Ecological Engineering 17 (2001) 55 – 62

www.elsevier.com/locate/ecoleng

Using spontaneous succession for restoration of

human-disturbed habitats: Experience from Central Europe
Karel Prach a,*, Petr Pyšek b
a
Faculty of Biological Sciences, Uni6ersity of C& eské Budějo6ice, and Institute of Botany,
Academy of Sciences of the Czech Republic, Branišo6ská 31, CZ-370 05 C& eské Budějo6ice, Czech Republic
b
Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice u Prahy, Czech Republic

Accepted 28 July 2000

Abstract

Total vegetation cover, cover of woody species, and participation of dominants in later (pre-forest) successional
stages of 16 successional seres starting on bare ground in various human-disturbed habitats (Czech Republic, Central
Europe) were compared and evaluated from the viewpoint of restoration ecology. Continuous vegetation was formed
before the 15th year of succession in all the seres studied. The establishment and expansion of woody species tended
to be easier under moderate environmental conditions and was retarded in rather extreme habitats (dry, wet, nutrient
poor, acid). The later-successional dominants were all native species and some rare and endangered species were
recorded in these stages. The main conclusion is that spontaneous succession can be relied upon in restoration
projects except in the case of extreme, especially toxic substrata. To leave a site to allow spontaneous processes to
revegetate it is especially advantageous if the disturbed site is small, surrounded by natural vegetation, and if site
conditions were not principally altered by the disturbance. Spontaneous succession is cheap and spontaneously
revegetated sites usually exhibit higher natural value. © 2001 Elsevier Science B.V. All rights reserved.

Keywords: Vegetation; Succession; Human-disturbed habitats; Total cover; Woody species; Restoration

1. Introduction Dettmar, 1996; Bradshaw, 1997). The present pa-

Spontaneous succession of vegetation is receiv-
ing increasing attention in various restoration
projects, especially in developed countries such as
the United States, United Kingdom, The Nether-
lands, and Germany (Luken, 1990; Rebele and
* Corresponding author.
E-mail addresses: prach@bf.jcu.cz (K. Prach),
sek@ibot.cas.cz (P. Pyšek).
per is based on experiences from Central Europe
(Czech Republic) where technocratic approaches
still prevail in the restoration of degraded land. In
this region, there are many sites disturbed by
various mining and building activities but the
money available for restoration of these habitats
is often limited. Large disturbed areas are there-
fore often abandoned so that spontaneous succes-
py-
sion will start, which represents an interesting
phenomenon and object of ecological studies. At

