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Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
1
Substrate quality and spontaneous revegetation of extracted peatland:
case study of an abandoned Polish mountain bog
E. Zając1, J. Zarzycki2 and M. Ryczek1
1Department of Land Reclamation and Environmental Development,
2Department of Ecology, Climatology and Air Protection,
Faculty of Environmental Engineering and Land Surveying, University of Agriculture in Kraków, Poland
_______________________________________________________________________________________
SUMMARY
If peatland is left without any restoration treatments after mechanical peat extraction ceases, the process of
secondary transformation of peat continues. The resulting changes in peat properties severely impede the
recovery of vegetation on cutover peatland. The aim of this study was to assess how secondary transformation
of peat affects spontaneous revegetation, and the relative importance of different factors in controlling the re-
establishment of raised bog species on previously cutover peat surfaces. The study was conducted on two
sectors of a raised bog in southern Poland where peat extraction ended either 20 or 30 years ago. Where the
residual peat layer was thin (~ 40 cm or less) and the water table often dropped into the mineral substratum,
the development of vascular plants (including trees) was favoured, and this further promoted the secondary
transformation of peat. In such locations the vegetation tended towards a pine and birch community.
Revegetation by Sphagnum and other raised bog species (Eriophorum vaginatum, Vaccinium uliginosum,
Ledum palustre, Oxycoccus palustris) was associated with thicker residual peat and higher water table level
which, in turn, were strongly correlated with hydrophysical properties of the soil. A species - environmental
factor redundancy analysis (RDA) showed that any single factor (of those considered) was not important in
determining the revegetation pattern, because of their intercorrelations. However, water table level appeared
to be the most important abiotic factor in determining the degree of soil aeration and, consequently, the stage
of secondary transformation attained by the peat.
KEY WORDS: peat extraction, cutover peatland, secondary transformation, peat quality
_______________________________________________________________________________________
INTRODUCTION
Peatlands are globally important ecosystems with
multiple roles in the natural environment. They serve
as water and carbon reservoirs and are known for
their specific biodiversity (Minayeva et al. 2017).
They are also natural archives of data on
palaeoenvironmental, climatic and hydrological
changes as well as human impact (e.g. Joosten &
Clarke 2002, Chambers & Charman 2004). Since the
beginning of the 19th century, the global area of mires
and peatlands has declined significantly due to
climate change and human activities, especially
drainage for agriculture and forestry. The largest
losses have occurred on the European continent,
where the remaining mire area is about 52 % of the
former extent of mires (Joosten & Clarke 2002). In
Poland (central Europe), mires cover about 0.6 % of
the country and their loss in relation to former extent
is estimated at 84 % including about 4 % of peatland
degraded by peat extraction (Bragg & Lindsay 2003).
It is especially important to restore extracted areas on
former raised bogs because only 4.4 % of all the
mires in Poland belong to this type (Ostrowski et al.
1995).
The first requirement for mechanical extraction of
peat is drainage, which is usually provided by open
ditches at about 20 m spacing that are gradually
deepened as peat extraction proceeds. The next step
involves removal of the acrotelm, which is a singular
hydrologically self-regulating component of a natural
peat bog (Ingram 1978). Thus, peat extraction
drastically affects both the peat and the vegetation,
and some of the changes are irreversible.
After removal of the acrotelm the denser (more
decomposed) peat of the catotelm is exposed, and this
in turn undergoes changes. The transformation of
peat under the aerobic conditions imposed by
lowering of the water table is known as secondary
transformation (primary transformation takes place
during the initial peat formation process) (e.g. Kalisz
et al. 2015). The effects of drainage on the physical
properties of peat are shrinkage, compaction,
oxidation and consequent peat subsidence
(Eggelsmann 1986, Lipka et al. 2017). Following
drainage, bulk density increases and pore spaces
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
2
decrease in size (Price 1997); soil moisture content,
specific yield and hydraulic conductivity decline; and
the amplitude of water level fluctuations increases
(Price et al. 2003, Van Seters & Price 2001). The
water retention capacity of peat is enhanced by
compaction, but the availability of water to non-
vascular plants may become limited (Price &
Whitehead 2001). Intensified oxidation of the
organic matter may result in nutrient regime changes
in the peat and groundwater (Wind-Mulder et al.
1996, Andersen et al. 2010) and may significantly
enhance acidity (Juckers & Watmough 2014). It also
results in a rise in CO2 emissions from cutover
peatland (Wilson et al. 2013).
When peatlands are left without any restoration
treatments (i.e. ‘abandoned’) after peat extraction
operations are terminated, spontaneous revegetation
may be observed. The revegetation process is
influenced by multiple factors, the most important
being the degree of damage to the peat bog, the
thickness of residual peat and its nutrient content, the
water level and water sources, the surface
topography, and the time that has elapsed since peat
extraction ceased (Wheeler & Shaw 1995).
