ISSN 1062-3590, Biology Bulletin, 2019, Vol. 46, No. 5, pp. 500–509. © Pleiades Publishing, Inc., 2019. Russian Text © The Author(s), 2019, published in Izvestiya Akademii Nauk, Seriya Biologicheskaya, 2019, No. 5, pp. 533–543.

Ground (Coleoptera, Carabidae) and Zoodiagnostics of Ecological Succession on Technogenic Catenas of Brown Coal Dumps in the KAFEC area ( Krai) V. G. Mordkovicha and I. I. Lyubechanskiia, * aInstitute of Systematics and Ecology, Siberian Branch, Russian Academy of Sciences, Novosibirsk, 630091 *e-mail: [email protected] Received March 4, 2019; revised April 15, 2019; accepted April 15, 2019

Abstract—The population of ground beetles on artificial catenas of dumps of brown-coal pits in (dump age of one month, seven years, and 25 years) was investigated. The material was collected in the eluvial, transitional, and accumulative positions of each catena. The species diversity and activity of ground beetles after 25 years of succession in all catena positions do not reach the state of natural communities. The rate and direction of succession of the ground taxocenes differed significantly depending on their relief position. The appearance of species typical of zonal forest–steppe ecosystems began after seven years of suc- cession. In the accumulative position, succession developed more slowly than in the eluvial and transitional positions, where ruderal carabid species were gradually replaced by meadow mesophilous, and then by mesoxerophilous species. After 25 years, the taxocene of ground beetles in the accumulative position became similar not to the herb, but to the wood communities of the forest–steppe.

DOI: 10.1134/S106235901905008X

INTRODUCTION Interest in the study of biota successions on dumps, in particular, of coal enterprises, arose in the 1960s The most important reason for the reduction of (Dunger, 1968). Data collected over half a century in biota diversity is the change in the “spatial matrix of different regions (mainly on brown coal dumps in East the habitat” (Margalef, 1992; Hanski, 2010). This is , the , the Soviet Union, and facilitated by the incorporation of man-made habitats the United States) have revealed a number of patterns inhabited by ruderal biota of depleted composition of universal ecological significance. In the territories with random cenotic organization and disordered occupied by dumps, ecosystems are formed, only dynamics into the landscape background of the biosphere partly resembling the natural ones, but preserving the (Parmenter and Macmahon, 1987; Dunger et al., 2001; zonal orientation of succession development. How- Hendrychov et al., 2012). Coal mining, which, according ever, they almost never reach the state of climax bio- to Rosstat data, from 2002 to 2013 increased by 37% geocenoses of the corresponding biome (Burykin, (http://www.gks.ru/bgd/regl/B08_14p/Main.htm), 1985; Titlyanova et al., 1993). makes a significant contribution to the distortion of The transformation of the basic properties of eco- the evolutionarily established system of habitats. The systems arising on the dumps is primarily due to the most negative environmental consequences result state and activity of the living phase, succession from coal mining using an open method, which is changes of vegetation and invertebrates, and the inten- accompanied by the destruction of the natural land- sity of the biological cycle. The soil forms more slowly scape and its replacement by extensive quarries and than biota. The qualitative composition of the techno- waste dumps with the burial of natural ecosystems zem exchange cations is determined primarily by the under the multimeter overburden. The technology of properties of the underlying rock, i.e., overburden. coal mining provides for land reclamation: technical, The process of humus accumulation dominates over i.e., leveling or terracing of the hilly-ridge relief of mineralization, but it develops very slowly: 0.5 mg of dumps, and biological, i.e., artificial sowing of grass or C/g per 1 g of the substrate in the first 30 years. Later, woody plants on technozems. However, in practice, the intensity of humus accumulation decreases five- land reclamation activities are usually either carried fold. A primitive binomial soil is formed with a thin out many years later or not at all. As a result, chaotic humus-accumulative horizon that lies directly on the relief changes and the formation of surrogate ecosys- parent rock. Artificial application of a fertile humus tems occur. layer (10–20 cm) to the dump surface does not restore

