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Effect of reintroduced manual mowing on biodiversity in abandoned fen meadows

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Wetlands have recently become of high environmental interest. The restoration effects on habitats like fens are one of the main topics of recent restoration ecology, especially due to their interconnection with other ecosystems. We studied the manual mowing effect on abandoned fen using the response of three study taxa: diurnal butterflies, flower-visiting beetles and vascular plants. Our results showed that butterflies seems to be quickly-responding indicator taxon for evaluation and that restored management had a positive effect on both species richness and composition of this insect group. The results indicated that the manual mowing effect could be rapid. In comparison with the surrounding landscape, we found that: (i) the manually mowed site was most similar to strictly protected area, (ii) some species of high conservation value could reach higher abundance in restored than protected site, and (iii) manual mowing could bring a new type of habitat (i.e., spatial heterogeneity) compared to the other management types (abandonment, conservation and agri-environmental mowing). The main implication seems to be optimistic for practice: The manual mowing of long-term abandoned fen is leading to the creation of habitat with high conservation value in a relatively short time.
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Biologia 70/1: 113—120, 2015
Section Zoology
DOI: 10.1515/biolog-2015-0009
Effect of reintroduced manual mowing on biodiversity
in abandoned fen meadows
Jakub Horak1&LenkaSafarova2
1Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamycka 1176,CZ-16521 Prague 6,Czech
Republic; e-mail: jakub.sruby@gmail.com
2East Bohemian Museum in Pardubice, Zamek 2,CZ-53002 Pardubice, Czech Republic
Abstract: Wetlands have recently become of high environmental interest. The restoration effects on habitats like fens are
one of the main topics of recent restoration ecology, especially due to their interconnection with other ecosystems. We studied
the manual mowing effect on abandoned fen using the response of three study taxa: diurnal butterflies, flower-visiting beetles
and vascular plants. Our results showed that butterflies seems to be quickly-responding indicator taxon for evaluation and
that restored management had a positive effect on both species richness and composition of this insect group. The results
indicated that the manual mowing effect could be rapid. In comparison with the surrounding landscape, we found that: (i)
the manually mowed site was most similar to strictly protected area, (ii) some species of high conservation value could reach
higher abundance in restored than protected site, and (iii) manual mowing could bring a new type of habitat (i.e., spatial
heterogeneity) compared to the other management types (abandonment, conservation and agri-environmental mowing).
The main implication seems to be optimistic for practice: The manual mowing of long-term abandoned fen is leading to the
creation of habitat with high conservation value in a relatively short time.
Key words: agri-environmental schemes; diurnal butterflies; Lepidoptera; flower-visiting beetles; Coleoptera; nature con-
servation; multi-taxa approach; vascular plants
Introduction
Human influence in the landscape is still rising and
often leads to a call for the restoration of former
ecosystems and reintroduction of traditional manage-
ment techniques (Hobbs & Harris 2001). Furthermore,
Roberts et al. (2009) argued that our planet’s future
may depend on the maturation of the discipline known
as restoration ecology. Former predictions indicated
that ecosystems will take centuries to recover from hu-
man disturbances (Jones & Schmitz 2009). However,
the main goal is not necessarily to restore or mitigate
the impacts of human activities to an ecosystem to
a pristine, pre-human ideal (Roberts et al. 2009), es-
pecially in the light of recent knowledge that many
types of damaged ecosystems are able to go through
rapid recovery (Jones & Schmitz 2009). It may be still
questionable what types of ecosystems are more or less
restorable (Moreno-Mateos et al. 2012), what kind of
changes in communities of organisms are caused by rein-
troduction of traditional management techniques and
how effective restoration or similar activities could be
(Hobbs & Harris 2001; Jones & Schmitz 2009).
