POLISH JOURNAL OF ECOLOGY
(Pol. J. Ecol.)
61
1
13–22
2013
Regular research paper
Agnieszka KOMPAŁA-BĄBA1*, Wojciech BĄBA2
1
Department of Geobotany and Nature Protection, Silesian University, Jagiellońska 28,
40-032 Katowice, Poland, *e-mail: [email protected] (corresponding author)
2
Department of Plant Ecology, Institute of Botany, Jagiellonian University, Lubicz 46
31-512 Kraków, Poland
THE SPONTANEOUS SUCCESSION IN A SAND-PIT –
THE ROLE OF LIFE HISTORY TRAITS AND SPECIES HABITAT
PREFERENCES
ABSTRACT: The prediction of species response to human activity is of great interest in
contemporary restoration ecology. The purpose
of the article was to analyse which species life history traits and species habitat preferences are important during succession after the abandonment
of mining activity in a sand-pit. During a 15-year
period (1996–2010), 176 phytosociological relevés were placed within vegetation patches of different ages and divided according to soil moisture,
thus forming two series of chronosequences that
ranged from 0 to ca. 50 years. The datasets were
analysed using both DCA/CCA ordinations and
regression trees. The successional gradient, apart
from the theoretically predicted replacement R by
C strategists, revealed the occurrence of stresstolerant ruderals and competitive ruderals in the
early successional stages. However, differences
were observed between the wet and dry series. On
dry soils the ruderals, anemochorous and windpollinated species dominated in the early phases.
In the middle successional phases, a preponderance of anemochorous species (nanophanerophytes, light or semi-shade demanding species),
nitrogen-poor and competitive ruderals or species
typical for nitrogen-rich soils was recorded. In
the late successional stages, species with both the
ability of vegetative and generative reproduction
appeared. On the other hand, on wet sites in the
early successional phases, species with vegetative
growth, hydrophytes, chamaephytes and stresscompetitors prevailed. Later in the succession,
journal 33.indb 13
they were replaced by insect-pollinated species in
nitrogen rich habitats and stress-tolerant ruderals
on less fertile habitats. Finally, competitors started
to prevail. Knowledge of the environmental conditions of a given site, the ecological processes and
species biology can assist in achieving the desired
goals or in initiating or enhancing succession on
some disturbed sites.
KEY WORDS: life history traits, habitat preferences, succession, ordination, sand-pits, regression trees
1. INTRODUCTION
Man-made disturbances caused by the
mining industry lead to drastic changes in
both the abiotic and biotic elements of a given
area and often results in the total destruction
or depletion of vegetation or animal communities, thus their restoration is of prime interest
to ecologists and conservationists. However, in
spite of the accumulation of knowledge on successional processes (Fa lińska 2003, Hobbs
et al. 2007, Pickett et al. 2005, 2008, Wa l ker et al. 2010, Prach and Wa l ker 2011),
the ecological restoration of vegetation on
post-industrial areas is still a challenging task
(Ho d ačová and Prach 2003, B ar t ha et al.
2004, Ř ehoun ková and Prach 2006, 2008,
2013-04-30 10:04:53
14
Agnieszka Kompała-Bąba, Wojciech Bąba
Prach and Hobbs 2008, Prach and Wa l ker 2011). On the other hand, the spontaneous
colonisation of species provides the opportunity to follow the changes in species composition and structure through time and in space.
The colonisation of a given area can be
connected with many factors which reduce
the initial, potential species pool via biotic and
abiotic filters: climatic conditions, availability and diversity of local site conditions (the
amount of available resources, light, water
holding capacity, granulometric composition,
soil reaction and moisture), propagule availability and interactions with other species.
The dispersal mode (both long-distance
and small-scale dispersability) to open sites
can also limit species-richness (Z ob el 1997,
Pos ch lo d et al. 2005). Communities which
are created during colonisation can be spatially heterogeneous. Such spatial heterogeneity is the result of a rich variety of the kinds
and intensities of disturbances that affect
colonisation and the surrounding landscape.
Species possess a suite of abilities that enable
them to exploit available resources, grow,
persist and reproduce (B azzaz 1996). Life
history traits, relative growth rate (RGR), life
span, stress tolerance, herbivory and predator defence are some of the specific features
that determine differential performance
(Vitouš ek et al. 1998). A lack of the necessary traits can mean that species cannot pass
through some environmental filters of changing environmental conditions and get to given sites in spite of being present in the local
species pool. Knowledge about the life history
traits of species and their habitat preferences,
which are responsible for their colonisation
success, seems to be very helpful in preparing
successful restoration projects on a variety of
wastelands (van Andel and Arons on 2006,
Ř ehoun ková and Prach 2008, 2010, Trop ek et al. 2010, Prach and Wa l ker 2011).
The purpose of the article was to analyse:
• what is the pattern of species life history traits along a successional gradient in a sand-pit,
• what are the habitat preferences of a
species during succession,
• whether such patterns are similar to
those observed in other types of postindustrial wastelands (e.g. gravel sandpits, abandoned quarries, spoil heaps).
journal 33.indb 14
2. STUDY AREA
The study was performed in 1996–2010
in the “Kuźnica Warężyńska” sand-pit, which
is situated in the southern part of Poland
[50°21’–50°24’N, 19°11’–19°13’E]. It comprises an area of 870 ha. Most of the soils
are loose sands with an acidic soil reaction
(pHH2O = 5.0–6.0), a low capacity of the sorption complex, a low retention capacity, poor
in nutrients and susceptible to drying. Other groups of sands that were deposited here
mainly contained a calcareous gravel admixture and locally silty-argillaceous inter-layers.
