Applied Vegetation Science 17 (2014) 193–200
SPECIAL FEATURE: ECOLOGICAL RESTORATION
Vegetation succession in restoration of disturbed sites
in Central Europe: the direction of succession and
species richness across 19 seres
ra Rehounkov
, Kamila Lencova
, Alena Jırova
, Petra Konvalinkova
,
Karel Prach, Kla
a
Ondrej Mudrak, Vojtech Student, Zdenek Vanecek, Lubomır Tichy, Petr Petrık,
Petr Smilauer
& Petr Pysek
Keywords
Ordination; Restoration; Species number;
Succession; Target species
Nomenclature
Kubát et al. (2002)
Received 26 February 2013
Accepted 12 July 2013
Co-ordinating Editor: Lawrence Walker
Prach, K. (corresponding author,
[email protected]),
, K. (klara.rehounkova@
Rehounkov
a
gmail.com),
, K. ([email protected]),
Lencova
, A. ([email protected]),
Jırova
, P. (petra.konvalinkova@
Konvalinkova
cmsterk.cz),
Student,
V. ([email protected]),
Vane
cek, Z. ([email protected]) &
Smilauer,
P. ([email protected]): Faculty of
e
Science USB, Branisovska 31, CZ-37005, Cesk
jovice, Czech Republic
Bude
, K. , Jırova
, A. &
Prach, K. , Rehounkov
a
Mudrak, O. ([email protected]):
Institute of Botany, Academy of Sciences of
the Czech Republic, Dukelska 135, CZ-37981,
, Czech Republic
Trebon
, L. ([email protected]): Department of
Tichy
Botany and Zoology, Masaryk University,
Kotl
arsk
a 2, CZ-61137, Brno, Czech Republic
Petrık, P. ([email protected]) &
Pysek, P. ([email protected]): Institute of
Botany, Academy of Sciences of the Czech
Republic, CZ-252 43, Pr
uhonice, Czech Republic
Py
sek, P.: Department of Ecology, Faculty of
Science, Charles University in Prague, CZ-128
44, Prague 2, Czech Republic
Abstract
Questions: (1) How do seres differ with respect to vegetation changes? (2)
What are the directions of succession? (3) How do species numbers change? (4)
How do target species, i.e. those typical of natural and semi-natural vegetation,
participate in succession? (5) Are spontaneously developed successional stages
acceptable from the point of view of ecosystem restoration?
Location: Extracted peatlands, bulldozed sites in forests destroyed by air pollution, an emerged bottom of a water reservoir, corridors of former Iron Curtain,
artificial fishpond islands and barriers, sedimentary basins, spoil heaps from
mining, stone quarries, forest clearings, road verges, sand and gravel-sand pits,
ruderal urban sites, river gravel bars and abandoned arable fields, located in
various parts of the Czech Republic in Central Europe.
Methods: Phytosociological releves were recorded in 10–25 m2 plots located in
the centre of representative successional stages defined by their age, ranging
from 1 to 100 yrs. In total, we obtained 2392 vegetation samples containing 951
species. We performed DCA ordination to compare 19 seres. Desirable target
species were considered as those representing (semi)-natural vegetation and all
Red List species.
Results: The seres studied are more similar in their species composition in the
initial and early stages, in which synathropic species prevail, than in the later
stages when the vegetation differentiates. This divergence is driven mainly by
local moisture conditions. In most cases, succession led to woodland, which usually established after ca. 20 yrs. In very dry or wet places (with limited presence
of woody species) open vegetation developed, often highly valuable from the
restoration and conservation point of view. The total number of species and the
number of target species increased in the majority of seres with successional age.
Conclusions: The vegetation in the sites studied formed a continuum along a
moisture gradient and by successional age. The individual seres largely overlapped in their species composition; the sere identity was not significant. Spontaneous succession usually proceeded towards woodland, except at very dry or
wet sites, and generally appeared to be an ecologically suitable way of ecosystem
restoration of disturbed sites because target species became dominant over time.