Restoration and traditional management tech-
niques appear to be more ecosystem friendly than
present prevailing farming methods. Agricultural in-
tensification and abandonment of less-productive and
less-accessible land have led to the declines of taxa as-
sociated with fens and similar areas (Prach 1993). Poli-
cies regarding these areas as threatened fauna and flora
habitats assume that a restored ecosystem may replace
losses in structure and function in relatively short time
frames (Zedler & Callaway 1999). Zedler (2000) argued
that the restoration of wetlands takes more than wa-
ter, e.g. because hydrological regimes of wetlands are
highly complex and the wetland restoration is mainly
driven by the mitigation of damage to them by agricul-
tural intensification. Nearly the same situation is with
more or less humid grasslands (Jones & Schmitz 2009).
The drainage of these sites has eliminated highly valued
ecosystem functions, leading to regulations that require
compensation (Zedler 2000).
Recent studies have dealt with the recovery of com-
munities of species that prefer humid or wet grasslands
(Zimmermann et al. 2005; Billeter et al. 2007; Hedberg
& Kotowski 2010; Valkó et al. 2012). Several meth-
ods and techniques were used in management amend-
ments, like tree or shrub removal, hay or species re-
introduction, sowing of ingenuous propagules or sod
cutting (Wynhoff 1998; Mouquet et al. 2005; Lepš et al.
2007; Klimkowska et al. 2010; Sundberg 2012). Prob-
ably the easiest and softest method of restoration of
formerly intensively-managed grasslands is diversifica-
tion of mowing regimes (Grill et al. 2008; Cizek et al.
2012), which can also be applied after the more radical
methods.
c
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114 J. Horak &L.Safarova
Hand mowing was the most traditional way of
producing hay and management of grasslands. Mow-
ing of meadows (using invention of scythes) is known
from Iron Age (1,000 B.C. –0; e.g., Horák et al. 2012).
Presently, this management type appears to be obsolete
way of producing hay. On the other hand, this manage-
ment type has been evaluated only rarely (Valkó et al.
2012) and it would be useful to know the differences
between hand- and machine-mowing effects, in order to
understand what had been lost due to modernisation
(Humbert et al. 2009).
Several taxa or individual species associated with
fens and grasslands appear to be decreasing their abun-
dances due to abandonment of traditional management
types (Konvicka et al. 2008). Butterflies, beetles and
plants belong to well-studied taxa and their responses
to disparate environmental factors often indicate the
actual condition of the studied environment (Benes et
al. 2002; Valkó et al. 2012). The arthropod-plant inter-
action in grassland habitats is very high. Larvae of but-
terflies and beetles mostly forage on the tissues of plant
species and the presence and abundance of adults well
reflects larval activity as in other insect taxa (Steffan-
Dewenter & Tscharntke 1997).
Our questions regarding the manual mowing effect
on wetland communities were: Are there differences in
effects of reintroduced manual mowing for butterflies,
beetles and plants? Is manual mowing in an abandoned
fen able to create a species rich habitat compared to
surrounding landscape? And, if yes, is only a species
rich habitat created or is another fragment of disparate
habitat mosaic established?
Material and methods
Study area and its history
Wetlands of Sruby (49.9877 N; 16.1946 E) are part of the
former large area between the riparian corridor of the me-
andered Loucna River and forested lower ridge, which is
a part of the eastern-Bohemian forests (Czech Republic)
between the cities of Chocen and Hradec Kralove (Fal-
tysová et al. 2002). Most parts of the Sruby rural envi-
ronment are comprised of former forests, which were de-
forested at the beginning of the 14th century, when the
first permanent settlement was established here (Sedlacek
1908). As most of the area was on low-productive hydromor-
phic (i.e., gley) soils with high moisture, the main source
for living was forestry and pasture combined with the es-
tablishment of several ponds in the beginning of the 16th
century (Láska 1936). Growing wealth and contemporary
trends in land use during the period after World War I
(1914–1918) (Václavík 1869) led to drainage of most of
the rural environment of Sruby (Láska 1936). Most parts
of the study area were then used as arable land, alter-
nating with temporary grasslands (D. Kurkova, pers. com-
mun.). Nature monument Vstavacova louka was established
in 1989 in response to agricultural intensification and the
use of heavy duty technologies (Faltysová et al. 2002).
