Ecological Engineering 189 (2023) 106916

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Ecological Engineering
journal homepage: www.elsevier.com/locate/ecoleng

The type of soil amendment during farming affects the restorability

of peatlands
Ming Wang a, b, c, d, Yixiao Wang a, c, d, Shengzhong Wang a, c, d, *, Ming Jiang b, d,
Guodong Wang b, d, **
a
Key Laboratory of Geographical Processes and Ecological Security in Changbai Mountains, Ministry of Education, School of Geographical Sciences, Northeast Normal
University, 5268 Renmin Street, Changchun 130024, PR China
b
Key Laboratory of Wetland Ecology and Environment, Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences, 4888 Shengbei Street,
Changchun 130102, PR China
c
State Environmental Protection Key Laboratory of Wetland Ecology and Vegetation Restoration, Institute for Peat and Mire Research, Northeast Normal University,
Changchun, 5268 Renmin Street, Jilin 130024, PR China
d
Jilin Provincial Joint Key Laboratory of Changbai Mountain Wetland and Ecology, Department of Science and Technology of Jilin Province, Changchun, 5268 Renmin
Street, Jilin 130024, PR China

A R T I C L E I N F O A B S T R A C T

Keywords: Peatlands have declined dramatically in the past century due to agricultural cultivation. The main goal of this
Soil seed bank study was to assess the impact of soil amendment during farming on the restorability of peatlands. We compared
Restoration potential the soil seed banks between the natural peatland and farmed peatlands with two soil amendment types (sand-
Peatland restoration
amended and silt-amended) in the Changbai Mountains, China. We found that the seed density and species
Soil amendment
Changbai Mountains
richness in silt-amended peatland were higher than the natural peatland, and they were the lowest in the sand-
amended peatland. Carex species are foundational species of these wetlands and these dominated both in
standing vegetation and soil seed banks of the natural peatland. Carex spp. occurred in the seed banks of silt-
amended peatland soils in low seed density, but were absent in the sand-amended peatland. The seed density
of peatland species was highest in the surface 10 cm soil depth in the natural peatland, while it was highest in the
10–20 cm soil depth in the silt-amended peatland. Redundancy analysis identified that soil C:N, soil water
content, and soil organic carbon explained most variance in seed bank composition. The result illustrates that soil
amendment types affect soil, and change soil seed banks and revegetation potentials of peatlands. The silt-
amended peatland had a higher revegetation potential than the sand-amended peatland. Removal of surface
silt mineral soil is necessary to reduce the weed seeds and promote the recolonization of peatland species in the
silt-amended peatland during restoration.

1. Introduction
Peatlands have been lost dramatically in the past century due to
agricultural cultivation (Hallema et al., 2015). Agricultural cultivation
not only degrades the native vegetation but also changes the hydro­
logical conditions and soil physicochemical characteristics in peatlands
(Berglund and Berglund, 2010; Kløve et al., 2010; Heller and Zeitz,
2012; Leifeld et al., 2019). In particular, organic soils are very vulner­
able to farming (Kalisz et al., 2015; Wang et al., 2021a). Peatland
development has obstacles such as high content of organic acids, high
soil water content, and low bulk density. Therefore, besides the removal
of aboveground vegetation and lowering of the water table, mineral soils
are usually filled in peatlands during farming to increase soil load-
bearing capacity and balance pH (Zakharova et al., 2020; Kalisz et al.,
2021; Wang et al., 2021a). The mineral materials included fine sands,
silts, and also clay, among which sands and silts were the most frequent
soil amendments (Zaidel'man et al., 2001). These materials can be ho­
mogenized with surface horizons of peatlands during cultivation or can

* Corresponding author at: Key Laboratory of Geographical Processes and Ecological Security in Changbai Mountains, Ministry of Education, School of
Geographical Sciences, Northeast Normal University, 5268 Renmin Street, Changchun 130024, PR China.
** Corresponding author at: Key Laboratory of Wetland Ecology and Environment, Northeast Institute of Geography and Agroecology, Chinese Academy of Sci­
ences, 4888 Shengbei Street, Changchun 130102, PR China.
E-mail addresses: szwang@nenu.edu.cn (S. Wang), wanggd@iga.ac.cn (G. Wang).

