Characteristics of vegetation and soils at studied plots in the Sysola River valley.
River floodplains are unique nature landscapes. In contrast to zonal communities on watersheds, soil biota of river floodplains is studied in less degree. The research was conducted in the floodplain forests in the European North‐East of Russia and showed high diversity of soil biota in alluvial forest soils. Floodplain forest soils are inhabited by 70 species of micromycetes, 53 genera of Nematoda, 60 species of Collembola, and 110 species of large invertebrates. Alluvial meadow soils with stable moisture and temperature conditions are characterised by high species diversity of micromycetes, nematodes and large invertebrates. Collembola prefer alluvial soddy soils. Soil microorganisms, meso‐ and macro‐fauna can essentially increase taxonomic diversity and number in alluvial meadow‐boggy soils at warming autumn.
- European North‐East of Russia
- Sysola River floodplain
- aspen‐birch forests
- alluvial soils
- soil microorganisms
- meso‐ and macro‐fauna
River floodplains are the most widely distributed habitats in the world, occurring from tropical to polar regions and from deserts to rainforests. Depending on their specific area of interest, various scientists view floodplains quite differently. Many ecologists perceive floodplains primarily as ecotonal extensions of river channels. Scientists who study plants and soils view floodplains as distinct habitats rather than simply ecotones between rivers and uplands. In fact, floodplains are a mosaic of sub‐habitats (some aquatic, some terrestrial, some wetland), and diverse sub‐habitats potentially supporting a unique biota. While most work focuses on the aquatic biota of floodplains, the terrestrial component is being increasingly recognized .
Due to annual snowmelt floods, river floodplains have a specific ‘terraqueous’ regime  which creates particularly special conditions for vegetation cover, soils and soil organisms. Duration and regime of floods respond for the fact that floodplain soils largely differ from watershed soils not only concerning morphological structure and physical‐chemical properties of soil profiles but also by life activity of soil biota which is involved into plant residues decomposition in terrestrial ecosystems. In contrast to zonal communities on watersheds, soil biota on river floodplains is less studied. In Russia, the role of soil biota on river floodplains was intensively studied in the 1960s–1980s of the twentieth century [3–6]. This was conditioned by the fact that floodplain soils were treated as to be used in engineering and agriculture. In the twenty‐first century, floodplain landscapes gained high attention in both Russia and European countries [7–12]. These studies aim at identifying the dependence in population number of soil organisms from flood regime [13–15]; adaptations of invertebrates to moisture deficient or excess [16–18]. Complete descriptions of soil biota in river floodplains are lacking, and existing research tends to focus on a few groups of soil organisms. It is known that floodplains contribute significantly to the biodiversity of the world because so many species occur solely in floodplains or at least rely heavily on floodplains to satisfy important ecological needs .
Globally, the soil invertebrate fauna has been extensively researched only at a few temperate‐zone floodplain ecosystems, mainly in the USA, Europe and Australia. The distinctive feature of European floodplains is that they have been transformed for centuries. As a consequence, 95% of riverine floodplains have been lost . But it is known that soil biota in floodplain communities has higher diversity indices as compared to that in zonal communities. And geologically ancient soils (Central Amazonia) with long‐term cycles of always alternating aboveground and water phases are inhabited with endemic invertebrates whereas young ecosystems (Central Europe)—with eurytopic species . Floodplain forest ecosystems are key habitats for rare invertebrates, including the representatives of the postglacial period . Overall, the knowledge of the soil biota in the northern floodplain communities remains incomplete and requires a sustained taxonomic and ecological research effort to provide better estimates of species diversity, distribution and evolutionary history.
