- FENS AND FLOODPLAINS OF THE TEMPERATE ZONE - 157
Applied Vegetation Science 9: 157-162, 2006
© IAVS; Opulus Press Uppsala.
This Special Feature focuses on lowland fens and ﬂood
plains. In this introduction we discuss the most important
mire-related terms, present status, threats and conservation and
restoration attempts. Floodplains and especially lowland fens
are rare and vulnerable ecosystems. They are highly threatened
all over the world because of direct conversion to agricultural
land and especially the lack of appropriate management and
altered catchment hydrology. Finally we present a framework
for the conservation and restoration of these ecosystems. This
consists of (1) optimising abiotic conditions; (2) safeguarding
propagule availability of the target species; (3) creating and
maintaining conditions for (re)establishment of these species,
and (4) appropriate management to keep the conditions
Keywords: Biodiversity; Diversity; River ﬂoodplain; Species
richness; Water table; Wetland.
The present volume contains 16 papers on the
analysis, management and restoration of riparian
wetlands and lowland fens, 14 of which were presented
at the 7th INTECOL international wetlands conference,
held from 25-30 July 2004 in Utrecht, The Netherlands.
The theme ﬁts the growing recognition of the value of
wetland ecosystems worldwide.
The importance of river ﬂoodplains as spawning
grounds for ﬁsh and as productive pasture for domestic
cattle is recognised since long by local people. Still, the
general attitude of marshes being wastelands comes to
expression in wetland-related words such as ʻswampʼ
and ʻmudʼ which deﬁnitely have negative connotations
in most languages. However, this attitude started to
change in the 1960s and 1970s. People began to realise
that these ecosystems did possess important values that
could be expressed even in economic terms (Pearce 1993;
Costanza et al. 1997; van den Berg et al. 2004).
Recently the biodiversity value of certain wetland
Fens and ﬂoodplains of the temperate zone:
Present status, threats, conservation and restoration
van Diggelen, Rudy1*; Middleton, Beth2; Bakker, Jan1; Grootjans, Ab1 & Wassen, Martin 3
1Community and Conservation Ecology Group, Biological Sciences, University of Groningen, PO Box 14, 9750 AA Haren,
The Netherlands; 2U.S. Geological Survey National Wetlands Research Center, 700 Cajundome Boulevard, Lafayette, LA
70506 USA; 3Copernicus Institute for Sustainable Development and Innovation, Environmental Sciences, Utrecht Univer-
sity, PO Box 80.115, 3508 TC Utrecht, The Netherlands;
*Corresponding author; E-mail firstname.lastname@example.org
types was recognised. Wetlands contain a large number
of adapted organisms (Mitch & Gosselink 2000) and, for
instance, calcareous fens are among the most species-
rich ecosystems of the temperate zone which also
harbour many endangered species (Wassen et al. 2005).
In addition to biodiversity there are other reasons why
wetlands are important. Mires play an important role in
the worldʼs carbon balance. Accumulating mires store
several 10 000s of kg C per ha per year (e.g. Asada et al.
2005), whereas drained mires lose even larger amounts of
CO2 (Wösten 1997) and other greenhouse gases (Flessa et
al. 1998; Groffmann et al. 2002). The role that wetlands
play in the puriﬁcation of water (Olde Venterink et al.
2006, this issue) and in dampening the effect of a large
variation in precipitation (Bragg & Lindsay 2003) is
important in many places, especially low-lying areas.
Wetlands also provide resources such as game, ﬁsh, reed
and wood (Bragg & Lindsay 2003).
Terminology and deﬁnitions
In the literature on wetland ecology there is much
confusion about terminology and especially about the
distinction between wetland, mire, fen, fen meadows and
similar terms. This calls for a section with deﬁnitions on
how we use some of these terms in this issue.
ʻWetlandsʼ are deﬁned as ʻareas of marsh, fen,
peatland or water, whether natural or artiﬁcial, permanent
or temporary, with water that is static or ﬂowing, fresh,
brackish or salt, including areas of marine water the
depth of which at low tide does not exceed six metresʼ
(Convention on Wetlands in Ramsar, Iran, 1971).
There are essentially two schools of thought with
respect to the deﬁnition of mires. The ﬁrst one states that
a ʻmireʼ is a peatland together with peat communities
(Godwin 1941), sometimes called a peat-producing
ecosystem (Godwin 1956; Gore 1983; Joosten & Clarke
2002) or an area that supports at least some vegetation
known to form peat, and usually includes a peat
158 VAN DIGGELEN, R. ET AL.
deposit (Bragg & Lindsay 2003). The second view
includes also some non-peat-producing ecosystems
such as groundwater-fed calcareous fens (Ratcliffe 1964;
Mörnsjö 1969) where the main deposit consists of tufa.
