Paludiculture can support biodiversity conservation in rewetted fen peatlands

Abstract

Paludiculture, the productive use of wet or rewetted peatlands, offers an option for continued land use by farmers after rewetting formerly drained peatlands, while reducing the greenhouse gas emissions from peat soils. Biodiversity conservation may benefit, but research on how biodiversity responds to paludiculture is scarce. We conducted a multi-taxon study investigating vegetation, breeding bird and arthropod diversity at six rewetted fen sites dominated by Carex or Typha species. Sites were either unharvested, low- or high-intensity managed, and were located in Mecklenburg-Vorpommern in northeastern Germany. Biodiversity was estimated across the range of Hill numbers using the iNEXT package, and species were checked for Red List status. Here we show that paludiculture sites can provide biodiversity value even while not reflecting historic fen conditions; managed sites had high plant diversity, as well as Red Listed arthropods and breeding birds. Our study demonstrates that paludiculture has the potential to provide valuable habitat for species even while productive management of the land continues.

Full text

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Paludiculture can support
biodiversity conservation
in rewetted fen peatlands
H. R. Martens
1, K. Laage
2, M. Eickmanns
2, A. Drexler
2, V. Heinsohn
2, N. Wegner
2, C. Muster
2,
M. Diekmann
1, E. Seeber
2, J. Kreyling
2, P. Michalik
2 & F. Tanneberger
2*
Paludiculture, the productive use of wet or rewetted peatlands, oers an option for continued land use
by farmers after rewetting formerly drained peatlands, while reducing the greenhouse gas emissions
from peat soils. Biodiversity conservation may benet, but research on how biodiversity responds
to paludiculture is scarce. We conducted a multi-taxon study investigating vegetation, breeding bird
and arthropod diversity at six rewetted fen sites dominated by Carex or Typha species. Sites were
either unharvested, low- or high-intensity managed, and were located in Mecklenburg-Vorpommern
in northeastern Germany. Biodiversity was estimated across the range of Hill numbers using the
iNEXT package, and species were checked for Red List status. Here we show that paludiculture sites
can provide biodiversity value even while not reecting historic fen conditions; managed sites had
high plant diversity, as well as Red Listed arthropods and breeding birds. Our study demonstrates
that paludiculture has the potential to provide valuable habitat for species even while productive
management of the land continues.
Peatlands contain massive stocks of carbon, storing over twice the amount of carbon in the biomass of all the
world’s forests, despite covering only 3% of the Earth’s land surface1. However, these ecosystems have historically
faced, and continue to face, enormous pressure and widespread degradation2,3. Once drained, peatlands emit
substantial amounts of greenhouse gases (GHGs) through peat mineralisation and are currently responsible
for approximately 5% of all anthropogenic GHG emissions4. Within Germany specically, more than 95% of
peatlands are degraded from drainage, with the majority being used for crops (21%) or meadows/pasture (60%),
and this degradation contributes to 7% of Germany’s total GHG emissions5,6. Substantial further emissions from
drained peatlands could be prevented by rewetting7.
While the need for rewetting is urgent, it is not possible to simply return all degraded peatlands into protected
wilderness areas, as rural livelihoods are dependent on continued production from these areas8. Paludiculture—
the productive use of wet or rewetted peatlands9—has been developed as a method for enabling rewetting while
allowing farmers to continue working their land, though with an alternative land use. Paludiculture can take
many forms, and in northeastern Germany can include harvesting common reed (Phragmites australis), sedges
(Carex spp.), cattail (Typha spp.), or alder (Alnus glutinosa), and pasture with water bualo (Bubalus bubalis)8.
e biomass from these sites can be used for feedstock or biofuel8. Unlike conventional agriculture on drained
peatland, paludiculture prioritizes preservation of the peat body9 and can contribute to the Paris Agreement
targets (warming below 2°C) through reduced GHG emissions8,10,11. To preserve the peat body and allow for
carbon sequestration, specialized mowing equipment adapted to wet conditions is used, and water levels are
kept at or above ground level year-round8. Deeply drained peatlands are especially good candidates for paludi-
culture, as they are unlikely to return to a historic state even aer restoration9,12. Continued production on this
land is an equitable approach, enabling farmers to remain on the land, and local communities to steward their
own peatland resources1,13.
