Steppe as a plant community. Successions of steppe phytocenoses of the European forest-steppe Structure and composition of steppe phytocenoses medicinal species
Stretching for considerable distances from north to south and from west to east, the steppes on the territory of our country do not remain the same in terms of floristic composition and structure. In the forest-steppe zone, forbs take a significant part in the formation of steppe phytocenoses, among which, along with steppe xerophytes, meadow species are common.
The nature of the grass stand of the steppe is influenced by animal world, in particular the activity of mouse-like rodents.
Steppe soils are predominantly various chernozems, which are replaced by chestnut soils in the south. Steppe plants have a high ash content, and the humus layer, whose thickness can reach 80-100 cm or more, is rich in mineral compounds, including calcium salts, and has a lumpy structure.
On the whole, the areas occupied by steppe vegetation are characterized by a continental climate; the degree of its continentality especially increases from west to east.
Steppe plants develop a deep root system covering a large volume of soil.
Adaptations to high temperature and lack of moisture are also manifested in the internal structure of organs: in cell sclerification, in an increase in the number of veins and stomata per unit area, etc.
Among the plants of the steppe there are those that “run away” from drought, completing the development cycle at a more favorable time of the year.
A greater number of species bloom in the first half of summer, during the best water supply for plants.
Ballista plants are common in the steppe, which have adaptations to keep fruits and seeds from spontaneous fall; but along with this, properties have developed that contribute to the scattering of the primordia and their removal from the mother's organism.
An even greater effect on the distribution of seeds and fruits is achieved in tumbleweed plants.
The steppes are characterized by a large species diversity, and it is richer in the northern steppes than in the southern ones.
In the northern steppes, with a large species and shoot saturation of the herbage, seven tiers are distinguished; the lower one is composed of species with shortened and creeping shoots; plants with tall erect shoots emerge in the upper tier. There are no clear boundaries between the individual tiers, and in each of them there is practically no dominant species, which distinguishes the northern steppes from the southern ones.
In the meadow steppes, the above-ground layer of green mosses is often well developed.
Great saturation of the herbage becomes possible in the presence of plants of different life forms. Plants differ from each other not only in the nature of the underground organs and the ability to root shoots, but also in the form of the shoot, the method of their renewal, the duration of the growing season of the monocarpic shoot before the transition to flowering, and the position of the shoot in space.
Among the cereals there are long rhizome, loose bush, rhizome and loose bush, dense bush.
According to their ecological characteristics, in particular in relation to water, the plants of the northern steppes are also different: among them are numerous xeromesophytes and mesoxerophytes.
In terms of the abundance of species and their participation in the herbage in the northern steppes, forbs predominate, and among grasses, horse and loose broad-leaved forms predominate.
Changes in the herbage take place not only in space, but also in time.
Multi-tiered addition of the steppe phytocenosis in air environment supplemented by tiered placement of underground shoots and root systems in the soil.
In the southern steppes, in contrast to the northern steppes, the role of cereals increases and, accordingly, the participation of forbs in the composition of the herbage decreases.
From rhizomatous cereals in the southern steppes grow wheatgrass, creeping wheatgrass, hairy wheatgrass. The bluegrass is a sharp-leaved - kotneischno - friable bush grass - characteristic of meadow steppes.
With the advancement from north to south, the steppes become less colorful, floristically poorer, the density of vegetation decreases, the edificatory role of large-turf grasses increases, the aerial parts of plants do not always close, the layered structure is less complex, the xeromorphic organization of long-term vegetative plants increases, the role and participation in the addition of the herbage of short-vegetating plants, the number of drought-resistant shrubs and semi-shrubs increases, and the summer break in the growing season becomes more pronounced.
It is necessary to preserve the entire diversity of species, with all their varying properties and traits, in protected areas and sanctuaries, and to preserve the gene pool created by nature.
Steppes are the main value for which the reserve was created. The steppes presented on its territory belong to the northern, or meadow. This means that they are located on the northern limit of the distribution of steppe vegetation.
Among other types of steppes, meadow steppes have suffered the most from human development. The main spaces once occupied by them have turned into arable land. All areas of meadow steppes that have survived in Eastern Europe, lying on watersheds (plakors), can now be counted on the fingers. The Central Black Earth Reserve includes the largest of them - the Streletskaya (730 ha) and Cossack (720 ha) steppes. Other more or less large surviving sections of the meadow steppes of Russia are the Yamskaya steppe of the Belogorye reserve (Belgorod region, 410 ha), the Kuncherovskaya forest-steppe and the Poperechenskaya steppe of the Volga forest-steppe reserve (Penza region, together 450 ha).
The Streltsy and Cossack steppes represent the most typical meadow steppes that have never been plowed (virgin) in their best expression. These steppes avoided plowing due to the fact that from the 17th century they were in the communal use of the archers and Cossacks and were intended only for haymaking and, in part, grazing. They have survived to this day, because. in 1935 they became part of the Central Black Earth Reserve, created thanks to the efforts of Professor V.V. Alekhin, who made a huge contribution to the study of the steppes of the entire Central Chernozem region and especially the Kursk region.
The conservation of steppe areas is not yet a complete solution to the problem of preserving their biodiversity. Meadow steppes retain their basic qualities only when the above-ground phytomass is alienated. The main role in this process in pre-agricultural times (before the beginning of agricultural development of the land by man) was played by large herd ungulates that grazed in the steppes in significant numbers: wild horses - tarpans, saigas, tours. Such large rodents as ground squirrels and marmots were found in abundance, as well as some herbivorous birds: bustards, little bustards, etc. Steppe fires apparently played a significant role, destroying rags. Steppe vegetation in modern absolutely reserved conditions, i.e. with complete non-interference of man in the ongoing processes, gradually gives way to the meadow, the introduction of tree and shrub species is observed. The main reason for this is the intensive accumulation of rags and perennial non-decomposing bedding, the so-called "steppe felt". This is due to the absence of large phytophage animals - consumers of green phytomass, which, dying annually, falls on the soil surface. Under the influence of the litter, the temperature, water and light conditions of the upper soil horizons change. Under these new conditions, long-rhizomatous meadow species become more competitive, and steppe forbs gradually fall out of the herbage; the structure of the vegetation cover changes, the species richness decreases. In order to avoid such undesirable changes, it is necessary to find a suitable replacement for the impact on the herbage of the steppes that wild animals and steppe fires previously had. Such measures can be haymaking or grazing of domestic animals, or a combination of both: mowing, and then grazing after the aftermath. When choosing a protection strategy, one should be guided by the goal of conserving maximum biodiversity. This goal is best achieved by combining different modes, where each of them contributes. Currently, the steppes of the reserve are maintained thanks to human activities: haymaking with different mowing periods and different rotations and grazing of livestock with a moderate load. The haymaking regime has options: annual mowing, haymaking with a five-year rotation, when the site is mowed in a row for four years, and in the fifth year “rests” to replenish the seed bank in the soil, haymaking with a ten-year rotation and grazing after aftermath (nine years of mowing and rest for the tenth year). Immediately after the establishment of the reserve, special experimental areas were also allocated - absolutely reserved areas where there is neither mowing nor grazing. On the main area of the upland steppe in the Central Chernozem Reserve, a hay rotation regime is used.
At the beginning of the 20th century, on the plains of the Streltsy and Cossack steppes, only mowed variants of the meadow steppes were presented. It was they who were proposed for conservation as having outstanding characteristics, which are now listed as the main "reference" for the northern steppes. "Kursk botanical anomaly" was called by Professor V.V. Alekhine these steppes.
The meadow steppes of the reserve are characterized by a rapid change of colors, an outstanding species saturation and richness of the floristic composition, a dense herbage in which several species play a significant role at once, therefore these steppes are called polydominant. Many species of steppe plants grow here, which have become rare outside the reserve due to the destruction of their habitats and are included in the Red Book of the Kursk Region (2001). In the reserve, the populations of these species, as a rule, are quite numerous and reliably preserved. In the steppes of the Streltsy and Cossack areas, such rare steppe plants grow: thin-leaved peony, pinnate feather grass, beautiful, narrow-leaved and pubescent, leafless iris, whitish hyacinth, Sumy cornflower, spring adonis, yellow flax, perennial, veiny, purple goat, etc.
At the beginning of the last century, the steppes had a clearly expressed forb character, i.e. in the herbage, dicotyledonous plants predominated both in terms of their role in aspects and the number of species, and in terms of weight in hay. Grasses also played a very large, but less noticeable role in the composition of the herbage compared to forbs. Among grasses, the predominance of species with more or less wide leaves, as well as the dominance of rhizomatous and loose-shrub types (non-soddy), was noted, which, in combination with an abundance of dicotyledons, allowed V.V. Alekhine (1934, p. 28) to call the northern steppes steppes of “colorful forbs with broad-leaved grasses”.
The northern steppes are characterized by a rapid change of physiognomic pictures (aspects) vegetation, associated with the successive flowering of different plant species, which is one of the distinguishing properties of meadow-steppe phytocenoses. In the meadows to the north of the forest-steppe zone and in true steppes to the south of it, the colorfulness of grass communities decreases. The change of aspects in the Streltsy steppe was first described in 1907 by V.V. Alekhin (1909). Later, this description was included in many popular science, methodological, and reference publications to characterize the "classical" picture of colorful changes occurring in the vegetation cover of meadow steppes. “Such a change of phases is undoubtedly the result of the adaptation of steppe plants in the sense of their distribution in different parts of the growing season: each species has found a certain place for itself, without being strongly constrained by others and less competing with them” (Alekhin, 1934, p. 23).
After the snow melts, which usually occurs in the steppe at the end of March, the brown background of last year's grass dominates. In mid-April, the first flowering species begin to appear, the most noticeable of which is the open lumbago, or sleep-grass with large purple flowers. Almost simultaneously with it blooms spring adonis, or adonis. This species is more abundant and, in combination with Siberian grits, forms a bright golden-yellow aspect of the steppe by early May. Yellow tones continue to dominate in mid-May, but now due to the flowering of other species: spring primrose and Russian broom. By this time, young grass is already growing well, creating a fresh green background. Against this background, by the end of May to replace yellow flowers bright white and purple spots of blooming forest anemone, milky white ranks and leafless iris come. At the beginning of June, the lilac-blue aspect of meadow sage and thin-leaved peas sets in, early grasses also bloom: pinnate and pubescent feather grass, downy oatmeal. By mid-June, the picture becomes very colorful, because. blooms at this time maximum amount species of herbs and most of the cereals. These are such species as mountain and alpine clover, common leucanthemum, purple goat, elecampane hard-haired, blood-red geranium, common meadowsweet, coastal brome, etc. Later, by the end of June, pink will become the predominant color - this is sandy sainfoin in large numbers; a noticeable role is also played by the real bedstraw with yellow inflorescences smelling of honey. The herbage reaches its greatest height and density, the time for haymaking is approaching. Starting from July, the steppe is already noticeably fading, most species fade, the rising cereals obscure the remaining colors. However, some species only now, in the midst of summer, carry their flowering shoots high, which are clearly visible against the background of the steppe becoming straw-colored: Litvinov's larkspur with blue flowers, black hellebore with dark cherry flowers. In the unmowed areas of the steppe, a straw-brown background from dying grass shoots remains until late autumn. In mowed areas, many species have secondary flowering, some plants even manage to give a second crop of seeds in favorable years. All new flowering species can be observed until mid-October. Secondary flowering, however, cannot be compared with the normal one in richness of colors and quantity. flowering plants.