0925-8574/01/$ - see front matter © 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 9 2 5 - 8 5 7 4 ( 0 0 ) 0 0 1 3 2 - 4
56 K. Prach, P. Pyšek / Ecological Engineering 17 (2001) 55–62
the same time, the spontaneous processes may
lead to desired developments which are often the
targets of restoration activities. Such targets in-
clude erosion prevention, enhancement of wildlife
and naturalness and social-economic benefits.
In previous studies, we compared a number (15)
of successional seres starting on bare ground and
running in various human-disturbed habitats. The
comparison was focused on the rate of succession,
changes of species traits during succession, and
characters of successional dominants (Prach et al.,
1993, 1997; Prach and Pyšek, 1994a,b, 1999). In
the present study, the data are used to answer the
following question: Under which circumstances
(given by site origin, history, and intended pur-
pose) and in which habitats (i.e. under which
ecological conditions) can spontaneous succession
be considered as a potentially useful tool in
restoration projects? We concentrated on the
changes in total vegetation cover, the role of
woody species, and characteristics of later-succes-
sional dominants because these aspects of succes-
sion are eminently important for achieving
restoration targets.
2. Material and methods
Cover data were obtained from published case
studies and unpublished records for 16 sites with
successional seres that had started from bare
ground in human-disturbed habitats located in the
western part of the Czech Republic. The duration
of the seres was between 12 and 76 years. In the
majority of seres, the cover of each species present
was estimated annually (by phytosociological
relevés and/or the point-quadrat method) in per-
manent plots fixed immediately after creation of
the site. In some seres, comparable stages of dif-
ferent age were used to infer the course of succes-
sion, or this method was combined with annual
sampling. Particular seres were described in detail
elsewhere (Prach et al., 1993; Prach and Pyšek,
1994a,b; Prach et al., 1997; Prach and Pyšek,
1999) and are thus only briefly characterised here.
Primary vs. secondary seres are distinguished by
abbreviation in brackets (PS=primary succes-
sion, SS= secondary succession).
The following types of disturbed sites were
studied:
(a) Large spoil heaps (PS) from open-cast
brown coal mining. The duration of the succes-
sion for which the species exchange was recon-
structed was 40 years (data from Prach, 1987 and
unpubl.). More than 90% of the spoil heaps area
had a characteristic successional sequence with
initial stages with various annuals and biennials
persisting for 8 years and followed by stages
dominated by perennial forbs (especially Tanace-
tum 6ulgare, Artemisia 6ulgaris, Cirsium ar6ense).
After 16 years, grasses (especially Calamagrostis
epigejos and Arrhenatherum elatius) expanded
with scattered shrubs and trees (esp. Sambucus
nigra and Betula pendula). In this stage, the suc-
cession seemed to be arrested for a long time. In
wet depressions with a water table near the soil
surface and occasional flooding events, the early
successional dominant Typha latifolia was fol-
lowed by Phragmites australis after 11 years of
succession.
(b) Abandoned sand pit (PS) for which 24 years
of succession were documented (Kočár and Prach,
unpubl.). Immediately after the start of the suc-
cession, Pinus syl6estris established and gradually
formed a closed pine woodland with a sparse herb
layer.
(c) Extracted peatland (PS); (20 years of succes-
sion; Bastl, unpubl.); two localities were consid-
ered separately, differing in the water table
position. Molinia coerulea formed a dense stand at
the drier site, whereas Juncus effusus prevailed at
the wet site. Both sites developed a woodland
vegetation with Betula pendula, Pinus syl6estris,
and Salix spp.
(d) Reclaimed sites in areas deforested due to air
pollution, formerly covered by Norway spruce
(Picea abies) (18 years of succession, Pyšek, 1992).
To facilitate the replanting of spruce, the sites
were bulldozed which resulted in plots with grass
cover and topsoil removed (PS) and mounds
formed by the dumped material (SS). If left with-
out planting, two grasses, A6enella flexuosa and
Calamagrostis 6illosa, gradually formed a compact
cover in both habitat types. The former species
prevailed in the parts with topsoil removed, while
the latter was dominant on the mounds.
 K. Prach, P. Pyšek / Ecological Engineering 17 (2001) 55–62
(e) Barriers around a newly constructed fishpond;
(15 years of succession, Prach, unpubl.). Two
particular seres were distinguished, i.e. on barriers
formed by sandy subsoil (PS) where a pine-birch
wood established early in succession, and on or-
ganic topsoil (SS) where Alnus glutinosa expanded
with Phalaris arundinacea dominating in the
understory.
(f) Ruderal urban sites in the town of Plzeň;
(succession of 12 years). Particular seres were
distinguished according to the nutrient status, i.e.
poor (PS), moderately rich (SS) and rich in nutri-
ents (SS), data from Pyšek (1978). Succession at
the nutrient-poor sites started with stages of rud-
eral species, after which Populus tremula gradually
expanded. At the moderately rich sites, Sambucus
nigra and Salix caprea dominated in later stages;
the former species dominated also in the nutrient-
rich sites.
(g) Exposed bottom of a destroyed water reser-
6oir (SS); (succession of 12 years, Frantı́k, Os-
bornová and Prach, unpubl.). A thin wood of
Betula pendula with very compact Calamagrostis
epigejos sward developed rapidly at this site. Veg-
etation in a specific acidic zone differed by the
absence of birch.
(h) Oldfields (SS). Three seres were distin-
guished according to the soil moisture conditions,
i.e. xeric (duration 76 years), mesic (55 years) and
wet (32 years) (data from Osbornová et al., 1990,
and Prach, unpubl.). A compact grassland domi-
nated by Poa angustifolia and Festuca rupicola
and with only very sporadic occurrence of shrubs
developed in the xeric sites. In the mesic sites,
grasses such as Arrhenatherum elatius were over-
grown by Prunus spinosa and then by a very dense
stand of Crataegus species. The expansion of
Fraxinus excelsior started after 50 years of succes-
sion. In wet sites, Petasites hybridus rapidly ex-
panded and was succeeded by Phragmites
australis later on.
The following parameters were used here to
describe the course of succession:
(a) Total vegetation cover was taken from vege-
tational relevés and the point-quadrat sampling.
(b) In each sere, the total cover of woody
species was calculated as the sum of percentage
covers of all woody species present in a given
year.
57
(c) Later-successional dominance was expressed
as the sum of the maximum values of species
cover achieved in particular seres after the 10th
year of succession. Later-successional dominants
(i e. those of pre-forest successional stages) were
selected by ranking the species according to this
value.
3. Results
Increase in the total vegetation cover was rapid
in many seres (Fig. 1) and continuous vegetation
was formed before the 15th year of succession in
all the seres studied. The vegetation cover was
rather lower only in very wet depressions in spoil
heaps and in nutrient-poor sites (e.g. the sand pit).
The increase in cover was relatively fast in seres
corresponding to secondary succession, with soil
seed bank and organic topsoil present at the onset
of succession, which was the case on abandoned
bare soils such as old fields and some ruderal
urban sites. Most primary seres developed much
more slowly on deposits created by mining activi-
ties and sites with removed topsoil.
More variable trajectories can be seen in the
increase of the cover of woody species (Fig. 2).
They were missing from succession or their partic-
ipation was negligible in xeric and wet oldfields
and in wet depressions on spoil heaps from coal
mining. The highest cover of woody species was
recorded in mesic oldfields, in a sand pit, and on
dumps around the newly constructed pond. In all
these seres, close woods were formed.
The twenty most dominant forbs and grasses
are listed in Table 1 (nomenclature from Roth-
maler et al., 1982). Fifteen woody species (8
shrubs, 7 trees) reached a cover of at least 1% in
any of the seres (Table 1), whereas only 9 attained
a cover of \ 10% in at least one sere, and 4 of
them reached a cover of over 50% (Betula pen-
dula, Pinus syl6estris, Sambucus nigra, and
Crataegus sp.). The largest numbers of woody
species appeared in mesic oldfields (9), on urban
sites with a moderate amount of nutrients (6), and
in the abandoned peat-extracted area (6). Only
one woody species was recorded on the emergent
bottom (Betula pendula) and at the nutrient-poor
58 K. Prach, P. Pyšek / Ecological Engineering 17 (2001) 55–62

