ISSN 2617-6106 (print) ISSN 2617-6114 (online) Agrology, 2(4), 205‒208 AGROLOGY doi: 10.32819/019029

Оriginal researches Spatial Organization of the Vallonia Pulchella (Muller 1774) Ecological Niche in Sod-lithogenic on Loesses-Like Clays in the Nikopol Manganese Ore Basin

Received: 04 September 2019 A. K. Umerova Revised: 09 September 2019 Bohdan Khmelnytskyi Melitopol State Pedagogical University, Melitopol, Ukraine Accepted: 10 September 2019

Bohdan Khmelnytskyi Melitopol State Abstract. The influence of edaphic and phytoindication parameters on the spatial organiza- Pedagogical University, Hetmanska Str., 20, tion of the micromollusc Vallonia pulchella (Muller 1774) ecological niche was experimentally Melitopol, 72312, Ukraine investigated. The field experiment was conducted in June 2015 at the research polygon within the Nikopol Manganese ore basin (sod-lithogenic soils on loam loesses-like clays). A promi- Tel.: +38-096-057-17-84 sing area of research is the issue: what exactly edaphic factor and phytoindication parameters E-mail: [email protected] is the most important determinants of micromolluscs distribution. The experimental polygon was consisted of 105 samples located within 7 transect (15 samples each). The Vallonia pul- Cite this article: Umerova, A. K. (2019). chella average density was 2.54 ind./sample. The average penetration resistance of the was Spatial organization of the Vallonia pulchella found as a result of the experiment studies to increase with depth down the profile. The analy- (Muller 1774) ecological niche sis of aggregate fractions showed that the number of molluscs is unstable and varies in the in sod-lithogenic soils on loesses-like clays in the Nikopol Manganese Ore Basin. Agrology, range of 5.17% to 25.45%. The snails were found to prefer the aggregate fractions 1‒2 mm or 2(4), 205‒208. doi: 10.32819/019029 in less extent – the 0.5–1.0 mm. The elevated level of carboxylic salts was detected by means of phytoindication methods as well as the insignificant content of free nitrogen in neutral soil. In the paper the soil and vegetation properties were considered as a indicators of the living or- ganism habitats. It allowed to assess the Vallonia pulchella ecological niche marginality and specialisation. The results obtained may be usful in the evaluation of artificial soil structures.

Keywords: vegetation; soil penetration resistance; aggregate fractions; terrestrial molluscs; population dynamics; population.

Introduction origin, after the disappearance of the geographical isolation, due to resettlement from a refugium after climatic changes, during The biosphere transformation is due to anthropogenic influ- the temporary colonization of micro stations as a result of the ence. This process has reached a global scale and impacts on all displacement by invasive species. Thus, ecological niche optima ecosystems, that has led to the structural and functional changes may be presented by the integral variables such as marginality in communities. The study of the ecological niche theory is one and specialization axes and maybe plotted in geographic space of the urgent problems of the modern ecology, indicating different by means of habitat suitability index reproduction (Yorkina, levels of the organization of living (Pianka, 1981). The theory of Maslikova, Kunah, & Zhukov, 2018). ecological niche describes the value of species in the community. The spatial distribution of land-snail species and individuals This theory distinguishes the concept of “population”, “communi- has been extensively investigated (Myšák, Horsák, Svobodová, ty” and “ecological system” (Smyntyna, 2013). Hutchinson (1957) & Cernohorsky, 2013; Faly, Kolombar, Prokopenko, Pakhomov, proposed a more formal, quantitative approach based on this theory. & Brygadyrenko, 2017). Mollusc populations may be relatively He conceived of the niche as a hyper-volume in multi-dimensio- evenly distributed or highly aggregated (Kralka, 1986; Locas- nal environmental space delimiting where stable populations can be ciulli & Boag, 1987). Studies at a large scale level have made it maintained (Kearney, Simpson, Raubenheimer, & Helmuth, 2010). possible to determine the role of edaphic factors in the spatial The Ecological Niche Factor Analysis (ENFA) is based on the as- distribution, abundance, and diversity of molluscs communities sumption that a species are not randomly distributed with respect to (Nekola & Smith, 1999; Juřičková et al., 2008; Szybiak, Błoszyk, ecogeographic variables (Hirzel & Guisan, 2002). Koralewska-Batura, & Gołdyn, 2009). The researchers found ef- The concept of the ecological niche is based on a study of the fects on the distribution of species that are commonly spatially relationship of organisms in natural communities. The analysis of structured as community also have spatial structure (Dray, Leg- its structure makes it possible to identify the specific behaviou- endre, & Peres-Neto, 2006). These patterns may be associated ral, ecological and morphological traits of organisms. With this with the distribution of microhabitats (Kralka, 1986; Hylander, in mind, is maybe better understood as a historical process of the Nilsson, Jonsson, & Gothner, 2005). The main edaphic factors formation of species. The research of the ecological niche theory that affect the lives of molluscs, mainly include the soil chemical is popular and important. Now the object of study of ecology is composition and ions and moisture (Čejka& Hamerlík, 2009) and worth the following question: how many ecological niches can calcium gradient (Hubricht, 1985). The spatial scale and the hi- accommodate biogeocoenosis and how many of these species can erarchy of factors acting on molluscs are of special interest (Bo- exist together. Also necessary to find out the value of a competi- han, Glen, Wiltshire, & Hughes, 2000; Myšák et al., 2013; Nekola tion in the divergence of organisms for ecological niches and the & Smith, 1999). The habitat is characterized by the presence of formation of communities. resources and conditions for given species in some territory, as The pedobionts ecological niche can occupy for various rea- a result of which the colonization of this territory becomes pos- sons: as a result of the overlapping of the faunas from different sible, including its survival and reproduction (Hall, Krausman,