Changes of colorful pictures can vary by year: there are “feather grass” years, when from mid-May to mid-June the steppe resembles a swaying silvery sea, and there are years when the feather grass aspect is not expressed at all. Most other species also form well-marked aspects not annually. The change in aspects over the years is associated, on the one hand, with fluctuations meteorological conditions, and on the other hand, with the frequency of flowering inherent in many herbaceous plants. By highlighting certain phases or aspects, we greatly simplify the observed phenomena. In fact, each phase contains dozens of flowering, fading and blooming plants, which in general creates an extremely complex picture. The steppe changes its appearance not only from day to day, but it does not remain unchanged during the day, because. some species open their inflorescences in the morning, and with the onset of the hottest time, they close until the next day. These are, for example, plants such as purple goat, oriental goat. Other species open their flowers for only a few hours, and then their petals fall off (flaxes are perennial and veiny).
In absolutely protected areas, the development of plants in spring is noticeably delayed due to the large amount of dead plant residues, which contribute to the accumulation of a larger supply of snow, which later melts. The vegetation is significantly inferior in variety of colors and saturation of colors to the mown areas of the steppe. A number of species with bright colors and large inflorescences avoid unmowed areas; here you can rarely find meadow sage, sandy sainfoin, purple goat, blush and many other species common and abundant in the mowed and grazing steppe.
Higher plants can be divided into biomorphs according to the characteristics of the general structure, multiplicity of fruiting and life expectancy: trees, shrubs, semishrubs and semishrubs, perennial grasses, annuals. According to the composition of the main biomorphs, the meadow steppes are characterized by the dominance of perennial grasses capable of multiple fruiting throughout their life - these are polycarpics. So, among the main components of the herbage in the Streletskaya steppe, about 80% fall to their share. There are very few ephemeroids among them; plants that have time in a short spring period bloom and give fruit, after which their above-ground organs die off, and bulbs or tubers remain in the soil: whitish hyacinth, Russian hazel grouse, blushing goose onion. Such a rapid development of ephemeroids is an adaptation to having time to take advantage of the spring moisture reserves in the soil before it begins to dry out; this life form is represented in the more southern variants of the steppes much more widely than in the meadow ones, where drought and heat are not so frequent. In second place are perennial and biennial herbs that bear fruit once in a lifetime and die off after that - these are monocarpics; they make up about 10% of the species composition of upland steppes. The role of ephemeral annuals is small both in terms of the number of species and abundance; are found in a small number of shaggy grains, northern breakwater, ferruginous gerbil and etc . Also, semi-shrubs and semi-shrubs play a small role, in which the lower parts of the stems do not die off in winter, these are plants such as Marshall's thyme, some types of wormwood. In the upland steppe, the spread of tree and shrub vegetation is retarded by mowing. In the absence of mowing (pasture and absolutely reserved regimes), trees and shrubs are represented by a fairly large number of species, and some of them are very numerous (thorns, pears, apple trees, hawthorns, wild roses, etc.).
Steppes are open spaces where strong winds often blow. Under such conditions, the distribution of fruits and seeds with the help of the wind is the most successful way to conquer new territories. In the forest-steppe zone, open areas of grassy vegetation are combined with massifs of forests, with thickets of shrubs that prevent the distant settlement of steppe plants, and among them there are not so many species whose fruits are equipped with effective flying devices. Such plants are called anemochores, they primarily include feathery feather grasses, the fruits of which (caryopses) are equipped with awns up to 40-50 cm long. By the time of ripening, these awns become distinctly feathery, due to which the fruits can be carried by the wind over distances of up to 100 m or more. Such a form of plants as tumbleweed is very interesting; it is represented by a small number of species. In plants of this form, by the time the seeds ripen, the aerial part takes the form of a ball, which breaks off at the root neck and rolls over with the wind, spreading the seeds along the way. The most prominent representative of this form in the Central Black Earth Reserve is the Tatar katran. The steppe slopes in the Bukreeva Barma area, where it grows in significant abundance, during its mass flowering are covered with large white balls and look as if a herd of sheep is grazing on them (Photo). Other representatives of this form are trinia multistem, common cutter. In very many species, the flight properties of seeds or fruits are weakly expressed; the role of the wind is reduced to the fact that it only shakes the stems of these plants and thus promotes seeding. In this case, the seeds scatter from the mother plant by only tens of centimeters (Levina, 1956). The fruits of some species, when ripe and dry, crack, the seeds are scattered around with force (thin-leaved peas, milky-white rank, etc.); such plants are called autohoras. The radius of expansion is also measured only in tens of centimeters or a few meters. The distribution of seeds and fruits with the help of animals (zoochory) in the steppe apparently plays a subordinate role (Levina, 1965), which, however, increases when woody plants with fruits edible for animals are introduced into the steppe; richer than others are myrmecochores - plants whose fruits are taken away by ants (scented and rocky violets, crested source, nun).
Due to the high floristic richness, the uniform distribution of many species and their large abundance, the meadow steppes are characterized by an extremely high species and specimen saturation. Species or floristic saturation is the number of species in a certain area. V.V. Alekhin (1935) registered up to 77 species of vascular plants per 1 m 2 and up to 120 species per 100 m 2 in the Streltsy steppe. “Such saturation of the Streltsy steppe is absolutely exceptional and represents a kind of “vegetative Kursk anomaly” (Alekhin, 1934, p. 65). Later, censuses on meter-sized platforms made by V.N. Golubev (1962a), gave even more striking results. On six surveyed meters, 87, 80, 61, 77, 80 and 84 species were recorded. Apparently, such a high species saturation of vascular plants is not found anywhere else in the temperate zone.
Trying to find an explanation for the "plant Kursk anomaly", V.V. Alekhin wrote that “there may be a connection between the exceptional richness and antiquity of a given territory, since the Kursk steppes lie on the Central Russian Upland, which was not under the glacier” (1934, p. 65).
ON THE. Prozorovsky (1948), objecting to V.V. Alekhin, emphasized that the high species richness of the Kursk steppes is explained by a particularly favorable combination of climatic conditions in this zone, and not by the antiquity of the territory that did not experience glaciation, as evidenced by the gradual change in species richness in an easterly direction, which manifests itself both in the territory, the former and not under the glacier.
G.I. Dokhman (1968, p. 97) believed that the optimal hydrothermal and edaphic conditions of existence in the forest-steppe lead to maximum saturation with individuals, i.e. to high specimen saturation, and the high number of species per unit area "should be partly explained by the heterogeneous quality of the microenvironment, which makes it possible for ecologically heterogeneous plant species to settle on a unit area."
A.M. Semenova-Tyan-Shanskaya (1966), who also noted that the species saturation of meadow steppes and steppe meadows of the forest-steppe differs from all herbaceous watershed communities of the Russian Plain, saw the reasons for this phenomenon in the variable nature of moisture, which explains the existence of ecologically different species in small areas : drought-resistant steppe, real meadow and forest-meadow mesophytes, as well as meadow-steppe plants most characteristic of the forest-steppe in a broad sense.
A.M. Krasnitsky (1983) explained the reasons for the signs of the botanical anomaly in the Streltsy steppe by the mode of anthropogenic protection - mowing. However, mowing alone would not lead to such indicators in any natural conditions. The species richness of the Kursk meadow steppes, which is unique for the Holarctic, can apparently be explained only by a combination of the above-mentioned reasons: natural-historical, physical-geographical, and anthropogenic.
Mowing weakens the competitive power of the dominant species, as a significant part of the assimilating organs is alienated; this deprives them of their leading positions in the interception of light. After mowing, new ecological niches are formed, as a result of which such a large number of species of vascular plants can grow together on a small area, while the role of each dominant species separately is not very high, i.e. the degree of dominance in mowed meadow steppes is low, and most grass stands are characterized by polydominance; the projective coverage of dominants, as a rule, does not exceed 10-15, and more often it is at the level of 5-8%.
The richness of the floristic composition and the high species saturation of the upland meadow steppe entail a complex vertical structure. The herbaceous layer is characterized by high density; soil uncovered by plants can be seen only by ejections of mole rats or other smaller rodents. The projective cover of plants can reach 90-100%, on average not less than 70-80%. The herbage in the period of its maximum development (June - early August) is usually divided into several sublayers (from 4 to 6 herbage sublayers have been identified by different researchers). Layering changes during the growing season: it becomes more complex (the number of sublayers increases) from early spring to summer and becomes simpler by autumn. The highest sub-layer, composed of coastal brome, high ryegrass, rough cornflower, cutweed gill, mealy mullein and other plants, exceeds 100 cm in a wet year. A typical terrestrial layer, consisting mainly of one type of green moss - thuidium spruce, which can cover more than half of the surface soil.
The layering of herbage is accompanied by underground layering. According to the depth of root penetration, all plants can be divided into three groups: small-rooted (up to 100 cm), medium-rooted (up to 200 cm) and deep-rooted (over 200 cm). It must be said that not all researchers share this point of view. There is also a directly opposite view: in the meadow-steppe communities, there is no true layered structure in the underground parts of the communities.
The uppermost layer of soil, most densely intertwined with roots, forms a dense sod, which well protects the soil from erosion. The total depth of the root layer reaches a record depth of 6 m, and possibly more (Golubev, 1962b). The exceptionally high depth of penetration of the roots of meadow steppe plants is determined by the properties of the soil: good aeration and porosity, sufficient moisture in the lower horizons, starting from 1.8 m, deep groundwater, lack of salinity, etc.
The total underground phytomass in the meadow steppes exceeds the above-ground phytomass by 2-3 times, the main mass of roots and rhizomes is located in the soil layer at a depth of 0-50 cm. In the total above-ground phytomass, green and dead (rags and bedding) parts are distinguished. According to the results of many years of research in the Streletskaya steppe, the green part of the aboveground phytomass ranged from 16 to 62 centners/ha in the hay rotation regime, averaging 32 centners/ha, and the total aboveground phytomass - from 21 to 94 centners/ha, on average - 49 centners/ha. ha. Under an absolutely reserved regime, the green part of the aboveground phytomass ranged from 23 to 55 centners/ha, averaging 37 centners/ha, and the total aboveground phytomass - from 50 to 135 centners/ha, averaging 91 centners/ha (Sobakinskikh, 2000) . Thus, under an absolutely protected regime, the total aboveground phytomass almost doubles, but this increase is mainly due to the dead part.