https://doi.org/10.1016/j.ecoleng.2023.106916
Received 4 June 2022; Received in revised form 22 January 2023; Accepted 29 January 2023
Available online 2 February 2023
0925-8574/© 2023 Published by Elsevier B.V.
M. Wang et al.
be partially leached into the deeper soil horizons (Kalisz et al., 2021).
Silt amendment hampered soil organic matter oxidation and CO2
emissions and had a stabilizing effect on SOM (Smolczynski et al., 2021;
Kalisz et al., 2021). Sand-amendment improved soil physical properties
for farming (Zakharova et al., 2020), but also increased the loss of peat
organic matter by 18%–28% through CO2 emission (Zaidel'man et al.,
2001). Therefore, the influence of soil amendment on soil varied
depending on soil amendment type.
In the restoration of disturbed habitats, the soil seed bank is an
important source to reestablish vegetation (Middleton, 2003; Wang
et al., 2021b). The number of viable seeds in the seed bank along with
seed dispersal determines the success of restoration practices (Moham­
med and Denboba, 2020; Wang et al., 2021b). If the seeds of key
structural dominants are absent from seed banks, farmed fields can be
difficult to restore to their original wetland type (Middleton, 1999;
Wang et al., 2015). Tussock sedges are foundational species in the
peatlands. Carex schmidtii dominates peatlands in the Changbai Moun­
tation and forms tussock structures that provide a carbon storage func­
tion (Wang et al., 2021c) and engineer communities by enhancing
microtopography and supporting biodiversity (Wang et al., 2019).
Therefore, a better understanding on the mechanisms determining seed
survival in soils, especially those wetlands dominated by tussock sedges,
is essential not only to understand plant community structures but also
to improve the strategies and techniques for peatland restoration. The
value of soil seed banks in restoration varies greatly according to
wetland type and human disturbance (Middleton, 2003; Wang et al.,
2016; Wang et al., 2020).
The Changbai Mountains include one of the largest peatlands in
China, with an area of 463.31 km2 and a SOC storage of peatlands of
47.64 Tg (Ma et al., 2013). Unfortunately, since the 1950s, large areas of
peatlands in this region were farmed as paddy fields. The area of peat­
lands has been greatly reduced and the original hydrology has been
destroyed (Li, 2013; Wang et al., 2019), resulting in a dramatic decline
in biodiversity and ecological services (Wang et al., 2020; Xu et al.,
2021). Recently, the Chinese government issued the National Wetland
Fig. 1. Site locations for seed bank samples in the Changbai Mountain region.
2
Ecological Engineering 189 (2023) 106916
Protection Law, which states that restoring the peatlands in China ac­
cording to their types and degradation status. The Jilin Provincial
Government developed plans for peatland restoration of >6000 ha of
farmland in the Changbai Mountains, and the Forestry Department
began restoring peatlands there. Our study compared the composition
and density of soil seed banks of natural peatlands to farmed peatlands
with two soil amendment types (silt-amended and sand-amended). We
asked the following research questions: (1) How does the composition of
seed banks change with the two soil amendment types and what is the
value of farmed seed banks for restoring the vegetation of peatlands? (2)
Which environmental factors influence the composition of soil seed
banks in these natural and farmed peatlands?
2. Materials and methods
2.1. Study site
We selected one undisturbed natural peatland and two farmed
peatlands (silt-amended and sand-amended) located in the west of the
Changbai Mountain (Fig. 1). This region is characterized by a temperate
continental monsoonal climate with a frost-free period of approximately
138 days per year. Annual mean precipitation in this area is 737.4 mm
and annual mean temperature is about 5 ◦ C (Qiao, 1993). The natural
peatland site locates in Huinan County, Tonghua City, Jilin Province
(126◦ 16′ 58′′ E, 42◦ 18′ 19′′ N, 560 m a.s.l.). The dominant species is the
