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Bryophytes and vascular plants on peat extraction sites - which factors influ-
ence their growth?
Zarzycki Jan, Zając Ewelina, Grzegorz Vončina
PII: S1617-1381(22)00160-1
DOI: https://doi.org/10.1016/j.jnc.2022.126287
Reference: JNC 126287
To appear in: Journal for Nature Conservation
Received Date: 7 July 2022
Revised Date: 29 September 2022
Accepted Date: 1 October 2022
Please cite this article as: Z. Jan, Z. Ewelina, G. Vončina, Bryophytes and vascular plants on peat extraction sites
- which factors influence their growth?, Journal for Nature Conservation (2022), doi: https://doi.org/10.1016/
j.jnc.2022.126287
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© 2022 Published by Elsevier GmbH.
Bryophytes and vascular plants on peat extraction sites - which factors influence their growth?
Zarzycki Jan1, Zając Ewelina2, Grzegorz Vončina3
1Department of Ecology, Climatology and Air Protection, University of Agriculture in Kraków,
Kraków, Poland
2Department of Land Reclamation and Environmental Development, University of Agriculture
in Kraków, Kraków, Poland
3Pieniny National Park, Krościenko, Poland
*Corresponding author: Jan Zarzycki; Department of Ecology, Climatology and Air Protection,
University of Agriculture in Kraków, al. Mickiewicza, 24/28, 30-059, Kraków, Poland, e-mail:
jan.zarzycki@urk.edu.pl
Abstract
Peat extraction leads to the formation of areas with altered habitat conditions in comparison to
natural peatlands. Restoration of the peat-formation process in these areas is very difficult and
requires the creation of suitable conditions for the growth of peatland species. The aim of the
study was to analyse the habitat requirements of bryophytes and vascular plants growing on
sites of peat extraction (30 and 40 years after extraction was terminated) and to determine
whether the water level influences the growth conditions of plants directly or indirectly through
changes in the peat physical, hydraulic and chemical properties. Analysing all factors together
revealed that the average water level had a decisive influence on bryophytes, but a statistically
significant increase in the percentage of variation explained was obtained by taking into account
other parameters as well (proportion of macropores, carbon content, and pH). In the case of
vascular plants the analysis showed that the water table coefficient included the effects of all of
the other factors analysed, and taking them into account did not increase the percentage of
variation explained. The two groups of plants use different resources of the environment.
Key words: raised bog; bryophytes; restoration; peatland water table
Introduction
Natural peatlands play an important role on a global scale as areas which not only retain vast
amounts of water (Liu et al., 2022), but also accumulate large amounts of carbon (Scharlemann
et al., 2014), thereby contributing to climate regulation. In this context, as functioning
ecosystems that have not been drained, peatlands help to slow climate change (Leifeld &
Menichetti, 2018). Apart from their impact on the abiotic environment, peatlands increase
biological diversity (Minayeva et al., 2017), creating habitats for many narrowly specialized
species of plants (Vítovcová et al., 2022) and animals, especially invertebrates (Buchholz et al.,
2009; Vítovcová et al., 2022).
In Europe, peatlands occupy a total area of 593,727 km2 (Tanneberger et al,. 2017) and are
present mainly in the northern and eastern parts of the continent. In Poland they occupy only
149,504 km2, which is 4.79% of the area of the country (Tanneberger et al., 2017). Raised bogs
are particularly rare, accounting for only 4.2% of all peatlands in Poland (Ostrowski et al.,
1995). Fed entirely by precipitation, ombrogenous raised bogs form characteristic domes,
which rise above the surrounding water table. This means that they can be formed only in areas
with a positive water balance, i.e. where precipitation exceeds evapotranspiration. For this
reason they are concentrated in the cool temperate zone between the 50th and 70th parallels
north. At lower latitudes they are present in some places in upland and mountainous areas
(Proktor, 1995). Natural peatlands, including raised bogs, have a diplotelmic structure. They
consist of the top acrotelm layer, a few dozen centimetres thick, which is within the reach of
fluctuations in the water level, and the catotelm, which is permanently saturated with water and
makes up most of the peat deposit. The acrotelm is the layer with the highest biological activity,
especially the growth of peat-forming vegetation (Proktor, 1995). The removal of the acrotelm
ends the peat-formation process.