Over the past century, some changes have occurred in the vegetation of the Streletskaya steppe. A decrease in the participation of a group of dicotyledonous plants in the structure of grass stands of the meadow steppe, which determined the high colorfulness of the meadow steppes at the beginning of the century, was noted. The abundance of broad-leaved grasses has increased significantly, among them the coastal rump still plays the largest role, but relatively recently high ryegrass has invaded the upland steppes from meadows and edges and has gained a strong position; its generative shoots can reach a height of 1.3-1.5 m in wet summer. Angustifolia bluegrass, downy oatmeal, Syreyshchikov's bent grass, cocksfoot, steppe and meadow timothy are quite abundant.
Of the coarsely sod grasses, feather grass is the most characteristic and abundant, narrow-leaved and pubescent feather grass are less common; from small sod - fescue, thin-legged comb.
In the first half of the last century, a special characteristic feature of the meadow steppes was the high proportion of low sedge, the tufts of which were found on almost every square meter. V.V. Alekhin considered it an indispensable member of the northern steppes, he even wrote about meadow steppes with low sedge undergrowth. In the second half of the 20th century, its abundance and occurrence decreased markedly in the upland steppes.
The abundance of whitish hyacinth is also reduced. If earlier it was mentioned that this species took part in the formation of aspects together with adonis and primrose, now it is difficult to count several dozen flowering specimens per hectare.
All observers, until the late 1980s, noted Popov's forget-me-not aspect. S.S. Levitsky (1968) wrote that the mass flowering of forget-me-not sometimes gives some parts of the steppe such a bright blue color that from a distance these places can be mistaken for water spaces reflecting the azure sky. To date, this species has lost its role in the creation of the aspect and is now recorded in the steppe only in small numbers.
While some species are reducing their abundance, others are increasing it. Above, we have already mentioned the mass introduction of high ryegrass, which in the first half of the 20th century was completely uncharacteristic of upland steppe grass stands. The second half of the 20th century in the Streletskaya steppe is characterized by the appearance in some places of the Siberian grain aspect, before that it was known that it was rare in the steppe, only a few curtains were noted. Rough cornflower has also become more widespread.
The horizontal structure of the vegetation cover is complex; it is difficult to single out individual communities (phytocenoses) in it, since the herbaceous vegetation is characterized by a continuum, i.e. smooth transitions of some communities to others, which is explained by rather homogeneous environmental conditions on the upland, the richness of the species composition and the predominance of species with a wide ecological amplitude. However, on the other hand, the meadow steppes are characterized by complexity, due to the well-developed microrelief and the complexity of the soil cover. On microelevations with various outlines, in a circle up to 1 m or more, up to 20-40 cm high, as a rule, groups develop with a large participation of dry-loving (xerophilous) plants. In small gently sloping rounded depressions, called saucers, more moisture-loving (mesophilic) species are more abundantly represented. The heterogeneity of the vegetation cover is more pronounced under an absolutely protected regime. The mowing steppe is characterized by a uniformly diffuse distribution of most plant species, which leads to a monotonous pattern in the vegetation cover, because mowing is a powerful leveling factor.
The classification of plant communities of meadow steppes is also associated with problems due to the rich species composition, polydominance, and the difficulty of distinguishing between meadow steppes and steppe meadows. Until recently, the ecological-phytocenotic approach to classification, mainly based on the consideration of dominants, prevailed. This led to the identification of a large number of small and inexpressive plant associations, often differing only in the ratio of the abundances of the same predominant species, which can vary greatly not only from place to place, but also within the same community from year to year and even within one growing season.
IN Lately the floristic approach began to be used more and more widely. Its application to classify the vegetation of the Streletskaya steppe made it possible to classify all communities of the upland mowing part into one association (Averinova, 2005).
It can be said that now the vegetation of the upland meadow steppes of the reserve is mainly represented by forb-broad-leaved grass communities with a significant participation of densely tufted grasses and legumes. Among the forbs, the following species are especially abundant: spring adonis, spring primrose, multi-flowered ranunculus, green strawberry, common meadowsweet, meadow sage, Kaufman's mytnik, rough cornflower, real bedstraw, common cutter, mountain hornwort, etc. Of the legumes, the most prominent role is played by: clover mountain and alpine, thin-leaved peas, sandy sainfoin, etc.
Meadow-steppe vegetation is represented not only on the plains of the Streletskaya and Cossack steppes, but also on the slopes of ravines (beams) with a predominantly southern exposure, where it often has a more steppe character than the upland steppe itself due to the greater aridity of such habitats. On the southern slopes, plant groups can be found, which include species that are not found in the upland conditions of these areas and are of a more xerophilic nature. Vegetation no longer forms a continuous cover, in some places the subsoil is exposed. Mainly confined to the southern slopes are drooping sage and hairy feather grass, as well as sickle-shaped volodushka, Russian muzzle, white broomrape, Siberian istod, chamomile aster, kachim tall and some other plants. It is for the southern slopes that the presence of thickets of steppe shrubs, the so-called dereznyaks, is typical, consisting mainly of steppe cherries, low almonds, called beaver, blackthorn, less often meadowsweet (spirea) Litvinov, and some types of wild roses. At the beginning of May, when the blackthorn and almonds bloom at the same time, some of the slopes become very picturesque due to the combination of white, pink and green. Dereza itself (shrubby caragana), from which the name of these thickets comes, is currently found on the territory of the reserve only in the Barkalovka area. On the northern slopes, phytocenoses have many mesophilic species in their composition and the vegetation approaches meadow. Outside the Central Black Earth Reserve, the remains of steppe vegetation are still preserved precisely along the slopes of the ravines and along the steep banks of the rivers, i.e. in places inconvenient for plowing.
Meadow-steppe vegetation can be restored on the site of arable land, if there are favorable conditions for this: the proximity of virgin steppes, which act as sources of seeds, suitable topography and soil, and the use of haymaking. There are positive examples of such restoration in several areas of the reserve, but this is not a quick process. If it is possible to destroy the steppe ecosystem in a matter of hours by plowing, then it will take decades for nature to restore. So, on the Cossack site there is an old 70-year-old deposit "Far Field" with an area of 290 hectares. At present, the vegetation on its mowed areas is represented by meadow-steppe communities, which, in their properties and appearance, are close to virgin steppes. However, even after such a long period of time, experts note some differences between these restored communities and those that were not subjected to destructive anthropogenic impact. In that part of the Dalnee Pole deposit, where the regime of absolute conservation was practiced, areas of steppe vegetation with well-developed feather grass communities have also recovered, but there is already a significant introduction of shrubs and trees, meadow and even forest species. At the Bukreeva Barma site, a 40-year-old fallow with an area of 20 ha is an example of a relatively fast and successful restoration of feather grass steppes on the slopes of the southern exposure with a close-to-surface occurrence of Cretaceous deposits. Under such drier conditions, the total phytomass decreases, a less significant layer of litter is formed, and pinnate feather grass gains an advantage in comparison with more mesophilic broad-leaved grasses that predominate on upland areas (shore and awnless rump, high ryegrass, meadow timothy grass, etc.).
Where there are no suitable conditions for the natural restoration of the steppe, steppe vegetation can be recreated using specially developed methods. The Zorinsky site became part of the CCHZ in 1998; more than 200 hectares were occupied by former arable land, which by the time the reserve was established was gradually overgrown with weed-meadow vegetation, and part of the land was still used for arable land. The possibilities of restoring the steppe vegetation here in a natural way were very limited, because. very few sites were preserved where steppe species grew, and the set of these species was rather poor.
In order to create more favorable conditions for the restoration of steppe vegetation on fallow lands and arable land, in 1999 the staff of the reserve conducted an experiment on 6 hectares to restore the steppes using a grass-seed mixture from the virgin Streletskaya steppe. This mixture was harvested by mowing different areas at several times, so that seeds of species that ripen at different times could get into it, and then applied to the experimental area. This recovery method was developed by D.S. Dzybov and was called the agrosteppe method.
Over the years since the experiment, specimens of more than 80 plant species have been found, about which there is reason to say that they appeared from the introduced material, including 46 species that were not previously part of the local flora, of which 23 species were noted on the experimental area. - These are rare steppe plants from the list of the Red Book of the Kursk region (2001). Such species as coastal brome, slender-legged comb, perennial flax, sandy sainfoin became quite widespread in the experimental area, bloom and bear fruit well. The first specimens of feather grass began to enter the generative phase in 2002; by now, there are hundreds of fruit-bearing tufts of feather grass and narrow-leaved feather grass.
In general, we assess the results of this experiment as modest, since it was not possible to achieve a close resemblance of the reconstructed communities with those represented in the Streletskaya steppe. If in the future steppe species become fixed in the plant communities of the Zorinsky site, become their significant components and spread far beyond the experimental area, then the experiment will justify itself.
In 2010, on the area of 7 hectares of the former potato field on the Streletsky site, a new experiment was started to recreate the meadow-steppe vegetation: on half of the field, a wide-row sowing of several types of pinnate feather grass was carried out; in the future, the aisles are planned to be sown with seeds of steppe forbs. This method was developed by V.I. Danilov and is used to restore the historical appearance of the Kulikovo field landscape in Tula region. In the second half, the agro-steppe method will be applied again.
The text was prepared by Ph.D. T.D. Filatova
- Specialty HAC RF03.00.05
- Number of pages 184
Chapter I. Literature review on the problem of the Oka flora. b
Chapter 2. Characteristics natural conditions and vegetation of the study area.
Chapter 3. Method of work.
Chapter 4
A. Microclimatic and soil studies.
B. Identification of steppe sithocenoses and their mapping. W
B. Floro-geographical analysis of steppe phytocenoses.
D. Ecological and phytocenotic features of steppe phytocenoses. A. Ecobiomorph composition. b. Ecological and phytocenotic composition. V. Age structure of cenopopulations of edi fi mushers. d. Species saturation. e. Horizontal structure. e. Vertical structure. and. The rhythm of phenological development. h. Physiognomy. And. weather dynamics.
Chapter 5. Discussion of research materials.
Recommended list of dissertations
Successions of steppe phytocenoses of the European forest-steppe: on the example of the Central Chernozem Biosphere Reserve. V.V. Alekhine 2006, candidate of biological sciences Avanesova, Anna Aleksandrovna
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Introduction to the thesis (part of the abstract) on the topic "Steppe phytocenoses of the Oka valley in the Moscow region and their origin"
The protection of nature and the rational use of natural resources is the most important economic task of our socialist state. This problem is given extremely great attention in the USSR, which is reflected in a number of important documents of our era: the Program of the CPSU, decisions of party congresses, government decrees, in the Constitution of the USSR. In the light of these decisions, a comprehensive study and protection of unique natural objects that are experiencing a strong anthropogenic impact in the modern era of scientific and technological progress acquire an exceptionally large role. An example of this is the unique for the Moscow region steppe flora of the Oka valley, the study of which over the past 120 years has been the subject of extensive scientific literature, which has to a large extent polemical character. N.N. Kaufman (1866) suggested that these plants were brought here by the course of the Oka River from the southern provinces, V.I. Taliev (1897) defended the opinion about their anthropogenic origin, D.I. Litvinov (1899) put forward a hypothesis about the relict nature Oka flora, preserved in some areas of the Central Russian Upland from the pre-glacial period. Despite having a large number scientific publications the problem of the origin of the Oka flora still remains unresolved, since there are still works proving either a relict (A.K. Skvortsov, 1969), or an anthropogenic (N.A. Kostenchuk and A.N. Tyuryu-kanov, 1980) character . Unfortunately, all this vast scientific literature approaches the solution of the problem of the origin of southern plants in the Oka Valley mainly from botanical and geographical positions, which is undoubtedly one-sided.