Unmanaged drainage after the Velvet Revolution in 1989
caused the rising level of soil moisture of arable land and
the worsening of crop farming, which led to the establish-
ment of secondary fen vegetation. Southern part of the
study area was abandoned in 1992 (J. Matousek, pers. com-
mun.).
Recently the non-forested part of the study area con-
sisted of (i) agri-environmental machine-mowed wet mead-
ows (agri-environmental thereinafter), (ii) strictly protected
fen meadows (conservation thereinafter), and (iii) aban-
doned fen (abandonment thereinafter), (iv) which was
partly manually mowed and hay was carried away in 2010
and 2011 (manual mowing thereinafter).
Study taxa, design and methods
Vascular plants (thereinafter plants) were counted in per-
centage cover scale between the first (2010) and second man-
ual mowing in the beginning of June 2011 within 32 plots
of 1 ×1 m. Each pair of 16 plots was distributed approxi-
mately in two lines with the distance of 5 m going through
particular treatment (manual mowing and abandonment)
and at regular distances of 20 m to the next pair. The dis-
tance to the edges was >10 m. The distance between plots
in each pair was 20 m for inner pairs and 30 m for outer
pairs.
Diurnal butterflies and burnet moths (Lepidoptera,
thereinafter butterflies) and flower-visiting beetles (Coleo-
ptera, thereinafter beetles) were studied using a time-limited
(16 min) survey method. Each treatment consisted of eight
survey walks in the same location as plots for the sampling of
plants. The edges as transition zones were omitted approx-
imately the same as in plants. All butterflies and beetles
were recorded during nine visits from the beginning of May
to the middle of September 2011, reflecting their phenolog-
ical activity in the conditions of eastern Bohemia (Horak
et al. 2013). All visits were only during fine weather con-
ditions (sunny, no wind, temperature 20
C) and during
10–11 AM or 2–3 PM.
For the analysis of total species richness of each study
taxa and comparisons among treatments, we used sample-
based species rarefactions (Mao Tau function) with 95%
confidence intervals (Colwell 2006) and the Chao estima-
tor (Chao 1984). Analyses were computed in EstimateS 8.2
(Colwell 2006). The number of randomisations was set at
1,000.
We focused on predictors which were testable only
within a limited spatial scale, because the study was con-
ducted in a limited grassland landscape surrounded by
forests and crop fields. Un-replicated treatments were the
only option for our study (Oksanen 2001). We controlled
this problem using randomised techniques for species rich-
ness data (Gotelli & Colwell 2001) and a set of coordinates
(x, y, xy, x2,y
2) as spatial co-predictors for the multivariate
analysis of species composition (Horak 2013) in comparison
with traditional statistical methods as it is recommended
by Oksanen (2001). Namely, the comparison of species rich-
ness of study taxa in study treatments was computed using
paired t-test for dependent variables with normal distribu-
tion in Statistica 7.
Another aim was investigated using an analysis fo-
cused at identifying species composition and response to
each treatment, which was carried out using multivariate
statistical methods provided by CANOCO for Windows ver-
sion 4.5 (ter Braak and Šmilauer 2002). Redundancy anal-
ysis (RDA), a constrained linear ordination method, was
used to solve our task with 9,999 unrestricted permutations
under the full model. The resulting ordination diagram was
created in CanoDraw 4.14 (ter Braak & Šmilauer 2002). In
this case we used data from the time of the peak activity
of wetland butterflies (i.e. four summer visits). As a tradi-
tional statistical method for comparison, we used ANOVA
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Reintroduction of manual mowing 115
Table 1. Check-list of butterflies in Wetlands of Sruby.