Introduction
Numerous different seres have been described from various parts of the world, often in remarkable detail (Walker
& del Moral 2003). There is, nevertheless, still a lack of
studies that compare a number of seres over a broad
geographic scale in a rigorous, quantitative way (Prach
et al. 1993, 2001, 2007, 2013; Anderson 2007; Pr
evosto
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193
Succession in disturbed sites
K. Prach et al.
et al. 2011), largely because the use of different methods
ehounmakes cross-study comparisons difficult (Prach & R
kova 2006). This limitation can be overcome by sampling
different seres in the same way, using a comparable size of
plots and compatible methods of vegetation sampling. Data
that match these criteria can also be extracted from different published and unpublished sources. Such comparable
data make it possible to identify common and general
trends in successional development among seres in a
meta-analysis (Stewart et al. 2008). Quantitative comparisons across various seres, such as those presented in this
paper, provide an overview of processes taking place in
developing vegetation in a broader context. This knowledge is useful in both theory and practice. The theoretical
side mainly includes general questions about successional
development towards potential vegetation, convergence
or divergence of succession, the rate of succession or
changes in species diversity and species traits during succession (Walker & del Moral 2003). Practical aspects concern potential insights for ecological restoration (Walker
et al. 2007; Prach & Walker 2011).
In the Central European country of the Czech Republic
it has been estimated that, in past decades, about 2% of its
area (about 1800 km2) was at least temporarily exposed to
spontaneous succession that started on bare ground. About
0.8% originated from recent mining activities conduced in
the second half of the 20th century (Prach et al. 2011),
and about 0.6% is abandoned arable land (Hrazsk
y 2006).
The rest includes mostly habitats disturbed by construction
activities. Restoration may be desirable at some of these
sites.
We have two contrasting options when it comes to
restoring disturbed land: (1) to rely on spontaneous succession, or (2) to adopt technical measures. The latter
approach has prevailed in post-mining sites in the Czech
ys & Branis 1999) and generally involves levRepublic (St
elling of the surface, spreading of organic material rich in
nutrients and sowing of commercial seed mixtures or
planting trees in regular rows. Similarly, road banks, sedimentary basins and various sites disturbed by construction
works are often reclaimed by technical means. But is this
really needed in all situations? Can desirable vegetation be
attained by spontaneous development within a reasonable
time frame? Spontaneous succession can, to a certain
extent, be manipulated to reach the target stage. A continuum thus exists between technical reclamation and spontaneous succession (Prach & Hobbs 2008). In this paper,
we compare 19 spontaneous seres, each subjected to a different kind of disturbance (Table 1). We focus on similarities in vegetation of different seres, directions of succession
and species richness. Special attention was paid to the role
of target species (defined as representatives of natural and
semi-natural vegetation in a given region; Van Andel &
194
Aronson 2012); these species are desirable from the restoration point of view and their participation can be used to
assess succession from the restoration perspective.
Our goal was to answer the following questions: (1)
how do the seres under study differ with respect to vegetation changes over time; (2) what are the directions of succession, and can they be generalized across different
environments; (3) how does species richness change over
the course of succession, and are there any general trends;
(4) how do target species participate; and (5) are spontaneously developed successional stages acceptable from the
point of view of ecosystem restoration?
Methods
Study sites
The successional sites considered in this study are located
in the Czech Republic, but the range of habitats makes
them representative of Central Europe. We sampled 19
habitats in which it was possible to identify seres. Twelve
of the seres have already been described, at least partly, in
separate studies (see Table 1 for description and references) and used for partial analyses (Prach et al. 1993,
1999, 2001, 2007, 2013); seven are newly included into
this study. The seres differed in the number of samples, the
number of sampled sites, the geographic area in which
they occurred, and successional age. The differences in
these characteristics resulted from the availability of successional stages and our capacity to observe them. All but
one of the seres (on a river gravel bar) were triggered by
human disturbances and started mostly on bare ground;
the only exceptions were forest clearings created in
Norway spruce plantations, but even here the cover of
herb layer at the beginning of succession was usually low.
We identified successional stages that were homogeneous,
with a known year of the site’s creation and left to undisturbed development since succession had started.
