Science of the Total Environment 922 (2024) 171218

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Science of the Total Environment

journal homepage: www.elsevier.com/locate/scitotenv

Freshwater wetland restoration and conservation are long-term natural

climate solutions
Lukas Schuster a, *, Pierre Taillardat b, Peter I. Macreadie a, Martino E. Malerba a
a
School of Life and Environmental Sciences, Deakin University VIC 3125, Australia
b
NUS Environmental Research Institute, National University of Singapore, Singapore 117411, Singapore

H I G H L I G H T S G R A P H I C A L A B S T R A C T

• Freshwater wetland restoration is an

effective way to turn degraded wetlands
from net carbon sources into carbon
sinks.
• Restored wetlands have a net cooling
effect on the climate on average 141 to
525 years after restoration.
• The radiative climate benefit of resto­
ration does not meet the timeframe set
by the Paris Agreement to limit warming
by 2100.
• The conservation and protection of nat­
ural freshwater wetlands should be pri­
oritised over wetland restoration.

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

Editor: Kuishuang Feng Freshwater wetlands have a disproportionately large influence on the global carbon cycle, with the potential to
serve as long-term carbon sinks. Many of the world’s freshwater wetlands have been destroyed or degraded,
Keywords: thereby affecting carbon-sink capacity. Ecological restoration of degraded wetlands is thus becoming an
Radiative forcing increasingly sought-after natural climate solution. Yet the time required to revert a degraded wetland from a
Net radiative cooling effect
carbon source to sink remains largely unknown. Moreover, increased methane (CH4) and nitrous oxide (N2O)
Greenhouse gas fluxes
emissions might complicate the climate benefit that wetland restoration may represent. We conducted a global
Net ecosystem exchange
Carbon sequestration meta-analysis to evaluate the benefits of wetland restoration in terms of net ecosystem carbon and greenhouse
Freshwater wetland gas balance. Most studies (76 %) investigated the benefits of wetland restoration in peatlands (bogs, fens, and
Teal carbon peat swamps) in the northern hemisphere, whereas the effects of restoration in non-peat wetlands (freshwater
Restoration marshes, non-peat swamps, and riparian wetlands) remain largely unexplored. Despite higher CH4 emissions,
Conservation most restored (77 %) and all natural peatlands were net carbon sinks, whereas most degraded peatlands (69 %)
Nature-based solutions were carbon sources. Conversely, CH4 emissions from non-peat wetlands were similar across degraded, restored,
and natural non-peat wetlands. When considering the radiative forcings and atmospheric lifetimes of the
different greenhouse gases, the average time for restored wetlands to have a net cooling effect on the climate
after restoration is 525 years for peatlands and 141 years for non-peat wetlands. The radiative benefit of wetland
restoration does, therefore, not meet the timeframe set by the Paris Agreement to limit global warming by 2100.

* Corresponding author.
E-mail address: l.schuster@deakin.edu.au (L. Schuster).