Before excavation of sand the level of the
ground water table extended from 1 to 6.5 m.
It was oligotrophic or mesotrophic and contained Ca, Mg ions (Krz a k l e w sk i 1999).
The history of the sand-pit was reconstructed on the basis of direct field observations and the records of mining company,
including the forest valuations. The mining
activity in the area started in 1967 and was
conducted until 2005. Because the excavation
of the sand was not carried out simultaneously in the whole area, patches which differ
in reference to the time of abandonment were
created. The abandoned area had not been reclaimed, so spontaneous vegetation could develop as a result of succession. A preliminary
analysis of the vegetation data (not shown)
revealed a strong differentiation of species
composition according to the soil moisture
gradient, thus in the further analysis, we considered two series of sample plots on dry and
wet soils. Each of them was divided into successional stages taking into account: (i) the
year of abandonment and (ii) the presence of
spontaneous vegetation.
Three successional stages were distinguished based on arbitrary age classes: young
(1–10 years), middle (11–25 years) and late
(>25 years). Based on these classes, we assessed the differences in the species traits of
early, middle and late successional species.
3. MATERIALS AND METHODS
In total 176 phytosociological relevés
(5 m ´ 5 m) were recorded in the field using
the Braun-Bl anquet (1964) approach. Basic life history traits (strategy type, life forms,
type of reproduction, self-sterility, pollen vec-
2013-04-30 10:04:53
The spontaneous succession in a sand-pit
tor, dispersal and diaspore weight) and habitat preferences (Ellenberg indicator values for
light (L), moisture (M), fertility (N) and soil
reaction (R) (E l lenb erg et al. 1991; explanations in Table 1) were considered. Life history
traits were compiled from existing databases
such as: Diasporus (B onn et al. 2000), Gr ime
et al. (1988), BIOPOP (Jackel et al. 2006), BIOLFLOR (Fran k and K lotz 1990, K lotz
et al. 2002), PHANART (L ind acher 1995).
Because species can belong to several classes
of particular trait and habitat preferences in
order to include complete information, fuzzy
coding was used (L epš and Šmi l auer 2003).
To evaluate which species traits can be important during a particular stage of the succession
of the sand pit, the ordination (CANOCO 4.5,
ter Braa k and Šmi l auer 2002) and regression tree methods (R package, R Development
Core Team, 2011) were performed. A preliminary analysis of species data (not shown)
revealed that gradient length exceeded 4 SD,
thus the unimodal relationship between the
species characteristics and time was considered and DCA and CCA ordinations were
used. The CCA was carried out using MOISTURE (which divided the sites according to
their Ellenberg indicator values) as only one
explanatory variable. The scores of the species
on the ordination axis were then used as the
response to the age of site. Next, Detrended
correspondence analysis (DCA) with species
traits and habitat preferences as a “species
data” was performed. The explanatory “environmental variables”: species response to the
MOISTURE and the age of site (AGE) were fitted ex post as supplementary passive environmental variables (L epš and Šmi l auer 2003).
Regression trees were used to determine
which species with what traits and habitat
characteristics occurred in a given successional stage on “dry” and “wet” series. The
length of the trees was controlled by the best
trade-off between the size of the tree and explained deviance via pruning, an analogue of
variable selection in regression (Brei man et
al. 1984, Thui l ler et al. 2003).
4. RESULTS
Overall, 321 vascular plant species were
found – 229 on dry and 253 on wet series. The
number of species on both series decreased
journal 33.indb 15
15
Fig. 1. DCA ordination performed on life history
traits (life forms, strategy, pollen vector, dispersal mode, reproduction, self-sterility) and habitat
preferences of species (light, moisture, nitrogen,
soil reaction/base saturation).
Abbreviations: life form (Th – therophyte, Ch –
chamaephyte, G – geophyte, Hy – hydrophyte, Na
– nanophanerophyte, Me – megaphanerophyte, H
– hemicryptophyte); C, R, SR, CR, CS, S, CSR –
strategy type; reproduction (Sv – mostly seeds, sv
– seed and vegetative, Vs – mostly vegetative); pollen vector (wind, ins - insect); dispersal type (anemo – anemochory, epizoo – epizoochory, endoz
– endozoochory, autoc – autochory), self-sterility
(S-C – self-compatible, S-I – self-incompatible);
Ellenberg Indicators Values: soil reaction (acid
1–5, neu - neutral 6–7, basic 8–9; light (shade 1–3,
semsha - semi-shade 4–6, L - light 7–9), moisture
(dry 1–4, mesic 5–8, wet 9–11), nitrogen (poor
1–3, intermediate 4–6, rich 7–9).
compared to the young and late stages: from
152 to 96 on the “dry” and from 165 to 126 on
the “wet” series.