tussock-forming Carex schmidtii. Sanguisorba tenuifolia, Thelypteris pal­
ustris, and Lythrum salicaria are main associated species. The peat
thickness in the study site ranges from 80 to 100 cm. The silt-amended
peatland site locates in Dayishan Town, Huinan County, Tonghua City,
Jilin Province (126◦ 13′ 10′′ E, 42◦ 16′ 35′′ N, 456 m a.s.l.). The peat aver­
ages 0.5–1 m in thickness in the study site. Carex schmidtii was the
dominant species before farming. In 1980s, most of the peatlands in this
region were drained and filled with silt soil to increase soil load-bearing
capacity and balance pH for agricultural purposes. As part of the soil
amendment process, about 10 cm of silt were covered on the surface of
M. Wang et al.
the peatlands. The sand-amended peatland was located in Sipeng town,
Tonghua County, Tonghua City, Jilin Province (125◦ 34′ 44′′ E,
41◦ 51′ 22′′ N, 560 m a.s.l.). The peat thickness in this study site ranges
from 0.6 to 1.0 m. Peatlands in this region were drained and intensively
reclaimed to paddy fields in 2000s, with introduction of sand in the
plowed horizon and mixing with peat when plowing.
2.2. Seed bank sampling
Sampling was conducted immediately after snowmelt (mid-April
2019) to assess the seed bank present at the time of spring germination.
At each site, 5 plots were randomly chosen (>1 km distant). Within each
plot, 5 quadrats (10 × 10 × 30 cm, length × width × depth) at least 20 m
apart were randomly chosen. The soil samples were sampled in three 10
cm soil sections. Soil samples were placed in plastic bags by depth and
site. A total of 225 soil samples were collected (3 sites × 5 plots/site × 5
quadrats/plot × 3 depths/quadrat).
2.3. Germination of soil seed banks
The soil samples were taken back to a greenhouse and sieved through
a coarse sieve to remove gravel, plant litter and roots from the soils. Soil
samples at the same layer (0–10, 10–20, 20–30 cm) from the 5 quadrats
within each plot were mixed thoroughly to create one sample per plot.
Each soil sample was spread as an even 2-cm layer in pots (25 × 25 cm)
filled with 8 cm of washed vermiculite. Pots were placed in a tank (150
× 200 cm) assigned to a moist soil treatment. The pots were all watered
from the bottom every week. Newly emerged seedlings were identified,
counted, and removed from the pots. Seed germination assays continued
for 6 months. Nomenclature follows Fu (1995).
2.4. Site environment and field standing vegetation survey
Standing field vegetation of seed bank sampling areas in the natural
peatland (25 plots) was sampled on 15 July 2020. To locate each plot,
starting in the center of the site, a 1-m2 quadrat frame was thrown from
the previous plot in a random direction (~10 m distant). Within each
plot, species name, density, height and percentage cover of each species
were recorded.
To quantify differences in soil properties, soil samples were
randomly collected in the three peatland sites. Five soil cores (diameter
5 cm, depth 30 cm, 10 cm interval) were taken from the seed bank
sampling areas in each site to measure soil water content (SWC), soil
bulk density (BD), soil organic carbon (SOC), soil total nitrogen (TN),
soil total phosphorus (TP), and soil pH. SWC was measured by the
gravimetric method (Jackson et al., 2000). BD was determined using the
volumetric core method. SOC was measured with the dichromate
oxidation method (Kalembasa and Jenkinson, 1973). TN and TP were
determined by the Kjeldahl method and Molybdenum blue method,
respectively, on an automated analyzer (Smartchem140, AMS-Alliance,
French). Soil pH was measured at a soil-to-water ratio of 1:10 using a
glass electrode (PHS-3E meter with E-201-C electrode, Leici, China).
2.5. Data analysis
Two-way analysis of variance (ANOVA) with Duncan's test were used
to examine the effects of site type, soil depth, and their interactions on
soil properties (SWC, BD, SOC, TN, TP, C:N, and pH), seed density and
species richness of soil seed banks, and the number of species in common
