Altai Mountain Country is situated in the borderland between two subcontinents – Northern and Central Asia and stretches from 44°30? to approximately 53–54° bor. lat. and from 80°30? – 82° – 92°30? – 95° – 98° orient. long. It’s a square of about 550 000 km2 bound within the territories of Russia, Kazakhstan, China, and Mongolia.
The huge, complicated mountain elevation of Altai and Altaids (so-called mountains of Bordering Dzungaria), on one side, presents the south-western outpost of great mountainous systems of Southern Siberia and Northern Mongolia, but, on the other side, this mountain country is closely connected in Dzungaria as well as in the South-West Mongolia with even more colossal mountainous systems of Mountainous (High) Asia, in particular, with Tian-Shan.
The Altai Mountain Country is the highest elevation of all the mountainous systems of Northern Asia. The highest peaks are the two-headed Belukha (4506 and 4400 m) in the Katunsky range and the mountain knot Tavan-Bogdo-Ula (5 divine mountains with highest peaks of 4374, 4360 and 3981 m). Six more ranges and mountain massives have peaks with a height of more than 4000 m (Munkh-Khirekhan, Tsast-Ula, Northern-Chuysky Alps, Kharkhira, Sutai-Ula, Turgen-Ula), and other 10 ranges and mountain massives of Altai and Saur have peaks exceeding 3700–3900 m. A remarkable peculiarity of AMC is the multistage construction of the whole elevation with three basic levels of planation as well as a presence in the central cupola-shaped elevation of the extensive high-mountain plateau at the altitude of 2200–2400 m, and also presence of two-step system of the flat intermountain valleys at the altitude of 1400 and 1800–1900 m, and, finally, a combination of sublatitudinal and submeridianal (up to meridianal) direction of ranges. This peculiarity is connected to the fact, that the massives of ancient peneplene still present at the different altitudes and in different regions of AMC, which were exposed to multiple block elevation, and only at the end of the tertiary and during the quaternary periods were renewed by recent tectonic activity and by ancient congelation, stronger than the last one. This is why the uppermost sites of numerous ranges of Altai Mountain Country are presented by the flat watersheds with separate peaks or massives both recently appeared and rather ancient. This is especially characteristic to the Russian Altai, the southern part of Mongolian Altai, upper parts of Tarbagatai and Adzh-Bogdo. Dominance of massive block elevations in the tectonic structure of Altai is also emphasized by the presence at the north-west periphery of Altai and also in Zaissan depression and, partly, in Barun-Khurai depression, of the front terrace at the altitude of neighbouring plains – to an average of 300–340 m in the north and west of the whole country, and about 450–500 m in Zaissan and Barun-Khurai depressions. In the eastern part of AMC the pedestal of the front foothills and, sometimes, of rather considerably and even sharply elevated high mountain massives are higher, especially in the direction from the north to the south. This is quite evident, because it is formed by the levels of intermountain valleys – Minusinsk depression (lowest altitude 240 m), Tuva depression (640 m), Ubsunur depression (743 m), Depression of the Great Lakes (1028 m), Shargyn-Gobi (948 m), Transaltai Gobi (1000 m).
Altai Mountain Country is also an important watershed. The major part of the upper (mountainous) basin of Ob and Irtysh, the minor part of the basin of Yenisey (big left branches – Abakan and Khemchik), and basins of rivers flowing into the closed depressions of Kazakhstan, Dzungaria and Mongolia, are situated here. The latter are big depressions of Balkhash (contemporary flowing of Ayaguz hardly reaches the lake), Alakol-Sassykol, northern part of Dzungarian Gobi (Urungu-Noor lake, which just recently has been isolated from the basin of Irtysh), a part of the Depression of the Great Lakes (Khara-Us-Nuur), and also borders of Barun-Khurai depression, closed depressions with lakes Achit-Nuur, Ereg-Nuur and Tsagan-Nuur in the north and Alag-Nuur in the south. The biggest lake of the region – Uvs-Nuur – gets a minor part of it’s basin from Altai mts.
Altai Mountain Country is covered with a dense web of rivers (somewhere these rivers flow through the big ranges). However, contemporary congelation of Altai is rather low. The main centre of congelation in Altai is a mountain knot Tavan-Bogdo-Ula, where the biggest valley glaciers are situated (chiefly in Mongolia). Other large massives of congelation are the glaciers of Belukha, Northern-Chuysky Alps, Southern-Chuysky Alps, Mest-Ula, to a less degree – glaciers of Turgen-Ula and Mongun-Taiga. The ice cupolas of Tsast-Ula, Sutai-Ula and Tsengel-Ula are apparently remnants of the past, much stronger, congelation . Congelation of high mountain massive Munkh-Khirekhan is very peculiar: small glaciers in narrow valleys exist as a result of gradual degradation of pendent glaciers, not forming a continuous ice cover of the tops. Some other ranges also have small glaciers and firn fields. In general, however, glaciers form only an insufficient part of drain of the biggest rivers, flowing from Altai mts. The annual melting of the rather thick snow cover plays a greater role, especially in northern and western parts of Altai, as well as in the knots of interception of wet air masses (orientation of which is mostly south-western). In this case annual precipitations (with a considerable role of hard precipitations during the cold season) amount up to 1500–1600 mm (and, sometimes, more) per year. Durability of snow cover (and snowdrifts, not melting during the warm season) is, evidently, highest at the northern slopes. But, despite of so high humidity of ranges, intercepting wet air masses, there are a lot of regions with extremly low amounts of precipitations in Altai, such as sites of extensive intermountain depressions both low and high (Zaissan depresion – 120–200 mm, Chuyskaya “steppe” and Achin-Nuur depression – up to 200 mm with some localities of 110–150 mm). But the high mountain plateau Ukok in the central part also gets only about 280–300 mm per year.
The climate of Altai Mountain Country is sharply continental, and, because of absolutely open exposure to migrations of arctic air masses, is very severe. However, highly complicated relief considerably changes climatic conditions in different parts of the region. In general, the local climate is determined by the position of the locality regarding the altitudinal zonality, which differs in various parts of Altai. In winter, owing to the inversion, a great deal of localities of the middle montane belt are warmer than sites on the neighbouring plains; at the same time, localities, situated in the high montane belt (especially in landscapes in which cool air masses can stay for a long-time), can have very low temperatures, both winter and summer. The average January temperature in Gorno-Altaisk (377 m above sea level) is about -13°C, whereas in Bijsk (205 m) – less than -16°C, in Katon-Karagai (1060 m) on the Bukhtarma – -14.8°C, but in Zyryanovsk (449 m) – -22.2°C. Average January temperatures in Chuyskaya “steppe” (1750 m), and in adjacent sites of Mongolian Altai (Tsagan-Gol) are -31 – -32°C, and an absolute minimum in Kosh-Agach is -60,2°C. At the same time, the average January temperature, registered by the high mountain station Akkem in vicinities of the Belukha, is -17°C. Summer temperatures in AMC, in comparison to neighbouring plains, are lower even in the low montane belt, and with the increase in the altitude they decrease considerably. However, amount of vegetational temperatures still reaches 1800–2000°C up to 800 m a. s. l. in northern regions, and up to 1000–1200 m – in southern, but then decreases strongly upwards. It’s quite obvious, that conditions both for growth of wild plants and agriculture are very different in different parts of AMC; they are most favorable in northern and north-western foothills (up to the middle montane belt), well provided with precipitations and heat, where the frost-free period lasts about 120–140 days and more, and summer drought is almost absent. Many regions of Altai get considerable amoumt of the sun radiation, and often in this case the main limiting factors are either early summer frosts, or insufficient (withou irrigation) amount of precipitations. On the other hand, some peculiarities of Altai climate are favorable for animal industries. In forest regions it is possible because of the high productivity of hay meadows; in southern regions, having almost no forest, it is possible due to the small amount of snow in winter, which provides the opportunity of winter grazing without distant migrations. Long-term (during many ages) animal industries have been strongly influenced and changed the nature of many regions of Altai.
Soil cover of Altai Mountain Country is extremelly diverse, and in many regions has still not been described and classified completely. Complexity of the soil structure depends on zonal factors of relief and vegetation (which, obviously, differ in defferent parts of this vast territory), but it especially increases for the reason of differences in types of altitudinal zonality in different parts of AMC. Complicated combinations of light chestnut and desert (brown carbonate and typical brown) soils are developed in southern regions of Altai (mostly in Zaissan and Barun-Khurai depressions). Almost the same combinations, but without typical brown soils, are peculiar to some highly located depressions – they are common in several parts of Chuyskaya “steppe”, in Tsagan-Gol and Achit-Nuur depressions, in the lowest part of the depression between Darvijn-Nuruu and Alag-Khirekhan ridges. These soils are often salty enough, and usually they are combined with typical solonchaks. In highly located depressions with snowless winters these soils are formed on the permafrost base, which provokes uptake of salts onto the surface and formation of carbonate and gypsum crusts. Different variants of chestnut soils – from light and typical to dark on higher altitudes – are well developed in the southern, steppe regions of AMC. Mountain variants of chestnut soils have still been poorly investigated and in some cases, probably, they have not been not separated from mountain brown carbonate soils under the steppe bushes. In the low montane belt of Northern Altai chernozem soiles are formed: typical chernozems – under steppes, mountain leached chernozems – under meadow-steppes and, partly, under park birch forests, sometimes typical solods are formed under small insular groves or at the tops of ravines. Specific fat meadow chernozems are usually formed in river valleys, sometimes at very low altitudes (about 450 m); as a rule, they are peculiar to the more flat ravine slopes. Specific variants of meadow-chernozem soils are developed also under high grass communities in the lowest part of subalpine step of forest regions of Altai. Somewhere in the lowest part of the forest belt in the northern and western regions of Altai mountain podzol chernozem-like soils under aspen and birch forests are formed. Specific mountain slightly podzol dark forest soils (in the past considered to be close to dark-brown forest soils) are mostly developed in forest regions of Altai with dominance of montane “chernevaya” taiga. Genuine podzol (dark, dark-grey, and grey forest) soils are formed under the montane taiga of Central Altai. The less podzoled variants are developed under the park Larix sibirica forests (and Pinus sibirica-Larix sibirica forests in the lowest part of subalpine step). Podzols under Larix sibirica forests in the Inner Altai are formed on the permafrost base. At the northern slopes in valleys and also in the upper part of the forest belt, forest soils are often represented by peat variants. High mountain soils of AMC are especially diverse and original, and investigated rather poorly and disproportionally. There are different types of mountain meadow-steppe, mountain subalpine meadow soils, formed without influence of permafrost (peat mountain-meadow, meadow-podzol and mountain soddy podzol meadow, meadow chernozem-like soils). There are also different variants of high mountain meadow, peat high mountain meadow, and cryophytic meadow-steppe soils, developed under the influence of permafrost. Mountain tundra soils (raw, gley and gley-like, peat soils) are formed in the northern part of Altai and also somewhere in the high mountain belt. It is obvious, that vertical rows of soil formation processes also strongly depend on the structure of the parent rock as well as on the mobility of the substratum. This diversity is emphasized by the complexity of relief and differences in climate of different parts of AMC. As a result, zonal (altitudinal) types of soils are represented here by numerous variants of soil texture, characteristics of chemical compound, water regime, etc. It is necessary to note, that the relationship between vegetation and soils, formed under it, is not so obvious (and observable) in these conditions. Moreover, there is a good deal of substrata with almost inobservable processes of soil formation, but they are also used by plants. There are taluses, placers, rock streams (“kurums”) both on the tops and slopes, rocks, brims of moraines, etc. In all these cases it is also possible to say, that processes of soil formation have still not been investigated.
Vegetational cover (basic characteristics)
Because of the exceptional diversity of environmental conditions, vegetational cover of Altai Mountain Country is also highly diverse in its’ different parts, and in some of them it is exceptionally diverse.
The principal features of distribution of vegetation are determined here, first of all, by latitudinal, zonal and sectoral position of the territory, and also by differences in the system of altitudinal zonality in different parts of AMC, which differ in their localization both in the system of zonality and within AMC.
The northern part of Altai is situated in the subzone of northern forest-steppe, but through the forests of Kuznetsky Alatau (its’ western macroslope) it is connected also with southern plain taiga of Tshulym-Ketsky taiga region. A comparatively small part of Western Altai is connected directly with southern plain forest-steppe of Ob-Irtysh watershed, whereas the main part of Western Altai has a connection with the steppe zone (under specific conditions of eastern part of Central Kazakhstan Highlands). Northern-Turan-Dzungarian desert types become a zonal element only at the far south of Altai (southern slopes of Tarbagatai, Chinese and Mongolian Altai). However, at the western border of Altai some important factors of the influence of this giant mountain system on adjacent plains are visible. First of all, it results in a decrease of subzonal limits of forest-steppe and zonal limits of steppe and northern deserts southward (in the area of influence of mountains and in complicated conditions of low mountain massives). This phenomenon is even more expressed in the east of AMC, where, under conditions of the “rain shadow” on the highly elevated plains, the Gobi-type deserts (in the south), then desert steppes (northward) and, finally, dry steppes and complexes “semideserts, dry steppes and genuine steppes” (in an insular conditions) are developed. This considerable shift of zonal limits nortward is, of course, noteworthy, but not unique: at the same latitudes the same shift northward is to be found in the Turgai Hollow and in the north-eastern vicinities of Central Kazakhstan Highlands.
The fact that Altai Mountain Country is situated on the border of two sectors of Northern Asia – West Siberian-Middle Asian-Himalayan and extreme continental Central Siberian-Central Asian-Indochinese – is also of a great importance. In general this border runs next to 90° orient. long., but in Altai it is more complicated: it starts in an almost latitudinal direction along the Sayan range and Tchulyshman plateau, then – south-westward amidst Kuraiskaya and Chuyskaya steppes to the Ukok plateau – then, passing Tavan-Bogdo-Ula, eastward, to the Tsengel-Ula and Munkh-Khirekhan, south-eastward to Alag-Khirekhan, then again south-westward between Baitag-Bogdo and Ikh-Khavtgijn-Nuruu, and then – southward and westward through the ranges of Eastern Tian-Shan to the 80° meridian. This broken line represents the western limit of most continental phenomena (in high mountain conditions) on the Asian continent.
Latitudinal and zonal borders also define difference between two basic systems of altitudinal zonality in AMC – with and without well developed forest belt (or belts), although the difference between these systems are not reduced to only this characteristic. The general composition of flora, vegetation, and formations of vegetation also display marked differences of these.
General characteristics of flora
Flora of vascular plants of Altai Mountain Country includes at the present time no less than 2700 species, and description of new species as a result of both critical revisions of complicated groups and the discovery of new taxa, are which sometimes of great interest, is still going on. Flora of Altai in general is rather complicated, formed on the base of at least three greatly differing floras – Boreal Euro-Siberian (Northern European-Ural-Siberian), Steppe (contact Boreal-Ancient Mediterranean, geographically – Kazakhstan-Altai-Dzungarian), and Ancient Mediterranean (the most complicated, including, according to leading complexes, Turanian, Mountain Asian, and Central Asian components).