The DCA analysis indicated the most important life history traits and habitat preferences of species in the course of succession
(Fig. 1). The eigenvalues of the first and second axis were 0.68 and 0.35, respectively. The
variable AGE represents the gradient, which
disregarding the differences between the
dry and wet series, is mostly in agreement
with the theoretical pattern of successional
replacement. On the young sites, a preponderance of therophytes, R (R, CR, SR) and
CSR, anemochorous, light-demanding and
self-compatible species typical for ruderal
and segetal communities was observed. Later
in the succession, they were replaced by the
2013-04-30 10:04:53
16
Agnieszka Kompała-Bąba, Wojciech Bąba
Fig. 2. Regression tree analysis used for predicting colonisation success in dry (A) and wet series (B) in
the “Kuźnica Warężyńska” sand-pit performed on life history traits (life form, strategy, pollen vector,
dispersal mode, reproduction, self-sterility, diaspora weight) and habitat preferences of species.
Abbreviations: C, R, SR, CR, CS – strategy type; life form (Ch – chamaephyte, Hy – hydrophyte, Na
– nanophanerophyte), Ellenberg Indicators Values: nitrogen (poorN 1–3, intN 4–6, richN 7–9); reproduction (Sv – mostly seeds, sv – seed and vegetative, Vs – mostly vegetative), pollen vector (wind,
insect); dispersal type (ane – anemochory, endoz – endozoochory, aut – autochory), diaspora weight
(LightD – light diaspore, HeavyD – heavy diaspore)
Achi mill – Achillea millefolium, Agro capi – Agrostis capillaris, Alis pla – Alisma plantago-aquatica, Anth
vuln – Anthylis vulneraria, Aren serp – Arenaria serpyllifolia, Arte camp – Artemisia campestris, Betu pend
– Betula pendula, Bide fron – Bidens frondosa, Cala epig – Calamagrostis epigejos, Call vulg – Calluna
vulgaris, Care acut – Carex acutiformis, Care flav – Carex flava, Care oede – Carex oederi, Cent umbe –
Centaurium umbellatum, Cori lept – Corispermum leptopterum, Cory cane – Corynephorus canescens, Coro
vari – Coronilla varia, Crep bien – Crepis biennis, Dauc caro – Daucus carota, Desc case - Deschampsia caespitosa, Desc flex – Deschampsia flexuosa, Digi isch – Digitaria ischaemum, Dros rotu – Drosera rotundifolia,
Eleo palu – Eleocharis palustris, Epip hell - Epip helleborine, Equi vari – Equisetum variegatum, Erig acri
– Erigeron acris, Eupa cann – Eupatorium cannabinum, Fest ovin – Festuca ovina, Fest rubr – Festuca rubra,
Heli numm – Helianthemum nummularium, Hern glab- Herniarnia glabra, Hiera pilo – Hieracium pilosella,
Holc lana – Holcus lanatus, Isol seta – Isolepis setacea, Jasi mont – Jasione montana, Junc arti – Juncus articulatus, Junc bufo – Juncus bufonius, Junc effu – Junc effussus, Koel glau – Koeleria glauca, Lotu corn – Lotus corniculatus, Lyco clav – Lycopodium clavatum, Lyco inun – Lycopodiella inundata, Lyco euro – Lycopus
europaeus, Lysi vulg – Lysimachia vulgaris, Lyth sali – Lythrum salicaria, Meli alba – Melilotus alba, Medi
journal 33.indb 16
2013-04-30 10:04:57
The spontaneous succession in a sand-pit
hemicryptophyte, epi- and endozoochorous, semi-shadow, wind-pollinated species
and finally by C-strategists, mega- and nanophanerophytes. Moreover, the differences between the dry and wet series were detected.
On the dry sites, the species typical of acid,
nutrient-poor sites, chamaephyte, stresstolerant, self-incompatible species prevailed,
while the wetter sites were dominated by
stress-competitors and species that prefer
nutrient-rich and neutral soils with predominant clonal growth.
The regression trees analyses revealed a
more detailed, hierarchical relationship between the life history traits and habitat preferences. The species in the early successional
stages on dry soils were ruderal, mostly anemochorous, wind pollinated and if not anemochorous then confined to nutrient-poor
habitats, if not nutrient poor then endozoochorous (Fig. 2A). In the middle successional phases (11–25 years), a preponderance
of anemochorous species was observed, both
nanophanerophytes, light or semi-shade demanding species. If not anemochorous, then
nitrogen-poor and competitive ruderals or
species that are typical of nitrogen-rich soils
and chamaephytes (Fig. 2A). In the late successional stages, the main difference on dry
sites was the presence of species with the ability to both vegetative and generative reproduction.
On the other hand, species which colonised the young wet sites were mainly clonal
plants, chamaephyte, hydrophyte or competitive stress-tolerators. If they reproduce mainly by seed, they have either heavy or light
diaspores (Fig. 2B). Later in the succession,
they were replaced by insect-pollinated species that prefer habitats richer in nitrogen or
stress-tolerant ruderals on soils with a lower
amount of nitrogen and finally with species
having a strong competitive ability.
17
5. DISCUSSION AND CONCLUSIONS
Vegetation changes during succession
in abandoned sand-pits are associated with
both habitat conditions (light, soil moisture,
nitrogen content, soil reaction) and life history traits, which enable species to colonise,
coexist, compete or regenerate after some
disturbances (Gr i me 2002, P y wel l et al.