between natural peatland vegetation and soil seed banks. Significance
was determined at an α level of 0.05. Data were log-transformed to
satisfy the assumptions of the ANOVA. All statistical analyses were
conducted using SPSS 17.0.
We used the similarity index to compare the species similarity be­
tween the vegetation and seed banks, and used importance value to
investigate species importance in aboveground vegetation (Perry and
3
Ecological Engineering 189 (2023) 106916
Hershner, 1999).
Similarity index : I = 2c/(a + b) × 100%
Importance value : IV = RD + RC + RH,
where a and b are the number of species present in aboveground vege­
tation and soil seed bank, respectively. ‘c’ is the number of species
present both in aboveground vegetation and soil seed bank. The
importance value (IV) which is the sum of relative density (RD), relative
coverage (RC), and relative height (RH) of the plant community.
Redundancy analysis (RDA) was conducted to explore relationships
between species composition of soil seed banks and environmental
factors in natural and farmed peatlands using CANOCO 5.0. Variables
were log-transformed and centered to equalize the weight of variables
with ranges of different orders of magnitude. Nonmetric Multidimen­
sional Scaling (NMDS) with the Bray-Curtis coefficient of dissimilarity
matrix was used to assess the similarities of seed bank composition be­
tween the three peatland sites in CANOCO 5.0.
3. Results
3.1. Soil environmental factors between peatland sites
Soil characteristics differed between the three peatland sites. The soil
pH and BD in the sand-amended peatland were higher than the natural
and silt-amended peatland types (Table 1). The SWC, SOC, TN and TP in
natural peatland were higher than the silt-amended and sand-amended
peatlands. The C:N in silt-amended peatland was higher than the natural
and sand-amended peatlands. The values of pH, BD, SWC, SOC, TN, and
TP in the silt-amended peatland were generally between those of the
natural and sand-amended peatlands.
3.2. Composition, richness and density of soil seed banks
A total of 49 species of seeds germinated from the soils. There were
23 species in natural peatland, 33 species in silt-amended peatland, and
26 species in sand-amended peatland (Fig. 2). The species composition
differed between the three peatland sites. Carex species included
C. schmidtii, C. rhynchophysa and C. pseudocuraica in soil seed banks of
the natural peatland (Fig. 2). In silt-amended peatland, the Carex species
occurred with low seed density. The dominant species were Lindernia
procumbens, Pycreus flavidus, Eriocaulon robustius, and C. schmidtii. In
sand-amended peatland, Carex species were absent from the soil seed
banks, the dominant species were Monochoria korsakowii, Pycreus flavi­
dus, and Alisma plantago-aquatica (Fig. 2).
Site type and soil depth were related to seed density and species
richness of the soil seed banks (Table 2). The seed density and species
richness in silt-amended peatland were higher than the natural peatland
(Fig. 3a, b), and the seed density and species richness in the sand-
amended peatland were lower than the natural peatland (Fig. 3a, b).
The seed density and species richness decreased with the increasing soil
depth in the three peatland types (Fig. 3a, b). As the dominant species in
the aboveground vegetation, the seed density of Carex spp. was highest
in the natural wetland, while they were absent from the soil seed bank in
the sand-amended peatland (Fig. 3c). The seeds of Carex spp. were
mainly present in the top 10 cm soil depth in the natural peatland, while
they mainly retained in the 10–20 cm soil depth in the silt-amended
peatland (Fig. 3c, d).
3.3. The relationship of seed bank composition with environmental factors
RDA showed that the environmental factors explained 57% of the
total variation in species composition (Fig. 4). The variation explained
by the first axis in the RDA was 36.8%, whereas that explained by the
second axis was 14.2%. C:N explained most variation in species
M. Wang et al. Ecological Engineering 189 (2023) 106916

