Ecological Significance of Life Forms of Plant Species of Floristic Complexes of the Ketpen Range

The article provides an analysis of life forms of 9 floristic complexes of the Ketpen ridge on the basis of many years of research materials, where life forms were studied in each floristic complex according to the systems of K. Raunkiaer and I.G. Serebryakov. An analysis of life forms according to the system of K. Raunkiaer established the dominance of hemicryptophytes in cryophilic-meadow (74.8%), petrophilic (72.5%) and deciduous-forest (80.3%) floristic complexes of the Ketpen ridge, which accounted for the total number of species of complexes. The second large group in terms of the number of species corresponded to cryptophytes, which predominate in the cryophilic-meadow (65.3%) and psammophilic (45.9%) floristic complexes. Distribution of species according to life forms according to I.G. Serebryakov demonstrated a clear predominance of polycarpic in the meadow complex amount - ing to 82.5%, deciduous-forest – 81.7%, psammophilous – 75.9% and steppe – 79.4% floristic complexes of the total number of species of complexes. An analysis of the meadow floristic complex of the Ketpen ridge according to Serebryakov showed that taproot plants account for 128 species or 20.5%, short-rhizome – 18.4%, long-rhizome – 11.3%, turf-rhizomatous – 4.8%. An analysis of the life forms of the psammophilic floristic complex revealed the prevalence of therophytes (40.7%). It was found that the spectrum of life forms according to Raunkiaer and Serebryakov emphasizes the peculiarities of the floristic complexes of the Ketpen ridge.


MATERIALS AND METHODS
The main factual material for the work was the data collected by the author during expeditionary research on the territory of the Ketpen ridge in 2001-2021. The object of the study corresponded to the floristic complexes of the Ketpen ridge. The aim of the study was to study the life forms of floristic complexes in this region. In field studies, the method of route floristic survey was used. In this case, the classical ecological-geographical and morphological methods were employed. As a basis for the analysis of plant species by life forms, the classification of I.G. Serebryakov and K. Raunkiaer were used, developed in the most detail for angiosperms and conifers. [I. Serebryakov, 1962Serebryakov, , 1964K. Raunkiaer, 1934]. In each floristic complex, on the basis of twenty

RESULTS AND DISCUSSION
On the basis of the results of many years of floristic research, the life forms of 9 floristic complexes of the Ketpen ridge were studied. On the Ketpen ridge, the following types of high-altitude landscape vegetation can be distinguished: desert, desert-steppe, meadow-forest, deciduous-forest, coniferous-forest, cryophilic-meadow, and shrub. Their selection as floristic complexes is quite natural, since these types of high-rise landscape vegetation are zonal for the study area, the vegetation cover, which is characterized by complexity and mosaic, i.e. the heterogeneity of the studied flora is obvious. In view of the foregoing, the following floristic complexes (FCs) are distinguished in the flora of the Ketpen ridge: 1. Desert floristic complex -(DFC). The above-mentioned floristic complexes of the natural flora of the Ketpen Ridge unite species that, according to their ecological and cenotic characteristics and the nature of their distribution, tend to be botanically and geographically homogeneous natural territorial complexes (landscapes) ( Table 1). An analysis of life forms according to K. Raunkiaer (Raunkiaer, 1934) shows a clear predominance of hemicryptophytes, which make up 67.7% of the total number of plant species of the floristic complexes of the Ketpen ridge, where it was revealed that the biomorphic structure of the study area in descending order is as follows: Hc-K−T-Ph-Ch and in general does not differ from the structure of the entire Northern Tien Shan [Rubtsov, 1963], where the leading positions are occupied by perennial herbaceous forms, with the dominance of taproot species, and cryptophytes (C) account for 21.1%, therophytes (T) account for 5.4%, chamaephytes (Ch) -3.0%, phanerophytes (Ph) -1.5%. This ratio is typical for the mountainous regions of Middle Asia (  Table 3). Among the polycarpic herbs, tap-root (caudex) plants prevail, which are characteristic of the climatic and edaphic conditions of the region, where most of them belong to the subclass of tap-root polycarpic (531 species; 28.1% of the total flora). It should be noted that tap-rooted polycarpic in the floristic complexes of the Ketpen ridge are very diverse. A small part of them develop turf, shortened shoots or have bright adaptive features in the structure of root systems. Thus, only 17 species have storage root thickenings. In second place is a group of life forms