Journal of Landscape Studies 1 (2008), 169 – 187
Received: 10 October 2008; Accepted: 30 October 2008; Published online: 12 November 2008
Journal of
Landscape
Studies
Invertebrate communities in man-made and spontaneously
developed forests on spoil heaps after coal mining
Markéta Hendrychová *1,2, Miroslav Šálek 1, Andrea Červenková 1
/1Czech University of Life Sciences Prague, Faculty of Environmental Science, Department of Ecology,
Kamýcká 129, Praha 6 – Suchdol, Czech Republic
/2Brown Coal Research Institute, j.s.c., Department of Environment and Landscaping, Budovatelů 2830,
Most, Czech Republic
Abstract
The areas remaining after open-cast brown coal mining in the North Bohemian Brown Coal Basin, Czech Republic, are a
remarkable landscape phenomenon. Many of these sites have been reclaimed. Technical reclamation has often been carried
out, forming new terrains and spreading fertile rocks, followed in places by biological reclamation (forestry or agriculture).
In much of the territory, a return to more natural stands is a matter for future decades. However, we are still not sure which
kind of management is best for non-productive functions of the landscape, particularly ecological functions. The objective
of this study is to determine the effects of environmental characteristics on the structure and diversity of invertebrates on
spoil heaps 18 to 40 years after open-cast brown coal mining, including differences between heaps following technical and
biological reclamation and heaps that have developed under spontaneous succession. Special attention is given to indicator
animals, which reflect relationships with soil attributes and plant diversity: epigeon and ground dwelling beetles
(Carabidae), invertebrates inhabiting ground-above vegetation (Heteroptera bugs), and snails and slugs (Gastropoda). In
general, higher species diversity and more abundant taxa were found on non-reclaimed sites under spontaneous succession.
These localities also provided more suitable habitats for rare species, or indicate sites of higher natural value. The most
significant environmental variables affecting invertebrates on spoil heaps were: slopes (negative microclimatic effects of a
north-west facing slope), moisture, herb cover, and proportion of birch (Betula pendula) in the forest. These outcomes can
be applied in restoration management of landscapes after open-cast brown coal mining.
Key words: Post-mining landscapes; succession; reclamation; invertebrates; Carabidae; Heteroptera.
1. Introduction
Teams of scientists have been involved in studies
of landscapes where brown coal was extracted in
open-cast mines over a considerable period of time.
Various ways of restoring impacted landscapes
have been applied. Štýs et al. (1981), Štýs and
Braniš (1999) and Sklenička et al. (2004) have
dealt with the general principles of reclamation.
Until now, most studies carried out in the Czech
Republic and elsewhere in the world have
concentrated on soil development (Šourková et al.,
2004), but have rarely taken into account
interactions with soil biota (Frouz et al., 2007;
Frouz, 2008), with above-ground communities
(Řehoř et al., 2006), or with selected taxa (61%
studies; Ruiz-Jaen and Aide, 2005), eventually
analysed habitat development during the first few
years after disturbance (reviewed by Majer, 1989).
A comprehensive study by Řehoř et al. (2006)
analyses the methodology for technical reclamation
and application of fertilisable rocks, and describes
the pedological characteristics of newly formed
anthropogenous soil profiles. A methodology for
* Corresponding autor; E-mail: [email protected]
Available online at: www.centrumprokrajinu.cz/jls/
169
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
applying fertilisable rocks in various types of
locations is broadly elaborated in this paper. A
review of pedological and biological studies was
published by Hendrychová (2008). Many plant
communities, but only a few animal assemblages,
were described in detail and compared with natural
successions on reclaimed dumps (Majer, 1989).
Studies by Prach et al. (2001), Hodačová and Prach
(2003) and Prach (2003) offer practical knowledge
from the field of plant ecology and management of
damaged sites. Vojar (2006) summarises what is
known about animal colonisation in post-mining
landscapes. Animals are suitable indicators of the
success of reclamation, as they reflect the
properties of the whole ecosystem by means of
diverse interactions (Majer, 1998). Animal life
cycles integrate a wide range of abiotic and biotic
variables (Parmenten et al., 1991), predetermining
them to properly quantify the restoration success in
disturbed areas (for colonisation of post-coal
mining habitats by microorganisms and
invertebrates, see Hutson, 1980; Parmenter and
MacMahon, 1987; Simmonds et al., 1994;
Wheather and Cullen, 1997; Pižl, 2001; Růžek et
al., 2001; Tajovský, 2001; Langcore, 2003 and
Majer, 2005, by amphibians Galán, 1997; Vojar,
2000; by birds Bejček and Tyrner, 1980; Bejček
and Šťastný, 1984, and by small mammals Bejček,
1981; Halle, 1993 and Rathke and Bröring, 2005).