Table 1
Soil environmental characteristics in each soil layer (0–10, 10–20, 20–30 cm) in the natural, silt-amended and sand-amended peatlands. The different capital letters
represent significant differences between the three sites within the same soil layer. Different lowercase letters represent significant differences between the three soil
layers within the same peatland site. (p < 0.05). SWC: soil water content, BD: soil bulk density, SOC: soil organic carbon, TN: soil total nitrogen, TP: soil total
phosphorus, C:N: carbon to nitrogen ratio.
Site Soil layer (cm) pH SWC (%) BD (g/cm3) SOC (g/kg) TN (g/kg) C:N TP (g/kg)
Ca
Natural 0–10 (5.11 ± 0.01) (505.64 ± (0.28 ± 0.03)Ca (344.07 ± 8.96)Ab (23.34 ± 0.40)Aa (14.73 ± 0.20)Bc (1.57 ± 0.06)Aa
12.43)Aa
10–20 (5.02 ± 0.05)Cb (332.66 ± 3.18)Ac (0.30 ± 0.00)Ca (317.99 ± 0.38)Ab (19.05 ± 0.05)Ac (16.69 ± 0.04)Bb (1.39 ± 0.01)Ab
20–30 (4.90 ± 0.00)Cc (405.03 ± 5.55)Ab (0.27 ± 0.01)Ba (372.68 ± 3.68)Aa (20.55 ± (18.14 ± 0.28)Ba (1.23 ± 0.04)Ab
0.16)Ab
Silt-amended 0–10 (5.63 ± 0.01)Ba (146.18 ± 3.38)Bb (0.43 ± 0.03)Ba (196.69 ± (10.61 ± 0.16)Bb (18.60 ± (1.01 ± 0.04)Ba
13.14)Bb 1.27)Aa
10–20 (5.54 ± 0.04)Ba (153.83 ± 7.56)Bb (0.43 ± 0.03)Ba (229.71 ± (11.76 ± 0.71)Bb (19.49 ± (0.95 ± 0.05)Ba
15.84)Bb 0.26)Aa
20–30 (5.24 ± 0.05)Bb (206.26 ± 3.57)Ba (0.32 ± 0.04)Ba (373.72 ± 5.04)Aa (16.74 ± 0.59)Ba (22.41 ± (1.08 ± 0.03)Ba
0.61)Aa
Sand- 0–10 (6.17 ± 0.03)Aa (57.21 ± 1.27)Cb (1.00 ± 0.01)Aa (41.18 ± 0.51)Cb (3.55 ± 0.05)Cb (11.59 ± (0.85 ± 0.01)Cc
amended 0.03)Cb
10–20 (6.22 ± 0.05)Aa (51.74 ± 1.62)Cc (1.05 ± 0.01)Aa (42.49 ± 0.29)Cb (3.54 ± 0.02)Cb (12.00 ± (0.91 ± 0.01)Bb
0.03)Cb
20–30 (5.79 ± 0.16)Ab (76.91 ± 2.05)Ca (0.75 ± 0.03)Ab (106.74 ± 9.54)Ba (7.48 ± 0.44)Ca (14.20 ± 0.50)Ca (1.00 ± 0.03)Ba

