Diversity of ground-dwelling arthropods on overburden dumps after coal mining
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Ecologica Montenegrina 61: 68-87 (2023)
This journal is available online at: www.biotaxa.org/em
https://dx.doi.org/10.37828/em.2023.61.8
Diversity of ground-dwelling arthropods on overburden dumps after
coal mining
SERGEY L. LUZYANIN1*, ANNA A. RESENCHUK2, MARIA O. OSIPOVA3 &
DMITRY A. SIDOROV4
1,2,3,4
Kemerovo State University, Krasnaya, 6, Kemerovo, 650000, Russia;
1
https://orcid.org/0000-0001-9293-4377; 2 https://orcid.org/0000-0001-5146-6650;
3
https://orcid.org/0009-0002-9473-6125; 4 https://orcid.org/0000-0002-5057-7436;
*Corresponding author. E-mail: sl_luzyanin@mail.ru
Received 28 January 2023 │ Accepted by V. Pešić: 7 March 2023 │ Published online 11 March 2023.
Abstract
Technogenic landscapes formed by coal mining are unique model sites for studying primary succession. We studied the
formation of arthropod communities on rock dumps at coal mine dump sites of different ages and in natural
environments. Using the pitfall traps, 5 classes of arthropods were found, with the domination of Insecta and Arachnida.
The species composition for most of the studied groups of invertebrates was determined, the Berger-Parker dominance
and Shannon species diversity indices were calculated. It has been established that the colonization of dumps depends
not only on the age of recultivated lands, but also on habitat conditions (soil characteristics, vegetation parameters). At
the beginning of the succession, dumps are massively populated by small-sized species that are tolerant of extreme
conditions and have high dispersal abilities. Further development of vegetation entailed a change in the general
structure of the ground-dwelling of arthropods communities. Discriminant analysis showed that the macrofauna of the
compared sites differs statistically significantly in the space of discriminant functions. The areas at the foot of the dump
and the natural biotope are characterized by the maximum differences; the greatest similarity was found between the
young and old dumps. A significant contribution to the overall discrimination is made by the beetles of the families
Geotrupidae, Staphylinidae, Elateridae, as well as centipedes (Chilopoda), and terrestrial bugs (Hemiptera). This study
showed that communities of ground invertebrates are good indicators for monitoring the restoration of technogenically
disturbed lands.
Key words: primary succession; Coleoptera; Araneae; Myriapoda; Oniscidea; open-pit coal mine.
Introduction
The Kuznetsk coal basin (Kemerovo region – Kuzbass, Western Siberia) is one of the most important coal
mining regions of Russia. Open mining activity significantly transforms the local landscape, has a negative
impact on local ecosystems and communities of organisms. A key feature of such territories is the presence
of extensive overlays of extracted rock materials in dumps.
It is established at the legislative level that all technogenically disturbed lands must be reclaimed.
Rehabilitation of post-technogenic territories is aimed at restoring landscapes disturbed or completely
Ecologica Montenegrina, 61, 2023, 68-87
LUZYANIN ET AL.
destroyed by mining, including the restoration of functional and sustainable ecosystems (Hüttl & Gerwin
2005; Macdonald et al. 2015). Two-stage reclamation actions are generally used: (1) technical reclamation
provides for planning, slope formation, removal and application of a fertile soil layer, the installation of
hydraulic and reclamation structures, as well as other works that create the necessary conditions for the
implementation of measures to restore soil fertility; (2) biological reclamation, the application of organic and
mineral fertilizers, sowing and planting of plants, and plant care (Hildmann & Wünsche 1996; Hendrychová
et al. 2012).
Unfortunately, nowadays there is no precise data on how much land in Kuzbass has been reclaimed
and how much still needs recultivation. According to preliminary data, the area of disturbed lands in the
region is about 170 thousand hectares (1.77% of the total area of Kuzbass), of which no more than 40% have
been reclaimed at present (Fotina et al. 2022).
Landscapes formed as a result of coal mining are quickly populated by microorganisms, plants and
animals. An important role in the rehabilitation of these territories is playing soil fauna, which is actively
involved in decomposition, nutrient cycling, and soil formation (Kirmer & Mahn 2001; Prach & Pyšek 2001;
Swift et al. 2004; Bröring & Wiegleb 2005). It is important to note that the study of the dynamics of the main
characteristics of the pedobiont community can be used as a bioindication of degradation or restoration of
ecosystems (Van Straalen & Verhoef 1997).
Depending on body size, soil organisms can be divided into groups: microfauna (< 100 µm) –
including protozoa and nematodes, mesofauna (100 µm – 2 mm) – acari, springtails, diplura, symphylans,
enchytraeids, and macrofauna (> 2 mm) – molluscs, spiders, insects, earthworms, woodlice (Swift et al.