Peatland degradation has resulted in substantial loss of biodiversity14. Fens in particular have lost biodiver-
sity due to a reduction of traditional management, both from abandonment and intensication of agriculture
through drainage and eutrophication15,16. Peatlands with a history of agricultural use have become adapted to
regular disturbance, leading to declines in biodiversity when management is abandoned15. Biodiversity loss may
occur from eutrophication in drained and rewetted peatlands due to past agricultural use and the mineraliza-
tion of peat13. In these cases, mowing of fens may be essential for reducing eutrophication and maintaining
biodiversity17,18. Without mowing or other forms of management, rewetted fens may be dominated by a few tall
OPEN
1University of Bremen, Bremen, Germany. 2University of Greifswald, Greifswald, Germany. *email: tanne@
uni-greifswald.de
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and competitive species, resulting in a loss of low growing plants, rare species, and those with a low competitive
ability14,19–21. Paludiculture sites are likely to have greater fen biodiversity and more wetland species compared
to their drained state22. Even agricultural or open landscape species may benet from peatland rewetting and
management due to the subsequent opening of vegetation structure16,22.
ere is a need to understand how biodiversity responds to paludiculture and how to maximize outcomes
for biodiversity conservation. Rewetted peatlands have been found to create novel ecosystems that dier in their
plant and spider biodiversity compared to historical peatlands12,23. Especially lacking is an understanding of the
response of biodiversity to dierent intensity levels of paludiculture22. In this study, we assessed the biodiversity
of plants, breeding birds, carabid beetles and spiders using both quantitative and qualitative methods. Six sites
located in northeastern Germany were studied in 2021 and 2022. ese sites varied in their dominant vegetation
type, either Carex or Typha species, and in their land use intensity, either unmown, mown occasionally, or mown
annually. Biodiversity was compared across sites to assess quantitative diversity and Red List status was used to
assess qualitative diversity values. We demonstrated that paludiculture sites can host high vegetation diversity
and critically endangered breeding birds, as well as spiders and carabids of conservation concern. Each taxon is
expected to respond dierently to management, indicating the need for a multi-taxon perspective to understand
the impact of paludiculture on the biodiversity of rewetted peatlands.
Results
A total of 78 plant, 18 breeding bird, 55 carabid, and 73 spider species were identied. A total of 32 Red Listed
species (3 plants, 7 birds, 12 carabids, and 10 spiders) were present; all but three of these (spiders) occurred in
managed peatlands. Most Red List species present were those associated with wetlands (28), or open landscapes
(3 breeding birds). Carex sites generally had higher mean vegetation coverage than Typha sites; sites ranged from
80-100% mean coverage to 60–80%, respectively. Trees and shrubs were almost never present, and bryophytes
were only occasionally encountered. Litter cover was generally high (> 85%) except for the high intensity Typha
site which had minimal litter. A full species list is available as a supplementary le.
Quantitative analysis
e iNEXT package, developed by Chao etal.24, was selected for the quantitative analysis because it both quan-
ties sample completeness and provides diversity estimates across the range of Hill numbers. Sample coverage
values, which are a measure of sample completeness, were generally close enough to 1.0 (or 100% complete) to
enable interpretation of iNEXT results, except for breeding birds. e newly developed high intensity Typha
cropping site had signicantly higher predicted plant diversity across the range of Hill numbers, while the low
intensity Typha site had signicantly lower diversity. e managed Carex sites had signicantly more plant
species than the unmanaged site (Fig.1). Results for breeding birds generally showed insucient sample cover-
age for interpretation (coverage maximum 0.75). e high intensity Typha site had signicantly fewer carabid
species: the site had one third of the estimated species richness of any other site. e spiders in the unharvested
Carex site had around 60% higher Shannon and Simpsons diversity than other Carex sites, and higher species
richness in the unharvested Typha site. All other sites were similar in their quantity of spider species. Vegetation
and spiders responded oppositely to management; plant diversity generally increased in mown sites, but spider
diversity decreased.