Conditions may vary widely between and within
sites, and this affects the rate and pattern of
revegetation. The most desirable outcome is
reinstatement of the original plant community
including, in the case of a raised bog, an acrotelm
composed mainly of Sphagnum mosses. However,
extracted areas are hostile and highly challenging
habitats for recolonising mire plants, mainly because
of poor water availability, exposure to desiccation
and erosion, lack of diaspores (Quinty & Rochefort
2003), and high acidity accompanied by low nutrient
content in the cutover surface peat (Salonen 1994).
Moreover, Sphagnum species are particularly
sensitive to difficulties of acquiring water by transfer
from the finer-textured cutover peat (McCarter &
Price 2015).
Early recognition of the potential for regeneration
of a specific site may be helpful when planning
ecological restoration activities aimed at encouraging
the development of target plant communities by
modifying environmental conditions (Campbell et al.
2000). In order to determine the local potential for
spontaneous revegetation, it is essential to know
which factors are important for species typical of
raised bogs. We hypothesise that the secondary
transformation of peat after drainage results in
changes in quality of the uppermost soil layer which
might in turn affect the establishment of bog species.
The water-holding capacity index W1 proposed by
Gawlik (e.g. 1992) is a quantitative characteristic of
water retention by the soil that reflects changes in the
physical, hydrophysical and chemical properties of
peat when subjected to drying (Gawlik 2000,
Sokołowska et al. 2005), and can be used to
quantitatively assess the secondary transformation
stage of peat.
The aim of this study was to investigate: 1) how
secondary transformation of peat on an area of
extracted peatland affects revegetation; 2) the relative
importance of a range of factors in controlling the
establishment of raised bog species; and 3) whether
the W1 index, as a single measure of secondary
transformation, may be a useful predictor of
revegetation trajectories for extracted areas.
METHODS
Study site
The research was carried out on the Bór za Lasem
bog, which is located in Czarny Dunajec commune in
southern Poland (Figure 1). It is one of a group of 27
peatlands belonging to the European Ecological
Network Natura 2000. These bogs were formed
within the Orava - Nowy Targ Basin, which is a
depression flanked to the north and south by
mountain ridges. The climate of the basin is
moderately warm with some local peculiarities
(Kondracki 2011). For the part with bogs, Olszewski
(1988) gives a mean annual air temperature of
+ 5.5 °C (highest and lowest monthly means: + 16 °C
for July, - 6 °C for February) and total annual
precipitation 750–825 mm, which is considerably
less than in the surrounding higher-altitude areas but
much greater than in the Polish lowlands. Basic
weather characteristics were recorded in 2016 with a
Davis Vantage Pro 2 weather station located near the
study site (49° 25' 31.33" N, 19° 48' 42.24" E;
Figure 1). At this location, mean daily air
temperature between June and August (2016) was
15.4 °C, with daily maximum 37.0 °C and daily
minimum -2.9 °C. The total precipitation recorded
during the same period was 375 mm.
The Bór za Lasem bog was initiated by
paludification of a sparingly permeable clay
substratum under the influence of shallow
groundwater, and subsequently developed into a
raised bog. This is reflected in the peat stratigraphy
by the presence of strongly decomposed and
transitional (poor-fen) Sphagno-Cariceti peat near
the base, overlain by Eriophoro-Sphagneti and
Eusphagneti peats formed under conditions of
ombrogenous water supply. Average degree of peat
decomposition within the deposit, determined by a
microscopic method (e.g. Tobolski 2000), is 30 %
and average ash content is 2.2 %. Average thickness
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
3
Figure 1. Map of the Bór za Lasem peat bog showing the two investigated sectors of the extracted area
(adapted from Łajczak 2006) (a), and location map (b).
of the deposit is about 1.8 m, and its maximum
thickness is 3.65 m (Lipka & Zając 2014).
The current area of the bog is 55 ha. Almost the
entire perimeter of the bog dome has been intensively
exploited by local people for centuries. After World
War II, industrial extraction of peat by the block-
cutting method commenced, using heavy machinery.
The extracted area is bisected by a railway
embankment. The northern sector (Sector A, ~16 ha)
is bordered by the railway to the south and by forest
and grassland to the north, while the southern sector
(Sector B, ~ 8 ha) is delimited by the railway to the
north and a surviving part of the bog dome to the
south (Figure 1). Peat was cut on Sector A from the
beginning of the 1960s until the beginning of the
1980s, then operations moved onto Sector B where
extraction continued until the early 1990s (Mr
Bogusław Sroka, Peat Production Plant “Bór za
Lasem” in Czarny Dunajec, personal communication
2016). Thus, peat extraction ceased approximately
ten years earlier in Sector A than in Sector B. The site
was subsequently left untouched. Across both sectors
there is a network of secondary ditches spaced at
about 20 m that discharge water into main ditches.