500 GROUND BEETLES (COLEOPTERA, CARABIDAE) AND ZOODIAGNOSTICS 501 the basic physicochemical properties typical of natural terized by a high level of species richness and abun- soils. The technozem remains a habitat with an dance and a wide range of environmental preferences. extremely short time of formation and erasure of soil In addition, it is the most numerous component of the properties (Titlyanova et al., 1993; Topp et al., 2001; zooedaphon, confined to the soil surface and its upper Shugalei, 2009). Nevertheless, the application of a fer- horizon, and therefore the most accessible to study. tile layer to the reclaimed surface accelerates the for- The study polygon was the dumps of the brown mation of vegetation and has a positive effect on the coal mines of KAFEC located within the - soil fauna (Frouz et al., 2007). , -Yenisei, and basins of the The number of species in the phytocenoses of coal Middle Siberian Plateau. Land rehabilitation at mining dumps, which is extremely low initially (2–4), KAFEC began in 1973. This process includes several grows with time. However, species that are character- successive stages. The first stage is the removal of istic of climax stages of succession, which appear, at least “overburden,” i.e., the rocks overlying coal seams, the once, from the first year of succession, rarely become the offloading and dumping of the rock of which leads to dominant community even after 20–30 years. But rud- the formation of the dump. The dumps are artificial eral species retain dominant positions in the commu- hills of light and medium loams mixed with aleurolite nity not only at the pioneering stages, but also after rubble, 15–25 m high, alternating with depressions of 10 years or more. The phytocenoses do not reach the relief between them. Physiologically, the terrain species richness of typical meadows or forests for 50 resembles the natural ridge-hollow relief of the forest– years (Brändle et al., 2000). During the succession of steppe latitude zone of Siberia. The second stage (rec- vegetation, the biological cycle becomes increasingly lamation) involves leveling the relief and forming a closed and the rate of exchange of total carbon and plakor raised above the rest of the terrain. At subse- nitrogen in the ecosystem slows down (Titlyanova and quent stages of reclamation, a 25- to 50-cm layer of Mironycheva-Tokareva, 1988). natural soil is poured over the parent rock. The final stage is biological rehabilitation, i.e., natural or accel- Of all living organisms, heterotrophs become erated regeneration of the vegetation cover on dumps. involved in the biological circulation of dumps faster than others (Dunger, 2008). Microarthropods In the 1980s and early 1990s, on the KAFEC terri- (springtails, oribatid mites) appear in the substrate of tory, a large team of biologists and soil scientists under dumps in the first days after dumping. Their number the guidance of A.A. Titlyanova conducted compre- and diversity increase multifold with the age of the hensive studies of the biodiversity and the biotic circu- dumps (Babenko, 1982; Stebaeva and Andrievskii, lation, in which one of the authors of this paper was a 1997). The data on the structure and dynamics of the participant. The results of these works were presented population of macroarthropods (mesofauna) are in the collective monograph “Successions and the much more limited. Unlike the protozoans, microar- Biological Cycle” (Titlyanova et al., 1993). For vari- thropods, and juvenile spider individuals that fall into ous reasons, materials on macroarthropods were not dumps by passive wind transport, macroarthropods included in it. This gap, although very late, is partly penetrate into the dumps actively. The most diverse filled in by this article. and abundantly represented in the dumps are beetles, The aim of this work is to study the succession of mainly staphylinids and ground beetles (Mordkovich the taxocene of ground beetles and to assess the and Kulagin, 1986; Dmitrienko and Shaimuratova, impact of hilly ridge terrain and catena organization of 1986; Mordkovich, 1994). It is assumed that omnivo- habitats on the α- and β-diversity parameters in the rous species are the most successful colonists (Par- dumps of coal mining, as well as to obtain answers to menter and Macmachon, 1987). The high species the following questions: diversity of beetles on reclaimed lands may be due to the diversity of habitats arising from the hilly-ridged How different are the composition and variety of relief of the dump. These micro-habitats can serve as a ground beetles in the technogenic ecosystems of the refuge for rare species, sometimes reaching high abun- Kansk-Achinsk fuel and energy complex (KAFEC) dance there (Brändle et al., 2000). As dumps over- from those in the natural ecosystems of the “parent” grow, the total number of beetles may experience sig- forest–steppe biome? nificant fluctuations against the background of a grad- Does the organization of communities of ground ual decrease in the degree of dominance of ruderal beetles on technogenic dumps show signs of classical species (Frouz et al., 2008; Luzyanin et al., 2015). ecological succession or is it an arbitrary fluctuation of However, the general patterns of the dynamics of the the diversity and abundance of randomly introduced beetle taxocene on reclaimed lands remain poorly species? studied and the features of beetle communities in the Does the taxocene development of ground beetles first days and weeks after the appearance of a dump attain a state characteristic of the climax stages of eco- are completely unexplored. logical succession in the forest steppe? Does the 20- Ground beetles (Carabidae) were chosen as a year period prescribed by the technological standards model group, because this group of beetles is charac- of planned recultivation (Motorina, 1986) suffice?

