Applied Vegetation Science 11: 99-106, 2008
doi: 10.3170/2007-7-18329, published online 12 November 2007
- drainage
© IAVS; Opulus Press and initial effects of hydrological restoration on rich fen vegetation
Uppsala.
Long-term effects of drainage and initial effects of
hydrological restoration on rich fen vegetation
Mälson, K.1*; Backéus, I.1,2 & Rydin, H.1,3
1Department of Plant Ecology, Evolutionary Biology Centre, Uppsala University, Villavägen 14, SE-752 36 Uppsala, Sweden;
2E-mail Ingvar.Backeus@ebc.uu.se; 3E-mail Hakan.Rydin@ebc.uu.se;
*Corresponding author; Fax +46 18553419; E-mail kalle.malson@ebc.uu.se; Url: www.vaxtbio.uu.se
Abstract
Questions: What vegetational changes does a boreal rich fen
(alkaline fen) undergo during a time period of 24 years after
drainage? How is plant species richness affected, and what are
the changes in composition of ecological groups of species? Is
it possible to recover parts of the original flora by rewetting the
rich fen? Which are the initial vegetation changes in the flora
after rewetting? What are the major challenges for restoration
of rich fen flora after rewetting?
Location: Eastern central Sweden, southern boreal vegetational
zone. Previously rich fen site, drained for forestry purposes
during 1978-1979. The site was hydrologically restored (re-
wetted) in 2002.
Method: Annual vegetation survey in permanent plots during
a period of 28 years.
Results: There were three successional stages in the vegeta-
tional changes. In the first stage there was a rapid (< 5 years)
loss of rich fen bryophytes. The second step was an increase of
sedges and early successional bryophytes, which was followed
by an increase of a few emerging dominants, such as Molinia
caerulea, Betula pubescens and Sphagnum spp. After rewet-
ting, there are indications of vegetation recovery, albeit at slow
rates. Depending on, for instance, initial species composition
different routes of vegetation change were observed in the flora
after drainage, although after 24 years, species composition
became more homogenous and dominated by a few species
with high cover.
Conclusion: Major changes have occurred after changes in
the hydrology (drainage and rewetting) with a severe impact
on the biodiversity among vascular plants and bryophytes.
Several rich fen bryophytes respond quickly to the changes in
water level (in contrast to vascular plants). The recovery after
rewetting towards the original rich fen vegetation is slow, as
delayed by substrate degradation, dispersal limitation and
presence of dominant species.
Keywords: Bryophyte; Long-term study; Mire; Rewetting;
Succession; Wetland restoration.
Nomenclature: Karlsson (2002) for vascular plants; Halling-
bäck et al. (2006) for bryophytes.
- 99
Introduction
Sweden is the country within the European Union
with the largest area of rich fens (ca. 100 000 - 150 000
ha or 2-3 ‰ of the total land area; Sundberg 2006). Many
of these rich fens have been affected by drainage during
the last two centuries – during the 19th century mainly
for agricultural purposes and during the 20th century for
forestry (Vasander et al. 2003). Ca. one million ha of the
peatland area in Sweden have been drained for forestry
and at a rough estimate equally large areas have been
drained for agriculture (Rydin et al. 1999).
Drainage may alter the peat chemistry (for example
lowered pH and leakage of cations; see Nauke et al. 1993)
and biological attributes (invasion of dominant species)
as a result of the combination of lowered water table and
peat decomposition. The species composition may also
respond to changes at the landscape level, such as altered
water levels and outflow of groundwater in the surround-
ings (Grootjans & van Diggelen 1995; van Diggelen et
al. 2006). Also, increased outflow of leached chemicals,
for example Ca-ions, have been observed from drained
peat (Nauke et al. 1993). Other factors, for example
reduced management such as haymaking (Moen 1995),
eutrophication and acidification (Sjörs & Gunnarsson
2002) may also cause drastic changes in the rich fen
flora. It has been suggested that vegetation changes after
drainage in mire ecosystems resemble effects of global
warming (Laine et al. 1995). The effect of drainage on
mire vegetation has been reported to be more drastic in
minerotrophic than in ombrotrophic sites (Heikkilä &
Lindholm 1995). Concerning acidification, the effects
are more obvious in less buffered systems (Wheeler
& Shaw 1995) and, hence, these effects are more pro-
nounced in moderately rich fens than in extremely rich
fens with higher buffering capacity (high concentration
of calcium), and the effects are also small in naturally
acid poor fens and bogs (Gunnarsson et al. 2000).
Rich fens are very important from a conservation
point of view and act as biodiversity hotspots in the
landscape (Wassen et al. 2005; van Diggelen et al. 2006).
100 Mälson, K. et al.
Rich fens contain a large proportion of threatened spe-
cies of various groups of organisms (including vascular
plants, bryophytes, land snails and insects): 160 rich
fen species occur in the Swedish Red list and among
these 74 are considered as threatened (Gärdenfors 2005;