Most of the secondary ditches are currently
overgrown with vegetation and some of them are
blocked. The main ditches discharge water mainly
during floods associated with major rainfall events
and spring thaws.
Study plots
Studies in the extracted sectors of the Bór za Lasem
bog were conducted during the years 2015 and 2016.
Within each sector, twenty 5 m × 5 m study plots
were set out in a W-shaped transect (40 study plots in
total). The plots were arranged in a regular pattern but
they were placed to avoid ditches, standing water and
dense groups of trees.
Analysis of peat
At each study plot the mean thickness of residual peat
was measured with a soil probe and surface peat
samples were taken for laboratory analysis. The peat
samples were collected from 0–10 cm depth.
Samples of undisturbed structure were collected in
small metal rings (250 cm3, two per plot, 80 in total)
and placed in plastic bags for transport. A further
(disturbed) sample was collected into a plastic bag
from each of the 40 plots and subsequently divided
into two parts for analysis (i.e. all analyses were
duplicated). To determine the type of peat that was
left at the surface when extraction ended, additional
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
4
samples were collected from just below the surface
layer that had subsequently degraded, at eight
randomly selected locations in each sector.
Degree of peat decomposition was estimated by
von Post’s method (von Post 1924). Peat type was
determined by a microscopic method based on plant
macrofossil analysis (Kac et al. 1977, Tobolski
2000). To determine bulk density (ρb) plus
volumetric (θv) and saturated (θs) moisture content,
the samples with undisturbed structure were dried to
constant weight at 105 °C. For θs, the samples were
soaked with water for three weeks before drying, and
gravimetric moisture content was converted to
volumetric basis using bulk density. Volumetric
shrinkage (Sv) was estimated as the difference in
volume of the sample between saturated and oven-
dry (105 °C) state divided by its volume in saturated
state. Volumes were calculated from measurements
of sample height plus top and bottom diameter (mean
of four measurements with a micrometer for each
dimension; Oleszczuk et al. 2003). Ash content (A)
was determined by a loss on ignition method (6 hours
at 550 °C). Specific density (ρ) was calculated from
ash content (A) using the equation 𝜌 = 0.011 ∙ 𝐴 +
1.451 (Okruszko 1971) and total porosity (n) was
calculated from specific density and bulk density. pH
and electrical conductivity (EC) were measured in a
1:10 (weight of soil : volume of liquid) mixture using
a potentiometric method; EC was measured in
distilled water only, while pH was measured in both
distilled water and 1 M KCl. The resulting EC values
were corrected for H+ ions (Sjörs 1950). Total carbon
(C) and total nitrogen (N) were determined using a
CNS analyser (LECO CNS-200), and the mineral
nitrogen forms nitrate (NO3-) and ammonium (NH4+)
using a flow injection analysis (FIAstar 5000, FOSS)
method. Available phosphorus (P) was determined
by the Egner-Riehm method based on soil extraction
with calcium lactate solution acidified with
hydrochloric acid (Lityński et al. 1976).
The stage of secondary transformation of peat was
assessed on the basis of water-holding capacity index
(W1) according to the procedure proposed by Gawlik
(1996). This index is the quotient of the (minimum)
water-holding capacity of the soil when absolutely
dry and its (maximum) water-holding capacity in
fresh condition. It was determined as the water
capacity of a soil sample dried at 105 °C divided by
that of a sample in fresh (field) condition. The soil
samples were divided into two batches. Samples that
were to remain in fresh state were soaked with
distilled water for seven days, while their
counterparts were dried to constant weight at 105 °C
before soaking (also for seven days). Then each
sample was centrifuged at 1000 × g for one hour, at
an ambient temperature of 10 °C. The water content
of each soil sample was determined gravimetrically,
then the W1 index was calculated. The values of W1
were classified as follows (Gawlik 2000):
W1 = 0.36–0.45: I - initial secondary transformation;
W1 = 0.46–0.60: II - weak secondary transformation;
W1 = 0.61–0.75: III - medium secondary transformation;
W1 = 0.76–0.90: IV - strong secondary transformation;
W1 > 0.90: V - completely degraded.
Water table coefficient
A dipwell made from PVC pipe (ø 50 mm) was
installed at each study plot. The wall of the dipwell
was perforated for one-third of its length and coated
with a geotextile screen to prevent silting. Water table
depth (cm below peat surface) was measured
manually every two weeks between April and
November 2016. The degree of drying of the cutover
peat layer was expressed as a coefficient calculated
as (mean water table depth ÷ thickness of residual
peat in cm). Thus, a coefficient of zero indicated that
the water table was level with the peat surface and a
coefficient of unity meant that there was no water
table within the peat layer.
Vegetation survey
The vegetation inventory was carried out by
estimating the cover of plant species on the plots
using a decimal scale (Londo 1976), separately for
the tree, shrub, and herb layers. The cover of
bryophytes and bare peat was recorded in 0.5 × 0.5 m
subplots located in the four corners of each plot.