1979; Briones 2014). In the composition of the soil fauna, geobionts are distinguished, obligate inhabitants
of the soil, and ground-dwelling invertebrates, inhabitants of the soil surface and its litter (Menta 2012).
Different research focused on revegetation in post-mining areas (Moreno-de las Heras et al. 2008;
Singh 2011; Kupriyanov & Manakov 2016; Novianti et al. 2017) as well as soil fauna (Dunger et al. 2001;
Frouz et al. 2008, 2013; Dunger & Voigtländer 2009; Boyer & Wratten 2010). The authors note that the
colonization of newly formed initial landscapes depends on the distance of the dump from natural cenoses,
the structure of vegetation and the species composition of plants in the areas, as well as abiotic
environmental factors (Bröring & Wiegleb 2005).
Comprehensive data on the structure of arthropods communities at different stages of recovery are
required for an assessment of environmental changes in the emerging post-technogenic ecosystems. The
main goal of our study is to investigate the ground-dwelling macrofauna communities of arthropods rock
dumps formed after coal mining, as well as to conduct long-term monitoring of the disturbed areas
recolonization by this group of invertebrates.
Materials and Methods
1. Research Area
The study was conducted on the Kedrovsky coal mine (Kemerovo region, Russia) (Fig. 1). According to the
natural zoning, the study area is located on the border transition zone between the northern forest-steppe of
the Kuznetsk Depression and the northwestern subtaiga of the Kuznetsk Alatau (Pisarenko 2014).
Overburden dumps after coal mining were selected, which differed from each other in the time of
formation and are at different stages of recovery: Kedr1 (2006), and Kedr2 (1998) (Fig. 2). The dumps are
composed of a heterogeneous mixture of sandstones, siltstones, coal particles, and a slight inclusion of loess-
like loams. The fractional composition of the surface layer of embryozems (0–10 cm) is dominated by stony
(particle size 3–10 mm) – an average of 22–26.5%, and gravel (1–3 mm) – 16.1–22.2% of the fraction, 9.3–
19.6% of blocky fractions (> 10 mm) are also present, and the proportion of fine earth (< 1 mm) is 5.1–9.3%.
After the dumps were formed, the technical (leveling the profile of the dump) and biological
(planting 5–7-year-old pines) stages of reclamation were performed on them. On the site Kedr1
(55°30'38.56"N, 86°04'00.44"E) there are row plantations of Scots pine (Pinus sylvestris L.), and a birch
(Betula pendula Roth) is occasionally found. Sainfoin (Onobrychis arenaria (Kit. ex Willd.)), cocksfoot
(Dactylis glomerata L.), and Kentucky bluegrass (Poa pratensis L.) predominate in the herbage. On the
dump, where the Kedr2 site is located (55°30'31.46"N, 86°04'12.43"E), by the time of the study, forb-
legume-grass meadows had formed with the inclusion of trees and shrubs represented by sea buckthorn
(Hippophae rhamnoides L.), bird cherry (Padus avium Mill.), Scots pine (Pinus sylvestris), birch (Betula
pendula), etc.
Ecologica Montenegrina, 61, 2023, 68-87 69
ARTHROPODS ON OVERBURDEN DUMPS AFTER COAL MINING
Figure 1. Location of the study site.
Site Kedr3 (55°30'29.70"N, 86°04'52.64"E) is located at the foot of the dump (the age of the dump is
about 20–25 years). The site presents a heterogeneous area of birch forest with areas occupied by trees and
shrubs, as well as open glades with forest meadows.
The control site (Kedr4, 55°33'26.17"N, 86°10'02.75"E), has no human activity, it is located 3 km
north of the dumps under study. It is a fragment of a sparse aspen-fir forest with a small admixture of birch
and spruce (also called boreal forest, or taiga). It has the type of vegetation that existed in the study area
before the start of active coal mining.
2. Sampling Procedures
The sampling campaigns were conducted for six consecutive years (2013–2017, 2022) from May to August
(a period which the maximal activity of arthropods).
The number of ground-dwelling arthropods was counted using the standard soil-zoological method –
pitfall traps. Were used plastic cups with a neck diameter of 70 mm, filled with a fixing liquid (4% acetic
acid). Traps were placed in each section in a linear transect of 10 pieces (one trap for every 10 meters). This
distance is necessary to obtain a representative sample. The traps were taken collected every 7–10 days; all
individuals collected from one site were considered a sampling unit.
The collection samples were identified into the systematic groups (classes, orders, families, etc.).
Most of those collections were identified as species, except for true bugs, earwigs, harvestmen, and some
families of beetles (Curculionidae, Dermestidae, Tenebrionidae). The number of captured specimens of
above-ground arthropods were converted per 10 trap-days to obtain relative values (dynamic density). In
total, more than 110 000 specimens of ground-dwelling arthropods were counted.