Sundberg 2006). Even moderate hydrological changes
in these ecosystems may have a drastic, negative impact
on the biodiversity, not least among vascular plants and
bryophytes, but continuous, long-time monitoring of
these processes are still lacking.
The recent interest and focus on restoration of
mires (Pfadenhauer & Klötzli 1996; Pfadenhauer &
Grootjans 1999; Vasander et al. 2003) also necessitates
evaluations of the vegetation – will the mire vegetation
recover and how long will it take? Several restoration
studies have been performed in wetland ecosystems in
western/central Europe (for a review see Pfadenhauer
& Klötzli 1996), but the dissimilarities in wetland types
and species composition often make comparisons with
Nordic mire ecosystems difficult. A long-term study was
performed in a fen meadow affected by drainage in The
Netherlands (Grootjans et al. 2005), with low similari-
ties in species composition to rich fens in the Nordic
countries. Even though there are large mire areas in the
Nordic countries that are directly or indirectly highly
influenced by drainage, only few scientifically evalu-
ated restoration projects have been carried out in these
countries (although several studies of vegetation changes
after drainage have been performed; see Backéus 1981).
There is a big need for restoration of rich fens from a
national point of view; 67 % (1067 objects) of the highest
classed rich fens in the Swedish wetland inventory are
affected by drainage (Sundberg 2006). There is also a
need for long-term vegetation monitoring, not only for
vascular plants but also for bryophytes, since the latter
are often better indicators of changed mire hydrology
than are vascular plants.
Comparison of drainage and restoration effects across
regions is facilitated if studies are based on ecological
groups, rather than on individual species. The use of
functional groups has been discussed extensively during
recent years, including their use in different biotopes
and vegetation systems (for example Smith 1997), their
usefulness for vegetation and biodiversity monitoring
(Rusch et al. 2003) and for management and restoration
(Hobbs 1997; Brown 2004). We use a similar approach
and describe the changes in ecological groups of species,
based on life form and affinity to rich fens.
In this study, we investigated the immediate and long-
term post drainage floristic responses in permanent plots
in a rich fen in central Sweden during a period of 24 years
with the aim to document time scales of changes in spe-
cies composition and in abundance of characteristic rich
fen species and other ecological groups. Thereafter, we
hydrologically restored (re-wetted) the fen and followed
the vegetation changes during four years to test if the
vegetation reverts back towards original composition,
and, if so, at what rate.
Material and Methods
Drainage and restoration experiment
The study site is in Gästrikland (east central Sweden;
60°51' N, 17°10' E), i.e. in the southern-boreal vege­
tation zone (Sjörs 1999). The mean temperature for the
nearest meteorological station (Gävle-Åbyggeby) for
January is –5.1°C and for July 15.4°C, and the annual
precipitation is 642 mm (Anon. 2005). The whole mire
complex, including the rich fen parts which are the focus
of this study, was drained for forestry purposes during
1978-1979 with one main ditch (1 m deep, 2 m wide)
and several smaller feeder ditches which all drain into a
small lake north of the study site. The bedrock in the area
is acidic, but the morainic soils in this coastal area are
influenced by calcareous material from nearby bedrock in
the Bothnian Bay that was deposited here during the last
glaciation. This calcareous material noticeably influences
the minerotrophic waters passing into the mire and also
influences the ecological properties of the mire vegeta-
tion. No water-chemical data from the pristine mire stage
are available. The pH in the main ditch close to the plots
was 6.4 (measured in May 2002) and the Ca concentra-
tion was 18 mg/l (measured in August 2005).
Still undisturbed mires in the vicinity of the experi-
mental site contain moderately rich fen vegetation but
also poor fens and bog communities. The vegetation
before drainage was dominated in the wetter parts by
brown moss species (mainly Amblystegiaceae s.l. such as
Scorpidium spp. and Campylium stellatum) and sedges,
such as Carex rostrata, C. panicea and C. livida.
Hydrological restoration by blocking of the main
ditch was performed in December 2002. Blocking the
main ditch was done at four positions (separated by a
distance of 50 m) along the main ditch by establishment
of a wooden dam in combination with peat and mineral
soil that was compacted by an excavator 3 m upstream
of the wooden dam. The altitudinal differences between
adjacent dams did not exceed 20 cm. In the investigated
part of the mire the water level rise after hydrological
restoration was approximately 175 mm in a dip well 4
m from the ditch.
Vegetation analysis
Nine subplots (1 m × 1 m) were arranged in each of
two plots (5 m × 5 m and 6 m × 10 m), ca. 10 m from
 - drainage and initial effects of hydrological restoration on rich fen vegetation - 101

