3.1. Structure of coenopopulations of Alhagi pseudalhagi
The location of the studied cenopopulation of
A. pseudalhagi and the conditions of its existence in the arid conditions of the Atyrau region are presented in
Figure 1. For the first time, a comprehensive description of the structure of cenopopulations, species composition, and its resources depending on the position of the relief is given. Data on the distribution of species by ecological groups showed that in the cenopopulations of plants with
A. pseudalhagi, xerophytes (53.9%), mesoxerophytes (15.8%), and mesophytes (14.3%) dominate. This fact is also confirmed by the ratio of ecological groups of plants according to humidification conditions. For example, the predominant number of species belongs to xerophytes, accounting for more than half of the species, while other ecological groups occupy a small proportion of the populations.
Cenopopulation 1 (Limonium suffruticosum (L.) Kuntze – Alhagi pseudalhagi (M.Bieb.) Desv. ex Wangerin – Herbaxerophytica) is located in the vicinity of Imankara Mountain. The total projective cover (hereinafter referred to as TCC) of vegetation was 40-50%. Relief plain with a slight elevation difference, soils - loamy, brown, with numerous outcrops of chalky rocks. The surveyed area is used for cattle grazing in spring. Vegetation consisted of two tiers: shrub, 40-55 cm (Atraphaxis spinosa L.), and herbaceous, 15-35 cm. The basic summarized species in a population are Limonium suffruticosum, Alhagi pseudalhagi, Ephedra distachya L., Anabasis salsa (C.A. Mey.) Benth. ex Volkens, Atraphaxis spinosa L., Ferula nuda Spreng., Centaurea scabiosa L., Scabiosa isetensis L., Tanacetum santolina C. Winkl., Kochia prostrata (L.) Schrad. and others. In the described population all age groups of A.pseudalhagi with dominance of middle-aged generative plants. The status of cenopopulation 1 can be characterized as stable, and capable of self-renewal.Soil type: flat area in front of the chalk mountains of Imankara, soils - loamy, dry, grey-earth, heavily gravelly, of a basic nature.
Cenopopulation 2 (Alhagi pseudalhagi - Salsola foliosa (L.) Schrad. - Limonium gmelinii(Willd.) Kuntze) is located in Zhangyr river valley. TCC was 75%. Soils are chestnut, and loamy. Vegetation forms 3 tiers: woody, 120-150 cm (Elaeagnus angustifolia L.), shrub, 70-90 cm (Tamarix laxa Willd.), and herbaceous, up to 50 cm high. The following species are part of the population: Cynodon dactylon (L.) Pers., Salsola orientalis S.G. Gmel., Plantago major L., Trifolium fragiferum L., Polygonum aviculare L., Limonium gmelinii (Willd.) Kuntze, Echinochloa crus-galli (L.) Beauv., Xanthium strumariumL., Solanum dulcamara L., Potentilla spuria A.Kern., etc. The area is actively exploited for livestock grazing, which leads to degradation of the vegetation cover; the degree of degradation is 50-55%. The population is normal, young, and dominated by pre-generative and young generative individuals.Soil type: soil is light chestnut, loamy, moist, without the presence of rocky elements. This area is flooded in the spring.
Cenopopulation 3 (Alhagi pseudalhagi - Herba varia) is located in the Coneu River valley. TCC was 75%. Soils are light- chestnut, loamy, in some places salty. Vegetation is degraded by 25-30% due to active cattle grazing. The vegetation is composed of 3 tiers (one shrub, 60-70 cm high, and 2 herbaceous, 30-50 and up to 20 cm high). The following species were found in the population: Alhagi pseudalhagi, Tamarix laxa Willd., Salsola foliosa (L.) Schrad., S. orientalis S.G. Gmel., Persicaria amphibia (L.) Delarbre, Butomus umbellatus L., Plantago major L., Mentha arvensis L., Trifolium fragiferum L., Zygophyllum fabago L., Limonium gmelinii (Willd.) Kuntze, etc. Cenopopulation 3 is A.pseudalhagi of the normal type, young with a predominance of young generative individuals. Soil type: soil is light chestnut, loamy, moist, without the presence of rocky elements. This area is flooded in the spring.