Fig. 2. Relative abundance of species germinated from soil seed banks in five plots of the natural, silt-amended and sand-amended peatlands.

composition (23.4%), followed by SWC (15.8%), SOC (8.4%), TN

Table 2
(5.3%), and pH (1.6%) (Fig. 4; Table 3).
A summary of analysis of variance (ANOVA) on the effects of site type and soil
NMDS analysis indicated that natural peatland types were located in
depth for seed density and species richness of soil seed banks.
the upper left side of the ordination graph, whereas all the sand-
Seed density Species richness amended peatland types were in the lower left side of the ordination
df F P value df F P value graph (Fig. 5). The silt-amended peatland types were in the middle right
Site type 2 42.22 <0.01 2 122.19 <0.01 side of the ordination graph (Fig. 5).
Soil depth 2 19.22 <0.01 2 44.51 <0.01
Site type * Soil depth 4 4.49 <0.01 4 1.60 0.20

4
M. Wang et al.
Fig. 3. Seed density (a) and species richness (b) of soil seed banks in the natural, silt-amended, and sand-amended peatlands. (c) Seed density of Carex species in the
soil seed banks and its proportion to the total seed density in the natural and silt-amended peatlands.
3.4. Similarity between the soil seed banks and aboveground vegetation
Twenty-one species were recorded in the vegetation survey of the
natural peatland (Table 4). The natural peatland was dominated by
C. schmidtii (Table 4). The similarity index between aboveground
vegetation of the natural peatlands and the soil seed banks in the nat­
ural, silt-amended and sand-amended peatlands were 34.8%, 24.5%,
and 9.3%, respectively.
Species of vegetation and seeds in the seed bank were most similar in
the natural peatland (Fig. 6a). The number of common species between
natural peatland vegetation and the soil seed banks in the silt-amended
peatland was higher than the sand-amended peatland (Fig. 6a). The
proportion of seed density of common species to the total seed density
was >50% in the three soil depths in natural peatland, 10–30% in the
silt-amended peatland, while it was <10% in the sand-amended peat­
land (Fig. 6b).
4. Discussion
The cultivation of wetlands can reduce the number of seeds in soils
because of artificial drainage and/or direct vegetation removal (Wang
et al., 2016). Seeds of wetland species become uncommon in cultivated
fields because the seeds lose viability and are not replaced by standing
wetland vegetation (Middleton, 1999). Seed viability is affected by
storage conditions, dry laboratory storage of Carices resulted in reduced
viability by as much as 95% even when stored in cold conditions (Leck
and Schütz, 2005). In this study, the species richness and seed density of
soil seed banks in silt-amended peatlands were higher than the natural
peatlands, while the species richness and seed density in sand-amended
peatlands were lower than the natural peatlands (Fig. 3). In the silt-
amended peatland, the soils not only keep a portion of seeds of peat­
land species, but also have additional upland and agriculture weed
species such as Eriochloa villosa and Echinochloa crusgalli. These species
are common in paddy field and spread rapidly (Fried et al., 2018), which
5
Ecological Engineering 189 (2023) 106916
result in the increasing seed density in the soil seed bank of these species
in the silt-amended peatland.
A principal determinant of seed survival in soil is burial depth
(Bekker et al., 1998; Burmeier et al., 2010; Egawa, 2017). Seed survival
in soil generally decreases with an increase in burial depth (Houle et al.,
2001; Bonis and Grillas, 2002). Most viable seeds are concentrated in the
upper few centimeters of the soil (Bekker et al., 1998). Therefore, most
of the seed bank studies in wetlands focus on the upper 10 cm layer of
soils (Wang et al., 2016; Davies et al., 2018; Zhao et al., 2021). However,
in peatlands, viable seeds could remain at a depth up to 50 cm (McGraw,
1987). In northern peatlands, cool, wet, and anaerobic conditions with
slow microbial activity are beneficial for seed survival. The anaerobic
conditions increase with the soil depth increased (McGraw, 1987). In
this study, we also found viable seeds at a depth of 20–30 cm in both the
natural and silt-amended peatlands. The seed density of Carex spp. was
highest in 10–20 cm in the silt-amended peatland (Fig. 3); However,
seed density was very low in the deeper soil layer in sand-amended
peatland and Carex spp. disappeared in the soil seed banks (Fig. 3).
This pattern indicated that the effects of farming on soil seed banks were
limited at the top soil layer (0–10 cm) in the silt-amended peatland,
which may mean that the covered seed banks cannot germinate (Leck
and Schütz, 2005). However, the mixing of sands damaged the seed
bank in deep soils, perhaps by changing soil moisture conditions. In
addition, the number of common species between aboveground vege­
tation of the natural peatland and the soil seed banks in silt-amended
peatland was higher than the sand-amended peatland (Fig. 6a).
The seed density of common species was the highest at 10–20 cm soil
layer in silt-amended peatland (Fig. 6a) indicating that more peatland
species are retained in the soil seed banks in silt-amended peatland,
especially in the deeper soil layer (10–20 cm) under the mineral soils.
Therefore, knowing the type of soil amendment for agriculture is
important to determine the best way to restore the old field to wetland.
When restoring the silt-amended peatland, removal of surface silt min­
eral soil maybe an effective way to reduce the weed seeds and promote
M. Wang et al. Ecological Engineering 189 (2023) 106916

Fig. 5. Non-metric multidimensional scaling analysis (NMDS) ordination graph

using a Bray-Curtis coefficient of dissimilarity. Natural, silt-amended, and sand-
amended peatlands are represented by green circles, yellow circles, and red
triangles, respectively. (For interpretation of the references to colour in this
figure legend, the reader is referred to the web version of this article.)