Bryophytes were identified at genus level. Species
that are characteristic of the classes Scheuchzerio-
Caricetea nigrae and Oxycocco-Sphagnetea
according to Ellenberg et al. (1992) were considered
to be raised bog species.
Statistical analysis
The data for soil properties were tested for normality
using the Shapiro-Wilk test (p > 0.05). Variables that
did not follow normal distributions were subjected to
log transformation. For some variables, this did not
provide the expected result so Spearman's non-
parametric correlation coefficient was used to
evaluate correlations. Although not replicated, we
assessed differences in peat properties between
Sectors A and B using a parametric t test for normally
distributed variables and a non-parametric U-Mann
Whitney test for variables with non-normal
distributions. Results were deemed significant at
α = 0.05 but were interpreted with caution in view of
the pseudoreplication. The statistical analysis was
performed in Statistica 12 (2016 version, Dell Inc.).
The comparison of plant species composition
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
5
between the two sectors was based on the frequency
of occurrence and mean plant cover on the plots. The
Shanonn–Wiener index (Lepš 2005) was used as the
metric of species diversity. Analyses of vegetation
and abiotic factors affecting the occurrence of plants
were carried out using multivariate methods,
Detrended Correspondence Analysis (DCA) and
Redundancy Analysis (RDA), in Canoco software
(ter Brak & Smilauer 2002). The linear method
(RDA) was used with a gradient length of 3.0 SD
(standard deviation) in the DCA analysis. The
significance of partial and marginal effects in RDA
analysis was calculated on the basis of a Monte Carlo
permutation test. In marginal analyses, only one
factor at a time was included as an environmental
variable. This provided information about the
importance of specific factors without reference to
their correlations with other variables (Lepš &
Šmilauer 2003). In partial analyses, each factor was
tested as an environmental variable using the other
factors as covariables.
RESULTS
Peat characteristics
When peat extraction ended, the cutover surfaces in
the two sectors consisted of different types of peat. In
Sector B the surface peat was composed of
cottongrass-Sphagnum (Eriophoro-Sphagneti) peat
with degree of decomposition H4–H6 on the von Post
scale. In the older Sector A, a layer of pine-
Sphagnum (Pino-Sphagnum) peat and sapric peat
with pine wood residues (H4–H8) had been exposed.
One-third of the study plots in the latter sector
featured a compressed moss layer at depth 5–6 cm, as
well as traces of fire.
The average thickness of the residual peat layer
differed significantly between Sectors A and B
(Table 1), as did water level (p < 0.1; Mann-Whitney
U-test). A significant (p < 0.05) negative correlation
(r = - 0.771) was observed between residual peat
thickness and water table depth. Water table
depth also significantly (p < 0.05) correlated with
θv (r = - 0.701), W1 (r = 0.650), Sv (r = - 0.492),
NH4+ (r = - 0.518) and corrected EC (r = 0.667).
In general, between April and November 2016 the
water table was lower in Sector A than in Sector B
(Table 1). From June to August it dropped to
13–22 cm below the base of the peat layer in study
plots with peat thickness ~ 40 cm or less. This was
recorded for 17 % of the plots in Sector B and 58 %
of the plots in Sector A. In the rest of the plots (where
the water table always remained within the peat
profile), water table depth was 3–37 cm (mean
25 cm) in Sector A and 1–42 cm (mean 16 cm) in
Sector B. The water table coefficient indicated that
the peat in Sector A was markedly drier than that in
Sector B (Figure 2). It is likely that this caused
differences between the sectors in some physical and
hydrophysical properties of the uppermost 10 cm of
peat (Table 1). Values of n, Sv, θv and θs were all
higher in Sector B while values of W1 were higher in
Sector A. On the basis of W1 index, the secondary
transformation class of top-layer peat in Sector A was
‘weak’ to ‘strong’, whereas Sector B featured less-
transformed peat, between ‘initial’ and ‘medium’. As
far as chemical properties are concerned, the sectors
differed in NH4+ content, pH and corrected EC. NO3-
outweighed NH4+, although the difference was
markedly lower in Sector A. Both sectors were
characterised by low content of available P and
extremely low pH, with higher values of both
attributes in the older Sector A (Table 1).
Vegetation characteristics
For all study plots the mean cover of raised bog
species was 46 %, while the mean cover of species
not usually associated with raised bogs was 85 %.