Qualitative analysis
Across all sites, most of the species identied were typical for wetlands (74%). Sites did not reect a historic
mire state since they had few mire-specic species. Species of conservation concern were found from all taxa;
the species of greatest concern and mire-specic species have been listed (Table1)28–31. Additionally, thirteen
threatened species and eight near threatened species were present at managed sites (complete list of Red List
species available as a supplementary le).
Discussion
Quantitative analysis showed no consistent diversity response to the intensity of use of rewetted fen peatlands,
regardless of dominant vegetation type. Qualitative results demonstrated that all sites, and, consequently, all
land use intensity levels, were providing habitat for Red List wetland species. Given that intensive grassland on
drained peatlands does not provide habitat for fen communities32, our ndings underline that paludiculture
can support fen biodiversity and conservation better than a drained state. Additionally, management supported
higher vegetation diversity then an unharvested wet state. However, birds, arthropods, and plants all varied in
their biodiversity between sites and management intensity, thus supporting the need for variation of land use
intensity in the landscape, as also suggested by other studies33.
Quantitative analysis
Managed Carex sites all had similarly high vegetation diversity values. In contrast, the unharvested Carex site had
signicantly lower diversity and had highly uniform and tall vegetation. Tall vegetation can restrict the growth
of light-dependent species in fens34,35. is study, like others, found that mown sites have the capacity to host
higher plant species richness than unmown sites34,36–41. Despite its isolated location and recent rewetting, the
high-intensity Typha site had signicantly higher diversity then other sites. However, given the site was recently
established (2019), species diversity may change over time. Typha-low had the lowest diversity values, which
may be attributed in part due to the high proportion of ruderal plant species (Urtica dioica, Cirsium arvense)
compared to other sites.
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Figure1. Coverage based biodiversity extrapolations for dierent taxa comparing paludiculture intensities for
Carex and Typha as target species. Estimate of sample completeness is given as sample coverage which is used
to standardize samples according to the iNEXT.4 package. Diversity results are extrapolated and interpolated
across the range of Hill numbers24. us, diversity at each site is compared using species richness, which is
biased towards rare species, Shannon diversity, biased towards common species, and Simpson’s diversity, biased
towards dominant species. Sites are compared at equal sample coverage, given as the coverage maximum
(double the smallest sample size), where a sample coverage of 1.0 for Simpson’s diversity indicates 100% of
dominant species are predicted to have been found24. Here, vegetation is compared at a maximum coverage of
0.95, and carabids and spiders both at 0.99. Bird results are not provided due to insucient coverage (coverage
maximum of 0.75). Shown are 83.4% condence intervals, whose non-overlap indicates a signicant dierence
at a lpha = 0.0525–27.
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e high intensity Typha site had signicantly lower carabid diversity than all other sites. A contributing
factor may be the low willingness of carabid specialist species to cross unfavorable terrain, reducing the chance
to disperse to new areas42. is site was rewetted only two years before our observations and is a hydrologically
isolated fen in a landscape dominated by drained peatlands used as pasture. e other sites that were studied had
been rewetted around twenty years prior (Table1). A study of a Sphagnum paludiculture site found that during
the rst three years aer rewetting, spider community structure changed considerably, but aer three years the
overall community structure remained stable43. To better support carabid species, connectivity to other peatlands
should be restored42, and it may take time for stable populations to form. Species re-introduction may be help-
ful and has been used for example in the partially successful reintroduction of the fen ra spider (Dolomedes
plantarius) in the UK44. However, the presence of rare and threatened species in the study sites indicates that
species assemblages are establishing in a positive trajectory. Results from the high intensity site vary between
all groups and show both signicantly more plant and less carabid beetle diversity than all other sites; diverging
diversity values between carabids and plants were also found by Görn & Fischer45 emphasizing the importance
of multi-taxon studies.