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How do the direction, speed, and order of succes- The tendency of formation and accumulation of sion shifts differ on different elements of the techno- humus corresponded to the direction of change of the genic relief and on dumps of different ages? vegetative cover. The concentration of organic carbon (Corg) and its stock in the substrate of dumps is two orders of magnitude less than in native forest–steppe MATERIALS AND METHODS soils. The accumulation of humus increases with the Studies of the soil population were carried out in age of the dump in all positions, and in the accumula- the summer of 1986 in naturally overgrown dumps one tive position of the 25-year-old dump, it reaches val- month, seven years, and 25 years of age, located inside ues close to natural meadow soils (Table 1) (Titly- a quarry on the territory of the Nazarovo coal mine, 10 anova et al., 1993). The stock of phytomass also km southwest of the town of Nazarovo, Krasnoyarsk increases with the age of the dumps. The total number krai. Within each dump, a chain of habitats was con- of species of higher plants on the 7-year-old dump is sidered: eluvial positions at the top of each hill, transi- 66 and on the 25-year-old, 79. In its eluvial and tran- tional positions on the slope and accumulative ones in sitional positions, there are associations of tall weeds the depression between the hills. A total of nine habi- with a huge biomass and high projective cover. A prim- tats in the dumps were examined. The selected ages of itive grass and rush community is formed in the over- the dumps correspond to the conditional stages of lev- moistened accumulative positions of the 7-year-old eling of the artificial relief and overgrowing of the sub- dump. On the 25-year-old dump, all three positions of strate, first with ruderal and then with characteristic the relief differ by the composition of the dominants. vegetation of natural meadows of the surrounding A community richer in meadow grasses is formed in zonal forest–steppe landscape. the accumulative position of the 25-year-old dump. Precipitation erodes friable loam dumps, displaces As a control for comparison with the dumps, two the thin fractions of the substrate down in the profile, typical natural landscapes of the Central Siberian for- and enriches reduced terrain elements with them. At est–steppe were selected in their surroundings: a the same time, there is an increase in the rubble con- steppe meadow in the eluvial–transitional position of tent of the summit substrate and especially of the hill a natural catena with meadow chernozem soil and forb slopes. By seven years, the height of the hill has and graminoid mesoxerophytic vegetation (Stipa pen- decreased from 25 to 22 m, and by the age of 25 years, nata, Poa pratensis, Onobrychis arenaria, Potentilla to 20 m. The slope steepness decreases from the origi- longifolia, Artemisia commutata) and a clay–pebble nal 40° to 35° in seven years and to 30° in 25 years. The beach in the floodplain of the Chulym River with removal of aleurite from the top and the slope of the hardly any higher vegetation. hill causes a decrease in the absorptive capacity in the In the habitats studied, ground beetles accounted initial period of soil formation and inhibits its develop- for 80–90% of the total species richness and abun- ment, whereas in the depressions of the terrain, it is dance of macroarthropods. The ground beetles were the reverse (Titlyanova et al., 1993). The profile of the accounted for using the method of soil traps (plastic dump can be considered a technogenic catena, which cups with a diameter of 6.5 cm). The censuses were is in the process of active formation, but already with performed by ten traps in each biotope over feve days differentiation of positions into eluvial, transitional, during the summer maximum of activity and accumulative. (July 21–27, 1986). During the period of mesofauna accounting, the For each biotope, the Menhinick’s species richness temperature of the upper 5 cm of soil was measured index was calculated. Its value is <1, corresponding to using Savinov’s soil thermometers and the moisture low species diversity with a high abundance of the pro- content of soil samples by weighing and subsequent dry- nounced dominant, and can be a sign of the primitive ing. The temperature difference in the different positions organization of a community that is in the pioneering of the catena was minimal (2–4°C) on the 7-year-old stage of succession. For a community of a forest– dump, and on the 25-year-old one, it increased to steppe biome at the middle stage of development, val- 10°C. The habitats of the accumulative positions of the ues of Menhinick’s index equaling 1–2 are character- dumps of one month of age turned out to be the cool- istic, and for climax communities, even higher values est, while the habitats of the eluvial positions on the (Mordkovich, 2017). Using cluster analysis, we stud- dumps of any age, but especially the 25-year-old one, ied the similarity of the populations of ground beetles were the warmest (Table 1). The soil moisture in the studied biotopes. The binary (Simpson’s coeffi- increased from top to bottom along the slopes of cient) and quantitative similarity coefficients (Dice dumps of any age (1.5 times on the 7-year-old and four index) were calculated. Clusters were constructed times on the 25-year-old dumps). The differences in using the unweighted pair group method with the the hydrothermal regime of soils in the eluvial and arithmetic mean (UPGMA) with the calculation of transitional positions were minimal in the first seven bootstrap coefficients (n = 1000). The calculations years of succession and doubled in the catena of were performed using the PAST V.2.17 program 25 years of age (Table 1). (Hammer et al., 2001).