https://doi.org/10.1016/j.scitotenv.2024.171218
Received 14 August 2023; Received in revised form 23 November 2023; Accepted 21 February 2024
Available online 27 February 2024
0048-9697/© 2024 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
L. Schuster et al.
The conservation and protection of natural freshwater wetlands should be prioritised over wetland restoration as
those ecosystems already play a key role in climate change mitigation.
1. Introduction
Despite only covering an estimated 5–8 % of the Earth’s surface
(Mitsch et al., 2013), wetlands (such as freshwater marshes, peatlands,
and swamps) have a disproportionately large influence on the global
carbon cycle. Wetlands emit 20–25 % of global methane emissions, yet
they have the potential to serve as long-term carbon sinks (Mitsch et al.,
2013; Mitsch and Mander, 2018; Rosentreter et al., 2021). At the global
scale, freshwater wetlands are estimated to sequester between 0.7 and
1.75 Pg C year− 1 (0.7 to 1.75 × 109 t C year− 1) (Lal et al., 2018).
Importantly, teal carbon ecosystems like freshwater wetlands retain
large parts of the sequestered carbon in their soils for centuries to
millennia as anoxic soil conditions slow microbial decomposition
(Richardson and Vepraskas, 2001). Overall, the carbon stored within
freshwater wetland soils has one of the highest carbon densities among
all terrestrial ecosystems, constituting one-third of the world’s total soil
carbon stocks (Cole et al., 2007; Kayranli et al., 2010).
In recent centuries, competition for space due to increased urbani­
sation and agriculture has caused a decline in the extent and the con­
dition of freshwater wetlands. An estimated 21 % of the global
freshwater wetland area (3.4 million km2) has been lost due to drainage
and land-use conversion between 1700 and 2020 (Fluet-Chouinard
et al., 2023). Crucially, the degradation of natural wetlands is causing
the release of significant quantities of previously stored soil carbon as
carbon dioxide (CO2) and, to a lesser extent, methane (CH4), effectively
turning natural wetlands from carbon sinks into carbon sources (Lal and
Pimentel, 2008; Lane et al., 2016).
With global warming intensifying, the preservation of natural wet­
lands and the restoration of degraded ones have become increasingly
popular natural climate solutions to help mitigate climate change
(Bossio et al., 2020). Unlike other terrestrial ecosystems, freshwater
wetlands (particularly peatlands) play a key role in reducing atmo­
spheric carbon as their carbon storage potential does not reach satura­
tion (Vepraskas and Craft, 2016). Beyond their role as natural climate
solutions, freshwater wetlands offer several other benefits, such as
providing habitat for a range of species, improving water quality,
increasing water security, and protecting from floods (Chausson et al.,
2020; Thorslund et al., 2017). Freshwater wetlands are, therefore, a
comprehensive nature-based solution to mitigate the impacts of climate
change.
There are several ways to restore degraded wetlands, including
rewetting, active revegetation, and fencing. Rewetting is a common
management intervention that is used to reinstate the natural hydro­
logical connectivity of wetlands previously disconnected from their
original waterway (Limpert et al., 2020; Meng et al., 2020). This
approach often relies on redesigning the hydrology of degraded wet­
lands (e.g., digging out soil or sediment to create hummocks and tus­
socks) and the blocking of ditches and drains to deliver water to dried
wetland areas to stimulate the development of wetland vegetation
(Baldwin et al., 2018; Kayranli et al., 2010; Schwieger et al., 2021).
Nevertheless, extended periods of dry conditions may cause the deple­
tion of seed banks, thus preventing the return of native species of
wetland plants following rewetting (Zedler, 2000). Restoration through
rewetting is therefore more effective when combined with active
revegetation, which involves the planting of seeds or plantlets of native
wetland species in the restored wetlands (Spieles, 2022). In cases of
severe wetland degradation, restoration efforts may also include the
removal of non-native plant species such as farmed trees or the addition
of topsoil to increase nutrient and substrate availability (Scott et al.,
2020; Spieles, 2022). The fencing approach, conversely, is a passive
approach to restoring wetlands through livestock exclusion. The
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Science of the Total Environment 922 (2024) 171218
presence of livestock can have several adverse effects on freshwater
wetland ecosystems. For example, high-intensity grazing can reduce
vegetation cover, thus substantially reducing a wetland’s potential for
carbon sequestration and storage (Morris and Reich, 2013). Similarly,
the presence of livestock can increase greenhouse gas emissions due to
an increased nutrient input from manure (Bonetti et al., 2022). Limpert
et al. (2021) found that excluding livestock grazing within wetlands in
south-eastern Australia significantly increased soil carbon concentra­
tions and lowered carbon emissions. Although these restoration tech­
niques are well established, the time required to return a degraded
wetland to its natural conditions and functions remains largely unclear.
Restoring degraded wetlands often produces an increase in methane
(CH4) and nitrous oxide (N2O) emissions, complicating the assessment
of wetland restoration as a natural climate solution (Gutknecht et al.,
2006; Malerba et al., 2022; Serrano-Silva et al., 2014; Van Cleemput
et al., 2007). Importantly, for freshwater wetlands to be an effective,
long-term climate solution, the carbon they sequester and store must be
greater than the CH4 and N2O they emit in terms of radiative forcing.
Recent models predict that the increased CH4 and N2O emissions
following restoration should be fully compensated by the concomitant
increase in the net CO2 uptake within 40–80 years after restoration
(Günther et al., 2020; Zou et al., 2022). Empirical long-term studies
investigating the greenhouse gas and carbon sequestration dynamics of
restored wetlands, however, are rare. Nevertheless, wetland restoration
will likely generate an initial warming effect on the environment as CH4
and N2O emissions overpower the cooling effects from carbon seques­
tration. As the warming potentials of CH4 and N2O in the atmosphere
decrease over time, restored wetlands will switch from having a net
radiative warming to having a net cooling effect on the climate (Nyberg
et al., 2022). It is thus important to consider the true effects of CH4 and
N2O to accurately assess the role of wetland restoration for climate
change mitigation.
One way to estimate the net radiative effects of restored wetlands is
to use the ‘switchover time’ framework (Neubauer, 2014; Neubauer and
Megonigal, 2015). The switchover time accounts for the time since CH4
and N2O emissions and CO2 sequestration have occurred and thus allows
determining how long it takes for a restored wetland to have a net
radiative cooling effect on the climate (Mitsch et al., 2013; Taillardat
et al., 2020). Considering the time since emissions have occurred is
particularly important since different greenhouse gases vary in their
atmospheric perturbation lifetimes (Neubauer and Megonigal, 2015;
Pierrehumbert, 2014). For example, it takes 12.4 years for a molecule of
CH4 to be oxidised to CO2, whereas the breakdown of N2O into nitrogen
and oxygen through photolysis takes 121 years (Myhre et al., 2013).
Other greenhouse gas metrics – such as the global warming potential
(GWP), sustained GWP, or GWP* – are unable to estimate the time
required for an ecosystem to have a net radiative cooling effect on the
climate as such metrics fail to consider the time since greenhouse gas
emissions have occurred and rather integrate the effect of greenhouse
gas breakdown in the atmosphere for a fixed period of time (i.e., 100
years under the United Nations Framework Convention on Climate
Change, UNFCCC). Estimating the switchover time of a restored wetland
based on empirically measured CO2 sequestration and CH4 and N2O
emission rates is a powerful approach to evaluate the radiative role of