River floodplains are ‘oases of life’ in the northern regions. Due to the warming effect of the river waters, highly productive grass‐forb meadows and deciduous forests with grassy ground cover, which atypical for the watershed landscapes of the northern part of the taiga zone, are formed in the valleys of the boreal rivers . At the same time, priority attention of ecologists has been paid to the identification of biodiversity and structure of soil biota in the coniferous forests occupied watersheds [22–24]. Alluvial soils of the northern river floodplains are studied fragmentarily in this respect, particularly soil biota of floodplain aspen‐birch forests [9, 10]. This was conditioned by the fact that morphological and physico‐chemical properties of alluvial soils as well as ways of preservation and increase of soils fertility of floodplain meadows were studied firstly at the European northeast of Russia. The high importance of river ecosystems in shaping migration flows of substances in landscapes, the role of floodplain soils as biogeochemical barriers to the migration of chemicals, the specifics of vegetation cover at the floodplain terraces of the boreal zone and the importance of floodplains in maintaining of soil biota biodiversity are the conditions which are important to identify the features of formation of not only soil and plant cover at floodplain landscapes of the North but also the biodiversity of soil invertebrates and microorganisms, playing a leading role in the transformation of plant residues in terrestrial ecosystems. In this study, we focused at the variation of the soil biota in a river floodplain system with natural hydrological conditions and well‐preserved forests that have not been modified by human disturbances. We assumed that a riverine landscape in the natural state exhibits a high level of complexity across a range of scales, which might contribute significantly to the species pool. So, the purpose of this study was to obtain new data about species diversity, number and structure of soil biota in the Sysola River floodplain located at the European North‐East of Russia and to identify ecological and functional interlinks between alluvial forest soils in the taiga zone and soil biota.
2. Materials and methods
The studies were conducted in the Sysola River valley, middle taiga, Komi Republic and European North‐East (Figure 1). The Sysola River (395 km long) is one of the largest left tributaries of the Vychegda River (1131 km) which, in turn, is a tributary of the Severnaya Dvina River (744 km), the White Sea basin. It is a typical plain river, which is occupied by meadows. Our researches were carried out in aspen‐birch forest, which is located in middle part of floodplain terrace, low course of the Sysola River. This forest was divided into three plots which take different positions in relief of floodplain and greatly differ by ecological conditions as snow‐melt water inundation period, ground water depth, soil type, plant composition in ground cover, etc. The plots form a natural ecological row along with increasing soil moisture content: Plot 1 (ridge top, high floodplain level) → Plot 2 (even part of floodplain, mean level) → Plot 3 (deep inter‐ridge depression, low level). Spring flood regime in the Sysola River is unstable. High ridge tops soils do not became inundated or stay under water for a short period of time (1–1.5 weeks), while inter‐ridge depressions sometimes stay under water for one and a half month or two months.
Morphological, physical‐chemical soil properties and their hydrothermal regime were studied in accordance with the accepted methods [25, 26]. Reference sections for the morphological description of soil horizons and sampling were laid at key plots. Names of soil types and horizons indices are given according to the Russian standards . Soil moisture content was identified gravimetrically, soil temperature—with an electronic transistor digital thermometer TET‐TS11 (Russia) and loggers DS1921G (Russia). Carbon content was measured by the gas‐chromatography method with CNHS‐analyzer (Carlo Erba, Italy), pHKCl—potentiometrically with glass and silver‐chloride electrodes at soil:solution ratio of 1:2.5 for mineral and 1:25 for organic horizons, hydrolytic soil acidity (Ha)—by titration using CH3COONa solution, exchangeable cations (Ca2+, Mg2+)—by driving with NH4Cl solution followed by atomic‐absorption identification at Hitachi 180–60, and texture—by the Kachinsky method with dispergation and boiling in the presence of NaOH.
Number of the principle ecologic‐trophic groups of microorganisms was assessed by inoculation of solid nutrition media . We identified concentration of ammonificators (beef‐peptone agar), oligonitrophilous (Aeshbi's medium), nitrifying (Vinogradsky's medium), and denitrifying (Giltai's medium) bacteria; microorganisms using mineral nitrogen compounds (starch‐ammonia agar); oligotrophic (starvation agar) and pedotrophic (soil agar) microorganisms. Saccharolytic microscopic fungi were assessed using acid Chapek's medium, cellulose‐decomposing fungi—Getchinson's medium, and oligotrophic fungi —starvation agar. Total microorganisms and micromyces number was stated in CFU/g a.d.s. (colony‐forming units per 1 g of absolutely dry soil). The microorganisms’ biomass carbon was estimated by the rehydration technique on the base K2SO4 extracts  in fresh samples of forest litter (A0, 0–3 cm deep) and humus horizon (A1, 3–15 cm deep). Samples were collected four times during vegetation period in threefold to fourfold replication. Taxa of micromycetes were identified after their extraction as pure cultures using Chapek‐Dox medium with help of manual books for the identification of different taxonomic groups of micromycetes, interactive keys, and information Internet site (http://www.indexfungarum.org).