Analysis
Sample plots of 10–25 m2 were placed in the centre of the
above stages. We combined the space-for-time substitution
approach and permanent plot research (Pickett 1989)
because some sites were observed over multiple years
whilst others were observed for only 1 yr (compare figures
in the 3rd and 4th column in Table 1). We identified all
vascular plants present in each sample plot and visually
estimated their percentage cover (Kent & Coker 1992). In
some cases, the cover was estimated using semi-quantitative scales (Braun-Blanquet, Domin) and then transformed into percentage cover according to Van der Maarel
(1979). The following successional stages were delimited
and used for calculation of centroids (see below):
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Succession in disturbed sites
K. Prach et al.
Table 1. Main characteristics of seres under study, listed approximately according to decreasing site moisture following their order in Fig. 1. In the figure,
each sere is referred to by its number (1–19).
Sere
Location in the
Czech Republic
No. of
samples
No. of
sampled
sites
Age of
stages [yrs]
Data sources and references
1. Extracted peatlands
2. Bulldozed sites in forests destroyed
by air pollution
3. Emerged reservoir bottom
4. Corridors of former Iron Curtain*
5. Artificial fishponds, islands and barriers
SW
W
267
14
267
12
1–100
1–20
Konvalinkova & Prach (2010); Bastl et al. (2009)
Pysek (1992); Prach et al. (2001)
W
SW
S
12
146
108
1
146
80
1–12
7–35
1–126
6. Sedimentary basins
7. Spoil heaps from uranium mining
8. Acidic stone quarries
9. Forest clearings
W and N
Central
Central
Central and N
21
85
135
62
21
85
135
53
1–41
7–32
1–86
1–107
10. Road verges
11. Spoil heaps from black coal mining
12. Spoil heaps from brown coal mining
S
Central
W
44
88
147
44
88
82
1–14
10–100
4–55
13. Sand and gravel-sand pits
14. Spoil heaps from brown coal mining
15.Urban ruderal sites
16. River bars
Various parts
NW
W
NE and S
233
75
36
70
233
75
3
41
1–75
1–45
1–12
1–150
17. Abandoned fields
Various parts
288
181
0–91
18. Basalt quarries
NW
441
441
1–80
19. Limestone quarries
E
120
120
0–35
K. Prach, unpublished data; Prach et al. (2001)
Springar (1995); Kostel (2000)
K. Prach, unpublished data; V. Student,
unpublished
data; Rejmanek & Rejmankova (2002)
K. Prach, unpublished data; Kovar (2004)
Dudıkova (2007)
Trnkova et al. (2010)
P. Smilauer,
unpublished data; P. Petrık,
unpublished data
Litvin (2000)
H. Dvor
akova, unpublished data; Prach et al. (2013)
K. Prach, unpublished data; O. Mudrak, unpublished
data; Frouz et al. (2008); Mudrák et al. (2010)
Rehounkov
a & Prach (2006, 2008, 2010)
Prach (1987); Hodacova & Prach (2003)
Pysek (1978); Prach et al. (2001)
cek,
K. Prach, unpublished data, Z. Vane
unpublished data
Osbornova et al. (1990); Prach et al. (2007);
Jırov
a et al. (2012)
Novak & Prach (2003); Nov
ak & Konvicka (2006);
Prach et al. (2013)
L. Tichy, unpublished data
*The corridors of the former Iron Curtain were created along the state border during the communist dictatorship. It was a fenced strip of land about 6 m in
width, kept without vegetation by continuous mechanical disturbance and herbiciding. It was abolished in 1989.
(1) initial, 1–3 yrs; (2) early, 4–10 yrs; (3) middle, 11–
25 yrs; (4) late, 26–40 yrs; (5) old, >40 yrs.
We carried out a detrended correspondence analysis
(DCA) of these vegetation samples (phytosociological
releves) in the form of a matrix comprising 2392 samples
and 951 species. Isolines, indicating the total number of
species and the number of target species, were included
into the resulting diagrams. They represent response surface fitted with a loess model (see Cleveland & Davlin
1988). Influence of the successional age as an explanatory
variable of species composition was tested with a permutation test in canonical correspondence analysis (CCA), with
999 permutations. In the case of repeated sampling, plot
identity and elsewhere the habitat type were used as covariables defining permutation blocks. Prior to the ordinations using the CANOCO application (Microcomputer
Power, Ithaca, NY, USA), we logarithmically transformed
the data and down-weighted rare species. The use of the
unimodal methods was justified by the length of the gradi
ent, which reached 8.96 SD units (see Leps & Smilauer
2003).