BIOLOGY BULLETIN Vol. 46 No. 5 2019 GROUND BEETLES (COLEOPTERA, CARABIDAE) AND ZOODIAGNOSTICS 503 Legumes, grasses, and sedge Carex capitate, P. pratensis, C. epygeios, T. pratense 1993) , and grasses repens, E. Equisetum pratense, Trifolium pratense, millefolium Achillea et al. ions of the catena. The soil temperature and humidity The soil temperature of theions catena. E. repens, Poa pratensis, Calamagrostis epygeios Typha angustifolia, Typha E. repens, T. farfara, C. setosum Dump age* Dump T. farfara, T. vulgaris, A. setosum, C. Elytrigia repens of the KAFEC Nazarovo coal mine dumps (Titlyanova mine coal dumpsof the(Titlyanova Nazarovo KAFEC Tussilago farfara, Tussilago vulgaris, Artemisia Cirsium setosum 1 month (0) 7 years (66) 25 years (79) 0 0 0 1337 1006 1098 2339 1848 2860 71 – 213 435 225 825 1790 2200 3208 EL TR AC EL TR AC EL TR AC 14.1 13.3 25.2 18.7 18.4 28.1 8.9 18.8 36.4 23.5 20.5 19 27.5 26 25 30.7 28 20.6 2 C ° 2 Ecological characteristics of technogenic ecosystems content, % content, 0.11 –, g/m 0.33 0.75 0.38 1.38 3.58 5.50 8.02 Parameters org org С Dominant species g/m Phytomass, Communities Single plants Weeds Weeds Grasses and rush Grasses Forbs, legumes, Soil temperature Soil temperature cm), (0–10 Soil humidity % cm), (0–10 С Table 1. posit accumulative and (AC) (TR) transitional, plants. (EL) Eluvial, vascular of of species the(*) The number parentheses give are given for the time of sample collection (July 1986). (July of sample the time collection for are given