The most comprehensive study on post-mining
animal succession (Nichols and Nichols, 2003) was
carried out in Southwest Australia, but the results
may not be applicable in the Czech Republic, as
the spontaneous succession in central Europe may
be driven by different environmental conditions
(Prach, 2003).
Reclaimed areas develop more rapidly from
the initiation stage. Reclamations minimize
geomorphological processes such as destructive
erosion, which is particularly burdening in unstable
localities (Hüttl and Gerwin, 2004). On the other
hand, spontaneous succession on damaged sites
with extreme conditions is slower, and these sites
can act as refuges for endangered or rare species
that are sensitive to competition, eutrophisation
(Prach, 2003) or require specific climatic
conditions (e.g., an overheated surface). Therefore,
areas left to natural self-development are a good
alternative to technical reclamation from the
perspective of plant communities that may
approximate in time to communities not directly
170
damaged by mining (Prach, 2003). We may expect
that animal communities also behave in a very
similar way.
The aim of this study was to analyse the data
on invertebrate communities inhabiting forested
spoil heaps after coal mining in Northern Bohemia.
Stands of spoil heaps achieve maximum vegetation
cover after 20-30 years, and experience only small
changes afterwards (Wiegleb and Felinks, 2003).
At the same successional stages, the animal
communities inhabiting reclaimed stands with tree
formations came to resemble the communities on
non-disturbed areas adjacent to mines (Holl, 1996;
Hüttl and Weber, 2001). We therefore focused on
medium successional stages (18-45 years old)
inhabited by groups of invertebrates that are
sensitive to disturbances and have a bioindicator
value, such as carabid beetles, bugs and terrestrial
molluscs (Majer, 1989; Lange and Mwinzi, 2003).
We examined the effects of microhabitat attributes
on these invertebrate taxa in the light of differences
between reclaimed and successional stands.
2. Material and Methods
2.1 Study site
The territory of interest is situated between Most,
Litvínov, Bílina and Chomutov in North-West
Bohemia, Czech Republic (50°28′-50°33′N,
13°30′-13°43′E). Most of the study localities lie in
the Brown Coal Basin of Most (Demek, 1987)
bordered by the Krušné Hory Mts. (formed of
ortho-gneisses) to the north and by the České
Sředohoří Mts. (formed of basalt and clinkstone) to
the south. The basin is filled with clay and sand
sediments with thick beds of brown coal.
Anthroposoils dominate in the reclaimed and nonreclaimed areas in the study region, while
cambisols, vertisols and luvisols occur widely in
the surrounding areas. The Bílina River with its
inflows (strongly regulated due to mining) forms
the hydrological core line of the territory. The
Krušné Hory Mts. fall sharply into the basin,
creating a strong anemo-orographic climate effect
in the basin (strong rain shadow and N-W winds).
The Most region as a whole is a part of the hot
climatic zone within the phytogeographic
Podkrušnohorská Basin division (Culek, 1996).
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
The vegetation zone of the region is an upper
hill country belt, alias gradus (supra)collinus
(Skalický, 1988). The complex of succession
stages on anthropogenic sites would be the most
frequent
potential
vegetation
formation
(Neuhäuslová,
1998).
Flood-plain
PrunoFraxinetum forests would naturally develop along
the streams and rivers. The flora of the bioregion is
formed by expansive ruderal species and
neophytes. Natural forest communities are rare,
and are restricted to a few sites around water
bodies and in the foothills of the Krušné Hory Mts.
Plantations on reclaimed spoil heaps, pioneer tree
species and non-forest greenery in villages are
usual. Some specialist species colonise the early
stages of non-reclaimed spoil heaps, in this way
indicating the forest-steppe character of these
habitats (Sládek, 1990). The fauna of this bioregion
is of Hercynian origin, with an impact of West
European elements.
2.2 Study plot selection
The overburden dumps after brown coal extraction
located in the Brown Coal Basin of Most were
mapped and divided into two categories according
to their history: (1) technically and silviculturally
reclaimed (Reclamations) and (2) spoil heaps
developing
under
spontaneous
succession
(Successions), naturally colonised by organisms
from neighbouring areas. In total, 7 and 8 study
plots, each 100 m x 100 m were selected in the
Reclamation
and
Succession
categories,
respectively, based on orthophotomaps and
additional field inspections. The selection criteria
included >18 years since the principal disturbance
(brown coal mining), >30 m from the dump edge
(to minimize the edge effect), >500 m between two
neighbouring study plots (to avoid risk of
pseudoreplications), and ~50% share of greenwood
(to reduce the open/forest habitat proportion
effect). The positions of the selected study plots
were drawn into topographical map using GPS
(Appendix 1). Within each selected plot, three
locations were chosen to represent three types of
microhabitats reflecting the gradient in the
vegetation floors: A – semi-open forest, B – closed
forest canopy without shrubs, C – dense and closed
forest canopy with shrubs (Appendix 2).