Family Species No. of individuals
Hesperidae Carterocephalus palaemon (Pallas, 1771) 4
Hesperidae Thymelicus sylvestris (Poda, 1761) 16
Lycaenidae Lycaena dispar (Haworth, 1802) 10
Lycaenidae Lycaena phlaeas (L., 1761) 5
Lycaenidae Lycaena tityrus (Poda, 1761) 3
Lycaenidae Phengaris nausithous (Bergstr¨asser, 1779) 16
Lycaenidae Phengaris teleius (Bergstr¨asser, 1779) 8
Lycaenidae Polyommatus amandus (Schneider, 1792) 1
Lycaenidae Polyommatus icarus (Rottemburg, 1775) 64
Lycaenidae Thecla betulae (L., 1758) 1
Nymphalidae Aglais urti cae (L., 1758) 3
Nymphalidae Aphantopus hyperantus (L., 1758) 39
Nymphalidae Araschnia levana (L., 1758) 17
Nymphalidae Argynnis adippe (Denis & Schifferm¨uller, 1775) 1
Nymphalidae Argynnis aglaja (L., 1758) 2
Nymphalidae Argynnis paphia (L., 1758) 3
Nymphalidae Boloria dia (L., 1767) 1
Nymphalidae Coenonympha pamphilus (L., 1758) 115
Nymphalidae Inachis io (L., 1758) 14
Nymphalidae Issoria lathonia (L., 1758) 1
Nymphalidae Maniola jurtina (L., 1758) 115
Nymphalidae Melanargia galathea (L., 1758) 18
Nymphalidae Pararge aeger ia (L., 1758) 2
Nymphalidae Vanessa atalanta (L., 1758) 3
Nymphalidae Vanessa cardui (L., 1758) 1
Pieridae Colias hyale (L., 1758) 6
Pieridae Gonepteryx rhamni (L., 1758) 53
Pieridae Leptidea reali Reissinger, 1989 8
Pieridae Pieris brassicae (L., 1758) 34
Pieridae Pieris napi (L., 1758) 157
Pieridae Pieris rapae (L., 1758) 21
Zygaenidae Zygaena filipendulae (L., 1758) 36
Zygaenidae Zygaena loti (Denis & Schifferm¨uller, 1775) 1
Zygaenidae Zygaena viciae (Denis & Schifferm¨uller, 1775) 2
Table 2. Check-list of beetles in Wetlands of Sruby.
Family Species No. of individuals
Buprestidae Anthaxia nitidula (L., 1758) 1
Cantharidae Rhagonycha fulva (Scopoli, 1763) 167
Cerambycidae Pseudovadonia livida (F., 1776) 2
Cetoniidae Cetonia aurata (L., 1758) 13
Cetoniidae Oxythyrea funesta (Poda, 1761) 37
Cetoniidae Protaetia marmorata (F., 1792) 1
Cetoniidae Valgus hemipterus (L., 1758) 1
Chrysomelidae Clytra quadripunctata (L., 1758) 1
Coccinelidae Coccinella septempunctata L., 1758 31
Coccinelidae Harmonia axyridis (Pallas, 1773) 7
Curculionidae Larinus turbinatus Gyllenhal, 1835 3
Melyridae Malachius bipustulatus (L., 1758) 1
Oedemeridae Oedemera femorata (Scopoli, 1763) 5
Oedemeridae Oedemera virescens (L., 1767) 1
for dependent variables with normal distribution in Statis-
tica 7.
Results
In total, we recorded 34 species of butterflies (Table 1),
14 beetle species (Table 2) and 90 species of plants (Ta-
ble 3).
Species rarefactions
With respect to differences between manual mowing
and abandonment we observed 29 species of butterflies,
14 beetle species and 90 species of plants.
Rarefactions were made separately for each taxo-
nomic group (Fig. 1). They did not reach their asymp-
totes. However, the Chao estimator approached the to-
tal number of species in the case of butterflies and
plants (Figs 1A, B). This suggested that most of the
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116 J. Horak &L.Safarova
Table 3. Check-list of plants in Wetlands of Sruby.