Table 1. Species in the two plots divided into ecological groups based on life form and affinity to rich fens in the region.

Rich fen sedges: Herbs: Frangula alnus Pellia sp.

Carex livida
Carex panicea
Rhynchospora fusca
Trichophorum alpinum
Other sedges:
Carex chordorrhiza
Carex echinata
Carex lasiocarpa
Carex limosa
Carex nigra
Carex rostrata
Eriophorum angustifolium
Eriophorum vaginatum
Rhynchospora alba
Grasses:
Calamagrostis canescens
Deschampsia flexuosa
Molinia caerulea
Cirsium palustre Myrica gale Pohlia cf. nutans
Drosera intermedia Picea abies Polytrichastrum longisetum
Drosera rotundifolia Pinus sylvestris Polytrichum commune
Epilobium palustre Rubus idaeus Riccia sp.
Equisetum fluviatile Salix caprea Splachnum sp.
Galium palustre Salix cinerea Warnstorfia exannulata
Hammarbya paludosa Salix myrsinifolia
Menyanthes trifoliata Salix repens Rich fen Sphagnum:
Potentilla erecta Vaccinium oxycoccus Sphagnum centrale
Ranunculus flammula Sphagnum contortum
Selaginella selaginoides Rich fen bryophytes: Sphagnum platyphyllum
Solidago virgaurea Bryum pseudotriquetrum Sphagnum subfulvum
Trientalis europaea Campylium stellatum Sphagnum subnitens
Utricularia intermedia Scorpidium cossonii Sphagnum warnstorfii
Viola palustris Scorpidium scorpioides
Viola riviniana Other Sphagnum species:
Other bryophytes: Sphagnum capillifolium
Woody species: Aulacomnium palustre Sphagnum cuspidatum
Alnus glutinosa Brachythecium sp. Sphagnum fallax
Betula pubescens Dicranella cerviculata Sphagnum girgensohnii
Calluna vulgaris Mnium sp. Sphagnum papillosum
Sphagnum squarrosum

the main ditch. Each plot was investigated annually in Results

July-September 1979-1997 (except 1985 and 1996). The
floristic monitoring was resumed in 2002 (before the hy-
drological restoration) and continued to 2006. One person
(I. Backéus) took part in all vegetation analyses. All vas-
cular plants and bryophytes were recorded, and their cover
in percent was estimated according to the following scale:
0-1 %; 1-2 %; 2-3 %; 3-4 %; 5-10 %; 10-20%; 20-30%
etc. The cover of the shrub layer (0.5-2 m) and tree-layer
(higher than 2 m) was estimated five times during the
investigation period (1979, 1984, 1989, 1995 and 2006)
as vertical projection above each subplot.
Statistical analyses
Multivariate analyses were performed with the
CANOCO software, version 4.51 (ter Braak & Šmilauer
2003). Detrended Correspondence Analysis, DCA, was
used. Detrending was performed by segments with no
further transformation of data. Average cover values
of subplots were used for analysis of changes among
all investigated years. For detailed ordination within 4
selected years, individual cover values for all subplots
were used.
Classification into ecological groups
To evaluate the effects of drainage and restoration
and facilitate comparisons with other studies (with a
different set of species), the species were divided into
nine ecological groups (Table 1) based on life form and
affinity to rich fens in the region (for example Rydin et
al. 1999).
A few species determined the major changes in spe-
cies composition throughout the investigation period
(Fig. 1). The decrease and local extinction of rich-fen
bryophytes, mainly Scorpidium scorpioides in plot 1
and Campylium stellatum in plot 2 occurred rapidly
after drainage. Following their decrease, several rich
fen sedges increased. In both plots, C. panicea peaked
at 5-10 years after drainage (Fig. 1).
At the final vegetation analysis before rewetting
(2002), Molinia caerulea clearly dominated among the
vascular plants. In plot 1, the species was absent in 1979,
but covered > 50% in 2002. In plot 2, cover increased
on average from 7% in 1979 to > 50 % in 1992. After
hydrological restoration in December 2002, Molinia
peaked in 2003 in Plot 1 and then decreased, while its
reduction started already in 2003 in plot 2 (Fig. 1).
A unidirectional movement along axis 1 in the DCA
ordination for each year can be seen (Fig. 2; Table 2).
The changes were especially rapid in 1982-1985. The
movement of the plots towards higher loadings along
axis 2 in these DCA ordinations fairly well coincides
with the major expansion phase of Molinia caerulea.
For both plots, a reduction of the movement or even a
minor transition back along axis 1 was observed after
hydrological restoration.
In a second ordination, the within-plot floristic vari-
ation was evaluated at four selected years after drainage.
Selected years were 1979 (first year after drainage), 1988,
1997 and 2006 (4 years after rewetting). Plot 1 initially
had a uniform vegetation among all subplots (seen by
the tight cluster of subplots of 1979 in the ordination;
Fig. 3), dominated by Scorpidium scorpioides and a few
102 Mälson, K. et al.