AGROLOGY | Volume 2 | Issue 4 205 A. K. Umerova Spatial organization of the Vallonia pulchella (Muller 1774) ecological niche in sod-lithogenic soils on loesses-like clays in the Nikopol Manganese Ore Basin

& Morrison, 1997). The identify the characteristics of the envi- ‒ Table. On the test site is a characteristic monotonic increase in soil ronment that make the place suitable for the species existence penetration resistance with increasing soil depth. Its average value the choice of habitats (Calenge, 2006). This is the called “envi- is increased from 3.66 MPa to 9.28 MPa to the surface at a depth of ronmental standard” species (Gilyarov, 1965) the need for each 50 cm down the profile. The highest coefficient of variation is ob- species in a certain complex environmental conditions. The aim served at a depth of 0‒5 cm (37.44%), and at 15‒20 cm, it is reduced of our work is to study the spatial organization micromolluscs V. 4,3 times (8.7%). The acquires stable value at the level of 30‒50 cm, pulchella (Muller 1774) population in terms of the edaphic prop- ranging (18.85‒17.39%) ‒ Table. erties and properties of the vegetation cover in the sod-lithogenic In the test site the average values of soil penetra- soils on loam loesses-like clays in Nikopol manganese ore basin. tion resistance was greater than the critical value for the growth of plant roots (3.0‒3.5 MPa), starting with 0‒5 cm Material and methods of soil layers. In the according to penetrometer indicators: 3 MP indicates soil compaction, which counteracts root The studies were conducted at the research station within- growth (Faechner, Pyrcz, & Deutsch, 2000). Also, may be Nikopol Manganese ore basin in June 2015. Tests were done in that this soil layer formed by the most appropriate condi- sod-lithogenic soils on loesses-like clays. The polygon consisted tions according to the criterion of penetration resistance for of 7 transects. Each transect was made up of 15 sampling points. the growth of herbaceous plants. Exceeding the limit to the The distance between rows within the polygon was 3 m. The level of soil penetration resistance of plant roots (3 MPa) calculation soil invertebrates were conducted by soil samples di- (Zhukov, Andrusevich, Lapko, & Sirotina, 2015) to was 7.84. sassembly. Samples consisted of single blocks of soil, 0.25×0.25 m The coefficient of variation of this value is significant and deep, dug out quickly. Measurement of soil mechanical imped- amounts to was 18.48%. ance was carried out in the field using a hand penetrometer Ei- The analysis aggregate structure indicates that the predomi- jkelkamp, to a depth of 50 cm at intervals of 5 cm. The ave- nant fraction is 1‒2 mm (25.45%), they determine the resis- rage error of the measurement results of the device is ±8%. tance of soils to erosion under the influence of water. Some- The measurements were made by a cone with a cross-sectional what lower, given the relative proportion, fractions 2‒3 mm and dimension of 1 cm2. Within each measurement point, the me- 3‒5 mm, the other fractions (<0.25 mm; 0.25‒0.50 mm and chanical impedance of the soil was made in a single replication. 0.5‒1.0 mm. 5‒7 mm. 7‒10 mm and > 10 mm) are at the level Measurement of the electrical conductivity of the soil in situ, 5.17‒7.83%. It was found, that molluscs V. pulchella are sen- the sensor HI 76305 was used (Hanna Instruments, Woodsocket, sitive to the soil aggregate structure: the highest number – ac- RI). This sensor works in conjunction with the portable device counted for aggregate fractions of 1‒2 mm, in less extend – HI 993310. Apparatus estimates the total electrical conductivity 0.5‒1.0 mm (due to exposure to sunlight and drying mollusc). of soils, i.e. soil total conductivity of air, water and particles. De- Observed the stable numbers of V. pulchella with growth ag- vice measurement results are shown in the soil saturation units gregate fractions (5‒7; 7‒10; >10 mm). On the zonal micro- salts solution (Pakhomov, Konovalova, & Zhukov, 2010). The ag- mollusk distribution, may be influenced uneven distribution of gregate structure was evaluated by Savinov’s dry sieving method minerals, root exudates and dendrite. (Vadunina & Korchagina, 1986). The percentage content of such The use of vegetation as an indicator makes it easier to obtain fractions was established: <0.25; 0.25–0.5; 0.5–1; 1–2; 2–3; 3–5; results on the status of various components of the ecosystem. 