2003, Frou z et al. 2008, Ř ehou n kov á and
Pr a ch 2010).
The results of our research confirmed,
given in other papers, the significant role of
moisture as a main environmental factor that
is responsible for the development, growth
and a large amount of vegetation variability
of vegetation, as well as the increase of wet
grasslands and woods (F i e r ro et al. 1999,
Ř ehou n kov á and Pr a ch 2006).
The general pattern of growth forms
during succession in a post-mining landscape from therophytes to hemicryptophytes
and phanerophytes observed in this study
(shows by the DCA; Fig. 1) was reported in
previous works (Brow n 1991, Huston and
Sm it h 1987, Pr a ch et al. 1997). However,
Ř ehou n kov á and Pr a ch (2010) found
that in the early stages of the colonisation of
abandoned gravel-sand pits chamaephytes
were more successful in comparison to therophytes. According to Woź n i a k (2010) the
significant participation of chamaephytes on
young spoil heaps was probably connected
with the presence of Chamaenerion palustre with a higher frequency and abundance.
Moreover, undemanding hemicryptophytes
and chamaephytes were also recorded during
the first stage of succession on the moraine of
the Aletsch glacier, which was explained by
harsh habitat conditions (nutrient poor, acidic soils) (S chu lz e et al. 2002). Wi e g l eb and
Fel i n ks (2001) found that therophytes were
only of minor importance in a post-mining
landscape, even in pioneer stands in con-
Fig. 2. cont.
falc – Medicago falcata, Moli caer – Molinia caerulea, Oeno bien – Oenothera biennis, Orth secu – Orthilia
secunda, Phra aust – Phragmites australis, Pinu sylv – Pinus sylvestris, Plan lanc – Plantago lanceolata, Popu
trem – Populus tremula, Pota pect – Potamogeton pectinatus, Rume aceto – Rumex acetosella, Sagi nodo –
Sagina nodosa, Sali capr – Salix caprea, Sali purp – Salix purpurea, S. repe – Salix repens, Sali rosm – Salix
rosmarinifolia, Sang mino – Sanguisorba minor, Scle pere – Scleranthus perennis, Seta viri – Setaria viridis,
Sisy alti – Sisymbrium altissimum, Soli cana – Solidago canadensis, Soli gram – Solidago graminifolia, Thym
serp – Thymus serpyllum, Thym pule – Thymus pulegioides, Trig palu – Triglochin palustre, Tuss farf – Tussilago farfara, Typh angu – Typha angustifolia, Typh laxm – Typha laxmanii, Verb lych – Verbascum lychnitis.
journal 33.indb 17
2013-04-30 10:04:58
18
Agnieszka Kompała-Bąba, Wojciech Bąba
trast to hemicryptophytes, which dominate
on all study sites. Woźni a k (2010) recorded
a high participation of therophytes on the
10–30 years old spoil heaps but hemicryptophytes were the most frequent dominants
there. Prach et al. (1994) examined changes
in species traits during succession on urban,
post-industrial wastelands and detected that
hemicryptophytes and geophytes did not
exhibit any clear trend in the data set. Other research showed that replacement from
terophytes to hemicryptophytes during succession takes place only on poor quality substrates (broken waste (<15 mm – by product
of uranium extraction), whereas on better
quality substrate, no clear trend in life forms
was detected (Mar t ine z-Ruiz and Mar rs
2007).
In relation to the dispersal mode, anemochory and zoochory were the most frequent
dispersal mechanisms on the studied sandpit even in the early stages of succession on
the dry series, which is similar to research
conducted on uranium mine wastes (Mart ine z -Ruiz and Mar rs 2007). However,
Wieg leb and Felin ks (2001) did not detect
any change in the predominance of individual
dispersal mechanisms in post-mining landscapes. In contrast, on wet sites, zoochorous
species started to play an important part in
later stages of succession. Hydrophytes could
extend their colonisation range regardless of
successional age, because as a result of mining activity, the area of the sand-pit was covered with drainage ditches, whose numbers
and sizes were constantly changing. K hate r
et al. (2003) discovered that limestone quar­
ries were dominated by annuals or opportunistic chamaephytes that spread mainly via
antropochory, which probably enables them
to disperse in large quantities and to establish themselves in open sites successfully.
Ř ehoun ková and Prach (2010) found that
in the case of succession in gravel sand-pits,
a kind of replacement between the types of
dispersal could not be found because anemochory, zoochory and hydrochory appeared to
be more or less neutral to succession.
L atzel et al. (2011) examined ruderal
urban sites and spoil heaps found that seed
dispersal by water was the most important
mode of dispersal at the beginning of succession even in areas distant from wetlands, and
journal 33.indb 18
that the prevalence of anemochory increased
slightly during succession.
Our results, similar to those obtained
in other types of wastelands (quarries, mine
spoils), indicate that ruderal species with
broad ecological amplitudes can prevail in the
initial seral stages on dry sites, whereas more
specialised wetland species occur on the wet
sites and canals crossing the area of a sandpit (B orgegård 1990, Ř ehoun ková and
Prach 2006). Woźni a k (2010) found that
species with an S, CR strategy occur more frequently on heaps where spoil substrates were
stored irregularly, whereas the number of species with S, CR was comparable regardless of
the method of storage. P y wel l et al. 2003
found that short-lived, early flowering ruderals with a heavy allocation to reproduction
mainly colonised open, undisturbed habitats.