Raised bogs play an important role in preserving biodiversity due to the habitat conditions
prevailing in them. Their extremely low nutrient levels and highly acidic pH favour the
development of specialized plant communities characterized by an abundance of Sphagnum
species (Bragg, 1995). The species diversity (α-diversity) of these communities is generally not
high compared to other natural and semi-natural ecosystems, but the species present in them
are not usually found in other habitats, and this increases the total local diversity (β and γ-
diversity).
The surface area of peatlands is decreasing due to drainage, mainly for agricultural purposes
(Swindles et al., 2019), but also as a result of extraction of peat, which has been exploited for
centuries in numerous sectors of the economy (Tanneberger et al., 2021). The habitat conditions
of post-mining peatland areas differ from those of undisturbed areas. Peat extraction decreases
the total thickness of the deposit and involves the removal of the acrotelm layer together with
the seed and spore bank. The acrotelm layer with its high water storage capacity is replaced by
a layer of highly decomposed peat with lower water storage capacity, resulting in greater
fluctuations of the water level (Schouwenaars, 1993). Mineralization of peat and the release of
nitrogen compounds alter the trophy of the habitat, which is particularly unfavourable in the
case of oligotrophic peatlands. The direction of succession in post-mining areas is strongly
influenced by topography, which depends on the extraction method. The surface of the block-
cut area is uneven, with elevations and depressions, while in the milled peat area (currently
predominant) the surface between ditches is flat. This creates varied starting conditions for
regeneration, which are much more difficult in the case of milled peat fields (Wheeler and Shaw
1995, Triisberg et al., 2011). Natural regeneration processes in mechanically mined areas are
very slow (especially in case of milled peat fields), and due to the altered habitat conditions the
succession process usually leads to the formation of alternative communities (Lavoie et al.,
2003; Vítovcová et al., 2022). Restoration of peat-formation processes is possible and
necessary, because it can limit emissions of greenhouse gases, especially carbon dioxide
(Wilson et al., 2016), regenerate typical plant communities, and increase biodiversity (Gonzales
et al., 2014). As a rule, however, this requires active measures to be taken. Attempts at
reclamation of raised bogs are made in many countries (Minayeva et al., 2017; Purre & Ilomets,
2021), but they are not always successful due to the many interdependencies between
environmental factors (Salonen, 1994). The impact of environmental factors on a macro scale
(elevation above sea level, climate, precipitation) is large (Benavides & Vitt, 2014), but local
conditions may be decisive (Howie et al., 2020). Successful restoration of a peatland requires a
good understanding of the effects of various environmental factors on individual groups of
plants and their mutual relationships (Lavoie et al., 2003). This applies to both bryophytes,
which make up the bulk of peat-forming vegetation, and specialized species of vascular plants.
These two groups differ in their structure and physiology, so they react differently to
environmental gradients. The relationships between vascular plants and bryophytes have been
studied in forest ecosystems (Vellak et al., 2003), grasslands (Ingerpuu et al., 2005), and
peatlands (Malmer et al., 2003, Hajkova & Hajek, 2004). Post-mining areas of peat bogs,
however, have different habitat conditions than natural ecosystems, and thus the relationships
between these groups of plants may be different as well. Numerous studies (Wilson et al., 2016;
Konvalikova & Prach, 2014) have shown that the main essential factor for the development of
typical raised bog vegetation is a sufficiently high water level, but this is not always enough
(Gonzales et al., 2014; Purre & Ilomets, 2021). Other important factors are the physical and
chemical parameters of the peat layer (Purre & Ilomets, 2018) and the presence of a seed bank
of peat-forming vegetation in the vicinity (Poschlod, 1995).
At the study site of the present study, the termination of extraction by the block-cutting method
was followed by the appearance of microhabitats differing in the thickness and degree of
secondary transformation of the remaining peat, the water level, and – depending on how much
time has passed since the termination of peat extraction – in the advancement of succession.
Studies on the implications of peat substrate depth, quality (hydrophysical and chemical
properties) and water level for the spontaneous revegetation of two post-mining peat fields at
different succession stages have been published in Zając et al., 2018a and Zając et al., 2018b,
while the relationships between microclimatic conditions and water level fluctuations at the
same study sites were the subject of work by Zarzycki et al., 2020.
The aim of the study was to analyse differences in the habitat requirements of bryophytes and
vascular plants growing in post-mined peat areas and to determine whether the water level
affects the growth conditions of plants directly or indirectly through changes in the physical,
hydraulic and chemical properties of the peat.