restored freshwater wetlands in climate change mitigation.
In this work, we compiled published data and performed a global
meta-analysis to determine the effects of freshwater wetland restoration
on (1) the greenhouse gas exchange (CO2 emissions or ecosystem
respiration Re, CH4 and N2O fluxes) and net CO2 sequestration (net
ecosystem exchange NEE, which takes into account CO2 emissions and
CO2 uptake through photosynthesis), (2) the net ecosystem carbon
L. Schuster et al.
budget (NECB), which allowed us to determine whether wetlands are
net carbon sinks or sources, and (3) the net ecosystem radiative balance
of restored wetlands. To do so, we compiled data from studies
comparing greenhouse gas fluxes and carbon sequestration rates be­
tween restored wetlands and at least one degraded and/or natural
wetland. We then calculated the switchover time of restored wetlands to
determine the time required for these wetlands to have a net cooling
effect on the climate after restoration.
2. Methods
2.1. Literature search
We systematically searched the Web of Science database Core
Collection in September 2022 using the following search terms in the
‘topic’ field (title, abstract, and extended keywords): ((greenhouse gas*
OR GHG* OR “carbon dioxide” OR CO2 OR “net ecosystem productivity” OR
NEP OR “net ecosystem exchange” OR NEE OR methane OR CH4 OR
“nitrous oxide” OR N2O or flux* OR emission* OR “global warming po­
tential” OR GWP OR CO2e OR CO2-e OR CO2equivalent OR CO2-equiva­
lent) AND (wetland* OR peat* OR bog* OR fen OR fens OR swamp* OR
mire* OR floodplain* OR marsh* OR freshwater marsh*) AND (restor* OR
re-establish* OR rewet* OR re-wet*) NOT (salt marsh OR saltmarsh OR
mangrove OR constructed OR lake OR reservoir OR tidal OR mesocosm* OR
laboratory)).
The initial search returned 1037 publications after excluding edito­
rials and retracted publications. We then screened all publications by
title and abstract and selected suitable studies that (1) appeared to assess
the impact of wetland restoration on greenhouse gas fluxes, and (2)
quantified greenhouse gas fluxes in situ. During the first-level screening,
we removed 712 publications that did not match these criteria, leaving
313 records to be assessed as full-text publications. During the second-
level screening, we only retained publications that included at least
one control wetland (natural and/or degraded), leaving 64 studies. We
were only interested in studies that used a paired experimental design (i.
e., where treatment and control wetlands had similar eco-
geomorphological characteristics and were located in similar
geographic and climatic regions), leaving 63 studies that had appro­
priate, extractable data for this meta-analysis.
2.2. Data extraction
From each study, we extracted yearly ecosystem respiration (Re in g
CO2 m− 2 year− 1), net ecosystem exchange (NEE in g CO2 m− 2 year− 1,
where a negative NEE indicates net CO2 uptake, a positive NEE indicates
net CO2 emissions from a given wetland) and net fluxes for methane
(CH4 in g CH4 m− 2 year− 1) and nitrous oxide (N2O in g N2O m− 2 year− 1)
for restored, degraded, and natural wetlands. In cases where studies
reported daily fluxes measured during the growing season only (42
studies), we used linear regressions calibrated with data from studies
that reported both daily and yearly emissions to convert these estimates
to yearly values (8 studies; Table S1). For studies that repeatedly
measured greenhouse gas fluxes over time (16 studies), we calculated
the average flux across the measurement years and reported the average
wetland age since restoration.
We extracted additional information from each publication including
wetland type, study location (latitude and longitude), annual mean
temperature, annual mean precipitation, wetland age (time since
restoration in years), hydrology, and the dominant vegetation present at
each study site. If the latitude and longitude were not reported, we
obtained these data using Google Maps according to the site information
described in the study. Similarly, if climatic data was not reported, we
used the National Aeronautics and Space Administration (NASA) Lang­
ley Research Center (LaRC) Prediction of Worldwide Energy Resource
(POWER) Project according to the reported location of the study.
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Science of the Total Environment 922 (2024) 171218
2.3. Statistical analyses
Overall, 76 % of all studies included in our meta-analysis investi­
gated the effects of wetland restoration in peatlands such as bogs (45
studies), fens (3 studies) and tropical peat swamps (1 study), whereas
only few studies focused on freshwater marshes (9 studies), riparian
wetlands (3 studies), or non-peat swamps (2 studies). We therefore
categorised these wetland subtypes into two overarching wetland types:
‘peatlands’ (bogs, fens, and peat swamps) and ‘non-peat wetlands’
(freshwater marshes, riparian wetlands, and non-peat swamps) to be
able to test for any differences in the effects of wetland restoration on
greenhouse gas fluxes between these wetland types.
To compare greenhouse gas fluxes between restored and control
wetlands (i.e., degraded vs. restored or natural vs. restored wetlands)
and wetland types (peatlands vs. non-peat wetlands), we used linear
mixed-effect models using the R package ‘nlme’ v.3.1-162 (Pinheiro and
Bates, 2023). We log10-transformed the response variable (yearly Re,
NEE, CH4, or N2O fluxes) where appropriate to conform with assump­
tions of linearity. When standardised residuals showed unequal vari­
ances, we included treatment-specific variance coefficients in the model
(function varIdent). We also included study ID as a random effect in all
analyses. If we found a significant treatment × wetland type interaction,
we used the R package ‘emmeans’ v.1.8.6 (Lenth, 2023) to conduct
tukey-adjusted post hoc tests.
To determine the net ecosystem carbon budgets (NECB) of restored,
degraded, and natural wetlands, we converted their average NEE (in kg
CO2 m− 2 year− 1) and CH4 emissions (in kg CH4 m− 2 year− 1) to kg C m− 2
year− 1 using the relative molecular weight of C in CO2 (27.27 %) and
CH4 (75.19 %), respectively, and calculated NECB as the sum of average
NEE (kg C m− 2 year− 1) and CH4 emissions (kg C m− 2 year− 1). Note that
our NECB estimates do not consider any aquatic export of carbon from
the freshwater wetlands as none of the studies included in our meta-
analysis reported such estimates.
To test whether the NEE of restored wetlands changes with wetland
age after restoration, we extracted data from studies that repeatedly
measured NEE of restored wetlands over several years and fit a linear
mixed-effects model with NEE (log10-transformed) as the response var­
iable, wetland age (in years) as the predictor variable, and wetland ID as
a random effect in the model.
2.4. Switchover time calculations
To calculate the switchover time of restored peatlands and non-peat
wetlands, we only considered wetlands that readily had a NEE < 0 (n =
37 out of 51 restored peatlands, n = 1 out of 1 restored non-peat
wetland), as wetlands with a NEE > 0 (CO2 emissions > CO2 uptake)
cannot have a negative radiative balance (Taillardat et al., 2020). We
used linear mixed effects intercept models for peatlands and non-peat
wetlands, respectively, with NEE, CH4, or N2O fluxes as the response
variable and study ID as the random effect in all analyses. We then used
the intercept estimate from each model to calculate the cumulative
radiative forcings (CO2 sequestration + CH4 emission + N2O emission)
of restored wetlands over time using equations presented in Neubauer
(2014) and Neubauer and Megonigal (2015). Briefly, we first calculated
the instantaneous radiative forcing of CH4 and N2O emissions over time
by taking into account their atmospheric lifetimes, and the decrease in
radiative forcing over time due to net CO2 uptake (i.e., NEE < 0). We
then summed the radiative forcings of CO2 sequestration, CH4 and N2O
emissions to estimate the switchover time.
We performed all analyses in R v.4.2.2 (R Core Team, 2013). For data
visualisation and plotting, we used the R package ‘ggplot2’ v.3.4.0
(Wickham, 2016).
L. Schuster et al. Science of the Total Environment 922 (2024) 171218

