205Natural Revegetation and Restoration of Drained and Cut-overRaised Bogs in Southern Germany — a Comparative Analysis
of Four Long-term Monitoring Studies
Peter POSCHLOD1*, Christina MEINDL1, Jan SLIVA2, Udo HERKOMMER3, Matthias JÄGER4,
Ulrike SCHUCKERT5, Andreas SEEMANN1, Anja ULLMANN3 and Teresa WALLNER1
1

Institute of Botany, Faculty of Biology and Preclinical Medicine, University of Regensburg,
D-93040 Regensburg, Germany
2
Department of Ecology, Technical University of Munich, D-85350 Freising-Weihenstephan, Germany3
agl Ulm (Working group Landscape Ecology Ulm), Marlene-Dietrich-Strasse 1, D 89231 Neu-Ulm, Germany
4
Institute of Animal Ecology and Tropical Biology, Faculty of Biology, University of Würzburg,
D-97074 Würzburg, Germany
5
Im Weizen 47, D-71636 Ludwigsburg, Germany*corresponding author

Abstract
Mires and peatlands have become the focus of nature conservation and restoration management policy
due to their multiple ecological functions such as accumulation, disposal, retention and conservation, as wellas their possible role in global climate regulation. In this paper we present the results of four long-term studies on spontaneous as well as directed revegetation of peat-mining areas in raised bogs situated in the foothills of the Alps in southern Germany. The results clearly show that(1) peat mining techniques affect spontaneous revegetation, peat-cutting being a technique which may
allow peat-forming vegetation to reestablish itself after abandonment of peat-mining. In contrast, peat
milling leads to mono-dominant successional stages of certain species that remain stable in their respective
stages over decades,(2) hydrological conditions after renewed inundation (rewetting) affect the reestablishment ofpeat-forming vegetation. In our case study, rewetting caused the flooding of large peat-mined areas, resulting in the extinction of trees, shrubs and dwarf shrubs. However, it also supported the establishment of floating mats consisting mainly of fen species which may initiate, in the long-term, the reestablishment of theformer peat-forming raised bog vegetation and(3) reintroduction of vascular plants typical of raised and transitional bogs by sowing and planting may
accelerate the establishment of a dense vegetation cover as a prerequisite for the establishment of other species. However, the establishment of a Sphagnum cover with typical peat-forming species such as Sphagnum
magellanicum has not been successful on a short- or mid-term basis and remains a major challenge inrestoring cut-over raised bogs.
Key words: mires, peatlands, peat-mining, re-wetting, succession

1. Introduction
Mires have multiple functions on global, regional
and local scales (Poschlod, 1994; Kapfer & Poschlod,
1998; Joosten & Clarke, 2002; Joosten, 2003). Globally,
they play an important role in the carbon cycle (accumulation function; Gorham, 1991, 1995; Franzén, 1994;
Roulet, 2000), regionally they buffer pollution by nutrients, heavy metals, etc. (disposal function; Succow &Jeschke, 1986), as well as mitigating heavy rainfall
events (retention function; Schmeidl et al., 1970;
Global Environmental Research
11: 205-216 (2007)
printed in Japan

Ingram, 1983). Locally they provide habitats for a lot of
specifically adapted, ice-age relict, rare and threatened
species (conservation function; Kapfer & Poschlod,1998). Furthermore, mires are an archive of climate,vegetation and even pollution history (archive function;
Frenzel et al., 1991; Görres, 1991).
Despite these multiple functions, mires have beenstrongly affected by agriculture, forestry and industry in
all countries of the northern temperate and boreal zones.
However, in southern Germany (Table 1), melioration ofmires for agricultural purposes started only at the end of©2007 AIRIES206

