meadows (Kuznetsova and Sterzynska, 1995); forests exposed to industrial ... botanical gardens, could equal those of natural commu- nities (Table 1, Fig. 1).
Russian Journal of Ecology, Vol. 34, No. 4, 2003, pp. 248–254. Translated from Ekologiya, No. 4, 2003, pp. 281–288. Original Russian Text Copyright © 2003 by Kuznetsova.
New Approaches to the Assessment of Structural Organization of Communities in Springtails (Hexapoda: Collembola) N. A. Kuznetsova Moscow State Pedagogical University, ul. Kibal’chicha 6, korp. 5, Moscow, 129243 Russia Received May 22, 2002
Abstract—Trends in the diversity of structural organization of soil animal communities are analyzed using an example of the taxocenes of springtails, small soil arthropods. Different methods for describing the structure of communities are considered. On the basis of parameters characterizing specialization and stability of communities, several patterns of their organization in springtails are distinguished. It is shown that the new approach offers new opportunities in forecasting community dynamics under the effects of natural and anthropogenic factors. Key words: pattern organization of communities, specialization, stability, springtails.
Historically, several stages of research into the structure of soil fauna can be distinguished. Studies performed in the early 20th century and dealing with the attempts to describe the sets of species characteristic of different habitats can be regarded as the first stage. At the second stage (the mid-20th century), researchers began to analyze the species structure, seasonal dynamics, and spatial heterogeneity of soil animal communities. Thereafter, attention was focused on the classification of communities based on identification of their types and associations by the criteria of the most numerous or typical species (Kuznetsova, 1988; Ponge, 1993). To date, specialists have accumulated a great amount of information on these communities, which requires new approaches to its interpretation. In particular, this concerns the analysis of their structural diversity, which may be the subject matter of research in this field at the fourth stage. Studies on springtails have shown that their natural communities (taxocenes) are multispecific groups organized on the basis of resource partitioning and competition between species, and their dynamics are generally predictable (Takeda, 1987; Chernova et al., 1989; Hägvar, 1994; Bengtsson, 1994). However, springtails inhabit virtually all soil types in all natural zones (Petersen and Luxton, 1982) and successfully colonize even severely disturbed habitats. Apparently, the communities developing in such a broad range of biotopes must differ in their organization pattern, as well as in species composition. The purpose of this work was to find adequate methods for revealing possible trends in the structural diversity of springtail communities.
MATERIAL AND METHODS To accomplish this purpose, it was necessary to consider springtail communities formed in a broad spectrum of habitats, from natural to anthropogenically transformed and artificial. The data generalized in this work concern the following habitats: (1) coniferous forests growing along a gradient of soil moistening (Kuznetsova, 1988; Kuznetsova and Krest’yaninova, 1998; Chernova and Kuznetsova, 2000); broadleaf forests (Kuznetsova, 1994); meadows (Kuznetsova and Sterzynska, 1995); forests exposed to industrial pollution (Kuznetsova and Potapov, 1997); urban green areas, from park forests to lawns and single trees growing in the openings in pavement (Kuznetsova, 1994, 1995; Krest’yaninova and Kuznetsova, 1996; Sterzynska and Kuznetsova, 1997); and (7) municipal waste dumps (Bugrov et al., 1996). Studies were performed in the southern taiga and broadleaf–conifer forest subzones of Eastern Europe (Russia, Belarus, and Lithuania). All samples were taken in 10–20 replications using the standard procedure. Springtails were collected using Thullgren funnel extractors. The species accounting for more than 12.4% of the total springtail abundance on the scale proposed by Engelmann (1978) were regarded as dominant. The species that were dominant both in each series of samples and in the total samples collected over the study period were named main dominants (Chernova and Kuznetsova, 2000). RESULTS AND DISCUSSION Parameters of species diversity. The generally accepted view is that species diversity is high in natural
1067-4136/03/3404-0248$25.00 © 2003 MAIK “Nauka /Interperiodica”
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Table 1. Parameters characterizing species diversity of springtail communities in different types of habitats (S is the number of species in the series of samples, H' is the Shannon index, and ND is the number of dominant species) Habitat Bilberry spruce forests Wood-sorrel spruce forests Sphagnum pine forests Lichen pine forests Oak forests Linden forests Black alder forests Meadows Forest parks and botanical gardens City gardens Boulevards and lawns Isolated trees Municipal waste dumps
