The article presents the study results made on the Adonis wolgensis coenopopulation in Northern Kazakhstan (2018–2019). The materials have been gathered in the course of field research taking into account the literary data. The age structure of the coenopopulations is provided on the basis of detailed route studies. There have been studied eight coenopopulations located in Akmola, Pavlodar, and Kostanay regions. A. wolgensis is a short-stem herbaceous polycarpic of Northern Kazakhstan. A. wolgensis coenopopulations are found in three main habitats: steppe, meadow-steppe, and forest communities. The study of A. wolgensis coenopopulations in the latitudinal direction from Ekibastuz in the east to the Kostanay region in the west has revealed a high recovery index in steppe ecotopes. In other coenopopulations it is less than one. It is especially low in forest habitats. Most coenopopulations have a normal character according to the spectrum of age conditions, and they are maturing. This fact proves no negative impact of environmental and anthropogenic factors on the structure of coenopopulations and ensures their recovery and preservation.
Introduction
Adonis wolgensis Stev. is in the Red Book of Kazakhstan (category III). Dying species) [1]. The species is protected in Ukraine, the Republic of Moldova, Bulgaria, Romania [2–5], as well as in 11 regions of the Russian Federation.
Material and research methods
Age structure is one of the essential signs of cenopopulation, as it provides the ability of the population system to maintain itself and to determine its stability [7]. The study of Adonis wolgensis Stev.'s age structure was held in Northern Kazakhstan in 2018–2019. There have been studied 8 coenopopulations of A. wolgensis located in Akmola, Pavlodar, and Kostanay regions. The area of maximal distribution of A. wolgensis was determined for each CP using GPS; 25 reference sites (1 m2) were made inside the CPs on which the number of all A. wolgensis specimens was counted and their age stages were determined.
The age stages were determined according to the guidelines of T.A. Rabotnov [8], A.A. Uranov [9–10], L.B. Zaugolnova and O.V. Smirnova [7], Plant coenopopulations... [11].
The type of coenopopulation was determined according to T.A. Rabotnov [8] and to the «delta-omega» classification of L.A. Zhivotovsky [12].
The age index of a coenopopulation (Δ) is calculated according to the formula
Δ = Σ Kimi ⁄ Σ Ki,
i.e. ΣKi is a sum of the plants of all age stages; mi is the age of individual plants [13].
The efficiency index was defined by the formula ω = Σ piei,
i.e. pi = ni/n is the share of plants of an i-state in this population, ni is the absolute number of i-state plants, n = Σ ni is the total number of plants, ei is energy efficiency [13].
The recovery index (I) was calculated by the formula
I = Σj→v ⁄ Σ g1→g3,
i.e. Σj→v is the sum of plants of all age stages of the pre-generative period; Σg1→g3 is the sum of plants of all age stages of the generative period [13].
Results and discussion
A. wolgensis is a short-stem herbaceous polycarpic plant of Northern Kazakhstan represented by three morphotypes: steppe, meadow-steppe, and forest ones. They all have significant morphological differences in the height of shoots and the form of leaf blades.
CP-1. Pavlodar region, near Ekibastus, 51.66494°N, 75.28173°E, h = 412 m.a.s.l. Interslope lowland, northern slope, adonis-fescue community. The population area is 200 m2, total projective cover (TPC) is 60 %, the TPC of A. wolgensis is 40 %, the community has 24 species. The vegetation is formed by Adonis wolgensis, Alyssum desertorum, Androsace maxima, Artemisia austriaca, Festuca valesiaca, Potentilla humifusa, Ranunculus polyrhizos, Seseli ledebourii, Valeriana tuberosa.