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the 18th century when the human population increased,
which in turn resulted in the need to produce more food.
A famous South German example is the drainage andcultivation of the large geogenous “Donaumoos” near
Ingolstadt from 1790, where elector Karl Theodor established new settlements and introduced potato cultivation(Pfadenhauer et al., 1991). Only at the end of the 19thand beginning of the 20th century, large-scale peatmining started in ombrogenous mires, first for fuel (railways, heating, etc.) and locally for litter (Poschlod,1990). From the 1960s on, peat was mined on a large
scale only for horticultural purposes (Göttlich, 1990).
Today, there are no mires left which have not been influenced by any type of the above-mentioned land-uses.
Furthermore, mires were, until recently, affected byatmospheric acid depositions, namely acidic sulphur(Ferguson et al., 1978; Ferguson & Lee, 1980) followedby an ombrotrophication of minerotrophic mires (Zoller& Seeldorf, 1989). Nitrogen (Lütke-Twenhöven, 1992a,b;Frankl, 1996) and dust (Pakarinen, 1977) are also
deposited in mires. Nitrogen depositions in ombrogenous
mires may result in a better growth of peat followed bysurface desiccation and tree invasion (Frankl, 1996) but
also in a shift towards species which are not able toform peat (Lütke-Twenhöven, 1992a,b), especially
species from the Sphagnum recurvum group. In Southern
Germany, the occurrence of Sphagnum angustifolium
was not reported from raised bogs before the 1980s(Poschlod, 1990).
Ombrogenous mires of southern Germany, which aremostly situated in the foothills of the Alps where precipitation is higher than 1,000 mm/year, have been
mainly affected by drainage followed by peat mining.
Only small areas were afforested or put to agriculturaluse. Therefore, these peatlands today often represent a
mosaic of drained, small or large peat mining sites withmore or less intact areas of peat forming communities.
Due to the above-mentioned multiple functions it is now
an aim of the federal not only to maintain more or lessintact areas – permission for drainage and peat mining
have not been granted since the end of the 20th century– but also to restore drained and mined areas (Pfadenhauer,
1988). However, is restoration management necessary atall? Comparing successional stages of abandoned peatmining areas in nine peatlands in the foothills of theAlps, Poschlod (1990, 1994) suggested that vegetation
development without management depends on (1) thepeat mining technique that was used, with peat cutting
being more suitable for the reestablishment of peatforming vegetation than peat milling, and (2) the thickness and quality, also (3) water quality as well as (4) thehydrological regime. Restoration management methods
have been deduced from these comparative studies(Poschlod, 1990) which involved rewetting of heath andforests in dry peat- cut areas as well as manipulation of
the surface topography, rewetting, sowing of seeds andtransplanting of plant material on peat-milled areas(Maas & Poschlod, 1991; Sliva, 1997; Sliva &Pfadenhauer, 1999; Pfadenhauer & Grootjans, 1999).
Until now, the hypotheses concerning successfulvegetation development without management were
never proven through long-term observations of thesesites. Only short-term studies existed which did notallow for adequate assessment (Pfadenhauer & Klötzli,
1996; Gorham & Rochefort, 2003; Vasander et al.,2003). However, long-term monitoring of restorationsites is necessary to confirm that restoration management practices are appropriate.
In this paper we present new results (“Wurzacher
Ried,” “Wieninger Filz,” “Wendlinger Filz”), and reconsider already published results (“Kendlmühlfilz”;
Sliva, 1997) regarding the following questions and casestudies in ombrogenous peatlands in the foothills of theAlps:(1) Is spontaneous revegetation in abandoned peatmining areas towards peat-forming vegetation (including peat growth) possible without restoration
management (case studies “Wieninger Filz,”

Table 1 Historical and current impacts by agricultural, forestry and industrial use and the current situation of mires/peatlands inSouthern Germany (Pfadenhauer, 1988; Poschlod, 1996). Nomenclature of mire/peatland types according Joosten and Clarke(2002). In bold – names of case study mires/peatlands in this paper.
Type of mires
Geogenous mires (developed from rising ground-water after theice age – primary mire type: water-rise mires, secondary type:
percolation mires) in the valleys of the Danube and rivers joining
it (e.g., “Donaumoos” at Ingolstadt, “Osterried” near Ulm)
Large complexes of geo- and ombrogenous mires in former
glacier basins (primary mire type: → terrestrialisation mires,
secondary type: → raised bogs) in the foothills of the Alps (e.g.,“Kendlmühlfilz” southeast of Munich, “Wurzacher Ried” southof Ulm)Small geo- and ombrogenous mires in small basins within
end/ground moraine landscape (developed mostly from lakes;
primary mire type: → terrestrialisation mires, secondary type:→ raised bogs) in the foot-hills of the Alps (e.g., “Wieninger
Filz” near Traunstein, “Wendlinger Filz” southeast of Munich)
Small geogenous mires on slopes (sloping mires, spring mires) inground/end moraine landscape (developed from springs) in thefoothills of the Alps

Historical and actual impacts and actual situation
Cultivated as early as in the 18th or beginning of the 19th century(“law of derelict land”); partly peat cutting for land owners’
purposes; today mostly intensive agriculture (corn, potatoes orgrassland)
Heterogenous character of more or less intact centres of the raised
bog part (however, often affected by drainage), large areas of peatmining (formerly by peat cutting, today by peat milling); along theborders, agricultural use (formerly mostly litter meadows once cut
only in autumn, today intensive grasslands cut three to five times
per year)
Large numbers of this type, therefore relatively many more or lessintact but also intensively agriculturally used (or left fallow);
ombrogenous peatlands often mined at the beginning of the 20th
century, mostly through peat cutting by hand or machine; borders
mostly cultivatedFormerly used as litter meadows, today drained and intensively
used as grasslands or abandoned, resulting in an overgrowth byshrubs such as Frangula alnus