Nevertheless, numerically Boreal species (forest, meadow, and steppe) prevail in the flora of Altai. There are widespread Holarctic and Eurasian species, but also a great number of Asian elements – Siberian-Mongolian, Kazakhstan-Siberian, Ural-Siberian and Siberian species, including representatives of such a genera as Limnas, Macropodium, Stevenia, Mertensia, Patrinia, etc. The spectrum of leading families also reflects mainly the Boreal-Asian character of this flora – C?mpositae, Gramineae, Fabaceae, Cyperaceae, Rosaceae, Cruciferae, etc., as well as a good number of species of Carex, Salix, and presence of all Northern-Eurasian species of Ericaceae (in Middle, and, moreover, in Central Asia number of Carex decreases considerably, Ericaceae either absent or represented by a single species). But, at the same time, numerous families and genera, typical for the Ancient Mediterranean, are also represented well enough. There are ancient xerophytic groups – Ephedraceae (number of Ephedra in AMC is the highest in Northern Asia), Chenopodiaceae, Zygophyllaceae, Frankeniaceae, Plumbaginaceae. There are also Ancient Mediterranean families Tetradiclidaceae, Cynomoriaceae, genera Calligonum, Reaumuria, Chondrilla, Middle-Central Asian types of C?mpositae – Cancrinia, Brachanthemum, Ajania, Kaschgaria, etc. But there is also the Eastern Mediterranean-Mountain Asian family Biebersteiniaceae, mostly Eastern Mediterranean genera Eremurus, Ixiolirion, Gymnospermium, Leontice, Cicer, Ferula, Middle Asian and Middle Asian-Himalayan Megacarpaea, Leiospora, Calophaca, Schrenkia, Lindelofia, Mountain Asian Paraquilegia, Sibiraea, Gueldenstaedtia, Allardia, Leibnitzia, Altai-Tian-Shan Alfredia, etc. Flora of AMC also includes a considerable number of Asian genera (predominantly steppe) – Rhammatophyllum, Ptilotrichum, Galitzkya, Amblynotus, Craniospermum, Rhinactinidia, and Asian-American genera (also connected in Eurasia chiefly with steppes) – Thermopsis, Brachyactis, in any measure – also nothern species of Stroganowia, etc. There are also rather numerous, mostly Asian, genera, which have representatives in European mountains, but absent both in the Urals and in the Caucasus (Callianthemum, Leontopodium, etc.).
Altai is also a territory where areas of distribution of more locally spread genera are joined – there are Southern Siberian-Northern Mongolian Sajanella, Hansenia, Lithosciadium, Ferulopsis (all – Umbelliferae), Altai-Dzungarian Microstigma (Cruciferae), Stenocoelium, Ledebouriella (Umbelliferae), Altai-Dzungarian-Tian-Shan Taphrospermum, subendemic genus Pachyneurum (Cruciferae).
It is certain, that different parts of Altai Mountain Country possess floras, which differ in their composition (richness and originality). It is important, that latitudinal differences in the composition of these floras are not so clear as differences, caused by the abovementioned sectoral (submeridianal) border. And these differences are not only a consequence of contemporary climatic and edaphic conditions, because they were developing historically, over a long space of time. The most bright they are in the low mountain and and montane (middle mountain) belts of two sectors of AMC, both of which have their own original relic floristic sets.
The western sector (which conditionally could be named “forest sector”) contains relic set of nemoral elements, connected with mountain (“chernevaya”) taiga (and with “chern”, i. e. linden forests). More precise selection allows to connect with “chernevaya” taiga and with “chern” itself the following species: Tilia sibirica (natural only at the northern periphery of Altai), Dryopteris filix-mas, D.dilatata, D. expansa, Polystichum braunii, Oreopteris limbosperma, Brachypodium sylvaticum, Elymus pendulinus s. l., Festuca altissima, F. gigantea, F. extremiorientalis, Poa remota, Carex sylvatica, C. arnellii, C. hancockiana, Gagea granulosa, Asarum europaeum, Pseudostellaria borodinii, Actaea spicata, Cimicifuga foetida, Anemone umbrosa, A. reflexa, Eranthis sibirica, Corydalus nobilis, Eutrema integrifolium, Cardamine impatiens, Chrysosplenium filipes, Ch. ovalifolium, Geranium robertianum, Viola mirabilis, Circaea lutetiana, Epilobium montanum, Osmorhiza aristata, Sanicula europaea, S. uralensis, Myosotis krylovii, Pulmonaria mollissima, Stachys sylvatica, Galium odoratum, G. paradoxum, G. triflorum, Campanula latifolia, C. trachelium, Codonopsis clematidea, and, probably, Parapyrum anemonoides, Carex otrubae, Limnas stelleri. (It is of great importance to note that Mitella nuda is still not found in Altai, and forest species of Mertensia are also absent here).
Besides this complex, connected with “chernevaya” taiga (which, apparently, was initially connected with Fagus forests, i. e. it was, following Yu. D. Kleopov, “fagetal”), a good number of species, initially connected with former distribution of Quercus forests, also occur in AMC. This “quercetal” set of species includes here at least such a species as: Malus sieversii, R?sa pimpinellifolia, R. platyacantha, R. potentillaeflora, Cotoneaster multiflora, Lonicera microphylla, L. tatarica, L. micrantha, Daphne altaica, Sibiraea laevigata, Clematis integrifolia, Aconitum anthoroideum, Delphinium dictyocarpum, Adonis villosa, Thalictrum petaloideum, Paeonia hybrida, Gymnospermium altaicum, Corydalis schanginii, Dentaria sibirica, Hesperis pseudonivea, Lychnis chalcedonica, Potentilla rupestris, Astragalus glycyphyllus, A. unilateralis, Euphorbia pilosa, Dictamnus angustifolius, Alcea froloviana, Stelleropsis altaica, Brunnera sibirica, Scrophularia altaica, Galium (Cruciata) krylovii, Leibnitzia anandria, Ligularia robusta, Alfredia cernua, Tulipa altaica, T. heteropetala, T. patens, Fritillaria verticillata, F. pallidiflora, Erythronium sibiricum, Allium tulipaefolium, etc. (more detailed in: Kamelin, 1998). “Quercetal” complex is more diverse in contemporary ecological and coenotic characteristics of species, their distribution within AMC and general distribution. In contemporary conditions these species are connected in Altai with bush, bush-steppe, or, more rarely – with communities of “chernevaya” taiga or with Pine forests. Species of this “quercetal” complex, as well as species of subnemoral forests of “chernevaya” taiga, are of Euro-Siberian (more western), and Siberian-Eastern Asian (less numerous in “quercetal” complex) distribution. But this complex contains more species, common with Middle Asia, and more original elements (Kazakhstan-Altai, Altai, Altai-Dzungarian). Some of these species survived only at the far south of AMC (in Tarbagatai).
Tarbagatai (and, to a less degree, South-Western Altai) is the only territory, where one more complex of a relic nature is present. It is a “pra-shibliak”, the Middle Asian complex, connected at the present time mostly with steppe bushes. Species constituting this complex, are: Cerasus tianschanica, Atraphaxis laetevirens, A. decipiens, Calophaca soongorica, Amygdalus ledebouriana, species of Astragalus (sect. Calycocystis), Inula macrophylla, Saussurea amoena, Ferula soongorica, Astragalus sieversianus, Alcea nudiflora, Eryngium macrocalyx, Scabiosa songarica, Hypericum scabrum, H. elongatum, Haplophyllum perforatum, H. latifolium, Stroganowia intermedia, Eremurus altaicus, Allium robustum, Ferula gracilis, Scabiosa austroaltaica, Rheum wittrockii, Solenanthus circinnatus and others.
The eastern sector (which conditionally could be named “forestless sector”) has a very bright, well distinguished set of relic Northern-Central Asian (Tuvinian-Mongolian) montane-semidesert species (including strongly separated endemics). They are: Chenopodium frutescens, Kochia krylovii, K. mela-noptera, Salsola abrotanoides, S. gemmascens subsp. subglabra, S. monoptera, Nanophyton grubovii, Gypsophila desertorum, Corydalis stricta, Potentilla astragalifolia, Astragalus polozhiae, A. monophyllus subsp. gubanovii, Oxytropis mongolica, Zygophyllum melongena, Z. pterocarpum, Euphorbia mongolica, E. potaninii, Limonium klementzii, Artemisia schischkinii, Cancrinia krasnoborovii, Asterothamnus heteropappoides, A. poliifolius, Saussurea ceterachifolia, S. klementzii, Frankenia bucharica subsp. tuvinica and others. An important feature of this complex is the far penetration of species northward (in Chuyskaya “steppe” and Tuva). Widespread Middle-Central Asian species Reaumuria songarica also penetrates considerably to the north.
One more relic Central Asian desert complex is the complex of Ancient Gobian desert elements, represented in the south and most south-eastern part of Altai Mountain Country. It includes: Ephedra przewalskii, Iljinia regelii, Gymnocarpos przewalskii, Zygophyllum xanthoxylon (Z. kaschgaricum), Z. potaninii, Kaschgaria komarovii, Micropeplis arachnoidea, Nitraria roborowskii, and tugai poplar Populus diversifolia. This complex is impoverished enough in ACM, and many of Ancient Gobian species have the northern limits of their distribution to the south of AMC.
At the same time, many common Turanian-Dzungarian and Turanian-Gobian desert species, which are not relic here, have spread to the southern part of AMC. There are: Ephedra glauca, Iris tenuifolia, Atraphaxis spinosa, Rheum nanum, Anabasis brevifolia, Turanian-Dzungarian A. aphylla, A. elatior, A. salsa, A. eriopoda, Atriplex cana, Haloxylon ammodendron, Kalidium schrenkianum, K. foliatum, K. cuspidatum, Sphaerophysa salsula, Cynomorium songaricum, Cistanche salsa, Asterothamnus centraliasiaticus, Artemisia mongolorum, Karelinia caspia, Saussurea salsa, S. davurica, S. laciniata, Scorzonera pusilla, S. pseudodivaricata, etc., and also some of widespread psammophytes: Salsola arbuscula, Calligonum gobicum, C. mongolicum, Alhagi kirghisorum subsp. sparsifolia, Heliotropium ellipticum, Cancrinia discoidea, Chondrilla leiosperma, montane-tugai species of Tamarix and others. Some species of this group are of more local distribution (Altai-Dzungarian, Dzungarian, Tian-Shan-Dzungarian): Anabasis pelliotii, Nanophyton mongolicum, Potentilla imbricata, ?hesneya grubovii, Zygophyllum neglectum, Ajania achilleoides, A. grubovii, Brachanthemum mongolicum, Jurinea mongolica, Saussurea grubovii, S. popovii, Scorzonera mongolica, S. grubovii. Some of them are relics, but with non-desert relationship, and their age is quite different.
The original set of psammophytic species is peculiar to the sound dunes alongside the rivers Irtysh and Urunghu, Zaissan and Barun-Khurai depressions: Eremurus inderiensis, Allium caespitosum, Calligonum aphyllum, C. rubicundum, C. crispum, C. dissectum, C. ebi-nuricum, C. pumilum, Agriophyllum squarrosum, Astragalus ammodytes, A. burtschumensis, A. candidissimus, A. steinbergianus, A. urunguensis, A. roseus, A. gebleri, A. kendyrliki, Oxytropis aciphylla, Hedysarum scoparium, Rubia rezniczenkoana and others. They are all of quite different origin, the most part of them are distinctly separated from related species, and sometimes they represent even sets of closely related races.
A considerable number of Turanian-Middle Asian species, which find here a northern limit of distribution, represent a part of an exceptionally rich and combined set of flora of dry and desert steppes of low mountain and montane belts (these steppes cover rather a wide range of altitudes in the “forestless” sector). The same set of species is peculiar to semidesert petrophytic communities of steppes and Dzungarian deserts. These species are: Lepidium lacerum, Megacarpaea megalocarpa, Astragalus scabrisetus, Eremostachys molucelloides, Lagochilus diacanthophyllus, Artemisia sublessingiana, Cousinia astracanica, Helianthemum soongoricum, Iris songarica, etc. There are also Central Asian desert-steppe species in a strict sense, spread westward to Tarbaratai (Allium polyrhizum, A. bidentatum, Zygophyllum rosowii, Amblynotus rupestris), and also Kazakhstan-Altai-Dzungarian, Dzungarian, Tian-Shan-Dzungarian, Altai-Dzungarian and Altai elements (Allium subtilissimum, Atraphaxis compacta, Astragalus borodinii, Ferula dissecta, F. ferulaeoides, Limonium chrysocomum subsp. semenovii, Lagochilus bungei, Echinops integrifolius, Jurinea chaetocarpa, Scorzonera curvata and others). Flora of coloured clay elevations in the Zaissan depression is enriched by bright endemic species or species at the northern limit of distribution (Sisymbrium subspinescens, Zygophyllum latifolium, Ferula potaninii, Limonium coralloides,L. rezniczenkoanum, Calligonum junceum, etc.). Limestone communities of low mountain and montane belts also contain endemics and species at the limit of their distribution (Arenaria potaninii, Asperula kryloviana, Pyrethrum kelleri, etc.). Some similar species are peculiar to certain regions of Central Altai (lower course of Chuya and Katun valley from the Chuya mouth to Ursul). These are Caragana altaica, Gueldenstaedtia monophylla, Arnebia guttata, Brachanthemum krylovii, B. baranovii and some others.
The high mountain flora of “forest” Altai (and Saur) also differs greatly from the flora of “forestless” Altai, Tarbagatai, and Baitak-Bogdo. It’s enough to point out that the first flora contains a set of elements of “subalpine coloured tall herbaceous vegetation”, which includes rather a good number of relics, whereas another flora (mostly) contains a relic complex of high mountain Asian shrubs, subshrubs and cushions. These complexes were characterized in detail in one of my earlier publications (Kamelin, 1998). Moreover, these differences are displayed even at the highest mountain step (in alpine-nival belt): the first region bears alpine tundra and “goltsy” floristic complexes, the second – cryopetrophytic and cryoxeropetrophytic complexes.
Thus, it is quite obvious, that the flora of Altai Mountain Country is a contact (joint) flora at the borders of highly ranged elements of floristic division.
Types of vegetation
Diversity of natural conditions and considerable richness of flora lead to development in the Altai Mountain Country of a highly diverse vegetation. The most detailed picture of diversity of the basic (highest) syntaxa of vegetation is provided by using the florogenetical (“genetical”, ecoligical-historical) method of subdivision and classification of types of vegetation (Sukachev, 1942, 1943; Ovczinnikov, 1947, 1948, etc.; Lavrenko, 1956, 1961). These approaches of subdivision of types of vegetation (florocoenotypes) were also developed by me earlier in some publications, dedicated to the problem of classification of the basic (highest) syntaxa of vegetation of Middle Asia and Mongolia, and vegetation of some regions of Altai was also partly taken into consideration (Kamelin, 1979, 1987, 1998, 2002). A more detailed classification of types of vegetation of AMC is presented in this paper.
The process of evolution of plants on the Earth runs at least on three levels of organization of the plant world – populational-specific, coenotic, and biotic levels. In this connection it is possible to discern three joint components – phylogenesis, phylocoenogenesis, and biotogenesis (florogenesis in the world of plants). These processes are joint, connected and parallel, they differ in both their quantitative characteristics and intensity. Coenotic objects of different structure are involved in the phylocoenogenesis. They are: communities of individuals of one species and of numerous species (in their interactions within areas of different square); communities (phytocoenosises) (in their interactions with each other within areas of different square and different periods of time); groups of communities, ranged coenotically, topologically, and genetically in their natural habitat. Complicated interactions of coenotic objects are also determined by: the process of development of habitat of plants and plant communities; the processes of phylogenesis (microevolution in populations and different types of speciation); compositions of biotas (and floras) within certain territories and periods of time. All the processes, constituting phylocoenogenesis, are rather slow, but huge bulks of organisms of different biology and ecology (individuals of plant, animal, fungi organisms, and microorganisms) are involved. These interactions of bulks of living organisms of different biology and ecology within spaces of exceptionally diverse, but limited in its resources inert nature, and living nature, during a long time create coenotic structures of life, observable for us, in which plants play a leading role both as main producers of organic substance and as a basic structural component of biocommunities. Structural characteristics of communities depend not only on a general set of plant species, but, mostly, on an ecobiomorphic composition of edificators (number of types of biomorps is much lower than the number of plant species). This is, firstly, why the number of basic coenophylums (lines of evolution of coenotic structures) is rather low.
Types of vegetation (florocoenotypes) used by the author for phylocoenogenetical classification present, first of all, modern cut (station) of these basic coenophylums, which differ from each other considerably in diverse vegetational cover. Presenting a certain degree of phylocoenogenesis (process of historical development of phytocoenosises within a certain territory, inhabited by flora of certain type), types of vegetation change the environment in a special way and keep their composition and structure of constituting communities during a long period of time. In the phylocoenogenetical classification types of vegetation are interpreted as follows: totality of modern formations of vegetation, formed within territories with certain type of flora, edificators of which, during a long time and under the influence of rather stable environmental conditions (conditions of both inert and living nature), obtained in the processes of joint evolution common adaptive properties (requirements of temperature and water regime, of climate and soils, main edaphic requirements, phytocoenotic compatibility and competitive abilities, structure of ecological niches, overlapping environmental resources, structure of biomorphs, layering of communities, etc.).