Sometimes springtails, acarid mites, and other soil invertebrates fell into pitfall traps. All of them
were excluded from the data analysis, since we consider them an accidental capture. The main method for
accounting for these groups is soil samples followed by forcing on a Tullgren funnels extractor (Crossley &
Blair 1991), which is not used in our work. In addition, we did not consider ants when calculating the total
dynamic density and abundance of ground-dwelling arthropods. These are insects living in eusocial colonies,
70
LUZYANIN ET AL. with a clear distribution of roles in the family. Forager routes may be in close proximity to pitfall traps and, as a result, be filled with them, which will lead to a large distortion of the data (large error). Figure 2A-B. Study sites: (A) Kedr1, (B) Kedr2. Ecologica Montenegrina, 61, 2023, 68-87 71
ARTHROPODS ON OVERBURDEN DUMPS AFTER COAL MINING
Figure 2C-D. Study sites: (C) Kedr3, (D) Kedr4.
72
LUZYANIN ET AL.
3. Microenvironmental parameters
Soil samples were taken annually at each site in three repetitions using the envelope method, consisting of 5
separate point samples (200 g of the top soil layer to a depth of 0–10 cm below the litter layer). For chemical
analyses, a combined sample was prepared by mixing point samples taken from one sample site. Trial sites
are laid in areas with a homogeneous soil and vegetation cover. Soil samples were delivered to the laboratory
of the agrochemical service testing center "Kemerovsky" (Kemerovo) within 24 h, where they were
analyzed.
Since the object of study in our research was above-ground arthropods, we measured the air
temperature precisely on the soil surface using automatic recorders Hygrochron Temperature/Humidity
Logger iButton DS1923‑F5. The Hydro-thermal Coefficient of Selyaninov (HTC) was calculated to assess
the level of moisture in the territory. It is determined by the ratio of the amount of precipitation (r) in mm for
the period with average daily air temperatures above 10 °C to the sum of temperatures (∑t) for the same
time, reduced by 10 times:
The lower the hydrothermal coefficient, the drier the area.
Description of phytocenoses were characterized according to Lavrenko & Korchagin (1976). Areas
10 × 10 m were randomly marked on the study sites. In this square, the coverage of forbs, grasses, and
woody vegetation was visually assessed, expressed as a percentage. For greater accuracy of the description,
several areas for description were laid at each site. The obtained values of the determined parameters are
shown in Table 1.
Table 1. Parameters measured at the dumps and control site (mean ± SE).
Study sites
Parameter
Kedr1 Kedr2 Kedr3 Kedr4
Vegetation cover (%) 57.05 ± 11.08 75.67 ± 3.00 87.5 ± 4.34 96.26 ± 2.18
Turf extent (%) 3.98 ± 1.25 51.08 ± 2.27 31.25 ± 2.17 20.01 ± 0.01
Soil pH 9.01 ± 0.08 8.34 ± 0.16 6.47 ± 0.04 5.56 ± 0.15
Soil organic carbon (%) 1.95 ± 1.11 4.94 ± 0.47 10.37 ± 0.04 6.61 ± 0.6
Total Nitrogen (%) 0.13 ± 0.04 0.29 ± 0.03 0.53 ± 0.01 0.35 ± 0.01
T (oC) 17.27 ± 3.44 16.56 ± 3.61 16.33 ± 3.28 15.02 ± 3.88
HTC 0.14 ± 0.09 0.15 ± 0.12 0.14 ± 0.1 0.12 ± 0.1
4. Data Analysis
The ground-dwelling arthropods assemblages were characterized by taxonomic composition, abundance, and
dynamic density. To assess alpha diversity, the Berger-Parker dominance index (d) and the Shannon species
diversity index (H`) were calculated.
To examine the influence of environmental factors on the abundance of ground-dwelling arthropods,
we used one-way ANOVA followed by Tukey's HSD test separately for each independent variable.
Variables were tested for normality and homogeneity of variance using the Shapiro-Wilk and Levene’s tests.
Invertebrate abundances were log-transformed before statistical analysis to achieve uniformity of variance.
Discriminant analysis was carried out to determine the features that best discriminate objects belonging to
different groups.
All statistical processing of the material was carried out using the software package Statistica 10. In
statistical analyses a minimum probability level of p < 0.05 was accepted.
Results
Analysis of the collected material showed that arthropods of the five classes take part in the formation of the
taxonomic composition of the ground-dwelling population: Malacostraca (Order: Isopoda), Arachnida,
Ecologica Montenegrina, 61, 2023, 68-87 73
ARTHROPODS ON OVERBURDEN DUMPS AFTER COAL MINING
Chilopoda, Diplopoda, and Insecta (Fig. 3, Appendix). The dynamic density of the study macrofauna was 2
times lower that at the foot of the dump and in the control (Fig. 4). The dominant classes in all areas of the
study were insects and arachnids, which account for 97.1% of the total collection of arthropods. Insects are
represented by three orders: Dermaptera, Heteroptera and, Coleoptera; arachnids – Araneae and Opiliones.