For the evaluation of taxonomic composition and number of nematodes, soil samples (5 cm in diameter, 5 cm deep) were collected in sevenfold replication monthly from June till August. Totally, 63 samples were collected. Nematodes were extracted from soil using modified Bermann funnels, heat‐killed and fixed in 4% formaldehyde. In each soil sample, at least 100 individuals were identified to the genera level using a Leica DM4000 B inverted microscope. Nematodes were identified following the taxonomic keys [30–32]. The abundance of nematodes was recalculated per 100 cc of soil. Nematodes were assigned to six trophic groups (bacterivores, fungivores, root‐fungal feeders, plant parasites, omnivores and predators), according to classification . In total, 190 soil samples (5 × 5 × 5 cm) were collected for identification of Collembola. Samples at each plot were collected in fivefold replication monthly from June till September 2003–2005. Extraction of Collembola was done in Berlese‐Tulgren funnels. Quantitative accounting of large invertebrates was done by hand using of soil samples (25 × 25 × 5 cm). In total, 380 soil samples were collected by analogy to the sampling procedure of Collembola at the same time but in 10‐fold replication at every plot. Characterization of soil organisms was done using general ecologic indices as occurrence frequency, relative abundance of species (P, %), species richness (S), Shannon's diversity (H') and evenness (J') indices, Simpson's index of dominance (DSM), Chekanovsky‐Sjerensen index of similarity (Ics). The obtained data were processed by standard methods of statistics using Microsoft Excel, STATISTICA 6.0 and PAST 3.1.
3. Characterization of plant and soil cover
River valleys of the taiga zone have two forest types, particularly birch and aspen forests . From position of the Russian classification, these forests are considered as floodplain cycle of plant associations. They are divided into two, herbaceous—stone bramble and hair grass sedge series. Each of the two series is composed of two associations. Communities of the herbaceous—stone bramble series (
|Parameter||Plot 1||Plot 2||Plot 3|
|Plot location||ridge top||even part of floodplain||deep inter‐ridge depression|
|high level||mean level||low level|
|Tussocks up to
15–20 cm high
|Level of groundwater||Deeper than 2.5 m||About 1.5 m||0.75 m|
herb‐stone brumble forest
stone brumble forest
grass sedge forest
|Stand age||I layer—VI
II layer—IV age class
II layer—IV age class
II layer—IV age class
|I layer—20–22 m
II layer—14–18 m
|I layer—20–22 m
II layer—14–18 m
|I layer—20–22 m
II layer—14–18 m
|Moss cover (TPC, %)||It is not expressed||It is expressed weakly,
(TPC up to 3%)
|It is expressed weakly,
species are registered
(TPC up to 10%)
|Alluvial soddy layered
soil on sandy alluvium
|Alluvial meadow soil
on clay alluvium
soil on clay alluvium
|The structure of the soil
Soil mineral horizons at Plot 1 have favourable moisture conditions for soil organisms – 40–60% of total moisture capacity (TMC). But forest litter suffers from moisture deficiency for practically whole summer period. Forest litter moisture degree equals 20–40% of TMC in summer. Plot 2 has best moisture conditions (within 40–60% of TMC) in upper soil horizons only towards July. In the other summer months and in autumn, soil moisture degree is 60–80% of TMC. At Plot 3, soil is seriously overmoistured due to close ground water occurrence. Additionally, this plot is under water for a long period of time during spring snow‐melt period. But towards in July, it also decreases in soil moisture content of forest litter to 40–60% of TMC. It resists 60–65% of TMC to the end of vegetation period. Lower mineral horizons remain overmoistured towards late autumn (80–90% of TMC). The study soils are acid, base‐unsaturated with a strongly decreasing organic carbon profile distribution (Table 2). At 20–30‐cm depth, organic carbon content was 0.5–0.9% and from 4.5–4.8 (Plot 1, Plot 2) to 3.0% (Plot 3) in humus horizon. In direction from Plot 1 to Plot 3, soil acidity in forest litter increases. In autumn, forest litter acidity in all soil types decreases due to fresh plant residues. Thus, there is a clear trend of worsening in living conditions of tree waste decomposing soil biota going from Plot 1 to Plot 3. Ecological conditions of biotopes undergo serious changes in direction from ridge top to inter‐ridge depression. These changes respond for differences in qualitative and quantitative composition of plant waste, intensity of its mineralization and humification processes, structure of soil humus horizons. Upper organic soil horizons of alluvial forest soils are highly unstable by moisture content and heat provision. They are the principle habitat for invertebrates. Even overmoistured (meadow‐boggy) soils have favourable conditions for soil biota life activity some time during summer‐autumn.