Number of S
H'
ND
16613 10531 2382 3067 6227 2039 2326 1232 20704
30 (25–35) 30 (27–32) 29 (14–36) 22 (19–25) 26 (17–33) 18 (14–21) 27 (19–38) 12 (6–18) 22 (16–32)
3.20 (2.50–3.50) 2.78 (2.70–2.85) 3.83 (3.55–4.20) 2.85 (2.55–3.30) 3.12 (2.76–3.37) 2.88 (2.65–3.16) 3.28 (2.82–3.64) 2.32 (0.75–3.40) 2.94 (2.22–3.82)
2–4 1–3 1–2 2 1–3 2–4 3–4 1–3 2–3
4532 2979 9282 3618
21 (19–24) 19 (13–24) 15 (2–18) 9 (2–15)
2.84 (2.16–3.60) 2.77 (2.02–3.42) 2.36 (0.3–3.21) 1.58 (0.33–2.53)
1–3 2 1–3 1–3
biotopes
samples
individuals
4 3 4 3 5 5 4 11 12
68 48 59 45 94 110 53 176 376
4 6 6 6
149 149 469 61
communities and low in anthropogenic communities. In springtails, however, a number of exceptions to this rule were observed. Thus, the Shannon index of diversity for the springtail communities of wood-sorrel spruce forests, which are most abundant and rich in species, averaged only 2.78 bit. Conversely, species richness and diversity in the communities of anthropogenic biotopes, such as park forests and plantations in botanical gardens, could equal those of natural communities (Table 1, Fig. 1). These data confirm the hypothesis that the highest diversity is characteristic of communities existing under conditions of environmental disturbances of medium strength (Connell, 1978). Number of dominants. Both natural and anthropogenically disturbed communities of springtails can be mono- or polydominant. Thus, in the springtail communities of natural forests, dominance belonged to one species (bilberry pine forest, the Darwin Nature Reserve), two species (wood-sorrel spruce forest, Moscow oblast), and three species (green moss spruce forest, Moscow oblast), with this pattern remaining unchanged from year to year (Chernova and Kuznetsova, 2000). In urban cenoses, springtail communities included one to four dominant species (Table 1). Rank distribution of species. In the communities markedly differing in their organization, the curves of rank distribution of species were often virtually identical (Fig. 2). Connectedness. This parameter, which characterizes actual interspecific interactions, is very important for the assessment of community organization (Margalef, 1992). To estimate it for the communities of organisms belonging to the same trophic level, a matrix of correlations between the parameters of abundance of RUSSIAN JOURNAL OF ECOLOGY
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individual species is calculated, with the significant values of correlation coefficients (both positive and negative) regarded as indicators of the existing interspecific connections. On this basis, connectedness is calculated as described (Mikhailovskii, 1983; Litvinov and Panov, 1998). This approach to the study of communities in springtails has not yet produced any conclusive results: Correlation analysis of 57 springtail communities in conifer forests showed high spatio-temporal lability of interspecific connections in this group (Kuznetsova, 1987). Stability. Our studies have shown that structurally stable and fluctuating communities can be distinguished in springtails (Chernova and Kuznetsova, 2000). In the former, the relative abundance of dominant species changes from year to year by a factor of no more than 3, and its variation coefficient is below 50%. Among the latter, there are two variants: weakly fluctuating communities with a set of dominants limited to a few species, whose ratio can markedly change from year to year, and strongly fluctuating communities with a large and variable set of dominants. The springtail communities of mesophytic conifer forests are usually of the stable type; in dry, overmoistened, and disturbed forests, meadows, and urban areas, different variants of the fluctuating types are found (Table 2). There is also an interesting but relatively rare variant of quasi-stable communities, which retain their general structure irrespective of anthropogenic impact (Kuznetsova and Potapov, 1997). Specialization. Most communities include four categories of species: (1) the species specialized to live in 2003
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Number of species S 40
Shannon index, bit H' 4
30
3
20
2
10
1
0
1
2
0
3
1
2
3
1
2
3
(b) 4
30
3
20
2 10 0
1 1
0
3
2
Fig. 1. Changes in (S) species richness and (H') diversity of springtail communities along the gradients of anthropogenic influences: (a) industrial pollution of bilberry pine forests: (1) weak, (2) moderate, and (3) heavy; (b) urbanization: (1) forests and meadows; (2) park forests, botanical gardens, and city gardens; (3) boulevards, lawns, and isolated trees growing in the openings in pavement.