2.3 Sample collection
Standard sampling methods were used to find out
the occurrence of particular invertebrate groups.
The collections were timed to the periods of
reproduction (May - August) and increased activity
(sunny days between 10:00 a.m. and 3:00 p.m.) of
the studied animal groups. Epigeon and insects
from the soil surface were collected using passive
pitfall trapping. One pitfall trap with ethylene
glycol was installed in the centre of each location
and exposed for one month. The species inhabiting
the herbs were swept by a net. Net-sweepings
combined with beating (Růžička, 2001) were
conducted in a set of 30 sweeps in a diameter of 20
m around the pitfall trap and additional beatings
from all trees and shrubs in this radius. Snail and
slugs were collected individually from herbs and
tree logs, and also by sieving leaf litter (8 litres)
and using paper traps with constant collection
intensity on all plots. Additional environmental
variables were recorded concurrently to
characterize the habitat structure within this 20-m
circle: Humidity (dry, semi-wet and wet),
percentage cover (%) of vegetation layers (E0 –
mosses, E1 – herbs, E2 – shrubs, E3 - trees),
dominance of particular tree species, mean height
of the herb cover, species richness of herbs, slope
(degree), microclimate (mild or severe accordingly
to the prevailing southerly or northerly winds),
proportions of dead wood (low, medium, high),
leaf litter (height in cm) and type of
microtopography (flat, asperity < 20 cm, asperity >
20 cm).
2.4 Data processing and statistical analyses
The data sets were divided and analysed in three
sections, in accordance with the sampling method
and the indicator groups of the monitored animals:
(1) Invertebrates from pitfall traps (epigeon) were
classified into families (or in a few cases into an
order). The indicator group of ground-dwelling
beetles (Carabidae) was identified into species; (2)
Invertebrates net-swept from vegetation were
classified into families (in a few cases into an
order). The indicator group of bugs (Heteroptera)
was identified into species; (3) Slugs and snails
171
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
4.0
H´: KW-H [1; 39] = 8.75, p = 0.003
3.5
Index of diversity H´
3.0
2.5
2.0
1.5
1.0
0.5
0.0
Succession
Reclamation
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Type of management
Figure 1. Shannon-Wiener index of diversity (H´) of invertebrates from pitfall traps on non-reclaimed successions
and reclaimed sites. KW – Kruskal-Wallis test.
(Gastropoda) were classified into species. For the
list of identified taxa and their abbreviations (used
in the ordination diagrams), see Appendix 3.
The sum of all collections obtained from one
trap, sweeping set and snail collection in a single
microhabitat within a plot was taken as a basic unit
(sample) for subsequent analyses. Ground beetles
(Carabidae) and bugs (Heteroptera) were used as
indicators reflecting the level of habitat
development due to their abundance, species
richness and knowledge of their ecology. Ground
beetles were sorted into groups according to niche
breadth (Hůrka et al., 1996) as relict (R), adaptabile
(A) or eurytopic (E) and according to their
preference for a wet, semi-arid or arid habitat
(Hůrka, 1996). Bugs were sorted as phytophagous
or zoophagous according to their food, and as egg
or adult survivors according to their over-wintering
strategies (Fauvel, 1999; Zurbrügg and Frank,
2006).
The index of diversity was calculated using the
Shannon-Wiener formula H´= – Σpiln pi, where pi
is the proportion that the ith species (taxa)
contributes to the total number of individuals of all
species (Krebs, 1999). Multivariate analyses and
visualization were performed using the CANOCO
172
software package (ter Braak and Šmilauer, 2002),
and basic statistical procedures (correlations and
non-parametric testing) were performed in
STATISTICA ver. 7.0. First, we verified that the
effect of the study localities on the composition of
invertebrate guilds was non-significant (all p <
0.05), and we excluded this factor as a potential
source of pseudoreplications from the analyses.
Second, we examined the inter-correlations (r >
0.6) between all quantitative explanatory variables
(microhabitat characteristics). A tight relationship
was found only between the mean height of the
herbs and the herb layer cover, so that only herb
cover was taken into account in the subsequent
analyses as an underlying factor representing the
herb layer. Finally, seventeen non-correlated
environmental variables were tested using the
Monte Carlo test with 4999 permutations. As the
factors were selected by manual forward selection,
the Bonferonni rule was applied in stating the
significance level (α = 0.05/17 = 0.003) (ter Braak
and Šmilauer, 2002). The significant factors were
entered as covariables in the subsequent analyses to
test the effect of site history (Reclamation/
Succession).