Formation of vegetation, in this case, can be indicated as totality of plant associations, in a dominant (basic) layer (or in several layers, equal in their influence on the structure of phytocoenosis) of which edificator is a single (monodominant formation) species or a group of ecologically close, often in communities mutually substituting species (polydominant formation). For within extensive terrirories and, especially as in the case of Altai Mountain Country, in conditions of the contact of different floras, types of vegetation (florocoenotypes) can differ sharply in predominant development within different ranges of thermal-solar zones and corresponding altitudinal belts, it is important to distinguish groups of joint florocoenotypes, interacting with each other more than with florocoenotypes from other groups, in accordance with general characteristics of climatic range of their development. Because the territory in question is situated at the border of Boreal and Ancient Mediterranian floristic subkingdoms of Holarctis, florocoenotypes, peculiar to these subkingdoms only, form naturally two totalities of types of vegetation of the highest rank – Boreal and Ancient Mediterranian vegetations.
Types of vegetation (florocoenotypes) of Altai Mountain Country
Boreal vegetation. (Florocoenotypes of cold-temperate and temperate floras of Boreal subkingdom, penetrating into Ancient Mediterranian subkingdom through the mountains only).
Group of cryohumid types of vegetation.
Taiga (Peucedrymion holarcticum, Acciculi-silvae). Zonal type of vegetation of cold-temperate Holarctis (in AMC – in mountains only), arisen polytopically and polychronically in different parts of Holarctis on the similar base, in Siberia – on the base of prataiga communities of different composition, mostly in the second part of Neogene. There are formations of microtherm kryomesophytic coniferous trees (usually with well developed on-soil cover of forest mosses, but sometimes also with insufficient role of mosses), representing in AMC a variant of Ural-Siberian fratria (group) of formations of vegetation (Sochava, 1945). Three types of taiga are distinguished in Altai.
Subtype of dark coniferous taiga (Acciculi-silvae sciophilae): mono- and polydominant formations of Pinus sibirica, P. sibirica – Picea obovata, P. sibirica – Larix sibirica, Picea obovata, Picea obovata with Abies sibirica, P. obovata – Larix sibirica, and Saurian form of P. obovata.
Subtype of lihgt coniferous taiga (Acciculi-silvae illuminatae): rare in Altai monodominant formations of Larix sibirica (with a well developed moss layer or with swamping).
Subtype of subalpine light forests and elfin woods (Acciculi-silvae collucatae et humistratae): subalpine light forests of Pinus sibirica, more rarely Pinus sibirica – Larix sibirica, elfin woods Pinus sibirica, Abies sibirica).
“White forests” (Leucodrymion holarcticum, Albi-silvae). Type of vegetation of cold-temperate and temperate Holarctis (in cold-temperate regions – in mountains only), arisen polytopically and polychronically in different parts of Holarctis simultaneously with the development of prataiga communities and Pine forests from the end of Neogene to the Pleistocene. It is represented by formations of microtherm, more rarely mesotherm deciduous hydromesophytic and mesophytic trees and shrubs, in high mountains – cryomesophytic low trees and shrubs; in Altai – of different fratrias. At the present time there are several separated original subtypes in AMC.
Subtype of hygromesophytic flood plain forests – “uremas” (Albi-silvae inundatae). Polydominant formations of Populus alba, P. nigra, Salix triandra, S. alba, S. viminalis, Betula pendula, Padus avium and others (of Ural-Siberian fratria), Populus laurifolia, Betula pendula s. l., B. microphylla, B. kellerana, Salix rhamnifolia, S. rorida, S. ledebouriana (of Altai-Dzungarian fratria). Most often – monodominant formations of Hippophae rhamnoides, Populus pilosa, Betula turunica (of different fratrias).
Subtype of “kolki white forests” (“small insular white groves”) (Albi-silvae silvistepposae). Predominantly monodominant formations of Betula pendula subsp. truncata, B. alba s. str., more rarely – Populus tremula (with participation of bushes), developed mostly in combinations with montane meadow-steppe, more rarely – with steppe meadows (predominantly – communities of Ural-Siberian fratria).
Subtype of “white taiga” (Albi-silvae austro-sibiricae). Monodominant formations, developed only at the northern periphery of AMC. Products of long-term successions of taiga, more rarely – subclimax communities of Betula pendula subsp. truncata, B. alba subsp. krylovii (all of Ural-Siberian fratria, but highly diverse).
Subtype of “aspen forests” (Albi-silvae tremulosae). Monodominant (more rarely – with Betula sp. sp.) communities of formations of Populus tremula, mostly in successional rows of “chernevaya” taiga (of Ural-Siberian fratria).
Subtype of “elfin woods” (Albi-silvae fruticosae). Poly- and monodominant communities of formations of Betula tortuosa, B. humilis, B. fusca, Salix krylovii, S. recurvigemmis, S. sajanensis, S. coesia, S. kochiana, more rarely – S. alatavica. Monodominant (more rarely – with Salix sp.) communities of formations of Betula rotundifolia. In conditions of Altai Mountain Country are chiefly spread in high mountains, more rarely – in the upper sites of river valleys (the most part – of Ural-Siberian fratria of formations).
Meadows (Mesopojon holarcticum, Prata). Type of vegetation of cold-temperate and temperate Holarctic floras, appeared polytopically and polychronically in different regions on the base of forest (especially of riverian forest) communities of the complex of floras of “Turgai type”, then – on the base of Boreal floras from Miocene to Holocene. It comprises communities of formations of microtherm mesophytic grasses, including diverse rootstock grasses, forming thick sod. As a rule, there are polydominant communities, edificators of which change one another in long-term cyclic changes. In conditions of Altai and, partly, southward, – mostly anthropogenic communities, developed instead of felled forests, including riverian forests, and only partly in flood plains – natural, indigenous. They are highly diverse, and sometimes it is not possible to distinguish different subtypes (indigenous flood plain (riverian) meadows, second post-forest meadows, meadows of forest-glades, bushy meadows, swamp meadows, saline meadows, etc.) Polydominant formations of Zerna inermis, Dactylis glomerata, Poa pratensis, Festuca pratensis, F. arundinacea, F. rubra, Alopecurus pratensis, Elytrigia repens and others, mostly mono- and oligodominant formations of Phalaroides arundinacea, Calamagrostis epigeios, C. pseudophragmites, Agrostis stolonifera – Poa pratensis, Alopecurus songoricus – Poa sibirica, etc., as a rule, with a rich set of herbs. Post-forest meadows and spacious forest-glades (“yelanis”) – communities of Festuca arundinacea, Elymus gmelinii, Trisetum sibiricum, Calamagrostis langsdorfii, or Brachypodium pinnatum, Phleum phleoides, Poa angustifolia, Carex macroura, Iris ruthenica, or Helictotrichon pubescens, Poa angustifolia, P. sibirica, Zerna inermis, Stipa sibirica (in all the cases with a rich set of herbs). Steppe meadows with Helictotrichon pubescens, H. asiaticum, Phleum phleoides, Koeleria cristata, Medicago falcata, Vicia unijuga, V. amoena, Carex macroura, etc. Saline (solonetz and solonchak) meadows with Hordeum brevisubulatum, Glaux maritima or Leymus paboanus, and a great deal of others. As a result of former phytomeliorations, (in different periods) and cultivation of forage crops even in indigenous meadows, some cultivated forms of Trifolium pratense, Onobrychis sativa, etc. are rather common and, sometimes, abundant.
Swamp meadows and swampy meadows in different regions contain many species of sedges, including Carex songorica, C. disticha, C. dichroa, C. diluta, C. polyphylla, C. muricata, C. orbicularis s. l., Deschampsia caespitosa, Calamagrostis obtusata, etc. These meadows are often overgrown with different species of willows, etc. Hydrophytic meadows with Phragmites australis, Filipendula ulmaria, etc.
Mesophilous mountain grass communities (Mesocoryphion oreoasiaticum, Montanoherbeta). Florocoenotype of temperate Holarctic floras, appeared on the base of forest floras of “Turgai type”during Neogene and Pleistocene in AMC in montane and high mountain belts, rarely – lowers, along the river valleys. Communities of formations of oligotherm and microtherm mesophilous perennial herbs (with insignificant role of Gramineae and Carex sp. sp.). Formations of Geranium collinum, G. albiflorum, G. saxatile, Alchemilla sp. sp., especially A. krylovii, A. cyrtopleura, A. sibirica, A. rubens, A. rubricaulis, A. schischkinii, A. bungei, A. circularis, etc., polydominant formations with Potentilla sp. sp., especially P. chrysantha, P. asiatica, Polygonum bistorta s. l. (P. nitens, P. alopecuroides), P. viviparum, and also Phlomoides alpina, Ph. oreophila etc. In many cases these formations are connected through intermediate variants with meadows and communities of tall herbs and, apparently, at the present time are stabilized by grazing. The most part of communities is connected with Ural-Siberian and Altai-Dzundarian fratrias.
Sazes, saze swamps (communities of habitats with high and permanent water table in arid regions) (Sasophorbion oreoasiaticum, Fontipaludes montanae). Florocoenotype of temperate and warm-temperate floras, developed on the South Boreal base in Pleistocene. In AMC – mostly in high mountain belt. It comprises communities of formations of microtherm, more rarely – mesotherm hygrophytic herbs (predominantly monocotyledonous), in combinations with mosses – Carex pseudofoetida, C. orbicularis, C. altaica, C. capitata, C. bigelowii subsp. ensifolia, Juncus triglumis, with Ranunculus pseudohirculus, Saxifraga hirculus, Potentilla gelida subsp. euroasiatica, Pedicularis rhinanthoides, in lower belts – with Ranunculus longicaulis, Allium schoenoprasum. Hygrophilous mosses: Tomenthypnum nitens, Calliergon giganteum, Cratoneurum filicinum, Mniobryum albicans, Brachythecium rivulare, Paludella squarrosa, etc. Species of Sphagnum, Drepanocladus are absent, which in normal for High Asian fratria.
Herbaceuos swamps (Coryphiophorbion holarcticum, Pratopaludes). Type of vegetation of cold-temperate, temperate, more rarely – mountainous warm-temperate Holarctic floras, appeared in the second half of Neogene and in Pleistocene. In AMC – mostly in high mountain belt (especially in combinations with dwarf shrub formations (“yerniks”), Cobresia alpine formations and, partly, with alpine tundras; all belong to the Ural-Siberian fratria, but with an influence of High Asian fratria). Communities of formations of microtherm and mesotherm hydrophytic perennial herbs (predominantly sedges) in combinations with green mosses – formations of Carex atrofusca, C. melanantha, C. coriophora, C. capillaris, C. orbicularis, C. caespitosa, C. dichroa, C. ensifolia (as a rule, polydominant) in combinations with Eriophorum angustifolium, E. humile, more rarely – with Deschampsia caespitosa s. l., Poa tibetica, Saxifraga hirculus and moss carpets, formed by Aulacomnium palustre, Dicranum sp. sp., Drepanocladus, Camptothecium nitens, Hylocomium proliferum, more rarely – with forest mosses Pleurozium schreberi, Dicranum undulatum, very rarely – with Climacium dendroides, and sometimes also with Betula humilis, B. rotundifolia – in high mountain belt. Presence of Vaccinium uliginosum, Ledum palustre is very seldom. At the highest altitudes these formations include Carex dichroa, C. caespitosa, C. gracilis, C. disticha, C. pallescens, C. orbicularis, Aulacomnium palustre, Helodium blandowii, Meesia triquetra and forest mosses in combination with swamp birch forests (“sogras”), and also with Potentilla fruticosa, Salix krylovii, Betula humilis, more rarely – with Potentilla fruticosa, Sibiraea laevigata. Species of Sphagnum are extremelly rare.
Peatbogs (Phorbium holarcticum, Paludes turfosa). Type of vegetation of cold-temperate and temperate Holarctic floras, appeared, probably, to the end of Paleogene, but maximally widespread in Pleistocene. It is an exceptionally rare phenomenon in Altai Mountain Country. Communities of formations of hygrophytic sphagnum mosses with insignificant participation of green mosses and cryophytic and psychro-hygrophytic shrubs, rarely – with long-rhyzomatous polycarpics and other obligatory helophytes (ivsectivorous rosette polycarpics, etc.); in Altai – mostly of Sphagnum warnsdorfii, S. fuscum, S. magellanicum, S. squarrosum, Drepanocladus sp. sp., with Carex pauciflora, Eriophorum angustifolium, and almost without indigenous helophytes, which are very rare in Altai, and, moreover – in Saur and Tarbagatai. These formations have developed either in valleys of lakes or on contact with taiga larch forests (and spruce forests), where they seem to be of rather young age, but somewhere in high mountain belt there are also relic (Upper Pleistocene) peat hills on the permafrost base with partly restored process of sphagnum peat formation.
Rose bay shrublets (Rhodothamnion eurasiaticum, Rhodoreta). Type of vegetation of cold-temperate and temperate mountain Eurasian floras, appeared, probably, in Neogene and widespread in Pleistocene. It is exceptionally rare phenomenon in Altai Mountain Country. Communities of formations of Rhododendron aureum (Rh. chrysanthum), separated rarely at the upper limit of distribution of polydominant taiga communities (Pinus sibirica forests) in the most North-East of Altai. Absence of commuities of Rh. adamsii and Rh. parvifolium in Altai is one of the most bright and important characteristics, separating the flora of Altai and Altaids from the flora of Sayan mountain system.
Alpine (or alpine-like) herbaceuos carpets (Kryonanocoryphion eurasiaticum, Alpotapeta). Type of vegetation of temperate mountain floras of Eurasia (analogous types are also present in Pacific sector of North America), arisen polytopically and polychronically from Neogene, but most widespread from Middle Pliocene to the Quarternary period. This type of vegetation is represented in AMC most completely in comparison with all other mountain systems of Northern Asia. These are communities of polydominant formations of psychrophilous and chionofilous mesophytic herbaceous polycarpics (predominantly dicotyls) – Viola altaica, Gentiana grandiflora, G. uniflora, Ranunculus altaicus, R. fraternus, Oxygraphis glacialis, Cerastium cerastoides, C. pusillum, Minuartia verna, M. arctica, Primula algida, Myosotis alpestris subsp. asiatica, Eritrichium villosum, Lagotis integrifolia, Veronica densiflora, Pedicularis amoena, P. violasceps, Erigeron altaicus, E. oreades, E. eriocalyx, with Astragalus alpinus, Oxytropis alpina, Potentilla nivea, Aquilegia glandulosa, Delphinium sauricum, Thalictrum alpinum, Saxifraga nivalis, Claytonia joanneana, Allium platyspathum, Lloydia serotina, Poa altaica, Paracolpodium altaicum, etc. Almost monodominant communities of Sibbaldia procumbens (S. macrophylla), oligodominant communities with leading role of Callianthemum sajanense, C. alatavicum and others.
Mountain (alpine) tundras (Crymion boreale-holarcticum, Tundrae). Florocoenotype (more precisely – several poorly separated, but quite different in the set of prevailing life forms florocoenotypes) of cold-temperate and cold (Arctic) flora of Holarctis, arisen on the combined base from the end of Neogene to the Pleistocene (in the North of Eurasia – to the Holocene). In Altai Mountain Country – in the high mountain belt (but sometimes at rather low altitudes – from 1700–1750 m). Communities of formations of cryoxerophytic, kryomesophytic and psychrohygrophytic undershrubs (often creeping, mat, prostrate, etc.) and numerous lichen species, mosses, more rarely – perennial herbs.