of herbaceous monocarpic, which are richly represented in PsFC((43.1%), a small percentage of   (14.4%). The following groups of polycarpic are turf-rhizome grasses (3.0%), long-rhizome (2.5%), short-rhizome (0.84%). The second place among DFC life forms is occupied by monocarpic grasses, which make up 16.8% of the total number of species of the complex, among which annual long-term vegetative plants predominate -14.7%. The monocarpic are followed by the type of semi-woody plants (15.7%), with a clear dominance of semi-shrubs, which make up 9.3% of the total number of plant species of this complex. Most of the shrubs are represented in the families Chenopodiaceae (12 species), Asteraceae (7 species), Cistaceae (1 species), Fabaceae (1 species) and others. They are followed by the type of woody plants (15.7%), among which vegetatively immobile erect shrubs predominate -9.1% of the number of woody plant species. Shrubs predominate in families Fabaceae (7 species), Tamaricaceae (4 species), Chenopodiaceae (8 species), Polygonaceae (4 species), Ephedraceae (3 species), Nitrariaceae (1 species), Rhamnaceae (1 species). The desert floristic complex is represented in the study area by rubble-sandy pebble deserts. The species of this complex are distributed on desertified trails of foothill and foothill plains, rising to the lower belt (Artemisia terrae-albae, A. sublessingiana, A. heptopotamica, Anabasis salsa, Nanophyton erinaceum, Kochia prostrata, Haloxylon aphyllum, Krascheninnikovia ceratoides).
The psammophilic floristic complex (PsFC) has a limited distribution, represented by relict sands in the Kegen and Saryzhaz valleys at an altitude of 2200 m above sea level in the western part of the Ketpen range and represents a local type of vegetation. The psammophilous floristic complex forms 25 families, 77 genera, 108 species. An analysis of life forms according to K. Raunkiaer shows a clear predominance of hemicryptophytes, which make up 55.4% of the total number of species of the complex. Cryptophytes are the next group in terms of the number of species, accounting for 45.9%. The third group is 44 species of therophytes or 40.7%. A significant proportion of therophytes is explained by the consequence of the influence of anthropogenic impact on sandy cenoses. Chamaephytes (1.85%) and phanerophytes (1.8%) are represented by single species. Distribution of plant species of the psammophilic floristic complex (PsFC) by life forms I.G. Serebryakov showed the dominance of polycarpic herbaceous plants, which make up 75.9% of the species of the entire complex, the second place is occupied by monocarpic herbs (43.2%), and a small percentage is made up of tree and shrub forms -1.8%. The vegetation cover of relict sands is dominated by Elymus giganteus, which occupies the tops and slopes of sand mounds, and Medicago falcata, which occupies depressions between mounds. Xerophilic shrubs and subshrubs characteristic of the desert type of vegetation are practically absent, except for rare and single specimens of Kochia prostrata and Krascheninnikovia ceratoides. The central part of the sandy massif is noticeably higher than its periphery. Sandy ridges, sometimes up to 10 m high, are elongated from east to west, their slopes are steep. The natural arrangement of the ridges is disturbed by the deflation of the sands, which occurs under the influence of intensive grazing from nearby villages. As a result of deflation, sandy ridges overgrown with vegetation are deformed into bare or almost bare sand mounds of various sizes. Steppe floristic complex. Steppe floristic complex (SFC) is represented on the Ketpen ridge by steppe and mountain-steppe eco-geographical groups, the species of which are distributed mainly in the foothills, low mountains, middle and upper mountain belts, which is explained by the variety of orographic and soil-climatic conditions in the mountains. The steppes are occupied by macro-slopes of southern exposure, and also the southern slope of the Ketpen ridge. The Tien Shan steppes are more ancient in relation to the flat Kazakh steppes, as they already existed in the Paleogene peneplain of the Tien Shan, while most of the flat Kazakhstan was covered by the sea. The steppes in the Northern Tien Shan were formed in the Pleistocene on the basis of a cryophilic floristic and coenotic complex as well as constitute an independent center for the formation of steppe vegetation [Rubtsov, 1954]. The total number of species registered in the SFC is 520, which belong to 200 genera and 36 families. The analysis of life forms according to Raunkiaer showed the predominance of hemicryptophytes (75.2%) and therophytes (22.8%). Cryptophytes account for 18.9%, phanerophytes account for 10.9% of the species of the complex, and chamaephytes account for 4.2% of the flora of the complex (Table 4).