In contrast, Wieg leb and Felin ks (2001)
found that competitors and stress-tolerant
competitors predominate in the initial phases during primary succession, or if not, then
stress-tolerant ruderals and C-S-R strategists
prevail in vegetation types. Research conducted in a gravel-sand-pit showed that species
with an S strategy had the highest colonisation
success (B orgegård 1990, Ř ehoun ková
and Prach 2006). D öl le et al. (2008) exami�
ned changes in life history trait composition
during undisturbed old-field succession and
found that the annual stage was dominated by
ruderals and competitive species. The ruderals showed a significant decrease during succession and have no relevance in the pioneerforest stage. In contrast, competitive species
showed a clear increasing rate from almost
45 to 95%. R-strategists (mainly annuals)
constitute a major proportion of species that
were recorded after profound disturbances in
quarries. Whereas their number and cover decreased during colonisation, myrmecochorous and barochorous native species, mainly
clonally spreading perennial species or shrub,
appeared (S or S-R strategists) (K hater et al.
2003).
A set of specific traits referring to competition such as: C-strategy, high stature, ability
to spread laterally, higher demand for moisture and nutrients enables some species such
as: Calamagrostis epigejos, Solidago canadensis, Carex acutiformis, Phragmites australis to
become the dominants during the course of
2013-04-30 10:04:58
The spontaneous succession in a sand-pit
succession and means that those species are
better suited to the environment than minor
species (Prach and P yš ek 1999, Br z o sko
2000, L atzel et al. 2011). Species such as
Carex acutiformis, C. rostrata can occupy a
vast area in a relatively short period of time
because they form new shoots which grow
exponentially (Brzosko 2000, Pr a ch and
P yš ek 2001). Some results show that soil
moisture appears to be a factor significantly supporting the success of clonal species
(Prach and P yš ek 1994, 2001). Solidago
graminifolia started to spread in the sand-pit
area in sites, where the level of the ground
water table was high and as a result of a disturbance caused by soil disintegration and
the transport of rhizomes (Komp a ł a - B ąb a
and B ąb a 2006).
The type of reproduction was particularly
important in the early stages of succession in
a sand-pit on a wet series. Some species that
reproduced mostly by seeds (Centaurium erythraea, Sagina nodosa) occupied a small area
of bare sand, sometimes close to drainage
ditches. Some rare species such as Lycopodiella inundata, Lycopodium clavatum or shrubs
- Salix rosmarinifolia, propagate vegetatively.
The early stage of succession in undisturbed
old-fields was dominated by species which
reproduce exclusively or mostly by seeds,
which participation decreased significantly
over time. Species that reproduce both by
seeds and vegetatively in equal amount show
a slight increase as do species that rarely reproduce in a vegetative manner or mainly by
seeds. Mainly vegetative reproduction was of
no importance at any time of the successional
stages (D öl le et al. 2008).
According to L atzel et al. (2011), dispersal and persistence traits (clonality, vegetative
regeneration) could along with other factors
(regional species pool, habitat conditions),
be important determinants of the changes of
vegetation that occur during succession.
Scientific research conducted in quarries
and sand-pits may provide valuable habitats
(wetlands, peat-bogs) for many rare and endangered species and act as local centres of
floristic diversity in a given landscape (G e m mel l 1982, Prach and P yš ek 2001, Nov á k
and Prach 2003, Nová k and Konv i čk a
2006, Bzdon 2008, Komp ała -B ąb a and
B ąb a 2009, Trop ek et al. 2010, Tr n kov á
journal 33.indb 19
19
et al. 2010, Ř ehou n kov á and Ř ehounek
2011). Disturbances such as the removal of
accumulating litter, mechanical disturbances
of wet sand, lightly scored patches, periodic
grazing that constantly lead to the creation
of microhabitats may enable populations of
rare species to be saved and protected, among
them Lycopodiella inundata, Liparis loeselii
(Whe eler et al. 1998, Ř ehou n kov á and
Ř ehou nek 2011). The last species possesses
many traits such as: rapid regeneration from
seeds and spores, high rates of turnover of individual plants, short-lived plants (2–3) and
can grow on sites with low fertility but with a
relatively high and stable ground water table
(Jone s 2008, Whe el er et al. 1998). However, as a result of the cessation of frequent
disturbances caused by sand exploitation, a
thick moss layer was formed on some sites,
which has an influence on the Liparis loeselli
population and more expansive grass species
(Phragmites australis) started to encroach
(Komp a ła -B ąb a and B ąb a 2009).
As was already stated, knowledge of the
environmental conditions of a given site, on
ecological processes (dispersal, competition)
and species biology (life history traits) and
the functioning of an ecosystem (e.g. nutrient cycle) together with spontaneous succession can help in achieving the desired goals
or in initiating and enhancing the succession process in some disturbed sites through
the choice of appropriate species for restoration purposes (B ąb a et al. 2012, B ąb a ,
Komp ał a-B ąb a 2005, Prach et al. 2001,
Wi eg leb and Feli n ks 2001, K hater et al.