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(а) 30 123 150

20 100

10 50

0 0 1 month 7 years 25 years M

30 (b) 150

20 100

10 50

0 0 1 month 7 years 25 years M Dynamic density Species number, Menhinick’s index Menhinick’s number, Species

30 (c) 300

20 200

10 100

0 0 1 month 7 years 25 years S

Fig. 1. Change of species richness and dynamic density of ground beetles in the course of succession in the different positions of artificial catenas and in natural habitats. (a) Eluvial, (b) transitional, and (c) accumulative positions of a catena, (M) steppified meadow; (S) shallows. (1) Number of species, (2) dynamic density, ind./50 t.d., (3) Menhinick’s index multiplied by 10.

RESULTS Amara fulva. The accumulative position was domi- nated by the same species, but with the addition of In all the artificial and natural habitats studied, 56 subdominant Bembidion semipunctatum and species of ground beetles were identified, the total livida. dynamic density of which varied in different biotopes On the 7-year-old dump, overgrown with tall thick from a few to >100 ind./50 trap days (t.d.). Just a weeds by this time, the species diversity of ground bee- month after the rock was dumped, the species diversity tles slightly increased in the accumulative position, of ground beetles in the eluvial and transitional posi- 2.4 times in the eluvial position, and 4.4 times in the tions of the catena in the absence of higher vegetation transitional one, compared to the fresh dump (Fig. 1). reached 5–8 species, and the dynamic density was 63– A change of dominants took place. In the eluvial and 90 ind./50 t.d. (Fig. 1). In the accumulative position of transitional positions of the catena, along with the catena, on the border with the temporary reservoir, A. fulva, which continued to play the leading role, the diversity of ground beetles reached 11 species, and Bembidion femoratum, B. quadrimaculatum, Curtono- the dynamic density was 72 ind./50 t.d. The only dom- tus convexiusculus, C. aulicus, and Acupalpus meridi- inant in the eluvial and transitional positions was anus occupied equal positions. In the accumulative