Materials and methods
Study site
The study site was an area where peat extraction had been carried out on the raised bog Bór za
Lasem, situated in the Orawa-Nowy Targ Basin in southern Poland (Fig. 1). The basin is a
depression surrounded by mountain ranges to the north and south, with a moderately warm
climate, but with distinctive climate features such as pools of cold air and fog and warmer and
drier summers than in the surrounding terrain (Kondracki, 2011). Meteorological data for
research area were published elsewhere (Zarzycki et al. 2020). The current area of the bog is
55 ha, the average peat thickness is 1.8 m (max 3.65 m), the average degree of decomposition
is 30%, and the average ash content is 2.2% (Lipka & Zając 2014). The peat bog is protected
under the European Ecological Network Natura 2000 as part of the Orawa-Nowy Targ
peatlands (PLH120016, PLB120007).
Fig. 1 Location of the study area
In the 1960s peat extraction was begun on an industrial scale by the block-cutting method, using
heavy equipment. Two sectors exploited during different periods were formed (Fig. 1). In sector
A (about 16 ha), peat extraction was carried out until the early 1980s, while in sector B (about
8 ha) it was continued until the early 1990s. . Sector A had a deep water table with small
fluctuations and low peat thickness, while sector B had thicker peat and a high water level with
large fluctuations (Fig. 2). Variation in the physiochemical properties of peat between the
sectors is described in Zajac et al., 2018ab. After peat extraction was terminated, neither sector
was reclaimed, but left to natural succession (Zając et al., 2018). Sector A is covered with forest
communities dominated by Scots pine and birch (Betula pendula). Sector B is a mosaic of non-
forest communities with heather (Calluna vulgaris) and hare’s-tail cottongrass (Eriophorum
vaginatum), as well as communities with pine and birch saplings (Zając et al., 2018 a).
Fig. 2 Location of the ground water table in the warm half of 2017
Sampling
A total of 22 research plots 5 m x 5 m in size were set up in the two sectors of the post-mining
area (11 plots in each sector). In each plot the vegetation, peat thickness and quality, and
groundwater level were studied. The plots were distributed systematically, but ditches, larger
depressions and other atypical locations were avoided. The study was carried out on the same
plots where the hydrophysical and chemical properties of the peat had been analysed by Zając
et al. (2018a,b). In each plot the vegetation, peat thickness and quality, and groundwater level
were studied. The analysis of vegetation involved estimation of the percentage area cover by
vascular plants in the tree layer a (>5 m), shrub layer b (1.5 – 5 m) and herbaceous layer c (<1.5
m) was done in August/September 2019. The area cover by bryophytes (d) was estimated in
September 2019. Hemispherical photographs processed using ImageJ software (Smith &
Ramsay, 2018) were used to calculate the degree of shade (canopy openness %) in the plots.
In each plot the thickness of the peat was measured (to the mineral substrate), and in the 2017
growing season the water level was measured in dipwells at two-week intervals. The degree of
drying of the peat layer was expressed as the water table coefficient (WD) and calculated as
mean water table depth (W) ÷ thickness of residual peat (D) and thus WD illustrates the
relationship between the residual peat thickness and hydrological conditions, expressed as the
water table. The higher the WD value, the higher the degree to which the peat layer is dried up,
which affects the proportion of peatland species. When WD has a value of zero, the water table
is at the level of the peat surface, whereas a value of one means that the water table is below
the peat layer (Zając et al., 2018). Samples of disturbed and undisturbed structure (metal rings)
were collected from each plot from a depth of 0–10 cm for laboratory analyses. The analyses
included the water-holding capacity, dry bulk density, specific density, total porosity,
volumetric moisture content, saturated moisture content, volumetric shrinkage, maximum
hygroscopic moisture content, proportions of macro-, meso- and micropores determined from
a water retention curve (pF), ash content, pH (in H2O and KCl), electric conductivity, total
carbon, total nitrogen, mineral nitrogen forms (NO3- and NH4+) and available phosphorus
(P2O5). A detailed description of the methods of the laboratory analyses and their results are
published in Zając et al. (2018). Species of bryophytes and vascular plants in the plots were
recorded and quantified according to the scale adopted by Braun-Blanquet (1964). Species that
were difficult to identify in the field were collected for identification in the laboratory. The
species names of vascular plants were adopted according to Mirek et al. (2020) and the names
of bryophytes according to Hodgetts et al. (2020).