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
8000
Number of individuals
7000
6000
1561
5000
531
3322
4000
3000
3826
674
2000
1062
Insecta
Chelicerata
1000
1058
1409
Reclamation
Succession
Crustacea
Coleoptera
0
Type of management
Figure 2. Proportions of invertebrate groups caught in pitfall traps on reclaimed sites and non-reclaimed
successions.
3. Results
3.1 Pitfall traps
In total, 36 samples from 12 localities were
analysed (nine damaged traps were excluded). The
samples included the complete data on higher taxa
(3.1.1) and on the selected indicator group of
ground-dwelling carabid beetles (3.1.2).
3.1.1 Epigeon on the level of higher taxa
In total, 34 taxa (families or orders) formed by
13,541 individuals were distinguished. The
samples on sites under spontaneous succession
consisted of more individuals (mean + SD = 364 ±
298) and also of more taxa (23 ± 5) than the
samples on reclaimed sites (mean + SD = 332 ±
348 individuals and 18 ± 4 taxa). The index of
diversity reflecting both the number of taxa and the
abundances was significantly higher on the
spontaneously developing stands than on the
reclamations (Fig. 1). Testaceous animals
(Crustacea) were the most abundant group on the
reclaimed sites, while insects (Insecta) dominated
on the non-reclaimed sites (Fig. 2).
Relationships between the abundances of
epigeic taxa and environmental variables were
investigated by the method of redundancy analysis
(RDA). Microclimate was the strongest predictor,
explaining 22.5% of the variation in taxa on the
sites (Monte Carlo test: F = 3.26, p = 0.002). Other
factors, including type of microhabitat, were not
significant (all F < 1.28, p > 0.22). The effect of
the Reclamation/Succession stand was nonsignificant (F = 1.66, p = 0.091) in the subsequent
analysis, in which the significant microclimate was
used as a covariate.
3.1.2 Indicator group of epigeic ground beetles
In total, 31 species of carabid beetles with 1,010
individuals were identified. Significant differences
in numbers of species between reclaimed and
successional sites were found (Fig. 4). The
successional sites were inhabited by more
individuals (mean + SD = 19.7 + 13.3) than the
reclaimed sites (9.9 ± 14.4; Kruskal-Wallis test,
173
1.0
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
Arm
Scar
Blat Dipl
Hist
Luc
Elat
Derm
Staph
mclima-
Forf
Geot
Silph
Hym
Chry
Aphi Leiod
Ar
AcarOpil
Tett
Chil
Lep Onis
Pan
Col_L
Tromb
Curc
Auch
Acri
Het
Bra
Car
Colb
-0.8
Cocc
-0.4
1.0
Figure 3. Biplot of the redundancy analysis (RDA), displaying the positions of all epigeic taxa within the space of the
first two ordination axes (see Appendix 3 for full name of the taxa). Microclimate (cool expositions = mclima-) as the
most significant predictor is associated with the first (horizontal) axis.
16
Number of species: KW-H[1; 36] = 4.53, p = 0.033
14
Number of species
12
10
8
6
4
2
0
-2
Succession
Reclamation
Median
25%-75%
Non-Outlier Range
Outliers
Type of management
Figure 4. Species richness of carabid beetles caught on non-reclaimed successions and reclaimed sites (R). KW –
Kruskal-Wallis test.
174
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
Number of individuals per trap
14
12
10
8
6
4
2
Succession
Reclamation
0
Adaptabile
Eurytopic
Type of management
Figure 5. Distribution of epigeic carabid beetles on non-reclaimed successions and reclaimed sites as relict,
adaptabile and eurytopic reflecting the species niche breadth (after Hůrka et al., 1996).
K[1; 36] = 6.39, p = 0.012). Pterostichus niger was
the most frequent species on both reclaimed and
non-reclaimed sites. Among the sites with
spontaneous succession, however, there were
extreme localities inhabited by poor beetle
communities in terms of low species numbers and
abundances. They included the extreme acidic spoil
with phytotoxic moulds near the village of Braňany
and the former Saxonie open coal mine, where
there were rich coal additions in the mould.
However, several rare or scarce species were found
there (e.g., Brachinus crepitans).
Twice as many adaptabile species and three
times as many eurytopic species were detected on
the non-reclaimed sites as on the reclaimed sites
(Fig. 5). Species associated with arid or wet stands
(including less common species) were caught more
frequently in localities under spontaneous
succession than on reclaimed sites (Fig. 6).
Reclaimed sites were more often inhabited by
species of semi-arid habitats.
Relationships
between
species
and
environmental attributes were evaluated by
canonical correspondence analysis (CCA). Only
the presence of birch (Betula pendula – variable
Bet) as a dominant tree was revealed as a
significant predictor (Monte Carlo test: F = 1.86, p
< 0.002) and humidity as a marginally significant
predictor (wet, F = 2.10, p = 0.003). These two
variables explained 24.3 % of the variation in the
species data, and they were included in the
subsequent model as covariables to test the
Reclamation/Succession effect. However, this
effect was not significant (F = 1.48, p = 0.134),
suggesting that environmental characteristics such
as presence of birch and wet patches influenced the
community of ground-dwelling carabids more than
the site history (i.e., reclaimed or left to
spontaneous succession).