Figure 3. The ratio of ground-dwelling arthropods taxa in the study site.
Environmental factors influence the abundance of ground-dwelling arthropods (Table 2). Hydro-
thermal Coefficient of Selyaninov (HTC) and soil organic carbon have the greatest effect, the smallest effect
size is noted for the turf extent.
Table 2. Results of one-way ANOVA in some factors effect on the number of ground-dwelling arthropods.
Effect size
Factor Wilks Value F Effect df Error df p-level
(η2)
Vegetation cover 0.218 16.37 8 1330 0.18 0.000
Turf extent 0.181 25.72 6 1332 0.26 0.000
Soil pH 0.157 17.76 9 1323 0.21 0.000
Soil organic
0.134 19.59 9 1329 0.22 0.000
carbon
HTC 0.817 4.02 5 1334 0.05 0.000
The results of the discriminant analysis showed that the ground-dwelling macrofauna of the
compared sites differs statistically significantly in the space of discriminant functions (Wilks' Lambda:
0.271, approx. F(51.3927) = 42.35053, p < 0.00001) (Fig. 5). The cohesiveness of the distribution in space is
weak, and the Mahalanobis distance between the sample centroids was small. The areas at the foot of the
dump and control are characterized by the maximum differences (Squared Mahalanobis distance is 6.47); the
greatest similarity was found between the young (Kedr1) and old (Kedr2) dumps (0.98). The F-remove
statistics and Wilks' private lambda statistics show that beetles of the families Geotrupidae, Staphylinidae,
and Elateridae, also centipedes (Chilopoda), and true bugs (Hemiptera) make the greatest contribution to the
overall discrimination.
74
LUZYANIN ET AL.
Figure 4. Dynamic density of ground-dwelling of arthropods in the study site (mean ± SD).
1. Families of Coleoptera
Among the beetles on the dumps, representatives of the families Staphylinidae and Carabidae dominate
(93.3% of the total collection), that is anticipated because these families are considered the most numerous in
the world (Sharova & Makarov 2012; Salnitska et al. 2022).
1.1. Species of Carabidae
Seventy-two species of ground beetles of 22 genera and 13 tribes were recorded in the study areas of the
Kedrovsky coal mine. The predominant tribes in terms of species are Zabrini, Pterostichini, and Harpalini,
which account for 49.9% of the total number of species.
The species diversity of ground beetles at the Kedr1 site is 31 species. The Shannon index calculated
for this area shows the maximum value – H` = 4.03, and the Berger-Parker index, on the contrary, the
minimum value – d = 0.27 (at p < 0.001). This may how that there is the greatest uniformity in the
abundance of ground beetle species (Magurran 1988). The older dump (Kedr2) is characterized by the lowest
species abundance of ground beetles – 24 species (H` = 1.52, d = 0.73), among them dominated Pterostichus
magus (Mannerheim, 1825) and Carabus regalis Fischer von Waldheim, 1820 (50.4% and 20.8%
respectively). Ground beetle communities at the foot of dumps include 22 species (H` = 2.29, d = 0.48), and
37 species in the control area (H` = 3.44, d = 0.31).
Directly on the dumps dominated the meadow-steppe Calathus erratus (C.R. Sahlberg, 1827) (25.3),
the eurytopic Amara communis (Panzer, 1797) (22.8%), and the floodplain-coastal Bembidion gilvipes
Sturm, 1825 (10.1 %) and meadow Trechus secalis (Paykull, 1790) (7.6%). In the control site, Trechus
secalis and Pterostichus magus were noted most abundant. Were recorded only on dumps: Amara
bamidunyae Bates, 1878, A. montivaga Sturm, 1825, A. gebleri Dejean, 1831, Ophonus laticollis
Mannerheim, 1825, Harpalus luteicornis (Duftschmid, 1812), Microlestes minutulus (Goeze, 1777), and
Lebia chlorocephala (J.J. Hoffmann, 1803). In 2022, we noted a decrease in the species diversity and
abundance of ground beetles in the dumps compared with the initial stages of the study. The ratios of ground
beetle biotopic groups also change. First, on the sites located on dumps, eurytopic species dominate in terms
of species and abundance, as well as groups that prefer open habitats (steppe, meadow-steppe). Then, there is
a clear tendency for the appearance of ground beetles confined to meadow and forest biotopes, for example,
Poecilus versicolor (Sturm, 1824) (dynamic density in plots 1.51 ± 2.45 indiv. / 10 trap-days), Pterostichus
magus (5.14 ± 6.38), and Harpalus latus (Linnaeus, 1758) (1.57 ± 2.13).