Data analysis
To describe the relationships between the occurrence of bryophytes and vascular plants, the
statistical analysis took into account only vascular plants from layer c (herbaceous layer). The
experimental plots were located in such a way that woody species grew outside the plot, but
their crowns could directly shade the surface. For this reason trees and shrubs (layers a and b)
were included in the analysis as one of the habitat factors. Two vegetation subsets were used in
the analyses: 1) only bryophytes and 2) only vascular plants. Redundancy analysis (RDA) was
performed separately for each subset, with environmental factors treated as explanatory
variables. The effect of each variable was evaluated separately (marginal effect). The ‘forward
selection’ method was used to select the variables that best describe the differences in the
species composition, and the statistical significance of each variable was evaluated by the
Monte Carlo permutation test (conditional effect) (Lepš & Šmilauer, 2003). To determine the
main gradients of variation in species composition, detrended correspondence analysis (DCA)
of the vegetation data was performed with downweighting of rare species.
Ecological interpretation of the main gradients of variation was based on Ellenberg indicator
values (EIV) (Ellenberg et al., 1992) defining the habitat requirements of bryophytes and
vascular plants with regard to habitat moisture (F), pH (R), light (L) and fertility (N). Due to
the lack of a fertility indicator for bryophytes, values reported by Simmel et al. (2020) were
used.
The Pearson correlation coefficient was used to correlate the ordination scores obtained for the
DCA bryophyte axes and vascular plant axes with each other and with habitat parameters, i.e.
indicator values (EVI), the ratio of Sphagnum (Sp) to true mosses (B) (Sp/B), and the proportion
of cover of diagnostic species of vascular plants for raised bogs among all species of vascular
plants in layer c (Rb/V), as well as with the dominant species of vascular plants from all layers
(a, b, c).
Diagnostic species of bryophytes and vascular plants for raised bogs (Q11) were determined
according to the EUNIS Habitat Classification (Chytrý et al., 2020).
Multivariate analyses were performed using CANOCO 5.1 software (ter Braak and Šmilauer,
2018). Other analyses were carried out using the Statistica 13.1 software package. For
environmental variables that did not meet the condition of normal distribution, log
transformation was used. Only parameters with the highest weights were presented in the
results.
Results
Vegetation composition
The main species in both the tree layer (layer a) and the shrub layer (layer b) were Pinus
sylvestris and Betula pendula. Among vascular plant species in the herbaceous layer (layer c),
Eriophorum vaginatum, a diagnostic species for raised bogs (Q11), was present in all plots.
Other raised bog species recorded were Ledum palustre, and much less frequently, Vaccinium
oxycoccos. Dwarf shrubs of the family Ericaceae were usually predominant in this layer, mainly
Calluna vulgaris, Vaccinium uliginosum, V. myrtillus, and V. vitis-idea. Among bryophytes
(layer d), the most frequently recorded species were Pleurozium schreberi (in all plots) and
Dicranum scoparium and Brachytecium rutabulum. There were 5 diagnostic bryophyte species
for raised bogs (Q11): Sphagnum fallax, S. magellanicum, S. rubellum, S. capillifolium, and
Polytrychum strictum. The most common was Polytrichum strictum. Among 9 recorded species
of the genus Sphagnum, only S. fallax was fairly abundant. In total 42 species of bryophytes
were identified (Table 1).
Table 1. Vascular plant and bryophytes species recorded on experimental plots (N = 22).