Spider diversity results were unique compared to other taxa, as the unharvested Carex site had signicantly
higher Shannon and Simpson’s diversity than all other sites. Plants and carabids had moderate to very low diver-
sity values for this site. Studies on spiders in fens have found that mowing reduces litter and vertical vegetation,
and thus may reduce structure-dependent species like rare wetland spiders and some widespread species46,47.
Research on other invertebrate groups also found lowest species richness in recently mown reedbeds33. ese
factors may be contributing to high diversity values in the site without management. Higher diversity of spider
and bird species than carabids at the high intensity Typha site may relate to mobility, since some spiders have
“ballooning” capabilities and thus higher dispersal ability43.
Qualitative analysis
All sites had a high proportion of mire-typical and general wetland species which aligns with work by Tan-
neberger etal.22, who found that paludiculture sites host primarily species adapted to wet environments. However,
sites lacked indicators of a natural mire, since very few mire-specic species were identied. Rewetted peat-
lands have been found to dier in their plant diversity, hydrology, and geochemistry compared to near-natural
peatlands12. ese rewetted landscapes typically have tall graminoid plants, are eutrophic, and have a higher water
table12. Despite its recent rewetting and isolated location, the high intensity Typha site hosts Red List species
from all studied taxa. For example, northern lapwing populations have declined dramatically in the last thirty
years as their habitat has decreased from both intensication and abandonment of land use and may benet
from low or moderate management intensity16,48–50. Moreover, multiple bird species associated with landscapes
other than wetlands, including agricultural (Emberiza calandra) or open landscapes (Saxicola rubicola, Saxicola
rubetra), were breeding in the paludiculture sites indicating that such sites can indeed host at-risk species. is
is in accordance with other paludiculture projects43. While in restored fens it may be preferable to have a high
number of mire-specic species, this may not be the case for paludiculture sites. For example, if paludiculture
sites can provide habitat for endangered agricultural and open landscape species whose habitat is disappearing,
this may also be considered a positive eect of such land use.
Further research over multiple years and on many more sites is needed to understand the conservation and
biodiversity value of paludiculture as sites change. For example, a study by Valkama etal.38 showed that aer
several years, mowing signicantly decreased invertebrate abundance, but in the short-term (1–2 years) the sites
appeared unaected34. A study by Muster etal. on a Sphagnum paludiculture site noted that each successional
Table 1. List of species of conservation concernrecorded at the study sites. Mire-specic species, IUCN Red
List species, and the top two categories of the German Red List have been included. Taxa are indicated by the
symbol: plants ●, carabids █, spiders ▼, birds ♦.
Conservation status Mire-specic International red list German red list: threatened with
extinction German red list: highly
threatened
Carex-unharvested
▼ Carorita limnaea
▼ Diplocephalus dentatus
▼ Pirata piscatorius
▼ Carorita limnaea
▼ Centromerus semiater
▼ Diplocephalus dentatus
♦ Locustella naevia
Carex-low intensity
● Triglochin palustris
▼ Carorita limnea
▼Pirata piscatorius
▼ Dolomedes plantarius ♦ Gallinago gallinago █ Elaphrus uliginosus
▼ Carorita limnaea
▼ Dolomedes plantarius
Carex-high intensity ▼ Pirata piscatorius ♦ Gallinago gallinago
Typha-unharvested ▼ Pirata piscatorius
▼ Diplocephalus dentatus
♦ Locustella naevia
♦ Saxicola rubetra
Typha-low intensity ▼ Pirata piscatorius ♦ Vanellus vanellus
█ Elaphrus uliginosus
♦ Anthus pratensis
♦ Locustella naevia
♦ Saxicola rubetra
Typha-highintensity ● Juncus subnodulus
▼ Pirata piscatorius ♦ Vanellus vanellus ♦ Saxicola rubetra
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stage had dierent species, and even at early stages sites had high conservation value species, but not mire-typical
species43. In our study, all but the high intensity Typha site reect a long-term state, since rewetting occurred in
the early 2000s (Table2). Future work on paludiculture biodiversity should study multiple animal groups, as each
may respond dierently to management, and additionally, more multi-year studies are important to understand
succession, annual uctuations, and dispersal in newly established sites or according to mowing regime. Long
term monitoring of such paludiculture sites would provide more information on typical species and conservation
value at each successional stage, especially on sites that are not mown annually (low-intensity management),
where species composition may vary temporally. Many factors inuence the impact of mowing on biodiversity,
including the block size in when creating a mosaic of mowing regimes47, mowing technique and machinery51,
and time of year49. More sites and thus spatial replication are needed for a robust understanding of how these
factors inuence diversity at paludiculture sites.