Natural Revegetation and Restoration of Drained and Cut-over Raised Bogs in Southern Germany“Wendlinger Filz”)?
(2) Is rewetting management of drained and peat-cut
areas useful/successful (case study “Wurzacher
Ried”)?
(3) How useful/successful is restoration managementof milled areas not only through rewetting but
also through sowing and planting (case study“Kendlmühlfilze”)?
To answer these questions, we installed permanentplots (1) in abandoned peat-mining sites in whichobservations were made in 1986 and 2005 or 2006, (2)in a peat-mined area in 1992 before initiation of rewetting management, which started in 1993, with observations repeated in 2002 and (3) in large peat-milled fields
where sowing, planting and transplanting of peatforming plants were applied and vegetation development was monitored from 1987 to 1994.
As a measure of restoration success we use thereestablishment of peat or peat-forming vegetation or
species, since this parameter is connected to all the ecological functions of mires mentioned above – accumulation, disposal, retention, conservation and archive. Peatforming species in ombrotrophic mires in the foothills of
the Alps are the peat mosses Sphagnum magellanicum(up to 90% of the ombrogenous peat), S. papillosum,
S. capillifolium, S. cuspidatum (all less than 10% of theombrogenous peat) and the vascular plant Eriophorum
vaginatum (Paul & Ruoff, 1932). Under minerotrophic
conditions, sedges and other members of the family
Cyperaceae as well as “brown mosses” may generatepeat.

2. Study Sites, Restoration Management andMonitoring
2.1 Study sites without restoration management
after drainage and peat mining
The “Wendlinger Filz” is situated in southeastern
Bavaria about 40 km east of Munich (Fig. 1). It is araised bog in a depression on the end moraine of a
glacier of the last ice age which probably developed
from melting ice blocks. Peat was extracted only after
the Second World War by local farmers who milled thesurface to produce litter for cattle kept in stables (Fig.
2b). This was done until the 1960s. Very few areas remained under peat extraction until the mid 1980s. Afterabandonment, the open peat surface remained bare with
only slow establishment of a few dominant species(Poschlod, 1990). Seven transects along which permanent plots (4 m2 in size) were irregularly arranged were
established in 1986. Vegetation and hydrological parameters were recorded at each permanent plot in 1986 and2006. Additionally, all transects were levelled, peat
profiles were taken and peat composition studied bymacro-remains analysis in 1986.
The “Wieninger Filz” is also situated in southeastern
Bavaria near Traunstein (Fig. 1). Like the “Wendlinger
Filz,” it is one of the typical small raised bogs at theborder of an end moraine of the last ice age. In contrast

S

BadenWuerttemberg

207

Bavaria
M

W

K Wi
We

●

Fig. 1 Location of the study sites ( ) in Baden-Wuerttemberg
and Bavaria (Southern Germany).
K – Kendlmühlfilz, W – Wurzacher Ried,
We – Wendlinger Filz, Wi – Wieninger Filz,
S – Stuttgart, M – Munich.

to the “Wendlinger Filz,” peat extraction here was doneby hand cutting since the beginning of the 20th century
as fuel for the brewery “Wieninger.” The upper rooted
horizon of newly mined areas was deposited into thearea which had been exploited (Fig. 3a) so that most of
the typical raised-bog species were present on theformerly mined area after its abandonment. They
persisted either as vegetative parts or in the diasporebank (Poschlod, 1990, 1995). In 1960, peat-mining
ended. No management has occurred since, except thatthe central ditch was dammed during the winter season
to provide an open water surface in winter for the
traditional Bavarian winter sport “ice stick shooting”(Poschlod, 1990). In late summer of the following year
the down was opened to allow cutting of reeds.
The peatland was mapped first in 1986 along threelevelled transects (Poschlod, 1990). Peat profiles weretaken every 20 to 50 m. Permanent plots (4 m2 in size)
were irregularly arranged along these transects, accompanied by hydrological measurements (water level,water quality). Vegetation and hydrological parameters
were recorded in 1986 and 2005 at these permanentplots.
2.2 Study sites with restoration management afterdrainage and peat mining
The “Wurzacher Ried” is situated in Upper Swabia,
south of Ulm (Fig. 1). It represents one of the largest
raised bog complexes in southern Germany. Growth ofthe mire started after the last ice-age from a shallow lake
in a basin closed by the glacier’s end moraine. Originally, the mire consisted of seven raised bog shields (Fig.
4a; Schwineköper et al., 1991). Drainage and local
peat-mining started in 1730. In 1880, industrial peatmining by means of cutting started and was intensified
after the First World War in 1920 when fuel was limited.
In 1946, a glass industry was established in Bad

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a

b

c

d

Fig. 2 Wendlinger Filz: a – location of the milled fields (grey) in the peatland (black line = border ofthe peatland), red line = Transect 2; b – actually milled field (1985); c – milled fields at theeastern part of Transect 2 in 1986 with dominant Calluna vulgaris; d – milled fields of theeastern part of Transect 2 in 2006 with dominant Rhynchospora alba.

abFig. 3 Wieninger Filz: a – diagram of peat cutting (from Poschlod, 1996); b – peat forming vegetation in a former peat-cut
area in 2005 with Eriophorum vaginatum and Sphagnum papillosum (Permanent Plot 3).

Wurzach which was followed by further intensificationof peat mining (Gremer & Poschlod, 1991). However,despite several trials the different owners of the peatland
did not succeed at draining the entire peatland due to its
situation in the basin (Schwineköper et al., 1991; Krüger& Pfadenhauer, 1992). Today, only two of the sevenraised bog shields are left, with the largest central shield
being partly affected by peat mining and drainage. However, the largest part of the central shield remained unaffected, although a road has been cut through it (Fig. 4b).
Although peat-mining continued until 1995, part of the
Wurzacher Ried became a nature reserve in 1959 (morethan 4 km2). The protected area was increased in 1981 toa size of nearly 14 km2 and in 1996 to more than 18 km2.
In 1989, the Wurzacher Ried was awarded the European
Diploma, and a detailed plan was developed. One
measure of this management plan was the rewetting of
the drained and mined areas of the central shield.