|Horizon||Depth, cm||pHKCl||Ha**||Exchangeable bases||S****||C||N||C/N||Sum of particles|
|Ca2+||Mg2+||<0.01 mm||<0.001 mm|
|mmol/100 g soil||%|
|Plot 1. Ridge top, alluvial soddy soil|
|Plot 2. Even floodplain part, alluvial meadow soil|
|Plot 3. Deep inter‐ridge depression, alluvial meadow‐boggy soil|
4. Diversity and structure of microbe communities in the alluvial forest soils
Microbe communities of alluvial soils differ from those of watershed soils  because of inundation of floodplain terrace and covering the surface of floodplain terrace with alluvial deposits . Floodplain forest soils have a short bacterial profile. Bacteria are available within upper 60–70 cm at Plot 2 and Plot 3. Single colonies were identified at a depth of 100 cm at Plot 1. In direction from Plot 1 to Plot 3, oligonitrophilous and denitrifying microorganisms become abundant. Nitrifying, ammonificators and bacteria using mineral nitrogen compounds strongly decrease in number at Plot 3 (Figure 3). This normally proceeds in soils formed in conditions of excessive moisture content. Oligonitrophilous and oligotrophic microorganisms dominate at Plots 2 and 3 in the beginning of vegetation period. Ammonifiers, nitrifiers, denitrifiers, assimilators of mineral nitrogen compounds become abundant in July and towards the end of vegetation period. Their number and ratio significantly vary depending on soils type and weather conditions. When weather conditions are unfavourable (chilly weather with excessive rain precipitations), bacterial communities are presented by oligonitroflora. Unusually, warm weather with insufficient rain precipitations activates ammonifiers, assimilators of mineral nitrogen compounds and nitrifiers. Normally this increase in bacterial number also occurs in autumn when soil surface gets covered with plant waste products which decrease forest litter acidity. Microbial biomass carbon content in alluvial soils largely varies (Figure 4). Hydromorphic soils (Plot 2 and Plot 3) normally contain more microbe biomass carbon than automorphic soils (Plot 1). This situation is especially obvious in August and September when microbe communities in these soil types are highly active.
5. Micromycetes activity in the alluvial forest soils
Micromycetes are the first inhabitants of plant waste products in forests [34, 35]. Soils of coniferous forests are rich in both microscopic and basidial fungi; their mycelium largely permeates forest litter . Seventy‐three species of micromycetes (including sterile forms) of 18 genera inhabit soils under spruce forests on watersheds of the Sysola and the Vychegda Rivers . Leaf waste of birch‐aspen forests differs from that of spruce and pine forests by chemical composition and contains by 1.5–2 times more mineral elements as calcium, magnesium and nitrogen than pine or spruce needles . So, deciduous leaf waste presents a well decomposition object for bacteria. Our data showed that floodplain forest soils have 70 species of micromycetes. But taxonomic diversity of microscopic fungi is truly higher in floodplain soils (31 genera). Practically, one half of micromycetes in floodplain forest soils (25 species) are species which were previously identified neither in taiga forest soils , nor in floodplain meadow soils of the middle taiga zone . They are species of the Penicillum (
6. Communities structure and diversity of nematodes in the alluvial forest soils
Soil nematodes of the alluvial soils include 53 genera of 30 families. Number of nematodes varies between 635 inds./100 cm3 (Plot 3) and1105 inds./100 cm3 (Plot 1). Diversity of nematodes in the floodplain forests is higher than that in non‐flooded spruce (35) and pine forests (31 genera) in the study region (not‐published data). Number of nematodes in the alluvial soils is also a little bit higher than that under watershed forests. It is 55–239 inds./100 cm3 in pine forests  and 300 inds./100 cm3 in deciduous forests. The greatest number of genera is found for soil at Plot 2—42 genera and the lowest one is noted for soil at Plot 3—32 genera (Table 3). Plot 3 is characterized by lowest diversity of nematodes. It is probably explained by unfavourable life conditions (overmoisture) for them. Excessive moisture of soil results into its poor aeration and reduction of qualitative and quantitative vegetation composition. Consequently, the taxonomic similarity of nematodes at the study plots is not high and it is equal