Family Species No. of samples
Apiaceae Aegopodium podagraria 6
Apiaceae Angelica sylvestris 12
Apiaceae Chaerophyllum bulbosum 1
Apiaceae Chaerophyllum temulum 2
Apiaceae Heracleum sphondylium 1
Apiaceae Pastinaca sativa 6
Apiaceae Peucedanum palustre 12
Asteraceae Achillea millefolium agg. 19
Asteraceae Centaurea jacea 6
Asteraceae Cirsium arvense 26
Asteraceae Cirsium canum 23
Asteraceae Cirsium oleraceum 3
Asteraceae Eupatorium cannabinum 3
Asteraceae Leontodon hispidus 1
Asteraceae Leucanthemum vulgare agg. 1
Asteraceae Tanacetum vulgare 1
Asteraceae Tarax a cum sect. Ruderalia 7
Boraginaceae Symphytum officinale 19
Brassicaceae Cardamine sp.1 1
Cannabidaceae Humulus lupulus 2
Caryophyllaceae Lychnis flos-cuculi 6
Caryophyllaceae Myosoton aquaticum 1
Caryophyllaceae Stellaria graminea 9
Caryophyllaceae Stellaria media 1
Colchicaceae Colchicum autumnale 3
Cyperaceae Carex acuta 3
Cyperaceae Carex acutiformis 1
Cyperaceae Carex hirta 14
Cyperaceae Carex ovalis 1
Cyperaceae Carex pallescens 1
Cyperaceae Carex vesicaria 3
Cyperaceae Carex vulpina 2
Cyperaceae Juncus conglomeratus 4
Cyperaceae Juncus effusus 3
Equisetaceae Equisetum arvense 3
Fabaceae Lathy ru s pra tensi s 15
Fabaceae Lotus corniculatus 1
Fabaceae Trifolium pratense 1
Fabaceae Vicia cracca 8
Fabaceae Vicia cracca 1
Fagaceae Quercus robur 2
Hypericaceae Hypericum maculatum 4
Lamiaceae Galeopsis sp.1 2
Lamiaceae Gl ech oma hederac ea 7
Lamiaceae Mentha sp.1 2
Lythraceae Lythrum salicaria 8
Malvaceae Tilia cordata 1
Orobanchaceae Melampyrum nemorosum 2
Plantaginaceae Veronica chamaedrys 5
Plantaginaceae Veronica serpyllifolia 1
Poaceae Agrostis capillaris 1
Poaceae Agrostis stolonifera 8
Poaceae Alopecurus pratensis 23
Poaceae Arrhenatherum elatius 6
Poaceae Calamagrostis canescens 2
Poaceae Calamagrostis epigejos 17
Poaceae Dactylis glomerata 1
Poaceae Deschampsia cespitosa 12
Poaceae Elymus caninus 3
Poaceae Elytrigia repens 6
Poaceae Fes t uca prat e n s is 3
Poaceae Holcus lanatus 7
Poaceae Molinia caerulea 1
Poaceae Phalaris arundinacea 1
Poaceae Phleum pratense 1
Poaceae Poa palustris 10
Poaceae Poa pratensis 18
Poaceae Trisetum flavescens 1
Polygonaceae Persicaria amphibia 2
Polygonaceae Rumex acetosa 3
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Reintroduction of manual mowing 117
Table 3. (continued)
Family Species No. of samples
Primulaceae Lysimachia nummularia 10
Primulaceae Lysimachia vulgaris 3
Ranunculaceae Anemone nemorosa 1
Ranunculaceae Ranunculus acris 2
Ranunculaceae Ranunculus auricomus 6
Ranunculaceae Ranunculus repens 19
Ranunculaceae Thalictrum lucidum 1
Rosaceae Alchemilla sp.1 3
Rosaceae Potentilla anserina 8
Rosaceae Potentilla erecta 1
Rosaceae Potentilla reptans 3
Rosaceae Rubus sp.1 1
Rosaceae Sanguisorba officinalis 15
Rubiaceae Galium album 4
Rubiaceae Galium aparine 5
Salicaceae Salix sp.1 2
Scrophulariaceae Scrophularia nodosa 1
Urticaceae Urtica dioica 3
Valerianaceae Vale r ian a sp.1 1
Violaceae Viola sp.1 4
species of butterflies and plants were represented in the
analysis.
Beetles were thus excluded from subsequent anal-
yses as unsuitable taxa.