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position community, Bembidion semipunctatum main- R tained the dominant position, a significant number of S 7/EL 7/TR 1/TR 1/EL 25/EL 25/T 1/AC 7/AC 25/AC B. quadrimaculatum, B. lunatum, and Asaphidion palli- M pes appeared along with it. 1.0 At the 25th year of succession, when graminoid and forb meadow vegetation took hold in the eluvial and transitional positions of the catena, lower species diversity (5–6) and dynamic density (up to 19 speci- 0.8 mens/50 t.d.) of ground beetles were observed (Fig. 1). Calathus erratus, Amara equestris, and Poecilus fortipes 85 95 became dominant. In the accumulative position of the artificial catena, by the age of 25, a hygrophyte meadow with sedge and reed was formed. The species 0.6 67 diversity of ground beetles was 1.5 times higher, and the dynamic density was 1.6 times higher than on the 7- year-old dump. Pterostichus niger, Trechus secalis, Cara- bus violaceus, C. henningi, and C. regalis were dominant. index Dice 21 0.4 17 The course of succession in the eluvial and transi- tional positions of the dumps was very similar up to 43 25 years. Even at the monthly stage of succession, the values of Menhinick’s index ranged from 0.5 to 1, and by the 7th year, they reached 2, which indicates a rapid 0.2 30 pace of succession development in the KAFEC habi- 22 tats. However, by the 25th year, the complexity of the 20 taxocene of ground beetles decreased again (Menhi- 100 nick’s index was 0.8–1). The succession of the population of ground beetles Fig. 2. Similarity of the ground beetle population of the from the accumulative position of the catena pro- KAFEC Nazarovo coal mine dumps according to the Dice ceeded more consistently. Here, Menhinick’s index index. Bootstrap coefficients are given at the graph nodes slowly but continuously increased from 1.3 to 1.7 from (n = 1000). (1) Dump age of one month, (7) dump age of seven years, (25) dump age of 25 years, (M) steppified the monthly stage to the 7-year stage and further to the meadow, (S) shallows. (EL) Eluvial, (TR) transitional, 25-year stage. and (AC) accumulative positions of the catena. In the control biotopes (steppe meadow for eluvial and transitional positions and river shoal for the accu- the positions in the “young” dumps. The eluvial posi- mulative one), the number of species and the dynamic tions between themselves, as well as the transitional density of ground beetles were lower and the Menhniki ones, have a similarity of 100% at low bootstrap coef- index was higher than at the intermediate stages of ficients (51–60). succession on the dumps (Fig. 1). Cluster analysis of the population of ground beetles When studying the distribution of the dominant of the studied catena positions showed that all bio- species of ground beetles by the positions of different topes were divided into two parts: “old” biotopes age catenas, we were able to identify species the num- (25 years), together with the natural biotope of steppe bers of which were significantly higher in some bio- meadows and “young” biotopes (one month, seven topes. The dynamic density of A. fulva reached years), along with the constantly updated biotope of 78 ind./50 t.d. in the transitional position of the fresh river shoals. The similarity of the dumps to the “refer- dump, B. semipunctatum was abundant in the accumu- ence” biotopes was small, which may indicate four lative position of the fresh and seven-year-old dump independent clusters (Fig. 2). The studied biotopes are (19 and 40 ind./50 t.d., respectively), as well as in the grouped mainly by the degree of closeness in the suc- shallows (38 ind./ 50 l-s). C. convexiusculus was char- cession age. The eluvial and transitional positions are acteristic of the transitional position of the seven- more similar to each other (and with high bootstrap year-old dump, and C. erratus, of the 25-year-old values) than the accumulative ones. A tree constructed dump (62 and ind./50 t.d., respectively). These species from the values of the Simpson coefficient, which can be considered as indicators of the corresponding takes into account only the presence or absence of a stages of succession and the geomorphological posi- species, is characterized by a high level of similarity in tion of the biotope.