3. Results closed chamber technique, with 8 studies relying on eddy covariance

3.1. Data description
Our global meta-analysis revealed 63 studies that quantified the
greenhouse gas fluxes of 100 restored wetlands (including 17 wetlands
that were monitored repeatedly over time, resulting in a total of 124 site-
year flux measurements) paired with at least one degraded and/or
natural wetland (control; n = 57 degraded wetlands, n = 44 natural
wetlands; Tables S2 and S3). These studies were conducted across 69
locations worldwide, with 64 wetland sites (93 %) located in the
northern hemisphere, mainly in North America and Europe (Fig. 1). We
found that 79 % (50 out of 63) of all studies were published in the last 10
years (Fig. S1), indicating growing scientific interest in the effects of
freshwater wetland restoration on greenhouse gas fluxes. Most studies
reported CO2 emissions (ecosystem respiration Re; 46 studies) and
methane fluxes (44 studies). Nevertheless, there has been an increase in
the number of studies that quantified the net ecosystem exchange (NEE)
of restored wetlands in recent years (31 studies) to gain a complete
picture of the net greenhouse gas fluxes. Studies reporting N2O fluxes,
however, are still rare (16 studies; Fig. S2). Overall, 55 of the 63 studies
(87 %) measured greenhouse fluxes from freshwater wetlands using the
data.
The most common types of restored freshwater wetlands were bogs
(ombrotrophic peatlands; n = 79 restored wetlands) and freshwater
marshes (n = 12). Other types included fens (mineral-rich peatlands; n
= 3), peat (n = 1) and non-peat swamps (n = 2), and riparian wetlands
(n = 3; Fig. S3). Due to the low number of replication within most of
these wetland subtypes, we categorised all wetlands into two over­
arching wetland types: ‘peatlands’, which are characterised by peat soils
and typically dominated by mosses (e.g., bogs and fens) and ‘non-peat
wetlands’, which do not typically have peat soils and are dominated by
vascular plants (e.g., freshwater marshes and riparian wetlands) to be
able to uncover any potential differences in restoration effects between
these types. Regarding restoration techniques, the most common ap­
proaches to restoring degraded wetlands were rewetting (n = 64 peat­
lands; n = 9 non-peat wetlands) and rewetting combined with re-
vegetation (n = 14 peatlands; n = 4 non-peat wetlands). Some wet­
lands were restored using fencing (n = 4 non-peat wetlands; n =
0 peatlands) or were abandoned and left to recover on their own (n = 5
peatlands; n = 0 non-peat wetlands; Fig. S4). The age of restored wet­
lands since restoration ranged between 1 month and 65 years (Fig. S5).
Most restored peatlands (71 %, n = 59 out of 83) were 10 years or less in

Fig. 1. Global distribution of studies that assessed the net greenhouse gas fluxes (net ecosystem exchange NEE, methane CH4, and/or nitrous oxide N2O) of restored
freshwater wetlands paired with at least one control wetland (degraded and/or natural wetlands). Circles indicate studies that investigated greenhouse gas fluxes
from peatlands, triangles indicate studies that investigated fluxes in non-peat wetlands.