Ecologica Montenegrina, 61, 2023, 68-87 75
ARTHROPODS ON OVERBURDEN DUMPS AFTER COAL MINING
Figure 5. Ordination of the ground-dwelling of arthropods in the overburden dumps and control area in the plane of two
discriminant axes.
1.2. Species of Staphylinidae
Fifty-seven species of rove beetles belonging to 40 genera from 7 subfamilies (Omaliinae, Tachyporinae,
Aleocharinae, Oxytelinae, Steninae, Paederinae, Staphylininae) were found on rock dumps and in the control
zone. The vast majority of species belong to the forest (31) and eurytopic (20) biotopic groups. Zoophages
(31 species) and mixophages (15) predominate by the type of nutrition, 11 species had a saprotrophic type of
nutrition.
The species diversity of rove beetles increases in the process of succession. Twenty-eight species of
rove beetles (H` = 2.77, d = 0.18, p < 0.001) were recorded on the young dump (site Kedr1). The maximum
number of species (47) was noted in the Kedr3 site (H` = 2.95, d = 0.22, p < 0.001). Thirty-five species of
rove beetles were identified in the control plot. At the same time, for this site, we got the smallest value of
the Shannon index (1.04) and the maximum for the Berger-Parker index (0.82). Such results indicate to us a
decrease in diversity and an increase in the degree of dominance of one species. Indeed, Drusilla
canaliculata (Fabricius, 1787), one of the most common Palearctic rove beetles, dominated in all sites. This
is a eurytopic species that often lives near anthills (Salnitska et al. 2022). The proportion of the species in the
collections ranged from 19% at the Kedr1 site to 82% in the control (Kedr4). The myrmecophilous species
Oxypoda formiceticola Märkel, 1841 also dominated in the dumps (17% of the collections), while it was not
recorded in the control.
1.3. Other Coleoptera
In the order Coleoptera, in addition to the indicated dominant families, representatives of at least 8 more
families of beetles were noted (Fig. 6): Geotrupidae, Elateridae, Curculionidae, Dermestidae, Tenebrionidae,
Byrrhidae, Histeridae, Silphidae. The ratio of these families in the study areas was different. As can be seen
from Fig. 6, weevils, carrion beetles, and click beetles are the most common on the dumps, the share of
which in the collections is 68.1%. An increase in the proportion of the family Byrrhidae due to the
dominance of Byrrhus pustulatus (Forster, 1771) was noted in the old dump (Kedr2). At the foot of the
dump, there is a strong predominance in the collections of the family Geotrupidae (72.2%), which is
represented by the only species Geotrupes stercorosus (Hartmann in Scriba, 1791). In the control site
(Kedr4), beetles of the families Elateridae and Silphidae dominated (53.3% and 26.3%, respectively). The
76LUZYANIN ET AL.
click beetle’s community was dominated by Agriotes obscurus (Linnaeus, 1758) and Hypnoidus gibbus
(Gebler, 1847), while the carrion beetle’s community was dominated by Nicrophorus vespilloides Herbst,
1783, Silpha carinata Herbst, 1783, and Phosphuga atrata (Linnaeus, 1758). These species of these families
were also widespread in the dumps.
Figure 6. The ratio of beetle families in the study site (with the exception of Carabidae and Staphylinidae).
2. Araneae
On the study dumps, 70 species of spiders belonging to 32 genera from 13 families were recorded. The
greatest species diversity was shown by the families Lycosidae – 19 species, Gnaphosidae – 17 species, and
Linyphiidae – 14 species, which can be attributed to the ecological group of "hunters on the soil surface".
The most significant species diversity of spiders, 37 species (H` = 3.01, d = 0.24, p < 0.001), was recorded at
the foot of the dump (Kedr3). 33 species were recorded directly on the dumps (H` = 2.93, d = 0.21 for the
Kedr1 site and H` = 2.91, d = 0.19 for the Kedr2 site, at p < 0.001). The control zone is characterized by the
lowest species diversity of spider – 21 species (H` = 0.95, d = 0.84).
Wolf spiders (51%) are the most numerous in collections at dumps, among which spiders of the
genus Pardosa dominate (82% of the number of Lycosidae). Spiders of the families Gnaphosidae (29%) and
Linyphiidae (10%) are less numerous. A statistically significant positive correlation was established between
the dynamic density of Lycosidae and Gnaphosidae (r = 0.43, p < 0.05).
On the young dump (Kedr1), Pardosa agrestis (Westring, 1861) prevailed (18.7% of the total
number of spiders), a species confined to open or lightly forested habitats, as well as Pardosa prativaga
(C.L. Koch, 1870) (11.2%), characterized by wide ecological plasticity (Trilikauskas & Luzyanin 2018). The
sites Kedr2 and Kedr3 are characterized by the population of spiders with different biotopic preferences.