We classified target species according to Ellenberg et al.
(1991) into the following categories representing: (1) dry
open habitats (Festuco-Brometea, Trifolio-Geranietea,
Sedo-Scleranthetea), (2) mesic and dump grasslands (Molinio-Arrhenatheretea, Nardo-Callunetea), (3) wetlands
(Phragmitetea, Oxycocco-Sphagnetea, Scheuchzerio-Caricetea nigrae), (4) scrublands and woodlands (RhamnoPrunetea, Querco-Fagetea, Quercetea robori, VaccinioPicetea). Target species included all Red List species
(Proch
azka 2001), including those affiliated with other
habitats. Ruderals and weeds representing Bidentetea,
Chenopodietea, Secalietea, Artemisietea, Agropyretea,
Plantaginetea and Epilobietea (Ellenberg et al. 1991), and
all alien species (Pysek et al. 2012a,b), except Red List species (Danihelka et al. 2012; Grulich 2012), were considered as non-target, undesirable species.
Results
In the DCA ordination, individual seres are arranged along
the first ordination axis (k = 0.727), which approximately
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Succession in disturbed sites
K. Prach et al.
(a)
(b)
(c)
(d)
Fig. 1. DCA ordination. Envelopes (a) enclose the positions of all samples of each sere. Ordination of species is presented in (b) with passively projected
successional age (Age). Different colours are used for target species typical of wetlands (blue), woodlands (black), mesic grasslands (green) and dry
grasslands and other dry habitats (yellow); non-target, synathropic species and alien species are in red. Abbreviations of species names are composed of
the first four letters of the generic name and the first four of the epithet; for full names, see the Appendix S1. Only species with the highest weights (>2%)
are shown. DCA ordination with isolines (loess curves), demonstrating the average total number of species (c) and average number of target species (d) per
vegetation sample. The arrows connect centroids of case scores representing the youngest and oldest observed age category of each sere and thus
indicate the main direction of succession. The numbers of seres correspond with those in Table 1.
reflects site moisture conditions (Fig. 1a). On the right side
of the diagram are rather dry basalt and limestone quarries
in relatively warm regions; on the left are wet peatlands,
which are often found in mountainous areas. The second
ordination axis (k = 0.565) reflects successional age, as is
evident from the directions of the arrow in Fig. 1b. The
correlation coefficient (R) between the successional age of
samples and their score on the second ordination axis is
0.839 (P < 0.001). The successional age, as the only
explanatory variable, explained 19.3% of variability of the
vegetation data in the CCA ordination (F = 22.25,
P < 0.01). The identity of the sere appeared non-significant in the CCA analysis (F = 2.13, P = 0.072).
Some of the envelopes, which enclose samples of a particular successional sere, spread broadly along the first
ordination axis, thus comprising a wide range of dry to wet
habitats (typically sand and gravel-sand pits, stone quarries
196
or abandoned fields). Other envelopes were limited in
their extent along the gradient, indicating rather homogenous successional vegetation (typically bulldozed sites in
forest destroyed by atmospheric pollution, emerged bottom of water reservoirs and urban ruderal sites, all represented only by one or a few localities). Most seres appeared
in the centre of the first axis gradient. The fact that the
envelopes largely overlap indicates that there are strong
similarities among the seres, and the successional vegetation forms a continuum along the moisture gradient.
A comparison between Fig. 1a,b reveals that succession
generally runs from stages composed predominantly of
synanthropic species (i.e. ruderals and weeds, such as
Elytrigia repens, Cirsium arvense, Tussilago farfara, Artemisia
vulgaris, etc.), located in the upper part of the diagram in
Fig. 1b, towards a woodland, represented by species in
the bottom part of the diagram (e.g. Quercus spp., Fraxinus
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K. Prach et al.
excelsior, Acer campestre, Crataegus spp. and Poa nemoralis).
Species representing dry grasslands and other open and
dry habitats (dry sandy or sunny rocky sites) appear on the
right part of the diagram (e.g. Bromus erectus, Festuca rupicola, Brachypodium pinnatum and Sedum album), and species
representing wetlands and wet grasslands are on the left
(e.g. Eriophorum vaginatum, Molinia caerulea, Agrostis canina
and Juncus effusus). Species representing mesic grasslands
appeared in the centre of the diagram (e.g. Agrostis capillaris, Poa pratensis, Festuca rubra and Dactylis glomerata).