Table 4
Species that emerged in the aboveground vegetation plots and their importance
values in the natural peatland site.
Fig. 4. RDA ordination plot of the seed bank composition of the natural, silt- Family Genus Species Importance value
amended, and sand-amended peatlands constrained by environmental factors. Cyperaceae Carex Carex schmidtii 0.467
SOC: soil organic carbon; SWC: soil water content; BD: bulk density; TN: total Rosaceae Spiraea Spiraea salicifolia 0.088
nitrogen; TP: total phosphorus; C:N: carbon to nitrogen ratio. Thelypteridaceae Thelypteris Thelypteris palustris 0.081
Note:1-Carex schmidtii；2-Carex rhynchophysa；3-Carex pseudocuraica；4- Rosaceae Sanguisorba Sanguisorba tenuifolia 0.059
Schoenoplectus triqueter；5-Eleocharis wichurae；6-Cyperus fuscus；7-Deyeuxia Rosaceae Filipendula Filipendula palmata 0.051
angustifolia；8-Cyperus orthostachyus；9-Pycreus sanguinolentus；10-Pycreus Lythraceae Lythrum Lythrum salicaria 0.047
Apiaceae Ostericum Ostericum maximowiczii 0.046
flavidus；11-Bolboschoenus planiculmis；12-Schoenoplectus juncoides；13-Schoe­
Betulaceae Betula Betula ovalifolia 0.023
noplectus mucronatus；14-Agrostis divaricatissima；15-Echinochloa crusgalli；16-
Hypericaceae Hypericum Hypericum monogynum 0.02
Eriochloa villosa；17-Lythrum salicaria；18-Erigeron canadensis；19-Centipeda Poaceae Phragmites Phragmites australis 0.019
minima；20-Lycopus lucidus；21-Epilobium hirsutum；22-Alisma plantago-aqua­ Onagraceae Epilobium Epilobium hirsutum 0.018
tica；23-Juncus papillosus； 24-Juncus effusus；25-Murdannia keisak；26-Lin­ Onocleaceae Onoclea Onoclea sensibilis 0.017
dernia procumbens；27-Bidens radiata；28-Typha latifolia；29-Typha Cyperaceae Eleocharis Eleocharis wichurae 0.016
angustifolia；30-Persicaria sieboldii；31-Polygonum viscosum；32-Ludwigia pros­ Lamiaceae Lycopus Lycopus lucidus 0.008
trata；33-Rorippa globosa；34-Eriocaulon robustius；35-Onoclea sensibilis；36- Sapindaceae Acer Acer tataricum 0.008
Viola arcuata；37-Thelypteris palustris；38-Hypericum japonicum；39-Cyperus Polygonaceae Polygonum Polygonum sagittatum 0.007
Apiaceae Saussurea Saussurea japonica 0.007
nipponicus；40-Linnaea borealis；41-Salix myrtilloides；42-Cirsium maackii；43-
Violaceae Viola Viola arcuata 0.005
Allium thunbergii；44-Poa annua；45-Glyceria triflora；46-Glyceria spiculosa；
Apiaceae Cicuta Cicuta virosa 0.005
47-Monochoria korsakowii；48-Spiraea salicifolia；49-Scirpus orientalis. Ranunculaceae Caltha Caltha palustris 0.004
Caryophyllaceae Lychnis Lychnis fulgensp 0.003