Raised bog species (Eunis Q11 according to Chytrý et al. 2020) in bold. Average cover of:
bare peat 17%, layer a 32%, layer b 10%, layer c 58%, layer d 25%
Layer
Species
No of
occurrence
Layer
Species
No of
occurrence
Layer
Species
No of
occurrence
a
Betula pendula
7
c
Epilobium angustifolium
1
d
Lophocolea heterophylla
5
a
Pinus sylvestris
6
c
Frangula alnus
1
d
Rhytidiadelphus
squarrosus
5
a
Picea abies
1
c
Juncus effusus
1
d
Polytrichum longisetum
4
b
Betula pendula
13
c
Picea abies
1
d
Sphagnum cuspidatum
4
b
Pinus sylvestris
6
c
Rubus idaeus
1
d
Tetraplodon angustatus
4
b
Picea abies
3
c
Salix cinerea
1
d
Straminergon stramineum
3
b
Frangula alnus
2
d
Pleurozium schreberi
22
d
Polytrichum formosum
3
b
Salix caprea
2
d
Dicranum scoparium
21
d
Sphagnum capillifolium
3
b
Populus tremula
1
d
Brachythecium rutabulum
20
d
Sphagnum fimbriatum
3
c
Eriophorum vaginatum
22
d
Pohlia nutans
18
d
Sphagnum rubellum
3
c
Calluna vulgaris
21
d
Polytrichum commune
16
d
Sciuro-hypnum reflexum
2
c
Vaccinium uliginosum
19
d
Polytrichum strictum
15
d
Ceratodon purpureus
2
c
Ledum palustre
16
d
Dicranum polysetum
13
d
Dicranum montanum
2
c
Pinus sylvestris
16
d
Brachythecium
salebrosum
12
d
Bryum pseudotriquetrum
1
c
Vaccinium myrtillus
10
d
Fuscocephaloziopsis
connivens
10
d
Calypogeia integristipula
1
c
Vaccinium vitis-idea
10
d
Plagiothecium
curvifolium
8
d
Campylopus introflexus
1
c
Betula pendula
9
d
Sanionia uncinata
8
d
Cephalozia bicuspidata
1
c
Carex nigra
6
d
Sphagnum fallax
8
d
Plagiomnium undulatum
1
c
Dryopteris carthusiana
6
d
Dicranella cerviculata
7
d
Platygyrium repens
1
c
Vaccinium oxycoccos
4
d
Herzogiella seligeri
7
d
Ptilium crista-castrensis
1
c
Sorbus aucuparia
3
d
Sphagnum
magellanicum
7
d
Sphagnum compactum
1
c
Empetrum nigrum
2
d
Aulacomnium palustre
6
d
Sphagnum girgensohnii
1
c
Eriophorum
angustifolium
2
d
Hylocomium splendens
5
d
Sphagnum papillosum
1
c
Carex canescens
1
d
Hypnum cupressiforme
5
d
Tetraphis pellucida
1
Main factors influencing the species composition of vegetation
The RDA analysis of the separate effect of each of the environmental factors (marginal effect)
on the species composition of bryophytes showed that the factors with a significant impact were
hydrological parameters (minimum, maximum and average water level), shade, physical
parameters of the peat (proportion of macropores) and its chemical parameters (carbon content
and C/N ratio) (Table 2). The use of the forward selection procedure (conditional effect) in the
RDA showed that for the occurrence of bryophyte species the deciding factor was the average
water level, but a statistically significant increase in the percentage of variation explained was
obtained by also taking into account parameters of the peat such as the proportion of
macropores, carbon content, and pH in KCl. In the case of vascular plants, the factors
influencing the species composition (marginal effect) were also hydrological parameters
(minimum, maximum and average water level), peat thickness, and shade, and among the
chemical parameters of the peat, electrical conductivity, content of NH4, and the NO3/NH4 ratio.
Analysis of the conditional effect, however, showed that the water table coefficient included
the effect of all other factors analysed, so that taking them into account did not increase the
percentage of variation explained.
Table 2. Results of RDA based on the bryophytes subset and vascular plants subset for each
variable separately (Marginal effect) and after taking into account all variables (Conditional
effect). Statistical significance at *0.05; **0.01 (Monte Carlo permutation test)
Bryophytes
Vascular plants
Marginal
Conditional
Marginal
Conditional
Wmean
0.19**
0.19**
0.15**
0.03
macropores
0.11*
0.09*
0.09
0.03
C
0.19**
0.08*
0.08
0.05
pHKCl
0.08
0.06*
0.06
0.02
Wmax
0.12*
0.05
0.10*
0.03
EC
0.09
0.05
0.12*
0.02
peat depth
0.07
0.04
0.12*
0.07
mesopores
0.07
0.04
0.08
0.05
C/N
0.13*
0.04
0.07
0.02
N
0.03
0.04
0.03
0.02
W/D
0.14**
0.03
0.24**
0.24**
Wmin
0.15**
0.03
0.21**
0.02
Wamp
0.03
0.03
0.03
0.02
ash content
0.11
0.03
0.06
0.06
NO3/NH4
0.12
0.03
0.15**
0.05
micropores
0.04
0.03
0.06
0.02
shade
0.13*
0.02
0.10*
0.07
NO3
0.02
0.02
0.06
0.03
NH
0.08
0.02
0.11*
0.02
P2O5
0.05
0.02
0.05
0.00
Dependency of species composition of vegetation on environmental factors
The first DCA axis, interpreted on the basis of indicator values (EIV) (Table 3), was
strongly associated with fertility, moisture, reaction and shade. This axis was also correlated
with the occurrence of Eriophorum vaginatum, a vascular plant characteristic of raised bogs,
and with the Sp/B and Rb/V indices (Table 3). Ordination of plots and bryophyte species along
the first two axes on the DCA diagram (Fig. 3) showed that the occurrence of diagnostic
bryophyte species for raised bogs (Q11) was associated with the main gradient (first DCA
axis). They were accompanied by species with wider ecological amplitude, but typical for
peatland habitats, such as Sphagnum cuspidatum, S. fimbriatum, and Aulacomnium palustre.