CP-2. Pavlodar region, near Ekibastus, «Three birches» tract, 51.57517°N, 75.13187°E, h = 403 m.a.s.l. Eastern slope of the hill, feather-grass and fescue steppes on alkaline soils. The population area is 300 m2, TPC is 60 %, the TPC of A. wolgensis is 0.5 %, the community has 29 species. The vegetation is formed by Artemisia nitrosa, Astragalus testiculatus, Draba nemorosa, Festuca valesiaca, Limonium gmelinii, Ranunculus polyrhizos, Spiraea hypericifolia, Tulipa patens, Valeriana tuberosa.
CP-3. Pavlodar region, near Bayanaul, 50.2209° N, 75.8009° E, 406 m.a.s.l. The foot of the slope, the shrubs of a fescue-feather grass steppe. The population area is 500 m2, TPC is 90 %, the TPC of A. wolgensis is 0,5 %, the community has 29 species. The vegetation is formed by Allium nutans, Caragana frutex, Carex pediformis, Gagea granulosa, Poa angustifolia, Rosa spinosissima, Seseli libanotis, Stipa capillata, S. pennata, Thymus marschallianus.
CP-4. Akmola region, near Ereimentau, the Ereimentau mountains, 51.65717° N, 73.19056° E, h = 313 m.a.s.l. The foot of the slope is the alkaline meadow. The population area is 400 m2, TPC is 100 %, the TPC of A. wolgensis is 3.0 %, the community has 25 species. The vegetation is formed by Agropyron cristatum, Artemisia dracunculus, Carduus nutans, Centaurea scabiosa, Glycyrrhiza uralensis, Potentilla bifurca, Ranunculus polyanthemos, Rumex acetosa, Tamarix ramosissima.
CP-5. Akmola region, the Burabay region, near Kartakol, 52.93287° N, 70.49210° E, h = 431 m.a.s.l. Pine and birch forest. The population area is 300 m2, TPC is 60 %, the TPC of A. wolgensis is 0.5 %, the community has 20 species. The vegetation is formed by Achillea asiatica, Betula pendula, Calamagrostis epigeios, Equisetum hyemale, Filipendula ulmaria, Jacobaea erucifolia, Pinus sylvestris, Populus tremula, Rosa majalis, Solidago virgaurea,
CP-6. Kostanay region, near Scherbakovo, 53.20453° N, 64.21550° E. The sparse pine forest is growing on sands. Population area is 1000 m2, OPP is 30 %, the OPP of A. wolgensis is 0.5 %. The community has 22 species. The vegetation is formed by Agropyron pectinatum, Carex supinа, Genista tinctoria, Phragmites australis, Potentilla arenaria, Scorzonera ensifolia, Stipa capillata, Thymus serpyllum, Tulipa biebersteiniana, Veronica spicata.
CP-7. Kostanay region, near Ozerny, the Novonezhensky forestry, sq. 21, 52.43971° N, 64.09279° E, h = 213 m.a.s.l. The secondary psammophyte steppe is at the site of an old fire. Population area is 300 m2, OPP is 100 %, the OPP of A. wolgensis is 5.0 %. The community has 38 species: Artemisia campestris, Echinops sphaerocephalus, Gypsophila paniculata, Medicago falcata, Oenothera biennis, Otites wolgensis, Potentilla arenaria, Pulsatila uralensis, Sedum telephium, Xanthoselinum alsaticum,
CP-8. Kostanay region, near Zarechny, «Stone Lake», 52.28013° N, 63.76616° E, h = 134 m.a.s.l. The northern slope towards the lake, shrub thickets. Population area is 400 m2, OPP is 100 %, the OPP of A. wolgensis is 1.0 %. The community has 23 species: Amygdalus nana, Calamagrostis epigeios, Filipendula vulgaris, Potentilla bifurca, Rosa acicularis, R. spinosissima, Spiraea hypericifolia, Stipa capillata, S. pennata.
A. wolgensis coenopopulations are confined to three main habitats: steppe communities (CP-1, CP-2, CP-7), meadow-steppe communities (CP-3, CP-4, CP-8), forest communities (CP-5, CP-6). Many authors who studied the structure of A. wolgensis coenopopulations consider it to be a steppe species. Forest habitats are not peculiar to it [14–16]. Therefore, forest habitats should be considered secondary for A. wolgensis. In this regard, the presence of A. wolgensis in a pine-birch forest can be explained by some dynamic processes of «forest fighting with steppe» in the process of climatic fluctuations in the Kokshetau highlands.