Fig. 2. DCA ordination showing sample scores based on aver-

age values of species cover in the nine subplots within each
plot for all years (1979-2006, indicated by inserted numbers).
The scaling of axis is in S.D. units.

Several similarities in the composition of ecological

groups can be seen in the two plots (Fig. 4): early decline
of rich fen bryophytes followed by expansion of woody
species and rich fen sedges, and later their replacement
by grasses. The invasion and expansion of Sphagnum
and other bryophytes occurred only in plot 1.

Fig. 1. Change in cover (average cover of the nine subplots)

in some common species during the investigation period
(1979-2006). The selected species covered more than 5 % in
at least one year.

dominant sedges (C. rostrata, C. livida). This uniform

vegetation diverged rapidly and intensely after drain-
age. In plot 2, the initial vegetation was more diverse
(with several drier subplots and higher proportion of
woody plants and grasses) and in this plot we see a
convergence in the flora as time after drainage passes
by (Fig. 3; Table 2).
The tree and shrub layers played a small role only
during the first 5 years after drainage, but then shrubs
expanded and peaked in 1995. As the birches grew taller
they started to form an expanding tree layer and the shrub
cover decreased (Table 3).
A considerable turnover of species occurred in the Fig. 3. DCA ordination showing sample scores in the nine sub-
plots based on cover values in the two plots at four individual
first 10 years after drainage. The species richness during
years after drainage. Selected years were 1979 (first year after
the drainage period was highest in 1988. Species richness drainage), 1988 (ten years after drainage), 1997 (20 years after
was higher in 2006 (4 years after rewetting) than in 1997 drainage) and 2006 (4 years after rewetting). The scaling of
(before rewetting) (Table 4.). axes is in S.D. units.
 - drainage and initial effects of hydrological restoration on rich fen vegetation - 103

Table 2. Summary of properties of the axes in the DCA ordinations. Gradient lengths are given in S.D units. E/TI is the eigenvalue
of the specific axis divided by the total inertia.

Plot Figure Data No. No. Total Axis Gradient Eigenvalue

of samples of species inertia no. length E/TI

1 2a All years, subplots averaged 22 67 1.870 1 4.749 0.844 0.451
2 1.426 0.100 0.053

3a Selected years, all subplots 36 48 4.120 1 5,078 0.912 0.221
2 2.896 0.411 0.100

2 2b All years, subplot averaged 22 50 0.887 1 2.017 0.490 0.553
2 0.579 0.033 0.037

3b Selected years, all subplots 36 34 2.267 1 3.489 0.782 0.345
2 1.697 0.185 0.082

Discussion

We saw drastic changes in the rich fen vegetation

during the period of 24 years after drainage, and
the changes occurred in a stepwise manner. Earlier
studies stress the importance of detailed long-term,
continuous studies in permanent plots of vegetation
changes after drainage of mires (Silfverberg et al.
2005) and the lack of knowledge about restoration ef-
fects in drained forested mires (Heikkilä & Lindholm
1995). A long-term study of vegetation changes was
performed in a fen meadow affected by drainage in
The Netherlands (Grootjans et al. 2005). That study
shows low similarities in species composition with
rich fens in the Nordic countries, but there are several
similarities in changes of ecological groups of plants
on a community level.
In our study, a continuous, detailed description of
the vegetation changes in a rich fen after drainage but
also after rewetting is given over a time span that is
unusual for permanent plot studies of any kind.

