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Review
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 Mobius (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
0025-326X/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.marpolbul.2006.01.003
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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.
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.
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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.
26
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.
Table 1
Potential role of coastal benthic biotopes in maintenance of sh resources at the exposed coast of Lithuania, Baltic Sea
Biotope
Spawning ground
++
++
0
++
+
+
+
++
+
0
0
++
0
0
0
+
++
0
++
++ very important, + little important, 0 unimportant (based on Olenin and Daunys, in preparation).
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; Hiscock, 1995; Glemarec, 1997; Bek, 1997; Burkovsky and
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
27
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.
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