4
L. Schuster et al.
age, with a median wetland age of 6.75 years. Similarly, 59 % of restored
non-peat wetlands (n = 10 out of 17) were 10 years or less in age, with a
median wetland age of 9 years (Fig. S5).
3.2. Greenhouse gas fluxes: degraded vs. restored wetlands
On average, CO2 emissions (ecosystem respiration, Re) from restored
wetlands (peatlands: 1.12 kg CO2 m− 2 year− 1, n = 45; non-peat wet­
lands: 1.54 kg CO2 m− 2 year− 1, n = 9) were not significantly different to
emissions from degraded wetlands (peatlands: 1.4 kg CO2 m− 2 year− 1, n
= 31; non-peat wetlands: 1.89 kg CO2 m− 2 year− 1, n = 8; F1,59 = 1.78, P
= 0.19; Fig. 2a) in both peat and non-peat wetlands (treatment ×
wetland type: F1,58 = 0.01, P = 0.92). Restored peatlands, however, had
overall higher CO2 sequestration rates (more negative NEEs) than
degraded ones, which were on average small carbon sources (restored
peatlands: − 0.19 kg CO2 m− 2 year− 1, n = 35; degraded peatlands: 0.06
kg CO2 m− 2 year− 1, n = 29; F1,42 = 11.1, P = 0.002; Fig. 2b). Altogether,
the proportion of peatlands with a negative NEE (i.e., CO2 uptake
through photosynthesis > CO2 emissions) was more than twice as high
for restored peatlands (77 %, n = 27 out of 35) compared to degraded
ones (31 %, n = 9 out of 29; Fig. 2b). The overall higher CO2 uptake in
Fig. 2. (a) Ecosystem respiration (Re), (b) net ecosystem exchange (NEE), (c) methane (CH4) fluxes, and (d) nitrous oxide (N2O) fluxes from degraded vs. restored
freshwater wetlands. Black points represent the means ±95 % confidence intervals, coloured points are the raw data, asterisks indicate significant differences be­
tween treatments.
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Science of the Total Environment 922 (2024) 171218
restored peatlands, in turn, resulted in an average reduction of CO2
emissions of 0.25 kg CO2 m− 2 year− 1. Contrary, for non-peat wetlands,
we could not test for differences between degraded (n = 2) and restored
wetlands (n = 1) due to low data availability (mean NEE in restored non-
peat wetlands: − 0.16 kg CO2 m− 2 year− 1; degraded non-peat wetlands:
− 0.42 kg CO2 m− 2 year− 1).
For restored peatlands, we found a negative relationship between
NEE and wetland age since restoration (F1,15 = 4.51, P = 0.05; Fig. 3).
This finding suggests that restored peatlands with a NEE > 0 (CO2
emissions > CO2 uptake) should eventually transition to net CO2-
sequestering systems (NEE < 0) over time. Overall, 74 % (n = 26 out of
35) of all peatlands were net CO2-sequestering systems within 5 years of
restoration.
Methane fluxes from peatlands were on average 530 % higher from
restored wetlands (0.016 kg CH4 m− 2 year− 1, n = 38) than degraded
ones (0.003 kg CH4 m− 2 year− 1, n = 33; pairwise t-test: t = − 6.72, P
≤0.0001). Methane fluxes from non-peat wetlands, however, were not
significantly different between restored (0.0015 kg CH4 m− 2 year− 1, n =
7) and degraded wetlands (0.0014 kg CH4 m− 2 year− 1, n = 6; t = − 0.03,
P = 1; Fig. 2c). Conversely, nitrous oxide (N2O) fluxes from restored
wetlands (peatlands: 0.0003 kg N2O m− 2 year− 1, n = 8; non-peat
L. Schuster et al.
Fig. 3. Predicted lines of best fit for each restored wetland from linear mixed effects models (with 95 % confidence intervals) for the relationship between NEE and
wetland age (time since restoration in years) in peatlands. Data points represent observed values, each line represents a wetland with a common slope and a unique
intercept. The solid black line indicates the overall relationship between the NEE of restored wetlands and wetland age.
wetlands: 0.0002 kg N2O m− 2 year− 1, n = 4) tended to be lower than
fluxes measured from degraded ones (peatlands: 0.0007 kg N2O m− 2
year− 1; non-peat wetlands: 0.0005 kg N2O m− 2 year− 1, n = 8 for peat­
lands, n = 4 for non-peat wetlands; F1,11 = 2.9, P = 0.09; Fig. 2d).
3.3. Greenhouse gas fluxes: restored vs. natural wetlands
Average CO2 emissions tended to be lower from restored (Re; 1.19 kg
CO2 m− 2 year− 1, n = 42) than natural peatlands (1.45 kg CO2 m− 2
year− 1, n = 28; F1,48 = 2.9, P = 0.09; Fig. 4a). Natural peatlands,
however, had an overall higher CO2 uptake (− 0.19 kg CO2 m− 2 year− 1,
n = 24) compared to restored peatlands (− 0.11 kg CO2 m− 2 year− 1, n =
30; F1,38 = 4.29, P = 0.04; Fig. 4b), with all natural peatlands (n = 24)
sequestering more CO2 than they emitted (NEE < 0) compared to 67 %
(n = 20 out of 30) of restored wetlands that had a NEE < 0. Contrary,
due to no or low availability of Re (n = 3 for both restored and natural
non-peat wetlands) and NEE (no data available) measurements in non-
peat wetlands, we could not test for differences between restored and
natural non-peat wetlands (mean Re reported from restored non-peat
wetlands: 1.33 kg CO2 m− 2 year− 1; natural non-peat wetlands: 1.39 kg
CO2 m− 2 year− 1).
CH4 fluxes were similar across restored (peatlands: 0.02 kg CH4 m− 2
year− 1, n = 26; non-peat wetlands: 0.01 kg CH4 m− 2 year− 1, n = 5) and
natural wetlands (peatlands: 0.02 kg CH4 m− 2 year− 1, n = 19; non-peat
wetlands: 0.01 kg CH4 m− 2 year− 1, n = 5; F1,35 = 0.19, P = 0.66; Fig. 4c)
for both peatlands and non-peat wetlands (treatment × wetland type:
F1,34 = 0.23, P = 0.63). N2O fluxes in non-peat wetlands were also
similar between restored (0.0002 kg N2O m− 2 year− 1, n = 7) and natural
wetlands (0.0001 kg N2O m− 2 year− 1, n = 7; F1,8 = 0.32, P = 0.59;