We follow Joosten & Clarke (2002) and use the word
mire in a strict sense for peat producing ecosystems.
Mires can be subdivided into bogs, fens and
ﬂoodplain mires. The deﬁnition of ʻbogʼ has always been
straightforward: a bog is a mire system that is entirely
dependent on precipitation for its water and solutes (Du
There is more uncertainty about the deﬁnition of
fen. Originally this term was simply used to describe
those types of mires which are not bogs (Du Rietz
1954). Soon people felt the need to make a distinction
between poor fens and rich fens, mainly based on
vegetation composition. Wheeler (1988) included similar
vegetation types on mineral soils also in the deﬁnition,
whereas Wheeler & Proctor (2000) used pH – and also
productivity to some degree – as criteria. All mires with a
pH > 5.5 were deﬁned as ʻfensʼ. Recently there has been
a tendency to narrow the deﬁnition of fen to ground-water
fed wetlands (Bedford & Godwin 2003). We agree with
this approach and use the word fen to mean all mires
that are pre-dominantly fed by groundwater. Note that
wooded wetlands (sometimes called ʻcarrʼ) are included
in this deﬁnition of fen.
The third type of mire is a ﬂoodplain, sometimes also
called ʻﬂood mireʼ (Succow & Joosten 2001). We deﬁne a
ʻﬂoodplainʼ as a mire that is predominantly fed by surface
water. This deﬁnition includes the tall sedge fens and reed
swamps from the European literature (e.g. Wheeler &
Proctor 2000) and most of the sedge meadows from the
American literature (Curtis 1959). These communities
are open wetlands dominated by sedges and grasses, and
consist mainly of tussocks of large helophytes.
The semi-natural communities ʻfen meadowsʼ and
ʻwet grasslandsʼ have developed after human mani-
pulation with water tables. Fen meadows have usually
developed from undrained fens after a modest lowering
of the water table to increase productivity. Wet grasslands
is a more general term that could include fen meadows,
but in a strict sense is used to describe managed (drained)
Distribution and present status
At present there are no exact data available on the
distribution and status of temperate fens and ﬂoodplains.
Nevertheless some rough ﬁgures exist. Lappalainen
(1997) and co-authors estimated that an area of slightly
less that 4 million km2 of mires exist throughout the entire
world and another 2.5 million km2 for other wetlands.
The great majority of these peat deposits are found in the
northern hemisphere, especially in the boreal zone, and
consist mainly of bogs. The sparse data available suggest
that bogs cover a much larger surface area than do fens,
which also occur in most countries of the temperate zone.
A study by Bragg & Lindsay (2003), however, shows the
opposite to be true in eight countries in central-eastern
Europe. Here, the majority of the mires consist of fens
with 78% of a total of ca. 73 000 km2 (5% of the total
surface of these countries). The authors estimate that
43% of this surface is still in a nearly natural state.
The situation in North America is rather similar to
Europe. The great majority of mires consists of bogs,
and fens are mainly found in localized areas with
groundwater outﬂow. In the United States, these areas are
found especially in the glaciated Midwest and Northeast,
as well as portions of the Appalachian Mountains and
mountainous West (Bedford & Godwin 2003). Only
fragmented information exists on the present state, e.g.
Pearson & Loeschke (1992) estimated that approximately
40% of the fens in Iowa were lost. Bedford & Godwin
(2003) wrote: “Few estimates of loss and current extent
exist, but where estimates are available, they indicate
extensive loss, fragmentation, and degradation”.
Changes in the status
Probably the most important factor affecting fens
and ﬂoodplains is their conversion into agricultural
ﬁelds. These wetlands were drained from medieval times
onwards and used for grazing cattle and making hay. In
the 20th century, drainage technology developed to such a
level that even crop production became possible on these
wet soils, especially (but not only) in former communist
countries. This conversion has resulted in substantial
emissions of carbon dioxide and other greenhouse gases,
making peat drainage a signiﬁcant contributor to global
warming (Flessa et al. 1998; Groffman et al. 2002).