a given type of habitat, (2) the species specialized to live in other natural biotopes, (3) eurytopic species, and (4) the species characteristic of disturbed habitats (Table 3). The greater the proportion of specialized species, the higher the specialization of the community as a whole. The relative abundance of all biotope-specific species (forest species in the forest, meadow species in the meadow, forest and bog species in a forest bog, etc.) 1000
100
may be used as a measure of community specialization. With respect to the prevalence of a certain biotopespecific group, four categories of springtail communities can be distinguished: specialized, eurytopic, mixed, and ruderal communities. The specialized communities are characterized by the dominance of the corresponding species group (the conventional threshold is 40% of the total abundance) or, in some cases, two groups (e.g., the groups of forest and bog species in a pine–sphagnum bog). They are typical of the majority of natural forests, especially coniferous. In the eurytopic springtail communities (in most meadows of the forest zone, some broadleaf forests, and park forests), the eurytopic group prevails, and none of the specialized groups reaches the 40% abundance threshold. The modular communities include many ruderal or compost species, in addition to specialized and eurytopic species. For example, they are characteristic of most soils found in urban green areas. The communities with the prevalence of ruderal or compost species are classified as ruderal. They have been found in mounds of earth near construction sites (Malysheva and Chernova, 2000), piles of decaying plant residues (Chernova, 1977), and municipal waste dumps (Bugrov et al., 1996). Selecting parameters for characterization of community structure in springtails. The parameters commonly used by ecologists for characterizing the structure of communities are only partly applicable to springtails. Thus, the parameters of species diversity that have been successfully used, for example, in hydrobiological research (Alimov, 2000) are insufficiently informative in the case of springtail communities: their values are often similar in natural and disturbed habitats (Table 1) and strongly vary in the dynamics of the same community (Kuznetsova and Potapov, 1997; Chernova and Kuznetsova, 2000); they are also unsuitable for bioindication (van Straalen, 1998). The same applies to mono- and polydominance. 1000
1
100
2 3 4
10
1
10
1
3
5
7
9 11 13 15 17
1
1
5
9 13 17 21 25 29 33 37
Fig. 2. Curves of rank distribution of springtail species with respect to their abundance (the X axis shows the numbers of species arranged in order of decreasing abundance, and the Y axis shows the logarithm of abundance): (1) soil under an isolated tree (C) growing in an opening in the pavement in the center of Moscow (1993; 70 samples, 1136 springtails); (2) lichen pine forest in the Darwin Nature Reserve (1981–1983; 60 samples, 914 springtails); (3) city garden in Kolomenskoe, Moscow (1991–1992; 40 samples, 1685 springtails); and (4) sphagnum pine forest in the Darwin Nature Reserve (1981–1983; 60 samples, 1696 springtails). RUSSIAN JOURNAL OF ECOLOGY
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Table 2. Interannual variation in the proportion of main dominants in springtail communities from different biotopes (max/min is the ratio of maximum and minimum values of species abundance; V is the coefficient of variation in relative abundance by years, V = σ/x 100%. The range of values is shown when the number of main dominants is more than one) Community pattern with respect to temporal variability
Number of years
Max/min
V, %
5 5 5 3
2.0–2.7 1.4 1.5–2.9 1.4–2.8
32 12 17–41 14–46
3
1.2
10
5 3 5 3
9.8–13 1.4–30 2.5–9.4 3.5–25
60–99 19–85 36–108 40–75
5
3.9–9.5
60–68
3 3 3
2.4–41 1.4–62 3.2–38
28–67 11–122 37–71
Stable: wood-sorrel spruce forest bilberry pin forest green-moss spruce forest linden forest Quasi-stable: bilberry pine forest exposed to heavy pollution Weakly fluctuating: lichen pine forest bilberry pine forest exposed to moderate pollution upland herb–grass meadow boulevard Strongly fluctuating sphagnum pine forest isolated trees growing in the openings in pavement tree A tree B tree C
Table 3. Average percent ratios of biotopic springtail groups in different habitats, M ± m (a plus sign shows that the proportion of a species is less than 0.1%; the values exceeding 40% are boldface) Biotopic group Habitat
Pine–sphagnum bogs Conifer forests Broadleaf forests Park forests Meadows City parks and lawns Isolated trees Municipal waste dumps
Number of plots 4 19 14 12 11 12 8 6
specialized bog
forest
meadow
22.1 ± 8.7 + 0.3 ± 0.4 1.7 ± 2.6 0.7 ± 1.1 + 0 0
70.5 ± 11.1 64.3 ± 12.3 41.0 ± 11.7 27.8 ± 10.2 6.7 ± 4.2 4.4 ± 3.4 1.0 ± 0.8 3.4 ± 3.4
+ 0.6 ± 0.9 5.1 ± 5.0 7.4 ± 3.7 25.1 ± 12.6 24.5 ± 12.3 21.8 ± 11.6 6.3 ± 6.0
Species distribution by ranks in springtail communities requires further investigation, although several interesting generalizations have already been made (Hägvar, 1994; Berg and van Straalen, 2000). Connectedness is a promising index, but experience in its application to multispecific communities of the same trophic level, especially soil communities, is insufficient. To describe community organization in springtails, it is feasible to use two parameters, specialization and stability. Specialization of a community, being a paramRUSSIAN JOURNAL OF ECOLOGY
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eurytopic
ruderal + compost
7.4 ± 3.5 35.0 ± 12.1 52.9 ± 11.0 52.2 ± 11.6 65.5 ± 12.6 28.6 ± 13.1 35.6 ± 10.7 15.8 ± 17.8