Cenopopulation 4 (Alhagi pseudalhagi - Glycyrrhiza glabra L. – Herbaxerophytica) is described in Taisoigan sands. TCC was 50-55%. The relief is flat; soils are sandy, with clay outcrops in places. There are traces of cattle grazing. Vegetation degradation by 10-15% is observed. The cover is composed of 2 tiers: high grass, 40-65 cm, and low grass, 15-25 cm. The species composition includes the following species: Euphorbia seguieriana Neck., Melica taurica K. Koch, Agropyron fragile (Roth) P. Candargy, Achillea micrantha Willd., Kochia prostrata (L.) Schrad., Glycyrrhiza glabra L., Carex physodes M.Bieb., Arnebia decumbens (Vent.) Coss. & Kralik, Limonium suffruticosum (L.) Kuntze, etc. Cenopopulation 4 is normal, stable, medium-aged, and dominated by medium-aged generative plants.Soil type: Typical sandy massifs, at a depth of 40cm - wet loans.
When describing the population structure, we also took into account the age spectrum of
A. pseudalhagi individuals. In populations of perennial plants, all individuals are characterized by a set of biomorphic traits that determine their age differentiation. Determining age states (biological age) is much more important for population studies (
Figure 2).
Thus, the analysis of age spectra showed that сenopopulation 1 and сenopopulation 4 have a middle-aged and stable status, сenopopulation 2 and сenopopulation 3 are normal, young. These characteristics make it possible to recommend all studied populations for organizing the procurement of raw above-ground organs.
Age structure is one of the most important traits of сenopopulation. It reflects the vital state of the species in the cenosis, as well as such important processes as the intensity of reproduction and the rate of generation change. It shows the ability of the population system to maintain itself and the degree of its resistance to the influence of negative environmental factors, including anthropogenic impact.
3.2. Floristic composition of A. pseudalhagi cenopopulations
As a result of the analysis of herbarium material collected during field studies, 63 species from 54 genera and 26 families were observed growing as part of populations with
A. pseudalhagi. Systematical analysis showed that the leading families by species composition are Poaceae (12.7%), Asteraceae (12.7%), Chenopodiaceae (9.5%), and Fabaceae (9.5%) (
Table S1). The leading 4 families include 28 species and 23 genera, representing 44.4 and 42.6%, respectively, of the total flora composition.
Comparison of the species composition of сenopopulation showed that the greatest number of species was recorded for сenopopulation 1 - 25, the minimum for сenopopulation 4 - 21, while сenopopulation 2 and сenopopulation 3 have the same quantitative composition - 24 taxa each.
Significant differences in the species composition of A. pseudalhagi сenopopulation were noted. Thus, the maximum index of similarity in floristic composition was found between сenopopulation 2 and сenopopulation 3 - 0.122; the minimum index was between сenopopulation 1 and сenopopulation 2 - 0.042.
The obtained data can be explained by the fact that in the valley of the rivers Zhangyr and Coneu similar soil and climatic conditions and the degree of anthropogenic pressure are observed, while сenopopulation 1 and сenopopulation 4 are located in significantly different conditions. Thus, in the vicinity of Imankara Mountain 25 species grow, in the valley of the rivers Zhangyr and Coneu – 24 species each, on the sands of Taisoigan - 21 species. Significant degradation of vegetation cover in the valley of the Zhangyr and Coneu rivers due to anthropogenic pressure is noted, which is confirmed by a significant number of weed-ruderal elements (Onopordum acanthium L., Xanthium strumarium L., Cynodon dactylon (L.) Pers., Polygonum aviculare L., Convolvulus arvensis L., Echinochloa crus-galli (L.) P. Beauv., etc). However, grazing has no depressing effect on A. pseudalhagi, as this species is practically not eaten by domestic animals. The analysis of species by occurrence showed that points IV and V were noted only for Alhagi kirghisorum Schrenk and Limonium suffruticosum (L.) Kuntze (3.2% of the total number of species). Score III was recorded for 7 species or 11.1% (Artemisia terrae-albae Krasch., Salsola foliosa (L.) Schrad., Carex physodes M.Bieb., Euphorbia seguieriana Neck., etc.), and a score of II - 17 species or 26.9% (Artemisia arenaria DC., Peganum harmala L., Xanthium strumarium L., Tamarix laxa Willd., etc.). The main number of species had a score of I - 49 taxa or 77.8% (Allium sabulosum Steven ex Bunge, Atriplex cana C.A. Mey., Salsola orientalis S.G. Gmel., Arnebia decumbens (Vent.) Coss. & Kralik, Onosma stamineum Ledeb., Alyssum lenense Adams, Trifolium fragiferum L., etc.). The analysis of life forms showed the predominance of herbaceous perennials (33 species or 52.4%), the second position is occupied by minor shrubs (15 species or 23.8%), the third – by semi-shrubs (9 species or 14.3%). Shrubs account for 5 species (7.9%) and trees for 1 species (1.5%).