urban sites (Populus tremula) and none in wet
oldfields and in wet depressions on spoil heaps.
4. Discussion
The successional seres studied included a rela-
tively wide range of human-disturbed habitats of
the Central European landscape and may be con-
sidered as a representative sample of such sites in
the region. Hence, the results can be used for
some generalisations and discussed in the context
of restoration ecology. We are nevertheless aware
that under specific conditions, especially on ex-
treme substrata, the course of succession can be
different. Increase of vegetation cover on toxic
substrata may be much slower than found in the
present paper (Banásová, 1976). The increase in
cover was generally faster in habitats with a sec-
ondary succession, particularly on abandoned
bare soil. However, the distinction between pri-
mary and secondary seres seems to be rather
vague (Glenn-Lewin et al., 1992) as was shown
previously by using the same data as treated in
the present paper (Prach et al., 1997). Taking the
total vegetation cover as a measure, spontaneous
successional processes can obviously be effective
in restoration of non-extreme habitats.
The proportion of woody species in the vegeta-
tion varied remarkably (Fig. 2). A whole array of
factors may have played a role here, such as
abiotic site conditions, competition from herbs,
presence of the sources of diaspores, intensity of
vectors transporting diaspores and their timing,
animals and man activities, and purely stochastic
factors (Olsson, 1987; De Steven, 1991). The high
variability in the participation of woody species
can be observed even within one type of succes-
sion in the same region (Osbornová et al., 1990
for the oldfields). It seems that the establishment
and expansion of woody species tends to be more
rapid under moderate environmental conditions
than in rather extreme habitats such as dry, very
wet, nutrient-poor, and acidic sites (Prach and
Pyšek, 1994a). The establishment of woody spe-
cies was also retarded in secondary seres on highly
productive sites such as wet oldfields, where com-
petitive herbs immediately formed a dense, com-