2003, Pr a ch and Wa l ke r 2011). In comparison to technical restoration, ecological restoration enables a mosaic of various habitats
with prevalence of native species and lesser
cost of restoration. As a result, the floristic
(measured as the number of species, the prevalence of native species in the flora) and the
vegetation diversity of a given area increases.
Spontaneous and carefully manipulated succession has a great influence for effectively
restoring damaged ecosystems (Ho d a č ov á
and Pr a ch 2003, Pr a ch et al. 2001, 2007,
Prach and Hobbs 2008, Prach and Wa l k e r 2011).
ACKNOWLEDGEMENTS: We would like
to thank the anonymous reviewers for valuable
2013-04-30 10:04:58
20
Agnieszka Kompała-Bąba, Wojciech Bąba
comments on the manuscript, and Michele Simmons for proofing English version of the text.
6. REFERENCES
B ar t ha S., C amp etel l a G., C anu l lo R .,
B o dis J., Mucina L. 2004 – On the importance of fine-scale spatial complexity in
vegetation restoration studies – Intern. J. Ecol.
Env. Sci. 30: 101–116.
B azazz F.A. 1996 – Plants in changing environments: linking physiological, population, and
community ecology – Cambridge University
Press, New York, 320 pp.
B ąb a W., Komp ała-B ąb a A. 2005 – Do smallscale gaps in calcareous grassland swards facilitate seedling establishment? – Acta Soc. Bot.
Pol. 74: 125–131.
B ąb a W., Kurowska M., Komp ała-B ąb a
A., Wi lczek A., Długosz J., Szarejko
I. 2012 – Genetic diversity of the expansive
grass Brachypodium pinnatum in a changing
landscape: effect of habitat age – Flora, 207:
346–353.
B onn S., Pos ch lo d P., Tackenb erg O. 2000
– Diasporus. A database for diaspore dispersal
– concept and application in case studies for
risk assessment – Zeitschrift für Ökologie und
Naturschutz 9: 85–97.
B orgegård S.-O. 1990 – Vegetation development in abandoned gravel pits: effects – J. Veg.
Sci. 1: 672–682.
Braun-Bl anquet J. 1964 – Pflanzensoziologie
[Phytosociology] – SpringerVerlag, Wien, 865
pp.
Breiman L., Fr ie dman J., Oh ls e n R .,
Stone C. 1984 – Classification and regression trees – Wadsworth, Belmont, CA, 358 pp.
Brow n V.K. 1991 – The effects of changes in
habitat structure during succession in terrestrial communities – (In: Habitat structure
the physical arrangement of objects in space,
Eds: S.S. Bell, E.D. McCoy, H.R. Mushinsky) –
Chapman & Hall, London, pp. 141–168.
Brzosko E. 2000 – Zmiany liczebności populacji
roślin o różnych strategiach reprodukcyjnych w
procesie sukcesji [Changes of population numbers of plant with different reproduction strategies in the process of succession] – Wiad. Bot.
44 (3/4): 13–22 (in Polish with English summary).
Bzdon G. 2008 – Gravel pits as habitat islands:
floristic diversity and vegetation analysis – Pol.
J. Ecol. 56: 239–250.
D öl le M., B er n hardt-R omer mann M.,
Par t h A., S chmidt W. 2008 – Changes
in life history trait composition during un-
journal 33.indb 20
disturbed old-field succession – Flora, 203:
508–522.
E l lenb erg H., Web er H.E., D ü l l R ., Wir t h
V., Wer ner W., Pau liss en D. 1991 –
Zeigerwerte von Pflanzen in Mitteleuropa –
Scripta Geobot. 18: 1–248.
Fa lińska K. 2003 – Alternative pathways of succession: species turnover patterns in meadows
abandoned for 30 years – Phytocoenosis, 15
(N.S.) Arch. Geobot. 9: 1–104.
Fier ro A., Angers D.A., B e auchamp C.J.
1999 – Restoration of ecosystem function in
an abandoned sandpit: plant and soil responses to paper de-inking sludge – J. Appl. Ecol.
36: 244–253.
Fran k D., K lotz S. (e ds.) 1990 – Biologischökologische Daten zue Flora der DDR. 2nd ed,
Wissenschftliche Beitrage Martin Luther Universitat 32 – Halle, Wittenberg.
Frouz J., Prach K., Pižl V., Háněl L.,
St ar ý J., Taj ovský K, Mater na J., B a lí k
V., Ka lčí k J., Ř ehoun ková K. 2008 – Interactions between soil development, vegetation and soil fauna during spontaneous succession in post mining sites – Eur. J. Soil Biol.
44: 109–121.
G emmel l R .P. 1982 – The origin and botanical
importance of industrial habitats – (In: Urban
Ecology, Eds: R. Borkmann, J.A. Lee, M.R.D.
Seaward) – Blackwell Scientific Publications,
Oxford, London, Edinburgh, Boston, Melbourne, pp. 33–39.
Gr ime J. P. 2002 – Plant strategies, vegetation
processes, and ecosystem properties – John Wiley and Sons Ltd, Chichester, England, 417 pp.
Gr ime J.P., Ho dgs on J.G., Hunt R . 1988
– Comparative plant ecology. A functional approach to common British species – Unwin
Hyman Ltd., London, 748 pp.