Compared to bogs, fens are less often used for fuel
extraction, but the absolute surface used for this purpose
is probably a large threat, especially in somewhat more
southern regions where bog peat is rare. The total surface
affected is unknown, not in the least because this type
- FENS AND FLOODPLAINS OF THE TEMPERATE ZONE - 159
of peat extraction is not well-administered and mostly
carried out in small-scale excavations by one-man
companies. In bogs, large companies do most of the
Effects of hydrological changes
Fens are especially sensitive to relatively small
changes in the hydrological system. Human activities
such as groundwater abstraction, large-scale drainage
of the surroundings for agricultural purposes and use
of groundwater for irrigation lead to a diminished
groundwater ﬂow to the fen, even when conducted at
large distances from the fen. The effects of groundwater
abstraction may often not be visible in the water table
inside the fen, but it always leads to an increase in the
relative importance of rainwater and ﬁnally to acidi-
ﬁcation of the top layer (Wassen et al. 1996; van Diggelen
1998; Grootjans et al. 2006, this issue). This process may
take many decades (van Diggelen et al. 1996; van der
Hoek & Sýkora 2006, this issue) and go unnoticed for
a while (van Belle et al. 2006, this issue). The factors
that control the acidiﬁcation rate are not completely
understood yet, but the speed certainly depends on the
amount of acid produced (Kooijman & Paulissen 2006,
this issue) and the buffering capacity of the soil (van
Diggelen 1998; van Bremen & Buurman 2002).
Water tables in fens and ﬂoodplains are lowered due
to the more intensive drainage pressure, and this leads
to increased mineralisation rates. Most often increased
mineralisation rates result in higher biomass production,
but they may also lead to a shift in the limiting nutrient
(Wassen & Olde Venterink 2006, this issue; Higgins et
al. 2006, this issue; van Belle et al. 2006, this issue).
Productivity gradients and vegetation patterns change
accordingly, in response to altered competition intensity
for nutrients and light (Kotowski et al. 2006, this
The opposite situation as to water tables may occur as
well, although less often; fens and especially ﬂood-plains
sometimes become wetter because of changing water
regimes, building of dams and similar activities. Under
such conditions, the surface water component increases
and nutrient dynamics change, depending on water
chemistry and sediment load. Sedimentation is normally
the most important nutrient source in floodplains
(Olde Venterink et al. 2006, this issue), and increased
sedimentation leads to higher nutrient availability
(Werner & Zedler 2002) and affects vegetation patterns
through shifts in competitive interactions (Kotowski et
al. 2006, this issue). However, sedimentation rates vary
greatly between vegetation types with different structure
(Olde Venterink et al. 2006, this issue).
Conservation and restoration
Fens and ﬂoodplains are among the most species-
rich habitats but at the same time biodiversity decline
has been more intense in these areas than in many other
ecosystems. Conservation of existing fens and ﬂood-
plains and restoration of degraded ones, therefore, is a high
priority (Resolution VII.17 of the San José Conference
7th Meeting of the Conference of the Contracting Parties
to the Convention on Wetlands (Ramsar), 10-18 May
1999; Final Resolution Adopted at the 7th INTECOL
International Wetlands Conference, 25-30 July, 2004).
Van Diggelen & Marrs (2003) categorized essential steps
for conservation and restoration into four groups:
1. Establishing or re-establishing the necessary abiotic
2. Supplying (sufﬁcient) propagules of constituent spe-
cies of the target communities;
3. Creating and maintaining suitable conditions for the
(re-)establishment of target species;
4. Appropriate management to keep the conditions
Establishing and safeguarding necessary abiotic
conditions in affected wetlands almost always involves
raising the water table (Timmermann et al. 2006,
this issue), (re-)establishing the major water source
(rainwater, groundwater and surface water) for the
wetland under consideration and creating the necessary
productivity level regime (van Belle et al. 2006, this
issue). Rewetting is in itself technically not so difﬁcult to
achieve (Timmermann et al. 2006, this issue; Bodegom
et al. 2006, this issue), but conserving and/or restoring
the two other parameters may be much more difﬁcult.
Restoration is comparatively easy in ﬂoodplains,
especially along the edges of water bodies such as larger
rivers and lakes where the appropriate water type is
nearby. The productivity of the typical vegetation is high,
and this makes ﬂoodplain vegetation much less sensitive
to increased nutrient availability in water and air than are
low-production communities (Verhagen & van Diggelen
2006). In the case of fens, it is much more difﬁcult and
often impossible to conserve the necessary groundwater
feeding. The hydrology of the surrounding landscape
has often completely changed (Barendregt et al. 1995;
Grootjans et al. 2006, this issue), and groundwater is
replaced by rain or surface water. Critical parameters
such as productivity, limiting nutrient, light penetration
and pH shift outside the limits typical/necessary for fen
vegetation, and the characteristic species disappear (van
Bodegom et al. 2006, this issue).