Both typical forest dwellers (Alopecosa accentuata (Latreille, 1817), Xerolycosa nemoralis (Westring,
1861)) and species typical of open biotopes – Pardosa agrestis, Pardosa bifasciata (C.L. Koch, 1834), and
Xerolycosa miniata (C.L. Koch, 1834) were encountered here. The control site is inhabited by species found
on dumps. The most numerous are Alopecosa cuneata (Clerck 1757), Xerolycosa miniata, and Trochosa
Ecologica Montenegrina, 61, 2023, 68-87 77ARTHROPODS ON OVERBURDEN DUMPS AFTER COAL MINING
terricola Thorell, 1856.
3. Myriapoda
Class Chilopoda is a universal predator, leading a hidden lifestyle, plays an important functional role in the
trophic webs of soil animal communities in both natural and anthropogenic habitats (Wolters & Ekschmitt
1997; Kunah 2013; Klarner et al. 2017).
In the study sites, 8 species of this class were noted. Directly on dumps (Kedr1 and Kedr2), only
Lithobius sibiricus Gerstfeldt, 1859 was caught in pitfall traps; this is a trans-Siberian species distributed in
the Asian part of Russia and northern Mongolia (Nefediev & Farzalieva 2020; Nefediev et al. 2020b). As the
dumps are restored, an increase in the average dynamic density of Chilopoda is observed from 0.14 ± 0.6
indiv. / 10 trap-days in 2013 to 0.32 ± 0.57 indiv. / 10 trap-days in collections of 2022.
Lithobius ostiacorum Stuxberg, 1876, prevails in collections at the Kedr3 site. This is a Southern
Siberian boreal species, also found in Northern Mongolia (Nefediev et al. 2018; Nefediev et al. 2020a).
The control site (Kedr4) is characterized by the highest species richness of Chilopoda (7 species).
The collections are dominated by Lithobius ostiacorum (50.7%), Central Siberian L. vagabundus Stuxberg,
1876 (22.4%) and L. nordenskioeldii Stuxberg, 1876 (14.9%). L. sibiricus, L. princeps Stuxberg, 1876, and
L. curtipes C.L. Koch, 1847 were found singly. The species affiliation of one species (Lithobius sp.n.?) is
difficult until the study of additional material of specimens with similar diagnostic characteristics.
Millipedes (Diplopoda) are active saprophages involved during the decomposition process of plant
remains. In the study area, the habitat of 5 species of this class was previously established. The population of
millipedes of dumps was represented by the only species Altajosoma sp., which was also recorded at other
sites. Orinisobates sibiricus (Gulička, 1963), inhabiting the forest litter of small-leaved and mixed forests,
was also noted at the foot of the dump (Nefediev et al. 2018). The ecologically plastic species Altajosoma
bakurovi (Shear, 1990) dominated in this area (73% of the total collection of millipedes in this area). In the
control zone, Sibiriulus profugus (Stuxberg, 1876) (76.5%) dominated in the collections. In addition,
Schizoturanius clavatipes (Stuxberg, 1876) was recorded here. Both species live mainly in different types of
forests: small-leaved and mixed forests, and dark coniferous and black taiga (Nefediev et al. 2021).
4. Oniscidea
Woodlice (Crustacea, Isopoda, Oniscidea) are terrestrial crustaceans that are actively involved in soil
formation by processing plant litter (Aleksanov 2016). Only Trachelipus rathkii Brandt 1833, one of the
most widespread species of woodlice in Russia, was recorded in the study area (Kuznetsova & Gongalsky
2012). The highest dynamic density of this species was noted in the Kedr3 site, 4.78 ± 7.55 indiv. / 10 trap-
days, occasionally found in open dump areas.
Discussion
The newly formed technogenic landscapes, which include rock dumps, are a conditionally free space from
life with a set of new diverse ecological niches. The first "invaders" of these territories are microorganisms
(bacteria, unicellular algae), then vascular plants are colonized, and almost simultaneously with them, the
colonization of the animal population occurs, including various groups of arthropods (soil and ground)
(Mordkovich & Lyubechanskii 2019).
To occupy free ecological niches, all living organisms adapt to local environmental conditions,
competing for trophic levels with other species. It is the consistent formation of stable trophic relationships
that plays a key role in the post-technogenic rehabilitation of disturbed landscapes. In terrestrial ecosystems,
there are two food webs – grazing and detrital, the interaction of which ensures their functional unity (Shurin
et al. 2006). After the species is fixed in technogenic biogeocenoses, reproduction begins, its abundance
increases, and this species spreads in the biocenosis (Lasky 2019; Koffel et al. 2022).
Self-restoring technogenically disturbed territories, as well as those on which forest reclamation has
been carried out, represent a mosaic, horizontal habitat heterogeneity, with specific microclimatic conditions.