Main subtypes
Lichen and moss-lichen tundras (Lichenotundrae) – communities of polydominant formations of Cladonia sp. sp. (C. alpestris, C. rangiferina, C. grayi, C. gracilis, C. subulata, C. sylvatica, etc.), Stereocaulon alpinum, S. grande, Rhizocarpon distinctum, Dactylina arctica, etc., Alectoria sp. sp. (A. ochroleuca, A. nigricans, etc.), species of Parmelia (Arctoparmelia) with Nephroma arctica, Peltigera aphtosa and others, Cetraria cucullata, C. nivalis, C. laevigata, Collema ceraniscum, Ophioparma ventosa, etc., sometimes also with some moss species – Polytrichum juniperinum, Ceratodon purpureum, Polytrichastrum sexangulare, mostly “yernik”-forest Dicranum undulatum, Rhodobryum roseum, Campylium stellatum, Rhytidium rugosum, Hylocomium proliferum, etc. Some species of undershrub willows and a few species of herbs (Braya sp. sp., Papaver canescens, Minuartia ar?tica, species of Oxytropis, etc.) occur in these tundras.
Dryas tundras (Dryastundrae). The edificator species is a dwarf semi-creeping (“mat”) undershrub Dryas oxyodonta. These tundras are very close to lichen tundras and represent either stages of evolution of the latter or an initial communities, arisen as a result of colonisation of bare soils by Drias; only a few lichen species are able then to invase in this communities (mostly crustaceous lichens on the surfaces of big stones).
Lichen tundras can also be changed by low (undershrub) willow tundras (Nanosalix-tundrae) with main species: Salix berberifolia subsp. brayi and S. berberifolia subsp. berberifolia, S. rectijulis, S. vestita, or with prostrate and creeping (almost herbaceous) Salix turczaninovii, S. reticulata, S. nummularia. Number of moss and flowering plant species increases in these tundras (besides the above mentioned, there are: Cerastium pusillum, species of S?xifraga, Pedicularis, in the North-East of Altai – also Hansenia mongolica and others).
One more special subtype of this type of vegetation is moss (lichen-moss) tundras (Musci-tundrae), constituted by both tundra and forest (“yernik”) moss species (up to Hylocomnium splendens, Pleurozium schreberi), and swamp species: Aulacomnium palustre, Tomenthypnum nitens, Paludella squarrosa, and, more rarely, by some species of Sphagnum. Some special undershrubs – Empetrum nigrum s. l., Vaccinium myrtillus (high mountain forms in Western Altai), Arctous alpina, low willows, Spiraea alpina, Cotoneaster uniflora, and also herbs: Gentiana algida, Papaver canescens, Polygonum viviparum, P. alopecuroides, Saxifraga punctata, S. oppositifolia, Schultzia crinita, S. albiflora, Pachypleurum alpinum, Senecio resedifolius, Cardamine bellidifolia, etc. – are also connected with moss tundras.
A special variant of moss tundras is “medallion” tundras (in combination with Kobresia-Carex communities). They are developed in the territories with permafrostogenous spots (“medallions”) of bare soils, gleizated on the surface, almost without plants. Braya aenea usually occur in these habitats, at the edges of these spots some lichens also grow. Suppressed individuals of Betula rotundifolia (nearer drained small streams and sometimes at the edges of medallions) also occur.
This complicated set of tundras of Altai Mountain Country has still been investigated rather poorly, and dynamics as well as interactions of them is not yet clear.
Group of humid types of vegetation
“Chernevaya” taiga (Protopeycedrymion nemorale, Acciculi-Nemoreta). Florocoenotype of temperate Palearctic flora, arisen on the base of northern variants of “Turgai floras” at the border of Oligocene and Miocene (and up to the Pliocene) in southern regions of Northern Asia (from the Urals to Baikal). It has been greatly reduced during the Pleistocene and, at the present time, is kept only in refugiums, the greatest of which covers all the northern part of Altai Mountain Country (in low mountain and montane belts only, where the conditions are most favorable). Communities of formations of Abies sibirica, more rarely – Pinus sibirica-Abies sibirica and Abies sibirica-Pinus sibirica oligodominant communities with a considerable set of nemoral herbs (see above) and mosses (Thamnium neckeroides, Eurynchium angustirete, Fissidens taxifolius, Amblystegium varium, Cirriphyllum piliferum, etc.), whereas the role of typical taiga mosses is insignificant or they are just absent (Gudoshnikov, 1979). Typical rich communities of “chernevaya” taiga are developed in Altai in the biggest refugium of the “District of big chern” (Shishkin, 1937), in the Western Altai refugium (Kolyvan-Upper Aley-Tigirek), in some sites of Northern-Western Altai (basins of Anuj, Peschanaya, Belokurikha, montane belt of the Katun basin), in Kazakhstanian Altai, and, probably, in the northern part of Chinese Altai. This is the most characteristic ancient type of vegetation of Ural-Siberian fratria.
“Dark forests”, “chern” (Therodrymion nemorale, Nemore-Silvae). Type of vegetation of temperate and warm-temperate Eurasia, developed polytopically and polychronically on the base of floras of “Turgai type”, and in mountains of warm-temperate Eurasia – on the base of Neogene floras of “Middle Asian-Hymalayan” and “Turgai” types of polydominant broad-laeved forests. This type is represented in Altai Mountain Country only by small fragments of relic Tilia sibirica forests, most completely kept in Gornaya Shoria (Ural-Siberian fratria of formations) and by relic Malus groves at the southern slope of Tarbgatai (Mountainous Middle Asian fratria of formations), which have lost a typical set of herbaceous species.
Mesophilous deciduous shrubs (Mesothamnion palearcticum, Nemorefrutices). Type of vegetation of temperate and warm-temperate Palearctic (especially of mountainous territories), arisen polytopically and polychronically on the base of broad-laeved and mixed (coniferuos-broad-laeved) forests in Upper Neogene and Pleistocene, in some cases – in Holocene. Polydominant formations of Lonicera tatarica, L. micrantha, Rhamnus catarthica, Crataegus pseudosanguinea, Daphne altaica, Rosa pimpinellifolia, R. potentillaeflora, R. platyacantha, sometimes with Sambucus sibirica, Padus avium subsp. asiatica, Ribes atropurpureum, etc. Usually mono- or oligodominant formations of Lonicera microphylla, Cotoneaster megalocarpa, Sibiraea laevigata, Caragana arborescens. Polydominant (intermediate) formations of some of mesophilous deciduous shrubs in combinations with steppe shrubs – Amygdalus ledebouriana, Calophaca soongorica, Spiraea hypericifolia, Atraphaxis laetevirens. The most part of these communities obviously belong to Altai-Dzungarian fratria (but they all are connected in their origin with oak forests degradation in Neogene). Analysis of some characteristic species is given in one ot the previous papers (Kamelin, 1998).
Tall herbaceous vegetation (Megalocoryphion nemorale, Altiherbeta). Type of vegetation of warm-temperate mesophilous floras, arisen on the base of different floras of polydominant coniferuos-broad-laeved forests (warm-temperate and mountainuos subtropical) as a result of disintegration of lower layers of vegetation from the Upper Paleogene and, mostly, during the Neogene. In Pleistocene-Holocene in high mountain belts it has been turned into a relic condition, but some elements of this florocoenotype (predominantly through flood plain forests) enriched also Boreal florocoenotypes. Different subtypes of tall herbaceous vegetation are developed in AMC.
Subalpine tall herbaceous vegetation (Altiherbeta subalpinae). Polydominant formations of Leuzea (Fornicium) carthamoides, Saussurea frolovii, Ligularia macrophylla, L. altaica, Paraligusticum discolor, Heracleum dissectum, Angelica decurrens, Aegopodium kaschmiricum, Aconitum altaicum, A. leucostomum, Anemone crinita, A. schrenkiana, Hedysarum neglectum, H. theinum, Allium ledebourianum, A. platyspathum subsp. amblyophyllum, Sanguisorba alpina, Potentilla asiatica with Cicerbita azurea, Doronicum altaicum, D. oblongifolium, Achillea ledebourii, Solidago gebleri, Phlomoides alpina, Geranium affine, G. albiflorum, G. collinum, Macropodium nivale, Pleurospermum uralense, Gentiana fischeri, Corydalis pauciflora, Viola disjuncta, more rarely – Veratrum lobelianum, Erytronium sibiricum and others. Rather good number of these species are sharply separated and relic (mostly from Neogene floras), considerable part of them are connected with contemporary floras of Middle Asia and Hymalayas, another part – are ancient South Siberian types (Kamelin, 1998).
Forest tall herbaceous vegetation, or broad-leaved tall herbaceous vegetation (Altiherbeta sylvaticae). Polydominant formations of Angelica decurrens, A. sylvestris, Heracleum dissectum, Pleurospermum uralense, Anthriscus sylvestris, Cirsium helenioides, Saussurea latifolia, Ranunculus grandifolius, Thalictrum minus s. l., Cacalia hastata, Lilium martagon s. l., Potentilla chrysantha, Cardamine macrophylla, more rarely – Delphinium inconspicuum, D. elatum, Aconitum septenrionale, Crepis sibirica, Alfredia cernua, Trollius asiaticus, Veratrum lobelianum, Ligularia glauca, and also some grass spesies – Milium effusum, Dactylis glomerata, Poa sibirica, Calamagrostis obtusata, etc. (Subalpine tall herbaceous vegetation is almost free of grasses).
High mountain (alpine) meadow tall herbaceous vegetation (Altiherbeta montano-pratosae). Polydominant (with different combinations) formations of Aquilegia glandulosa, A. gubanovii, Trollius altaicus, Doronicum altaicum, Geranium albiflorum, Aconitum altaicum, A. ukokense, A. mirabile, Anemone crinita, Polygonum nitens, Sanguisorba alpina, Matricaria ambigua, Swertia obtusa, Polemonium pulchellum, and also Aconitum pallasii, Pyrethrum alatavicum, Swertia marginata, Phlom?ides oreophila, Hedysarum austrosibiricum, and grasses Festuca kryloviana, Trisetum altaicum, Antho?anthum odoratum subsp. alpinum, Alopecurus turczaninovii, A. songoricus.
Group of cryosemihumid types of vegetation.
Pine forests (Pitydrymion holarcticum, Pineta). Type of vegetation of temperate and mountainuos warm-temperate floras, arisen on the base of floras of “Turgai type” during Miocene and Pleistocene, comprising communities of formations of micro- and mesotherm xeromesophytic and heliophilous coniferous trees. In AMC it is developed chiefly in western regions and in the montane belt of northern periphery. Formations of Pinus sylvestris, more rarely – Larix sibirica. Communities of Pinus sibirica (with Larix sibirica) with on-soil cover of “yagel” (“reindeer moss” – species of Cladonia s. l. and Cetraria s. l.) are extremelly rare.
Pine forests of Altai Mountain Country are highly diverse: these are forest-steppe and taiga pine forests (according to the zonal-belt position); subclimax communities (variants of “montane pine forests”, especially rocky, and a part of forest-steppe pine forests – “herbaceous” pine forests); these are also communities of successional changes with Pinus sibirica, and, partly, with “chernevaya” taiga (“fern” pine forests of the Katun valley). In general these communities belong to Ural-Siberian fratria, but somewhere in the Inner Altai and in the north-east of AMC they can be viewed as an elements of Angarid fratria (larch forests with Rhododendron sp. or Fornicium carthamoides, and also boggy forests).
Juniper prostrate formations (Arceuthothamnion oreoasiaticum, Junipereta humistratae). Type of vegetation of mountainuos temperate and warm-temperate Eurasian floras, arisen polytopically and polychronically mostly as a derivative of pine forests, probably, from Middle Pliocene to Pliestocene. It is represented in AMC by two main phylums (subtypes).
There are communities of high mountain juniper prostrate formations (Junipereta altimontanae) with dominance of Juniperus sibirica or J. pseudosabina, or, sometimes, of both species, more rarely – with J. sabina (Mongolian Altai, partly Central Altai, Tarbagatai, Saur). There are also communuties of steppe juniper prostrate formations (Junipereta stepposae), formed only by Juniperus sabina (Western Altai, Tarbagatai, Saur, partly Mongolian Altai) or J. sabina with Lonicera microphylla, Cotoneaster roborowskii (the south of the Mongolian Altai, Baitak-Bogdo, Eastern Tarbagatai). The first subtype belongs, probably, to Ural-Siberian fratria, whereas the second – to Altai-Dzungarian one.
Relic kryoxerophilous shrubs (and subshrubs) of the Mountainous Asia (Cryoxerothamnion montano-asiaticum, Aridofrutices relicti). Type of vegetation, arisen on the base of montane forest floras of Mountainous Asia in Neogene as a result of intensive orogenesis. In AMC it is in a relic condition in semiarid and arid high mountain belt, more rarely – in montane (middle mountain) belt. Communities of mostly mesotherm kryoxerophilous shrubs, tall subshrubs, and mat semiunderground subshrubs. Mono- and polydominant formations of Potentilla salesoviana, Biebersteinia odora, Ribes graveolens, Lonicera hispida, Berberis sibirica, Grossularia acicularis, and also Artemisia rutifolia, Ribes heterotrichum, sometimes also in combination with Juniperus pseudosabina, J. sibirica, more rarely – J. sabina.
Meadow-steppes (Coryphiopojon uralo-sibiricum, Pratosteppae). A zonal type of vegetation of temperate Asia, developed on the Post-Turgai and Boreal-Angarid base begining from the middle of Pliocene, but mostly in Pleistocene in connection with the development of pine forests and “white insular forests” (groves). Communities of formations of microtherm xeromesophytic polycarpic herbs, especially of loose-bunch and short-rootstock grasses, bunch sedges and with a rich set of herbaceous dicotyls. This type is well developed in AMC from the low mountain belt up to high mountains, but only within the limits of forests; it is often an element of complexes of pine forests, groves, and expositional forest-steppes (in combinations with lihgt coniferous forests).
Polydominant formations of Helictotrichon schellianum, H. pubescens, H. desertorum, Koeleria cristata, K. altaica, Phleum phleoides, Poa angustifolia, P. urssulensis, Elymus gmelinii, Carex pediformis, C. humilis, Artemisia sericea, A. tanacetifolia, more rarely – Stipa pennata, Poa stepposa, with a richest set of herbs – Polygonum alpinum, Delphinium dictyocarpum, Aconitum anthoroideum, Potentilla tanacetifolia, P. longifolia, Filipendula vulgaris, Oxytropis pilosa, Onobrychis arenaria, Medicago falcata, Hypericum elegans, H. perforatum, Bupleurum multinerve, Peucedanum morissonii, Origanum vulgare, Dracocephalum ruyschiana, Phlomoides tuberosa, Galium verum, Achillea asiatica, Inula salicina, often also Fragaria viridis, Schizonepeta multifida, Iris ruthenica, etc.
The main part of these communities belong to Ural-Siberian fratria, but, partly, – also to Altai-Dzungarian one. Petrophilous meadow-steppes have a specific type of composition (and connections): the most common elements of these are Elytrigia gmelinii, Allium rubens, Vicia costata, Hedysarum gmelinii, Veronica pinnata, in Tarbagatai – Hyssopus ambiquus, Dracocephalum integrifolium, etc.
Kryoxerophilous meadow-steppes (Kryocoryphiopojon asiaticum, Frigidopratosteppae). Type of vegetation of montane floras of temperate Asia, developed on the South-Boreal base mostly in Neogene, widespread during Pleistocene (when it was aslo enriched by elements of other florocoenotypes), and, apparently, reduced in Holocene. It comprises communities of formations of kryoxerophilous polycarpic herbs, especially bunch grasses. There are polydominant formations of Festuca altaica, F. tristis, F. brachyphylla, Ptilagrostis mongolica, Helictotrichum asiaticum, Koeleria atroviolacea, Trisetum mongolicum with Saussurea schanginiana, Polygonum nitens, Papaver tianschanicum, Schultzia crinita, Leontopodium ochroleucum, Aster alpinus s. l., Myosotis alpestris, Gastrolychnis apetala, Swertia obtusa, Eritrichium villosum, Thermopsis alpina, and also with Allium bogdoicolum, A. amphibolum, Saussurea pseudoalpina, etc. As a rule, species of Kobresia, especially K. smirnovii, K. capilliformis, Carex stenocarpa are present in these communities; rather usually kryoxerophilous meadow-steppes can be met together with Kobresia and Carex communities over extensive squares in high mountain belt.