The non-peatland species usually found in various types of habitats were connected with
opposite site of the first DCA axis. The second axis was not correlated with any factor enabling
ecological interpretation. This indicates that the differences in species composition of
bryophytes arise from the existence of one main habitat gradient.
Table 3. Pearson’s correlation coefficient between DCA ordination site scores (first two axes)
for the bryophytes subset and vascular plants subset and selected environmental parameters and
area cover by dominant species of vascular plants
Variable
Bryophytes
Vascular plants
DCA_1
DCA_2
DCA_1
DCA_2
Ordination axes
Bryophytes DCA_1
-0.11
0.60**
0.21
Bryophytes DCA_2
0.06
-0.03
Vascular plants DCA_1
0.02
EIV for bryophytes
L
-0.69**
-0.11
-0.11
-0.14
F
-0.94**
0.08
-0.72**
-0.18
R
0.69**
0.30
0.69**
0.33
N
0.72**
0.25
0.51*
0.26
EIV for vascular plants
L
-0.01
0.06
-0.07
0.22
F
-0.30
0.12
-0.26
0.57**
R
-0.21
-0.29
-0.56**
-0.08
N
-0.17
-0.32
-0.53*
0.14
Ratios
Sp/B
-0.89**
0.13
-0.61**
-0.20
Rb/V
-0.75**
0.05
-0.66**
-0.47*
Dominanat vascular species cover
Eriophorum vaginatum
-0.86**
0.09
-0.70**
-0.35
Ledum palustre
-0.23
-0.37
0.07
-0.23
Calluna vulgaris
0.26
0.17
0.28
0.51*
Vaccinium myrtillus
0.34
0.05
0.42
0.62**
Vaccinium uliginosum
0.30
-0.06
0.59**
-0.19
Vaccinium vitis-idaea
0.29
-0.03
0.60**
-0.55**
Explanations: EIV – Ellenberg Indicator Values: L – light; F – moisture; R – reaction; N – fertility.
Sp/B – number of Sphagnum species to all bryophytes species; Rb/V – number of raised bogs vascular
species to all vascular species. Significance level: ** P<0.01; * P<0.05
Fig. 3 Ordination DCA scatter plot based on bryophytes species. Squares – plots; green circles
– raised bog species (Q11); grey circles – non raised bog species. Only species with highest
weights are presented. Wmean – mean water table level, C – total carbon, macropor –
proportions of macropores, pHKCl – pH in KCL
In the case of vascular plants the first DCA axis was correlated with the fertility and reaction
of the substrate as well as with the percentage of certain vascular plants, while the second axis
was correlated with moisture and the percentage of non-peatland species (Table 3).
As the water level coefficient increases, there is a marked increase in the proportion of non-
peatland vascular plant species, especially trees and shrubs, and a decrease in peatland species
(Fig. 4).
Fig. 4 Ordination DCA scatter plot based on vascular plant species. Squares – plots; green
circles – raised bog species (Q11); grey circles – non raised bog species, WD – water table
coefficient.
Only the main gradients of variation (first DCA axis) for bryophytes and vascular plants were
correlated with each other (Table 3). The different directions of the main gradients (association
with different factors) are evidenced by significant correlations (except for L) between the first
DCA axis of vascular plants and EIV for bryophytes, while there is no such relationship for the
first DCA axis of bryophytes and EIV for vascular plants (Table 3). The first DCA axis of
bryophytes is much more strongly correlated with parameters describing the proportion of
peatland species (Sp/B and Rb/V). There were no significant correlations between the EIVs of
vascular plants and bryophytes, the Pearson correlation coefficient was below 0.4 and p > 0.5.