Rewetting management was initiated in 1993. Monitoring of permanent plots (4m2 in size) that were regularly
arranged along three transects (Gremer & Poschlod,1991) and ran from the peat-mined to the drained part of
the raised bog shield was carried out in 1992, before theinitiation of the management, and in 2002. Hydrologicalparameters along these transects were measured to gain
baseline data for the management plan. After rewetting,hydrological measurements were only carried out in
ditches which were blocked.
The “Kendlmühlfilz” is one of the largest raised-bog
complexes in Bavaria, situated in a basin south of Lake
Chiemsee (Fig. 1). The basin was formed by the
Inn-Chiemsee Glacier during the last-ice age. Today, thepeatland represents a complex of arable fields, grasslands and afforested areas at its borders, with peatmining areas and a remaining part of the former raised
bog shield, which is partly affected by drainage

Natural Revegetation and Restoration of Drained and Cut-over Raised Bogs in Southern Germany(Fig. 4c).
Until the end of the 19th century peat was cut locallyor extracted for litter by farmers. Large-scale drainage
started only in 1885. Peat was cut for industrial purposes,
namely for salt production. After the First World War,peat was extracted from an area of 2.4 km2 by theBavarian peat company for fuel, first still by hand but
later by machine. In 1976, a private company started
peat extraction in an area of about 0.4km2 by milling for
horticultural purposes. Peat mining stopped in 1985
(Pfadenhauer et al., 1990).
Restoration of the milled areas, where ombrogenous
peat layers about one meter high remained, started
in 1986 by surface re-configuration of the nearly
1.5 km-long milling fields into polders, a method
described in detail by Eggelsmann (1987). Afterrewetting the polders, reestablishment of typical mirevegetation according to the peat and water quality in thepolders was initiated by planting and sowing specificvascular plant species in 1987 (Maas & Poschlod, 1991).
Later on, fertilizer was applied to accelerate vegetation
development, and Sphagnum species were reintroducedby transplanting small pieces of Sphagnum lawns (Sliva,
1997). The success of restoration management practices
was monitored in permanent plots of 100 m2 (vascularplants) and 1 m2 size (bryophytes) which were situatedalong a transect running along Field 6S (Fig. 8). Here,
we present only results of sowing and planting selectedvascular plants in the summer of 1987 (Carex rostrata,

209

Eriophorum angustifolium; only sown: E. vaginatum)and introduction of Sphagnum (S. magellanicum,
S. capillifolium – hummock/lawn species; S. cuspidatum– hollow species) sods in summer of 1992. For theentire experimental approach see Maas and Poschlod(1991) and Sliva (1997).
2.3 Vegetation and hydrological monitoring
Vegetation was recorded according to the dominancescales of Schmidt or Braun-Blanquet (Pfadenhauer et al.,1986). Nomenclature follows Ehrendorfer (1973) for
vascular plants and Frahm and Frey (1983) for bryophytes except for Sphagnum spp., which follow Daniels
and Eddy (1985).
Hydrological parameters were measured using plastic tubes of 1 or 2 meters length (Poschlod, 1990).
Measurements of water levels and water chemicalparameters such as pH and ions were carried out
monthly from May to September during 1985 in the“Wieninger Filz” and “Wendlinger Filz.” In 2005, measurements in the Wieninger Filz were made only every
other months in the same period. In 2006, hydrologicalparameters in the “Wendlinger Filz” were measured
only once, in September. To compare data from thedifferent years we used data from these months where
measurements were made in 1986 and 2005 and 2006,
respectively. Since in the “Wurzacher Ried,” measurements of water level and water chemical parameters
before and after rewetting were carried out at different

Fig. 4 Wurzacher Ried: a – historical situation (horizontal lines – ombrotrophic parts (raised bog shields), horizontal broken lines –minerotrophic parts); b – current situation (air photograph from Schuckert et al., 1997; c – unaffected part of the central
raised bog shield cut through by the road; m – mined areas of the central raised bog shield).
Kendlmühlfilz: c – infrared coloured aerial photograph from the central part in 1986: in the south the more or less unaffectedpart of the raised bog shield, in the north peat-mined areas (red: by peat cutting until 1971, black: by peat milling from 1976
to 1985).

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sites (see above), they were not evaluated for this paper.
In the “Kendlmühlfilz,” data on hydrology have alreadybeen presented by Sliva (1997). They will, therefore,
only be used for the discussion. Vegetation development
was analysed by applying a variance analysis tocompare changes in the dominance of single species
followed by a Mann-Whitney-U-test, available in
WinStat, and multivariate statistical methods, available
in the software package PC-Ord (McCune & Mefford,
1999) to compare changes in the vegetation.