Table 3
Results of redundancy analysis model of soil seed bank composition explained by dominant sedges (e.g., Carex schmidtii, C. lasiocarpa) and grasses (e.g.,
environmental factors. Calamagrostis angustifolia) survived as seeds if farmed for <5 years,
Factors Explains % pseudo-F p therefore fields farmed for short periods of time are the best candidates
for wetland restoration (Wang et al., 2015). Certain important structural
C:N 23.4 18.1 0.002
SWC 15.8 15.1 0.002 components (tussock-forming Carex) are not retained in seed banks
SOC 8.4 9.1 0.002 when farmed for 6–15 years (Wang et al., 2017). However, in this study,
TN 5.3 6.4 0.002 the revegetation potential of peatlands was not fully affected by the
pH 1.6 2 0.04 duration of farming. Although farmed for over 30 years, the key struc­
BD 1.5 1.8 0.068
TP 1 1.2 0.266
tural dominant Carex species remained in soils as seeds in silt-amended
peatland, which may be due to the protection of the covered silt soils.
SOC: soil organic carbon; SWC: soil water content; BD: bulk density; TN: total Submergence may provide suitable storage conditions for prolonged
nitrogen; TP: total phosphorus; C:N: carbon to nitrogen ratio.
viability of Carex seeds (Leck and Schütz, 2005). The increasing in
aerobic environment and sand soil generally lead to an increase in the
the recolonization of peatland species. Top soil removal with heavy decomposition of organic matter and a significant change in soil phys­
machine has been done in restoration in Europe and the U.S. (Klim­ ical and chemical properties (Zaidel'man et al., 2001). Therefore,
kowska et al., 2007). At the same time, management, such as the active although farmed for a shorter period, the soil environment in sand-
revegetation of dominant Carex species, must be done to restore the amended peatlands had changed dramatically. The result illustrated
native plant diversity in the sand-amended peatland. that soil amendment type affects the revegetation potentials of peat­
Former studies showed that the duration of farming significantly lands, and that the silt-amended peatland has a higher revegetation
affected the seed bank viability, thus it can be used as a general indicator potential than the sand-amended peatland.
of restoration potential of sedge meadows (Wang et al., 2017). Most

6
M. Wang et al.
5. Conclusion
Peatland soils are usually amended during farming in northern
China. We assess the impact of soil amendment on the restorability of
peatlands. We found that the type of soil amendment during farming
affected the potential of peatland restoration. The silt-amended peatland
had higher species richness and seed density than the sand-amended
peatland, and the seed density of native Carex spp. was highest at
10–20 cm depth in the peat of the silt-amended peatland. Therefore,
knowing the type of soil amendment for agriculture is important to
determine the best way to restore the old field to wetland. Our study
suggested that the silt-amended peatland has a higher revegetation po­
tential than the sand-amended peatland. It is important to remove the
added silt, particularly because there may be peatland seeds in the lower
layer useful for restoration. Our study elucidated the vegetation
restorability of Carex-dominated peatlands after farming based on a site-
scale vegetation survey and a short-term seed bank germination exper­
iment in the Changbai Mountains, China. However, before broad-based,
general conclusions concerning the impact of soil amendment on the
restorability of peatlands can be made, long-term research are needed in
a diverse range of peatlands that differ by hydrology, plant species
composition, and soil nutrients, among other factors.
Funding
We thank Dr. Beth Middleton for suggestions and English editing.
This study was supported by the National Natural Science Foundation of
China (41871081, 42077070, 41877075, U19A2042), the Natural Sci­
ence Foundation of Jilin Province (20200201213JC), and the Youth
Innovation Promotion Association CAS (2019234).
CRediT authorship contribution statement
Ming Wang: Investigation, Software, Data curation, Visualization,
Writing – original draft, Writing – review & editing. Yixiao Wang:
Investigation, Writing – original draft. Shengzhong Wang: Methodol­
ogy, Writing – original draft, Writing – review & editing. Ming Jiang:
Writing – review & editing. Guodong Wang: Project administration,
Conceptualization, Methodology, Funding acquisition, Writing – orig­
inal draft, Writing – review & editing.
Declaration of Competing Interest
The authors declare no competing financial interests.
7
Ecological Engineering 189 (2023) 106916
Fig. 6. (a) The number of common
species between the natural peatland
vegetation and the soil seed banks in
the natural, silt-amended, and sand-
amended peatlands. (b) The rate of
seed density of common species to the
total seed density in the natural, silt-
amended and sand-amended peat­
lands. Different capital letters indi­
cate significant differences between
three peatland sites within the same
soil layer, different lowercase letters
indicate significant differences be­
tween three soil layers within the
same peatland site.
Data availability
Data will be made available on request.
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