The purpose of this work is to resolve the issue of the origin of the Oka flora in the Moscow region from a phytocenological standpoint. During the study, it was necessary to solve the following tasks:
1) to study the current distribution of the main areas of steppe phytocenoses in the Oka Valley on the territory of the Moscow Region, to conduct geobotanical mapping of the most preserved sites and to compile a list of associations for them;
2) to study the microclimatic and soil conditions of the habitats of the steppe phytocenoses of this territory;
3) to identify the floristic, eco-biomorphic, ecological-phytocenotic and population composition of the components of steppe phytocenoses, their horizontal and vertical structure, productivity, rhythm of phenological development, seasonal and annual dynamics and, by comparing them with similar indicators of steppe plant communities in the Kursk region, decide the question is to what extent the steppe phytocenoses of the Oka valley in the Moscow region correspond to typical zonal meadow steppes of the European part of the USSR.
This work is based on the materials of our own field research in 1975-1982. The modern distribution of steppe phytocenoses in the Oka valley in the Moscow region was studied as a result of route studies in 1975-1978. Stationary phytocenological studies of the steppe phytocenoses of the Oka valley were carried out on the territory of the Drioka-Terrasny State Reserve on 26 standard accounting sites (100 m2 each) and a geobotanical profile (25 m x 500 m). On scale I: 500, geobotanical and soil mapping of the area of steppe phytocenoses of Doly (area of 36 ha) was carried out.
For ease of comparison, various methodological guidelines were used, on the basis of which similar studies of steppe vegetation were carried out in the Central Chernozem and other reserves of our country. As a result of the research, we have developed our own methods:
I. Studying the seasonal stages of the formation of the vertical structure of herbal phytocenoses;
2. Accounting for the flower coverage of aspectable species.
The scientific novelty of this work lies in the elucidation of the similarities and differences between the steppe phytocenoses of the south of the Moscow region with typical zonal meadow steppes of the European part of the USSR, which makes it possible for the first time to approach the problem of the origin of the Oka flora from phytocenological positions.
This work is also of undoubted practical importance, since on the basis of these studies, measures have been drawn up for the regime of keeping steppe phytocenoses under the conditions of a protected regime, which will allow for the long-term preservation of these valuable plant communities in the south of the Moscow region.
The main materials of the dissertation were presented at the All-Union Conference "Experience and Methods of Ecological Monitoring" (October 1978 - Pushchino, Moscow Region), at a joint meeting of the Moscow Branch of the All-Union Botany and the Botany Section of the MOIP (March 1981) and at the Alekhin Readings (Moscow branch of the Geographical Society of the USSR - April 1981).
The dissertation text occupies 184 typewritten pages (continuous numbering); it has 30 tables, 6 figures and consists of an introduction, five chapters and conclusions. The list of cited literature includes 123 titles.
Similar theses in the specialty "Botany", 03.00.05 VAK code
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Restorative dynamics of Eastern European meadow steppes: On the example of the Central Chernozem Biosphere Reserve. prof. V.V. Alyokhina 2005, candidate of geographical sciences Filatova, Tatyana Dmitrievna
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Dissertation conclusion on the topic "Botany", Danilov, Vladimir Ivanovich
The comprehensive studies of the steppe shitocenoses of the Oka valley B of the MOSCOW region and the conditions of their habitats carried out by the NSh "LI made it possible to draw the following conclusions: sloping exposures at the junction of the floodplain with the first floodplain terrace, To a lesser extent, they are found on the tops and southern slopes of sandy ridges and remnants, rarely affected by the floodplain reshsha, Steppe phytocenoses of the Oka valley are confined to soddy podzolized saturated high-humus sandy loam , which are characterized by dark gray (almost black) color, granular structure, high content of humus in the upper horizons (6-12%), slightly acidic reaction (pH of aqueous solution from 6.0 to 7.Z) and in the absorbed complex of horizon A, calcium over magnesium. The Oka River is distinguished by total solar radiation, which corresponds in its levels to more than one territory (the latitude of Kursk-Voronezh). This increased radiation balance favors the habitation of steppe plants here.4. A comparative flora and geographic analysis of the steppe phytocenoses of the Prioksko-Terrasny and Central Chernozem reserves showed that these plant communities are characterized by a similar composition of steppe species with the Pontic and Sarmatian habitat classes (84-85%). At the same time, in the steppe phytocenoses of the Oka valley, a larger percentage of species distributed in the Mediterranean and Central Europe is noted. Species saturation of steppe phytocenoses of the Oka valley in Moe- 159 -
in the KoB region (the number of species per I ijr and 100 arc) is significantly less than in the meadow steppes of the Kursk region (in the Oka valley, 37 species per I m, 78 species per 100 m; in the Streletskaya steppe, per I m*^ - 61 species, per 100 m^ 118); north,
6. Biomorphic analysis of the steppe phytocenoses of the Oka Valley in the Moscow region and the Central Chernozem Reserve showed their fundamental similarity both in the composition of aboveground and underground structures, the plants included in them. .9%, in the Central Chernozemny - 63.0%, underground structures of the brush-root series in the Prioksko-Terrasny Reserve 61.7%, in the Central Chernozemny - 53.4%.7. Analysis of plants by belonging and rpynnaiv! different duration of vegetation also emphasizes the fundamental similarity between the steppe phytocenoses of the Oka valley in the Moscow region and the Central Chernozem Reserve. The increase in the role of short-vegetative plants towards the south is explained by adaptation to the increasing dryness of the climate,
8. Steppe phytocenoses of the Oka valley in the Moscow region and the Central Chernozem Reserve are characterized by a predominant predominance in their composition of plants of the steppe and Obostein meadow ecological and phytocenotic groups over forest-meadow (steppe in the share of the Oka there are 55.8%, in the Kursk region - 73.8%; forest-meadow "In the Oka valley - 33.1%, in the Kursk region - 15.6%", This indicates that the steppe phytocenoses of the Oka valley have a more meadow nature.
rakter, which is undoubtedly due to their more northerly geographical position. Along with this, typical meadow phytocenoses of the Oka valley include 55.9% of forest-meadow species and 3% of forest-steppe species.9. The structure of the age composition of the populations of edificatory species in the steppe phytocenoses of the Prioksko-Terrasny and Central Chernozemny nature reserves is also generally characterized by similar indicators. However, the observed shift in the age spectrum of the pinnate feather grass maximum of individuals from the adult generative group (Streltskaya steppe) to the group of young generative individuals (valley
Oki), indicates a more accelerated rate of passage in the latter case by each plant through the cycle of age stages, which indirectly indicates that the ecotopes of the Oka Valley are approaching the ecological optimum for this plant.10. This is also evidenced by the horizontal structure of the distribution of sods of edificatory species in the steppe phytocenoses of both reserves. Sods of adult individuals of feather grass under hay conditions in the Oka Valley have a somewhat higher density per unit area compared to the Streletskaya steppe of the Central Chernozem Reserve. This indicates that the ecotopic and biocenotic factors of the Oka valley in the Moscow region correspond to the optimal habitat conditions for the main edificatory species of steppe phytocenoses,
11. The seasonal rhythm of the phenological development of the steppe phytocenoses of the Oka valley has one-peak curves for the main phases, which in type corresponds to the meadow-steppe vegetation of the middle zone of the RSFSR. reserve, indicators. However, typical meadow phytocenoses of the Oka valley are characterized by phenological
with a later culmination than those located in the neighborhood -
12, The seasonal change of aspects in the steppe phytocenoses of the Oka valley, as well as in the steppes of the Central Chernozem Reserve, is manifested in the mass flowering of one and those se aspectable species. The smaller number of steppe aspects in the Prioksko-Terrasny Reserve is explained by the gradual loss of some aspectable species as the steppe vegetation moves north. different from the steppe seasonal dynamics of aspectivity,
13, The structure of the aboveground biomass by biological groups of plants in the steppe phytocenoses of the Prioksko-Terrasny and Central Chernozem reserves have a fundamental similarity both in the growing seasons of individual years and in their weather dynamics. In the pinnate-feather grass-forb associations of the Prioksko-Terrasny Reserve, the proportion of densely bushy grasses in the above-ground biomass increases, while the share of leguminous plants, forbs, and rhomboid grasses decreases at the same time. In wet and cool years, the reverse trend is observed. Long-term studies of the dynamics of aboveground biomass in the steppes of the Central Chernozem Reserve are characterized by similar indicators,
1^, Numerous facts of finding steppe phytocenoses at considerable distances north and south of the Oka valley in the Moscow region (village of Zakharyino, Yasnogorsk district, Romanovo, Zaoksky district, Tula region, Argukovo settlement, Zaraisky district, Staraya village Sitnya, Stupinsky district, s, Zelenaya Sloboda and nose, Volodarsky Ramensky district, Moscow region, etc.) contradict the hypothesis
Kaufman 6 the drift of steppe plants from the young regions by the river." The fruits and seeds of these plants could not penetrate against the current of the rivers Sturgeon, Besputa, Skniga, Kashirka, Moscow and Pshsra to the above points,
15. The hypothesis of anthropogenic penetration of steppe plants into the Oka valley is contradicted by I) the growth here of a number of ore oshit plants, the ranges of which do not extend to the southeast of Russia, from where, according to the supporters of this hypothesis, the plants of the Oka flora were introduced; 2) close approach from the south to the Oka (along the basin
Sturgeon) of a continuous strip of chernozem soils (40-50 gal), an undoubted indicator of the former steppeization of this territory, as well as finds in this narrow strip of rupture not only spots of chernozems and chernozem-like soils, but also a number of areas with steppe phytocenoses and southern plants. 16. In addition, our research data on the schloro-geographical, ecobiomorphic, ecological-phytocenotic and population composition, horizontal and vertical structure, seasonal and annual dinshyaik, as well as soil and microclimatic characteristics of the habitat conditions of steppe plant communities of the Oka valley in the Moscow region are characterized by similar indicators with the zonal meadow fescue steppes of the Kursk and Voronezh regions. time degree of anthropogenic impact impact on the natural complexes of the Oka valley (laying roads, designing water intakes near the villages of Luzhkov and Nikiforov, plowing floodplain lands, intensifying grazing of agricultural animals), there is a need for special conservation of areas of steppe plant communities in the south of the Moscow region,
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168. Dae ty lis glomerata - - s p - - - _
170. Eryngium planum - sps-copi - - sol sol - so
171. Euonyraus verrucosa sp-copi s p - - - - _
172. Pestuca pratensis - _. _ - _ _ _ so
173. Festuca valesiaca - - - cop2 sp sp
174. Filipendula vulgaris - - &ol - sp sp-sol cop1 s
175. Fragaria viridis _ - - - sol sp-copi cop2 co
176. Galium verum - - sp un sol sp sp sp-sol
177. Geranium sanguineum - - sol sol - copi cop2
179. Helictotrichon pubescens - _ _ - - - sol
184. Phlomis tuberosa - -. sp - -sol sp
185. Pimpinella saxlfraga - - sol un sp sol sol
186. Pinus sylvestris with op 2 cop2 copi cop 2 - - -1. Plantago lanceolata
191. Sorbus aucuparia - sol sp sol - - -
192. Tanacetum vulgare - - sp - sol - - s
193. Thalictrum minus Mt - sp - - sol sp
195. Trifolium montanum - - - - sol sp sp
197. Yeronica chamaedrys - - sol - sol - sp
199. Urtioa dioica - - copi - - - -
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The adaptation of plants to coexistence in a community is expressed in the horizontal and vertical division of the phytocenosis into structural and functional elements that take various part in the transformation and accumulation of substances and energy.