|Genera, parameter||Studied soils|
|Plot 1||Plot 2||Plot 3|
|Number of genera||39||42||32|
|Density (inds./100 cm3)||1105 ± 185||745 ± 94||635 ± 90|
|Shannon diversity index (H')||2.30 ± 0.12||2.63 ± 0.13||1.93 ± 0.34|
|Simpson's index of dominance (DSM)||0.15 ± 0.03||0.12 ± 0.03||0.29 ± 0.12|
7. Diversity and structure of Collembola in the alluvial soils
Collembola of floodplain communities in the taiga zone of the European part of Russia are understudied in comparison with coniferous forests on watersheds. For example, 65 Collembola species are registered in the floodplain ecosystems of the Komi Republic  whereas 173 species are noted in the coniferous forests of the European part of Russia . The greatest number of species and stenobionts (species which exist only in particular habitats) are found at Plots 1 and 3 (Table 4). It is confirmed by the Shannon indices (2.70–2.94) that indicate their high taxonomic diversity. It is noticed that
|Species, parameter||Plot 1||Plot 2||Plot 3|
|Total number of individuals||21,574||13,642||8248|
|Total number of species (S)||45||35||39|
|Shannon diversity index (H')||2.94||2.14||2.70|
|Shannon evenness index (J')||0.48||0.41||0.51|
8. Diversity and structure of soil macrofauna in the alluvial soils
|Taxa||Plot 1||Plot 2||Plot 3|
|Total number of taxa||17||16||9|
|Total number of species||95||87||30|
Macrofauna of alluvial soils has a rich species composition (Table 5). Invertebrates in floodplain soils inhabit only the upper 30‐cm soil layer. About 80% of large invertebrates occupy the upper 0–10‐cm soil layer. Under the mark of 20 cm, there are only single individuals of earthworms . Six species of Lumbricidae are registered in alluvial soils.
|Species, parameter||Plot 1||Plot 2||Plot 3|
|Shannon diversity index (H')||3.50||4.38||3.84|
|Shannon evenness index (J')||0.69||0.86||0.74|
|Simpson's index of dominance (DSM)||0.16||0.07||0.12|
|Shannon diversity index (H')||4.55||4.72||2.58|
|Shannon evenness index (J')||0.87||0.94||0.86|
|Simpson's index of dominance (DSM)||0.06||0.03||0.18|
|Shannon diversity index (H')||2.95||2.64||-|
|Shannon evenness index (J')||0.82||0.88||-|
|Simpson's index of dominance (DSM)||0.16||0.20||-|
9. The impact of ecological conditions of the alluvial soils on soil biota
Deciduous birch and aspen forests in river floodplains cause development of specific biotopes. They, in turn, affect soil cover formation, species composition of plants, soil microorganisms, meso‐ and macro‐ fauna. But very similar floristic composition of plant communities within one series of floodplain forests evidences different tree species be not as pronounced edificators in specific floodplain ecotope conditions as those on watersheds. Ecological conditions considerably affect taxonomic diversity and number of biota in alluvial forest soils. The highest number of species and genera of micromycetes was found in alluvial meadow soil (Plot 2). Soil of this plot takes a transitional position between alluvial soddy soil (Plot 1) and alluvial meadow‐boggy soil (Plot 3). It is well moistured, not extremely dry or moist. The large number of micromycetes species is found in alluvial soddy soil with moisture deficiency. The lowest number of species is found in alluvial meadow‐boggy soil. Generally, complex of microscopic fungi is specific for each soil type. Taxonomic diversity of microscopic fungi in floodplain forest soils is higher than that in floodplain meadow and coniferous forest soils of the taiga zone. Nevertheless, species composition is practically the same. High taxonomic diversity of microscopic fungi in floodplain forests of the North could be conditioned by specific plant waste. Plant waste in spruce forests is homogenous and consists of moss and needle residues, so it has a specific biochemical composition and it is decomposed mainly by