5–7; 7–10; >10 mm, and plant roots. The phytoindication to seem most convenient for determining Based on geobotanical descriptions, phytoindicative assess- of environmental factors (Didukh, 1994; Didukh, 2000). Let us ment of environmental factors according to Didukh (2011) was analyze the relation V. pulchella for each of the considered envi- made. Didukh phytoindication scales (Didukh, 2011) include ronmental factors. Water regime (Hd) area favoura- edaphic and climatic scales. The edaphic phytoindication scales ble for sub-mesophytes (Didukh, 2011). The average variability include the regime (Hd), the variability of humidity of humidity (fH) of 6.05 and corresponds hydrocontrastophiles (fH), the soil aeration (Ae), the soil acidity (Rc), the total salt type, adapted to dry steppe or damp habitats which are formed regime (Sl), the carbonate content in the soil (Ca) and nitrogen in conditions of extremely irregular wetting of soil layer small content in the soil (Nt). The climatic scales include the parame- drenched by precipitations and melted waters (Didukh, 2011). ters of the thermal climate (thermal regime, Tm), humidity (Om), By soil acidity factor (Rc), it is established that the species under cryo-climate (Cr) and the continentality of climate (Kn). In addi- study is in comfortable conditions (neutral soils). In the study tion to these, the lighting scale (Lc) is indicated, which is charac- area, the salt regime (Sl) is a sub-glycotrophes. This causes car- terized as a microclimate scale. bonate salinity type, the cause of which is the lack of of Statistical calculations were performed using Microsoft Excel water due to the small amount of rainfall and high capacity for 2010 program. Estimation of the mean from assumptions about evaporation (Didukh, 2011). The carbonate content in the soil of the distribution of the random variable according to Poisson was the test portion (11.29) creates the most favourable conditions obtained by the formula (Shebanin, Melnik, Kramarenko, & Gan- for carbonatophiles. The content of nitrogen (Nt) in the soil in- ganov, 2008): dicates that the investigated areas dominated hemi-nitrophiles – plants which grow on soils moderately provided with mineral n D = −ln 0 , nitrogen (Didukh, 2011). Аeration mode creates favourable con- n ditions for hemi-aerophobes. Thus, soil aeration is sufficient to where D is the mean estimation; n0 is the number of samples without implement V. pulchella needs. Phytoindication parameter indi- the presence of the species; n is the total number of samples. cates that thermal climate (Tm) was 8.94, thus corresponding to the optimum for sub-mesotherms. Results The indicator humidity is associated with a flow rate of precipi- tation, evaporation and transpiration, , groundwater In total air-dry soil weighing 10,5 kg were examined, in which level (Konstantinov, 1968). Average microclimate value typically 266 specimens of Vallonia pulchella (Muller 1774) were found. for sub-aridophytes. The continentality of climate (Kn) determined Thus, the average density of this species in the sod-lithogenic soils by the effect on it of large areas of land and sea, climate-process on loam loesses-like clays was 2.54 ind./m2. With this in mind, the (atmospheric pressure, the nature of the circulation of the atmos- edaphic and vegetation properties can be considered as determi- phere, the concentration and distribution of moisture, the quantity nants of ecological space micromollusc V. pulchella (Muller 1774) of precipitation, temperature amplitude) (Didukh, 2011). Average