It should be noted that the steppe and semi-desert territories of Kazakhstan are characterized by the predominance in the floristic composition of populations of representatives of the families Poaceae, Polygonaceae, Asteraceae, Chenopodiaceae and Fabaceae. Thus, the Chenopodiaceae family is the most characteristic family for the steppes and deserts of the globe. The large number of species in this family indicates the presence of significant areas of saline or heavily saline areas. The species of this family mainly belong to the group of plants with summer-autumn vegetation. The Asteraceae family serves as a price-maker in the flora of arid areas such as steppes and deserts. The Poaceae family includes groups of species that play the role of edificators and subedificators of steppe and desert flora (genera Agropyron, Anisantha, Poa, Eremopyrym), as well as forming small areas of vegetation along water sources (Cynodon dactylon (L.) Pers.). The Turanian flora is characterized by the first place of the families Chenopodiaceae and Polygonaceae; a large number of species of the families Asteraceae and Fabaceae indicates the influence of the Mediterranean flora proper.
3.3. Morphological differences between cenopopulations and resource potential of A. pseudalhagi
Biomorphological studies of various cenopopulations made it possible to determine that the range of A. pseudalhagi in the Zhylyoysky, Kzylkoginsky, and Kurmangazinsky districts of the Atyrau region differ in a systematic structure, morphometric indicators and age spectrum. It is noted that previous studies of natural populations of A. pseudalhagi both in the Atyrau region and in Kazakhstan as a whole have not been carried out.
Morphometric indicators of individuals from 4 cenopopulations differed in the number of individuals per 1 m
2, the height of generative shoots, and their number per 1 individual.
Table 2 presents the morphometric parameters of
A. pseudalhagi in the studied populations of the Atyrau region. By the data obtained, the maximum number of individuals per 1 m
2 was noted for cenopopulations 2 (2.6 pieces) and cenopopulation 3 (2.2 pieces), and the minimum growth density was for cenopopulations 4 (0.5 pieces).
The main morphological difference between individuals of the four cenopopulations of
A. pseudalhagi is the height of the plants. In terms of growth indicators, the maximum values were noted in cenopopulations 3 – 30.5 cm, and the minimum in cenopopulations 1 – 24.4 cm (
Figure 3).
Accordingly, individuals in cenopopulation 3 with the maximum height of generative shoots had significantly the largest number of generative shoots per individual (6.1 pieces), and in cenopopulation 4 - the largest number of generative shoots (2.9 pieces). Сenopopulation 1 occupies an intermediate position in morphometric parameters between cenopopulation 2 and cenopopulation 4. Probably, the difference between the cenopopulations is due to differences in climatic conditions and the degree of anthropogenic load at the growing points.
The principle of basic coordinates showed the presence of similar basic parameters of coenopopulations between coenopopulations 2 and 3, and between 1 and 4 (
Figure 4). This group association is associated with the geographical proximity of these two groups. The most significant significant difference in the morphological parameters of individuals between different populations was observed for cenopopulations 1 and cenopopulations 4, while cenopopulations 2 and cenopopulations 3 have similar parameters.
The following indicators were correlated: Plant height, cm (PH); plant diameter, cm (PD) number of generative shoots (GS), pieces number of individuals per 1 sq.m, pieces (NI); Soil type (ST); Cenopopulation (CP). The correlation showed the positive influence of factors as the main ones for identifying and assessing the state of
A. pseudalhagi coenopopulations (
Figure 5).
The one-way factor analysis (ANOVA) showed the significance (P < 0.05) of the influence of the soil type (ST) factor on the plant indicator in different coenopopulations: plant height (PH) with a value of 4.16e-06; plant diameter (PD) with a value of 6.55e-13; number of generative shoots (GS) with a value of 3.79e-10; pieces number of individuals per 1 sq.m, pieces (NI) with the value 3.59e-06.
The dendrogram showed that there is a small mixture of individuals of coenopopulations 2, 3, and 4, as a special group that does not climb from the mountainous territory. The mountain's first coenopopulation is characterized by a low mixture. All populations differed significantly from each other in the number of individuals and in the growth of significant differences (
Figure 6).
And between cenopopulations 2 and 4, she pointed out differences from each other. There were significant differences in the number of generative shoots between cenopopulations 1 and 4 and cenopopulations 2 and 3. Probably, the difference between the cenopopulations was due to the difference in climatic conditions of the arid zone and various natural conditions for the growth of the species.