For the first time, we conducted a comprehensive study on major components of floodplain forests for middle taiga territory of European north‐eastern Russia. Alluvial soils were shown to distinguish by ecological habitat conditions, particularly ridge top, levelled floodplain part and deep inter‐ridge depression. Soils formed on different parts of floodplain terrace were revealed for different taxonomic composition of microorganisms, meso‐ and macro‐fauna. Floodplain forest soils of the middle taiga subzone are inhabited 70 species from 31 genera of micromycetes, 53 genera from 30 families of nematodes, 60 species from 39 genera and 13 families of Collembola, 110 species from 17 taxa of large invertebrates. Alluvial meadow soils with stable moisture and temperature conditions are most diverse by species composition of micromycetes, nematodes, and large invertebrates. Springtails prefer alluvial soddy soils. Towards the end of vegetation period, every alluvial soil type increases in number of bacteria, microscopic fungi, and soil invertebrates in forest litter and becomes inhabited by new species. This is related with fresh plant waste and appropriate moisture conditions. So, fresh plant residues is actively transformed at this period not only in soils at elevated places (with favourable water‐air regime at whole vegetation period) but also in overmoistured soils at inter‐ridge depressions. Alluvial soils under floodplain forests in the European North‐East of Russia are habitats with a high life density. In contrast with soils under coniferous forests on watersheds, alluvial soils in river valleys have a high taxonomic diversity of soil biota. Floodplain forest soils are habitats for rare species which cannot be met in soils under coniferous forests. As result of our research, 29 species of Collembola were noted at first for the Komi Republic, 2 species of Collembola were identified as new for science , 2 rare species of large invertebrates were included in the Red List of Komi Republic. The obtained data on quantitative and qualitative composition of micromycetes, nematodes, springtails and large invertebrates can be used for assessment of spatial‐temporal changes of floodplain soils under anthropogenic impacts.
The research was financially supported by the UrD RAS Presidium Program ‘Scientific Biodiversity Conservation Bases in Russia’ (projects: ‘Environmental Functions of Alluvial Soils and Biodiversity Formation of Floodplain Landscapes in the European North‐East of Russia', ‘Identification of Biodiversity Formation Mechanisms, Interdependences of Macro‐ and Microorganisms and their Role in Organic Matter Transformation in Soils of Floodplain Forests in the European North‐East of Russia', 'The relationship of biodiversity and biological production potential of the terrestrial ecosystems of the European Arctic with the peculiarities of formation of permafrost soils and dynamic aspects of their transformation in the modern climate conditions', 'The diversity of plant and soil cover on the UNESCO World Heritage area “Virgin forests of Komi republic'), the Grants of the Government of the Komi Republic and the RFBR (09‐04‐98808 r_sever_a) ‘Animal Population in Soils of Floodplain Ecosystems of the European North’, (16-44-110989 r_a) The creation of an information system “Soil fauna of Komi Republic”.
Batzer, D., Boix, D. (Eds.). Invertebrates in freshwater wetlands. An International Perspective on Their Ecology. New York, Dordrecht, London: Springer International Publishing Switzerland, 2016. 645 pp.
Dobrovoljskij, G.V. Soils of river floodplains in the central part of Russian Plane. Moskow, 1968. 298 p (In Russia).
Kryshtal, A.F. To study of soil and litter entomofauna dynamics in connection with the flood in the Dnepr river valley. Entomological Review, 1955. Vol. 34. No. 1. P. 120–139 (in Russia).
Geltser, Yu. A. About soil fauna in the Kljasjma River floodplain. Floodplain Soils of Russian Plane, M., 1963. Vol. 2. P. 141–145 (In Russia).
Striganova, B.R. Soil invertebrate complexes in the Dnestr River floodplain. Entomological Review, 1968. Vol. 47. No. 3. P. 360–368 (in Russia).
Perel, T.S. Differences of Lumbricidae organisation with their ecological features. Adaptation of Soil Invertebrates to Environment Conditions. M., 1977. P. 129–145 (In Russia).
Lessel, T., Marx, M.T., Eisenbeis, G. Effects of ecological flooding on the temporal and spatial dynamics of carabid beetles (Coleoptera: Carabidae) and springtails (Collembola) in a polder habitat. ZooKeys, 2011. Vol. 100. P. 421–446.
Busmachiu, G. Collembola (Hexapoda) from the riparian habitats of the Dniester River. Muzeul Olteniei, Craiova. Studii si comunicări. Stiintele Naturii., 2011. Vol. 27. No. 1. P. 63–70.
Taskaeva, A.A., Lapteva, E.M. The dynamics of Collembola communities in middle‐taiga flood plain forests. Povolzhsky ecologichesky zhurnal, 2012. No. 4. P. 426–436 (In Russia).