In the vertical direction, the plant community is divided into tiers. Tier- this is part of the layer of the plant community, to which the assimilating (leaves, stems) or storage (tubers, bulbs, roots) plant organs are confined. The layered structure is especially pronounced in forest phytocenoses (18). Here the upper tier (canopy) is formed by the crowns of the tallest trees; the second tier - lower trees or undergrowth; in the third tier there are shrubs (undergrowth); the fourth tier is made up of herbs and shrubs; in the fifth, ground, mosses, lichens, mushrooms are located.
The underground organs of plants - roots, tubers, bulbs, rhizomes, etc. plants, seedlings, mycorrhiza, etc. The topsoil is a special layer - the forest floor.
In the horizontal direction, phytocenoses are also divided into separate elements - trees, bushes, groups or "spots" of lichens, mosses, herbaceous plants, etc. Such structural elements communities are called synusia. They usually coincide with certain elements of the microrelief (hillocks, hummocks, depressions) and reflect the distribution of certain environmental factors within the phytocenosis (for example, spots of light-loving plants in the “windows” of a dense forest or groups of dry-loving plants on hummocks in the middle of a swamp).
The rhythm of the development of a plant community is expressed in its seasonal variability due to the different biological properties of its constituent species. This phenomenon, called tiering in time, has great importance for the maximum use of substances and energy in the phytocenosis and is determined by the climatic features of the habitat.
The change of one phytocenosis by another as a result of the influence of natural and anthropogenic factors is called successions. The result of countless changes and various successional processes is the modern diversity of the plant cover of the Earth.
On each fairly vast territory, there are many different plant communities belonging to different vegetation types. The totality of all phytocenoses of any area or geographical area called vegetation cover(or vegetation) of the area. The totality of all plant species that make up these phytocenoses is called the flora of a given territory. Thus, it is possible to distinguish, for example, the flora and vegetation of the Moscow region, the Caucasus, Central Asia, USSR, Europe or the whole globe.
In accordance with the predominance of a certain type of vegetation in the vegetation cover of communities, each territory belongs to a certain vegetation zone. Depending on climatic conditions, there are zones of tundra, coniferous forests, broad-leaved forests, steppes, deserts, tropics, etc. There are also transitional, intermediate zones (forest-tundra, forest-steppe, semi-desert, subtropics, etc.), as well as subzones and vertical belts in the mountains.
Each type of vegetation has its own characteristic features, according to which plant communities of this type differ from phytocenoses of other types.
tundra- a zonal type of vegetation, represented by communities of low shrubs, shrubs and grasses, and mainly mosses and lichens. A variety of adaptations of tundra plants allow them to endure severe frosts. However, in the harsh conditions of the North, with a short and cold summer, tundra plants grow very slowly, the roots do not penetrate deeply into the soil bound by permafrost, and the vertical extent of communities is very small. Therefore, the easily destroyed vegetation of the tundra requires special care.
Forests- zonal type of vegetation, uniting communities with a predominance of woody plants. The northern part of the forest zone is formed by coniferous forests (taiga), dominated by coniferous trees (spruce, pine, fir, larch). To the south, the taiga gradually turns into mixed, and then into deciduous forests. Communities of deciduous tree species (birch, oak, linden, maple, ash, aspen, etc.) are widely developed here. Forest phytocenoses have a large biomass, complex structure, and have a strong impact on the environment. The high water protection role of forest communities is well known. Forests are the most important source of timber and other raw materials for the national economy, as well as valuable food products (mushrooms, berries, etc.). At the same time, the forest is a habitat for many animals, birds and insects. The protection of forests is of great importance in human life.
Steppes- zonal type of vegetation, vast treeless spaces covered with perennial grasses growing on chernozem soils. Steppes are usually divided into southern - feather grass and northern - forb. Turf grasses, bulbous plants, and ephemera dominate in steppe communities.
A characteristic feature of the steppe communities is their adaptation to the dry summer period: the development of plants mainly occurs only in the spring. At present, the steppe communities have survived only in reserves, the rest of the territory is involved in agricultural production.
Deserts are a zonal type of vegetation with very poor sparse vegetation, consisting of wormwood, saltwort, saxaul and other drought-resistant plants. There are clay, saline, rocky and sandy deserts. Desert plants are characterized by various adaptations that allow them to conserve moisture (thorns, lignified stems, deeply penetrating roots). The deserts are full of ephemeral plants that develop only in the short period of early spring. Irrigation of deserts makes it possible to grow many valuable crops there (cotton, grapes, wheat).
meadows- type of vegetation that can occur in any zone. Meadow communities consist of perennial rhizomatous grasses, legumes, and numerous species of herbs adapted to conditions of moderate moisture. There are floodplain meadows (along the river valleys), upland (mainland) and high-mountain (alpine) meadows. The meadows are used for haymaking and grazing.
swamps- also a non-zonal type of vegetation, combining phytocenoses that develop under conditions of excessive stagnant moisture. There are raised bogs, in which mainly sphagnum mosses grow, and lowlands, in which sedges dominate. Swamp plants have specific features that allow them to live on poor, cold and waterlogged soils. A characteristic property of swamps is the formation of peat from dead plants.
Aquatic communities- a widespread non-zonal type of vegetation, including phytocenoses of fresh inland water bodies and sea shelves. Plants of aquatic communities have numerous adaptations for life in the water. At different depths, different communities develop, for example, coastal reed beds, then water lilies, pondweeds, hornwort, even deeper communities of algae, etc.
More artificial community or agrophytocenoses - unstable plant communities, most often of few species, created by man, mainly agricultural (fields, gardens, etc.). They can exist only if they are constantly taken care of, otherwise natural overgrowth and the formation of natural communities take place in these places.
Any plant community- an integral part of the landscape, an important link in the overall system of the biosphere. The use of plants in human life is natural and necessary, but this use of nature must be rational. Protecting plant communities means maintaining natural processes in them, helping to restore disturbed balance, and eliminating undesirable factors and consequences of human influence.
Botanical Garden- this is the concentration of collections of living plants - representatives of local, domestic and foreign flora, not only for their study, but also for the creation of new, more productive plant forms. The most important link in the work of botanical gardens is the acclimatization of plants, the adaptation of living organisms to new conditions of existence, to new biocenoses. Botanical gardens enrich the local flora with new representatives of the plant world. They are not only research, but also educational institutions that promote advanced ideas and achievements of botanical science.
There are 115 botanical gardens in our country, among them. V. L. Komarov in Leningrad, Kiev, Batum, etc. The largest in terms of area is the Nikitsky Botanical Garden in the Crimea (635.6 hectares and the steppe branch 454 hectares). The botanical garden of Moscow State University is recognized as the oldest scientific botanical institution in Russia. It was founded by decree of Peter I in 1706 as an apothecary's garden.
BOTANICAL GARDENS
There are also specialized botanical gardens. For example, in 1952, a living collection was created on an area of 45 hectares. medicinal plants All-Union Research Institute of Medicinal Plants (Moscow). It is a leading institution in the field of medicinal plant growing and the development of new medicinal products from plant materials. The collection includes almost 2.5 thousand plant species and is constantly updated with new species.
The most large-scale and significant among the botanical gardens of our country is the Main Botanical Garden of the USSR Academy of Sciences, established in 1945 (361 hectares). There are about 10 thousand plant species in its collection. On this base, extensive research is being carried out on various problems of botanical science, and many issues of protecting and enriching the plant world are also being resolved.
As a manuscript
AVANESOVA Anna Alexandrovna
SUCCESSIONS OF STEPPE PHYTOCOENOSIS OF THE EUROPEAN FOREST-STEPPE (on the example of the Central Chernozem Biosphere Reserve named after V.V. Alekhin)
Voronezh - 2006
The dissertation was completed at the Department of Botany of the Faculty of Natural Geography of the Kursk state university
Scientific adviser:
Official opponents:
Lead organization:
doctor of agricultural sciences, professor
Pruzhin Mikhail Konstantinovich
doctor of biological sciences, professor
Krylov Artur Georgievich
Candidate of Biological Sciences Terekhova Natalia Alekseevna
Voronezh State Biosphere Reserve
The defense will take place on October 20, 2006 at 2:00 pm at a meeting of the Dissertation Council D 212.038.05 at Voronezh State University at the address: 394006 Voronezh, st. University Square, 1. 59 auditorium.
The dissertation can be found in the library of the Voronezh State University at the address: 394006 Voronezh, st. University Square, 1.
Academic Secretary of the Dissertation Council
G.I.Barabash
GENERAL DESCRIPTION OF WORK
Relevance of the topic. The study of succession is not only of deep theoretical interest, but also of practical importance. It is also important to study the succession of the vegetation cover, which occurs as a result of the changes that a person is currently producing with his economic activity (Kamyshev, 1964).
Central Chernozem State Biosphere Reserve named after V.I. prof. V.V. Alekhin (Kursk region) is the basis for studying changes in virgin steppe phytocenoses under the influence of certain protection regimes. The meadow steppes of the Central Black Earth Reserve are unique plant communities that have almost disappeared throughout Eurasia due to plowing and human economic activity (Alekhin, 1936). The processing of biological material accumulated in the Central Black Earth State Biosphere Reserve is of great scientific importance in connection with its compliance with the UNESCO program "Man and the Biosphere" (MAB) and the Convention on the Biological Diversity of Wildlife Species adopted in 1992.
The successions of meadow-steppe phytocenoses under various protection regimes have not been sufficiently studied. It is necessary to know the direction of these changes in order to correct the processes occurring in them and optimize the steppe nature management (Lavrenko, 1959).