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DISCUSSION beetles, stenobionts of the natural forest–steppe are among the dominants. A comparison of the results obtained with the avail- able data on springtails and oribatid mites reveals a The order, rate, and direction of differentiation of fundamental difference in the trends of diversity of the ground beetle population depend not only on the micro- and macroarthropods as the dumps age. age of the dump, but also on its position in the relief (Fig. 2). In the eluvial and transitional positions that The occupation of dumps of one month of age by become warmer and drier year by year, the ruderals oribatids and collembolans initially occurs due to a few (B. femoratum and A. fulva), completely dominating species, the number of which is small. However, later, on the monthly dump, are replaced with meadow the abundance of some of them (the oribatid mites mesophiles on the seven-year-old dump (B. quadri- Opiella nova, Tectocepheus velatus) grows unevenly, maculatum, C. convexiusculus, C. aulicus, and making them the dominant community (Babenko, A. meridianus). On the 25-year-old, the dump where 1982; Dmitrienko, 1990; Titlyanova et al., 1993; Ste- soil moisture in the eluvial and transitional positions is baeva and Andrievskii, 1997; Bogorodskaya et al. , halved compared to that in the monthly dump (Table 1), 2010). In a four-year-old dump, the recovery of the the mesoxerophilous species characteristic of mead- invertebrate complex occurs more slowly. The com- ows in the natural landscapes of the Siberian forest– munity is dominated by springtails and predatory steppe seize the dominant positions in the ground bee- mites (Bogorodskaya et al., 2010). The third stage of tles population (Mordkovich and Lyubechansky, microarthropod succession is observed in dumps of 2010): C. erratus, A. equestris, P. fortipes, Harpalus seven to ten years of age. It is characterized by a tran- , and . . In the accumulative posi- sition to a multispecific community while maintaining nigritarsis H modestus the dominance of all the same ruderals as in the previous tions, where the soil becomes wetter with age (Table 1), stages of succession. In even older dumps (20–30 years amphibian species (Nebria livida and B. semipunc- old), the species richness increases several times, but tatum), which occurred along the edge of a temporary the number of species is unstable; ruderal species still reservoir of a month’s age, on the seven-year-old dominate (Titlyanova et al., 1993; Wanner and dump replaced meadow hygrophiles A. pallipes and Dunger, 2002). B. lunatum and meadow mesophiles C. aulicus, C. convexiusculus, and Clivina fossor (Table 2). Thus, the microarthropods with their passive pho- resis and the need to develop directly in the substrate The appearance of single specimens of the forest of a fresh dump are characterized by a consistent species (according to their topical preferences) in the increase in species diversity and abundance over 25– accumulative position of the seven-year-old dump, 30 years. However, no complete equivalence with the seemingly an accident, turns into a trend on the 25-year- composition and structure of the population of any of old dump, when forest and forest edge species become the zonal communities of the forest–steppe (meadow, dominant in the ground beetle population (T. secalis, meadow–steppe, or park–forest) is observed C. violaceus, C. henningi, and C. regalis) (Mordkovich (Babenko, 1982; Titlyanova et al., 1993; Frouz et al., and Lyubechanskii, 2010), although there are no signs 2008; Hendriková et al., 2012). of afforestation in the soil or in the vegetation cover (Titlyanova et al., 1993). Thus, the forest taxocene of Successions of populations of ground beetles, rep- ground beetles develops earlier than the corresponding resenting the category of macroarthropods, occur dif- vegetation appears, and one can see the difference ferently. In populating a dump in the first hours after between the succession on the dumps and the one tak- the dumping, imagoes of beetles massively invade the ing place in the natural biocenoses of the forest– dump from the surrounding ecosystems and without steppe. Apparently, it is necessary to recognize ground delay are included in the biological circulation of the beetles as forerunners of some, as yet unidentified, pro- fresh dumps. This is facilitated by the migration, cesses and properties that predict the appearance of a for- which is, unlike that of microarthropods, carried out est-type ecosystem in the habitat in the near future. with the help of special adaptations (developed wings, chemoreception for wet earth, limbs adapted for The change in ground beetles species with different movement not only on a flat surface, but also in a topical preferences is determined by the fundamental loose substrate), and also polyphagy. A high number restructuring of the structural and functional organi- of ground beetles is observed on the middle-aged zation of the communities forming on the dumps. The dump, while on the 25-year-old dump, it decreases. greatest contribution to this process is made by ceno- Among microarthropods, even after 25 years, eurybi- phile species with a high abundance and clearly differ- ont species dominate in the composition of the popu- entiating the geomorphological positions and stages of lation of dumps, as well as on young dumps; in ground succession.

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Table 2. Distribution of ground beetles in the studied KAFEC habitats (ind./50 t.d.) Compared Catena position Species habitats N 1/EL 1/TR 1/AC 7/EL 7/TR 7/AC 25/EL 25/TR 25/AC M S Acupalpus meridianus 00014800000022 Agonum ericeti 000013005009 A. gracilipes 000010000001 A. livens 000000001001 Amara apricaria 7611100000016 A. bifrons 000110000002 A. communis 000000005005 A. equestris 00013011300018 A. fulva 4078311017000000176 A. ovata 001100000002 A. parvicollis 000000000033 A. spreta 000000000303 A. tibialis 000000000303 Anisodactylus signatus 000112000004 Asaphidion pallipes 001016000008 Bembidion articulatum 000000000044 B. femoratum 102051100000028 B. litorale 00000000005555 B. lunatum 003024000009 B. properans 000000000808 B. quadrimaculatum 202178100003143 B. semipunctatum 0019004000003897 B. striatum 00000000004646 Calathus erratus 00000022460032 Cal. melanocephalus 1002240050014 Cal. micropterus 000002000002 Carabus aeruginosus 000000003003 C. henningi 000000005005 C. regalis 000000007007 C. violaceus 000000008008 Chlaenius nigricornis 000000000033 Clivina fossor 0013020030817 Curtonotus aulicus 0009400010014 Cur. convexinsculus 120156240000084 Dyschirius obscurus 00000000002424 Dyschirius sp. 000000000055 Elaphrus riparius 000000000066 Harpalus affinis 001440000009 H. distinquendus 000010000001 H.modestus 000000100001 H. nigrans 000000710008 H. rubripes 0002140130011 H. rufipes 000020016009