The total number of species exhibited a general tendency to increase or, in a few cases, remained more or less
equal over the course of succession (Fig. 1c) or decreased
(peatlands and forest clearings). The total number of species increased towards the dry end of the moisture gradient
represented by the first ordination axis. High total numbers
of species are typical of seres in dry and sunny habitats,
namely alkaline stone quarries and the majority of the
abandoned fields studied. The high numbers of target species were also present in stone quarries and abandoned
fields (Fig. 1d). Seres started with the lowest number of
target species in abandoned fields and at some of the mining sites. In peatlands the number of target species clearly
decreased. At sites bulldozed following forest destruction
by air pollution, in forest clearings and in corridors of the
former Iron Curtain, the numbers of target species
decreased only slightly. In all the other seres the numbers
of target species increased. Comparing Fig. 1c,d, one can
see that in the late stages the target species represent on
average about three-quarters of the total number of
species.
Discussion
The main direction of seres considered in this study is
towards re-establishment of woodland, which is the typical
potential vegetation in the study area (Neuh€auslov
a 2001;
Chytr
y 2012). Succession towards woodland is typical of
areas with relatively humid climates (Walker & del Moral
2003), which includes Central Europe. Reed or sedge wetlands generally establish over the course of succession in
the wettest sites, provided that the water table remains sufficiently high. These wet microhabitats are typically found
in low areas of spoil heaps (Prach et al. 1999), sand and
ehounkova & Prach 2006), extracted
gravel-sand pits (R
peatlands (Konvalinkova & Prach 2010), some stone quarries (Trnkova et al. 2010), artificial fishpond islets (Rejmánek & Rejmánková 2002) and exceptionally in abandoned
fields (Jırova et al. 2012). On the other hand, grassland or
shrubby grassland usually develop at dry sites in stone
quarries (Novak & Prach 2003; Tich
y 2012), on spoil heaps
(Prach 1987) or in abandoned fields (Jırova et al. 2012).
Woody species also usually do not establish on shallow,
rocky substrates in quarries (Nov
ak & Prach 2003; Tich
y
2012), on dry, sandy soils in certain sand and gravel-sand
ehounkov
pits, depending on the climatic region (R
a &
Prach 2006), or on sporadically toxic substrates (Kov
ar
2004). Both wetlands and grasslands are valuable from the
point of view of nature conservation and restoration
because many heliophilous species can establish and survive in such open habitats without being out-competed by
taller woody species. The importance of such open habitats
has been documented in detail – for plants, see papers cited
in Table 1; for other organisms, mainly arthropods, see
ehounkov
Benes et al. (2003), Tropek et al. (2010), R
a
et al. (2011) or Heneberg et al. (2013); Vojar (2006)
reports that wetlands developed at former mining sites are
crucial for the survival of amphibians in the Czech Republic.
The sites in this study formed a continuum along the
moisture gradient, and site moisture appeared here to be
more important for vegetation development than successional age. Unfortunately, it was technically impossible to
measure site moisture over all the studied sites. Site
moisture, temperature, nutrients, substrate structure and
acidity are most frequently reported, beside successional
age, as the most important environmental factors determining the course of succession (Walker & del Moral
2003). For Central European seres, including some of
those considered in this study, the macro-climate and soil
pH were identified as significantly influencing the development of vegetation (Prach et al. 2007). Regional and local
species pools, in addition to site conditions, influence community species pools in general (Zobel et al. 1998) and the
participation of species in succession in particular
ehounkov
(R
a & Prach 2008).
The majority of the seres can be considered as primary
succession, a few as secondary succession (abandoned
fields, forest clearings) or exhibiting features of both (artificial fishpond islands and barriers). In a previous study
(Prach et al. 2001) we showed that primary vs secondary
status of seres did not have a significant effect on the
course of succession, thus, we did not consider this status
as a factor in the present paper.
Our seres, representing a range of initial site conditions,
exhibited mostly increasing trends in the total numbers of
species and target species, a pattern usually found especially in primary seres (Walker & del Moral 2003). The
number of target species is perhaps a more important characteristic in ecological restoration than the total number of
species (Van Andel & Aronson 2012). From the restoration
point of view, species numbers are a more relevant indicator in older stages than in more ephemeral initial stages.