There was no difference between the two sectors in
the number of species per plot and species diversity
(Shannon-Wiener index). The cover of herbaceous
plants was significantly higher in Sector B, while
moss cover (with prevalence of true mosses) was
higher in Sector A. No differences were found for
shrub layer cover or bare peat, while tree layer cover
was higher in Sector A (Table 2). The raised bog
plants Sphagnum spp., Ledum palustre and
Oxycoccus palustris were more common in Sector B;
indeed, the last of these species was not found in
Sector A. Sphagnum (cover 12 %) occurred mainly
in the form of isolated cushions and did not form a
continuous acrotelm. Eriophorum vaginatum and
Ericaceae such as Calluna vulgaris, Vaccinium
uliginosum, V. vitis-idaea and V. myrtillus were
common in both sectors but Eriophorum vaginatum
cover in Sector B was double that in Sector A. Tree
species (Pinus sylvestris, Betula pendula) occurred in
all vegetation layers. However, Pinus sylvestris in the
shrub layer was associated with Sector B (Table 3).
Effect of peat quality on revegetation
The first and second axes of the DCA ordination
diagram shown in Figure 3 explain 16.9 % and 8.9 %,
respectively, of the variability in plant species
composition. The first axis reflects the main
differences in species composition. Thus, the left-
hand part of the diagram contains raised bog plants
such as Sphagnum spp., Eriophorum vaginatum,
Oxycoccus palustris and Ledum palustre while the
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
6
Table 1. Comparison of soil properties for the uppermost 10 cm of peat, between Sector A and Sector B. Significance was tested with a parametric t-test or a non-
parametric Mann-Whitney U-test; *comparison of medians with non-parametric Mann-Whitney U-test. Key: SD = standard deviation; n.s. = non-significant at p < 0.05;
b.g.l. = below ground level.
Soil properties
Units
Sector A
Sector B
Sector
A vs. B
min.
max.
mean
SD
min.
max.
mean
SD
Peat depth
cm
19
98
39.3
21.8
32
113
64.6
23.9
p < 0.001*
Total porosity (n)
%
78
89
84
3
77
91
86
3
p < 0.05
Bulk density (ρb)
Mg m-3
0.16
0.33
0.25
0.05
0.14
0.35
0.22
0.06
n.s.
Specific density (ρ)
Mg m-3
1.50
1.73
1.59
0.06
1.47
1.78
1.57
0.10
n.s.*
Ash content (A)
%
4.24
25.35
12.69
5.81
1.99
30.34
10.94
9.41
n.s.*
Volumetric shrinkage (Sv)
%
15.75
53.12
31.68
9.07
19.14
59.93
45.07
10.40
p < 0.001
Volumetric moisture content (θv)
vol.%
12.48
67.66
34.65
17.87
58.95
88.59
76.05
8.75
p < 0.001*
Saturated moisture content (θs)
vol.%
61.14
90.85
73.00
7.15
75.87
88.69
81.86
3.06
p < 0.001*
Water-holding capacity index (W1)
-
0.53
0.76
0.61
0.07
0.39
0.68
0.55
0.08
p < 0.05
Total C
%
47.45
58.14
53.79
2.81
33.84
63.07
55.57
6.71
n.s.*
Total N
%
1.38
2.02
1.70
0.17
1.19
2.02
1.65
0.21
n.s.*
C/N
-
25
39
32
3
27
48
34
5
n.s.
NO3-
mg kg-1
0.83
37.08
3.30
8.22
0.44
6.24
1.51
1.26
n.s.*
NH4+
mg kg-1
7.68
44.58
16.19
9.03
35.84
172.46
77.54
33.79
p < 0.001*
Available P
mg kg-1
10.09
48.77
23.30
9.98
2.62
41.94
17.92
9.70
n.s.
pHH2O
-
3.56
3.96
3.72
0.12
2.96
3.60
3.29
0.21
p < 0.001*
pHKCl
-
2.35
3.24
2.82
0.23
2.52
2.90
2.68
0.13
p < 0.05*
Corrected electrical conducticity (EC)
µS cm-1
119.00
252.08
177.79
38.34
60.62
264.00
134.58
50.35
p < 0.01
Water table depth
cm b.g.l.
50
0
30.4
7.0
51
0
17.9
5.9
p < 0.1*
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
7
right-hand part lacks raised bog species and is rich in
trees and true mosses (mainly Pleurozium sp.). The
plots located in the older Sector A are clustered in the
right-hand part of the diagram, whereas those in the
younger sector B appear in the left-hand part.
However, the central part of the diagram contains
plots from both sectors that harbour raised bog
species (Vaccinium uliginosum) and plants that are
typical for oligotrophic habitats but not peat-forming
(e.g. Calluna vulgaris).
The main gradient of change in species
composition correlated with water table depth and
soil factors (Figure 4) that reflect the secondary
transformation stage of the uppermost layer of
cutover peat. Plots with raised bog species were
characterised by higher water level, lower W1 and ρb,
higher n, θv, θs, Sv, NH4+ content, total C and C/N, as
well as greater residual peat depth. The second axis
correlated (weakly) only with available P, total N and
pH. The RDA analysis of the significance of
individual variables for plant species composition
showed that twelve of them (Table 4) were
significant when each was analysed as an individual
variable (marginal effect). The most important
attribute seemed to be water table depth, which
accounted for 35 % of the variation. However, the
strong intercorrelation of significant variables meant
that the effects of individual variables without the
influence of all other variables (partial effect) were
non-significant in all cases (Table 4).