0.1 ± 0.1 0.7 ± 1.0 0.7 ± 0.8 10.9 ± 5.8 2.0 ± 2.9 41.7 ± 17.2 41.6 ± 19.0 74.5 ± 24.8
eter of its structural organization, can possibly reflect its functional efficiency. Laboratory experiments with litter decomposition have shown that the quality and amount of humic acids formed in the litter are higher when it is inhabited by a group of springtails from the same biotope, rather than by their complexes taken from other biotopes (Chernova et al., 1989). Thus, specialization and stability reflect the structural (and, possibly, functional) and temporal aspects of community organization. 2003
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Table 4. Characteristics of the organization patterns of springtail communities (an example of the taiga and broadleaf forest subzones) Organization pattern Parameter
stable specialized
fluctuating eurytopic
Set of dominants Constant Number of potential Several dominants Spectra of life forms Constant and biotopic groups Predictability of comHigh munity structure Prevailing biotopic Forest Eurytopic group Stages of succession Late Intermediate Significant environ- Stable moistening conditions, no mental factors anthropogenic disturbance
Habitats
Mesophytic Broadleaf conifer forests forests
specialized
eurytopic
mixed
ruderal
Changes with time Many Change with time Low Forest or bog
Ruderal–compost Intermediate Intermediate Initial Initial Unstable moiste- Weak anthropo- Heavy anthro- Disturbed soils, ning conditions, genic impact pogenic imaccumulations no anthropogenic pact of organic dedisturbance bris Xerophytic and Anthropogeni- Most urban Waste dumps, hygrophytic cally transforsoils under earth piles conifer forests med forests, up- lawns land meadows
Using the combination of these two parameters, six patterns of community organization in springtails can be distinguished (Table 4). Some theoretically possible patterns (stable mixed and stable ruderal patterns) have not been found in nature. The ruderal communities cannot be stable, as they are confined to the initial stages of succession and, hence, are short-lived. The modular communities are formed under hard-to-predict conditions of urban soils, in which one biotopic group of springtails can gain an advantage over the others depending on random events: compost species actively develop in the presence of organic debris; ruderal species, upon an increase in recreational load; forest species, in fallen leaves; etc. However, it is possible to achieve the artificial quasi-stability of these communities by maintaining constant environmental conditions (Krest’yaninova and Kuznetsova, 1996). It is noteworthy that eurytopic species usually prevail in the springtail communities of natural broadleaf forests of Eastern Europe. This unexpected fact may be regarded as evidence that the corresponding phytocenoses are far from reaching the climax state. The type (pattern) of community organization in springtails is determined by external factors. Among them, the stability of moistening and the type of habitat (conifer or broadleaf forest, meadow, etc.) are most significant for natural communities, and the degree of anthropogenic impact, for anthropogenic communities. Knowing these characteristics of the environment, it is
Eurytopic
None
possible to predict the organization pattern and properties of the corresponding springtail community (Table 4). The same organization pattern may be formed with the involvement of diverse species and life forms. In this respect, the proposed categories differ from the types of communities, assemblages, etc. distinguished by the concrete sets of dominant or characteristic species. For example, the springtail community of the stable specialized type in Eastern European conifer forests is characterized by the prevalence of either Isotomiella minor, or I. minor and Parisotoma notabilis, or both these species together with Folsomia fimetarioides; in the first instance, euedaphic life forms are absolutely dominant; in other instances, dominance belongs to both hemiedaphic and euedaphic forms (Chernova and Kuznetsova, 2000). Takeda (1987) described another example of a stable springtail community from a mountain pine forest in Japan. In this case, the set of dominant species is absolutely different (Folsomia octooculata, Mesaphorura yosii, Tetracanthella silvatica, etc.). In the communities of a fluctuating specialized type, the dominant species of springtails may differ even in the neighboring phytocenoses of the same region. In the Darwin Nature Reserve, for example, the dominant group in lichen pine forests consists of Anurophorus septentrionalis, Xenylla brevisimilis, Desoria hiemalis, and Protaphorura stogovi; in sphagnum pine forests, it includes Sminthurides spp., Isotoma viridis, Pachyotoma crassicauda, and some other species (Kuznetsova and Krest’yaninova, 1998). The composition of fluctu-
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(b)
100 80 60 40
Relative abundance, %
20 0
1
2
3
100
1
(c)
80
2
3
4
I
60 II 40 III
20 0
1
2
3
4
5
Fig. 3. Spectra of biotopic groups of springtails in habitats along the gradients of different environmental factors: (a) gradient of moistening: (1) lichen, (2) bilberry, and (3) sphagnum pine forests; (b) industrial pollution: (1) control, (2) weak, (3) moderate, and (4) heavy pollution; (c) gradient of urbanization: (1) linden forests, (2) park forests, (3) boulevards, (4) isolated trees in the openings in pavement, and (5) municipal waste dump; (I) specialized species, (II) eurytopic species, and (III) species characteristic of disturbed habitats.