Kolesnikova, A.A, Taskaeva, A.A., Lapteva, E. M., Degteva, S.V. Vertical distribution of Collembola, Lumbricidae and Elateridae in alluvial soils of floodplain forests. Contemporary Problems of Ecology, 2013. Vol. 6. No. 1. P. 34–42.
Taskaeva, A.A. Springtails (Collembola) assemblages in floodlands of the taiga zone of the Republic of Komi. Entomological Review, 2009. Vol. 89. No. 8. P. 965–974.
Golovchenko, A., Dobrovo'skaya, N. Population density and the reserves of microorganisms in floodplain soils of the protva river. Eurasian Soil Science. 2001. Vol. 34, No. 12. P. 1300–1304.
Russell, D., Hauth, A., Fox, O. Community dynamics of soil Collembola in ains of the Upper Rhine Valley. Pedobiologia. 2004. Vol. 48. No. 5–6. P. 527–536.
Sterzyńska, M. Assemblages of soil Collembola in wetlands in the floodplains of some Polish rivers. Museum and Institute of Zoology PAS, Warszawa. 2009. 96 pp.
Leontieva, M.V., Dobrovol'skaya, T.G., Pochatkova, T.N. Influence of flooding regime on the taxonomic structure of soil bacterial communities. Bulletin of Moscow University. 17: Soil. 2005. No. 1. P. 36–40 (in Russia).
Sterzyńska, M., Pižl, V., Tajovský, K., Stelmaszczyk, M., Okruszko, T. Soil fauna of peat‐forming wetlands in a natural river floodplain. Wetlands. 2015. doi:10.1007/s13157‐015‐0672‐0.
Russell, D., Schick, H., Nahrig, D. Reactions of soil Collembola communities to inundation in floodplain ecosystems of the Upper Rhine Valley. Broll, G., Merbach, W., Pfeiffer, E.‐M. (Eds.). Wetlands in Central Europe: Soil Organisms, Soil Ecological Processes and Trace Gas Emissions. Görlitz: Springer, 2002. P. 35–70.
Tuf, I.H. Four‐year development of a centipede (Chilopoda) community after a summer flood. African Invertebrates. 2003. Vol. 44. P. 265–276.
Adis, J., Junk, W. Terrestrial invertebrates inhabiting lowerland river floodplains of Central Amazonia and Central Europe: a review. Freshwater Biology. 2002. Vol. 47. P. 711–731.
Kuhle, J.C. Spatial patterns of distribution of earthworms in a hardwood floodplain forest. Pizl V., Tajovsky K. (Eds.). Soil Zoological Problems in Central Europe. Ceske Budejovice, 1998. P. 125–134.
Degteva, S.V. Parameters of ecological space and floristic diversity of forest formations in the northeast of European Russia. Russian Journal of Ecology, 2005. I. 36. No. 3. P. 158–163.
Khabibullina, F.M., Kuznetsova, E.G., Vaseneva, I.Z. Micromycetes in podzolic and bog‐podzolic soils in the middle taiga subzone of northeastern European Russia. Eurasian Soil Science. 2014. Vol. 47. No. 10. P. 1027–1032.
Kudrin, A.A., Dolgin, M.M., Kolesnikova, A.A., Konakova, T.N., Taskaeva, A.A. Spatial distribution features of soils fauna in fine forests of the north taiga subzone (Komi Republic). Bulletin of the North (Arctic) Federal University. Series: Natural Sciences. 2014. No. 1. P. 72–83 (in Russia).
Kuznetsova, N.A. Organization of springtails communities. M., 2005. 244 p (in Russia).
Vorobyov, L.A. (Eds.). M. Theory and practice of chemical analysis of soils. GEOS Publicity 2006. 400 p (in Russia).
Vadyunina, A.F., Korchagina, Z.A. The study methods of soil and ground physical properties. M.: Vyssh.shkola, 1986. 345 p (in Russia).
Classification and diagnostics of the USSR soils. M.: Kolos, 1977. 224 p (in Russia).
Zvyagintsev, D.G. Methods of Soil Microbiology and Biochemistry. M.: MGU Publicity 1991. 304 p (in Russia).
Blagodatskiy, S.A., Blagodatskaya, E.V., Gorbenko, A.Y., Panikov, N.S. A re‐hydration method of determining the biomass of microorganisms in soil. Pochvovedenie(Russian J. Soil Science). 1987. No. 19. P. 119–126 (in Russia).