By density (pcs./100 m2) all populations are divided into three groups: high > 800 pcs./100 m2 (CP-1, CP-7), medium — 150–800 pcs./100 m2 (CP-3, CP-4, CP-6, CP-8), low < 150 (CP-2, CP-5). Steppe CPs (except CP-2) have high density, meadow-steppe ones are medium, forest CPs are low or medium (Table 1).
Table 1 Characteristics of A. wolgensis coenopopulation
No. |
TPC, % |
TPC A. wolgensis, % |
Area of CP, m2 |
Density of plants, pcs./100 m2 |
Number of plants in CP |
Δ |
ω |
I |
CP-1 |
60 |
40.0 |
200 |
1480 |
2960 |
0.47 |
0.62 |
0.65 |
CP-2 |
60 |
0.5 |
300 |
51 |
153 |
0.20 |
0.59 |
1.43 |
CP-3 |
90 |
0.5 |
500 |
169 |
845 |
0.30 |
0.77 |
0.32 |
CP-4 |
100 |
3.0 |
400 |
490 |
1960 |
0.29 |
0.75 |
0.32 |
CP-5 |
60 |
0.5 |
300 |
85 |
255 |
0.33 |
0.86 |
0.06 |
CP-6 |
30 |
0.5 |
1000 |
315 |
3150 |
0.32 |
0.81 |
0.31 |
CP-7 |
100 |
5.0 |
300 |
870 |
2610 |
0.22 |
0.64 |
1.07 |
CP-8 |
100 |
1.0 |
400 |
155 |
620 |
0.27 |
0.84 |
0.63 |
According to the age structure, mature (CP-1) and maturing (CPU-2) species grow only in steppe coenopopulations, all the rest are young (Table 2). By the «delta-omega» ratio, most populations are maturing, but CP-1 is the one which is transitive and CP-2 is young.
The recovery index (Table 1) in steppe ecotopes is very high 0.65–1.43. In other CPs it is less than one. It is especially low in CP-5 (forest habitats) where it is 0.06 (Table 1).
Table 2 Characteristics of A. wolgensis according to the age ratio and «delta-omega» (by Zhivotovsky, 2001)
No. |
Age ratio |
«delta — omega» |
CP-1 |
Mature |
Transitive |
CP-2 |
Maturing |
Young |
CP-3 |
Young |
Maturing |
CP-4 |
Young |
Maturing |
CP-5 |
Young |
Maturing |
CP-6 |
Young |
Maturing |
CP-7 |
Young |
Maturing |
CP-8 |
Young |
Maturing |
All the coenopopulations of A. wolgensis are normal with slight deviations. The exception is CP-2 located on the eastern slope of the hill in the lowland. There is a long-lasting snow cover that provides favorable conditions for resumption. As a result, a large number of young vegetative plants and CPs has antropogenous invasive character. Inconsistency of the CP is bound to an extremely small number of senile plants and seedings.
Conclusion
Thus, the study of A. wolgensis CPs in a latitude direction from Ekibastus in the east to the Kostanay region in the west has found that the majority of CPs has normal character and belongs to maturing CPs according to the spectrum of age stages. This fact proves no negative impact of environmental and anthropogenic factors on the structure of coenopopulations and ensures their recovery and preservation. The gathered data are consistent with the study results of A. wolgensis coenopopulations in the Saratov's region [17, 18].
The research was carried out within the framework of grant financing project of the Ministry of Education and Science, the Republic of Kazakhstan for 2018–2020. «Molecular genetic analysis of gene pools of rare plant species populations in Northern Kazakhstan» № AP05132458, number of the state registration is 0118RK00404.
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