Fig. 4d), whereas we could not test for differences in N2O fluxes between
restored and natural peatlands due to low data availability (n = 2 for
both restored and natural peatlands; average reported N2O emissions
from restored peatlands: 0.0009 kg N2O m− 2 year− 1; natural peatlands:
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Science of the Total Environment 922 (2024) 171218
0.0006 kg N2O m− 2
year− 1; Fig. 4d).
3.4. Net ecosystem carbon budgets (NECB)
On average, degraded peatlands were net carbon sources (0.02 kg C
m− 2 year− 1), whereas restored peatlands were net carbon sinks (− 0.02
kg C m− 2 year− 1), with 77 % (n = 27 out of 35) of restored peatlands
taking up more carbon than they emitted. All natural peatlands were net
carbon sinks (− 0.03 kg C m− 2 year− 1).
For non-peat wetlands, both degraded (− 0.11 kg C m− 2 year− 1) and
restored wetlands (− 0.04 kg C m− 2 year− 1) were carbon sinks. Note,
however, that these estimates are based on a very limited number of NEE
measurements that are available in the literature (degraded wetlands: n
= 2; restored wetlands: n = 1; natural wetlands: no data), resulting in
high uncertainty around these estimates.
3.5. Switchover time of restored wetlands
We estimated that, for restored wetlands that were net CO2-seques­
tering systems (NEE < 0), the average time required to switch from
having a net warming to a net cooling effect is 525 years for peatlands
and 141 years for non-peat wetlands after restoration (Fig. 5a, b). Due to
a sharp increase in CH4 emissions from peatlands after restoration (see
above), these wetlands will have a strong net radiative warming effect
on the climate for the first 268 years (up to ~4.65 pW m− 2), after which
they start to have a radiative cooling effect and turn into net radiative
sinks on average after 525 years. Contrastingly, non-peat wetlands have
a comparatively weaker net radiative warming effect on the climate for
the first 76 years (up to ~0.18 pW m− 2), turning into net radiative sinks
on average after 141 years. At the millennial time scale, however,
peatlands become more effective at cooling the atmosphere than non-
peat wetlands. For example, 1400 years after restoration, the cumula­
tive radiative forcing of peatlands is − 76.9 pW m− 2 compared to
L. Schuster et al.
Fig. 4. (a) Ecosystem respiration (Re), (b) net ecosystem exchange (NEE), (c) methane (CH4) fluxes, and (d) nitrous oxide (N2O) fluxes from restored vs. natural
freshwater wetlands. Black points represent the means ±95 % confidence intervals, coloured points are the raw data, asterisks indicate significant differences be­
tween treatments.
2
− 66.57 pW m− for non-peat wetlands (Fig. 5b).
4. Discussion
Our global meta-analysis revealed that the majority of studies (76 %)
investigating the effects of freshwater wetland restoration were con­
ducted in peatlands (bogs, fens, and peat swamps) in the northern
hemisphere (particularly in Europe, the US, and Canada; c.f. Fig. 1). Less
information is available on the effects of restoration in tropical peat­
lands, peatlands in the southern hemisphere, and global non-peat wet­
lands (e.g., freshwater marshes, non-peat swamps, and riparian
wetlands). Overall, our meta-analysis showed that most restored peat­
lands (77 %) were net carbon sinks, whereas most degraded peatlands
(69 %) were carbon sources. For those restored wetlands that were net
CO2-sequestering systems (NEE < 0), the average time before they have
a net radiative cooling effect on the climate (i.e., the switchover time) is
525 years for peatlands and 141 years for non-peat wetlands after
restoration. Our findings thus indicate that freshwater wetland resto­
ration effectively reduces carbon emissions at least from degraded
peatlands, with most peatlands (74 %) sequestering more CO2 than they
emit within 5 years of restoration. Nevertheless, it takes up to five
7
Science of the Total Environment 922 (2024) 171218
centuries for restored wetlands to become net radiative sinks. Although
the switchover time for peatlands is longer than for non-peat wetlands,
our analysis suggests that at the millennial time scale peatlands are more
effective at cooling the atmosphere than non-peat wetlands. While we
found that peatlands release, on average, more CH4, they also store more
carbon than non-peat wetlands. Hence, it takes longer for the net radi­
ative balance to become negative, but the net cooling effect is more
powerful over longer timescales.
The range of our estimated switchover times is greater than recently
reported estimates for restored wetlands (between 57 and 299 years)
(Taillardat et al., 2020). These previous estimates, however, ignored
N2O emissions and only accounted for CO2 sequestration and CH4
emissions. Nevertheless, our calculations omitted wetlands where CO2
emissions exceeded sequestration as we could not model the change in
NEE with increasing wetland age due to the limited availability of long-
term studies (>2 years). Restored wetlands that initially are net CO2
emitters may have longer switchover times than our estimates,
depending on the time it takes for wetland vegetation to establish and
turn these restored wetlands into net CO2-sequestering systems. Simi­
larly, our estimates do not consider the hydrological loss of carbon that
can contribute to off-site greenhouse gas emissions, which, in turn, can
L. Schuster et al. Science of the Total Environment 922 (2024) 171218