Such a ratio of life forms of chamephytes and therophytes in the steppe complex of the Ketpen range brings this steppe complex closer to the desert flora, where a high proportion of these groups is characteristic of arid zones (Poa bulbosa, Colpodium humile, Carex pachystylis, Tulipa kolpokovskiana, T. biflora, Euphorbia rapulum, Leontice incerta, Eremopyrum orientale, Trigonella arcuata, Strigosella africana, Tetracme quadricornis, Allysum desertorum, Meniocus linifolius, Lepidium perfoliatum, Koelpinia linearis). As it can be seen from the data (Table 4)  Meadow floristic complex (MFC). The meadow complex is well developed in the mountainous area and is widely represented in the upper and middle belts, where meadow vegetation is especially rich and diverse in the river valleys of the piedmont plains of the Ketpen ridge. This complex includes a number of families whose representatives are associated exclusively with meadow habitats. The meadow floristic complex is formed by 47 families, 230 genera, and 624 species. The biomorphological spectrum according to Raunkiaer shows a clear dominance of 2 types of life forms: hemicryptophytes, which make up 79.2% of the total number of species of the complex, and cryptophytes, 34,2%. Analysis of herbaceous forms according to Serebryakov's classification showed the dominance of herbaceous polycarpic (494; 82.5%), which is typical for temperate floras. Herbaceous monocarpic play a lesser role in the composition of the flora (115 species; 19.0%). Of the polycarpic taproots, there are 128 species or 20.5%, rhizomatous -76 species (12.1%), short-rhizomatous -115 species (18.4%), long-rhizomatous -71 species (11.3%), turf-rhizomatous -30 species, or 4.8%, tuberous -17 species, or 2.7%, onion -7 species, In the cryophilic-meadow complex, high-mountain alpine meadows belong to the primary meadows, which are considered as a long-term native vegetation type in the upper part of the high-mountain belt of the mountains of Middle Asia. Genetically, the alpine forms of plants turn out to be quite diverse in their origin, but in their overwhelming majority they gravitate towards the arctotertiary floristic complex. Most of the genera to which the majority of characteristic components of alpine formations belong are of Asiatic origin. They are secondary elements of mountainous mesophilic vegetation of various types (light forests, meadows, stony and rocky cliffs, etc.), which originally existed in a forest environment [Rubtsov, 1965].

Wetland and coastal-aquatic floristic complex (WCAFC)
The species of the wetland and coastal-aquatic floristic complex are confined to water bodies, along the banks and valleys of rivers, where they form aquatic, wetland and meadow (coastalaquatic) ecological-geographical groups. On the Ketpen ridge, this assemblage is also richly and diversely represented; it includes a number of families where representatives of this assemblage are associated exclusively with aquatic and coastal aquatic habitats. The total number of species registered with the WCAFC is 211 species, which belong to 101 genera and 39 families. The ratio of life forms of WCAFC species according to K. Raunkiaer demonstrates the absolute predominance of hemicryptophytes, which accounted for 61.5% of the total number of species of the complex, and cryptophytes (38.4), which is typical for aquatic and coastal aquatic communities. The ratio of life forms according to Raunkiaer and Serebryakov emphasizes the peculiarities of the floristic complexes of the Ketpen ridge.