3.2 Net-sweeping
In total, 45 net-sweeping samples entered into the
analyses.
3.2.1 Higher invertebrate taxa swept from
vegetation
175
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
Number of species per pitfall trap
16
14
5.667
12
10
3.190
8
3.190
6
4.048
4
5.238
2
2.714
wet
semi-arid
arid
0
Reclamation
Succession
Type of management
Figure 6. Wet, semi-arid and arid habitat preferences of ground beetles (after Hůrka, 1996) on reclaimed sites and
non-reclaimed successions.
2.8
H´: KW-H[1; 44] = 11.20, p = 0.001
2.6
2.4
Index of diversity H'
2.2
2.0
1.8
1.6
1.4
1.2
1.0
0.8
Succession
Reclamation
Median
25%-75%
Non-Outlier Range
Outliers
Extremes
Type of management
Figure 7. Shannon-Wiener index of diversity (H´) of invertebrates from vegetation on non-reclaimed successions and
reclaimed sites. KW – Kruskal-Wallis test.
176
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
The analysis included 1,918 individuals of 26
higher taxa (families or orders). Typically,
communities inhabiting non-reclaimed sites
showed significantly higher diversity than
communities on reclaimed sites (Fig. 7). The
proportions of major invertebrate groups were
similar on non-reclaimed and reclaimed sites, but
higher absolute numbers appeared on successional
stands. Bugs and dipterous (Diptera) dominated
there (Fig. 8). Herb cover (E1) was detected as the
best predictor of the guild structure (RDA, Monte
Carlo test: F = 4.56, p < 0.002). This factor was
then used as a covariate to examine the
Reclamation/Succession effect. However, this
effect was not significant (F = 2.32, p = 0.032).
3.3 Bugs as an indicator group on vegetation
Bugs were represented by 33 species, consisting of
604 individuals. The most frequent species was
Kleidocerys resedae. The results show a clear
difference between reclaimed and non-reclaimed
stands (75 individuals versus 529 individuals in
total, K[1; 45] = 9.98, p = 0.002; 11 species versus 30
species in total, K[1; 45] = 10.48, p = 0.001). The bug
guilds were also more diversified on sites
developing under spontaneous succession than on
reclaimed sites (Fig. 9).
In addition, the number of individuals
recruited from species over-wintering as
inseminated females (compared to species overwintering as eggs) was higher on successions than
on reclaimed sites (Fig. 10; Kruskal – Wallis test,
K = 8.86, p = 0.003). Sites of both types were
inhabited by phytophagous species rather than
zoophagous species (58.1% and 66.7% of
phytophagous individuals on successions and
reclamations, respectively) with non-siginificant
differences in proportions between successions and
reclamations (p = 0.13; Fig. 11). The proportions
of species classified in these food guilds also did
not differ significantly between reclaimed sites (5
phytophagous and 5 predator species) and
successions (21 and 8 species) (Test of proportions:
p = 0.20).
Neither of the tested environmental variables
contributed significantly in explaining the variation
in the structure of the bug guilds (RDA, Monte
Carlo test: all F < 2.604, p > 0.005). The effect of
Reclamation/Succession was also not significant
(F= 1.51, p = 0.186).
1600
172
1200
214
Number of individuals
1400
1000
405
800
Insecta
123
600
400
200
34
74
99
75
Heteroptera
472
Hymenoptera
Diptera
144
0
Chelicerata
82
22
Reclamation
Coleoptera
Succession
Type of management
Figure 8. Composition of the main invertebrate taxa on reclaimed sites and un-reclaimed successions.
177
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
Number of species: KW-H[1; 45] = 10.48, p = 0.001
10
Number of species
8
6
4
2
0
Median
25%-75%
Non-Outlier Range
-2
succession
reclamation
Type of management
Figure 9. Number of bug species on non-reclaimed successions and reclaimed sites. KW – Kruskal-Wallis test.
4
Mean number of species
3.5
3
2.5
2
1.5
1
0.5
Eggs
Adults
0
Reclamation
Succession
Type of management
Figure 10. Mean number of bug species wintering as adult females or eggs caught on reclaimed and non-reclaimed sites.
178
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
400
352
Number of individuals
350
300
250
200
176
150
100
50
43
31
Phytophagous
Zoophagous
0
Reclamation
Succession
Type of management
Figure 11. Numbers of caught bugs sorted into food guilds on reclaimed and non-reclaimed sites.