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Table 2. (Contd.) Compared Catena position Species habitats N 1/EL 1/TR 1/AC 7/EL 7/TR 7/AC 25/EL 25/TR 25/AC M S Loricera pilicornis 000002000002 Microlestes minutulus 100200000003 Nebria gyllenhali 000000000055 N. livida 001100000001324 Poecilus cupreus 021010000004 P. fortipes 000210822648087 P. versicolor 0001200090012 Pterostichus eximus 00000000053053 Pt. melanarius 000000006107 Pt. minor 000000001001 Pt. niger 00000000440650 Synuchus vivalis 1004040030012 Trechus secalis 00000000150015 Total individuals 63 9 0 72 95 135 87 19 62 142 119 217 1101 in catena (1) Dump age of one month, (7) dump age of seven years, (25) dump age of 25 years, (M) steppified meadow, (S) shallows. (EL) Elu- vial, (TR) transitional, and (AC) accumulative positions of the catena. (N) The total number of individuals of one species.

CONCLUSIONS process does not require 12–18 years, adopted today as the standard time for reclamation, but much more. Over time, the differentiation of environmental According to our data, 25 years is not sufficient for this. conditions in the different positions of dump relief, negligible at one month of age, gradually increases. The catena type of organization of dumps and frac- ACKNOWLEDGMENTS tional differentiation of the environment in space and time form a multitude of habitats of macroarthropods. The authors are grateful to A.A. Titlyanova, N.P. Mironycheva-Tokareva, V.S. Andrievskii, N.P. Kosykh, The speed and direction of the taxocene succession G.D. Mordkovich, and N.A. Afanas’ev (Institute of Soil of ground beetles differ significantly depending on the Science and Agrochemistry, Siberian Branch, Russian position of cenoses in the relief. In the eluvial and Academy of Sciences, Novosibirsk) for cooperation in joint transitional positions of technogenic catenas, separate research, and to B.M. Kataev (Zoological Institute, Russian species characteristic of the zonal ecosystems of the Academy of Sciences, St. Petersburg) and I.A. Belousov forest–steppe appear seven years after the beginning (All-Russia Institute of Plant Protection, St. Petersburg) for of the succession. The conditions in the accumulative help in the identification of ground beetles. positions change the most rapidly and in the direction the most comfortable for soil invertebrates. The suc- cession develops consistently, but more slowly than in FUNDING the eluvial and transitional positions. The species diversity and dynamic density of ground beetles after This study was supported by the Fundamental Scientific 25 years of succession in all positions of a technogenic Research Program (FRP) of the state academies of sciences catena do not become identical with natural commu- for 2013–2020 (project no. VI.51.1.7. (0311-2016-0007) nities. In 25 years, a ground beetle taxocene in the (АААА-А16-116121410123-1)). accumulative position acquires similarity not with the herb, but with woody communities of the forest– steppe, in contrast to phytocenosis and the soil. COMPLIANCE WITH ETHICAL STANDARDS The characterized patterns of change in the taxo- Conflict of interest. The authors declare that they have no cene population of ground beetles, consistent in time conflict of interest. and space, can collectively qualify as ecological suc- Statement on the welfare of . All applicable inter- cessions, as they approach a state close to climax states national, national, and/or institutional guidelines for the of natural forest–steppe communities. However, this care and use of animals were followed.

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