ating communities of mixed and ruderal types is even more variable. Due to the “insular effect,” the urban soils under neighboring trees growing in the openings in pavement may be inhabited by different sets of springtail species with the local prevalence of any group of life forms, as well as with different dominant species (Kuznetsova, 1994). Taking into account the proposed patterns of community organization, it is possible to better understand and predict probable transformations of springtail communities under the effects of natural and anthropogenic factors. This is illustrated by the following examples. (1) Changes in the springtail communities of conifer forests along a gradient of soil moistening. In the zonal mesophytic forests (spruce forests and bilberry and wood-sorrel pine forests), the optimum moisture supply provides conditions for the formation of stable specialized springtail communities (Kuznetsova and Krest’yaninova, 1998; Chernova and Kuznetsova, 2000). When hydrologic conditions in the habitat deteriorate (insufficient or excessive moistening), these communities lose their stability and can be classified as fluctuating. Irrespective of differences in the species composition of communities, specialized forms prevail throughout the moistening gradient (Fig. 3a). RUSSIAN JOURNAL OF ECOLOGY
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(2) Changes in the springtail communities of conifer forests along a gradient of industrial pollution. As pollution increases, fluctuations of species composition in the initially stable springtail community become stronger, with specialized forest species being gradually replaced by eurytopic species (Fig. 3b). It is noteworthy that the heaviest pollution results in the formation of a new, eurytopic community with a stable pattern of annual dynamics. Here, we are dealing with quasi-stability, because it develops on the basis of continuous anthropogenic influence (Kuznetsova and Potapov, 1997) (3) Changes in springtail communities along an urbanization gradient. The springtail community of a natural linden forest is of the stable eurytopic type. The forest species of springtails have almost disappeared from the majority of green areas (parks, boulevards, and lawns), whereas the meadow, eurytopic, and ruderal or compost species are fairly abundant there (Fig. 3c). If green areas are properly cared for (regularly watered and protected from recreational overload), these modular communities change from year to year not as strongly as could be expected and can be classified as weakly fluctuating (e.g., the community of a boulevard; see Table 2). However, the soil under isolated trees growing in the openings in pavement is inhabited by strongly fluctuating communities of the 2003
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mixed type (Fig. 3c). The compost and ruderal species are dominant in waste dumps (Bugrov et al., 1966). Thus, transformations of springtail communities upon any deviations in environmental conditions from the optimum have the same general trend: these communities gradually lose their stability and specialization. ACKNOWLEDGMENTS The author is grateful to N.M. Chernova for valuable comments on the manuscript. This work was supported by the Russian Foundation for Basic Research, project no. 02-04-49063; the program “Leading Scientific Schools,” project no. 00-15-97 885; and the State Research Program “Biodiversity.” REFERENCES Alimov, A.F., Elementy teorii funktsionirovaniya vodnykh ecosystem (Principles of the Theory of Aquatic Ecosystem Functioning), St. Petersburg: Nauka, 2000. Bengtsson, J., Temporal Predictability in Forest Soil Communities, J. Anim. Ecol., 1994, vol. 63, pp. 653–665. Berg, M.P. and van Straalen, N.M., Does Resource Partitioning or Habitat Properties Determine Collembola Community ^
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