3.4 Slugs and snails (Gastropoda)
4. Discussion
Most of the snails and slugs were registered in
small abundances insufficient for calculating a
diversity index. In total, 12 species and 256
individuals were found on 45 sites. The snail
Cepaea hortensis was the most frequent species on
both reclaimed and non-reclaimed sites. Reclaimed
sites were occupied by more species (KruskalWallis test: K = 3.9, p = 0.048) and individuals (K
= 4.33, p = 0.038) than successions. Tree cover
(E3) as the best predictor of the guild composition
(CCA, Monte Carlo test: F = 2.76, p = 0.002)
explained 24.5% of the variation in species data.
Site humidity was a marginally significant
predictor (F = 2.17, p = 0.023) and was thus not
included in the model. When tree cover was used
as a covariate, the effect of Reclamation/
Succession was also not found to be statistically
significant (F = 1.56, p = 0.112). However, some
species tended to prefer reclaimed sites
(hygrophilic species) while several others (e.g.,
acidiphilic species) tended to appear on nonreclaimed sites.
The main objective of this study was to analyse
invertebrate communities on stands that were
reclaimed or started with spontaneous development
18 to 45 years ago and are at the present time
composed of a microhabitat mosaic dominated by
forest formations.
The most important outcome of this study is
that the diversity of invertebrates was found to be
generally higher on spoil heaps under spontaneous
succession than on reclamations, both for epigeic
(ground-dwelling) groups and for invertebrates
activating on vegetation. Non-reclaimed sites
create more suitable habitats for ground beetles
(Carabidae) and bugs (Heteroptera). These results
are consistent with previous findings in the Czech
Republic that many plant and animal species
(including scarce or endangered species) prefer
areas without technical and biological reclamations
(Bejček and Šťastný, 1984; Prach and Pyšek, 2001;
Hodačová and Prach, 2003; Voženílková, 2003;
Novák and Konvička, 2006; Vojar, 2006).
However, the differences between communities of
179
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
slugs and snails inhabiting reclamation and
succession stands remain less clear.
Ground-dwelling invertebrates (individuals)
activated more on reclaimed sites than on
spontaneously developing sites. However, totally
55% of these animals belong to a single taxonomic
group – testaceous animals (Crustacea). It is known
that a small number of highly dominant species
form communities at the beginning of ecosystem
development, and the abundances balance in later
stages (Neumann, 1971; Hejkal, 1985; Vojar,
2006). Forests on technically reclaimed spoil heaps
seemed to resemble younger succession forest
stands from the perspective of invertebrates, and
the study sites under spontaneous succession were
more developed. Sometimes, the diversity of
ground beetles was found to be significantly higher
on early successional stages (Purtauf et al., 2004),
where species that are more tolerant to extreme
conditions may prevail due to their nocturnal
activity or larval dormancy (Hejkal, 1985). Loss of
some pioneer species during succession then
results from the top-down effect of newly settled
species (particularly predators or stronger
competitors), which is typical for later successional
stages (Andersen and Sparlink, 1997). Hejkal
(1985) related the lower abundances of ground
beetles on older spoils to denser vegetation cover,
which is more convenient for other groups of
predators, e. g., millipedes (Chilopoda) or rove
beetles (Staphylinidae), which are better
morphologically adapted to dense vegetation than
carabids. Prevalence of adaptabile ground beetles
implies that the areas after brown coal mining
(succession and reclamation) are well restored and
verge on a natural state (Hůrka et al., 1996). At the
beginning of site development, the succession of
invertebrate communities is faster on reclaimed
sites than on non-reclaimed sites, due to the
immediate human-induced changes that accelerate
reclaimed sites toward tree-stand habitats.
However, only a few other new species inhabit the
reclaimed localities afterwards, while more diverse
communities develop continually on sites with
spontaneous succession, being enriched by other
forest species. During development, reclaimed sites
stay at the same level, but there is a clear trend
toward an increase in species numbers and animal
individuals on spontaneously developing sites
(Červenková, 2008). In addition, reclaimed sites
may suffer from expansion of reed grass
180
(Calamagrostis epigejos), where the root systems
are dissected by silvicultural machinery, and
vegetative dispersion of this strongly competitive
plant creates very compact cover that blocks
subsequent development of the plant community
(Prach, 2003). A higher number of adult bugs
wintering on vegetation of spontaneously
developed sites may indicate better or more diverse
shelter on these stands (more organic matter, e.g.,
dead leaves, bark, leaf litter and stones).