Hobbs R .J., Wa l ker L.R ., Wa l ker J. 2007
– Integrating restoration and succession – (In:
Linking restoration and ecological succession,
Eds: L.R., Walker, J.R. Walker, Hobbs R .J.) –
Springer, pp. 168–179.
Ho d ačová D., Prach K. 2003 – Spoil heaps
from brown coal mining: technical reclamation versus spontaneous revegetation – Restor.
Ecol. 11: 385–391.
Huston M., Smit h T.M. 1987 – Plant succession: Life history and competition – Am. Nat.
130: 168–198.
Jackel A.K., D annemann A., Tackenb erg
O., K le yer M., Pos ch lo d P. 2006 – BIOPOP: BioPop-funktionelle Merkmale von
Pflanzen und deren Anwendungsmöglichkeiten im Arten, Biotop- und Naturschutz
– Naturschutz und Biologische Vielfalt, 32:
1–168.
2013-04-30 10:04:58
The spontaneous succession in a sand-pit
Jones P. S. 1998 – Aspect of the population biology of Liparis loeselii (L.) Rich. var ovata Ridd.
Ex Godfery (Orchidaceae) in the dune slacks
of South Wales, UK – Bot. J. Linn. Soc. 126:
123–139.
K hater C., Ar naud M., Mai l let J. 2003
– Spontaneous vegetation dynamics and restoration prospects for limestone quarries in
Lebanon – Appl. Veg. Sci. 6: 199–204.
K lotz S., Kü hn I., D urka W. 2002 – BIOLFLOR – Eine Datenbank mit biologisch-ökologischen Merkmalen zur Flora von Deutschland. Schriftenreihe für Vegetationskunde 38:
1–334 – Bundesamt fur Naturschutz, BonnBad Godesberg.
Komp ała-B ąb a A., B ąb a W. 2006 – Solidago
graminifolia (L.) Elliott on anthropogenic sites
of the Silesian Upland Poland – Biodiv. Res.
Conserv. 3–4: 329–332.
Komp ała-B ąb a A., B ąb a W. 2009 – Threatened and protected species in the Kuźnica
Warężyńska sandpit (Wyżyna Śląska Upland, S
Poland) (In: Rare, relict and endangered plant
and fungi in Poland, Eds: Z. Mirek, A. Nikel) –
W. Szafer Institute of Botany, Polish Academy
of Sciences, Kraków, pp. 259–268.
Krza k le wsk i W. (e d.) 1999 – Projekt techniczny rekultywacji leśnej części wyrobiska górniczego Kopalni Piasku “Kuźnica Warężyńska”
S.A. [Project of technical reclamation into forest of the open-cast “Kuźnica Warężyńska” S.
A.] – Project, Kraków (in Polish).
L atzel V., K limeš ová J., D ole ža l J., P yš ek
P., Tackenb erg O., Prach K. 2011 – The
association of dispersal and persistence traits
of plant with different stages of succession in
Central European man-made habitats – Folia
Geobot. 46: 289–302.
L epš J., Šmi l auer P. 2003 – Multivariate analysis of ecological data CANOCO – Cambridge
University Press, Cambridge, 269 pp.
L ind acher R . (e d.) 1995 – Phanart – Datenbank der Gefasspflanzen Mitteleuropas Erklarung der Kennzahlen, Aufbau Und Inhalt
(Phanart, Database of Centraleuropean Vascular Plants, Explanation of codes, Structure
and Contents) – Veroffentlichungen Geobotanischen Institut der ETH Stiftung Rübel Zurich.
Mar t ine z-Ruiz C., Mar rs R .H . 2007 – Some
factors affecting successional change on uranium mine wastes: insight for ecological restoration – Appl. Veg. Sci. 10: 333–342.
Nová k J., Prach K. 2003 – Vegetation succession in basalt quarries: patterns on a landscape
scale – Appl. Veg. Sci. 6: 111–116.
Nová k J., Konv ička M. 2006 – Proximity of valuable habitats affects succession pat-
journal 33.indb 21
21
terns in abandoned quarries – Ecol. Eng. 26:
113–122.
Pickett S.T.A., C adenass o M.L. 2005 –
Vegetation dynamics (In: Vegetation Ecology,
Ed: E. van der Maarel) – Blackwell, Oxford,
United Kingdom, pp. 172–198.
Pickett S.T.A., C adenass o M.L., Meiners
S.J. 2008 – Ever since Clements: from succession to vegetation dynamics and understanding to intervention – Appl. Veg. Sci. 12: 9–21.
Pos ch lo d P., Tackenb erg O., B onn S. 2005
– Plant dispersal potential and its relation to
species frequency and co-existence (In: Vegetation Ecology, Ed. E. van der Maarel) – Blackwell, Oxford, United Kingdom, pp. 147–169.
Prach K., P yš ek P. 1994 – Clonal plants –
what is their role in succession? – Folia Geobot. Phytotax. 29: 307–320.
Prach K., P yš ek P. 1999 – Using spontaneous
succession for restoration of human-disturbed
habitats: Experience from Central Europe –
Ecol. Eng. 17: 55–62.
Prach K., P yš ek P. 2001 – How do species
dominating in succession differ from others?
– J. Veg. Sci. 10: 383–392.
Prach K., P yš ek P., B ast l M. 2001 – Spontaneous succession in human-disturbed habitats : a pattern across seres – Appl. Veg. Sci. 4:
83–88.