This heterogeneity of the microenvironment has a positive effect on the species diversity of invertebrates
(Topp et al. 2001; Šálek et al. 2010).
Different research focused on the restoration of soil fauna on overburden dumps after coal mining.
Significant data have been collected from the coal regions of Germany (Dunger et al. 2001; Topp et al. 2001;
78LUZYANIN ET AL.
Dunger & Voigtländer 2009) and the Czech Republic (Frouz et al. 2006; Roubíčková et al. 2013). In
addition, the succession of communities of soil invertebrates is considered in the technogenically
transformed territories of Russia (Luzyanin & Eremeeva 2018; Mordkovich & Lyubechanskii 2019;
Luzyanin & Blinova 2022), Brazil (Toso et al. 2020), Poland (Madej & Kozub 2014; Schwerk 2014), Italy
(Ottonetti et al. 2006), and other countries. The authors note that in the newly formed territories, a peculiar
population of soil invertebrates is formed, in which eurybiont species predominate, penetrating dumps from
adjacent slightly disturbed cenoses. It has been established that among the community of terrestrial
invertebrates inhabiting technogenic landscapes, in terms of species diversity and abundance, representatives
of the orders of spiders and Coleoptera stand out first of all. It is also noted that successions of vegetation
cover in post-technogenic territories affect the structure of arthropods communities.
At the dumps of the Kedrovsky coal mine, we found a ground-dwelling of arthropods complex
consisting of various taxonomic groups. The noted invertebrates belonged to zoophages (for example, many
ground beetles), saprophages (woodlice, carrion beetles, carpet beetles), phytophages (pill beetles, some true
bugs), and animals with labile feeding behavior (polyphages) (various species of beetles).
Our studies have shown that almost all groups of ground-dwelling of arthropods are represented at
the initial stages of recovery, which is the same as in the control group. The initial colonization of arthropods
occurs very quickly during the first years after restoration. Such a high diversity of taxa may indicate
successful restoration of the ecosystem (Majer 1989). This can be confirmed by the detection of ground
beetle and carrion bettles larvae, and some other beetles species in our collections from dumps, despite the
method unsuitable for their collection and accounting (pitfall traps). In addition, juvenile forms of spiders
and centipedes have been registered, while there is evidence that very young juveniles Lithobius sibiricus
have a small migratory activity, in contrast to imaginal forms (Dunger & Voigtländer 1990).
The biodiversity and abundance of the considered taxa are not the same at different stages of
restoration. Young sites are dominated by species that prefer open habitats and are also well adapted to
anthropogenic impacts (Luzyanin & Blinova 2022). Thus, for example, the previously noted dominant
species of ground beetles Calathus erratus and Amara communis are widespread in the urbanized cenoses of
the city of Kemerovo (Eremeeva & Efimov 2006). The mass species of Chilopoda, L. sibiricus, also has a
high ecological plasticity and easily adapts to living in anthropogenic habitats (Nefediev et al. 2020a). The
species of woodlice that we captured, Trachelipus rathkii, is considered as a eurytopic species adapted to
disturbed biotopes (Ferenţi & Dimancea 2012).
Post-mining sites are often quite dry, which provides a good opportunity for colonization by
meadow-steppe and steppe species, which we observed in our study. At the same time, as noted above, the
mosaic nature of the habitat in these areas contributes to the emergence of microstations with meso- and
hygrophytic conditions. These areas are inhabited by invertebrates, which are demanding for medium and
high soil moisture. For example, we found the dominant species of ground beetles Bembidion gilvipes and
Trechus secalis on reclaimed dumps, which are confined to floodplain-coastal biotopes in the Kemerovo
region (Luzyanin et al. 2022). Such sites with high humidity occupied by various moss synusia, which retain
moisture and create favorable conditions for the habitat of these species.
Soil characteristics of post-technogenic territories are of great importance in the succession of biota
(Ufimtsev 2017; Kupriyanov et al. 2021). Soil parameters such as nutrient concentration, pH, and moisture,
affect the species richness of invertebrate communities (Frouz et al. 2006). For example, a sharp decrease in
soil pH reduces the diversity and abundance of soil fauna taxa. More basic soil is a favorable factor for
pedobionts (Lavelle et al. 1995; Manu et al. 2022). In general, our studies are consistent with these findings.
We can add that the dynamic density of ground-dwelling arthropods and pH values are negatively correlated
with each other.
Soil moisture has a noticeable effect on the distribution and abundance of some species of ground
beetles (Kagawa & Maeto 2014) and rove beetles (Irmler & Gürlich 2007). Ansgar Poloczek et al. (2021)
note that the occurrence of Lithobius ostiacorum also depends on the moisture content of the substrate
(positive correlation), which is typical for forest cenoses, in contrast to open habitats. In addition, there is
evidence that Trachelipus rathkii lives at air humidity close to 100% (McQueen 1976).