Methods
Site selection
e study sites are in the state of Mecklenburg-Vorpommern in northeast Germany (Fig.1, Table1). Site bounda-
ries were delineated by barriers (roads, open water bodies, ditches) or by transition to a new mowing regime or
vegetation type. Sites were selected for their vegetation type, either Carex or Typha, and had dominant species
of either Typha latifolia, or Carex acuta, C. acutiformis, and C. disticha. All sites have a history of deep drainage
and subsequent rewetting in the early 2000s as permanent grassland paludiculture52, except for the high intensity
Typha site, which was rewetted in 2019 and developed as a cropping paludiculture site with planted Typha. e
study locations varied in their connectivity with surrounding natural fen habitat; the Carex sites are all three
similarly close to peatlands that were only slightly drained (north of Neukalen and on the eastern side of Lake
Kummerower) (Fig.2), Typha-unharvested and Typha-low were surrounded partly by agriculture and partly by
other rewetted peatlands, and the Typha-high site was isolated, surrounded by drained peatland used as grassland
and the Teterower Peene river, and rewetted in 2019 (Table2). High intensity sites were harvested completely
every year, and low intensity sites were mown every two to three years, in some years only mulched (without
biomass removal). e sites are in a temperate climate and experience a mean temperature of 9.5°C, with around
735mm of annual precipitation, with most of this falling in the summer months52. Site selection was limited since
few paludiculture areas have been established thus far and more replicates were not readily available, especially
for managed sites. Additionally, further sampling would have demanded too many resources and would have
been beyond the scope of the current study. erefore, our study had replicates within each site, but did not have
true replicates for management intensity. However, geostatistical analysis of fen peatlands has demonstrated that
spatial autocorrelation is rarely present53,54. is suggests that the spatial replicates within each of our six sites
can be treated as independent and their variation is representative for their respective vegetation type.
Data collection
Vegetation data was collected in 2022, and breeding bird, carabid, and spider data in 2021. Water level classica-
tion is based on water level measurements taken at a representative permanent monitoring well located at each
site measured from April 2021 until February 2022. Water levels are classied based on Couwenberg55, adapted
from Koska56.
Vegetation was surveyed in late June and early July of 2022. Plots were placed using stratied random sam-
pling and number of plots varied due to dierences in the size of each site (Table2) (Carex-unharvested: 6,
Carex-low and high: 10, Typha-unharvested: 20, Typha-low: 18, Typha-high:22).Two by two-metre plots were
placed at regular intervals along a transect running through the site center. Additional plots were placed at
random if multiple vegetation zones were present. Edges, open water, and areas heavily trampled by mowing
near site entrances were avoided, resulting in a small reduction in sampling area. Cover values of each species
were estimated as percent coverage at < 1% coverage and intervals of 10%. ese values were then converted into
Table 2. Site descriptions, use, and history. Water level class calculated from summer 2021 and winter 2021/22
median water level based on water level classication from Couwenberg55, adapted from Koska56. Water level
of 4+ may preserve peat (depending), while levels of 5+ and 6+ are peat preserving or even peat forming57.
Mean vegetation height was taken as an average across the entire site, all other values are from a single point at
the site in 2022. Amplitude gives the dierence between the minimum and maximum water level during the
recorded period. pH data was collected in 2021.
