaete Pygospio elegans. (Below) Provisional scheme showing the functional role of and interrelations between biotopes at the exposed coast of the Baltic Sea.

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elsewhere. Often production of the benthic macroalgae rst is transformed into detritus and only then is consumed by macrofauna. The coastal zone, as a transitional system, depends on many external factors: trophodynamic processes in the open sea, deposition of spat of benthic animals from plankton, river discharge, transportation of bottom sediments, etc. In most cases, only part of a biogeochemical or reproductive cycle takes place within the coastal marine ecosystem, while the whole cycle involves a much larger ecosystem or, even goes beyond the limits of the hydrosphere (Ducrotoy and Olenin, 2003). These peculiarities of coastal ecosystems should be taken into account when studying their functional organisation so that, in our opinion, the use of the biotope approach may give insight into spatial as well as functional structure of the coastal zone. Physico-chemical peculiarities of a coastal habitat determine the diversity of species, as well as the functional diversity, allowing the presence of certain functional groups and restricting the existence of others. Hence, a biotope may be viewed not only as a structural unit convenient for mapping of a coastal zone but also as the site with its own processes. These processes will change according to the biotope. For example, along the very exposed Baltic Sea coast, active biosedimentation is possible only on large boulders covered by attached colonies of blue mussels below the breaker zone; production of macroalgae takes place only within the euphotic zone on large stones; herring spawning occurs on stony bottoms with macroalgae (Fig. 2, Table 1). Reise (1985) dened trophic types of North Sea coastal tidal ats according to their role in primary production, decomposition of detritus and consumption of biomass. Bek (1997) used the biotope approach to study peculiarities of distribution and life cycles of macrobenthos at the White Sea littoral. Once their biological characteristics have been taken into account, biotopes dier not only in their appearance (exterior) but also in their functions, which they perform in coastal marine ecosystems: production, storage and distribution of organic material; reproduction of biological resources; modication of bottom sediments, etc. Thus, in their extended denition, biotopes can be considered as functional units of a coastal marine ecosystem.

Complementary approaches to the use of biotope methodology are necessary. The classication of benthic animals and macroalgae into functional groups (cf. Padilla and Allen, 2000; Pearson, 2001) oers interesting perspectives. When establishing and recognising functional or morphological groups, relatively few species attributes are of importance in determining the structure of communities. For example, the categorisation of algal species simply by body plan can give substantial insight into the community structure (Tobin et al., 1998). Attributes used to identify the groups are often shared between taxonomically distant species (Steneck and Dethier, 1994). This eliminates the noise at the species level to give a more continuous description. Similar conclusions were reached by biologists working on guilds of invertebrates (Hily and Bouteille, 1999) and shes (Elliott and Dewailly, 1995). Thus, the biotope concept can be adapted to t in a functional approach to the ecology of coastal marine ecosystems (Text Box 2). Text Box 2. Functional aspects of biotope research in coastal marine ecology: biotopes as components of the ecosystem and structuring aspects of dominant organisms, the spatial scale of biotopes in relation to their physical boundaries and their individual characteristics, the temporal scale relating to the changes in the distribution of biotopes within the ecosystem over time, connections between biotopes within the ecosystem demonstrating processes and functions, constraints (natural or anthropogenic disturbances) on the ecosystem behaviour and how biotopes translate such changes.

5. Conclusions and perspectives The language of science is a living entity where the meaning of words (especially in English) changes with

Table 1 Potential role of coastal benthic biotopes in maintenance of sh resources at the exposed coast of Lithuania, Baltic Sea Biotope 1. Shallow hollows between the shoreline and the rst sand bar with decomposing algal mats (in summer time) 2. Mobile sand in the upper part of the slope with amphipods and mysids 3. Boulders in the swash zone with green algae 4. Stony bottoms with red algae Furcellaria lumbricalis 5. Boulder reefs with dense blue mussel colonies 6. Gravel and pebble bottoms with rare macrofauna and no macroalgae 7. Sandy bottoms with bivalve Macoma balthica and polychaete Pygospio elegans Spawning ground 0 0 0 ++ + + 0 Fish fry shelter 0 + + ++ + 0 0 Fish fry nursery ++ ++ 0 ++ 0 0 0 Adult sh feeding ground 0 + 0 + ++ 0 ++

++ very important, + little important, 0 unimportant (based on Olenin and Daunys, in preparation).

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usage. Nonetheless, as with all sciences, ecology needs precision in its work and the terminology needs to support theory. Unfortunately, this is not the case with the concept of the biotope, the meaning of which has evolved in several directions during the last 20 years. However, the concept, as it is used in the early 21st century in coastal marine ecology, has a heuristic value and a biotope is now recognised as a fundamental organizational unit of coastal ecosystems (cf. Reise, 1985; Ducrotoy et al., 1989; Connor, 1995; His marec, 1997; Bek, 1997; Burkovsky and cock, 1995; Gle Stoliarov, 2002; Ducrotoy and Olenin, 2003). The concept of the ecosystem remains more dicult to use as, very often, its properties and boundaries are abstract. The ecosystem conceived as a network of its biotopes is easier to circumscribe because the individual biotopes provide an adequate scale for the study of the ecosystem properties, in space and time. They also t into the intermediate scale concept of landscapes. Changes in numbers of populations and processes linking the physical and the biotic components are approachable through the use of pilot-stations at biotope level. Biotopes help to reconcile the divisive controversy between the population-community view (networks of interacting populations) and the process-function approach (biotic and abiotic components). In particular, the use of functional groups may help to further divide ecosystems and help to assess dynamics at complementary levels. The concept of the biotope links with other levels of biodiversity in the ecosystem and integrates its various functions. However, further research needed at biotope and lower hierarchical levels includes the modelling of relationships between biotopes in relation to the overall behaviour of the ecosystem. The quantication of uxes between various compartments, at biotope level and lower, is another avenue to explore uses of photography and geographical information systems. Applications to management could lead to interesting socio-economic considerations such as the sustainable exploitation of natural resources or the search for new sheries. Acknowledgements The EU Concerted Action BIOMARE (Implementation and Networking of large-scale long-term Marine Biodiversity research in Europe) provided an opportunity for starting a discussion and a reection on biotopes and their use in functional ecology. This work was also supported by the EU Projects CHARM (Characterisation of the Baltic Sea Ecosystem: Dynamics and Function of Coastal Types) and ELME (European Lifestyles and Marine Ecosystems). Authors are indebted to colleagues of GEMEL (Groupe dEtude des Milieux Estuariens et Littoraux) and CORPI (Coastal Research and Planning Institute, Klaipeda University) who provided support to work in North Western French estuaries and in the Lithuanian coastal zone. The authors gratefully acknowledge the valuable discussions with Karsten Reise (AWI, Island of Sylt, Germany), Jan

Marcin Weslawski (Institute of Oceanology, Sopot, Poland) and Chingiz Nigmatullin (AtlantNIRO, Kaliningrad, Russia). Special thanks go to Mike Elliott (IECS, Hull University, UK) and David Connor (UK Joint Nature Conservation Committee) who suggested important changes in the paper and made the nal language check. References
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