Fig. 4. Vegetation composition divided into ecological groups

Table 3. Average cover (n = 9 in each plot) of shrub and tree
layer (0.5 - 2 m and > 2 m, respectively) measured in per-
cent, analysed in a vertical projection above each individual
subplot.
of species at four different years during the investigation period
(1979, 1988, 1997 and 2006).
Table 4. Species turnover at four occasions during the inves-
tigation period.

Cover (%) 1979 1988 1997 2006

1979 1984 1989 1995 2006
Plot 1
Plot 1 Total number of species 14 22 15 27
Tree layer 0.3 0.3 0.5 9.2 12.6 Number of original (1979) species remaining - 9 6 6
Shrub layer 0.3 0.0 7.5 26.0 4.7 New species in comparison to previous column - 13 2 14

Plot 2 Plot 2
Tree layer 5.3 5.3 0.5 18.1 19.5 Total number of species 14 16 13 18
Shrub layer 0.0 2.6 6.6 10.7 1.4 Number of original (1979) species remaining - 7 6 6
New species in comparison to previous column - 9 4 7
104 Mälson, K. et al.
Changes in dominance and species richness after
drainage
The changes in species composition occurred in
three major steps. The first step was a major decrease of
several species (mainly brown mosses) during the initial
5 years after drainage, similar to the changes reported
by Silfverberg et al. (2005). After this stage, we saw a
stage of high species turnover and an increase of rich
fen sedges (such as Carex panicea) and woody species
(e.g. Myrica gale) in plot 2, which again declined after
ca. 10 years. The third stage, with major emergence of
dominants (Molinia and Betula), started at different
times after drainage, but after almost 30 years the species
composition was quite similar in the two plots.
The rapid loss of brown mosses is noteworthy from
a biodiversity point of view and indicates the direct
influence of aeration of the peat for these groups of
species. The loss of brown mosses occurred during the
first 5 years after drainage. These changes were most
pronounced in the wettest parts, and species in these
microhabitats have earlier been reported to be the first
species to disappear after drainage (Laine et al. 1995).
It is noteworthy that species richness peaked during
the period of drainage, with the highest number found
after about 10 years. At this stage, species from several
groups of vascular plants and bryophytes did occur in
the mire. Species from the pristine stage in combination
with ruderal colonists and forest species all contributed
to the high species richness, as has also been shown in
other studies (Vasander 1987).
Spatial variation in effects
The difference in the vegetational change between
the plots most probably reflects the fact that the surface
of plot 1 initially was drier. The vegetation there was
initially very homogeneous. The vegetation changes took
different routes, mainly during the first period (1979-
1997) after drainage, although the species composition
and distribution between ecological groups converged
later on. Spatial heterogeneity will always occur and
may be promoted by an expanding shrub and tree cover.
Knowledge of different routes and rates of changes after
hydrological interaction is needed to understand in which
phase of degradation the fen is. When this is understood.
it is easier to suggest correct restoration measures.
In plot 1 the vegetation in the subplots diverged
through time, while in plot 2 the subplots converged,
which most probably reflects the different initial condi-
tions. This indicates that different parts of individual
mires may respond to drainage depending on initial
species composition. The most drastic species turnover
will be seen in the wetter parts, where the rich fen bryo-
phytes dominate and respond directly to the lowered
water table (see Rydin et al. 1999; Hedenäs & Kooijman
1996). These conditions also clearly promote the invasion
and expansion of Sphagnum and Polytrichastrum (see
below for their effects).
The emergence and expansion of dominant species
Molinia caerulea has been reported in earlier studies
to expand in, for example, heathlands in western Europe
as an effect of increased nitrogen availability, and it seems
to be a dominant grass excluding other species (Berendse
& Elberse 1990). As shown in our study, the establish-
ment of Molinia may take some time, but then the ex-
pansion is very rapid. Several studies have discussed the
increase of Molinia, and the reasons therefore. Annual
or biennial scything reduces Molinia (Moen et al. 1999)
and dry periods are expected to lead to an expansion of
the species (Boeye et al. 1999). This has been shown
to be partly due to effective use of phosphorus in drier
habitats in comparison with wetter sites (El-Kahloun et
al. 2000). Molinia also has a wide pH tolerance (Taylor et