Name Mowing intensity Area (ha) Year drained Yea r R ew et te d Water level class pH Mean vegetation
height (cm) Vegetation height
SD (cm) Water level
amplitude (cm)
Carex-unharvested None 1.0 1925 2002 5+ 9.3 92.5 8.2 62.7
Carex-low intensity Infrequent 3.5 1925 2002 5+ 8.7 80.0 8.6 57.2
Carex-high
intensity Annual 2.5 1925 2002 5+/6+ 9.5 75.0 25.2 57.1
Typha-unharvested None 16.5 1967 2005 4+/5+ 8.9 120.0 40.6 61.1
Typha-low intensity Infrequent 5.8 1940 2001 5+ 8.5 90.0 21.5 11.0
Typha-high
intensity Annual 9.0 1935 2019 6+ 8.8 82.5 55.8 20.0
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presence-absence data to t the format required by the iNEXT package. Species were identied using Streeter
etal.60 and names veried using Euro + Med PlantBase61.
Breeding birds were surveyed following the breeding bird territory survey method outlined by Südbeck etal.62.
Surveys were conducted over ve mornings starting 30min before sunrise and two evenings starting 30min
aer sunset. All birds singing, calling, and all those engaged in behavior indicating breeding within the site were
recorded using QField and mapped using QGIS. Breeding pairs were determined based on their behaviour and
the time of year62. Surveys were conducted at the end of March, end of April, middle of May (one evening, one
morning), beginning of June (evening survey), middle and end of June. Sites were surveyed over three days each
time, always with a minimum of seven days between each survey round. e order of sites surveyed, and the
route taken while surveying was altered each time.
Carabid beetles and spiders were collected using pitfall traps (six per site) and additional oating traps
were placed at the three Typha sites to collect arthropods due to high water level. Pitfall traps were made from
a standardized colorless transparent reusable plastic cup63. Cups were held in place using tent pegs. Floating
traps were constructed using a cup surrounded by a Styrofoam ring and were weighted to keep the cup rim at
surface level64. ese were set within a polypropylene pipe, diameter of 15cm and length of 100cm to hold
traps in place. Each pipe had several 5cm diameter holes to allow arthropods to enter and was plugged on
the upper end to prevent rainwater and debris from entering. Sampling cups had a diameter of 8cm, depth of
Figure2. Map of sites in Macklenburg-Vorpommern, Germany58,59. Sites are labelled by their dominant
vegetation type, Carex (C), or Typha (T), and the land use, including unharvested (UH), low intensity (LI), and
high intensity (HI). e majority of sites were located near Neukalen but the high intensity Typha site (T-LI) was
located approximately 70 km east near Anklam.
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10cm, and contained a solution of ethanol, water, glycerin, and acetic acid at a ratio of 4:3:2:1 and unscented
soap65. Locations of traps were recorded with GPS and marked with bamboo sticks and were spaced 10m apart
and at least 20m away from site boundaries. Five sampling periods occurred in spring (April–June) and three
in autumn (September and October) for a total of eight. Each sampling period lasted 14days. Identication for
carabids was done following Müller-Motzfeld66 and nomenclature using Schmidt etal.67 . Spider identication
and nomenclature followed Nentwig etal.68.