Kryoxerophilous grass carpets (Kobresia communities) (Nan?coryphion oreoasiaticum, Frigidotapeta). Type of vegetation of mountain floras of temperate Asia, arisen polytopically from the end of Neogene up to Pleistocene. At the present time this is the most important high mountain zonal florocoenotype of Mountainous (High) Asia. It includes communities of kryoxeromorphic polycarpic herbs, bunch species of the genus Kobresia, some Carex species, but in most cases – without or with an insignificant role of grasses. In Altai Mountain Country it is predominantely spread in the central, strongly elevated part, where the highest ranges are situated. Polydominant formations of Kobresia myosuroides s. l. (K. bellardii), K. capilliformis, K. simplicifolia subsp. subholarctica, K. smirnovii, Carex stenocarpa, C. rupestris with C. melanantha, C. ledebouriana, C. coriophora, Allium schoenoprasum s. l., A. karelinii, Saxifraga hirculus, S. flagellaris subsp. macrocalyx, S. oppositifolia, more rarely with few grasses – Poa alpina, Trisetum mongolicum, Anthoxanthum odoratum subsp. alpinum.
Two main variants of Kobresia communities – with low and high role of herbaceous dicotyls (the latter are developed on the soils with better drainage) – are always to some extent connected (and only role of each differ in different regions).
Kryoxerophilous cushion plant formations (Pycnocoryphion oreoasiaticum, Pulvineta). Type of vegetation of montane temperate and warm-temperate floras of highest Asian mountains, arisen on the mixed base of Turgai floras derivatives and Ancient Mediterranean mountainous florocoenotypes (mostly juniper stands) from Neogene up to Pleistocene, and then reduced (in the northern regions). In AMC it form combinations with Kobresia communities and kryophilous meadow-steppes, as well as with high mountain variants of steppes. Communities of formations of microtherm kryoxeromesophilous and kryoxerophilous low cushion subshrubs and also herbs (perennial and annual). Formations of Stellaria petraea, S. pulvinata, Arenaria formosa, A. mongholica, Sibbaldia (Dryadanthe) tetrandra, Oxytropis komei, O. saposhnikovii, O. tschujae, Artemisia dolosa, A. pycnorrhiza, etc., with Potentilla biflora, Krascheninnikovia compacta, Eritrichium sp., etc.
Group of semiarid types of vegetation
Steppe shrubs (Xerothamnion stepposum, Steppofrutices). Type of vegetation of temperate and mountain warm-temperate floras of (mostly) Asia, arisen on the contact of degraded forests of Turgai type with Ancient Mediterranean prashibliaks simultaneously with steppe formation (or, in some regions, earlier). This florocoenotype comprises communities of oligotherm or mesotherm deciduous heliophilous xeromesophilous shrubs. In Altai Mountain Country it is spread in western and southern parts, usually not in the high mountain belt;and in Tarbagatai it is developed most completely. Formations (usually polydominant) of Spiraea hypericifolia, Caragana ?ygmaea, C. altaica, C. leucophloea, C. bungei, C. frutex, C. camilli-schneideri, C. spinosa (the lattar species is the most common in flood plains), Calophaca soongorica, Amygdalus ledebouriana, Cerasus tianschanica, Atraphaxis laetevirens, and also steppe petrophytic species Spiraea trifoliata.
Tragacanth formations (Tragacanthion mediterraneum, Tragacantho-fruticeta). Type of vegetation of warm-temperate Mediterranean flora, represented in AMC by the most northern (autochtonous) variants, arisen polytopically in Neogene as a result of intensive orogenesis in arid conditions, but on the similar base (juniper stands, and, southward, mostly prashibliak communities). In AMC it was strongly reduced in Pleistocene, and is represented now by combinations with steppes and Dzungarian deserts, including high mountain communities. Formations of Oxytropis tragacanthoides, O. hystrix (rarely, only Tarbagatai), O. aciphylla, very rarely – O. polyphylla, i. e. spinose cushion shrubs and subshrubs (mesotherm and kryoxerophilous).
Steppes (Xeropojon eurasiaticum, Steppae). Zonal type of vegetation of temperate and warm-temperate Eurasian floras, developed politopically and polychronically on the mixed Ancient Mediterranean and Ancient Boreal (Turgai, Angarid pine forest, etc.) base from Neogene up to Pleistocene, and also transformed cryogenically in mountains. In AMC it is common and peculiar almost to all altitudinal belts (in exception with extremelly kryophytic high mountains). Communities of microtherm (rarely – mesotherm) xerophilous (predominantly – scleroxerophilous) herbs, first of all – bunch grasses, but also Artemisia sp. sp., kryoxerophilous sedges, Allium sp. sp., and abundant set of dicotyls, especially from the Fabaceae family. Steppes in AMC, occupying diverse habitats, represent very heterogenous type of vegetation, developed within different fratrias and including several subtypes.
Subtype of shrub steppes (Steppae fruticetae). Formations of Stipa rubens, S. capillata with Caragana frutex, Spiraea hypericifolia (more rarely also with S. crenata), Stipa rubens, S. capillata with Calophaca soongorica, more rarely – with Amygdalus ledebouriana and others (of Altai-Dzungarian fratria). Formations of Stipa capillata, Poa stepposa – Artemisia glabella with Caragana pygmaea, Stipa capillata, Agropyron cristatum – Ajania fruticulosa with Caragana pygmaea (of Altai-Dzungarian fratria). Formations of Stipa orientalis, S. krylovii, Agropyron cristatum, Cleistogenes squarrosa (+Artemisia sp. sp.) with Caragana pygmaea, Stipa krylovii, Cleistogenes squarrosa, Artemisia frigida with Caragana pygmaea, Stipa glareosa, Agropyron cristatum, Artemisia frigida with Caragana leucophloea, etc. (of Tuvinian-Mongolian fratria).
Subtype of true steppes (Steppae magnicaespitosae). Formations of Stipa rubens, S. kirghisorum, S. capillata, S. lessingiana, derivative communities of Festuca valesiaca with Koeleria cristata s. l., Phleum phleoides, Poa stepposa, rarely communities of Stipa sareptana, Festuca pseudovina, F. rupicola (with Artemisia austriaca, A. sublessingiana, A. frigida, and even with dominance of these species in disturbed habitats). The most of these communities belong to Altai-Dzungarian fratria.
Subtype of Siberian-Mongolian polydominant firm-bunch grass steppes (Steppae parvicaespitosae multidominantae). The main belt-forming subtype of steppe vegetation for most parts of the AMC; it is common and plays an important role in Angarid frafria of formations. The classic variant is four-grass steppes (Festuca lenensis, Stipa krylovii, Cleistogenes squarrosa, Koeleria cristata subsp. macrantha) which mostly developed in regions, attributed to the Tuvinian-Mongolian province (in the second altitudinal belt). In some cases the role of Poa botryoides, Artemisia frigida, Carex duriuscula, different Allium species – A. tenuissimum, A. bidentatum, etc. is especially high. Another important variant is dry steppes with a high role of Agropyron cristatum in polydominant communities, and Festuca valesiaca, F. borissii, F. kryloviana instead of F. lenensis with different significance of Artemisia frigida, A. santolinifolia, sometimes – A. gracilescens, in disturbed communities – also with Potentilla acaulis. In the east of the region these steppes are mostly spread at rather low altitudes, in Central Altai – within more wide altitudinal range, whereas in Tarbagatai – usually in the upper patr of the steppe belt. Communities of both these polydominant variants with diminished number of edificators (two-three grass steppes with quite different combinations of edificators) are usually to be met in different conditions. Some other grass species can also take part in these communities – Agropyron tarbagataicum, Elytrigia gmelinii, Psathyrostachys juncea, Elytrigia nevskii, etc.
The most original are the high mountain variants of this subtype, such as high mountain Festuca steppes (Festuca kurtschumica) with Koeleria cristata, Phleum phleoides, Carex aneurocarpa, Artemisia frigida, Aster alpinus, etc., with a sufficient role of Stenocoelium atamanthoides and S. trichocarpum (the latter is endemic to Tarbagatai). There are also communities with Festuca tschujensis (Koeleria cristata s. l., Poa attenuata, Agropyron cristatum, Oxytropis, sp., Astragalus dilutus, Rhinactinidia eremophila, etc.), with Festuca kryloviana (Koeleria cristata s. l., K. altaica, Poa altaica, P. attenuata), with Phlomoides oreophila, Dracocephalum origanoides, Hedysarum roseum and others, also with Festuca valesiaca, F. tschuensis (Poa attenuata, Koeleria cristata s. l.) – Artemisia borotalensis, etc. And, finally, there are diverse dry steppe communities with Stipa orientalis.
Subtypes of subshrub – firm-bunch grass desert-like and desert steppes (Steppae subdesertosae). Formations of Stipa glareosa, S. orientalis, more rarely – S. gobica, S. klementzii, much more rarely – S. breviflora, S. consanguinea with Artemisia argyrophylla, A. caespitosa, A. frigida s. l., A. schischkinii, A. mongolorum, Allium polyrhizum, Elytrigia nevskii and subshrubs: Ajania fruticulosa, Asterothamnus heteropappoides, Kochia prostrata, Krascheninnikovia ceratoides, K. compacta, Nanophyton mongolicum, Rhinactinidia eremophila, Dracocephalum fruticulosum, more rarely – Anabasis brevifolia, etc. The disturbed steppes with Artemisia pectinata, A. macrocephala, A. intricata, etc. should also be attributed to this subtype. The most part of these communities belong to Turan-Dzungarian fratria, the rest – to Altai-Dzungarian one.
Subtypes of sandy steppes (Steppae ammophilae). Extensive spaces of sandy steppes in the Altai Mountain Country are sharply separated from each other and have different sets of floristic elements. Most western sandy steppes are formed by communities of Stipa dasyphylla, Festuca beckeri, Koeleria glauca, Agropyron cristatum (with Artemisia marschalliana, A. tomentella, Euphorbia seguieriana, and also Caragana frutex and, sometimes, Juniperus sabina). These are typical post-pine forest sandy steppes.
Elymus giganteus, Agropyron fragile, Chondrilla juncea, Astragalus ammodytes, and also Astragalus candidissimus appear on a more mobile sands; such a species as Caragana hololeuca, Astragalus roseus, Poa bulbosa are peculair to sandy habitats with light salinity (and with an increased gypsum content).
Sandy steppes of southern part of Achit-Nuur depression with Agropyron cristatum, A. fragile, Koeleria cristata s. l., Carex supina s. l. with Caragana bungei, somewhere – communities with Oxytropis aciphylla, are also quite original. A part of sandy steppes developed at the periphery of the Depression of the Great Lakes also include communities with Artemisia globosa. Even to the south, in the inner depressions at the border of Mongolian Altai, little sites of sands with Artemisia xerophytica, and, in the eastern part of southern slope of Mongolian Altai, inversed to Barun-Khurai depression, – with A. sphaerocephala, are developed. In both cases the grass base of these steppes is constituted by Agropyron cristatum, A. fragile, Koeleria cristata s. l.
Stony (petrophytic) steppes are even more diverse in the Altai Mountain Country, in which they cover a great altitudinal range. Petrophytic variants (sometimes highly original) are formed to a certain degree within each subtype of steppes. Since they connected with each other even less than sandy steppe communities, it is quite impossible to treat them as a uniform subtype. One part of these communities represent derivatives of not the steppes themselves, but of the steppe bushes (petrophytic steppes with Spiraea trilobata, Cotoneaster sp. sp), another one – of combined semidesert type complexes (petrophytic steppes with Krascheninnikovia ceratoides, or, in Tarbagatai, – with Hulthemia persica (=H. berberifolia), or, at the southern periphery of the Mongolian Altai, – with Nanophyton mongolicum). The high mountain petrophytic variants of steppes are especially diverse, in which numerous combinations of species from quite different florocoenotypes are formed (up to invasion of Dryas oxyodonta into the petrophytic variants of firm-bunch grass steppes).
At the same time, a considerable part of steppe communities, developed at the periphery (mostly western and southern) of the Altai Mountain Country, can be treated as a general reflection of an ancient, but, partly, modern topologically-successional rows of steppe communities transformation on their contact with Turanian-Dzungarian phryganoid formations (communities of northern-desert Turanian subshrubs), and even with previously well developed low mountain Iranian-Turanian phryganoid formations, which are absent now in AMC. Anyway, in Tarbagatai, partly in Kazakhstan Altai (Kurtchum and Narym ranges), in some sites of the northern part of Mongolian Altai and in several localities in the Central Altai we can observe steppe communities, enriched by southern in genesis subshrubs, undershrubs, undersubshrubs, as well as petrophytic communities of primary stages of colonisation of stony habitats by mostly petrophytic subshrubs with moderate participation of bunch grasses, which further turn into steppe communities, enriched by petrophytic phryganoid elements.
It could be possible to consider such communities as a special combined (contact) subtype of steppes – phryganoid steppes (Steppae suffruticulosae vel “phryganosae”). These are communities with species of the genus Brachanthemum – B. fruticulosum, B. krylovianum, B. mongolicum, Astragalus sp. sp. (sect. Cysticalyx) – A. cysticalyx, A. inflatus, A. scleropodius, A. vereszaginii, A. xanthotrichus, Helianthemum soongoricum, above mentioned communities of Hulthemia persica, some communities with Nanophyton mongolicum, also with Acanthophyllum pungens, communities with species of the genus Rhammatophyllum, rare steppe communities with Acantholimon tarbagataicum (or with A. alatavicum in Saur), etc. Unfortunately, all of them are not described at all.
In fact, more common steppe communities with increased role of some Labiatae species and petrophytic variants of them, absolutely homologous to the thyme communities of chalky soils of the Trans-Volga region and North-West Kazakhstan, are also not described. These are communities with leading role of Ziziphora clinopodioides s. l., some Thymus species, Hyssopus ambiquus, H. cuspidatus, Dracocephalum integrifolium, Scutellaria sieversii, etc. Variants of similar communities with Dracocephalum discolor are known up to the North-Western Altai.
These communities in many respects provide originality of the vegetation in Altai-Dzungarian fratria of formations and at its contacts with Middle Asian mountainous vegetation.
Ancient Mediterranean vegetation (Florocoenotypes of warm-temperate (and arid) floras of Ancient Mediterranean subkingdom).
Arid types of vegetation.
Turanian-Dzungarian subshrub deserts (desert phryganoids) (Eremo-phryganion turano-dshungaricum, Desertae turano-dshungaricae). Type of vegetation of temperate and warm-temperate folras, arisen on the Ancient Mediterranean and, partly, ancient xerophytic base in Neogene after the degradation of forest floras in northern parts of Turanian-Dzungarian sector of Ancient Mediterranean in Miocene. This is a zonal type of Northern Turan and Dzungaria, developed in the south of AMC (Zaissan and Barun-Khurai depressions, mostly at the periphery of the latter). Formations of micro- and mesotherm desert subshrubs, undersubshrubs, and short shrubs (sometimes prickly) – Artemisia terae-albae, A. gracilescens, A. schrenkiana, polydominant communities with significant role of Atraphaxis spinosa, Kochia prostrata, Krascheninnikovia ceratoides, Nanophyton mongolicum, Acanthophyllum pungens, communities of some species of Anabasis – A. salsa, A. aphylla, A. elatior, A. eriopoda, Atriplex cana, etc. Communities of Turanian-Dzungarian fratria, but developed in a close contact with communities of Altai-Dzungarian fratria (first of all, steppe).
Central Asian shrub-subshrub deserts (Eremo-phryganion centraleasiaticum, Desertae cenraleasiaticae). Type of vegetation of predominantly warm-temperate Asia, arisen on the Ancient Mediterranean and ancient xerophilous base of more southern (apparently, subtropical) type in Neogene (on the base of Paleogenic desert-savannic communities) and strongly reduced during Pleistocene. It is a zonal type of northern part of Central Asia, spread only in the far south of Altai Mountain Country (Barun-Khurai depression,at the border with Transaltai Gobi, and somewhere in the east of Saur-Tarbagatai foothills). Formations of Haloxylon ammodendron, Ephedra przewalskii, Zygophyllum xanthoxylon s. l. (Z. kaschgaricum), Sympegma regelii (sometimes with Kaschgarica komarovii), Iljinia regelii, Salsola abrotanoides, Reaumuria (Hololachne) songarica (fragmentary up to Chuyskaya “steppe” in Altai), Artemisia mongolorum s. l., derivative communities of annuals Halogeton glomeratus, Micropeplis arachnoidea, etc.