Petrophilic floristic complex
Petrophilic floristic complex (PFC) is quite widespread and well represented on the stony slopes, pebbles, and screes of the Ketpen ridge. The total number of species registered in the PFC is 845 species, which belong to 298 genera and 69 families. An analysis of the life forms of species of the petrophilic floristic complex according to Raunkiaer showed the predominance of hemicryptophytes -83.5% of the species of the complex. In second place in terms of the number of species are cryptophytes -32.6%, therophytes -21.2%. chamaephytes make up 13.8% and phanerophytes 9.7%; thus, the PFC is dominated by hemicryptophytes and cryptophytes. Analysis of PFC life forms according to Serebryakov established the predominance of polycarpic herbaceous plants -526 species; 72.5% of the total number of species of the complex. Of the type of polycarpic herbs, the dominant group is taproot herbaceous, characterized by the powerful development of underground plant organs, which number 230 species (27.2%). The following groups of polycarpic are short-rhizome 72 species (8.5%), long-rhizome (7.1%), turf-rhizome (4.0%), onion -39 species (4.6%), tuberous -12 species (1.4%). The second place among PFC life forms is occupied by herbaceous monocarpic, which play a much smaller role in the composition of the petrophilic flora -180 species, accounting for 25.3% from the total number of species of the complex, among which annual long-vegetating plants predominate -17,3%. Monocarpic are followed by the type of semiwoody plants (23.5%), with a clear dominance of shrubs and semi-shrubs making up 13.8% of the total number of plant species of this complex (Rhamnus cathartica, Ephedra equisetina, E. Intermedia, Cerasus tianschanica, Juniperus sabina, Atraphaxis frutescens). Shrubs are presented in families Rosaceae (19 species), Fabaceae (17 species), Chenopodiaceae (7 species), Salicaceae (7 species), Polygonaceae (5 species), Grossulariaceae (4 species), Ephedraceae (4 species). Subshrubs are contained in families Lamiaceae (6 species), Asteraceae (21 species), Convolvulaceae (2), Fabaceae (1) and others.

Coniferous forest floristic complex (CFFC)
The total number of species registered with the CFFC is 315 species, which belong to 159 genera and 54 families. The coniferous-forest complex (CFFC) was formed on the basis of Turgai, much more complex, polydominant forest cenoses, by differentiation and some simplification of them under the influence of later cooling and continentalization of climatic conditions. Spruce forests of the Tien Shan are a mountainous Middle Asian variant of taiga forests. Their separation from the general complex forest Turgai complex occurred simultaneously with the pleistocene uplift of the Tien Shan. The coniferous-forest complex of the Ketpen ridge is represented by boreal species, which is 80.3% of their total number. In the spectrum of life forms of the coniferous-forest complex, according to Serebryakov is dominated by herbaceous taproot plants -253 species, accounting for 70.0% of the total number of species. The second place in terms of the number of species is followed by short-rhizomatous plants, represented by 101 species (32%). Long-rhizome plants are represented by 40 species (12.6%), turf plants by 15 species (4.7%). There are 9 types of tubers (2.8%), onions -8 types (2.53%) (12.5%). The third place in terms of the number of species is occupied by shrubs -19 species or 8.2%. These include species common on the slopes Juniperus turkestanica, Lonicera altmannii, Rosa alberti and others.