Jairajpuri, M.S., Ahmad, W., Dorylaimida. Free‐living, Predaceous and Plant Parasitic Nematodes. E.J. Brill, Leiden, 1992. 458 p.
Bongers, T., De nematoden van Nederland. Nederlandse Natuurhistorische Vereniging. Utrect, The Netherlands, 194. 408 p.
Brzeski, M.W. Nematodes of Tylenchina in Poland and Temperate Europe. Muz. Inst. Zool. PAN. Warszawa, 1998. 397 p.
Yeates, G.W., Bongers, T., de Goede, R.G., Freckman, D.W., Georgieva, S.S. Feeding habits in soil nematode families and genera e an outline for soil ecologists. Journal of Nematology. 1993. Vol. 25. P. 315–331.
Dobrovol'skaya, T., Zvyagintsev, D., Chernov, I., et al. The role of microorganisms in the ecological functions of soils. Eurasian Soil Science. 2015. Vol. 48, No. 9. P. 959–967.
Berg, M.P., Kniese, J.P., Verhoef, H.A., Dynamics and stratification of bacteria and fungi in the organic layers of a scots pine forest soil. Biol. Fert. Soils. 1998. Vol. 26. P. 313–322.
Lindahl, B.D., Ihrmark, K., Boberg, J., Trumbore, S.E., Högberg, P., Stenlid, Ja., Finlay R.D. Spatial separation of litter decomposition and mycorrhizal nitrogen uptake in a boreal forest. New Phytologist, 2007. 173 pp.
Berger, T.W. Auswirkungen der Baumartenzusammensetzung auf den Waldbodenzustand von sekundären Fichtenwäldern und gemischten Fichten‐Buchenbeständen. Centralbl. Gesamte Forstw. 2001. 118, No. 4. P. 193–215.
Lapteva, E.M., Khabibullina, F.M., Vinogradova, Yu. A. Diversity of micromycetes in flood plain meadow soils. Mycology and Phytopathology J., 2009. Vol. 43. Issue 3. P. 200–206 (in Russia).
Sohlenius, B. Influence of clear‐cutting and forest age on the nematode fauna in a Swedish pine forest soil. Applied Soil Ecology. 2002. Vol. 19. P. 261–277.
Zhao, J., Neher, D.A. Soil energy pathways of different ecosystems using nematode trophic group analysis: a meta analysis. Nematology. 2014. Vol. 16. No. 4. P. 379–385.
Raschmanová, N., Kováč, L., Miklisová, D. The effect of mesoclimate on Collembola diversity in the Zádiel Valley, Slovak Karst (Slovakia). European Journal of Soil Biology. 2008. Vol. 44. P. 463–472.
Kuznetsova, N.A. Humidity and distribution of springtails. Entomological Review, 2003. Vol. 83. No. 2. P. 230–238 (in Russia).
Potapov, M.B., Taskaeva, A.A. Analysis of vicarious species Folsomia kuznetsovaesp.n. and F. bisetosaGisin (Collembola: Isotomidae). Russian Entomological Journal. 2009. Vol. 18. No. 1. P. 1–6 (in Russia).
Maculec, G. The effect of long term drainage of peat soil on earthworm communities (Oligochaeta: Lumbricidae). Polish Ecological Studies. 1991. Vol. 17. P. 203–219.
Pizl, V. Earthworm communities in Palava Biosphere Reserv (Southern Moravia) with special reference to the impact of floods. Pizl, V., Tajovsky, K. (Eds.). Soil Zoological Problems in Central Europe. Ceske Budejovice, 1998. P. 157–166.
Tajovsky, K. Impact of inundations on terrestrial arthropod assemblages in southern Moravia floodplain forests, the Czech Republic. Ekologia. 1999. Suppl. 18. No. 1. P. 77–184.
Tufova, J., Tuf, I.H. Survival under water – comparative study of millipedes (Diplopoda), centipedes (Chilopoda) and terrestrial isopods (Oniscidea). Tajovsky, K., Schlaghamersky, J., Pizl, V. (Eds.). Contributions to Soil Zoology in Central Europe I. Ceske Budejovice, 2005. P. 195–198.
Bayley, P. B. Understanding large river: floodplain ecosystems. BioScience. 1995. P. 153–158.
Tockner, K., Lorang, M. S., Stanford, J.A. River floodplains are model ecosystems to test general hydrogeomorphic and ecological concepts. River Research and Applications. 2010. Vol. 26. No. 1. P. 76–86.