Turanian psammophytic formations (psammophyton) (Psammo-eremophryganion turanicum, Psammodesertae turanicae). Type of vegetation of warm-temperate floras developed on the Ancient Mediterranean base in Neogene and then strongly transformed in Turan and Dzungaria (as a result of high speed of speciation) in Pleistocene-Holocene. Formations of mesotherm xerophytic and psammophytic shrubs, more rarely – subshrubs, as a rule, with psammophilous annual and perennial grasses as well as with other herbs from different families: Calligonum sp. sp. (C. rubicundum, C. crispum, C. affine, C. pavlovii – in Zaissan depression, C. mongolicum, C. gobicum, C. ebinuricum – in Dzungaria and Barun-Khurai depression), Ephedra stenosperma, Hedysarum scoparium, Astragalus gebleri, more rarely – Krascheninnikovia ewersmanniana, Haloxylon ammodendron with grasses: Leymus giganteus, Agropyron fragile, Stipa consanguinea, Stipagrostis pennata, and also Allium caespitosum, Astragalus candidissimus, A. roseus, Soranthus meyeri, Eremurus inderiensis, more rarely – Carex physodes, Rubia rezniczenkoana, Heloitropium acutiflorum and others.
In the past typical psammosavannic species Ammodendron bifolium (A. sieversii) was spread in the southern part of Bukon sands, but it was not a community-forming species (as well as Stipagrostis pennata or Carex physodes). Secondary open communities are formed by species of Agriophyllum, Corispermum, Schismus arabicus, Chondrilla ambigua, etc. This type belong to Turanian-Dzungarian fratria of formations.
Halophytic formations (Halophyton turano-centroasiaticum, Salineta). Type of vegetation of temperate and warm-temperate (and hot-temperate) floras, arisen on the ancient xerophytic and, partly, Ancient Mediterranean base in Neogene. It includes communities of mesotherm (partly oligotherm) xerophytic and halophilous undersubshrubs, perennial caudex herbs, more rarely – shrubs, rather often – halosucculents. Formations of Kalidium cuspidatum, K. gracile, K. foliatum, Hal?cnemum strobilaceum, Suaeda salsa, S. corniculata, S. physophora, Limonium suffruticosum, L. coralloides, Nitraria sibirica, N. schoberi, and also Salsola rosacea, species of the genus Climacoptera, rarely – Tetradiclis tenella, etc.
Relic tall bunch-grass formations (“tussocks”) (Potamomegalopojon asiaticum, Macrogramineta). Type of vegetation of warm-temperate and subtropical Asian floras, arisen in north-subtropical regions of Asia in Paleogene, then turned into a relic condition and spread a little bit from Neogene to Pleistocene, and partly conjugated with other florocoenotypes. Primary communities of this type was constituted by tall xerophilous bunch-hillocky (tussock), usually strongly close-up grasses in the upper layer, and saline meadow small-bunch or rootstock grasses and herbs among the grass hillocks; they has been almost completely degraded to the present time. The main present-time formations of this type are the Achnatherum splendens-stands (“tchievnics”), derivative formations – Iris lactea brakes. Probable relic type – Messerschmidia sibirica s. l.
Tugais (bottomland complexes with forests, bushes and meadows in river valleys) (Xeropotamodrymion asiaticum, Inundato-Silvae deserto-asiaticae). Type of vegetation of warm- and hot-temperate arid floras, arisen on the ancient xerophilous and the Ancient Mediterranean base in the east part of the Ancient Mediterranean at the turn from Paleogene to Neogene. Within the region in question it is spread only at the southern periphery (at the border with Dzungarian deserts), where presented by small fragments. Formations of mesotherm hydrophilous trees and shrubs – Populus euphratica s. l. (mostly subsp. diversifolia), Elaeagnus oxycarpa, Tamarix sp. sp. In the pure state tugais are almost absent in Kara Irtysh and Urunhu valleys, and small fragments of these formations are only developed in the lower reaches of small rivers, found their end in sandy dunes.
Relic elements of “coloured clays” vegetation (Phryganoeremion iranoturanicum). This type of relic (subtropical-Mediterranean) vegetation of “coloured clays” includs communities of mesotherm xerophytic and gypsophilous shrubs, subshrubs, undersubshrubs, and caudex herbs; it is widespread in the Middle Asia, but almost absent in Central Asia, in spite of the fact that “coloured clays”with high salt and gypsum content are rather common here. But rather good number of relic elements, connected directly with this type of vegetation, are also present in Central Asia and at the periphery of the Altai Mountain Country (from Chuyskaya “steppe” up to Tuvinian Altai and Zaissan depression), although they extremelly rare form communities (and even if these communities are rather close up, but predominantly monospecious). The most part of these elements are of respectable (usually Neogene) age, which survived in few refugiums; sometimes they are narrowly or more widespread endemics to the AMC. Besides “coloured” clays and sands (colours of which are highly diverse – white, pinkish, pale, bright red, brick red, even violet, and, consequently, their geochemical characteristics are also sharply different), relic types of this origin also colonise secondary habitats (alluviums, washed off both slopes and horizontal surfaces, and friable chalky soils with gypsum layers). These are remarkable Chenopodium frutescens in the north (which forms sometimes true communities with 2–5 more species), Corydalis stricta (spread within great altitudinal range, but most common on gypsum clays at 2000–2200 m) and, apparently, Zygophyllum melongena, peculiar to saline soils at the altitudes from 1500 up to 2400 m. In Zaissan depression and at the periphery of Barun-Khurai depression these are Calligonum junceum, Atraphaxis compacta, Arthrophytum subulifolium (Zaissan depression only), Sisymbrium subspinescens, Zygophyllum subtrijugum, Euphorbia blepharophylla, Hedysarum splendens, H. kamelinii, Artemisia (Turaniphytum) eranthema, at the border with Transaltai Gobi – Chesneya grubovii and some other similar types.
Semiarid types of vegetation
Ancient Mediterranean semiarid types of vegetation are not present in Altai Mountain Country in the pure state, but Ancient Mediterranean semiarid elements take an active part in formation of steppe shrub communities and shrub steppes at the flat foothill slopes of Southern Altai and Tagbagatai and, sometimes, they even form communities with dominance of typical elements of high grass mountain semisavannas (more rarely – of Iranian-Turanian short-grass semisavannas and shibliak communities, but in combinations with steppe species) in the herb layer. These are, for instance, remarkable communities of Atraphaxis laetevirens, A. virgata, sometimes Amygdalus ledebouriana and Spiraea hypericifolia with dominance in the herb layer of Botriochloa ischaemum, Poa bulbosa, Carex turkestanica, Hedysarum songaricum, Hypericum scabrum, Artemisia sublessingiana, with Fritillaria verticillata, Tulipa altaica, Ixiolirion tataricum, and also with ephemers Anisantha tectorum, Bromus japonicus, Trigonella orthoceras, Alyssum desertorum, A. linifolium, etc., but also with Stipa capillata, Artemisia frigida, Hyssopus ambiquus, Ziziphora clinopodioides, Scutellaria sieversii, Euphorbia pachyrrhiza and others. These are also communities of Inula macrophylla (I. grandis), Alcea nudiflora, Astragalus sieversianus, Ferula soongorica, with Poa bulbosa, Hypericum scabrum, Convolvulus pseudocanthabrica, Eryngium macrocalyx, Haplophyllum perforatum, Oedibasis apiculata, Stroganowia intermedia, S. sagittata, Allium robustum, etc., with an insignificant role of steppe shrubs and typical steppe spesies – Stipa capillata, Artemisia sublessingiana, Koeleria cristata, etc. at the foothills of southern macroslope of Tarbagatai.
Azonal types of vegetation, peculiar to both Boreal and Ancient Mediterranean subkingdoms
Hydrophilous and hygrophilous vegetation (Hydrophyton)
Hygrophilous grass and herbaceous communities (Hygrocoryphion asiaticum). In Altai Mountain Country this florocoenotype is represented mostly by formations of plurizonal or Boreal species of grasses, sedges, and other hydrophytic herbs. These are formation of Phragmites australis, communities of Schoenoplectus, Bolboschoenus, Typha species, diverse species of Carex, Juncus, Beckmannia syzigachne, Glyceria plicata, Scolochloa festucacea, etc.
Submerged plant communities (Hygrocoryphion asiaticum). This type of vegetation is more widespread in AMC and constituted by the Boreal (mesotherm) and more thermophilic species. These are formations of Nymphaea candida, N. tetragona, Nuphar luteum, Nymphoides peltatum, numerous species of the genus Potamogeton, Ceratophyllum demersum, Myriophyllum verticillatum, M. spicatum, Hippuris vulgaris, Najas marina, Caulinia minor, saline water Zannichellia pedunculata, Ruppia spiralis, etc. High species diversiry demonstrate genera Batrachium and Ranunculus, presented by species, occuring in different altitudinal belts and in different types of reservoirs. More southern types are Potamogeton malainus (Zaissan), Batrachium rionii (several localities), Hydrilla verticillata.
Free-floating hydrophyte vegetation (Hydromegaloplankton asiaticum) is also rather rich, but moustly because of a great number of reservoirs at the periphery of the Altai Mountain Country. These are thickets of Lemna minor, L. trisulca, L. turionifera, rather rare Hydrocharis morsus-ranae, Stratiotes aloides, Utricularia sp. sp. It is important, that both in the north and in the south the more thermophilic Trapa natans s. l. (T. sibirica, T. kasachstanica, T. zaissanica, T. pectinata) is spread, Salvinia natans grows in Zaissan depression, and Utricularia australis – both in Zaissan and Barun-Khurai depressions.
Petrophilous vegetation (Petrophyton) is represented in AMC by several not completely distinguishable florocoenotypes, in general similar to Middle Asian and, partly, South Siberian andNorth Mongoliam petrophilous vegetation, bur highly diverse in detail.
Xerolitophytic vegetation (Xerolithophyton asiaticum) is a vegetation of warm rocks, represented by microcommunities of Silene turgida, Ptilotrichum tenuifolium, Chamaerhodos altaica, Sedum hybridum, Eritrichium altaicum, Asperula paniculata, A. kryloviana, Galium coriaceum, Thymus, sp. sp., Dendranthema sinuatum, Brachanthemum baranovii, Saussurea jadrinzevii, Stevenia incarnata, Allium clathratum, A. eduardii, also Woodsia asiatica, W. ilvensis, etc. – in Altai, Allium galanthum, A. subtilissimum, A. saxatile, Galitzkya spathulata, Pseudosedum lievenii, Sedum albertii, Rosularia platyphylla, Seseli incanum, Hyssopus cuspidatus, Thymus roseus, Artemisia glabella, A. salsoloides and others – in western massifs of low mountains, and also in Tarbagatai, Dracocephalum fruticulosum, etc. – at the eastern periphery of Altai. The diversity of litophytes is not high in Altai, but considerable part of them are not widespread and even endemic species.
Xerochasmophytic vegetation (Xerochasmophyton asiaticum) is a vegetation of warm and dry taluses. Obligatory chasmophyric species are not numerous – Silene incurvifolia, Peucedanum hystrix (also peculiar to more wet slopes), Dracocephalum peregrinum, Valeriana martjanovii, Chondrilla leiosperma, Ch. brevirostris, sometimes Scrophularia incisa, Sterigmostemum (Oreoloma) violaceum, etc.
Xeropetrophytic vegetation (Xeropetrophyton) is a vegetation of stony and skeleton slopes and dry tops, which does not form a close-up cover. These communities are more diverse and usually include a lot of species: Silene altaica, S. alexandrae, S. gubanovii, Stellaria amblyosepala, S. dichotoma, S. gypsophiloides, Gypsophila patrinii, Orostachys spinosa, O. thyrsiflora, Potentilla rupestris, Seseli buchtormense, Onosma gmelinii, Lagochilus bungei, Helianthemum soongoricum, Convolvulus gortschakovii, Pyrethrum kelleri, Echinops integrifolius, Syreitschikovia tenuifolia, Ligularia robusta, Youngia altaica, Y. tenuifolia, more rarely – Ephedra intermedia, etc.
Mesolitophytic vegetation (Mesolithophyton sibirico-oreoasiaticum) is a vegetation of rocks with normal moistening (at least temporarily). These are microcommunities of Asplenium septentrionale, A. ruta-muraria, A. viride, A. trichomanes, Cystopteris altajensis, Camptosorus sibiricus, Woodsia heterophylla, Polypodium sibiricum, Lepisorus albertii, Parietaria debilis, Corydalis capnoides s. l., Aquilegia lactiflora (Tarbagatai only), Thlaspi cochleariforme, Sedum populifolium, S. ewersii, Saxifraga sibirica, Hackelia deflexa, Campanula dasyantha and some others, sometimes also communities of Bergenia crassifolia.
Cryopetrophytic vegetation (Kryopetrophyton oreoasiaticum) is a vegetation of cold and dry stony habitats, often belt-forming in high mountains. This is a highly diverse florocoenotype (in fact – several types), peculiar to the Mountainuos Asia, well developed and represented by numerous variants in the Altai Mountain Country.
Kryoxeropetrophytic communities, which usually do not form a close-up cover, include Minuartia verna, M. biflora, Arenaria formosa, Smelowskia calycina, S. pectinata, Chorispora bungeana, Ch. songorica (Tarbagatai), Potentilla biflora, Papaver pseudocanescens, Oxytropis schrenkii, O. chionophila, O. saurica, Eritrichium villosum, Dracocephalum bungeanum, Saussurea pricei, S. glacialis, Valeriana petrophila, some Draba species, etc. These communities are most common in the southern parts of AMC, and spread there (together with other cryophytic types of vegetation) at the highest altitudes up to the upper level of vegetation.
The most original complexes of this florocoenotype can be designated as “pseudotundras”. These forms of high mountain vegetation are developed on plateau-shaped ridges in conditios of an intensive blowing out of snow and, consequently, permafrost formation in thin soil layers. The structure of these “pseudotundras” resemles the strusture of “polygonal”, or “medallion” tundras: an extensive plant-free spots are framed by the skeleton edges with plants, sometimes of rather numerous species. The most common species are: Carex rupestris, Paracolpodium altaicum, Stellaria petraea, Draba altaica, D. fladnizensis, D. oreades, Braya siliquosa, B. rosea, Oxytropis oligantha, O. komei, Rhodiola coccinea and some others. True alpine tundra elements are absent, in exception with diverse and coloured lichen species.
Since numerous rock streams at the tops and under massive unmelting snowdrifts; moraines, weakly fastened by plants; and taluses at the edges of moraines are very common at the high altitudes of Altai Mountain Country, a special variant of cryochasmophytic vegetation, peculiar to such a type of movable habitats, is also developed in AMC. These are communities of Cerastium lithospermifolium, Stellaria martjanovii, Callianthemum angustifolium, C. alatavicum, C. sajanense, Aconitum rotundifolium, Parrya (Neuroloma) stenocarpa, Leiospora excapa, Taphrospermum altaicum, Astragalus tschujensis, Veronica densiflora, Dracocephalum stamineum, D. bungeanum, Youngia nana, etc., in Tarbagatai also with Allium carolinianum, A. oreophilum. Some other species – Saussurea involucrata, S. orgaadayi, Viola biflora and also Chamaenerium latifolium, reaching high altitudes via the edges of small streams, – prefer more large bulks of stones (with better drainage). One of the most ecologically plastic species of this set is Allardia (Waldheimia) tridactylites.
At last, a small group of species, strictly peculiar to rocky habitats, forms cryolitophytic vegetation, including Paraquilegia anemonoides, P. microphylla, Corydalis grubovii, Valeriana fedtschenkoi and some others (species of Draba, Youngia).