3. Results
3.1 Sucession in milled areas without restoration
management – the “Wendlinger Filz” case study
Vegetation within the milled areas developed
towards mono-dominant stages of vascular plant species.
The water level as well as pH, which depended on theremaining peat layer, determined vegetation development. In 1986, Eriophorum vaginatum was dominant at
wet (annual average water level between 10 to 0 cm
below surface) and acidic sites, Rhynchospora alba at
wet acidic to subneutral sites and Carex rostrata,
Eriophorum angustifolium or Phragmites australis at
wet or flooded sites with subneutral to neutral pH. On
all dry sites (average water levels lower than 10 cm
below the surface), Calluna vulgaris dominated(Poschlod, 1990; Fig. 2c). In 2006, no changes could beobserved. The same species were still dominant at therespective sites and no Sphagnum species had invaded
except at flooded sites with Carex rostrata where
Sphagnum cuspidatum became dominant and formed aclose layer. No peat growth could be observed. However,many of the sites formerly dominated by Calluna
vulgaris were invaded by Rhynchospora alba, the latterbecoming the more dominant species despite the factthat water levels remained the same as in 1986 (Figs. 2d,
5).
3.2 Succession in peat-cut areas without restoration
management – the “Wieninger Filz”case study
Vegetation development in the “Wieninger Filz” differed strongly from that in the “Wendlinger Filz.” In1986, non-mined areas were affected by drainage whichbecame more obvious in 2005 when the dominance ofindicators of drained ombrotrophic peatlands such as
Pinus mugo or Calluna vulgaris increased. Dependingon the water level and water and peat quality, peat forming vegetation was already established in 1986, eitherwith Eriophorum vaginatum and Sphagnum papillosum
or with Scheuchzeria palustris, Sphagnum papillosum,
S. cuspidatum and S. angustifolium. In 1986, fresh peat
layers of more than 80 cm could be observed. Due tocontinuing peat growth only a few permanent plots withpeat forming vegetation (Fig. 6 – Permanent Plots 1a, 1b,
3 and 4) could be found in 2006. In the case of Permanent Plots 1a and 1b, where apart from Eriophorum
vaginatum, Sphagnum angustifolium was also dominant,there was no increase in the peat layer. However,in Permanent Plots 3 (Fig. 3b) and 4, Sphagnum
magellanicum and especially S. papillosum contributed
to further peat growth of 28 and 30 cm, respectively,
since 1986.
3.3 Succession in peat-cut areas after rewetting
management – the “Wurzacher Ried” case study
Rewetting management of the peat-mined areas in
the Wurzacher Ried caused not only raised water levels
within drained as well as mined areas, but extensive
flooding of the peat-cut areas. It was followed by large

Fig. 5 Wendlinger Filz: Ordination of the permanent plots which were classified in 1986 as stages dominated by Calluna vulgaris(DCA, main matrix: 12 plots, 27 species; second matrix: 12 plots, 27 species [highly correlated species, red; R² > 0,30] and4 abiotic parameters [WLev = water level; pH; Cond = conductivity; Ca = Calcium content mg/l water]; however, R² for all
abiotic parameters was < 0,30).
Pic abi = Picea abies, Cal vul – Calluna vulgaris, Vac uli – Vaccinium uliginosum, Eri vag – Eriophorum vaginatum,
Rhy alb – Rhynchospora alba, Dic cer – Dicranella cerviculata, Leu gla – Leucobryum glaucum.

Natural Revegetation and Restoration of Drained and Cut-over Raised Bogs in Southern Germany

211

Fig. 6 Wieninger Filz: Ordination of selected permanent plots recorded in 1986 and 2005, abiotic parameters and plant species
correlated (DCA, main matrix: 22 plots, 65 species; second matrix: 22 plots, 65 species [highly correlated species, red; R² > 0,30]
and 4 abiotic parameters [blue; WLev = water level; pH; Cond = conductivity; Ca = Calcium content mg/l water]; R² > 0,30);
green circle – peat forming vegetation at high water level (WLev) in former peat-cut areas with Eriophorum vaginatum (Eri vag),
Sphagnum magellanicum, S. papillosum, S. angustifolium a.o.; red circle – vegetation at low water level with dominant Pinus
sylvestris, Picea abies and Calluna vulgaris in the peat-cut area (Permanent Plots 2a and 2b) or with dominant Pinus mugo and
Calluna vulgaris on drained sites in the unmined area; brown circle – alder carr vegetation with Alnus glutinosa (Aln glu),
Frangula alnus (Fra aln), Typha latifolia (Typ lat), Angelica sylvestris (Ang syl), Caltha palustris (Cal pal), Lycopus europaeus(Lyc eur), Peucedanum palustris (Peu pal) a.o.

