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Marine Pollution Bulletin 53 (2006) 20–29
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The concept of biotope in marine ecology and coastal management
Sergej Olenin
b
a,*
, Jean-Paul Ducrotoy
b
a
Coastal Research and Planning Institute, Klaipeda University, H. Manto 84, 92294, Klaipeda, Lithuania
Groupe d’Etude des Milieux Estuariens et Littoraux, 115 quai Jeanne d’Arc, 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 Möbius (1877). Initially it determined the physical–chemical 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 classification 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
Scientific terms have a life of themselves as they appear,
develop, and change content according to emerging scientific 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 definition of a
biotope was used in the framework of the European programme Biomar-Life: it combines the concepts of habitat
and community for defining 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 defined 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 field. 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: [email protected] (S. Olenin).
0025-326X/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.marpolbul.2006.01.003
In 1877, Möbius (in Keller and Golley, 2000) was
requested by fisheries managers to examine an oyster bank,
S. Olenin, J.-P. Ducrotoy / Marine Pollution Bulletin 53 (2006) 20–29
which failed to produce as much as expected. Writing that
the oyster bank was a ‘‘Biocönose’’, or a ‘‘Social Community’’, he founded the basis of ecology. He defined 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 Möbius’ 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 Möbius,
coined the new term (‘‘biotope’’) to define 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 first definition of
ecosystem. Followers introduced complementary notions
describing the physical conditions and groups of plants
and animals living there and finally 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 field of related concepts
Evolution of the biotope concept can be better defined
in the context of relative or accompanying ideas. The
important issue is the spatial scale, since by definition,
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 specific 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 structural–functional analysis based on biotopes
or comparable spatial units (Reise, 1985; Kuznetsov, 1980;
Burkovsky and Stoliarov, 2002).
21
Different derivatives of the word landscape (seascape,
benthoscape, benthic landscape, sea floor 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 differences 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 classification, these authors proposed a
new term, ‘‘benthem’’, as a derivate of ‘‘benthal’’ and
‘‘system’’.
In many papers (cf. Reise, 1985; Glémarec, 1997; Olenin, 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 reflecting changes in climatic and oceanic
conditions through time and space. The first mention of
‘‘facies’’ in a marine science paper dates from 1913, by Zernov. His definition of the facies related to ‘‘an area of the
sea floor, homogeneous by its natural conditions and occupied by a characteristic community of marine organisms’’.
Presently, there are more than 200 various definitions 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
classification.
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
22
S. Olenin, J.-P. Ducrotoy / Marine Pollution Bulletin 53 (2006) 20–29
Committee, working on a classification of the coastal marine environment, produced a new definition of the biotope
(Connor, 1995; Hiscock, 1995): ‘‘Biotope = habitat + community’’, broader than under its former accepted definition
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 defined 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 identifiable
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 definition, 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 definition of biotope/habitat whereas the earlier definition of
ecosystem does take into account this energy aspect. However, strictly speaking (according to its original definition),
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 parts—the 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 differs 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 fit
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 scientific and applied environmental literature (CORINE, 1991; HELCOM, 1998; EUNIS, 2005;
Connor et al., 2004). For instance, the Internet search
engine for the scientific literature SCIRUS (www.scirus.com) gives more than 1700 links to journal articles, in
which the ‘‘biotope’’ term is used in the fields 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 environment’s 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) identified three structural
parameters in relation to the ecological significance of a
particular habitat: its heterogeneity, its complexity, and
the scale at which the habitat is defined. 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 benthic–pelagic 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.
S. Olenin, J.-P. Ducrotoy / Marine Pollution Bulletin 53 (2006) 20–29
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 Beklemishev’s 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.
Text Box 1. Beklemishev’s (1961, 1973) postulates of
biotope homology:
(1) Homological biotopes are formed under the
influence 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 defining
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 definition of a community:
it is a species assemblage occupying a well-defined physical
structure—the habitat. However, it would seem difficult 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 classifications, benthic biotopes are identified 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 affect 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 classification of biotopes is not only useful but also necessary from a methodical point of view.
23
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 1980–1990s,
with many European Directives being promulgated, law
has become a driving force for ecology (Ducrotoy and Elliott, 1997; Elliott et al., 1999). Classifications were the most
widely used aspects of biotopes. The EU CORINE biotope
classification was developed in the 1980s and used to derive
the ‘‘habitats’’, meeting the requirements of the EU Habitats Directive (EU, 1992). Because of significant shortcomings in its structure, the CORINE classification remains
very broad and alternatives were proposed. The marine
biotope classification was published by the Joint Nature
Conservation Committee (JNCC) in the United Kingdom
(Connor et al., 2004). In France, the Zones Nationales
d’Intérêt Scientifique, Faunistique et Floristique (ZNIEFF)
classification was developed (Dauvin et al., 1996); similar
activity took place in Lithuania (Olenin et al., 1996). A
regional international classification 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
classification 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
effectively defined 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 classification 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 classification system
and its use for coastal mapping, includes identification of
coastal types as the complexes of interrelated neighboring
biotopes. This step gives the coastal typology a robust scientific 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 classification systems
developed for the coastal zones of Europe (see above).
4.2. Monitoring and surveillance
The application of an international classification system
should offer an opportunity for monitoring changes in
24
S. Olenin, J.-P. Ducrotoy / Marine Pollution Bulletin 53 (2006) 20–29
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 classification procedure and its relevance to the coastal typology (modified 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 Europe’s 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 refine existing classifications to
ensure they are sufficiently accurate for monitoring changes
in the long term.
Operational monitoring and surveillance provide tools
for understanding and assessing natural and humaninduced disturbances and produces scientific 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 classifications were
not devised as a monitoring tool per se but were designated
to promote consistent interpretation of data. Assuming
that the biotope classification is ecologically sound, complementary methodologies will need to be introduced at a
S. Olenin, J.-P. Ducrotoy / Marine Pollution Bulletin 53 (2006) 20–29
suitable scale if certain designated sites should be surveyed
using such classifications.
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
25
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 different 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 first 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
S. Olenin, J.-P. Ducrotoy / Marine Pollution Bulletin 53 (2006) 20–29
elsewhere. Often production of the benthic macroalgae first
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) defined trophic types of
North Sea coastal tidal flats 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 differ 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; modification of bottom sediments, etc. Thus,
in their extended definition, biotopes can be considered
as functional units of a coastal marine ecosystem.
Complementary approaches to the use of biotope methodology are necessary. The classification of benthic animals
and macroalgae into functional groups (cf. Padilla and
Allen, 2000; Pearson, 2001) offers 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 fishes (Elliott and Dewailly, 1995). Thus, the biotope
concept can be adapted to fit 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 fish resources at the exposed coast of Lithuania, Baltic Sea
Biotope
Spawning ground
Fish fry shelter
Fish fry nursery
Adult fish feeding ground
1. Shallow hollows between the shoreline and the first
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
0
0
++
0
0
+
++
+
0
++
+
+
+
++
+
0
0
++
0
0
0
+
++
0
0
0
0
++
++ very important, + little important, 0 unimportant (based on Olenin and Daunys, in preparation).
S. Olenin, J.-P. Ducrotoy / Marine Pollution Bulletin 53 (2006) 20–29
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; Glémarec, 1997; Bek, 1997; Burkovsky and
Stoliarov, 2002; Ducrotoy and Olenin, 2003).
The concept of the ecosystem remains more difficult 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 fit 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 quantification of fluxes 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 fisheries.
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 reflection 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
d’Etude 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 final language check.
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