2. Determine the direction of succession of steppe phytocenoses with RAS
3. Identify stages of succession of steppe phytodenoses during RAS and RPK.
Scientific novelty of the work. For the first time, the stage of light forests in phytocenoses with an absolutely reserved regime was described, and the stages of vegetation succession were identified under an absolutely reserved regime of protection and a regime of periodic mowing. The dynamics in the expansion of trees and shrubs in all unmowed areas of the Streltsy section of the Central Black Earth Reserve was revealed. Meadow formation was established in areas with the PKK regime and afforestation in areas with RAS in the steppe phytocenoses of the Streletskaya steppe.
Practical significance. The materials presented in the dissertation summarize the accumulated experience in the protection of meadow steppes in the Central Chernozem Reserve, allow you to choose rational ways protection of meadow steppes, confirm the assessment of the absolutely reserved regime given by N.S. Kamyshev (1965), as a regime leading to the loss of meadow-steppe phytocenoses and the emergence of light forests.
Approbation of work. The main provisions of the dissertation were presented and discussed at scientific conferences "Phytocenoses of the northern forest-steppe and their protection" (Kursk, 2001), "Flora and vegetation of the Central Chernozem region" (Kursk, 2002, 2003, 2004), "Anthropogenic impact on flora and vegetation" (Lipetsk , 2001), "Studying and protecting the nature of the forest-steppe" (Kursk, 2002), II International Conference on anatomy and morphology of plants (St. Petersburg, 2002), "Historical landscape. Nature. Archeology. History "(Tula-Kulikovo field, 2002).
To date, the character of virgin upland steppe phytocenoses of the Streletskaya steppe has been studied (Alekhin, 1936; Lavrenko, 1956, 1991; Kamyshev, 1961, 1964; Semenova-Tyan-Shanskaya, 1966; Dokhman, 1968; Mirkin, 2000; Zozulin, 1959). Changes in the flora of the Streltsy section of the Central Chernozem Reserve are reflected (Alekhin, 1909, 1940; Nosova, 1973; Safonov, Sobakinskikh, Pruzhin, Sapronova, 1998; Zolotukhin, Zolotukhina 2000, 2002). The data on productivity and species richness in RAS and RPK were analyzed (Alekhin, 1909; Prozorovsky, 1940; Radulescu-Ivan, 1965; Semenova-Tyan-Shanskaya, 1966; Utekhin, 1977; Sobakinskikh, 2000). A definition of succession is given (Mirkin and Rozenberg, 1998; Rabotnov, 1992) and changes in steppe phytocenoses under different protection regimes are considered (Alekhin, 1925, 1926, 1934, 1940, 1951; Prozorovsky, 1940; Radulescu-Ivan, 1965; Petrova, 1990; Sobakinsky, 2000). The issues of biomorphological features of plants of the Streltsy steppe (Prozorovsky, 1940; Pokrovskaya, 1940; Levina, 1956; Zozulin, 1959; Golubev, 1962; Golubeva, 1964, 1965; Persikova, 1966; Paderevskaya 1977, 1979; Zhmykhova, 1979; Zhmykhova, Filatova, 1997; Filatova, 2000).
Based on the generalized literature data, the purpose and specific objectives of the study were formulated.
Chapter 2. Area, materials and research methods
The object of the study was the meadow-steppe phytocenoses of the Streltsy section of the Central Chernozem Reserve under the conditions of a periodical (ten-year rotation) haymaking regime and an absolutely reserved regime. A section of the Streletskaya steppe with a periodic mowing regime until 1959 was mowed annually. Since 1959, it has been transferred to a 4-year hay rotation, and since 1992 - to a 10-year grazing after-after. Thus, this site has gone through all forms of periodic mowing. An absolutely reserved regime on the Streltsy site was introduced in 1935.
Aspectivity was studied at the Streletsky site under conditions of periodic mowing and conservation regime from April to October 2001-2005. according to the method of I.N. Beideman (1974). As a result of route
observations, the main aspects of steppe phytocenoses were recorded.
Standard geobotanical descriptions were carried out in 2001-2005. on test plots of 100 m2 using the methods of "Field Geobotany" (1959-1976).
The abundance of plants was considered according to the Drude scale, since All previous researchers used this method in the reserve. According to this scale, seedlings of trees and shrubs included in the grass layer were taken into account in absolutely protected areas. Vegetation was classified according to the ecological-morphological (dominant) principle (Yaroshenko, 1953).
Species saturation of steppe phytocenoses was determined on meter squares in triplicate (Whittaker, 1980). The biomorphological analysis of species is based on the system of life forms of Raunkier (Raunkier, 1937) with additions for the analysis of the vegetation of the Central Russian forest-steppe by M.I. Paderevskaya (1977, 1979) and data on biomorphological structures identified by V.N. Golubev (1962). Moisture indices of plant communities were calculated using the scales of L.G. Ramensky (Ramensky et al., 1956)
The determination of the productivity of steppe phytocenoses with an absolutely protected regime and a regime of periodic mowing was carried out by the value of the aboveground phytomass during its maximum development, when at least 80% of vascular plant species pass into the generative phase (phenological maximum). The cuttings were taken on these plots two weeks later, taking into account the lag in the development of plants under the conditions of the unmowed regime (Sobakinskikh, 1996).
To determine the value of biological productivity, cuts were taken using an accounting frame of 0.25 m2 in 8 repetitions. Alienation was carried out at the level of the soil with the cutting of all dense turfs. Litter and moss were collected in separate bags. All aboveground phytomass was differentiated by us into green and dead parts. In the green part of the aboveground phytomass, fractions were distinguished - economic and botanical groups: turf grasses, rhizomatous and loose shrub grasses, sedges, legumes, forbs and mosses. The dead part of the cut was subdivided into rags and bedding. Rag - dead shoots that have retained a connection with the mother plant. Litter (felt) is dead plant residues that form on the surface of the soil. Dry samples were subjected to weight analysis (Utekhin, 1977).
The distribution of trees and shrubs was studied on all unmown areas. Mapping of tree and shrub species was carried out on sample plots of 500 m2 and 10,000 m2, the size of which provides, according to the method of VN Sukachev (1957), the identification of features of forest phytocenoses. A re-mapping of the profile (6 ha) laid down by V.D. Sobakinsky in 1980-1981, which takes place in
meridional direction from the observation fire tower to Petrin log. Horizontal projections of crowns of trees and shrubs and their heights were recorded for the listed areas. As the main criterion for identifying the ratio of the species composition of trees, shrubs and their cumulative role in phytocenoses, preference was given to the area of crown projections - projective cover. The projections of the crowns of continuous thickets of shrubs were plotted along the contours on the plan.
When identifying the main stages of successions of steppe phytocenoses, the time boundaries were determined on the basis of more complete descriptions of phytocenoses by researchers. The classical description of the communities of 1933 (Alekhin, 1936) was taken as the initial state of the vegetation. The stages of succession were distinguished according to the main indicators of species composition, species saturation, the ratio of economic and botanical groups, dominants of the main associations, moisture indices of L.G. Ramensky (1956), chronoclines (Mirkin, 1978). The name of the succession stages was given in accordance with the dominant association.
For data analysis, the method of mathematical statistics was used using programs Microsoft Excel 2002 Biostat.
All Latin plant names are given according to S.K. Cherepanov (1995).
Chapter 3
Studies of meadow-steppe phytocenoses of the Streletskaya steppe showed that changes took place depending on the protection regimes. To assess the current state of vegetation under periodically mowed and unmowed regimes, phytocenoses of the western and central parts of the Streletskaya steppe were studied.
The western part of the Streletskaya steppe is the most visited: an ecological excursion trail passes through it. Numerous studies are carried out on this territory by botanists, soil scientists, microbiologists, zoologists, and climatologists. The central part of the Streletskaya steppe is distinguished by less disturbance by humans, but by a greater influence of animals, which introduce weed and forest species into these phytocenoses.
Aspective pictures are basically similar throughout the territory of the Streltsy steppe. There is a difference in the time of the onset of phases of aspects in the areas with RAZ and RPK. The development and flowering of plants on the unmowed steppe is delayed by 2 weeks. The reason for this is the effect of the litter on developing plants by holding the snow cover for a longer period and creating a barrier to penetration. sunlight. 11 aspects currently highlighted
time (Filatova, 2002), are noted throughout the steppe, some fragmentary. An aspect of Pulsatilla patens (L.) Mill, and Carex humilis Leyss. expressed only in Selikhov bushes, and the aspect of Adonis vernalis L. - in a pasture near a watering place, Anemone sylvestris L. forms an aspect in single places. Myosotis popovii Dobrocz. is rare, and therefore nowhere its aspect was noted. In spring, at the end of April, a common yellow aspect is given by Prímula veris L. and Draba sibirica (Pall.) ThelL The aspect of Salvia pratensis L. is distributed throughout the steppe. Tragopogon orientalis L. s.l. during flowering does not give a colorful aspect. Few blooms of Stipa pennata L. and Echium russicum J.F. Gmel.. Variegated multicolor aspects are observed from the beginning of summer until the end of June. At the end of June, eared grasses (Bromopsis riparia (Rehm.) Holub, Bromopsis inermis (Leyss.) Holub, Arrhenatherum elatius (L.) J. et C. PreslJ) overlap the multicolored herbs and create a general straw-brown background throughout the mowed steppe.
The weight participation and ratio of the main botanical and economic groups of plants in grass stands show that cereals predominate both in non-mowed phytocenoses and in phytocenoses with periodic mowing, and in both cases, rhizomatous grasses prevail (Fig. 1).
EZ mowed steppe □ some May steppe
Rice. 1. The ratio of the mass of economic and botanical groups in phytocenoses of the mowed and unmowed steppe
Zd - turf grasses, zkr - rhizomatous and loose shrub grasses, p - forbs, b - legumes, o - sedges, m - mosses, c - rags, p - litter.
Species saturation in unmowed areas is two times less (24 species per 1 m2) than in mown areas (57 species per 1 m2).
Comparison of the lists of species of steppe phytocenoses with an absolutely reserved regime of the present time and a description of the steppe,
cited by V.V. Alekhin (Alekhin, 1936, 1951) shows that annuals and
juveniles (Acinos arvensis (Lam.) Dandy, Androsace septentrionalis L., Campanula patula L., Draba nemorosa L., Erigeron acris L., etc.), whose seed renewal is difficult due to the litter. Perennial rosette plants also fall out (Echium russicum, Plantago lanceolata L., Plantago major L., Plantago media L.). and also Carex humilis, Festuca valesiaca Gaudin s.I, Amoria repens (L.) C. Presl., Centaurea sumensis Kalen, Draba sibirica (Pall.) Thell, Gagea erubescens (Bess.) Schult.et Schult. Fil., Hyacentella leucophaea (C. Koch) Schur, Polígala comosa Schkuhr, Sedum acre L., etc., which cannot tolerate shading by tall grasses and litter.
Forbs are represented singly and scattered, only Galium verum L. s.I and Agrimonia asiatica Juz. rarely found in abundance. Cirsium setosum (Willd.) Bess, Urtica dioica L. are represented locally in great abundance.