vegetation changes: however, they were not on the peatridges between the peat-cut areas, but in the peat-cut
areas, where a large-scale die-off of trees, dwarf shrubs
and other indicators of drainage or degradation such as
Molinia caerulea was noticeable due to flooding (Table2). Reestablishment of or increase in peat-forming
raised-bog species such as Eriophorum vaginatum andSphagnum species occurred only through the establishment of floating mats (Fig. 7, Table 2). In contrast,
Sphagnum magellanicum lawns which had already been
established before the water level rose, disappeared after
rewetting due to flooding (Table 2). Where peat wasmined into the geogenous peat layers or the former
drainage ditches touched the mineral underground,peat-forming fen species such as Carex rostrata or
Phragmites australis became established or spread(Table 2).
3.4 Succession on milled areas after rewetting
management and reintroduction of vascularplants and Sphagnum spp.–the “Kendlmühlfilz”
case study
Establishment of success differed between sowing
and planting vascular plants, planting being more successful than sowing (Fig. 8a). Planting of Carex rostrata
achieved a cover of 100% in four years after planting
and it managed to become established along nearly thewhole transect, except in the middle of the first half andat the end of the second half of the transect during themonitoring period (Fig. 8a). Eriophorum angustifolium

became established after sowing and planting except on
the first planting plot. In contrast to Carex rostrata, its
occurrence remained restricted to the sown and plantedplots even seven years after introduction (Fig. 8a).
Eriophorum vaginatum became established only on thesown plot but was more successful in establishing itself
in the first half of the transect, where no introduction
management took place (Fig. 8a). Transplantation success of Sphagnum sods differed strongly as well.
Irrespective of their ecology, all three species showedsimilar behaviour within the same plots. Sods increased
in diameter and cover in five plots (Fig. 8b) but died infour other plots.

4. Discussion
4.1 Spontaneous revegetation
Our comparative analysis of peat-cut and peat-milledareas without restoration management clearly shows thatthe peat-mining technique affects the spontaneous reestablishment of peat-forming vegetation. Peat-forming
vegetation became established only in peat-cut areas but
not in milled areas, as was already stated by Poschlod(1990) in a analysis of nine peatlands in the foothills of
the Alps. Monitoring of the formerly mined peatforming areas in the Wieninger Filz indicates that inpeat-cut areas (1) the process of peat-formation iscontinuing and (2) that the accumulation rates areenormous, about 30 cm in 20 years, confirming theassumptions of Poschlod (1990). In the Wendlinger Filz,

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Table 2 Significant changes in dominance of potential peat forming species in peat-mined areas of the Wurzacher Ried after
rewetting management (Indicator species: first group – peat-forming bog species; second group – peat-forming fen
species; third group – species of dry/degraded sites; Habitat: PR – un-mined peat ridges between the peat-cut areas;
PCA – peat-cut areas; Significance (Mann-Whitney-U-test; p < 0.05): ↑ – increase; ↓ – decrease; - no significance).
Indicator species
Eriophorum vaginatum
Sphagnum magellanicum
Sphagnum papillosum
Sphagnum capillifolium
Sphagnum cuspidatum
Carex rostrata
Phragmites australis
Trees and dwarf shrubs(Betula spec.; Vaccinium myrtillus;
V. vitis-idaea a.o.)Molinia caerulea

Habitat
PR
PCA
PR
PCA
PR
PCA
PR
PCA
PR
PCA

Significance
↑
↓
↑
↑

PR
PCA
PR
PCA
PR
PCA

↑
↑
↓

Remarks
Establishment of floating mats
Decrease due to flooding
Establishment of floating mats
Highest increase of all peat-forming species;
establishment of floating mats
Increase due to flooding
Increase due to flooding
Decrease due to flooding

PR
PCA

↓

Decrease due to flooding

Eriophorum vaginatum – 1992

Eriophorum vaginatum – difference between 1992 and 2002

Sphagnum magellanicum – 1992

Sphagnum magellanicum – diff. 1992 and 2002

Fig. 7 Wurzacher Ried: Transect levelling profile, location of the permanent plots and cover (% – y-axis) in 1992 as well as changes
in cover (%) of Eriophorum vaginatum and Sphagnum magellanicum from 1992 to 2002.

however, the original peat forming vegetation has notreestablished even more than 40 years after mining
activities ended of mining.
Studies in North America recently reported by
Lavoie et al. (2003) and Poulin et al. (2005) have also
shown that Sphagnum establishment occurs only inblock-cut (= peat-cut) peatlands whereas at milled sites
only vascular plants were able to establish. Lavoie et al.
(2003) saw as a positive example the invasion of cotton
grass (Eriophorum vaginatum) as a positive development which in our opinion, however, represents no

success, as 20 years of succession at the Wendlinger Filz
were insufficient for the establishment of Sphagnum
species in a successional stage of Eriophorum vaginatum.
Not even has the cover of Eriophorum vaginatum increased from 1986 to 2006. The only effect which could
be measured was the invasion of Rhynchospora alba
in formerly Calluna vulgaris-dominated successional
stages. In all milled and vacuum harvested areas,
Sphagnum could reestablish only under flooded conditions. After 20 years, these Sphagnum mats stillcontained only Sphagnum cuspidatum or in rare cases