Data analysis
General analysis was done in R69 using RStudio70 and the package tidyverse71 and visualization done using
viridis72, ggrepel73, gt74, MetBrewer75, and ggplot276. Several methods of biodiversity analysis were utilized,
given that no one method has been found to be entirely eective or representative of site diversity. Quantitative
biodiversity analysis was made using iNEXT77,78, iNEXT.4steps79, and devtools80. e iNEXT package provides
diversity estimates across the range of Hill numbers and thus across the range of sensitivity to species abundance
and was used following Chao etal.24. e method is based on the work of Hill81 who found that species richness,
Simpson’s diversity and Shannon’s diversity can be placed on a continuum of diversity measures based of their
bias towards rare species. is continuum approach is more robust than using any of these diversity estimates
individually since each are biased and when used alone may provide contrasting results24,82,83. iNEXT method
enables comparison using sample completeness rather than sample size, allowing for comparison between dif-
fered sized sites without having to reduce to the smallest sample size for comparison24,84. e method for sample
completeness estimation is formulated on the codebreaking work of Allan Turing during WWII and estimates
the amount of information that is unknown to quantify what is known, given the frequency that something
appears exactly once or exactly twice84. e iNEXT.4steps package provides analysis in four steps, as suggested
by the name, but only two of these were utilized for this analysis. Sample coverage (step 1) and non-asymptotic
coverage-based rarefaction and extrapolation (step 3) were the focus, since they provide analysis of sites with
uneven sampling intensity. Step two (asymptotic and empirical diversity) has been le out, since samples were
insuciently complete to detect true diversity, and step four (evenness) was also omitted, since a lack of replicates
resulted in large and inconclusive condence intervals24. Samples were bootstrapped 50 times (the packages
default) to estimate 83.4% condence limits which were used to determine signicance of dierences between
the land use intensities. Condence intervals were set based on research that demonstrates non-overlap of 83.4%
condence limits correspond with approximately an alpha of 5%26,27.
Species were also evaluated qualitatively, both concerning their endangerment status and their typical habitat
preference using literature for northeast Germany. Mire-specic plant species were identied using Hammerich
etal.85 and mire-specic spider species using Martin86. Furthermore, area-specic literature was used to deter-
mine the typical habitat for each species (vegetation60,87, breeding birds88–90, carabids91, and spiders92,93). e goal
of this classication was to determine if paludiculture sites were attracting wetland species, or if the sites continue
to host mostly species associated with traditional agricultural land, generalists, or other habitat types. National
level Red List information was obtained from the German Red List Center for plants94, birds31,95, carabids96, and
spiders28. International information comes from the IUCN Red List website97.
Conclusion
e approaches taken in this study provide a multi-taxon view of biodiversity in the selected paludiculture sites
by using four dierent taxa and both a qualitative and quantitative approach for assessing biodiversity. All sites,
irrespective of management intensity, hosted species with high national and international conservation value,
indicating that not only protected “wilderness” sites but also paludiculture sites can provide refuge for endan-
gered species. However, these sites did not resemble natural fen conditions and had few mire-specic species but
did contain primarily wetland species. e site with greatest management inuence (Typha-high intensity) had
both the lowest and the highest qualitative biodiversity values depending on the taxon. us, further research is
needed to understand long-term biodiversity trends in these novel ecosystems, and many more sites should be
established and studied to create a more robust understanding of the factors shaping biodiversity in paludiculture
sites. Since responses varied between taxa, management should aim to provide a habitat mosaic with variation
in management intensity. Also from a biodiversity perspective, eorts towards rewetting and management of
degraded peatlands should continue, since it has been demonstrated that this land use supports high biodiversity
and species quality compared to a drained peatland.
Data availability
All data generated or analysed during this study are included in the supplementary information les of this
published article.
Received: 17 July 2023; Accepted: 9 October 2023
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Acknowledgements
We thank the landowners and land managers for allowing us access to their land for our data collection. is
research was funded through the 2019-2020 BiodivERsA joint call for research proposals, under the BiodivClim
ERA-Net COFUND programme, and with the funding organisation VDI-VDE.
Author contributions
H.R.M. wrote the main manuscript text and analyzed the data. E.S., J.K., P.M., F.T. designed and supervised the
experiment. H.R.M., K.L., M.E., A.D., V.H., N.W., and C.M. collected data and identied species. All authors
reviewed the manuscript.
Funding
Open Access funding enabled and organized by Projekt DEAL.
Competing interests
e authors have no competing interests as dened by Nature Research, or other interests that might be perceived
to inuence the results and/or discussion reported in this paper.
Additional information
Supplementary Information e online version contains supplementary material available at https:// doi. org/
10. 1038/ s41598- 023- 44481-0.
Correspondence and requests for materials should be addressed to F.T.
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