Anthropogenic (created or transformed by human) vegetation (Agrophyton). The territory of Altai Mountain Country was settled by human from an ancient time. Already paleolitic man residing in the most favourable regions of Altai and Tarbagatai, had obviously visited high mountain areas, probably, for the purposes of getting better raw material for constructing more perfect stony instruments. According to V.I. Molodin’s and Yu.F. Kiryushin’s opinion, numerous tribes have been residing here in the late Bronze epoch; they were mostly cattlemen. Since even the ancient metal instruments production required a great deal of charcoal for metal melting, these cattlemen, undoubtedly, were able to prolong the time of pasturing by using fire and, moreover, to expand areas of pastures with the same method (burning out the forests). It is almost obviously, that in times of the Pazyryk culture (about 3000 years ago) Altai cattlemen were assotiated with farmers at the periphery of Altai, but it is also logical to suggest that any forms of agriculture had appeared here much earlier. In fact, Altai has never been mastered to such an extent as its periphery along the rivers, flowing down from high mountains, but cattleman population was rather great also several thousands years before Christ, and, moreover, no less than for last 2.5 thousands years this territory was a centre connecting different cultures and, simultaneously, – it was an arena of struggle of powerful “The Great Steppe” state establishments beginning from the Ueges and the Gunns to the ancient Turks, Iran-language ethnoses of Xinjiang, Tanguts-Mongols and also ethnoses of Ancient and Medieval China.
It is a temptation for the botanist to find the cause of phenomenon of the forest-free nonth-facing slope of Tarbagatai in intensive human impact taking its begining from the activity of ancient cultures, having used and felled forests. It is also obviously that extensive spaces of spurce forests (“yelans”) in North-West and Central Altai (where taiga communities prevail) appeared not only in the Ancient Turks times, but were mastered much earlier. Nevertheless, nature of Altai, highly changed during many thousands of years, especially its vegetation, strongly disturbed in the last several hundred years, seems to be almost primary, still living according to natural rules, in comparison with Altai periphery (in spite of our precise knowledge on the cutting down of Western (“Mining”) Altai forests and Chuyskaya “steppe” flood plain forests, and further changes of “desert-steppe” territories of the latter provoked by overgrazing and melioration). Evidently, it is impossible to describe vegetation of this huge region without taking into consideration communities, changed or created by human (vegetation of fields, settlements, roadsides, etc.). It should be noticed, however, that these communities have not been investigated well enough and, moreover, not even described in special literature. In connection with this fact, only a general scheme of subdivision of types of anthropogenic vegetation with some preliminary notes, but without detailed characteristics of them is given here.
Agrocommunities of unirrigated cultures are spread in north-western and western foothills and low mountains of Kazakhstan, Chinese, in less degree – of Russian Altai and part of Western Tarbagatai. These are, first of all, sowings of cereals (spring wheat, more rarely – winter rye, more rarely – winter wheat, barley (both fodder and beer), very rarely – outs). These are also peas, buckwheat, some oil plants – Brassica napus (hybrids), in the past – Camelina, in experiments – Saflora. Weeds of these seedlings are mostly peculiar to European agriculture, but this set was strongly exhausted and changed during the last 50 years (the most common now are rhizome perennials – species of Sonchus, Cirsium, whereas specified annual weeds were almost eliminated; the role of Artemisia sieversiana increased). Weeds of intertilled crops (potatous, cabbage, more rarely – other crops, predominantly fodder), and forage crops (mais, sunflower) are also mostly European, but role of casual aborigen species was also increased.
Specific type of unirrigated cultures are atrificial hay meadows (usually hybrid forms of Onobrychis sativa, more rarely – Medicago sativa). Aborigen grass of unirrigated cultures of this area – millet – was practically superseded by other crops.
Agrocommunities of irrigated cultures are presented in small oasises in the South-Western Altai, in Chinese Altai, in the Bulugun oasis of Mongolian Altai, and in Tarbagatai. The set of crops is nearly the same, but weeds are mostly more specific (Dzungarian-Northern Middle Asian – Echinochloa crus-galli, etc. Setaria glauca, S. viridis s. l., species of Eragrostis, Bolboschoenus, Rumex similans, Heliotropium ellipticum, Lycopsis orientalis, and also – Goebelia alopecuroides, Dodartia orientalis, Acroptilon australe, Artemisia scoparia, etc.). Melon fields with water-melons and bad quality melons and with specific set of aborigen weeds are peculiar to the southern part of AMC. Specific flood plain meadows, exposed in the past to a radical melioration (now deserted) or partly changed by melioration (territories, where fodder herbs has been sown) are common in the northern parts of the region. In the case of radical melioration these were either monocultures or combinations of cultural forms of Trifolium pratense, T. hybridum, hybrid (blue) lucerne, peas and also Phleum sp., cultural forms of Bromopsis inermis, Dactilys glomerata, in Kazakhstanian Altai – Arrhenatherum elatius (now extinct). Species of Trifolium, Onobrichys, Dactilys have been used as a main fodder plants. The more exotic cultivares, propagandized in the past and, partly, successfully used, were Sorghum sudanense, Papaver somniferum, valuable melliferous plant Phacelia tanacetifolia.
Vegetation of settlements. As a rule, these are combinations of natural communities, constituted by aborigen species; ruderal communities (fixed to settlements), including species of different nature; successional communities in demutational rows of disturbed vegetation, formed on the base of invasive species; and also agrocommunities in the strict sense, existing continuously in settlements. It is possible to distinguish several general types of vegetation of settlements of Altai Mountain Country.
First of all, it is an urban vegetation (with insufficient role of kitchen garden agrocommunities, but usually with parks, gerdens (partly – outside the manors), and also with a high role of different industrial landscapes). As a rule, the urban floras are considerably enriched.
This is also vegetation of temporary gardens and summer residences, in Altai – mostly with kitchen garden crops, flower beds, and, in less degree, gardens. The number of the aborigen species is strongly decreased in these communities, but the set of cultivated and invasive species is very diverse.
Countryside vegetation of foothills and highlands includes permanent kitchen garden agriculture, gardens, flower beds, and original composition of ruderal vegetation, constituted by species, “runaway” from the past agriculture, invasive species (including carantine weeds), and widespread Euro-Siberian ruderal species (which are dominating). The invasion of Hordeum jubatum, Cardaria repens, C. draba, Cyclochaena xanthifolia (as Xanthium sibiricum before the latter) is going on now in the countryside of more southern regions; new localities of Conyza canadensis, etc. are also numerous there. A prominent feature of settlements of this type is an experience of the grapes agriculture, as well as the remnants of Humulus lupulus breeding, etc. Asiatic appearance of this vegetation is provided by the high role of Cannabis sativa s. l. in the set of ruderals. Because of the presence of Urtica cannabina, Atriplex sibirica, A. fera, Axyris sp. sp., Lepidium densiflorum, Chorispora sibirica, Ch. tenella, Leonurus quinquelobatus, L. glaucescens, Artemisia sp. sp., etc., the countryside vegetation of the middle altitudes in the forest part of AMC has more Asiatic base. The number of species of Artemisia is most considerable in settlements of the high intermountain depressions.
Vegetation of temporary seasonal (winter) stationts in high altitudes has more sharply expressed Asiatic (mountainous Asiatic) character because of presense of numerous ruderal representatives of Chenopodiaceae, (including annual Axyris, Kochia, Salsola species) and Artemisia (especially A. macrocephala, A. anethifolia, A. pectinata, A. mongolica, etc.). The originality of this flora is also emphasized by the rare species growing in high mountain seasonal settlements – Hedinia altaica, Zygophyllum melongena, etc.
Rather specific anthropogenous vegetation is peculiar to areas of gold mining with using of drags, bulks of raw material near the mines, road borders, where the role of aborigen apophytes is especially high (for example, Nepeta sibirica forms monodominant, in fact, communities along the roads, as well as Dracocephalum foetidum in more dry regions; natural Cuscuta europaea s. l. is often spread there, etc.). But it is necessary to emphasize once more, that the knowledge on anthropogenous vegetation of Altai Mountain Country are extremelly low.
Some general features of the vertical zonality
Such a rich set of types of vegetation in conditions of complicated mountain relief is reflected by the system of the vertical zonality. In fact, rows of changes of vertical zonality differ considerably in different parts of Altai Mountain Country because of differences in climatic charasteristics as well as for the reason of position in different structures of macrorelief. But plant communities are no less diverse within the limits of the single altitudinal belt. In some regions of AMC vertical zonality is now known well enough. There are two special generalizing monoghaphs, in which problems of botanical-geographical localization and subdivision of the main regions of Altai are considered on the base of srtuctural types of vertical zonality in different parts of this territory. One of these works comprises materials of numerous botanists, having worked in Russian Altai, and the author’s data on profile cuts, made in some parts of Altai (Ogureeva, 1980). The role of the author’s materials is especially high in another work on the botanical geography of Mongolian Altai (Volkova, 1994), but, unfortunately, some big mountain massifs were not included in this investigation. Vertical zonality in Tagbagatai has thoroughly been investigated and described in details by E. G. Stepanova (1962); in Kazakhstanian Altai it was characterized by V. I. Baranov (1929), P. P. Polyakov (1934), A. G. Borissova (1935), and then generalized in some inportant theoretical papers by R. A. Elenevsky (1938, 1940). But Kazakhstanian Altai is still a region, the vertical zonality of the vegetational cover of which is investigated most poorly; it ts also only outlined in Chinese Altai, and not described in big massifs Turgen-Ula and Khirekhan-Ula in the east of Altai. These problems of lack of theoretical analysis of materials on the vertical zonality of Altai Mountain Country were recently mentioned by me in the monograph on alalysis of some elements of Altai flora regarding its history.
The general scheme (Kamelin, 1998), presented below (fig.), includes 7 equal sectors (one of them is also subdivided onto 3 subsectors), rather sharply differing in vertical zonality. Different variants of interpretation and generalization of this subdivison, reflecting position of the whole territory situated at the border between different provinces of Boreal and Ancient Mediterranean floristic subkingdoms, meridianal border between sectors of Asia (mostly Non-Tropical), running, in particular, throuhg Altai, and regional specificity of vertical zonality, has also been proposed. The main conclusions drawn from different variants of generalization regarding the floristic subdivision of Altai, are the following. 1) The combined types of the vertical zonality dominate in AMC (with conjugation of different belts within the sectors). 2) Subdivision of the territory in question onto 3 basic regions – Boreal Mountainous South Siberian, Subboreal Steppe Mountainous South Altai-Dzungarian, and Steppe-Desert Central Asian – does obviously reflect most completely the provincial floristic peculiarities of different regions of Altai Mountain Country.
Both in flora and vegetation Altai Mountain Country belongs entirely to the Circumboreal regoin of the Boreal subkingdom, but to different subregions of the latter – Euro-Siberian (represented by the single province – Altai-West Sayan), and Steppe (represented by Altai-Dzungarian province, named before by the author as East Kazakhstanian-Altai-Dzungarian, see Kamelin, 1990, and Tuvinian-Mongolian province). Because of the “depression effect” the provincies of the Steppe subregion contain not only separate elements of Ancient Mediterranean flora, but also its exclaves – in Zaissan and Barun-Khurai depressions and at the eastern periphery – in the Depression of the Great Lakes. Even comparatively high number of mountainous Middle Asian elements in the flora of southern slopes of Tarbagatai and Baitak-Bogdo does not change the whole composition of the Altai flora, which is South Boreal. And even in conditions of high mountains we still don’t reach here the border of Mountainous Middle Asia, where it runs along the ridge (watershed) of Dzungarian Alatau (Goloskokov, 1984; Kamelin, 1973, 1990).
The natural subdivision of Altai Mountain Country, accepted in new “Flora of Altai” is based on these well distinguished provinces (fig. 1).
Circumboreal region of Holarctic
Euro-Siberian subregion
Altai-West Sayan province (A) includes (within the limits of Altai Mountain Country) Northern and Central (Russian) Altai, a part of the Gornaya Shoria, Abakan range, the east part of Russian Altai and the ranges of Western Sayan up to Yenissey. A considerable part of this territory was named by B. K. Shishkin “The District of the Big Chern”. In fact, the main feature of the province is a presense of the “chern” and well developed “chernevaya” taiga. But, at the same time, a mountain taiga without nemoral elements is also greatly developed in this area. This is also the only region in the whole Altai Mountain Country, where a swamp taiga is spread as well as small squares of Sphagnum swamps. Mountain pine forests and birch forest-steppe are common here; subalpine herbaceous communities, alpinogenous herb carpets and alpine tundras are developed at the high altitudes. An extent areas in Central Altai are covered with expositional forest-steppe communities with larch; insular “steppes”, usually surrounded by expositional forest-steppe communities, are well developed in some depressions. Relic fragments of dry and shrub steppes with Tuvinian-Mongolian elements and some specific endemic species are spread in the Katun valley, Chuya lowr course, the upper Ursul basin, and somewhere along Bashkaus and Tchulyshman.
The main differential types of the “chern” survived in the flora of the province, some of them are now endemics or subendemics: Tilia sibirica, Menispermum dauricum, Eranthis sibirica, Dentaria sibirica, Chrysosplenium ovalifolium, Brunnera sibirica (and about 40 species more, also common in Altai-Dzungarian province). Important pine forest species: Zygadenus sibiricus, Polygala tenuifolia, Phlox sibirica, Pyrola media, Dendranthema zawadskii; South Siberian-Mongolian-Dahurian Astragalus melilotoides, A. davuricus; peatbog Scheuchzeria palustris, Drosera rotundifolia, D. anglica, Rubus chamaemorus, Viola epipsiloides, Chamaedaphne calyculata, Andromeda polifolia, Vaccinium uliginosum, Oxycoccus palustris, O. microcarpus occur only in this part of Altai Mountain Country; the peatbog species are spread only in the northern part of this sector. The same distribution within AMC demonstrate Boreal species Schizachne callosa, Agrostis clavata, Calla palustris, Luzula pilosa, Maianthemum bifolium, Corallorhiza trifida, Carex chordorhiza, C. capitata, Viola mauritii, V. selkirkii, Rubus arcticus, Orthilia secunda, Hypopitis monotropa, Angelica palustris, Gentiana pneumonanthe, Lonicera xylosteum, etc. Thus, the majority of these species can only be met in the north and notrh-east of AMC. Such an important taiga and taiga-“goltsy” Siberian species, as Ribes procumbens, Rhododendron aureum, Campanula turczaninovii find there south-eastern limit of their distribution.
Besides above mentioned species, flora of the province contains about 40 species with more or less local distribution – endemics (Altai-West Sayan) and subendemics (Altai-Sayan), more rarely – species also spread insufficiently outside the limits of the province (in Kazakhstanian Altai or in the region of the Korgon range). These are, for instance, Aconitum biflorum, Delphinium retropilosum, Aquilegia borodinii, Ranunculus schischkinii, R. akkemensis, R. pseudomonophyllus, R. submarginatus, Stellaria imbricata, S. (Mesostemma) martjanovii, Silene turgida, Chrysosplenium filipes, Sedum populifolium, Caragana altaica s. str., Bupleurum martjanovii, Galium coriaceum, Scrophularia altaica, Dendranthema sinuatum, Saussurea jadrinzevii, S. serratuloides, etc. A typical differential element of subalpine herbaceuos communities of the province is also South Siberian-North Mongolian Hansenia mongolica.
Although this provice shares rather numerous features of flora and vegetation with the nieghbouring Sayan-Prebaikal province, situated eastward, the “chernevaya” taiga of the Sayan-Prebaikal type is quite different (important differential types – Mitella nuda, Mertensia stylosa). Moreover, the high mountain communities with rose bay shrublets (Rhododendron adamsii, Rh. parvifolium), and tundras with Phyllodoce caerulea, sharply separate these provinces from each other.