Figure 2. Variation of the water table coefficient (mean water table depth ÷ thickness of residual peat) during
the period April to November 2016, for Sector A (unfilled circles) and Sector B (filled circles).
Table 2. Diversity measures and mean cover (%) of plant species typical of raised bogs; other plant species
(not characteristic for raised bogs); the tree, shrub and herb layers; and bare peat, in the two extracted sectors
of the Bór za Lasem bog.
Attribute
Sector A
Sector B
p
Mean for
both sectors
Mean number of species per plot
11.5
10.6
0.23
11.1
Shanonn-Wiener index
1.8
1.6
0.13
1.7
Raised bog species cover
28
65
0.00
46
Non raised bog species cover
93
76
0.00
85
Tree layer cover
11
1
0.00
6
Shrub layer cover
10
10
0.95
10
Herb layer cover
44
80
0.00
83
Moss layer cover
23
11
0.00
17
Bare peat cover
18
22
0.37
20
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
8
Table 3. Frequency of occurrence (%) of plant species and mean plant cover (%) calculated only for plots
where the species was present. Only species with frequency > 50 % and raised bog species (bold) are presented.
Species
Sector A
Sector B
Frequency
Mean cover
Frequency
Mean cover
Oxycoccus palustris
0
0
35
4
Sphagnum spp.
32
4
55
12
Pinus sylvestris (shrub layer)
16
7
45
10
Ledum palustre
42
3
90
8
Eriophorum vaginatum
95
23
95
49
Pleurozium schreberi
89
19
75
9
Calluna vulgaris
84
17
80
24
Vaccinium uliginosum
84
18
80
13
Brachythecium spp.
68
2
60
4
Pinus sylvestris (herb layer)
68
1
50
2
Betula pendula (shrub layer)
58
11
45
8
Polytrichum spp.
84
10
80
6
Aulacomnium palustre
74
2
20
1
Betula pendula (herb layer)
53
2
35
1
Vaccinium myrtillus
53
7
35
3
Figure 3. Unconstrained ordination diagram (DCA) of plots (filled circles - Sector A, unfilled circles -
Sector B) and vegetation data. Only raised bog species (in bold) and species with the highest weight are
presented. Abbreviations of species names: Aul_pal = Aulacomnium palustre, Bet_pen_a = Betula pendula
in tree layer, Bet_pen_b = Betula pendula in shrub layer, Bra_spp. = Brachythecium species, Cal_vul =
Calluna vulgaris, Eri_vag = Eriophorum vaginatum, Led_pal = Ledum palustre, Oxy_pal = Oxycoccus
palustris, Pin_syl = Pinus sylvestris, Ple_sch = Pleurozium schreberi, Pol_spp. = Polytrichum species,
Sph_spp. = Sphagnum species, Vac_myr = Vaccinium myrtillus, Vac_uli = Vaccinium uliginosum, Vac_vit =
Vaccinium vitis-idaea.
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
9
Figure 4. Unconstrained ordination diagram (DCA) of all environmental variables (arrows) shown as the
passive ones. Abbreviations of variables: Water bgl = water table depth (below ground level), Depth = depth
of residual peat, n = total porosity, ρb = bulk density, ρ = specific density, A = ash content, Sv = volumetric
shrinkage, θv = volumetric moisture content, θs = saturated moisture content, W1 = water-holding capacity
index, C = total carbon, N = total nitrogen, NO3 = nitrate nitrogen, NH4 = ammonium nitrogen, P = available
phosphorus, EC = corrected electrical conductivity.
Table 4. Results of the constrained ordination (RDA). Marginal effects of the environmental variables and
their partial effects on the vegetation based on forward selection (Monte Carlo permutation test). Only
variables with significant marginal effect are presented.
Variable
Marginal effect
Partial effect
Explained
variability (%)
p
F
Explained
variability (%)
p
F
Water table depth
35
0.002
11.92
2
0.616
0.58
Volumetric moisture content
26
0.002
7.78
3
0.408
1.10
Corrected electrical conductivity
24
0.002
6.96
2
0.592
0.73
Sector
16
0.012
4.10
1
0.710
0.36
Total C
15
0.010
3.95
2
0.492
0.92
Water-holding capacity index W1
15
0.010
3.87
2
0.638
0.54
Peat depth
13
0.020
3.42
3
0.584
0.70
NH4+
13
0.022
3.18
2
0.486
0.93
Volumetric shrinkage
12
0.026
3.05
3
0.584
0.71
C/N
11
0.042
2.74
2
0.526
0.86
Total porosity
10
0.044
2.58
1
0.690
0.50
Bulk density
10
0.048
2.43
2
0.608
0.65
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
10
DISCUSSION
Because of their intercorrelations, the factors
investigated were not individually important in
determining the revegetation pattern on the extracted
parts of the Bór za Lasem bog. However, water table
depth seemed important as the abiotic factor that
determined the degree of soil aeration and,
consequently, the secondary transformation stage of
the peat. Therefore, it may be concluded that the
interactions of water table depth with peat soil
properties further promote the importance of water
conditions on extracted areas.