Species richness and manual mowing effect
The paired t-test showed that the species richness of
butterflies was significantly higher on manually mowed
than abandoned sites (t= 4.46; P<0.01) and the same
result was derived from rarefactions (Fig. 2A). Plants
showed a non-significant response (t= 1.03; P= n.s.)
and an almost opposite pattern in species rarefactions
(Fig. 2B).
As analyses on plants showed potentially biased
results (Oksanen 2001), thus we excluded plants from
further analyses.
Effect of manual mowing and surrounding habitats on
species richness and composition of butterflies
During this study, we recorded 28 species of butterflies.
Our results showed that the design using four manage-
ment types was significant (F= 6.98; P<0.01) ex-
plaining 36.66% of adjusted variance.
The rarefactions showed that there was only a
slight difference in the species richness of butterflies be-
tween conservation and manually mowed sites, while
abandoned site showed the lowest species richness
(Fig. 3).
The results of redundancy analysis of butterfly
data (first canonical axis: R2= 19.80%; F= 5.93;
P<0.01; all canonical axes: R2= 24.39%; F=
2.57; P<0.01) showed that some species assem-
blages were associated with disparate management
type – majority of wetland preferring species of con-
servation interest (Lycaena dispar,Phengaris nausit-
hous and P. telejus) were presented only in conser-
vation and manually mowed sites and only generalist
species showed an association with agri-environmental
type of management. Figure 3 also shows that con-
servation and manually mowed sites were quite sim-
ilar in species composition, while there were only a
few species that were more distributed in abandoned
fen.
Discussion
Butterflies
Our comparisons of three taxa showed that there were
differences among them and that, in our case, butter-
flies seem to be a quickly-responding indicator taxa for
the evaluation of manual mowing effects compared to
beetles and plants. Butterflies are more mobile and can
therefore potentially respond more quickly to the mow-
ing effect than plants. This also indicated that with-
out any hydrological restoration (Moreno-Mateos et al.
2012; Zedler & Callaway 1999), or any introduction of
species (Lepš et al. 2007; Klimkowska et al. 2010) in a
site that has been drained and abandoned for a long
time, mowing might increase butterfly species abun-
dance, but not restore the plant community (Hedberg
& Kotowski 2010). The main reason for plants might
be relatively short time compared to butterflies (Valkó
et al. 2012).
Beetles
Flower-visiting beetles were not suitable group for our
aims, even though they have been occasionally used
for the evaluation of disparate management activi-
ties (Noordijk et al. 2009). The most probable rea-
son was that they were relatively species poor and
nearly half of them were singletons and thus proba-
ble tourists (Novotny and Basset 2000), while only four
species reached a reasonable abundance. We also did
not observe beetle species associated with humid habi-
tats, probably caused by the study group of flower-
visiting beetles, which seems to be more generalis-
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118 J. Horak &L.Safarova
Fig. 1. Rarefactions and Chao estimators of total species richness
of observed (A) butterflies, (B) beetles, and (C) plants. Complete
data for species from all samples of manual mowing and abandon-
ment design are included. The solid lines show the sample-based
rarefaction, the two surrounding dashed lines 95% confidence in-
tervals and the upper grey dash-and-dotted lines are the Chao
estimators of the total number of species. Note that the x and y
axes are not to the same scale for (A), (B) and (C).
tic than other specialised groups (Rainio & Niemela
2003).
Plants
Vascular plants as sedentary organisms have a tendency
to lower dispersal ability, which most probably limited
them in potential quick responses to the management
activities in our study (but see Nathan 2006). Seden-
tary behaviour may cause their higher persistence in the
exploited landscape, but they probably need a longer
Fig. 2. Results of species rarefactions comparing the effect of man-
ual mowing and abandonment on the species richness of (A) but-
terflies and (B) plants. The solid lines (black for manual mow-
ing and grey for abandonment) show the sample-based species
rarefactions and the two surrounding dashed lines are 95% confi-
dence intervals. Note that the scales on x and y axes are not the
same for Figs B and C.
time for the response, e.g., restoration of their propag-
ules (Prach 1993). The same reason could influence
relatively low number of species that prefer habitats
influenced by water (e.g., wetland specialists). More-
over, specialized species could vanish before the man-
ual mowing was reintroduced. The next reason for non-
response of the plant community to the management
could be the fact that no hydrological restoration was
conducted (Zedler & Callaway 1999). As our results
showed, the response of plants to the restoration ef-
fect tended to be spatially biased (Oksanen 2001). The
response of wetland specialised species of plants was
nearly the same as the total species assemblage, al-
though there was no significant trend in their response.