A significant improvement in environmental conditions occurs when a dump of potentially fertile
rocks, i.e., loose Quaternary sediments (loess-like and cover loams and clay) and/or a fertile soil layer, is
applied to the surface of the rock mass. They have more favorable agrophysical and agrochemical properties
compared with the substrates of rock dumps (Belanov & Androkhanov 2013). Together with soil ground, soil
invertebrates can get to dumps; therefore, the expansion of soil fauna on such substrates proceeds faster than
Ecologica Montenegrina, 61, 2023, 68-87 79ARTHROPODS ON OVERBURDEN DUMPS AFTER COAL MINING
in more gravel ones (Frouz et al. 2007). Potentially fertile rocks were deposited on our model rock dumps,
which improved the conditions for the restoration of entomocomplexes, which is confirmed by studies that
were conducted on other coal mines in the Kemerovo region (Bespalov 2014; Bespalov & Androkhanov
2019).
The development of the vegetation cover has an important influence on the soil fauna. For example,
Babin-Fenske & Anand (2010) note that an increase in plant diversity does not necessarily increase terrestrial
insect diversity. We found positive correlations between the projective cover of plants and the dynamic
density of centipedes (especially millipedes), woodlice, and some beetle families (ground beetles and rove
beetles). As shown in Table 1, the projective vegetation cover increases as the dumps are restored. Not only
herbaceous vegetation, but also trees and shrubs are actively developing on these sites. This is accompanied
by the arrival of a large amount of leaf litter, deadwood, which are food for saprophages. Many soil
arthropods are involved in the destruction of the litter, accelerating the decomposition and formation of
humus (Dickinson & Pugh 1974). This is consistent with our studies, in which we observed an increase in
the dynamic density of centipedes and wood lice when moving from open areas of dumps to more forested
areas, which indicates their preference for forest litter. Some authors (Simmonds et al. 1994; Hendrychová et
al. 2012) indicate that the thickness of leaf litter has a significant impact on the species richness and
abundance of some terrestrial species (for example, ground beetles, spiders), since more prey is concentrated
in the litter, fluctuations in temperature and humidity are reduced, and there are also shelters from predators
(Uetz 1979).
Pine litter is a very unsuitable food for most soil saprophagous animals due to its hardness and
chemical composition. This significantly slows down the decomposition of the litter (Voigtländer &
Balkenhol 2006). The type of reclamation process is the same in all natural and climatic zones of Kuzbass
(planting pine, cedar) gives a low soil-ecological effect, since such reclamation forms soils in which there is
no profile differentiation to genetic horizons (Dvurechenskiy 2012; Dvurechenskiy & Seredina 2015). We
recommend using fast-growing hardwoods (birch, poplar) for forest reclamation of post-technogenic areas in
the Kemerovo region. In our opinion, this will significantly accelerate the flow of organic matter into
embryozems, which will improve the processes of soil formation and leads to the development of more
productive and economically valuable landscapes.
Conclusion
The conducted studies allowed us to obtain detailed information about the communities of ground-dwelling
arthropods in reclaimed rock dumps formed after coal mining. In the first years of their existence,
technogenic landscapes are rapidly colonized by various groups of invertebrates, among which ground
beetles and rove beetles dominate in the ground layer. We see that some species easily adapt to the existing
conditions at the initial phase of succession. Gradually, during biota recovery, qualitative and quantitative
changes occur in the structure of ground-dwelling arthropods communities. For example, dominant species
change, and the overall population growth, which was noted in the first decades of succession, gradually
fades away, and self-regulation occurs at the ecosystem level. The general trend in the restoration of post-
technogenic territories is directed toward the establishment of communities of the zonal vegetation type. This
study shows that communities of ground invertebrates are good indicators for monitoring the restoration of
technogenically disturbed lands. The results of the research can be useful in developing a plan for the
reclamation of such areas.
Acknowledgments
We thank our colleagues for their help in the species identification of various groups of ground-dwelling
arthropods: ground beetles – Roman Dudko (Institute of Animal Systematics and Ecology, Novosibirsk,
Russia), rove beetles – Alla Dietz (Institute of Biology, Komi Scientific Centre, Ural Branch, Russian
Academy of Sciences, Syktyvkar, Russia), spiders – Laimonas Triliskauskas (Institute of Animal
Systematics and Ecology, Novosibirsk, Russia), myriapoda – Pavel Nefediev (Altai State University,
Barnaul, Russia), woodlice – Konstantin Gongalsky (Institute of Problems of Ecology and Evolution named
after A. N. Severtsov RAS, Moscow).
80LUZYANIN ET AL.
Funding
This research received funding from the Russian Science Foundation (RSF) grant 22-24-20014, and a grant
from the Kemerovo Region – Kuzbass, agreement No. 07 dated 03.23.2022.
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