al. 2001) and is classified as a stress tolerant competitor
(Grime et al. 1988). Although we did not investigate the
physiological explanation for the expansion of Molinia,
our data indicate that it is a superior competitor under
drier conditions in rich fens.
The effects on species diversity should also be dis-
cussed in relation to the increase of grasses and woody
species, which probably caused decreased levels of light,
increased litter accumulation and desiccation of the upper
peat profile. Similar increase of grasses after drainage
in combination with reduced species richness has been
described from wetland ecosystems in The Netherlands
(Grootjans et al. 2005). There, natural compaction of the
peat layers led to rewetting some 16 years after drainage,
which caused a decline of the dominant grasses. This is
similar to the reduction of Molinia we found after the
hydrological restorations.
Effects of expanding bryophytes
In plot 1, the initial great loss of brown mosses was
later followed by an increase in other bryophytes, such
as Pohlia nutans and Polytrichastrum longisetum (stress
tolerant ruderals; cf. Grime 2002) that started after the
decrease of rich fen sedges, approximately ten years after
drainage. These bryophytes seem to be good colonizers
that explore the bare peat surface of decaying brown
mosses and sedges. The dominance by Polytrichastrum
will prevent recolonization of other bryophytes due to
their rapid spatial cover of bare peat patches. A similar
succession of bryophytes has been reported from for-
ested fens in Finland, where decreasing Sphagnum was
 - drainage and initial effects of hydrological restoration on rich fen vegetation -
replaced by colonizers established by spores (for example
Pohlia nutans) and then followed by forest floor species
(Laine et al. 1995).
One major obstacle for the recovery of the vegetation
is the increase of Sphagnum after rewetting in parts of the
mire. These mosses will change the chemical properties
of the water and surface and inhibit recolonization of
brown mosses. One such expanding species is Sphag-
num squarrosum, which has been reported to respond
positively to increased levels of nutrients (Kooijman &
Bakker 1995) and to replace species in minerotrophic mires
after eutrophication and acidification. It is one of the first
species that invade rich fens that have become drier and
more acid (Hedenäs & Kooijman 1996). Several brown
moss species (e.g. Scorpidium scorpioides and Campylium
stellatum), have the same pH range as Sphagnum squar-
rosum (Andrus 1986; Hedenäs 2003), but the presence of
Sphagnum will gradually lower the pH further (Clymo
1963, 1964). Sphagnum hummocks may also overgrow
the brown mosses (Kooijman et al. 1994).
Restoration
In addition to unsuitable chemistry and hydrology,
restoration may be unsuccessful because of dispersal
limitations. Since several rich fen specialists disappear
in just a few years after drainage, natural establishments
have to rely on long distance dispersal, dispersal from
remnant populations in the surrounding mire, or from
long-term persistent seed banks (Jensen 2004). Overcom-
ing the dispersal distances by the use of transplantations
of locally extinct species may be feasible (Mälson &
Rydin 2007).
Restoration of rich fen vegetation depends on the
severity of the damage (Grootjans & van Diggelen 1995),
the time that has passed since drainage (as is clear from
our results), the changes in chemical properties of the
peat and the possibilities to re-introduce (in our case Ca-
rich) water (Heikkilä & Lindholm 1995). A eutrophicated
and acidified surface peat may have to be removed from
the site to facilitate successful establishment of locally
extinct species, but it is also possible to add a small
amount of lime to the peat surface to promote brown
moss establishment (Mälson & Rydin 2007).
In summary, several actions may be needed for suc-
cessful regeneration of the rich fen flora. Rewetting is
the first step, and should preferably be more drastic
than in this study. Removal, perhaps repeated, of domi-
nant species (trees, shrubs, Molinia, Polytrichastrum,
Sphagnum) in combination with surface peat removal,
liming and reintroduction of rich fen specialists may also
be necessary. Otherwise the substrate may once again
degrade as an effect of invading dominants.
105
Acknowledgements. The study was financed by the Swedish
Environmental Protection Agency in 1980-1982 and 2002-
2006. We thank Sebastian Sundberg and two anonymous
reviewers for comments on the manuscript.
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Received 7 February 2007;
Accepted 23 May 2007;
Co-ordinating Editor: J. Pfadenhauer.