Fig. 1. Increase in the total vegetation cover in the studied successional seres (16). Abbreviations: Eb — emerged bottom; Bp —
bulldozed plots, and Bm — mounds in the area deforested due to air pollution; Ds — sandy, and Dp — peaty dumps around a
newly constructed pond; Sp — sand pit; Ur — urban ruderal sites rich, Up — poor, and Um — moderate in nutrients; PeP —
wetter, and PeB — drier extracted peatland; Ow — wet, Om — mesic, and Ox — xeric oldfields; Sh — spoil heaps, and Shw —
wet depressions on spoil heaps.
 K. Prach, P. Pyšek / Ecological Engineering 17 (2001) 55–62
Fig. 2. Changes in cover of woody species in the successional seres studied (16). Sum of covers of all woody species present in a given
year is shown. For the explanation of abbreviations see Fig. 1.
pact cover and did not permit woody species to
establish. Participation of woody species is usually
desirable in the restoration of previously forested
sites (Bradshaw and Chadwick, 1980; Jordan III
et al., 1987). Their establishment and subsequent
expansion can indicate gradual return of an ulti-
mately closed forest, typical of the temperate zone
(Ellenberg, 1988). The final physiognomy of spon-
taneous vegetation is usually determined by the
proportions between the graminoid and the
woody species. Spontaneous succession of woody
species may be expected in restoration projects at
sites where juvenile trees and shrubs are neither
limited physiologically nor by competition from
the herb layer, herbivory or other disturbances.
Some woody species recorded in this study (Acer
pseudoplatanus, Alnus glutinosa, Fraxinus excel-
sior, Pinus syl6estris, and Betula pendula) have
been used for artificial afforestation of disturbed
sites such as those created by large-scale mining
(S& týs, 1981).
Betula pendula took a prominent position
among the woody species recorded. Despite its
occurrence in a wide range of environmental con-
ditions (Ellenberg, 1988), its establishment early
in succession and its fast growth, Betula is mostly
59
not considered as a valuable amelioration species,
except for occassional cases where the species was
sown in areas deforested due to air pollution
(Pyšek, 1992). The only practical disadvantage of
this species is probably its possible role as a
source of allergic pollen (Kopecký, 1983).
In the present study, the period of 10 years
from the initial disturbance was used to define
‘‘later successional stages (i.e. pre-forest stages)’’.
Definitions of late successional species are gener-
ally relative and context-dependent (see Glenn-
Lewin et al., 1992). Most dominants listed in
Table 1 are important components of various
stabilised vegetation types in the Central Eu-
ropean landscape (Ellenberg, 1988). Their expan-
sion can be appreciated from the point of view of
restoration ecology. The only exceptions are
Artemisia 6ulgaris, Cirsium ar6ense, and Tanace-
tum 6ulgare, i.e. competitively strong dominants
of late successional herbal stages prior to the
establishment of scrubland (Pyšek, 1977, 1978;
Pyšek and Pyšek, 1991).
Competitive grasses and grass-like species pre-
vail among non-woody species of later succes-
sional stages. No alien species were present
among dominants, which can be considered as a
60 K. Prach, P. Pyšek / Ecological Engineering 17 (2001) 55–62