Between the initial and older stages, species richness usually fluctuates, with minima being reached when strong
competitors dominate (Peet 1978). Competitive exclusion
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Succession in disturbed sites
K. Prach et al.
is typically more common in secondary rather than in primary seres (Glenn-Lewin et al. 1992). Our Fig. 1c,d only
include general trends between the youngest and the oldest successional stages. Moreover, the seres differ in the
time over which succession was recorded, and some seres
were sampled in plots of somewhat different size, i.e. by
10–25 m2. Plot size can influence the absolute number of
species but is not likely to effect overall trends (Chytr
y
2001). In our case, the regression of the number of species
on the size of sampling plots was not significant (R = 0.12,
P > 0.05, n = 2392). Extracted peatlands are an exception
among our seres because they exhibited a clear decrease in
both number of total and target species over time. This pattern is probably caused by competitive exclusion under a
dense canopy of woody species, which forms gradually,
and/or an expansion of a strong dominant in the herb layer
in later successional stages, e.g. Phragmites australis or
Molinia caerulea (Konvalinkova & Prach (2010).
The establishment of a continuous vegetation cover
occurred within 1–15 yrs in all seres (see also Prach &
ehounkova et al. 2011), with its fastest forPysek 2001; R
mation in secondary seres, especially in abandoned fields
(Osbornova et al. 1990). Heavily eroded, stony, toxic or
ehounkov
exposed sites represent exceptions (R
a et al.
2011), but such sites were not sampled in our study. In
Central Europe it usually takes ca. 25 yrs of succession to
reach (semi)-natural vegetation, usually woodland, which
subsequently changes little in species composition (Prach
2003). Thus, the establishment of woodland does not necessarily need to be aided by planting trees, as is often practised in many disturbed sites subjected to technical
reclamation. Quite the contrary: it is sometimes desirable
from the ecological point of view to slow down the establishment of a woodland by grazing or by felling trees and
removing shrubs because open habitats are often more
ehounkov
valuable than wooded ones for biodiversity (R
a
et al. 2011). Only a slightly manipulated natural succession (Luken 1990) or completely spontaneous succession
should therefore be preferred over technical reclamation
in the habitats that we studied. Compared to technical reclamation, spontaneous succession results in the development of more diverse plant communities with species
composition closer to that of (semi)-natural vegetation
(Hodacova & Prach 2003), and represents a cheaper option
ehounkova et al. 2011).
(R
Restoration ecologists are sometimes worried about
invasions of alien species if a site is left to spontaneous
development. This is certainly justified in some parts of the
world (e.g. Luken 1997; Flory & Clay 2010; Saccone et al.
2010). In our case, the only alien species that may become
a serious invader during succession in disturbed habitats is
the black locust (Robinia pseudacacia), especially in certain
sand and gravel-sand pits and stone quarries in dry and
198
warm regions. Even at these sites, it was only observed if
an invasive population occurred within ca. 100 m
ehounkov
(R
a & Prach 2008). This species usually forms
dense, species-poor stands dominated by nitrophilous
species in the herb layer, which is not acceptable from the
restoration point of view (Kowarik 1992; Pysek et al.
2012a,b).
We conclude that spontaneous vegetation succession
appears to be a suitable means of ecosystem restoration for
disturbed sites under Central European conditions.
Acknowledgements
The study was supported by the following grants: GACR
P505/11/0256, RVO 67985939 and GAJU 138/2010/P.
The authors thank Lawrence Walker, Kristina AndersonTeixeira and anonymous reviewers for their comments,
Frederick Rooks for language revision, and former students
for providing us with their primary data. Petr Pysek was
supported by institutional resources of the Ministry of
Education, Youth and Sports of the Czech Republic
and acknowledges support of the Praemium Academiae
award from the Academy of Sciences of the Czech
Republic.
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Supporting Information
Additional supporting information may be found in the
online version of this article:
Appendix S1. List of species names used in Fig. 1b.
Applied Vegetation Science
Doi: 10.1111/avsc.12064 © 2013 International Association for Vegetation Science