Soil properties which explained plant species
composition (W1, θv, n, ρb, Sv, corrected EC, NH4+,
C, C/N) were associated with water table position, as
confirmed by partial DCA. Except for NH4+ content,
edaphic factors (e.g. pH, NO3- content, available P)
did not affect vegetation development at Bór za
Lasem, even though a role has been demonstrated in
other studies (Salonen 1994, Graf et al. 2008,
Konvalinková & Prach 2010). This was probably due
to the very small variation in these factors within the
two cutover areas investigated here. The values of the
soil properties tested indicated that secondary
transformation of peat was more advanced in
Sector A, which was abandoned about ten years
earlier than Sector B.
The revegetation patterns on the two extracted
sectors of Bór za Lasem bog were similar to those
observed on extracted areas of other peatlands in
Europe, e.g. by Poschlod et al. 2007 (Germany),
Konvalinková & Prach 2014 (Czech Republic),
Triisberg et al. 2014 (Estonia) and Orru et al. 2016
(also Estonia). In our study areas, the water table
often dropped into the poorly permeable mineral
(clay) substratum, especially where the peat layer
was thin (< 0.4 m). This favoured the development of
vascular plants including trees, whose roots can
access nutrients by growing into the rich mineral
substratum. Trees promote drying of the peat layer
through transpiration and interception of water
arriving as precipitation (Van Seters & Price 2001,
Limpens et al. 2014), and this further enhances the
processes that cause secondary transformation of
peat. Such conditions prevailed in Sector A, where
revegetation was tending towards a pine and birch
community. Common inhabitants of this sector were
species typical of coniferous forest such as
Vaccinium myrtillus, V. vitis-idaea and V. uliginosum,
along with mosses belonging to the genera Polytrichum
and Brachythecium, Pleurozium schreberi and
Aulacomnium palustre. These mosses are usually
associated with forest communities where trees are
providing considerable shade (Hedwall et al. 2017).
The revegetation by Sphagnum and other raised
bog species (Eriophorum vaginatum, Vaccinium
uliginosum, Ledum palustre, Oxycoccus palustris)
was associated with areas of deeper residual peat and
high water table, which in turn were strongly
correlated with hydrophysical soil properties such as
volumetric moisture content θv and water-holding
capacity index W1. High water level is a prerequisite
for Sphagnum re-establishment because it mitigates
the effect of radically differing hydrophysical
properties between the regenerating moss cover and
the cutover peat (McCarter & Price 2015). Sphagnum
lacks roots and vascular channels, so water uptake
and transport processes differ from those in vascular
plants (Schouwenaars & Gosen 2007). Price &
Whitehead (2001) formulated limit values to define
hydrological conditions suitable for the development
of Sphagnum, which included a mean water table
depth of 24.9 ± 14.3 cm below the ground surface,
θv > 50 % and soil-water pressure above - 100 mb.
Soil-water pressure was not evaluated in our study,
but θv and mean water table depth indicated
potentially favourable conditions for Sphagnum
development in the younger Sector B and
unfavourable conditions in Sector A. The difference
in θv between the two sectors was important. There
were serious consequences for regeneration of typical
raised bog plant species including, most notably,
Sphagnum spp. Reduced moisture content (and, thus,
high soil water tension) in the uppermost peat layer
inhibits Sphagnum development because these
mosses are unable to take up water from the substrate
when the pore-water pressure is below -100 mb
(Hayward & Clymo 1982 op. cit. Price 1997). On the
other hand, the root systems of vascular plants can
collect water from deeper soil layers and nutrients
from mineralised organic matter (Malmer et al. 2003)
and/or from the mineral substratum underlying a thin
layer of peat. Therefore, they are capable of
colonising extracted peatland areas at a faster rate.
In our study, DCA revealed correlations between
the establishment of Sphagnum and other raised bog
species and the water-holding capacity index W1.