Comparison of the descriptions of the mowed steppe we received with the classical one given by V.V. Alekhin (Alekhin, 1936, 1951) shows that phytocenoses with PKK have retained their species composition. 12 plants not found in our lists: Vicia cracca L., Acinos arvensis (Lam.) Dandy, Cerastium holosteoides Fries, Dianthus andrzejowckianus (Zapal.) Kulcz., Helichrysum arenarium (L.) Moench, etc.
The composition of cereals has changed due to the appearance of Arrhenatherum elatius, Bromopsis riparia in phytocenoses. The abundance of Festuca valesiaca, Carex humilis has decreased.
The species composition of forbs is preserved with some change in the abundance of individual species. We reduced the abundance of Onobrychis arenaria (Kit.) DC., Trommsdorfia macúlala (L.) Bernh, Filipéndula vulgaris Moench, Leucanthemum vulgare Lam., Tragopogon orientalis L., Valeriana rossica P. Smirn. Increased the abundance of Delphinium cuneatum Stev. ex DC., Primula veris. Salvia pratensis, Serratula lycopifolia (Vill.) A Kern..
An analysis of geobotanical descriptions of steppe phytocenoses with an absolutely protected regime provided a basis for the analysis of dropping out and invading plants on unmowed areas.
Most of the steppe phytocenoses with an absolutely reserved regime are meadow and meadow-steppe species, mesoxerophytes, xeromesophytes, semi-rosette juvenile and annual plants, plants of the lower tiers.
Most of the forest and meadow species, eumesophytes and
eurymesophytes, trees and shrubs, semi-rosette and horizontally rhizomatous perennial plants, plants of the upper tiers.
The similarity in the species composition of periodically mowed and unmowed phytocenoses is 42%-44% (according to the Jaccard coefficient), which indicates a significant difference between these communities.
An important change over the past decades is the expansion of trees and shrubs into communities with an absolutely protected regime (Fig. 2). In the descriptions of these sites, I.F. Petrova (1990) indicated only single shrubs. The appearance of these species is associated with the introduction of their seeds into steppe phytocenoses from nearby forests by mammals, birds and wind.
Rice. Fig. 2. Projections of crowns of tree and shrub species on the site of 500 m2 of the unmowed area of the Streletskaya steppe
over the past 20 years. Light forests create shading conditions under the crowns, which leads to the loss of plants adapted to open spaces.
A study of the abundance and frequency of occurrence of species in phytocenoses shows the wide distribution of Argentiierum elataus in the Streltsy area. N.I. Zolotukhin and I.B. Zolotukhina (2000) indicate that the rapid spread of this species began in the 1960s and is associated not only with the biological characteristics of this plant, but also with the mesophytization of the vegetation cover due to an increase in moisture over the past 20 years. Possibly, this distribution of Argentineum etata is connected with a change in the time of mowing: previously, the steppe was mowed from June 13, at present, haymaking activities are carried out in early July. Later mowing allows abundant seed maturation.
This assumption is supported by the work of Wilson and Clark (2001) on the impact of haymaking on the introduction and spread of Arrhenatherum elatius in steppe communities. The authors argue that the annual early summer haymaking (until June 26) for 5 years significantly reduces the projective cover and the abundance of ryegrass.
Thus, changes in the phytocenoses of the Streletskaya steppe have a certain direction, depending on the protection regime. Statement by I.F. Petrova (1990) on the completion of succession on unmowed plots is currently refuted by the presence of light forests. It is planned to replace these communities with forest phytocenoses. The haymaking regime ensures that the steppe areas are maintained unchanged. There is a change in the abundance of some species, the weight participation in the herbage of cereals and forbs.
Chapter 4
The successions mentioned above are due to irreversible changes in steppe phytocenoses, which occurred under the influence of changing interconnected ecological conditions in complex systems of plant communities of meadow steppes.
The complexity of the meadow steppes lies in the combination in them of plants that are different in their biological characteristics, which belong to different ecological groups, various ecological and phytocenotic elements, and various life forms.
During the time that has passed since the establishment of the reserve (1935), there have been changes in vegetation, leading to the replacement of some communities by others.
The non-mow regime created for the steppe phytocenoses in the reserve has never been characteristic of these communities. The pre-agricultural steppe was exposed to the influence of hoofed herbivores, and later people used the steppes for pastures and hayfields. Throughout the existence of the steppes, alienation of terrestrial phytomass and grazing were constantly taking place.
The absolutely reserved regime (removal of grazing and haymaking), as an exogenous anthropogenic factor, led to the emergence of new endogenous factors, which were expressed in:
1. Accumulation of rags and bedding.
2. Delay snow, its slow melting.
3. Changing the temperature regime.
4. Changing the moisture regime (Semenova-Tyan-Shanskaya, 1966).
Since changes in vegetation occur due to the interaction of phytocenosis and ecotope caused by the processes of reproduction and growth of plants, this succession process is an autogenous syngenesis (Mirkin, 1989).
Analysis of descriptions of communities, their characteristics, and values of biological productivity makes it possible to detect the features of changes in the main indicators. At the same time, stages in the development of phytocenoses are indicated, the boundaries of which are determined by the time of the most complete descriptions of phytocenoses (Table 1).
Table 1
Changes in the main indicators in phytocenoses with RAS
Stages of succession Species saturation Weight participation of grasses and forbs in herbage (%) Dominant associations *
100 m2 1m2 Forbs and legumes Grains
I (1935-1939) 101 44 62 38 Fescue-forb
II (1940-1964) 73 35 30.2 63.7 Coastal feather grass
III (1965-1981) 63 33 33.6 64.1 Ground reed grass
IV (1982-2005) 49 24 29.9 69.7 Ryegrass
*According to Alekhin (1935); Prozorovsky (1940), Zozulin (1955), Radulescu-Ivan (1965,1967), Dokhman (1968), A.M. Semenova-Tyan-Shanskaya (1966); Petrova (1990), Sobakinskikh (2000), Avanesova (2004)
The processing of descriptions according to ecological scales indicates changes in phytocenoses towards an increase in moisture levels (51.9 to 56 index), which indicates the transition of meadow steppes to fresh meadows. According to L.G. Ramensky (1956), meadow steppes are characterized by indices 47-52, while indices 53-63 characterize dry and fresh meadows.
For 70 years (until 2005), steppe grass stands with a reserved regime changed in the direction:
1. Decrease in the species diversity of phytocenoses, the number of species per 1 m2 and 100 m2 decreased by almost 2 times.
2. Changes in the ratio of the weight participation of the main plant species. The proportion of grasses increased in cuttings, while the proportion of herbs and legumes decreased.
3. Loss of annuals and juveniles, especially at the first stages of development.
5. Overgrowth and distribution of feather grasses, “hobble” (6070s), later limited, along with an increase in the presence of rhizomatous (coastal rump) grasses.
6. Increase in indicators of moisture.
7. The introduction of forest and meadow, weedy, edge species of herbaceous plants.
4.2. Expansion of trees and shrubs in the Streletskaya steppe
At the VI delegate congress of the All-Union Botanical Society in 1978, the director of the reserve A.M. Krasnitsky and botanists O.S. Ignatenko, B.C. Zhmykhova, V.D. Sobakinsky noted the absence of expansion of tree and shrub vegetation in the reserve under the reserve regime (Ignatenko et al., 1978).
Initially, shrub components were found in the steppe in the steppe phytocenoses of Cerasus fruticosa Pall., Chamaecytisus ruthenicus (Fish, ex Woloszcz.) Klaskova, Genista tinctoria L..
In the descriptions of steppe phytocenoses by various authors (Alekhin 1935, Dokhman, 1967, Semenova-Tyan-Shanskaya 1966, Radulescu-Ivan 1967, etc.), trees and shrubs were not indicated, with the exception of the steppe shrubs named above. In the work of I.F. Petrova (1990), who carried out a geobotanical survey of the Streletskaya steppe, trees and shrubs were not noted or indicated as single ones.
In all unmowed areas of the Streletskaya steppe, 13 species of trees and 17 species of shrubs were identified. The species found by us are given by N.I. Zolotukhin, I.B. Zolotukhina (2002) in the list of the flora of the Streltsy upland steppe.
On the meridional profile laid down by V.D. Sobakinskikh in 1980-1981 (Avanesova and Sobakinskikh, 2006), Pyrits pyraster (L.) Burgsd (1 sp.), Malus praecox (Pall.) Borkh (4 sp.), Rosa canina L. (2 sp.), Lonicera tatarica L. (1 specimen), Quercus robur L. (3 specimens), Populus trémula L. (3 specimens). On this basis, 1980 can be considered the beginning of the expansion of tree and shrub flora.
A re-survey of the profile showed a significant increase in the number of trees and shrubs. To date (2005), 46 specimens of trees of different ages have been identified on the profile (Quercus robur (4 specimens), Populus trémula (4 specimens), Ulmus glabra Huds. (4 specimens), Acer negundo L. (6 specimens) , Acer platanoides L. (1 sp.), Pyrus pyraster (L.) Burgsd (19 sp.), Malus domestica Borkh and Malus praecox (Pall.) Borkh. (7 sp.)) and 52
thickets of shrubs of various sizes (Louicera tatarica L.(17), Rosa canina L. and Rosa majalis Herrm. (4), Prunus spinosa L.(23), Rhamnus cathartica L.(3), Cerasus fruticosa (4), Crataegus curvisepala Lindm.(l) Mapping of sample plots of unmowed areas of the Streletskaya steppe and profile shows that the projections of crowns of trees and shrubs currently occupy up to 20% on them (Table 2).
The appearance of trees and shrubs in unmowed areas can only increase expansion, because. most of them are currently in the fruiting stage. The death of trees and shrubs
table 2
Projective cover of crowns of trees and shrubs on trial plots of the Streltsy plot (in %)
Unmown area No. 1 500 m2 Unmowed area No. 2 500 m2 Unmowed area No. 3 10,000 m2 Unmowed area No. 4 10,000 m2 Profile on unmowed area No. 2 64,000 m2
21,5 14 9 3,5 4,7
almost not observed. Established woodlands can further lead to the development of forest communities. The forest-steppe is characterized on the watersheds by the presence of both forest communities and meadow steppes. The history of steppe forestry, experience and observations of the development of unmown areas, which were summarized by N.F. Komarov (1951), talk about the possibility of afforestation of phytocenoses with a reserved regime. This point of view is confirmed by the work of L.I. Denisman (1967) based on the excavations of marmots in the Dubroshina tract adjacent to the steppe, in which the author claims that this territory was previously a steppe. Modern works on the Far field of the Cossack steppe also show the overgrowing of steppe territories with trees and shrubs (Ryzhkova, Ryzhkov, 2001)
There is another opinion about the resistance of steppe phytocenoses to overgrowing with trees and shrubs. It consists in the fact that light forests will not give way to forests, but the indigenous steppe phytocenoses will be restored. This point of view is based on ideas about the originality of the forest-steppe (Dokhman, 1967).