Steppe subregion
Altai-Dzungarian mountain province (KAD) within the limits of AMC includes south-western part of Russian Altai, Kazakhstanian Altai, Mongolian Altai (territories, attributed to the basins of Irtysh and Urunghu, some sites with well developed forest belt in the upper reaches of rivers of Kobdo-Gol basin), Tarbagatai, Semistai, Saur, and also Baitak-Bogdo with other ranges enclosing intermountain Barun-Khurai depression, the latter itself and also Zaissan depression. Outside the limits of the territory in question this province includes, evidently, highlands of Chingiz-Tau – Akshatau, and, probably, some parts of Semipalatinsk and Bayanaul-Karkaraly-Kent district of Central Kazakhstan Highlands, and Urkashar range in the Saur-Tarbagatai mountain system (the data on vegetation of which are sparce). The main feature of the province is a wide development of diverse shrub communities (both steppe and mesophilous) in lower mountain belt, and very common presense of the wide forest belt (spread within a great range of altitudes, increaring southward, and, in exception with the most southern mountains, with dark coniferuos forest trees). Another characteristic details are development of the subalpine tall herbaceous vegetation in the regions of interception of wet air masses as well as development of juniper prostrate formations at the high altitudes or prostrate forms of dark coniferuos forest trees, and diversity of kryopetrophytic communities. Steppes are mostly developed in the lower mountain belt (but somewhere in the most southern regions they also reach high altitudes). Semidesert and desert communities of Turanian-Dzugarian type, including psammophytic, gypsophilous-petrophilous, and halophytic ones, are developed in close-up intermountain depressions.
The flora of Altai-Dzungarian province is the richest in Altai Mountain Country. It contains a good number of nemoral-“chernevaya” taiga species (Anemone umbrosa, Parapyrum anemonoides, Eutrema integrifolium, Campanula latifolia, Codonopsis clematidea are peculiar only to this province), and also taiga species which are common in other regions of Altai. A relic “quercetal” set of species, connected predominantly with mesophilous bush communities, is also most complete here. Some species connect flora of the province with floras of oak forest-steppe regions of East (more rarely – Central) Europe: Daphne altaica, Clematis integrifolia, Gymnospermium altaicum, Potentilla rupestris and others. Another group includes species with different relationship, disrtibuted in mountain regions of the Middle Asia: Sibiraea laevigata, Dictamnus angustifolius, Stelleropsis altaica, etc. Obviously, flora of the province also contains a great number of steppe species, both widespread and endemic either to Central Kazakhstan or Altai-Dzungarian province. Some of them have disjunctive areas of distribution (with South Ural-Preural and Altai-Dzungarian fragments and disjunctions over the most part of Kazakhstan) as, for instance, Silene altaica, Astragalus unilateralis, Pedicularis interrupta, etc. Rather high diversity demonstrate also different meadow-steppe, shrub-steppe, high mountain species, connecting this region with different parts of mountain Middle Asia. The majority of such species occur in the southern slope of Tarbagatai and ranges, surrounding the Barun-Khurai depression, but some of these mountain Altai-Middle Asian species reach south-western foothills of Russian or Kazakhstanian Altai (Eremurus altaicus, Polygonatum roseum, Atraphaxis laetevirens and many others). Some eastern Ancient Mediterrannean species also find there the north-east limits of their distribution (Ixiolirion tataricum, etc.). The flora of Zaissan and Barun-Khurai depressions contains numerous Turanian, Turanian-Dzungarian, Northern Turanian-Dzungarian desert elements (Stipagrostis pennata, Carex physodes, Ammodendron bifolium, Heliotropium acutiflorum, etc.), and also Gobian desert-steppe and desert species (Allium polyrhizum, Zygophyllum xanthoxylon s. l., Z. potaninii, Gymnocarpos przewalskii and others).
The flora of the province is highly original: it includes more than 100 endemic species and at least 50 subendemic species (spread slightly outside AMC – in Kazakhstan Highlands, partly in Balkhash highlands or mountain massifs from Semistai southward to the Dzungarian Gates). Autochtonous group speciation on the base of quite separated ancestral types goes on here in some different genera and families of flowering plants. These are psammophytic species of the genus Calligonum (Polygonaceae) in Zaissan depression (5–7 endemic species); tragacanthoid species of the genus Caragana (3 endemic species: Caragana hololeuca, C. bongardiana, C. tragacanthoides) – in the same region; representatives of the section Cysticalyx of the genus Astragalus (up to 7 endemic species) – in ? Kazakhstanian, Chinese, and Mongolian Altai; species of the section Leucophysa ot the same genus (3 endemic races) – in Zaissan depression and in sandy valleys of Kara-Irtysh and Urunghu, and also in Barun-Khurai depression; species of Sterigmostemum (=Oreoloma) and Stroganowia (Cruciferae), Lagochilus (Labiatae) and some other groups. Another noteworthy endemic species are Limnas vereszaginii, Allium caespitosum, A. petraeum, A. robustum, Iris ludwigii, Atraphaxis canescens, Arenaria pentandra, Rosa schrenkiana, R. baitagensis, Amygdalus ledebouriana, Calophaca soongorica, Astragalus kendyrlyki, A. roseus (special section), A. gebleri (the most northern representative of the section Ammodendron), Oxytropis hystrix, O. spinifer, Hedysarum scoparium, H. splendens, Euphorbia blepharophylla, Stenocoelium trichocarpum, Ferula krylovii, F. potaninii, Acantholimon tarbagataicum, Limonium rezniczenkoanum, Mertensia pallasii, M. popovii, M. tarbagataica, Lappula cristata, L. rupestris, Craniospermum subfloccosum, Rindera ochroleuca, Lindelofia angustifolia, Scutellaria alberti, S. krylovii, S. sieversii, Phlomoides gymnocalyx, Paraeremostachys phlomoides, Eremostachys molucelloides, Hyssopus cuspidatus, Linaria hepatica, Asperula kryloviana, Rubia rezniczenkoana, R. dolichophylla, Echinops gordiaginii, E. integrifolius, Saussurea grubovii, S. cana, S. rigida, S. elata, Jurinea chaetodonta, Scorzonera vereszaginii, Chondrilla sajanensis, Ligularia kareliniana, L. schischkinii, Brachanthemum fruticulosum, B. mongolicum, Pyrethrum kelleri. A presense of subendemic genus Ledebouriella H. Wolff (Umbelliferae) with two poorly separated species – L. seseloides (Hoffm.) H. Wolff (Kazakhstan Highlands: southern part from the upper reaches of Sarysu to Chingiz-Tau) and L. multiflora (Ledeb.) H. Wolff (Zaissan depression – Tarbagatai – Saur) – also emphasizes high originality of the flora of the province and (in conditions of high specifisity of the whole flora) naturalness of it as a part of the flora of the Steppe subregion.
Tuvinian-Mongolian province (ZM, UM), as against the two previous provinces, is represented in Altai Mountain Country by a smaller part, whereas the main part of it stretches widely eastward. It includes Chuysky district of the Altai-Khangai subprovince of Altai-Sayan province (Revushkin, 1988), but without the Ukok plateau (i. e. Chuysky district in B. K. Shishkin’s sense (1937) and also, partly, in the sense of A. V. Kuminova (1960), but without the Kurai subdistrict), Tuvinian Altai (Mongun-Taiga), and the main part of Mongolian Altai (in exception with the main watershed ridge, ranges of the Urunghu basin and close-up depression in the central and eastern parts of Barun-Khurai). Finally, massifs of the south-eastern part of Mongolian Altai in between the Shargyn-Gobi depression, the east part of Dzungarian Gobi and Transaltai Gobi also belong to the province. They are: Adzh-Bogdo, Uertijn-Khuren-Ula, Burkhan-Budai-Ula, Khassagt-Khairkhan, and Khan-Taishirijn-Nuruu; the two latter are most close to the mountain system of Khangai.
Outside AMC the Tuvinian-Mongolian province covers the entire Depression of the Great Lakes; Ubsunur (Huvs-Nuur) depression with two ranges: Khan-Khukhei in the south and Western Tannu-Ola in the north; Central-Tuvinian depression; Eastern Tannu-Ola range; the lower course of Tessijn-Gol (without Sangilen plateau); and extensive territories of Western and South-Western Khangai in the basins of Dzabkhan (Zavkhan) and Khungijn-Gol, including the highest range Otgon-Tengri and the west parts of the watershed between the Selenga basin and basins of rivers, belonging to the Valley of Lakes, in exception with Tarvagatai-Nuruu range. An attribution of the south slope of Khangai as well as the highest sites of Gobian Altai to the province is a matter of further analysis, material for which is still not enough. But including into the province of both high ranges and spacious intermountain depressions (in particular, the most southern Shargyn-Gobi depression with development of Halohylon communities in its lowest parts) is justified completely by the whole floristic set of Depression of the Great Lakes, where the desert Gobian elements play insufficient role, and also by the fact that this flora (of Depression of the Great Lakes) is connected well enough with the mountain floras of the Khobdo basin via the floras of more highly elevated Achit-Nuur and Urek-Nuur depressions.
The main feature of the vegetation of the province is extremelly low role of forest communities in the vertical zonality of the overwhelming majority of its ranges. The only tree species is Larix sibirica, dark coniferous species are absent here, in exception with small areas with spurce, developed within the river valleys of the northern slope of Tannu-Ola, where also a stripe of Pinus sibirica in some sites of the timberline is present. All the other stripes with forest communities, spread in Mongolian Altai and Western Khangai, represent, in fact, remnants of the disturbed forest-steppe belt (A. V. Kuminova in her work, focused on Tuva (1985), calls this phenomenon “peristeppe”). It is mostly developed in Mongolia (at the higher altitudes than in Tuva) in such ranges as Turgen-Ula and Khairkhan-Ula, in the eastern part of the northern macroslope of Khan-Khukhei, in Otgon-Tengri (narrow stripe) and, strangely enough, – in the south-east of Mongolian Altai, inverted to the Shargyn-Gobi depression from the north, in Uertijn-Khuren-Ula mountains (Arshantijn-Nuruu) and Burkhan-Budai mountains.
In thorough correspondence with such a distribution of forests in altitudinal range, different variants of mountain steppes and shrub steppes prevail in the profile of altitudinal zonality of mountains of the province. In lower belts they are usually represented by the mountain variants of desert steppes in various combinations with specific semidesert and even North Gobian desert communities, desert-steppes and petrophytic desert-like steppes. In higher belts the steppes (mountain polydominant firm-bunch grass steppes), spread at the southern slopes, and meadow-steppes, combined with fragments of mesophilous shrub communities, are to be met together with specific high mountain complexes. The latter represent usually a typical “goltsy” formations or combinations of Kobresia and Carex communities with spots of kryopetrophytic and kryophytic cushion plant formations. In the north of the province (in conditions of the increased moistening) and at the highest altitudes where a firn fields and ice cups are developed, these complexes include tundra communities.
Because of the good development of steppes in the rows of the altitudinal zonality flora of the province is most close to the flora of Altai-Dzungarian province, but much more poor than the latter. At the same time, its originality is rather high, which is provided, first of all, by comparatively small, but important set of relic desert-steppe and desert species, considerable part of which are not only endemics to the province (mostly to its northern part), but also far-separated taxonomically. There are Chenopodium frutescens, Nanophyton grubovii, Salsola gemmascens subsp. subglabrata, Potentilla astragalifolia, Astragalus monophyllus subsp. gubanovii, Zygophyllum melongena, Euphorbia potaninii, Frankenia tuvinica, Limonium klementzii, Craniospermum tuvinicum, Saussurea ceterachifolia, Asterothamnus heteropappoides, and some more common species, occuring in Tuvinian-Mongolian province at the nortern limit of their distribution, often spread sparcely and having spacious disjunctions (Reaumuria songarica, Zygophyllum brachypterum, Z. rosovii, Potentilla imbricata, Convolvulus gortschakovii, Allium mongolicum, etc.).
Another group (rather considerable) comprises mountain-steppe species (often also with a local or disjunctive distribution and diverse relationship). The most part of endemics belong to the genera Astragalus and Oxytropis: Astragalus aksaicus, A. argutensis, A. changaicus, A. polozhiae, A. tephrolobus, Oxytropis acanthacea, O. borissovae, O. polyphylla (all tragacanthoid, but of quite different relationship), O. ladyginii, O. saposhnikovii, O. martjanovii, O. ulzijchutagi, O. macrosema, O. tschujae, O. alpestris, O. nivea, O. potaninii, O. rhizantha, O. komei, O. lanuginosa, O. mongolica, O. sutaica, O. physocarpa. Representing different sections, but usually with a group speciation, these species demonstrate development of corresponding groups in the flora of the province during a long time. The genus Potentilla displays the same picture: narrowly endemic species P. rigidula, P. inopinata grow here next to numerous closely related species. Another noteworthy endemics of the steppe fraction are Euphorbia tschuensis, Limonium congestum, Scutellaria grandiflora subsp. tuvensis, Pedicularis lasiostachys, Artemisia changaica, and, probably, Cancrinia krasnoborovii (of Mountain Asian relationship). Non-endemic, but differential for the province species of the steppe fraction are quite different: Arenaria mongholica, Erodium tibetanum, Euphorbia mongolica, Dracocephalum fruticulosum, Arctogeron gramineum and others.
Naturally, a number of mesophilous species which are differential for the province, is much smaller. These are, for example, Delphinium barlykense, Astragalus politovii, Swertia banzragczii, Gentiana atrata, Pedicularis moschata. Betula turunica, described from the western part of Ubsunur depression, represents sharply separated type, far from any other (numerous) hybrid races, growing there. The whole number of endemic species in no less than 80. The flora of the most southern part of the province is rather rich, since larch forests, diverse steppes, and spacious high mountain territories (the latters especially in Adzh-Bogdo and Uertijn-Khuren-Ula ranges) are present here. Moreover, the flora of this region is also enriched with Gobian elements (first of all, in Alag-Nuur depression and along the big sairs flowing down from Adzh-Bogdo and the main watershed ridge). These are different elements: Gymnocarpos przewalskii, Dontostemon crassifolius, Atraphaxis bracteata, Peganum nigellastrum, Sympegma regelii, Calligonum gobicum, around the springs – Rosa kaschgarica, Clematis intricata, C. orientalis. Rather numerous Gobi Altai (mostly) species grow here at the western limit of distribution (Ranunculus gobicus, Silene mongolica, Astragalus sancziri, Oxytropis fragilifolia, O. bungei, Seseli grubovii); some species are local endemics (Artemisia feddei subsp. arschantinica, Saussurea klementzii). Phlagrostis pelliotii, Stipa gobica, S. orientalis, Elytrigia nevskii, etc. are common in bush-steppe communities. Chesneya grubovii (species of Tian-Shan relationship) was found at the border with Altai-Dzungarian province. This South-Mongolian district (UM) is just outlined now, since the flora of the northern macroslope of Adzh-Bogdo is still investigated quite weakly. It is possible, that provincial limits should be improved here in the future.
All three characterized above provinces are strongly connected, in spite of the fact that they belong to two different subregions. These connections are rather complicated, what is evident from the following examples. Thus, the genus Taphrospermum, represented by Th. altaicum in alpine belt of all three provinces, unites Altai Mountain Country with the Middle Asia, genus Pachyneurum with the single species P. grandiflorum (distributed mainly in Tuvinian-Mongolian province, but also spread in high mountains of Central Altai and, partly, in Altai-Dzungarian province) is common in the most part of Khangai Mountain Country, in Central and Southern Sayans. The genus Microstigma (represented in Altai Mountain Country by 2 species: M. deflexum, occuring in Altai and Tuvinian-Mongolian provinces, and in Western Sayan, and narrowly Western Sayan endemic M. sajanense) demonstrates Central Asian connections.
In fact, the whole flora (of all three provinces) in the mountain conditions of such a great area is not homogenous. The present subdivision of the territory onto natural regions is based, firstly, on the data on general floristic composition and distribution of endemic and differential species. In many cases these regions correspond to the floristic districts, some of them could be united into subprovinces. But at this stage we refrain from this, since only complete review may result in changes of both some borders and characteristics of differences between the districts. The detailed description of the borders of districts (regions) to be used in characteristics of distribution of species is therefore only given. Probably, this new, improved scheme of more detailed subdivision of the territory will be presented immediately as the “Flora” is completed (based on more precise data).
R.V. Kamelin