very positive fact from the viewpoint of restora- long-persisting swards of these species are not
tion. Surprisingly, the occurrence of alien species desirable from the point of view of restoration
in the sites studied was generally rather rare. practices.
Their representation was higher only in several The endangered and retreating species recorded
urban ruderal sites but none of them ever reached in the studied seres belonged to two categories: (i)
a high cover (Bastl et al., 1997) despite the fact rare ruderal species occurring in initial stages
that they represent an important part of the re- especially on spoil heaps and in urban sites (e.g.
gional flora (Pyšek et al., 1995). Atriplex rosea, Silene dichotoma, Kochia scoparia);
Especially the competitive grasses can arrest and (ii) competitively weak species of later succes-
succession for a long time by forming a dense, sional stages which grew either in nutrient-poor,
compact sward. In this respect, Calamagrostis wet habitats (dumps around the newly con-
epigejos plays a prominent role in the present structed pond, sand pit, depressions on spoil
Central European landscape. This species is ex- heaps) or in dry sites such as xeric abandoned
panding agressively in many disturbed sites. In oldfields. In the oldest xeric oldfields, several spe-
our data set, it appeared with a high cover cies typical of natural steppe-like surrounding
( \ 25%) in 6 seres out of 16. Calamagrostis 6illosa communities were found (e.g. Pulsatilla pratensis,
Aster amellus, Bupleurum falcatum). The occur-
exhibits a similar behaviour in the mountain ar-
rence of rare and endangered species in the later
eas, inhibiting both artificial and spontaneous af-
successional stages is desirable from the point of
forestation of sites deforested due to air pollution
view of restoration, similarly as the occurrence of
(Pyšek, 1992) and abandoned mountain grass-
all the species representing components of natu-
lands (Prach et al., 1996). Extensive and probably
ral, stabilised vegetation. The presence of woody
species typical of late successional vegetation, e.g.
Table 1
Twenty most common herbs (forbs and gramioids) occurring Pinus syl6estris, Acer pseudoplatanus, Alnus gluti-
in late successional stages and all woody species with maxi- nosa, Frangula alnus, Crataegus spec. div., Rosa
mum cover of at least 1% recorded in at least one successional canina, Acer campestre, is particularly encourag-
serea ing, because some of them belong to the species
pool of the climax vegetation of the region.
Forbs and graminoids S% Woody species S%
In some cases, the vegetation type which was
Calamagrostis epigejos 226 Betula pendula 145 present in the sites prior the disturbance was
Phragmites australis 219 Sambucus nigra 118 restored more or less directly. This happened
Arrhenatherum elatius 176 Crataegus sp. div. 101 when species typical of later stages established
Artemisia 6ulgaris 136 Pinus syl6estris 95 immediately after the initiation of succession. We
Calamagrostis 6illosa 123 Prunus spinosa 50
found this phenomenon in disturbed, nutrient-
Deschampsia flexuosa 118 Fraxinus excelsior 40
Urtica dioica 91 Salix caprea 39
poor sites surrounded by natural vegetation,
Phalaris arundinacea 88 Alnus glutinosa 30 whose environmental conditions were not much
Petasites hybridus 88 Frangula alnus 30 changed by the initial disturbance (sand pit,
Agropyron repens 85 Rosa canina agg. 13 dumps around the new pond and extracted peat-
Tanacetum 6ulgare 63 Populus tremula 9 lands). Similar situations may be observed in
Festuca rupicola 63 Acer pseudoplatanus 8
some stone quarries, on spoil heaps from small-
Poa angustifolia 49 Acer campestre 5
Galium album 43 Salix cinerea 3 scale mining, on forest road banks, and on depo-
Cirsium ar6ense 41 Populus nigra 1 sitions of mud bulldozed from water bodies. It
Juncus effusus 40 appears that the smaller the disturbed site, the
Holcus lanatus 39 easier and more direct succession towards vegeta-
Poa palustris 38 tion similar in species composition to natural
Coronilla 6aria 38
Molinia coerulea 37
regional communities will take place. However,
despite some limitations, spontaneous succession
a
Sums of maximum covers (S%) which the species attained can be viewed as an acceptable and cheap restora-
after 10 years of succession in 16 seres is shown. tion measure in other disturbed sites as well.
 K. Prach, P. Pyšek / Ecological Engineering 17 (2001) 55–62 61