Natural Revegetation and Restoration of Drained and Cut-over Raised Bogs in Southern Germany

representatives of the Sphagnum recurvum group (S.
recurvum var. mucronatum, S. angustifolium). However,
the typical peat-forming bog species have not yet
invaded the Sphagnum cuspidatum mats and, therefore,
peat growth towards raised bog communities has not yet
occurred. This confirms hypotheses that all mining
practices that result in bare peat areas cannot beassumed to enable spontaneous revegetation towards theformer peat-forming raised-bog vegetation, at least notin southern Germany, but this has also been noted inother countries. Price et al. (2003) state that suitable locifor Sphagnum recolonization are absent at milled andvacuum-harvested sites. They attribute this fact to theloss of capillary water flow to the surface, resulting inquick desiccation of the surface peat layers and to thehuge extent of the mined sites which decreases thelikelihood of the arrival of plant propagules. However,there are more causes for non-successful spontaneous
re-vegetation of milled sites after abandonment. One of
the key factors is not only dispersal, but alsoestablishment and growth of the respective Sphagnum
species (Rochefort, 2000; Chirino et al., 2006). Neither
dispersal of spores nor establishment from spores havebeen observed so far, which is not surprising e.g. for thea213main peat- forming species in the foothills of the Alps,
Sphagnum magellanicum. Records of individuals with
spore capsules are extremely rare (Poschlod, 1990).
In contrast, vegetative fragments may be dispersed(Poschlod, 1995) and would be able to regenerate a new
Sphagnum plant (Poschlod & Pfadenhauer, 1989;
Spencer-Famous & Taylor, 2005). However, reestablishment of larger patches of peat-forming species at nonflooded sites was successful only after active reintroduction (Sliva, 1997; see below).
Conditions for successful spontaneous reestablishment of peat-forming vegetation in peat-cut areas havebeen summarised by Poschlod (1994; Table 3). The
main factor is the conservation of the surface layer as avaluable source for peat-forming species either vegetatively or in the propagule bank (Rhynchospora alba,
Sphagnum spec. a.o.); the latter, however, is maintained
only when stored under moist or wet conditions(Poschlod, 1989, 1990, 1995). Furthermore, the top spit
may be a more suitable microsite for germination andestablishment than a bare milled peat surface (Roderfeld,
1992; Roderfeld et al., 1993). Successful germinationand establishment in milled areas depend on seed sizeand growth rate, limiting the recolonization potential ofb

Carex rostrata

Sphagnum magellanicum

Eriophorum angustifolium

Sphagnum capillifolium

Eriophorum vaginatum

Sphagnum cuspidatum

Fig. 8 Kendlmühlfilz: a – development of cover of selected vascular plant species along the transect through milled field 6S from1988 to 1994. Outlined arrow: sown plots, solid arrow: planted plots; b – development of the diameter (cm; outlined dot) andcover (%; solid dot) of transplanted Sphagnum sods at site J (Sliva, 1997).
Table 3 Comparative analysis of two peat-mining techniques – peat cutting and peat milling (vacuum mining) – from a restoration
point of view.
Peat mining by cutting
Vegetation (rooted) horizon (or surface layer, top spit) is
maintained (potential for regeneration – whole plants,
seeds, spores and bud banks)
Mining vertically (1.5 to 3 m per year) → mined area isrelatively small
Mined area will be abandoned after one year – possibilityof fast regeneration also from adjacent areas
Mining can also be done “under water” – re-wetting
management in many cases not even necessary

Peat mining by milling / vacuum extraction
Vegetation (rooted) horizon is destroyed
Mining horizontally (vertically only 6 to 8 cm per year) → mined area isextremely large
Mining of the same area over several decades depending on the depth – nopossibility of fast regeneration because of the large mined areas anddestruction of most of the adjacent areas during the mining period
Mining only under dry conditions possible – huge efforts for rewetting
management necessary (including the construction of polders to minimize
erosion)

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P. POSCHLOD et al.