The most critical environmental factor
negatively influencing the epigeon in our study was
unfavourable microclimate, primarily caused by
slope orientation to the north-west with cool
prevailing winds. However, opposite results were
documented by Hawkins and Cross (1982). In their
study, north-facing spoils had a more positive
influence on the occurrence of invertebrates than
south-facing slopes. However, this was found in
the initial stages of succession, when the soil
surface is less overgrown by vegetation and thus
rather overheated. By contrast, our study sites with
forest formations in later succession stages had
rather moderate microclimatic conditions in
general. Another important environmental factor
positively influencing the occurrence of grounddwelling beetles was an increasing proportion of
birch forest, suggesting that birch, as one of the
earliest tree colonists, has a favourable impact on
the development of the early stages of ecological
succession. The composition of the net-sweeping
samples markedly reflects the cover of herbs (herb
floor E1), as the invertebrates caught by this
method are usually phytophagous insects that
consume plants or use them as shelter.
The effect of tree cover was most important
for slugs and snails. Eyre et al. (2003) revealed a
wet regime and plant cover as factors that
particularly influence these invertebrates. Snails
are strongly dependent on site humidity and also on
availability of calcium (Ložek, 1956). Some tree
species, such as maple (Acer) or ash (Fraxinus),
may improve the content of calcium in the soil or
leaf litter. On the other hand, some dendrophilic
molluscoids do not prefer birch stands resulting
from spontaneous succession (stands that are not
mature), due to the inappropriate chemical
character of the substratum. Our results suggest
that slugs without shells (e.g., Arion sp.), as well as
acidophilic or not strictly calciphilic snails (Ložek,
1955 in Kajerová-Rafajová, 2002), prefer
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
spontaneously developing sites, while hygrophilic
species tend to inhabit denser forest vegetation on
reclaimed stands.
Poor communities composed of few species
were observed in localities with extreme habitat
conditions (acidic soils or high salt content were
indicated by acidiphilic or halophilic plants or
fungi). On the other hand, some rare species
colonise these sites, expanding from the
neighbouring refuges of the České Středohoří Hills,
where there is a similarly hot microclimate. This is
consistent with the findings for several
thermophilic plants in the Most Region (Sládek,
1990). Disturbed sites with extreme conditions
where spontaneous succession proceeds can
therefore function as refuges for scarce species,
e.g., species sensitive to eutrophization (Prach,
2003) or species requiring specific soils or a
specific microclimate (e.g., an overheated surface).
Brändle et al. (2000) classified 10 out of 75 species
recorded on spoil heaps after brown coal mining in
Germany as regionally scarce. Also, high numbers
of uncommon or specialized spider species were
documented on spoil heaps in Germany (Mrzljak
and Wiegleb, 2000). Many rare species have to
displace from intensively cultivated farmland to
post-industrial areas (Konvička and Beneš, 2005),
and spoil heaps may thus play an important role in
these refugial translocations.
5. Conclusions
Several microhabitat characteristics (environmental
variables) including slope, herb layer cover, tree
cover and tree species composition had more
considerable effects on the invertebrate groups
under study than the history of the site (either
reclaimed or left to natural succession after
heaping). However, as the diversity and the
numbers of invertebrates was always generally
higher on spoil heaps under the spontaneous
succession, we conclude that these sites provide
better
ecological
conditions
than
most
silviculturally reclaimed spoil heaps. This is
especially due to differences within habitat
characteristics (microtopography, microclimate,
wet regime), diverse vegetation cover, complexity
of food webs or nutrient cycles (more species at
higher levels of food chains or with demands on
organic litter). In addition, localities with natural
development were inhabited by some species not
present on reclaimed sites.
We point out that leaving selected areas unreclaimed is a good alternative to the conventional
restoration process, which aims at higher biological
diversity in post-mining landscapes. The most
effective positive impacts are then expected on
sites situated near to natural centres of animal
dispersion and on sites with diverse soil conditions,
including patches of acidic or saline moulds and
uneven terrain with small depressions that support
water accumulation.
Acknowledgement
We thank the Grant Agency of the University of
Life Sciences in Prague, which funded the field and
laboratory work (Project IGA 41110131242125),
and the Brown Coal Research Institute in Most for
the loan of an off-road vehicle and for facilitating
entry through mining territory. We thank Pavel
Hendrych for his help in the field, and Jan Růžička,
Petr Kment, Oto Nakládal and Lenka Sirovičová
for classifying the invertebrates. We are obliged to
Robin Healey for language revision of the text.
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Appendix 1. Selection of study locations.
Appendix 2. Design of the experiment.
184
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Abbrev.