Discussion
The results of the study indicate that the occurrence of bryophytes and vascular plants in post-
mining areas of peat bogs depends on different factors. Similarly, different reactions of plants
to environmental factors were observed in undegraded peatlands in mountainous areas
(Bragazza & Gerdol, 1996; Hajkova & Hajek, 2004).
Relative to natural systems, however, the basic properties of the substrate are altered in post-
mining areas. The removal of the upper peat layer (the acrotelm) and the considerable decrease
in the thickness of the remaining deposit leads to the initiation of degradation processes. Soil
degradation resulting from drainage of peatlands significantly alters the physical and hydraulic
properties of peat soils (Liu & Lennartz, 2019). Depending on the water level and the resulting
rate of peat decomposition and mineralization, spatial variation in habitat conditions occurs.
This leads to better growth of bryophytes or vascular plants in different places. Due to the
different structure of these groups of plants and thus their different life strategies (Malmer et
al., 1994), their occurrence depends on different habitat factors.
In the case of bryophytes, a single main gradient of variation was found, corresponding to the
first DCA axis. It is associated with hydrological conditions, but also with the chemical and
physical parameters of the peat. A single main gradient of the species composition of
bryophytes was also reported by Hajkova & Hajek (2004). Bryophytes, especially Sphagnum,
have no root systems or well developed conductive tissues; they take up water and nutrients
with the entire surface of the body and therefore depend on a high water level in the substrate,
precipitation, or high humidity (van Breemen, 1995). The water level determines the redox
conditions in peat. A high water level, and thus poor oxygenation, inhibits the transformation
of organic matter. Access to air causes mineralization of organic matter, which increases access
to nutrients, mainly mineral forms of nitrogen (Andersen et al., 2011; Urban et al., 2018). The
physical parameters of the peat (its structure) change as well. The percentage of micropores
increases while that of macropores decreases (Rezanezhad et al., 2016; Liu & Lennartz, 2019),
which reduces infiltration and retention of water near the surface, and thus its availability for
Sphagnum mosses (Grosvernier et al. 1995, Okruszko et al. 1995), as they are unable to extract
water from peat with soil-water pressure below −100 mb) (Price & Whitehead, 2001). The main
direction of changes in the species composition of bryophytes is therefore away from very wet
habitats with low fertility and low pH and a higher proportion of macropores, i.e. conditions
similar to those prevailing on a raised bog (Proctor, 1995). On the plots with these properties
mainly Sphagnum mosses were present, which despite the differences in habitat requirements
between species (Rochefort, 2000; Robroek et al., 2007) have common traits (water retention
inside tissue/body, continues growth) that distinguish them from other bryophytes. Apart from
Sphagnum, mosses Polytrichum strictum and Aulacomnium palustre were recorded; these are
common species on raised bogs, which in post-mining areas can facilitate the growth of
Sphagnum (Grosvernier et al., 1995). At the other end of the gradient are mesophilic habitats,
where the water level was lower, and thus the degree of mineralization of organic matter, pH,
and fertility were higher. In these conditions various species of true mosses were present, having
relatively wide habitat amplitude and associated with forest habitats (e.g. Pleurozium schreberi)
(Hedwall et al., 2017). Similar changes in the occurrence of Sphagnum and brown mosses along
a gradient of moisture, fertility and pH were observed by Purre & Ilomets (2018).
Numerous studies (Lavoie et al., 2005; Bastl et al., 2009) have shown that the water level has a
decisive impact on the direction of succession in post-mining areas as well as at the edges of
natural peatlands (Bragazza et al,. 2005). In undisturbed raised bogs, where the hydrological
conditions are stable, the impact of the water level on the occurrence of Sphagnum is mainly
manifested in the case of extreme drops in the water level caused by droughts (Bragazza, 2008;
Walker et al., 2017).