Prach K., Hobbs R .J. 2008 – Spontaneous
succession versus technical reclamation in the
restoration of disturbed sites – Restor. Ecol. 16:
363–123.
Prach K., Wa l ker L.R . 2011 – Four opportunities for studies of ecological succession –
Trends Ecol. Evol. 26: 119–123.
Prach K., P yš ek P., Šmi l auer P. 1997 –
Changes in species traits during succession. A
search for pattern – Oikos, 79: 201–205.
Prach K., Sándor B., Yoyce C h. B., P yš ek
P., van Dig gelen R ., Wieg leb G. 2001
– The role of spontaneous vegetation succession in ecosystem restoration: A perspective –
Appl. Veg. Sci. 4: 111–114.
Prach K., Mar rs R ., P yš ek P., van Dig gelen R . 2007 – Manipulation of succession – (In: Linking restoration and ecological
succession, Eds: L.R., Walker, R.J. Hobbs.) –
Springer, pp. 121–149.
P y wel l R .F., Bu l lo ck J.M., R oy D.B.,
War man L., Wa l ker K.J., R ot her y P.
2003 – Plant traits as predictors of performance in ecological restoration – J. Appl. Ecol.
40: 65–77.
R D e velopment C ore Te am, P. 2011 – R – a
language and environment for statistical computing – Vienna: R Foundation for statistical
computing.
2013-04-30 10:04:58
22
Agnieszka Kompała-Bąba, Wojciech Bąba
Ř ehoun ková K., Prach K. 2006 – Spontaneous vegetation succession in disused gravelsand pits: Role of local site and landscape factors – J. Veg. Sci. 14: 583–590.
Ř ehoun ková K., Prach K. 2008 – Spontaneous vegetation succession in gravel-sand pits:
A potential for restoration – Res. Ecol. 16:
305–312.
Ř ehoun ková K., Prach K. 2010 – Life-history traits and habitat preferences of colonizing
plant species in long-term spontaneous succession in abandoned gravel-sand pits – Basic
Appl. Ecol. 11: 45:53.
Ř ehoun ková K., Ř ehounek J. 2011 –
Sand pits and gravel-sand pits – (In: Nearnatural restoration vs. technical reclamation of mining sites in the Czech Republic,
Eds: K. Řehounková K., J. Řehounek, K.
Prach) – University of South Bohemia in
České Budějovice, České Budějovice, pp.
51–67.
S chu lze E-D., B e ck E., Mü l ler-Hohenstein K. 2002 – Plant Ecology – SpringerVerlag, Berlin, Heidelberg, New York.
ter Braak C.J.F., Šmilauer J. 2002 – CANOCO
4.5 Reference manual user’s guide to Canoco
for Windows: Software for canonical community ordination (version 4.5). Microcomputer
Power, Ithacs, NY, US.
Thui l ler, W., Araúj o M.B., L avorel S.
2003 – Generalized models versus classification tree analysis: a comparative study for predicting spatial distributions of plant species at
different scales – J. Veg. Sci. 14: 669–680.
Tr n ková R ., Ř ehoun ková K., Prach K.
2010 – Spontaneous succession of vegetation
on acidic bedrock in quarries in the Czech Republic – Preslia, 82: 333–343.
Trop ek R ., Kad le c T., Kareš ová P., Spitzer
L., Ko č árek P., Ma lenovský P., B aňař
P., Tuf I. H., Hej d a M., Konv ička M.
2010 – Spontaneous succession in limestone
quarries – J. Appl. Ecol. 47: 139–147.
van Andel J., Arons on J. (e ds.) 2006 – Restoration ecology – Blackwell, Oxford, 319 pp.
Vitouš ek P.M., He din L.O., Mats on P.A.,
Fow nes J.H., Nef f J. 1998 – Within-system element cycles, input-output budgets, and
nutrient limitation. (In: Successes, limitations,
and frontiers in ecosystem science, Eds: M.
Pace, P. Groffman) – Springer-Verlag, Berlin,
pp. 432–451.
Wa l ker L.R ., Ward le D.A., B ardgett, R .D.,
C l arks on B.D. 2010 – The use of chronosequences in studies of ecological succession and
soil development – J. Ecol. 98: 725–736.
Whe eler B. D., L amble y P. W., G e es on
J. 1998 – Liparis loeselii (L.) Rich. in eastern
England: constraints on distribution and population development – Bot. J. Linn. Soc. 126:
141–158.
Wieg leb G., Felin ks B. 2001 – Predictability
of early stages of primary succession in postmining landscape of Lower Lusatia, Germany
– Appl. Veg. Sci. 4: 5–18.
Woźni a k G. 2010 – Zróżnicowanie roślinności
na zwałach pogórniczych Górnego Śląska [Diversity of vegetation on coal-mine heaps (Upper Silesia Poland)] – W. Szafer Institute of
Botany, Polish Academy of Sciences, Kraków,
310 pp. (in Polish, English summary).
Z ob el M. 1997 – The relative role of species
pools in determining plant species richness: an
alternative explanation of species coexistence
– Trends Ecol. Evol. 12: 266–269.
Received after revision October 2012
journal 33.indb 22
2013-04-30 10:04:58