Fig. 5. (a) Instantaneous radiative effects of CH4, N2O, and CO2 following a pulse addition based on the average estimated fluxes for each gas from restored wetlands
and the decay of CH4 and N2O over a 1500-year period. The CH4 curve includes any radiative effect by CO2 that was produced from the oxidation of atmospheric CH4.
fW refers to femtowatts (10− 15 W). (b) Relationship between the average cumulative radiative forcing of restored freshwater wetlands based on their radiative
balance (kgCO2:(kgCH4 + kgN2O); solid lines) and wetland age (time since restoration in years). The dashed lines indicate the wetland ages at which restored
peatlands and non-peat wetlands switch from having a net warming effect to a net cooling effect on the climate (switchover time). pW refers to picowatts (10− 12 W).

also affect the switchover time of a restored wetland. Contrary to
restored wetlands, most natural freshwater wetlands already have a
strong net radiative cooling effect on the climate due to their greater age,
which likely exceeds their estimated switchover time of 298–2184 years
(Taillardat et al., 2020). These findings highlight that restored fresh­
water wetlands only have a long-term net cooling effect on the climate.
Protecting natural wetlands should therefore be a priority as those
ecosystems already play a key role in carbon drawdown and climate
change mitigation.
We found that restored and natural peatlands had higher CO2 uptake
rates and higher CH4 emissions than degraded peatlands, whereas in
non-peat wetlands, CO2 uptake and CH4 emissions did not differ be­
tween degraded, restored, and natural wetlands. The proportion of
peatlands taking up more CO2 than they emitted (NEE < 0) was twice as
high for restored peatlands compared to degraded ones, whereas all
natural peatlands were net CO2-sequestering systems. Here, a higher
CO2 uptake rate indicates a higher gross primary productivity (GPP,
given that GPP = Re − NEE). A higher primary productivity along with
prevailing anoxic conditions, in turn, can drive higher CH4 emissions as
plants are the main carbon source for methanogenesis (Bastviken et al.,
2023; Updegraff et al., 2001; Vann and Megonigal, 2003; Whiting and
Chanton, 2001). When vascular plants are abundant, however, CH4
emissions can be hampered in several ways through the promotion of
CH4 oxidation. For example, oxygen can be transported through the
roots into the sediment or soil, floating vegetation can trap CH4 bubbles
before they reach the water surface, and high oxygen availability within
plant stems can reduce plant-mediated CH4 emissions (Bastviken et al.,
2023; Dušek et al., 2023). These processes may drive the comparatively
low CH4 emissions from restored and natural non-peat wetlands
compared to Sphagnum moss-dominated peatlands where vascular
plants are largely absent, although anaerobic CH4 oxidation within the
soil may still play an important role in regulating CH4 emissions from
such peatlands (Bastviken et al., 2023; Segarra et al., 2015).
Not all restored wetlands were net carbon sinks, but we found that
the NEE of restored peatlands declines with wetland age after restora­
tion. This finding suggests that restored peatlands that initially are net