In forest ecosystems the fundamental effect of trees is due to shading, through interception and
attenuation of shortwave radiation (Davies et al., 2019). In effect, the microclimate of the forest
interior is distinguished by smaller temperature amplitudes and higher humidity than in non-
forest areas (Chen et al., 1999). This creates favourable conditions for the growth of bryophytes,
which respond to the conditions prevailing on the soil surface, while soil fertility is particularly
important for vascular plants (Hokkanen, 2004). Vascular plants and bryophytes in grassland
also differ in their habitat requirements (Herben, 1987, Ingerpuu et al,. 1998). Compared to
vascular plants, bryophytes respond to factors of much shorter duration and greater fluctuations
(Herben, 1987). Moreover, they have different phenological development and are much more
abundant in cold, moist conditions (Al-Mufti et al., 1977). In the case of managed grassland, an
important factor differentiating the occurrence of bryophytes and vascular plants is fertilizer,
which negatively affects bryophytes but increases the biomass of vascular plants (mainly
grasses) (Boch et al., 2018).
In the case of vascular plants, no dominant gradient was observed. There were fewer factors
affecting species composition and the percentage of variation explained. Hydrological
conditions also play an important role in determining species composition, but in connection
with peat thickness. In our study, the most important parameter was the ratio of the average
water level to the peat thickness (water table coefficient). The influence of other factors, such
as peat parameters (both chemical and physical), was contained in the effect of the water table
coefficient. Tree shade, which also significantly influences the growth of plants in peatlands
(Limpens et al., 2014), was indirectly associated with the water table coefficient because low
peat thickness and a low water level are favourable to the growth of trees and shrubs (Holmgren
et al., 2015). Triisberg et al. (2014) also observed a significant effect of the thickness of the
remaining peat layer, which was associated with the values of other peat parameters (pH, ash,
nutrients content). For vascular plants, which take up water through their root system from
deeper layers of the substrate (Schouwenaars, 1993), a high water level is unfavourable because
root growth is inhibited by oxygen deficiency (Boggie et al., 1977; Wheeler & Shaw, 1995).
Vascular plants also make use of nutrients derived from organic matter mineralized in aerobic
conditions (Malmer et al., 1994). A thinner peat layer facilitates rooting of plants in the mineral
substrate. This means that analysis of the water table coefficient, taking into account changes
in its value over time after peat mining is terminated, can be useful in assessing whether a given
area has potential for spontaneous restoration or requires active restoration measures. The main
gradients of the species composition of bryophytes and vascular plants (first DCA axis) are only
partially correlated with one another. For bryophytes the main gradient is correlated with
fertility, the water level, and pH. For vascular plants the first axis is only correlated with fertility
and pH, and the second with the water level. This ecological interpretation of the axes and the
lack of correlation between the indicator values for bryophytes and vascular plants from the
same plots indicate that the two groups of plants utilize different resources of the environment
(Hajkova & Hajek, 2004). Individual species of vascular plants differ in their response to
environmental factors, as indicated by their correlations with the first or second DCA axis.
Among vascular plants, the main gradient of species composition for both bryophytes (first
DCA axis of bryophytes) and vascular plants (first DCA axis of vascular plants) is correlated
only with the occurrence of Eriophorum vaginatum. The different requirements of E. vaginatum
from those of other raised bogs species is also indicated by the correlation of the Rb/V index
with the second DCA of vascular plants. This diagnostic species for raised bogs is believed to
create favourable conditions for the growth of Sphagnum mosses (Grosvernier et al., 1995;
Rochefort, 2000; Pouliot et al., 2011; Kaštovská et al., 2018). The results of our study confirmed
that Eriophorum vaginatum protects Sphagnum and can be used as a companion plant in the
restoration process.
Conclusions
The occurrence of bryophytes in post-mining areas is in part determined by different habitat
factors than in the case of occurrence of vascular plants, due to their different life strategies.
The main factor influencing the species composition of bryophytes is the water level and the
associated degree of changes in the physical and chemical properties of the peat resulting from
degradation processes taking part under the conditions of a long-term decrease in the water
level.
The species composition of vascular plants in post-mining areas, apart from the water level, is
significantly influenced by the thickness of the peat layer remaining after extraction.
The use of the water table coefficient in peatland restoration practice can provide additional
information to be used in making decisions regarding the need for restoration measures.
Acknowledgments
The work was financed by the Ministry of Science and Higher Education in Poland for
Agriculture University in Krakow.
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Declaration of interests
☒ The authors declare that they have no known competing financial interests or personal
relationships that could have appeared to influence the work reported in this paper.
☐ The authors declare the following financial interests/personal relationships which may be
considered as potential competing interests:
