Distribution of Rh. rosea in East Kazakhstan
The study was carried out in the period 2015-2020 on the Ivanovo Ridge by stationary research methods with the laying of monitoring sites in the upper reaches of the Bolshaya Transverochka River (50 ° 19 '13.5" s. w., 83 ° 45 '11.0" w.d.). The studied species are distributed on the ridges of the Kazakhstan Altai and Tarbagatai.
Based on the long-term herbarium collections of the authors of this work stored in Altai Botanical Garden (further termed ‘Alt.’) and Astana Botanical Garden (further termed ‘Ast.’ ), as well as a result of the revision of the herbarium materials of the Moscow State University (MW) (Seregin, 2020) and the herbarium of the Institute of Botany and Phytointroduction (AA), the distribution of Rh. rosea in Eastern Kazakhstan was revealed. In addition the literature data are taken into account (Perrino et al.; Zair et al., 2020; Artemov, 2020; Kotuhov, 2005; Isayev, 1993; Zibzeyev, 2015; Kupriyanov, 2020) the distribution of Rh. rosea in the study region. The distribution map of the view is shown in Figure 1.
Specimens examined:— WESTERN ALTAI: was revealed: Ivanovski ridge: vicinity of Ridder mountain, Khorizovka river narrow, 29.VI.1936, Rubam & Mikhailova (АА); near the small liver, between blocks of moraine debris, 15.VI.2015, Kubentayev (Ast.); top of Poperechka river, along the river bank, 03.VII.2012, Kotukhov (Alt.); «Prohodnoj belok», along the damp rocky places, 10.VI.2016, Kubentayev (Ast); vicinity of Rodonovoi river, along the damp rocky places, 10.VI.2015, Kotukhov (Alt.); Ulbi ridge (northern faces of Kreslova mountain, Ridder region, 26.VII.1937, Kuznecov (АА, MW); Lineiski ridge: valley of Chernaya Uba river, south- western slope on the altitude of 1830 m., 03.VII.1998, Kotukhov (Alt.); Koksi ridge: in Latuniha river-valley, 27.VI.2003, Kotukhov (Alt.); in Chernaya Uba river-valley, 03.VI.2003, Kotukhov (Alt.); Ulbi ridge: Bolshoi Turgusun river-valley, 15.VI.2004, Kotukhov (Alt.); top of Tatarka river, 15.VII.2004, Kotukhov (Alt.); Ubi ridge: Belaya Uba river-valley, 12.VI.2006, Kotukhov (Alt.). IN SOUTHERN ALTAI: Chindagatui mountains: Southern Alai ridge, moist meadow in the down part of the slope, 1800m., 27.VII.1986, Ivaschenko & Utebekov (AA); Narym ridge: Koksar mountain, 7.VII.1973, Mikhailov & Stepanova (AA); Southern Altai-Tarbagatai ridge: Burkhat pass, northern slope at the top of the forest, 14.VII.1973, Isayev (АА); left bank of Karakaba river, northern slope, 2100 m., 28. VI.1988, Ivaschenko (АА); vicinity of Chernovoye village, in the upper part of the forest, 15.VI.2016, Kubentayev (Alt.); in the vicinity of the bridge across Karakaba river along the Austria road, damp rocky places, 18.VI.2016, Kubentayev (Alt.); southern-eastwards of Enbek village, at the top of a ridge in the alpine belt, 01.VI.2017, Kotukhov (Alt.); Sarymsakty ridge: Kumshybai, by the stream along the path to the waterfall, damp meadows, 22. VI.1986, Ivaschenko (AA); at the top of Solonechnaya river, 22.VIII.2010, Kubentayev & Zhumagul (Ast.); in the vicinity of Topkain river, alpine meadows, 10.VIII.2020, Kubentayev (Ast.); Western Listvyaga ridge: at the foot of Schebniuha hill, valley of mountain river, 10.VII.2019, Kubentayev (Ast.); Repnoie river head, Kubentayev & Zhumagul, 22.VII.2019; vicinity of Aksharbak village, Katon-Karagai region, Verhkatun riverhead, 22.VII.2020, Kubentayev (Alt.).
"Distribution of Rh. rosea in East Kazakhstan " was obtained by ArcMap
Ecological-biological and phytocenotic structure of populations Rh. rosea
The study of population of Rh. rosea was carried out at 4 loci: Ivanov ridge (4 population); 2) Sarymsakty ridge (2 Population); 3) Southern-Altai Tarbagatai ridge (2 Population); 4) Western Listvyaga ridge (2 Population) belonging to the territory of Kazakhstan Altai (22. Altai), according to the floristic zoning of Kazakhstan (Kazakhstan flora.1957). This species populates on wet moss-covered rocks, rocky hills, near snowfields, over growing morains, among moss along the river banks, in the upper limit of mossy cedar-larch forests (Fig. 2)
- Population (Macropodium nivale–Angelica archangelica–Rh. rosea ) is timed to the western slopes of shallow ravines of shallowed river beds. Population is studied Burkhat pass, Sarymsakty ridge, (49°07'49.9"N, 86°02'19.8"E) in the altitude of 1950-2050 m. Projective cover (PC) is 55%. Standing grass crop is formed diffusively, along the rock cracks, between the debris of blocks and in degradations where the fertile soil layer accumulates. There are Coptidium lapponicum, Aquilegia glandulosa, Sanguisorba alpina, Rumex acetosa, Bistorta elliptica, Trollius altaicus, Macropodium nivale, Geranium albiflorum and etc. in the community. The population of Rh. rosea as a rule, is represented by all the age states with predominance of generative adult species. The habitat conditions are livable to the development.
- Population (Carex stenocarpa+C. orbicularis+Rh. rosea +Dracocephalum grandiflorum) occurs in well overgrown steep moraine slopes, Sarymsakty ridge, Solonechny river (49°05'34.7"N, 85°29'10.3"E). Moraine hillocks are located on the steep northwestern slopes of the ridges in the altitude of 1900–2100 m above sea level with closely located snowfields. PC is 80-90%. Typical species for these communities are Festuса kryloviana, Allium schoenoprasum, Hedysarum neglectum, Anemonastrum narcissiflorum, Thalictrum alpinum, Papaver nudicaule, Neogaya simplex, Gentiana grandiflora, Pedicularis oederi, P. amoena, P. violascens, Oxytropis alpine, rare enough Gentiana algida, Papaver nudicaule, Festuca altaica. Betula rotundifolia should be noted among the shrubs, which forms separate small clumps. Favorable water and temperature conditions, high humus content in the soil determine the lush development of vegetation, which negatively affects the populations of Rh. rosea due to its low competitiveness.
- Population (Alchemilla gottsteiniana–Polygonum ellipticum+Rh. rosea ) is timed to smooth slopes of alpine meadows on the alpine meadow soils, Southern Altai Tarbagatai, Karakaby depression, KaraKaba river-valley (49°04'06.8"N, 86°05'14.8"E). Total PC is 85 %. The shrub layer, where Lonicera altaica, Potentilla glabra is rarely occurred, is poorly expressed, Spiraea media shrubs are singly noted. Koenigia alpina, Phleum alpinum, Valeriana dubia, Rumex acetosa Galium boreale, Iris ruthenica, Dracocephalum grandiflorum, Aster alpinus, Papaver nudicaule, Vicia cracca, Sedum hybridum, Pedicularis achilleifolia, Pachypleurum alpinum, Oxytropis alpina, Gentiana algida, Crepis chrysantha is often found in the phytocoenosis. The population of Rh. rosea in this type undergo degradation. Gradually expanding, meadow vegetation displaces Rh. rosea from familiar habitats. Ontogenesis is dominated by generative and senile individuals, seed reproduction is absent.
- Population (Rh. rosea +herbo variae) occupy excessively cold, moderately humid rocky peaks and southwestern slopes of weakly closed pressure moraine ridges in the altitude limit of 2000-2300 m. above sea level, Southern Altai Tarbagatai ridge (49°10'04.4"N, 86°16'07.3" E). PC is 30%. The vegetation cover is poorly developed and relatively poor in terms of species. The most common herbaceous plants are Сагех aterrima, С. orbicularis, Schulzia crinita, Micranthes punctata, Papaver croceum, Anemonastrum narcissiflorum, Salix rectijulis is relatively common among shrubs. In these population Rh. rosea is represented mainly by aging generative and deeply senile individuals. Analysis behind the developmental state of Rh. rosea in the upper limit of its distribution give grounds to consider these habitats as extreme.
- Population (Hedysarum neglectum+Carex orbicularis+C. aterrima+Rh. rosea ) occurs in weakly covered moraine ridges, Ivanov ridge, upper parts of Big Poperechka river (50°19'13.5"N, 83°45'11.0"E). They are usually along the northwestern microslopes, on the altitude of 2000-2300. PC is 70%. There are Rhodiola algida, Dryas oxyodonta, Bergenia crassifolia, Hedysarum neglectum, Pedicularis achilleifolia, Neogaya simplex, Oxytropis alpina, Pedicularis amoena, P. oederi and etc. in the community Hedysarum theinum, Saussurea alpina, Schulzia crinita, Luzula spicata occurs comparatively rare. Open areas are richly covered by Polytrichum juniperinum, Р. аlpinum mosses and species from the genus Bryo. Patches of lichens from the genus Cladonia are commonly found. Rh. rosea in the coating occupies no more than 5-6% of the total amount.
- Population (Deschampsia caespitosa + Senecio pratensis + Rhodiola rosea) observed on moderately humid stony-mobile fine-gravel slopes of moraine ridges, Ivanov ridge, passing snow-covered mountain peak (50°15'10.3"N, 83°31'31.8"E) (Fig. 1). It is found on the altitude of 1800-1900 m. above sea level. PC is not more than 40%. In this kind of conditions Rh. rosea grows far from drains, constantly face the lack of moisture. The vegetation cover is not poorly expressed, it is usually found Carex pediformis var. macroura, С. capillaries, Festuca borissii, Helictotrichon altaicum, Lagotis globosa, Callianthemum alatavicum, Saussurea alpina, Dracocephalum grandiflorum here in the community. Dryas oxyodonta, Thalictrum alpinum, Gentiana algida, Eritrichium villosum, Allium schoenoprasum, Pachypleurum alpinum, Crepis chrysantha are found very rarely. Plants do not form dense tangled vegetation. They are found in separate groups or single individuals. Betula rotundifolia, Cotoneaster uniflorus are rare among the shrubs, and Juniperus sibirica is individually found.
- Population (Rh. rosea –Trisetum altaicum–Deschampsia cespitosa) occupy cold waterlogged coastal lake habitats, Ivanov ridge, near Maloye lake (50°18'36.9"N, 83°44'44.7"E), on the altitude of 2000-2100 v. above sea level. population data is timed to northeastern the shores of permanent dammed lakes in the close proximity of the water. PC is 15-25 %. Tangles of Rh. rosea occupy a narrow coastal strip from the very edge of the water no more than 1.5–2 m wide. The vegetation cover is represented by separate plants or small groups of the communities, where Сarex aterrima, Deschampsia cespitosa, Festuca borissii, Trisetum altaicum, Phleum alpinum, Swertia obtuse, Primula nivalis, Rhodiola algida, Sanguisorba alpine, Caltha palustris, Bistorta vivipara, Allium schoenoprasum, Gentiana algidа are often found. Salix lanata and S. rectijulis are relatively rare. In the herbage, Rh. rosea is found relatively abundantly, the population is full-lived.
- Population (Salix lanata–Betula rotundifolia–Rh. rosea ) occupies moderately humid bushy tundra habitats, Ivanov ridge (50°19'36.4"N, 83°48'17.8"E). The communities with the participation of Rh. rosea are timed to northwestern steep slopes on the altitude limit of 2100 – 2200 m above sea level. PC is 50-60%. The herbage is less abundant. Carex aterrima, Trollius altaicus, Pedicularis oederi, Thalictrum alpinum, Macropodium nivale, Geranium albiflorum, Gagea serotine are found in phytocenosis. Betula rotundifolia tangles reach a height of 35-40 m, rarely 50 m in the communities. The population of Rh. rosea in this kind of population is believed extreme. Gradually expanding, tangles of birches crowd them out from their habitats.
- Population (Rh. rosea +Achillea ledebourii–Sanguisorba alpine) occupies coastal and excessive wet meadows, constant verges of the streams, cedar larch wood meadows on the altitude of 1700-1900 m. Western Listvyaga, upper part of Repnaya river (49°21'06.0"N, 85°41'54.8"E). Excessive moisture and light shading throughout the growing season create unfavorable conditions for the development of Rh. rosea .The vegetation cover is well formed, PC is 65–80%. Alchemilla altaica, Primula nivalis, Carex curaica, С. aterrima are often found in the community, while Carex orbicularis, Cerastium davuricum, Bistorta vivipara, Trollius altaicus, Deschampsia cespitosa, Allium schoenoprasum, Myosotis scorpioides, Veratrum lobelianum, Delphinium elatum, Caltha palustris are rarely found. Rh. rosea is timed to areas in the form of narrow ribbons of 1.5 – 2 m width along the coasts. There are no shrubs. In rare cases, Lonicera altaica is noted along the coastline. Rh. rosea forms small clumps on areas bare from grass. Generative individuals of Rh. rosea predominate in the population of this type.
- Population (Rh. rosea –Dichodon cerastoides–Allium schoenoprasum) occupy the shores of mountains streams, wells, drains which bear a temporary nature by the process of melting snow patches and which dry starting from the mid July, Western Listvyaga ridge, in the vicinity of Schebnuiha mountain (49°21'54.0"N, 85°44'58.9"E), on the altitude limit of 2000-2200 m. Rh. rosea plants on the boulders covered by the thick moss cover. The vegetation cover in the phytocenosis is poorly expressed. PC is 30-40%. The following species are found in the community: Primula nivalis, Carex orbicularis, C. aterrima, Bistorta vivipara, Pedicularis oederi, Deschampsia cespitosa, Micranthes punctata, Macropodium nivale, Sanguisorba alpina, Lagotis globosa, Gentiana algida. Populations of Rh. rosea are of normal type, young, full-lived. Habitat conditions can be considered optimal.
Table 1 shows the morphometric parameters of Rh. rosea in the surveyed population of Kazakhstan Altai. According to the data obtained, it was found that the highest number of individuals per 1 m2 was observed in P10 (0.75), P7 (0.68) and P1 (0.56), a relatively low number per unit area was noted in P8 (0.18 ), P 6 (0.21) and in P3 (0.23). For Rh. rosea , in populations with a high number of individuals per unit area, undersized, multi-shoot shrubs with large flowers are characteristic. In the population with a low abundance of Rhodiola per unit area, tall individuals are observed, with loose, low-shoot bushes and relatively small inflorescences. Undersized, multi-stem structure with large flowers prevail per square for Rh. rosea in populations with a high number of species. Tall individuals, with crumbly, low-running bushes and relatively small inflorescences prevail in the population with a low number of Rh. rosea per square. This pattern is explained by the habitat conditions. As a rule, species of Rh. rosea are relatively tall (45-50 cm), have crumbly low-running bushes (6-10 pcs) and small inflorescences (3-4.5 cm) in the forest belt and in tall grass communities. In open, poorly populated areas, and along the valleys of mountain streams, individuals are significantly undersized (20–25 cm), but have a multi-stem structure (30–50 pcs) and large flowers (5.2–6 cm) (Fig. 3). The correlation analysis of morphological and quantitative indicators of Rh. rosea between the studied populations is shown in Fig. 4.
Table 1. Quantitative and morphological indicators of Rh. rosea
Quantative and morphological indicators
|
Number of adult specimens per 1m2
|
Height of generative species at the time of flowering (cm)
|
The number of sprouts per a specimens (pcs)
|
Inflorescence diameter (cm)
|
P 1
|
М
|
0,56
|
24,42
|
21,41
|
5,25
|
SD
|
0,03
|
1,62
|
2,01
|
0,31
|
P 2
|
М
|
0,32
|
48,33
|
15,22
|
4,20
|
SD
|
0,01
|
1,28
|
1,61
|
0,26
|
P 3
|
М
|
0,23
|
35,52
|
6,91
|
4,35
|
SD
|
0,01
|
2,12
|
0,65
|
0,16
|
P 4
|
М
|
0,28
|
26,66
|
20,32
|
3,63
|
SD
|
0,01
|
2,91
|
1,71
|
0,18
|
P 5
|
М
|
0,42
|
47,22
|
28,31
|
5,39
|
SD
|
0,04
|
1,81
|
2,16
|
0,27
|
P 6
|
М
|
0.21
|
38.30
|
12.27
|
4,23
|
SD
|
0.01
|
1.21
|
1.63
|
0,15
|
P 7
|
М
|
0,68
|
31,19
|
36,55
|
5,46
|
SD
|
0,02
|
1,72
|
1,11
|
0,31
|
P 8
|
М
|
0,18
|
49,29
|
8,12
|
3,81
|
SD
|
0,01
|
2,33
|
0,69
|
0,25
|
P 9
|
М
|
0,33
|
45,61
|
6,06
|
3,22
|
SD
|
0,03
|
2,72
|
0,31
|
0,13
|
P 10
|
М
|
0,75
|
32,30
|
37,33
|
4,81
|
SD
|
0,03
|
1,44
|
2,82
|
0,33
|
Ontogenetic state Rh. rosea
The ontogenetic state of Rh. rosea was studied on Ivanov ridge, in the upper parts of Big Poperechka river (50°19'13.5"N, 83°45'11.0"E). Rh. rosea begins to vegetate under a cover of snow starting from mid-May to mid-June in the studied area and when the snow melts, it begins to grow rapidly. It blooms from mid-June to late July. The fruits ripen from August to September. It should be noted that the seasonal rhythm of the species development depends on the height of the location of the population. The species begins to grow from mid-May in the upper limit of the green belt at an altitude of 1700–1900 m. Rh. rosea grows in the second half of June in the alpine belt at an altitude of 2200-2400 m. On average, the growing season lasts 4 months.
The species in the surveyed area reproduces predominantly in a vegetative way due to the division of rhizomes and spread by melt water during the period of abundant snow melting, but in some places seed renewal is noted. Seeds are small, oblong. The seeds shape is curved-pin-shaped. The seeds surface is bare, longitudinally wrinkled. The seed scar is small, slightly protruding, basal, rounded. Seed color is from light brown to hazel. The length is 2,13±0,16 mm, Cv=15.7%; min-max – 1,65 – 2,78 mm, the width is 0,48 – 0,81 mm (0,59±0,07 mm, Cv=17.5%). The weight of 1000 seeds is 0,208 – 0,239 g. Once in the soil, ripen seeds undergo natural stratification over the next 7-8 months. The laboratory germination of Rh. rosea seeds in Petri dish at 18°C in three replications has shown 51%.
Plantlets (Fig.5. (p)) start appearing at the end of May and beginning of June. Emergence of seeding is above-ground. Seed leaf is light green, bare, succulent 3,2±0,09 mm long, 1,8±0,06 mm wide. The plates are oval-ovoid, on short petioles up to 2.8 mm long. Rounded at the apex, sharply tapered at the base turning into a short petiole. The hypocotyl is 3.2 ± 0.07 mm long, up to 1.1 ± 0.03 mm thick, pale green, the basal part is thickened, sharply passes into the embryonic (primary) root. The main root up reaches 2.3 ± 0.08 cm long by the time the cotyledons dry up with a significant number of lateral shortened suction roots. Cotyledons persist until mid or late July. In 2-3 months after germination of the seed, this age condition ends.
At the end of July and beginning of August the species turns into juvenile state (Fig.5(j)). Plants in this phase are characterized by the formation of a rosette of 2.6 ± 0.04 cotyledonous leaves, the presence of a crown bud and 1-2 axillary buds of an open type. The part of the growth sprout (rhizome) does not die off after the end of the vegetation season, but becomes perennial, from which the rhizome is subsequently formed. The seedlings end the juvenile phase at the age of 2-3 years and less often.
Immature phase (Fig.5((im)) is characterized by the growth of vegetative stems of normal type in the structure of medial sprout. The medial sprout of 7,2±0,12 cm height has 6,9±0,21 pieces of natural leaves. The leaves are set by turn, the leaf blade is oblong-ellipsoid at the base and smoothly tapers into a short petiole. The crown and lateral buds of the growing sprout are of a closed type. The growth part of the rhizome is 2.4 ± 0.2 cm in length and 0.6 ± 0.003 cm in thickness. Rhizome branching is observed. The root system is well developed in the horizontal projection 2.8 ± 0.08 cm and in the vertical projection 12.1 ± 1.56 cm (deepened). In the primordial state, plants are on average 2 vegetation seasons. In the future, the plants move to the next age state.
Virginal phase might be seen for 5-7 year and is characterized by the beginning of branching of the medial sprout of the rhizome with the development of a significant number of lateral sprouts of the first order with the development of stems on them. The plants in this age state are 14±1,31 cm in height. The rhizome has 3,6±0,07 pieces of stems of the first order. The medial sprout of the rhizome and some lateral sprouts are 6.2 ± 0.06 cm long at the base and 0.83 ± 1.2 cm across. The root system is well developed, 10.2 ± 1.8 cm in horizontal projection and 16 ± 2.2 cm in vertical projection. The primary root is about 1.8 ± 0.06 cm in thickness. The buds of the renewal of the medial and lateral sprouts are large, of a closed type.
The plants come into the phase of young generative species (Fig.5(g1)) at the age of 8-11 years. Generative stems are formed in medial sprout. Young plants usually generate 2,8±0,06 pieces of generative stems with a depleted inflorescence of 1,8±0,02 flowers and 8,2±2,6 vegetative stems in their first two years. Plants at the age of 8-10, more often 12 years old begin to form generative stems on the sprouts of the rhizome of the first order. The rhizome of the horizontal projection has a thickness of 3.2 ± 0.9 cm and a length of 16.3 ± 0.8 cm. The number of flowers in an inflorescence is 6.3 ± 1.8 pieces, a diameter of 4.8 ± 0.35 cm. The height of the plant is 30 ± 1.8 cm. In this age state, stems begin to form on the sprouts of second-order rhizomes. This age-related condition ends by 18-22 years.
The mature generative individuals (Fig. 5(g2)) include plants aged 22-30 years, characterized by a powerful development of 43 ± 2.1 cm in height. There is an intensive development of generative stems on the medial sprout of the rhizome and sprouts of the first and second orders. Such individuals have 35 ± 3.6 pcs of generative and 42 ± 2.8 pcs of vegetative stems. Inflorescence with 10.6 ± 1.7 flowers, 5.2 ± 0.35 cm in diameter. Abundant flowering and fruiting. Particulation and clone formation are observed.
The plants come into the phase of old generative species at the age of 30-40 years. There is a noticeable predominance of vegetative stems up to 68 ± 3.9 pcs and the formation of a significant number of weakened generative stems 52 ± 2.8 pcs in this age state. There are usually 6.2 ± 0.12 flowers, 3.8 ± 0.35 cm in diameter in the inflorescence. The beginning of the rhizome sprouts death and the formation of extensive foci of the main root necrosis and medial sprout of the rhizome are observed. Also, mass dying of rhizomes sprouts of the first order is typical in this age state.
Senile species (Fig. 5 (ss)) are very rare. It is pretty hard to define the age of this species. According to our data, it comes into this phase at the age of 50-55 years. In this state, extensive foci of necrosis appear almost along the entire length of the main rhizome and its disintegration into separate girders. There are frequent cases of dying of the adventitious roots of the first-order rhizome sprouts. The bushes easily break up into 3-6 clones, form new plants and spread over the population area.
Analysis composition of the flora of communities with Rh. rosea involvement
The spectra analysis of geographic elements of floras of various ranks, including floras of plant communities in the volume of specified classification units (composition of the floras) is one of the main tools of comparative floristry. Composition of the flora is a set of plant species which form communities of any rank and any type of vegetation. From this point of view, the composition of the flora represents the unification of historically and coenotically homogeneous groups of species within the syntaxon, which makes them the most important indicator of the vegetation cover from the level of a particular phytocenosis to altitudinal-belt units. This reveals the most important meeting points between floristry and geobotany.
As a result of data processing of field studies and herbarium collections, it was found that the composition of the flora of communities with the participation of Rhodiola rosea includes 140 species belonging to 39 families and 104 genera, which is 14% of the Altai highland flora (AHF), where 996 species of vascular plants from 325 genera and 80 families are registered (Table 2). Herbarium collections are kept in the herbarium of the Astana Botanical Garden and the Altai Botanical Garden.
Table 2. Floristic composition of Rh. rosea composition of the flora of Kazakhstan Altai
№
|
The species name
|
1 ⃰
|
2
|
3
|
4
|
5
|
Amaryllidaceae
|
1.
|
Allium schoenoprasum L.
|
III
|
C
|
Bbp
|
HP
|
euras.
|
2.
|
Allium flavidum Ledeb.
|
I
|
C
|
Bbp
|
MP
|
as.
|
Apiaceae
|
3.
|
Neogaya simplex (L.) Meisn.
|
II
|
HC
|
Tp
|
P
|
euras.
|
4.
|
Angelica archangelica L.
|
III
|
HC
|
Tp
|
М
|
as.
|
5.
|
Angelica decurrens (Ledeb.) B.Fedtsch.
|
II
|
HC
|
Tp
|
М
|
as.
|
6.
|
Bupleurum multinerve DC.
|
II
|
HC
|
Tp
|
MP
|
as.
|
7.
|
Sajanella monstrosa (Stephan ex Schult.) Soják
|
III
|
HC
|
Tp
|
P
|
as.
|
8.
|
Schulzia crinita (Pall.) Spreng.
|
IV
|
HC
|
Tp
|
P
|
as.
|
Asteraceae
|
9.
|
Aster alpinus L.
|
II
|
HC
|
Tp
|
X
|
holarc.
|
10.
|
Achillea millefolium L.
|
II
|
HC
|
Lrp
|
М
|
holarc.
|
11.
|
Saussurea alpina (L.) DC.
|
II
|
HC
|
Lrp
|
P
|
holarc.
|
12.
|
Senecio nemorensis L.
|
I
|
HC
|
Brp
|
М
|
euras.
|
13.
|
Solidago virgaurea L.
|
III
|
HC
|
Brp
|
М
|
euras.
|
14.
|
Hieracium virosum Pall.
|
II
|
HC
|
Tp
|
MX
|
euras.
|
15.
|
Crepis chrysantha (Ledeb.) Turcz.
|
I
|
HC
|
Srp
|
P
|
euras.
|
16.
|
Saussurea latifolia Ledeb.
|
II
|
HC
|
Lrp
|
М
|
as.
|
17.
|
Achillea ledebourii Heimerl
|
IV
|
HC
|
Brp
|
М
|
as.
|
18.
|
Frolovia frolowii (Ledeb.) Raab-Straube
|
II
|
HC
|
Tp
|
MP
|
as.
|
19.
|
Leuzea carthamoides (Willd.) DC.
|
II
|
HC
|
Tp
|
MP
|
as.
|
20.
|
Doronicum altaicum Pall.
|
IV
|
HC
|
Tp
|
P
|
as.
|
Berberidaceae
|
21.
|
Berberis sibirica Pall.
|
I
|
NF
|
S
|
MPt
|
cent.as
|
Betulaceae
|
22.
|
Betula glandulosa Michx. (=B. rotundifolia Spach)
|
III
|
NF
|
S
|
P
|
euras.
|
Boraginaceae
|
23.
|
Myosotis scorpioides L.
|
II
|
HC
|
Brp
|
H
|
holarc.
|
24.
|
Myosotis sylvatica Ehrh. ex Hoffm.
|
I
|
HC
|
Brp
|
М
|
euras.
|
25.
|
Eritrichium villosum (Ledeb.) Bunge
|
I
|
HC
|
Tp
|
P
|
euras.
|
Brassicaceae
|
26.
|
Cardamine macrophylla Willd.
|
II
|
C
|
Lrp
|
HP
|
as.
|
27.
|
Macropodium nivale (Pall.) W.T.Aiton
|
IV
|
HC
|
Tp
|
P
|
as.
|
Campanulaceae
|
28.
|
Campanula cervicaria L.
|
I
|
HC
|
Tp
|
М
|
euras.
|
Caprifoliaceae
|
29.
|
Patrinia sibirica (L.) Juss.
|
II
|
HC
|
Tp
|
XPt
|
as.
|
30.
|
Valeriana dubia Bunge
|
I
|
C
|
Brp
|
MX
|
as.
|
31.
|
Lonicera caerulea subsp. altaica (Pall.) Gladkova
|
II
|
MF
|
S
|
MP
|
as.
|
Caryophyllaceae
|
32.
|
Dichodon cerastoides (L.) Rchb.
|
I
|
H
|
Srp
|
P
|
holarc.
|
33.
|
Cherleria biflora (L.) A.J.Moore & Dillenb.
|
I
|
H
|
Tp
|
P
|
holarc.
|
34.
|
Sabulina verna (L.) Rchb.
|
II
|
HC
|
Tp
|
P
|
holarc.
|
35.
|
Sagina saginoides (L.) H.Karst.
|
II
|
H
|
Tp
|
P
|
holarc.
|
36.
|
Dianthus superbus L.
|
II
|
HC
|
Lrp
|
М
|
euras.
|
37.
|
Cerastium davuricum Fisch. ex Spreng.
|
III
|
HC
|
Tp
|
HP
|
as.
|
38.
|
Silene bungei Bocquet
|
I
|
HC
|
Tp
|
P
|
as.
|
Crassulaceae
|
39.
|
Hylotelephium telephium (L.) H.Ohba
|
IV
|
HC
|
Brp
|
М
|
euras.
|
40.
|
Phedimus hybridus (L.) Hart
|
III
|
H
|
Srp
|
XPt
|
as.
|
41.
|
Hylotelephium ewersii (Ledeb.) H.Ohba
|
II
|
H
|
Brp
|
MPt
|
as.
|
42.
|
Rhodiola algida (Ledeb.) Fisch. & C.A.Mey.
|
IV
|
H
|
Tp
|
P
|
Alt.
|
Cupressaceae
|
43.
|
Juniperus communis var. saxatilis Pall. (=J. sibirica Burgsd.)
|
II
|
NF
|
S
|
MP
|
euras.
|
Cyperaceae
|
44.
|
Eriophorum angustifolium Honck.
|
II
|
HC
|
Lrp
|
HP
|
holarc.
|
45.
|
Carex capillaris L.
|
V
|
HC
|
HS
|
H
|
euras.
|
46.
|
Carex aterrima Hoppe
|
IV
|
HC
|
Brp
|
HP
|
euras.
|
47.
|
Carex pediformis var. macroura (Meinsh.) Kük.
|
II
|
HC
|
Srp
|
MX
|
euras.
|
48.
|
Carex curaica Kunth
|
I
|
HC
|
Brp
|
H
|
as.
|
49.
|
Carex stenocarpa Turcz. ex V.I.Krecz.
|
IV
|
HC
|
Srp
|
P
|
as.
|
50.
|
Carex altaica (Gorodkov) V.I.Krecz.
|
I
|
C
|
Lrp
|
HP
|
Alt.
|
Ericaceae
|
51.
|
Vaccinium myrtillus L.
|
II
|
H
|
Ds
|
MP
|
holarc.
|
Euphorbiaceae
|
52.
|
Euphorbia pilosa L.
|
IV
|
HC
|
Tp
|
XPt
|
tur.
|
Fabaceae
|
53.
|
Hedysarum neglectum Ledeb. (=Hedysarum austrosibiricum B.Fedtsch.)
|
III
|
HC
|
Tp
|
М
|
euras.
|
54.
|
Oxytropis purpurea (Bald.) Markgr.
|
II
|
HC
|
Tp
|
М
|
euras.
|
55.
|
Trifolium lupinaster L.
|
IV
|
HC
|
Srp
|
MX
|
euras.
|
56.
|
Hedysarum theinum Krasnob.
|
III
|
HC
|
Tp
|
MP
|
as.
|
57.
|
Thermopsis alpina (Pall.) Ledeb.
|
II
|
HC
|
Lrp
|
P
|
as.
|
58.
|
Oxytropis alpina Bunge
|
III
|
HC
|
Tp
|
P
|
Alt.
|
Gentianaceae
|
59.
|
Swertia obtusa Ledeb.
|
III
|
HC
|
Lrp
|
М
|
as.
|
60.
|
Gentiana algida Pall.
|
II
|
HC
|
Brp
|
P
|
as.
|
61.
|
Gentiana grandiflora Laxm.
|
I
|
H
|
Tp
|
P
|
as.
|
Geraniaceae
|
62.
|
Geranium albiflorum Ledeb.
|
II
|
HC
|
Brp
|
М
|
as.
|
Juncaceae
|
63.
|
Luzula spicata (L.) DC.
|
I
|
HC
|
Brp
|
P
|
holarc.
|
Lamiaceae
|
64.
|
Dracocephalum ruyschiana L.
|
II
|
C
|
Srp
|
М
|
euras.
|
65.
|
Dracocephalum peregrinum L.
|
I
|
HC
|
Tp
|
XPt
|
as.
|
66.
|
Dracocephalum grandiflorum L.
|
III
|
HC
|
Srp
|
MP
|
as.
|
Liliaceae
|
67.
|
Gagea serotina (L.) Ker Gawl. (=Lloydia serotina (L.) Rchb.)
|
II
|
C
|
Bp
|
P
|
holarc.
|
Lycopodiaceae
|
68.
|
Diphasiastrum alpinum (L.) Holub
|
II
|
H
|
Cm
|
P
|
holarc.
|
Melanthiaceae
|
69.
|
Veratrum lobelianum Bernh.
|
III
|
HC
|
Brp
|
М
|
as.
|
Montiaceae
|
70.
|
Claytonia joanneana Schult.
|
I
|
HC
|
Tp
|
P
|
as.
|
Onagraceae
|
71.
|
Epilobium palustre L.
|
II
|
HC
|
Srp
|
HP
|
holarc.
|
72.
|
Epilobium angustifolium L.
|
I
|
HC
|
Tp
|
М
|
holarc.
|
Orobanchaceae
|
73.
|
Pedicularis oederi Vahl
|
III
|
HC
|
Tp
|
P
|
holarc.
|
74.
|
Pedicularis violascens Schrenk
|
I
|
HC
|
Tp
|
XPt
|
as.
|
75.
|
Pedicularis amoena Adams ex Steven
|
II
|
HC
|
Tp
|
P
|
as.
|
76.
|
Pedicularis achilleifolia Stephan ex Willd.
|
I
|
HC
|
Tp
|
XPt
|
as.
|
Papaveraceae
|
77.
|
Papaver nudicaule L.
|
III
|
HC
|
Tp
|
MP
|
as.
|
78.
|
Papaver croceum Ledeb.
|
I
|
HC
|
Tp
|
P
|
as.
|
Pinaceae
|
79.
|
Abies sibirica Ledeb.
|
I
|
MF
|
T
|
М
|
euras.
|
80.
|
Larix sibirica Ledeb.
|
I
|
MF
|
T
|
М
|
euras.
|
81.
|
Picea obovata Ledeb.
|
I
|
MF
|
T
|
М
|
euras.
|
82.
|
Pinus sibirica Du Tour
|
I
|
MF
|
T
|
М
|
euras.
|
Plantaginaceae
|
83.
|
Veronica densiflora Ledeb.
|
I
|
HC
|
Lrp
|
P
|
as.
|
Poaceae
|
84.
|
Deschampsia cespitosa (L.) P.Beauv.
|
IV
|
HC
|
Brp
|
H
|
cosm.
|
85.
|
Anthoxanthum monticola (Bigelow) Veldkamp
|
I
|
HC
|
Srp
|
М
|
cosm.
|
86.
|
Festuca rubra L.
|
III
|
HC
|
Brp
|
HP
|
holarc.
|
87.
|
Poa alpigena Lindm.
|
II
|
C
|
Lrp
|
М
|
holarc.
|
88.
|
Festuca borissii Reverd.
|
IV
|
HC
|
Brp
|
MP
|
holarc.
|
89.
|
Calamagrostis purpurea (Trin.) Trin.
|
II
|
HC
|
Lrp
|
H
|
euras.
|
90.
|
Elymus repens (L.) Gould
|
II
|
C
|
Lrp
|
М
|
euras.
|
91.
|
Alopecurus pratensis L.
|
II
|
C
|
Srp
|
М
|
euras.
|
92.
|
Dactylis glomerata L.
|
III
|
HC
|
Brp
|
М
|
euras.
|
93.
|
Helictochloa hookeri (Scribn.) Romero Zarco
|
II
|
HC
|
Brp
|
MX
|
euras.
|
94.
|
Phleum alpinum L.
|
II
|
HC
|
Brp
|
P
|
euras.
|
95.
|
Poa sibirica Roshev.
|
I
|
HC
|
Brp
|
MP
|
as.
|
96.
|
Paracolpodium altaicum (Trin.) Tzvelev
|
I
|
HC
|
Lrp
|
P
|
as.
|
97.
|
Trisetum altaicum Roshev.
|
II
|
HC
|
Brp
|
P
|
as.
|
98.
|
Festuca kryloviana Reverd.
|
IV
|
HC
|
Brp
|
P
|
as.
|
99.
|
Poa attenuata Trin.
|
III
|
HC
|
Brp
|
MX
|
Alt.
|
100.
|
Koeleria altaica (Domin) Krylov
|
I
|
HC
|
Brp
|
P
|
Alt.
|
Polygonaceae
|
101.
|
Bistorta vivipara (L.) Delarbre
|
III
|
HC
|
Brp
|
HP
|
holarc.
|
102.
|
Oxyria digyna (L.) Hill
|
II
|
HC
|
Lrp
|
MPt
|
holarc.
|
103.
|
Koenigia alpina (All.) T.M.Schust. & Reveal
|
II
|
HC
|
Tp
|
М
|
euras.
|
104.
|
Rumex acetosa L.
|
III
|
HC
|
Tp
|
М
|
euras.
|
105.
|
Rumex scutatus L.
|
II
|
HC
|
Tp
|
М
|
euras.
|
106.
|
Bistorta elliptica (Willd. ex Spreng.) V.V.Petrovsky, D.F.Murray & Elven
|
III
|
HC
|
Brp
|
P
|
euras.
|
Primulaceae
|
107.
|
Primula nivalis Pall.
|
I
|
HC
|
Brp
|
HP
|
as.
|
Ranunculaceae
|
108.
|
Ranunculus lapponicus L.
|
II
|
HC
|
Lrp
|
H
|
holarc.
|
109.
|
Caltha palustris L.
|
III
|
HC
|
Brp
|
H
|
holarc.
|
110.
|
Thalictrum alpinum L.
|
I
|
HC
|
Srp
|
P
|
holarc.
|
111.
|
Delphinium elatum L.
|
IV
|
HC
|
Srp
|
М
|
euras.
|
112.
|
Thalictrum flavum L.
|
II
|
HC
|
Brp
|
М
|
euras.
|
113.
|
Aconitum septentrionale Koelle
|
III
|
HC
|
Tp
|
М
|
euras.
|
114.
|
Trollius altaicus C.A.Mey.
|
II
|
HC
|
Srp
|
М
|
as.
|
115.
|
Anemonastrum narcissiflorum (L.) Holub
|
II
|
HC
|
Srp
|
MX
|
as.
|
116.
|
Aquilegia flabellata Siebold & Zucc.
|
IV
|
HC
|
Srp
|
MX
|
as.
|
117.
|
Callianthemum alatavicum Freyn
|
II
|
HC
|
Srp
|
P
|
as.
|
118.
|
Trollius lilacinus Bunge
|
I
|
HC
|
Brp
|
P
|
as.
|
119.
|
Aconitum apetalum (Huth) B.Fedtsch.
|
III
|
HC
|
Brp
|
MP
|
Alt.
|
120.
|
Aconitum glandulosum Rapaics
|
II
|
HC
|
Bbp
|
MP
|
Alt.
|
121.
|
Ranunculus altaicus Laxm.
|
IV
|
HC
|
Brp
|
P
|
Alt.
|
Rosaceae
|
122.
|
Dasiphora fruticosa (L.) Rydb.
|
II
|
NF
|
S
|
М
|
holarc.
|
123.
|
Spiraea media Schmidt
|
II
|
NF
|
S
|
М
|
euras.
|
124.
|
Alchemilla altaica Juz.
|
IV
|
HC
|
Srp
|
М
|
euras.
|
125.
|
Cotoneaster uniflorus Bunge
|
II
|
NF
|
S
|
P
|
euras.
|
126.
|
Sibbaldia procumbens L.
|
III
|
H
|
Lrp
|
P
|
as.
|
127.
|
Dasiphora glabrata (Willd. ex Schltdl.) Soják
|
II
|
NF
|
S
|
P
|
as.
|
128.
|
Dryas oxyodonta Juz.
|
III
|
H
|
Ds
|
P
|
as.
|
129.
|
Sanguisorba alpina Bunge
|
III
|
HC
|
Tp
|
P
|
as.
|
130.
|
Sibiraea laevigata (L.) Maxim.
|
II
|
NF
|
S
|
MX
|
Alt.
|
Rubiaceae
|
131.
|
Galium boreale L.
|
II
|
HC
|
Lrp
|
М
|
holarc.
|
Salicaceae
|
132.
|
Salix lanata L.
|
II
|
NF
|
S
|
P
|
holarc.
|
133.
|
Salix turczaninowii (Laksch.)
|
II
|
NF
|
Ds
|
P
|
euras.
|
134.
|
Salix rectijulis Ledeb. ex Trautv.
|
II
|
NF
|
S
|
P
|
as.
|
Saxifragaceae
|
135.
|
Saxifraga sibirica L.
|
I
|
HC
|
Brp
|
HP
|
as.
|
136.
|
Micranthes punctata (L.) Losinsk.
|
II
|
HC
|
Tp
|
HP
|
as.
|
137.
|
Bergenia crassifolia (L.) Fritsch
|
II
|
HC
|
Lrp
|
MPt
|
as.
|
Urticaceae
|
138.
|
Urtica dioica L.
|
I
|
HC
|
Lrp
|
М
|
euras.
|
Violaceae
|
139.
|
Viola biflora L.
|
II
|
HC
|
Srp
|
MP
|
holarc.
|
140.
|
Viola altaica Ker Gawl.
|
III
|
HC
|
Tp
|
P
|
as.
|
*1 – Occurrence of species: I – 0–20%, II – 21–40%, III – 41–60%, IV – 61–80%, V – 81–100%.
2 – The life forms are given (Raunkiaer): Meso phanerophytes – MF, Nano phanerophytes – NF, Hamefites-H, Hemicryptophytes- HC, Cryptophytes – C.
3 – The life forms are given (Serebryakov): T – tree; S – shrub; Hs – half-shrub; Ds – dwarfshrub; Lrp – long rhizomatous plant; Srp – short rhizomatous plants; Bbp – bulbotuberiferous plants; Bp – bulbous plants; Tp –taproot plants; Brp – brushy root plants; Tsp – tussock plants; Cm – club-moss.
4 – Ecological groups of plants in relation to the temperature, moisture and stonyness of the substrate: H - hygrophytes, HP - hygropsychrophytes, GM - hygromesophytes, M - mesophytes, MX - mesoxerophytes, MP - mesopsychrophytes, X - xerophytes, XPt - xeropetrophyte, P - psychrophytes, MPt - mesopetrophytes.
5 – The groups of areas are listed: cosm. - cosmopolitan; holarc. - Holarctic; euras. - Eurasian; as. - Asian; tur. - Turanian; Mtr - Mediterranean; Alt - Altai (endemics of the Altai-Sayan botanical-geographical province).