Table 2 Acknowledgements
Summary of advantages and disadvantages of using sponta-
neous succession in restoration programmes in the temperate
zone of Europe
We thank two anonymous referees for their
comments on the manuscript and Marek Bastl for
Positives Negatives help with data analysis. The study was partly
supported by the Grant Agency of the Czech
Low costs Republic (projects no. 206/94/0395 and 206/97/
Usually fast (up to 15 years) Succession may be arrested
0077) and by the grant MSM 1231-00004.
formation of a continuous in early stage by expansion
vegetation cover (except of competitively strong forbs
of toxic substrata) or graminoids
Often expansion of native, Certain dominants can act References
well adapted woody as source of allergenic
species pollen Banásová, V., 1976. Vegetácia medených a antimonových
Usually low cover of invasive species háld. Biol. Práce, Bratislava 22, 1 – 109 (in Slovak with
Often refugia for wildlife Occasional seed sources of German summary).
weeds Bastl, M., Kočár, P., Prach, K., Pyšek, P., 1997. The effect of
Usually higher natural value Usually lower productive successional age and disturbance on the establishment of
in comparison with value in comparison with alien plants in man-made sites: an experimental approach.
technically restored sites technically restored sites In: Brock, J.H., Wade, M., Pyšek, P., Green, D. (Eds.),
Plant Invasions: Studies from North America and Europe.
Backhuys Publishers, Leiden, pp. 191 – 201.
Bradshaw, A., 1997. Restoration of mined lands — using
natural processes. Ecol. Engin. 8, 255 – 269.
Bradshaw, A., Chadwick, M.J., 1980. The Restoration of
Land. The Ecology and Reclamation of Derelict and De-
graded Land. University of California Press, Berkeley.
5. Conclusions De Steven, D., 1991. Experiments on mechanisms of tree
establishment in old-field succession: seedling emergence.
Results presented here and other experience Ecology 72, 1066 – 1075.
Ellenberg, H., 1988. Vegetation Ecology of Central Europe.
from various successional habitats enabled us to Cambridge University Press, Cambridge.
tentatively summarise the advantages and disad- Glenn-Lewin, D.C., Peet, R.K., Veblen, T.T., 1992. Plant
vantages of using spontaneous succession in succession. In: Theory and Prediction. Chapmann and
restoration programmes in the temperate zone of Hall, London.
Jordan III, W.R., Gilpin, M.E., Aber, J.D., 1987. Restoration
Europe (Table 2). The positive effects prevail Ecology. In: A Synthetic Approach to Ecological Re-
especially if the disturbed site is small, surrounded search. Cambridge University Press, Cambridge.
by natural vegetation, and site conditions were Kopecký, K., 1983. Remarks on the changes of vegetation in
relation to allergic respiratory diseases — a case study
not principally altered by the initial disturbance.
from SW Prague. Preslia, 55, 149 – 163 (in Czech with
Among the venues of future research should be (i) English and German summaries).
a comparison of spontaneous and controlled Luken, J.O., 1990. Directing Ecological Succession. Chapman
revegetation; and (ii) course of succession in more and Hall, London.
Olsson, G., 1987. Effects of dispersal mechanisms on the initial
extreme environments. pattern of old-field succession. Acta Oecologica 8, 379 –
Information on spontaneous succession, based 390.
on both exact quantitative data and field experi- Osbornová, J., Kovářová, M., Lepš, J., Prach K., (Eds.), 1990.
ence, was summarised into an expert system Succession in Abandoned Fields. Studies in Central Bo-
hemia, Czechoslovakia. Kluwer Academic Publishers,
named SUCCESS. This tool enabled a tentative Dordrecht.
prediction of the rates and directions of succes- Prach, K., 1987. Succession of vegetation on dumps from strip
sion in various Central European disturbed habi- coal mining, NW Bohemia. Czechoslovakia. Folia Geobot.
Phytotax. 22, 339 – 354.
tats up to 50 years since the initial disturbance
Prach, K., Pyšek, P., 1994a. Spontaneous establishment of
(Prach et al., 1999). This expert system can be woody plants in Central European derelict sites and their
exploited in restoration projects. potential for reclamation. Restor. Ecol. 2, 190 – 197.
62 K. Prach, P. Pyšek / Ecological Engineering 17 (2001) 55–62
Prach, K., Pyšek, P., 1994b. Clonal plants — what is their role
in succession? Folia Geobot. Phytotax. 29, 307–320.
Prach, K., Pyšek, P., 1999. How do species dominating in
succession differ from the others? J. Veget. Sci. 10, 383–
392.
Prach, K., Pyšek, P., S& milauer, P., 1993. On the rate of
succession. Oikos 66, 343–346.
Prach, K., Lepš, J., Michálek, J., 1996. Establishment of
Norway spruce seedlings in a Central European mountain
grassland: an experimental study. J. Veget. Sci. 7, 681–684.
Prach, K., Pyšek, P., S& milauer, P., 1997. Changes in species
traits during succession: a search for pattern. Oikos 79,
201 – 205.
Prach, K., Pyšek, P., S& milauer, P., 1999. Prediction of vegeta-
tion succession in human-disturbed habitats using an ex-
pert system. Restor. Ecol. 7, 15–23.
Pyšek, A., 1977. Sukzession der Ruderalpflanzengesellschaften
von Gross-Plzen. Preslia 49, 161–179.
Pyšek, A., 1978. Ruderal vegetation of the city of Plzeň. Diss.,
Institute of Botany, Průhonice, (in Czech).
Pyšek, P., 1992. Dominant species exchange during succession
in reclaimed habitats: a case study from areas deforested
due to air pollution. For. Ecol. Manage. 54, 27 – 44.
Pyšek, P., Prach, K., S& milauer, P., 1995. Invasion success
related to plant traits: an analysis of Czech alien flora. In:
Pyšek, P., Prach, K., Rejmánek, M., Wade, P. (Eds.), Plant
Invasions: General Aspects and Special Problems. SPB
Academic Publishing, Amsterdam, pp. 39 – 60.
Pyšek, P., Pyšek, A., 1991. Succession in urban habitats: an
analysis of phytosociological data. Preslia 63, 125 – 128.
Rebele, F., Dettmar, J., 1996. Industriebrachen. O8 kologie und
Management, E. Ulmer Verlag, Stuttgart.
Rothmaler, W., et al., 1982. Exkursionsflora. Band 4. Volk
und Wissen Volkseigener Verlag, Berlin.
S& týs, S., 1981. Reclamation of Areas Disturbed by Mining of
Raw Materials. STN, Prague (in Czech).