typical raised-bog species (Campbell & Rochefort,2003).
4.2 Restoration management
As the case study “Wurzacher Ried” shows, rewetting alone will not be followed by the reestablishment ofpeat-forming vegetation, at least not in the short or
medium term, although succession after peat-mining but
before local rewetting management leads to the reestablishment of the original peat-forming vegetation. Due tothe extremely heterogeneous surface of the peat-minedarea with un-mined ridges and mined areas of up to
more than four meters in depth (Fig. 7) large parts of thepeat-mined areas were flooded. As Rochefort et al.
(2002) have shown, only shallow and short-term flooding may enhance the growth of most Sphagnum species.
Nevertheless, according to Schouwenaars (1982) andBeets (1993), the establishment of floating mats as insome of the peat-cut areas of the Wurzacher Ried may
initiate the establishment of peat-forming vegetation,
however, mostly of fen species. Some bog or transition
mire species such as Eriophorum vaginatum andSphagnum papillosum may use these floating mats toincrease their dominance (Beets, 1993).
Due to the slow recolonization of milled bare peatareas, sowing and planting of vascular plants as well as
reintroduction of vegetative Sphagnum propagules oreven transplanting Sphagnum sods were recommended
to initiate re-vegetation (Maas & Poschlod, 1991). As
already mentioned above, sowing was not successful but
planting, especially of clonal plants with long rhizomes,
has been a successful tool (Fig. 8, see also Sliva, 1997;
Sliva & Pfadenhauer, 1999). However, an appropriatehydrological management plan including blocking
ditches, constructing bunds and reconfiguring the surface is a prerequisite for applying these methods (Priceet al., 2003; Campeau et al., 2004). Although hydrological management was carefully planned in the case of theKendlmühlfilz, the reconfiguration of the surface resulted in upwelling of mineral water and accumulationof peat mud in some of the polders, followed by invasion of Phragmites australis and Typha latifolia but also
Carex rostrata. Nevertheless, in most of this area otherspecies have become established, Eriophorum vaginatum,
E. angustifolium and Carex rostrata being the mostsuitable to cover rapidly the bare peat areas. Transplantsof Sphagnum sods did not spread at all sites and manyeven died. If spreading occurred it was extremely slow
(Sliva, 1997; Sliva & Pfadenhauer, 1999). Therefore,
this technique will rarely be an option. Rochefort et al(2003) applied another technique, the spreading ofvegetative parts of Sphagnum spp. (collected from anatural bog) as a thin layer on bare peat surfaces. Afterthey were spread, the Sphagnum propagules are covered
with straw mulch, including phosphorus fertilization,
resulting in a low Sphagnum cover after three years.
Despite these first successful steps in Sphagnum
establishment, reestablishment of Sphagnum carpets still
remains a big challenge in bare peat areas.

4.3 Conclusions
The application of specific peat-mining techniques
determines the direction of spontaneous revegetation
after mining activities end. We compared revegetation ofabandoned areas mined by cutting and milling. However,there are more techniques which are applied throughout
temperate regions worldwide. Future work, therefore,should include comparisons of all techniques in order tounderstand the factors that influence spontaneous revegetation. Furthermore, it might also be useful not to
aim for the regeneration of pristine mires, but for theregeneration of certain functions as mentioned in the
introduction. E.g., Vasander et al. (2003) state that after
restoration, mined peatlands may return to a functional
status close to that of pristine mires within a few years,
taking into account, however, only the carbon-sink function. Some successional stages of milled areas where thewater level is at the surface may serve as suitable
habitats for many hollow species such as Drosera spp.
or Lycopodiella inundata, all species of the Red Data
Book of Germany (Korneck et al., 1996).
The above-mentioned aspects should be included ina new edition of “Wise use of mires and peatlands”(Joosten & Clarke, 2002). Restorability of ecosystems
is one of the central criteria in nature conservation(Riecken, 1992). However, this aspect is rarely considered before ecosystems are altered. Principles 5 and 6of the Convention on Biological Diversity state (5)“conservation of ecosystem structure and functioning, inorder to maintain ecosystem services, should be apriority target” as well as (6) “ecosystems must be
managed within the limits of their functioning,” but thishas been, until now, completely ignored in the case ofmires. Furthermore, Principle 11 says, “the ecosystemapproach should consider all forms of relevant information, including scientific, indigenous and local knowledge, innovations and practices.” However, since thisknowledge does not yet exist with respect to therestoration of mined raised bogs, such a principle does
not help. Even if the knowledge existed, who would be
responsible for collecting and reviewing it? This is in
our opinion another dilemma of the scientific community, that textbooks or handbooks which summarize
and review the state of art on actual problems are rarely
supported by independent funding institutions. We
should also take into account that mires or peatlands areindividual entities and existing knowledge on one mire
type or in one region cannot be transferred to anothermire type or another region or even to the same mire
type in the same region. As Price et al. (2003) state forpeat-mined areas: “No standard management prescription can be made, because each site presents uniquechallenges.” Therefore, the establishment of a databasewith data on peat harvesting techniques, long-termmonitoring research and scientific findings on the results of spontaneous and directed revegetation processesor restoration management would be helpful in deciding
on appropriate mining techniques when the aim after
peat mining is the reestablishment of the functions of a

Natural Revegetation and Restoration of Drained and Cut-over Raised Bogs in Southern Germany

pristine mire. However, it should be also clear to everyone working on the restoration of peatlands thattheoretical (scientific) considerations are often not suitable to be implemented into practice. This is especiallytrue for hydrological management practices, as the studies at the “Wurzacher Ried” and the “Kendlmühlfilz”
have shown.
Finally, further efforts should be undertaken to look
for alternatives to Sphagnum peat for its respective purposes (Schmilewski, 2001) and also for ways of
producing Sphagnum as a renewable resource, either inpeat-mining areas or even outside of mires and peatlands (Gaudig & Joosten, 2002; Grantzau & Gaudig,
2005; Krebs & Gaudig, 2005).

Acknowledgements
We thank the governments of Bavaria and BadenWürttemberg as well as all institutions which wereinvolved in these studies for their support. Special
thanks go to Prof. Jörg Pfadenhauer, Technical University of Munich, who initiated more than 20 years ago thefirst studies on wetland restoration; to the nature conservation centre in Bad Wurzach, which has been ensuring that continuous monitoring is taking place after theestablishment of rewetting management; and finally toKathrin Bylebyl, who corrected the English version of
the text.
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