Synthesis
Funds for restoration and conservation activities are
limited in rural agricultural landscapes, especially in
those which are not a part of protected areas. Thus, re-
cent limited and short-time funding called for rapid as-
sessment and suitable indicator organisms. In our study,
butterflies were the suitable group for both response to
the manual mowing effect and comparisons with the
surrounding landscape. From the point of view of this
study group, manual mowing of abandoned fen brought
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Reintroduction of manual mowing 119
Fig. 3. Abundance based pie symbols species-environmental plot visualising distribution of species with different response to the type
of management. Note that only species with more than five occurrences are shown. Species with black labels (Leptidea reali,Lycaena
dispar,Phengaris nausithous and P. telejus) are wetland specialists. Result of final values of species rarefactions with 95% confidence
intervals comparing the effect of abandonment, manual mowing, conservation and agri-environmental mowing on the species richness
of butterflies is in the upper right corner.
a species rich habitat, which also promoted endangered
species dependent on wetlands (e.g., Phengaris Blues).
In comparison with the surrounding landscape, butter-
flies were able to rapidly restore their populations or re-
colonise the restored fen, and their species richness was
(quite surprisingly) high, as in the long-term preserved
site. This was probably caused by environmentally-
friendly manual mowing with biomass removal (Cizek
et al. 2012). The highly similar and species rich fauna of
restored sites compared with preserved sites indicates
that the recolonisation of formerly abandoned sites is
the most probable driver, even in highly endangered
and sedentary Phengaris Blues (Fric et al. 2007) asso-
ciated with wetlands (Wynhoff 1998).
Conclusion
The most interesting information derived from our
study is that, in similar cases, the manual mowing ef-
fect is able to create a type of habitat that is, from a
species composition point of view, of conservation in-
terest, and secondly the effect of some species groups
(i.e., butterflies) could be relatively rapid.
Acknowledgements
We would like to thank the non-governmental organisation
Lesák, o.s. (www.lesak.eu) for logistics and the creation of
different management activities, D. Kurková, J. Stratílek, J.
Matoušek and J. Šimek for historical information, L. Bour-
don for English proof reading and two reviewers for sug-
gestions. This research was partly supported by the Grant
CIGA no. 20144302.
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... However in many places, this practice has become economically unsustainable where mechanisation is not possible for example due to difficult access and steep slopes (Török et al. 2011;Valkó et al. 2012). Conservation-driven mowing is an effective practice to maintain high species diversity in abandoned semi-natural grasslands, because it simulates traditional land management practices (Horak and Safarova 2015). It avoids the dominance of grasses and other graminoids and the often irreversible encroachment of shrubs and trees into open areas (Agnoletti 2007;Halada et al. 2011). ...
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Hypothetical models in the scientific literature sug- gest that ecosystem restoration and creation sites fol- low a smooth path of development (called a trajec- tory), rapidly matching natural reference sites (the target). Multi-million-dollar mitigation agreements have been based on the expectation that damages to habitat will be compensated within 5-10 years, and monitoring periods have been set accordingly. Our San Diego Bay study site, the Sweetwater Marsh Na- tional Wildlife Refuge, has one of the longest and most detailed records of habitat development at a mit- igation site: data on soil organic matter, soil nitrogen, plant growth, and plant canopies for up to 10 years from a 12-year-old site. High interannual variation and lack of directional changes indicate little chance that targets will be reached in the near future. Other papers perpetuate the trajectory model, despite data that corroborate our findings. After reviewing "trajec- tory models" and presenting our comprehensive data for the first time, we suggest alternative management and mitigation policies.