8
L. Schuster et al.
CO2 emitters (NEE > 0) should become net CO2-sequestering systems
like natural peatlands as wetland vegetation develops over time (Wad­
dington et al., 2010; Wilson et al., 2022). Freshwater wetland restora­
tion thus effectively turns degraded peatlands from net carbon sources
into carbon sinks over time, with older restored wetlands being more
important for climate change mitigation. If and how long it takes for
non-peat wetlands to turn from net carbon sources into carbon sinks,
however, remains unclear due to a limited number of studies reporting
the NEE of such wetlands.
Freshwater wetlands can be important carbon sinks that sequester
CO2 from the atmosphere and lock it away in their soils for centuries to
millennia due to anoxic soil conditions slowing microbial decomposition
(Mitsch et al., 2013; Richardson and Vepraskas, 2001). Contrastingly,
anoxic soils are typically absent in degraded wetlands. The carbon
captured through photosynthesis and stored in plant tissues is mostly
mineralised and re-emitted as CO2 and, to a lesser extent, CH4 (Xu et al.,
2019). Degraded wetlands may, however, take up more CO2 than they
emit (NEE < 0) due to periodic rainfall events or temporarily high
groundwater levels creating water-saturated soils (Fortuniak et al.,
2021; Robroek et al., 2009). Similarly, in degraded peatlands, deeper
soil layers may remain saturated, thereby maintaining carbon storage
and carbon sink capacity for up to several years after degradation
(Minkkinen et al., 2018). We found that most degraded wetlands were
net emitters of CO2, yet 31 % of degraded peatlands and all degraded
non-peat wetlands took up more CO2 than they emitted (NEE < 0).
Similar to restored and natural wetlands, these degraded wetlands can
have a net radiative cooling effect on the climate at least in the short
term. Nevertheless, natural and restored freshwater wetlands are also
key nature-based solutions that can help mitigate the impacts of climate
change. For example, they can mitigate floods and increase drought
resilience, and they are biodiversity hotspots for native and endangered
species (Chausson et al., 2020; Thorslund et al., 2017).
Given that freshwater wetland soils are saturated with water, we
expected to find higher N2O emissions from restored and natural wet­
lands. Specifically, anoxic conditions provide a suitable habitat for
denitrification in addition to aerobic nitrification, which can result in
increased N2O emissions (Van Cleemput et al., 2007). Instead, we found
that restored and natural wetlands tended to have overall lower N2O
emissions than degraded ones. Denitrification strongly depends on the
abundance of nitrate (NO−3 ) as an electron acceptor (Neubauer and
Verhoeven, 2019). In freshwater wetlands, plants compete with soil
microorganisms for nutrients including NO−3 (Hodge et al., 2000;
Kuzyakov and Xu, 2013). These strong interactions can lead to lower
levels of denitrification within wetland soils and hence lower N2O
emissions (Brummell et al., 2017; He et al., 2016). Since restored and
natural wetlands had relatively high primary productivities, increased
competition for NO−3 between plants and denitrifiers may have resulted
in the comparatively low N2O emissions from these ecosystems.
5. Limitations and future direction
Our meta-analysis provides a valuable first global assessment of the
effects of ecological restoration on greenhouse gas emissions and the
time required for restored wetlands to have a net cooling effect on the
climate when considering the exchange of the three most important
greenhouse gases (CO2, CH4, and N2O). However, the main limitation of
this work is the lack of continuous long-term studies (>2 years) quan­
tifying the effects of freshwater wetland restoration on greenhouse gas
fluxes and carbon sequestration. Additionally, we found that most
studies (87 %) were based on manual flux chamber measurements rather
than more accurate, continuous in situ measurement techniques such as
eddy covariance. Going forward, the continuous, long-term monitoring
of greenhouse gas balances of wetlands should be prioritised.
Another limitation is that most studies were conducted in the
northern hemisphere, particularly in Europe, the United States and
Canada (c.f. Fig. 1). We were therefore unable to test for differences in
9
Science of the Total Environment 922 (2024) 171218
restoration effects among different climates and across latitudes. Given
that primary productivity and methane emissions strongly depend on
environmental variables such as rainfall and temperature (Brinson et al.,
1981; Wang et al., 1996), restored wetlands in wetter and warmer cli­
matic regions may have overall higher CO2 uptake rates due to higher
primary productivities (yet possibly also higher CH4 emissions) and
potentially switch to having a net radiative cooling effect on the climate
sooner. The effectiveness of freshwater wetland restoration for climate
change mitigation purposes may therefore vary across latitudes. To test
this hypothesis, however, more studies in warmer regions and in the
southern hemisphere are needed.
We found a limited number of studies investigating the effects of
restoration particularly in freshwater marshes (c.f. Fig. S3) and using
fencing (c.f. Fig. S4). We were, therefore, unable to test for any differ­
ences in the switchover times across wetlands and restoration ap­
proaches. To better inform policy makers about the types of wetlands
that should be prioritised and the most effective restoration practices to
maximise climate benefits, more studies on the effects of different
restoration techniques in different types of wetlands are needed.
The relationship between NEE and the age of restored wetlands is
likely non-linear as primary productivity within any given ecosystem is
predicted to vary over time as vegetation develops and changes (e.g.,
Miller and Fujii, 2010). Due to the lack of long-term studies (>2 years
post restoration), however, we were unable to test for any non-linear
relationships. To capture this non-linear change in NEE over time,
more long-term studies that go beyond the immediate effects of resto­
ration are thus needed. Importantly, this change in NEE, along with
potential long-term changes in CH4 emissions can be incorporated into
switchover time calculations, making predictions about when a restored
wetland will have a net radiative cooling effect on the climate more
accurate.
Lastly, we estimated the net ecosystem carbon budgets (NECB) of
degraded, restored, and natural freshwater wetlands based on reported
net CO2 and CH4 fluxes. Ideally, NECB estimates would also consider any
aquatic carbon export from freshwater wetlands, including any lateral
transport of organic and inorganic carbon (Webb et al., 2019). However,
none of the studies that we included in our meta-analysis measured the
aquatic carbon export from freshwater wetlands likely due to the
extensive work that comes with estimating such fluxes or due to the
difficulties of estimating the effects of restoration on lateral carbon
fluxes when the restored wetland is part of a larger catchment area
(Evans et al., 2016).
6. Conclusions
Our meta-analysis showed that restoration is an effective way to turn
degraded wetlands from net carbon sources into carbon sinks, especially
in peatlands in the northern hemisphere, which represented 76 % of the
available data. Yet the average time for restored wetlands to have a net
cooling effect on the climate is 525 years for peatlands and 141 years for
non-peat wetlands. These results suggest that the radiative climate
benefit of wetland restoration does not meet the timeframe set by the
Paris Agreement to stabilise the temperature at 1.5 ◦ C above pre-
industrial levels by the year 2100. Wetland restoration is nonetheless
an effective way to reduce carbon emissions from degraded wetlands,
yet a long-term solution (~200–500 years) to mitigate climate change.
We therefore suggest that priority should be given to the conservation
and protection of natural freshwater wetlands as those ecosystems
already play a key role for carbon drawdown and climate change
mitigation.
CRediT authorship contribution statement
Lukas Schuster: Conceptualization, Data curation, Formal analysis,
Investigation, Methodology, Project administration, Validation, Visual­
ization, Writing – original draft, Writing – review & editing. Pierre
L. Schuster et al.
Taillardat: Formal analysis, Writing – review & editing. Peter I.
Macreadie: Writing – review & editing. Martino E. Malerba: Concep­
tualization, Supervision, Writing – review & editing.
Declaration of competing interest
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.
Data availability
All data included in our meta-analysis is available on Dryad
(https://doi.org/10.5061/dryad.vq83bk40p).
Acknowledgements
M.E.M. is supported by the Australian Government through the
Australian Research Council (project ID DE220100752).
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.scitotenv.2024.171218.
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