A_com
A_conv
A_macul
A_paral
A_simil
B_crep
C_conv
C_cor
C_erat
C_fus
C_gra
C_hort
C_intr
C_nem
H_rub
L_dep
L_ferr
M_min
N_big
N_germ
N_pal
O_schrau
Ox_obs
P_bicus
P_cup
P_mac
P_mel
P_nig
P_obl
P_ruf
P_vers
Species
Amara communis
Amara convexior
Amara makolskii
Abax paralelipipedus
Amara similata
Brachinus crepitans
Carabus convexus
Carabus coriaceus
Calathus eratus
Calathus fuscipes
Carabus granulatus
Carabus hortensis
Carabus intricatus
Carabus nemoralis
Harpalus rubripes
Licinus depresus
Leistus ferrugineus
Microlestes minutulus
Notiophilus biguttatus
Notiophilus germinyi
Notiophilus palustris
Ophonus schaubergerianus
Oxypselaphus obscurus
Panagaeus bipustulatus
Poecilus cupreus
Pterostichus macer
Pterostichus melanarius
Pterostichus niger
Pterostichus oblongopunctatus
Pseudoophonus rufipes
Poecilus versicolor
Abbrev.
ACAR
ACRI
APHI
AR
ARM
AUCH
BLAT
BRA
CAR
COC
COL_L
COLB
CURC
DERM
DIPL
ELAT
FORF
GEOT
HET
HIST
HYM
CHIL
CHRY
LEP
LIOD
LUC
ONIS
OPIL
PAN
SCAR
SILPH
STAPH
TET
TROMB
Taxon
Acarina
Acrididae
Aphididae
Araneae
Armadillidiidae
Auchenorrhincha
Blattodea
Brachycera
Carabidae
Coccinellidae
Coleoptera_larvae
Collembola
Curculionidae
Dermestidae
Diplopoda
Elateridae
Forficulidae
Geotrupidae
Heteroptera
Histeridae
Hymenoptera
Chilopoda
Chrysomelidae
Lepidoptera
Leiodidae
Lucanidae
Oniscidea
Opilionida
Panorpidae
Scarabaeidae
Silphidae
Staphylinidae
Tettigoniidae
Trombidiidae
Appendix 3. List of found taxa
a) Ground beetles (Carabidae) and other taxa of epigon
185
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
Abbrev.
ACAR
ACRI
APHI
AR
AUCH
BRA
CAR
COC
CURC
DIPL
ELAT
FORF
HET
HIPP
HYM
CHRYSOM
CHRYSOPA
KATER
LAR
LEP
NEM
NEUR
ODON
OPIL
STAPH
Taxon
Acarina
Acrididae
Aphidiinae
Araneae
Auchenorrhyncha
Brachycera
Carabidae
Coccinellidae
Curculionidae
Diplopoda
Elateridae
Forficulidae
Heteroptera
Hippoboscidae
Hymenoptera
Chrysomelidae
Chrysopidae
Kateretidae
Larvae
Lepidoptera
Nematocera
Neuroptera
Odonata
Opilionida
Staphylinidae
TROMB
Trombidiidae
Appendix 3. List of found taxa
b) Invertebrates from vegetation and bugs (Heteroptera)
186
Abbrev.
Ad_lin
Ael_ac
Carp
Cor_mar
Dic_ech
Elas_gr
Eur_ole
Graph_lin
Him_apt
Him_mirm
Kleid_res
Lyg_pra
Myrm_mir
Nab_bre
Nab_lim
Nab_pseud
Orius
Orthops
Pal_pras
Perit_gen
Phyt_aus
Pyrr_ap
Rhop_par
Rhop_sub
Scol_thom
Stag_bip
Sten_cal
Sten_leav
Stenot_bin
Stict_punc
Styg_cimb
Ting_amp
Species
Adelphocoris lineolatus
Aelia acuminata
Carpocoris sp.
Coreus marginatus
Dictyla echii
Elasmucha grisea
Eurydema oleraceum
Graphosoma lineatum
Himacerus apterus
Himacerus mirmicoides
Kleidocerys resedae
Lygus pratensis
Myrmus miriformis
Nabis brevis
Nabis limbatus
Nabis pseudoferus
Orius (Heterorius) sp.
Orthops sp.
Palomena prasina
Peritrechus geniculatus
Phytocoris austriacus
Pyrrhocoris apterus
Rhopalus parumpunctatus
Rhopalus subrufus
Scolopostethus thomsoni
Stagnomus bipunctatus
Stenodema calacaratum
Stenodema laevigatum
Stenotus binotatus
Stictopleurus punctatonervosus
Stygnocoris cimbricus
Tingis ampliata
M. Hendrychová et al.: Journal of Landscape Studies 1 (2008), 169 – 187
Abbrev.
Helix_p
Vit_pell
Aegop_mi
Trich_hi
Brad_fru
Lym_sta
Ario_hor
Arion_lus
Succ_put
Cepa_hor
Euom_str
Alin_bip
Species
Helix pomatia
Vitrina pellucida
Aegopinella minor
Trichia hispida
Bradybaena fruticum
Lymnaea stagnalis
Arion hortensis
Arion lusitanicus
Succinea putris
Cepaea hortensis
Euomphalia strigella
Alinda biplicata
Appendix 3. List of found taxa
c) Snails and slugs (Gastropoda)
187