Bog species were present on the study plots showing
lower W1 index values, i.e. where the peat was
transformed to a lower degree. This relationship is
most important for Sphagnum, which does not have
roots and is thus more highly dependent than vascular
plants on water conditions in the uppermost layer of
cutover peat. Thus, the W1 index may be useful not
only for evaluating the secondary transformation
stage of peat, but also as an indicator of the potential
for spontaneous regeneration of vegetation composed
of Sphagnum species on degraded bogs. However, in
view of the relatively low representation of
E. Zając et al. SUBSTRATE QUALITY AND REVEGETATION OF EXTRACTED PEATLAND
Mires and Peat, Volume 21 (2018), Article 12, 1–14, http://www.mires-and-peat.net/, ISSN 1819-754X
© 2018 International Mire Conservation Group and International Peatland Society, DOI: 10.19189/MaP.2017.OMB.310
11
Sphagnum on the peatland that we investigated here,
this suggestion must be explored in further research
before it can be verified.
There is a possibility that interventions to help
raise the water table in Sector B may promote further
expansion of Sphagnum. Local conditions such as the
proximity of the bog dome remnant and thus of a
diaspore bank (Konvalinková & Prach 2014), as well
as the small area (Triisberg et al. 2011, Kollman &
Rasmussen 2012), may favour the establishment of
bog species. Recovery towards raised bog might be
positively affected by the great abundance of
Eriophorum vaginatum, which is nearly two times
more common (with larger individual plants) in
Sector B. This species prefers areas of thick peat with
low ash content, predominance of NH4+ over NO3-
(Salonen 1994), water table depth no greater than
30–40 cm and θv > 70 % (Lavoie et al. 2005), which
is consistent with our results. Various studies have
suggested that plants such as Eriophorum (Soro et al.
1999), ericaceaous shrubs or young trees (Pouliot et
al. 2011a) and Polytrichum strictum (Groeneveld et
al. 2007) may facilitate Sphagnum colonisation by
improving the microclimate and shaping
microtopography.
We found a strong negative correlation between
water table depth and residual peat thickness.
Differences between these two environmental factors
were expressed in terms of a coefficient that clearly
illustrated the contrast in conditions between the two
sectors. In 2016, 80 % and 40 % of the residual peat
layer in Sectors A and B, respectively, was above the
average water table level (and, thus, usually
unsaturated) during the growing season. Where
residual peat thickness was less than 40 cm, the water
table was below the organic soil horizon for the entire
observation period. This suggested that a certain
depth of residual peat was required to stabilise the
water level, i.e. it helped to limit water table
fluctuations. However, further observations (in
progress) are needed to confirm this hypothesis.
Assuming that satisfactory peat thickness and degree
of peat decomposition are preconditions for
successful recovery of mire vegetation, it may be
concluded that the residual peat layer on the
investigated sectors was in general too thin and
decomposed. There is no strict threshold for the
minimum residual peat depth required for restoration
but the value that is most often recommended is at
least 0.5 m for well-decomposed peat (H ≥ 7) and
1.0 m for less-decomposed peat (H5–H7) (Wheeler
& Shaw 1995, Quinty & Rochefort 2003). Successful
restoration has been performed on Canadian sites
with less than 1.0 m depth of less-decomposed peat
(González & Rochefort 2014). On the other hand,
Poschlod et al. (2007) consider that the peat thickness
values stated above are insufficient in the conditions
of southern Germany, and Triisberg et al. (2014) state
that raised bogs with less-decomposed peat in the
boreo-nemoral region should be restored when
residual peat thickness is greater than 2.3 m.
Spontaneous regeneration of cutover bogs is
possible only under favourable conditions. It is also a
long-term process that may take more than a century
to complete (Pouliot et al. 2011b). Restoration of a
properly functioning hydrological system is a crucial
element of rehabilitation for any type of peatland
(Chimner et al 2017). There are many techniques for
improving hydrological conditions on extracted
peatlands (see, for example, Wheeler & Shaw 1995,
Price et al. 2003, Graf et al. 2012), but the effects of
a residual peat layer that is too shallow may be
sufficiently serious to prevent the restoration of a
raised bog. Thus, leaving behind only a shallow peat
layer at the end of peat extraction operations may be
justified only if the recovery of peat-forming bog
vegetation is not an objective for the peatland.
ACKNOWLEDGEMENTS
We thank the authorities of Gmina Czarny Duanjec,
especially Michał Jarończyk and Bogusław Sroka
from Peat Production Plant “Bór za Lasem” in
Czarny Dunajec for help and permission to conduct
the research. We also thank Ewa Zagrodzka and
Marek Turschmid for assistance with the field and
laboratory work. This research was carried out within
Projects DS-3331/KMIKŚ and DS-3337/KEKiOP
financed from a research grant allocated by the Polish
Ministry of Science and Higher Education.
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Submitted 08 Nov 2017, final revision 13 Jun 2018
Editor: Stéphanie Boudreau
_______________________________________________________________________________________
Author for correspondence:
Dr Ewelina Zając, Department of Land Reclamation and Environmental Development, Faculty of
Environmental Engineering and Land Surveying, University of Agriculture in Kraków, al. Mickiewicz a 24/28,
30-059 Kraków, Poland. Tel. 48126624015; E-mail; e.zajac@ur.krakow.pl