Even if the critical abiotic constraints lie within the
tolerance of the target vegetation, this vegetation will
not necessarily contain all target species, nor will they
quickly reappear. Many species do not form a long-
160 VAN DIGGELEN, R. ET AL.
persistent seed bank and have to rely on dispersal to reach
a site after local extinction. Other authors (e.g. Novak &
Prach 2003; Galatowitsch 2006, this issue) found a clear
relationship between immigration rate and isolation of
a site, which suggests that dispersal is a constraint for
many species. There are also differences between modes
of dispersal. Soons (2006, this issue) showed that wind
dispersal is of limited value for most wetlands and is
negatively affected by increased productivity. Dispersal
by water (in ﬂooded parts) and by large herbivores (in
grazed parts) seem to be more efﬁcient dispersal vectors
(van den Broek et al. 2005; Middleton et al. 2006a, this
issue). Of course the latter implies that sites must be
connected to each other by a water course or by moving
Species establishment after restoration, (or, in the case
of annuals, yearly establishment) is a process that shows
many similarities to that of alien species that have to
establish in existing vegetation. Presently, we are not able
to predict which species will re-establish, and whether
or not these species will be invasive. Nevertheless, we
do know some of the constraints for establishment.
Kotowski et al. (2006, this issue) showed that competition
for light was the major factor that kept many species out
of the high-productive zone, whereas Bartha et al. (2003)
showed that colonisation rates increased signiﬁcantly
after extreme weather events when total species cover had
decreased considerably. Chirino et al. (2006, this issue),
on the other hand, showed that the relative establishment
success was independent from weather conditions but
did depend on species identity. All these results suggest
that interspeciﬁc competition is a major bottleneck for
Unlike the term ʻnatural areaʼ suggests, most fens
and ﬂoodplains are not capable of surviving without
regular human intervention. We know from palyno-
logical evidence (e.g. van Diggelen et al. 1991; Succow
& Joosten 2001; Grootjans et al. 2006, this issue) that
large surfaces of base-rich fens were present in natural
landscapes at one time. We know also that all over
Europe the remaining ʻnatural fensʼ are in fact ʻfen
meadowsʼ that were slightly drained (Wassen & Joosten
1996) and used for hay production and sometimes
grazing. The same was true in North America, except
that grazing was much more common there than it was
in Europe. These activities declined in the second half
of the 20th century, and the effects were the same in
both continents: shrubs and large helophytes started to
invade the sites and gradually took over, outcompeting
the original fen vegetation with many light-demanding
species (Kotowski et al. 2006, this issue). The situation
is much less dramatic in the more productive ﬂoodplains,
but abandonment also leads to shrub invasion in these
areas (Jensen 1998).
Nature management tries to counteract these un-
wanted developments by applying certain management
techniques that mimic traditional agricultural manage-
ment. The objective of all techniques is to remove exces-
sive nutrients and create recruitment gaps for low-com-
petitive species. The techniques used include mowing
and removing hay (Slotte 2001), grazing (Littlewood et
al. 2006, this issue) and also burning (Middleton 2002).
Apart from many similarities, there are also considerable
differences between these techniques (van Diggelen &
Marrs 2003; Middleton et al. 2006b, this issue). Mowing
creates a spatially homogeneous situation that is highly
heterogeneous in time with high selection pressure on
species for exact timing of the life-cycle. Low-intensity
grazing, on the other hand, often results in a spatially
heterogeneous pattern that is stable in time because
herbivores create and maintain intensively used graz-
ing lawns adjacent to hardly used spots (Bakker et al.
1984). There are still many uncertainties about the role
of ﬁre in managing fens and ﬂoodplains. It is obvious
that prescribed burning removes dead vegetation and, at
least temporarily, results in an increase of biodiversity
but winter ﬁre does not control shrubs.
Although not exhaustive, this introduction should
point to many research questions concerning the
maintenance and restoration of fen and ﬂoodplain
biodiversity in the temperate zone. This special issue of
Applied Vegetation Science is a ﬁrst attempt.
Acknowledgements. We thank Jarita Davis for editorial
- FENS AND FLOODPLAINS OF THE TEMPERATE ZONE - 161
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