Deciduous forest floristic complex (DFFC)
The origin of mountain deciduous forests in Middle Asia dates back to the beginning of the Tertiary period, in the Oligocene. At the same time, the type of deciduous forests was formed due to the Turgai invasion and, to a greater extent, due to migration from East Asia, which could have occurred by the end of the Tertiary period, by the time of the Alpine orogeny [Rubtsov, 1956]. The deciduous-forest floristic complex on the Ketpen ridge is represented by boreal species, where floristic elements of the boreal type play a significant role: mountain middle-asian, mountain central-asian, palearctic, holarctic, which is 96.6% in DFFC. The formation of forests with the participation of Populus tremula L. in the Tien Shan is associated with vegetation migrations in the pleistocene. The formation of forests from Malus sieversii (Ledeb.), M. Roem., Armeniaca vulgaris Lam., and a small fragment of walnut forests (Juglans regia L.) are older and autochthonous in origin. The relict nature of these forests is proven by the fragmentation of their geographical area, species composition, geographical, ecological and floristic connection with the relict forests of the Western Tien Shan [Rubtsov, 1956]. The total number of species registered in the Deciduous-forest floristic complex (DFFC) is 184 species. They belong to 94 genera and 35 families. The share of shrubs in the deciduous-forest complex is 24,4% (45 species), the share of trees is 8,7% (16 species). Shrubs are richly represented in families Rosaceae (20 species), Salicaceae (7 species), Fabaceae (12 species), Caprifoliaceae (4 species), Tamaricaceae (2), Grossulariaceae (4), Celastraceae (1), Rhamnaceae (3). Trees are represented by families Rosaceae (6 species), Rhamnaceae (1), Salicaceae (4), Elaeagnaceae (1). Shrubs are represented by such species as: Crataegus songarica, Crataegus altaica, Berberis sphaerocarpa, Cotoneaster multiflora, Cotoneaster melanocarpa, Lonicera tatarica, Rosa spinosissima, Rosa platyacantha and others. The study of DFFC life forms according to the classification of K. Raunkiaer (1934) showed a clear predominance of hemicryptophytes -66.8% of the total number of species of the complex, constituent cryptophytes -36.2% and phanerophytes -33.1%. The remaining types are represented by an insignificant number of species: chamephytes -2.1%, therophytes -9.4%. The analysis of life forms according to Serebryakov for deciduous-forest flora showed the predominance of herbaceous plants (123; 70.6%). Of these, the vast majority belong to herbaceous polycarpic (106; 66,8%), which is typical for the floras of the temperate zone. Herbaceous monocarpic play a much smaller role in the composition of the flora (17 species; 9.4%).

CONCLUSIONS
Thus, the analysis and comparison of life forms of floristic complexes of plant species of the Ketpen ridge is heterogeneous, which reflects the specifics of floristic complexes, showing the features of plant adaptation to changing soil and climatic conditions during florogenesis. The ratio of life forms according to Raunkiaer and Serebryakov emphasizes the peculiarities of the floristic complexes of the Ketpen ridge. The basis of the flora of the Ketpen ridge is hemicryptophytes, which corresponds to the modern climate of the territory and the mountainous Middle Asian nature of the flora. The high content of cryptophytes, chamaephytes and therophytes shows the influence of arid ancient Mediterranean elements. An insignificant number of phanerophytes is explained not only by hydrothermal, climatic and geological conditions that limit the distribution as well as formation of coniferous-forest and deciduous-forest vegetation in the mountains of the Ketpen ridge, but also by human economic activity, which led to the destruction of forests. An analysis of the floristic complexes of the Ketpen ridge showed that the formation of floristic complexes, especially of the coniferous-forest, cryophilicmeadow, deciduous-forest upper and middle belts, proceeded for a long time. The flora of the Ketpen ridge occupies an intermediate position, between the mountain Middle Asian and Siberian-Altai flora, while a large percentage of boreal elements, on the one hand, indicates its connection with Siberia, western Altai, and, on the other hand, with Western and Central Asia. The settlement of plants in the study area proceeded from the north, northeast, and from the south.