206 AGROLOGY | Volume 2 | Issue 4 A. K. Umerova Spatial organization of the Vallonia pulchella (Muller 1774) ecological niche in sod-lithogenic soils on loesses-like clays in the Nikopol Manganese Ore Basin

Table. Determinants of ecological space micromollusc Vallonia pulchella (Muller 1774), n = 105

Confidence interval Characteristics Mean CV, % 95% ‒95% Soil penetration resistance in MPa at depth, cm 0–5 3.66 3.92 3.40 37.44 5–10 6.10 6.50 5.69 34.67 10–15 7.53 7.73 7.34 13.30 15–20 8.00 8.13 7.87 8.70 20–25 8.48 8.63 8.33 9.10 25–30 8.71 8.91 8.51 11.96 30–35 8.66 8.97 8.35 18.85 35–40 8.85 9.11 8.60 17.31 40–45 9.18 9.49 8.86 18.13 45–50 9.28 9.59 8.97 17.39 Aggregate structure, size of fractions, mm >10 7.49 8.08 6.89 41.51 7–10 6.95 6.28 5.62 28.97 5–7 7.83 8.18 7.49 23.14 3–5 18.93 19.85 18.01 25.40 2–3 16.97 17.34 16.59 11.66 1–2 25.45 26.02 24.88 11.66 0.5–1.0 5.17 5.61 4.73 44.73 0.25–0.50 6.6 7.19 6.01 46.80 <0.25 5.62 6.03 5.20 38.50 Didukh phytoindicator values Hd 10.20 10.46 9.94 13.32 fH 6.05 6.29 5.81 20.43 Rc 7.70 8.98 8.82 4.60 Sl 8.22 8.34 8.11 7.25 Ca 11.29 11.38 11.19 4.30 Nt 5.03 5.28 4.77 26.5 Ae 6.38 6.50 6.25 10.53 Tm 8.94 9.05 8.83 6.50 Om 11.81 11.94 11.69 5.60 Kn 8.91 9.18 8.65 15.34 Cr 7.45 7.71 7.19 18.21 Lc 8.80 8.83 8.77 1.90 value continental mode (Kn) (Didukh, 2011) was 8.91, which cor- live organisms with completely opposite requirements to the con- responds for hemi-continental type. The cryo-climate (Cr) inherent ditions of existence (Dmitriev, 1997). V. pulchella distribution in was hemi-cryophytes. The light (Lc) mode varies from 8.83 to 8.77. the soil profile is uneven. This is due to the variability of the en- The average score was 8.8, which is favourable for heliophytes vironment, causing structural and functional diversity (Demidov, (Didukh, 2011). Kobets, Gritsan, & Zhukov, 2013). Aggregate particles of different sizes have affected the number of V. pulchella: macroaggregates, Discussion which have high porousness, important for the penetration of water and air, for the development of microorganisms, small animals and Soil penetration resistance is an important mechanical property plant roots. This is the prerequisite for life support mollusc, their that can be used as an indicator of soil. It is found, that with increa- breathing, moving and obtaining nutrients. Small aggregate parti- sing penetration resistance of the soil occurs reduction in the amount cles form a system of small pore size, which adversely affects the of available oxygen in the soil air to the root system. This leads to a lives of micromollusc. Thus, soil aggregates of various sizes cause gradual decrease in the physiological activity of the plants, which in the stochastic dynamics of the spatial distribution of V. pulchella, turn affects the zonal mollusc distribution. which can regain an important role in maintaining According to scientific views, the process of formation of ag- and its functioning. gregate particles in the soil is regulated not only by the level of Phytoindication parameters allow distinguishing the environ- organic matter, but also its qualitative composition. In the soil can mental situation within the studies biocenosis. Place, where lived

AGROLOGY | Volume 2 | Issue 4 207 A. K. Umerova Spatial organization of the Vallonia pulchella (Muller 1774) ecological niche in sod-lithogenic soils on loesses-like clays in the Nikopol Manganese Ore Basin analyzed species, enriched carbonate salts, but poor in nitrogen in Hubricht, L. (1985). The distributions of the native land mollusks neutral soils. That is what determines the nature of the placement of of the Eastern United States. Fieldiana: Zoology New Ser: 24. variability of the test object since under these conditions decreases Field Museum of Natural History, Chicago. doi: 10.5962/bhl. soil nutrition. title.3329 Hutchinson, G. E. (1957). Concluding remarks. Cold Spring Har- Conclusions bour Symposium on Quantitative Biology, 22, 415–427. Hylander, K., Nilsson, C., Jonsson, B. G., & Gothner, T. (2005). 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