Thus, the further development of vegetation in unmowed areas will depend on the new ecological conditions created on them, the biology of plants growing in the community, especially on trees and shrubs that are strong in the competitive relation.
4.2. Stages of succession in areas with absolutely reserved
Based on the analysis and generalization of the available descriptions of steppe phytocenoses for different years (Alekhin, 1935; Prozorovsky, 1940; Zozulin, 1955; Radulescu-Ivan, 1965,1967; Dokhman, 1968; Semenova-Tyan-Shanskaya, 1966; Petrova, 1990; Sobakinskikh , 2000) and our own research results (Avanesova, 2004), the following stages of succession were identified (Fig. 3).
Stage I Fescue-forb (1935-1939) was distinguished on the basis of a comparison of the data of V.V. Alekhin (1935) and N.A. Prozorovsky (1940).
At the first stage of succession, the plots differ physiognomically. Vegetation cover becomes more monotonous and less colorful, aspects change, plants lag behind in their development, some species of forbs proliferate (^1ed¡cago
Gstadia stage II
III stage
Afforestation
o 5 10 15 20 25 30 35 40 45 50 55 60
succession time,
Rice. 3. Scheme of succession in areas with an absolutely reserved regime
falcata L., Adonis vernalis, Pulsatilla patens, Thalictrum flexuosum Bemh. ex Reichenb., Vicia tenuifolia Roth), there is a group distribution of Onobrichis arenaria (Kit.) DC., Senecio jacobea L., Linum perenne L..
Species saturation decreases from 54 to 44 species per 1 m2.
The coefficient of species saturation of rhizomatous and loose shrub grasses Bromopsis riparia, Bromopsis inermis is increasing. Festuca pratensis Huds., Phleum phleoides (L.) Karst..
The abundance of feather grasses is increasing, in some places their aspect is observed. The role of Festuca valesiaca Gaudin s.l., Carex humilis is decreasing. The abundance of some legumes is decreasing (Astragalus danicus Retz, Lotus corniculatus L. s.i.;.
The number of annuals and weeds is decreasing (Androsace septenirionalis, Arenaria uralensis Pall, ex Spreng, Leontodon hispidus L.J
There is a decrease in shoots per unit area (Myosotis popovii -38 per 1 m2, Salvia pratensis 24 per 1 m2).
Festuca valesiaca, Roa angustifolia L. are the dominant plants from cereals, Filipéndula vulgaris Moench from forbs.
Mosses are abundant, on every square meter. Litter accumulation begins.
Stage II Coastal-feather-grass (1940-1964) was distinguished on the basis of the generalization of the data of G.M. Zozulina (1955), D. Radulescu-Ivan (1965, 1967), G.I. Dohman (1968), A.M. Semenov-Tyan-Shanskoy (1966)).
At the second stage of successions, Bromopsis riparia becomes the common dominant. The abundance of Elytrigia intermedia, Bromopsis inermis, Calamagrostis epigeios (L.) Roth, Roa angustifolia, Helictotrichon pubescens, Stipa pennata is increasing. The weight participation of cereals in the herbage increases to 50-60%, the aspect of cereals is noted.
The spread and vitality of the feather grasses, which form an aspect, is intensifying. "Walking" occurs.
The not very common Arrhenatherum elatius appears. The abundance of Festuca valesiaca and Carex humilis continues to decrease.
The abundance of Vicia tenuifolia, Filipéndula vulgaris, Fragaria viridis (Duch.) Weston, Galium verum, Salvia pratensis is decreasing.
There are fewer spring flowering plants (Adonis vernalis, Pulsatilla patens).
The participation and abundance of legumes decreases, only Chamaecytisus ruthenicus plays a significant role in places.
At this stage of succession, the species saturation is 35 plant species per 1 m2.
The mosses are almost gone.
Litter accumulation is stabilized.
Stage III Veynikova (1965-1982) was identified based on the materials of I.F. Petrova (1990), V.D. Sobakinsky (2000).
At this stage of successions, a high degree of dominance of Calamagrostis epigeios is noted. Flat areas of uplands and watershed spaces are occupied by the ground-reed association (about 70%). In it, the subdominants are Roa angustifolia, Bromopsis inermis, Stipa pennata. Rump associations occupy 20% of the territory. Feather-grass associations are preserved only on microelevations, although Stipa pennata is distributed throughout the upland with
constancy 83% and occurrence up to 60-80%. The weight participation of cereals in the herbage is 3 times greater than that of forbs.
The number of colorful aspects is reduced to 4. Filipéndula vulgaris, Fragaria viridis, Vicia tenuifolia, Achillea millefolium L., Stachys officinalis (L.) Trevis, Hypericum perfortum L., Galium verum, Phlomoides tuberosa (L.) Moench., Salvia pratensis dominate among herbs. Species saturation is 33 species per 1 m2. Stage IV The development of woodlands with a predominance of the ryegrass association (since 1982) was identified based on the materials of V.D. Sobakinskikh (2000) and our data.
There is an expansion of trees and shrubs, which by 2005 occupy 6-20% of the territory, which has changed the face of the unmowed steppe. Calamagrostis epigeios, Poa angustifolia, Bromopsis inermis, Stipa pennata, Elytrigia intermedia dominate. Among the dominants is Arrhenatherum elatius, it is distributed in all associations, creating by mid-summer a common aspect in most of the unmowed steppe. Carex humilis has completely disappeared.
The introduction of forest, edge and meadow species is noted (Convallaria majalis L., Aegopodium podagraria L., Anthriscus sylvestris (L.) Hoffm., Pyrethrum corymbosum (L.) Willd, Veronica spuria L., etc.).
There was an increase in associations of weeds Urtica dioica, Cirsium setosum. Species saturation is up to 24 species per 1 m2.
Chapter 5
On the mowing steppe, during the existence of the reserve, environmental conditions have changed due to the following factors:
1. Removal of grazing in spring and autumn after the aftermath.
2. Changing mowing dates. If earlier the steppe was mowed from June 13, then during the conservation, the dates were postponed to the beginning of July.
3. The emergence of new mowing methods. If earlier they mowed with a scythe, and raked the hay with a rake, then later they switched to machine methods of harvesting.
Since changes in vegetation are caused by external anthropogenic action in relation to phytocenosis, these succession processes are related to allogeneic geitogenesis (Mirkin, 1989)
Phytocenoses with PKK in a somewhat modified form are close to the original ones described by V.V. Alekhin (1936) at the beginning of the 20th century. Changes in the main indicators also make it possible to distinguish the stages of succession in areas with periodic mowing (Table 3).
Processing descriptions on ecological scales shows that steppe phytocenoses with periodic mowing have changed the moisture index from 51.9 to 52.7 and are currently on the border between meadow steppes and dry, fresh meadows.
Table 3
Changes in the main indicators in phytocenoses with PKK
Stages of succession Species saturation Weight participation of grasses and forbs in herbage (%) Dominant associations*
100 k? 1 m2 Forbs and legumes Cereals
I (1935-1939) NO 54 62 38 Fescue - forb
II (1940-1964) 99 56 66.7 26.8 Fescue-shore-brim-forb
III (1965-1981) 97 55 55.3 44.7
IV (1982-2005) 87 57 43.8 52.5
*According to Alekhin (1935); Prozorovsky (1940), Zozulin (1955), Radulescu-Ivan (1965,1967), Dokhman (1968), A.M. Semenova-Tyan-Shanskaya (1966); Petrova (1990), Sobakinskikh (2000), Avanesova (2004)
While maintaining the species diversity of the steppe phytocenoses of the Streletskaya steppe, changes are noted in areas with a haymaking regime, which are expressed as follows:
1. An increase in the abundance of rhizomatous and loose shrub grasses and their weight participation.
2. Oppression of turf grasses.
3. Decreased abundance of Carex humilis.
4. Reducing the abundance of some plants from forbs.
5. Loss of annuals and juveniles, Miosotispopovii.
6. In the distribution of Bunios orientalis L., Rhinanthus angustifolius C.C. Gmell.
7. Decreased abundance of late flowering species, Veratrum nigrum L..
8. Increasing the moisture content of phytocenoses.
When mowing, the Streletskaya steppe remains multicolored, retaining its main aspects, and is characterized as a community close in its state to the original, noted during the creation of the reserve. The haymaking regime ensures the preservation of the species diversity of the Streletskaya steppe, noted at the beginning of the century in the descriptions of V.V. Alekhine.
5.2. Stages of succession in areas with periodic mowing
When identifying these stages of succession, more complete descriptions of phytocenoses for different years were considered (Alekhin, 1935; Prozorovsky, 1940; Zozulin, 1955; Radulescu-Ivan, 1965, 1967; Dokhman, 1968; Semenova-Tyan-Shanskaya, 1966; Petrova, 1990; Sobakinskikh, 2000, Avanesova, 2004). In areas with periodic mowing, 4 stages of successions are distinguished (Fig. 4).
Stage I Fescue-forb (1935-1939) was identified based on a comparison of the data of V.V. Alekhin (1935) and H.A. Prozorovsky (1940).
By 1939, the mown areas changed little in comparison with the original ones. Vegetation cover keeps species diversity and main aspects. Plants are ahead in their development of plants in areas with RAS. The vitality of many plants increases, especially in Festuca valesiaca, as well as rhizomatous grasses. Stipa pennata is increasing its distribution. The study of aspects shows an increase in the abundance of Myosotis popovii, Salvia pratensis, Ajuga genevensis L., Knautia arvensis (L.) Coult., Leucanthemum vulgare.
The moss cover in places occupies 100%. The weight participation of forbs in the herbage is greater (70%) than that of cereals.
1st stage II stage III stage IV stage
olugovenie
111111111111111" 0 5 10 15 20 25 30 35 40 45 50 55 60
Succession time, years
Rice. 4. Scheme of succession in areas with periodic mowing
II stage Fescue-coastal brome-forb. (19401965) was identified according to G.I. Zozulina (1955), D. Radulescu-Ivan (1965), V.D. Sobakinsky (2000).
The species composition changes slightly. Diversity dominates. Up to 77% are polycarpic perennials, annuals up to 9%. By weight, herbs make up 61-71%. The forbs define numerous colorful aspects. Salvia pratensis, Filipéndula vulgaris, Galium verum, Vicia tenuifolia, Onobrychis arenaria (Kit.) DC. bloom profusely. Festuca valesiaca, rhizomatous and loose shrub grasses (Bromopsis riparia, Koeleria cristata (L.) Pers.) are abundant. There are well-marked aspects of Stipa pennata. The presence of mosses is reduced.
Species saturation decreases to 56 species per 1 m2.
Stage III Forbs-coastal rump. (1965-1982) is described in the works of I.F. Petrova (1990), V.D. Sobakinsky (2000).
The areas under the Coastal Coastal Association are increasing. Participation in associations of Festuca valesiaca decreased: it left the groups of dominants. The weight participation of forbs is inferior to cereals. The presence of Carex humilis continues to decrease.
Of the forbs, Vicia tenuifolia dominates. Filipendula vulgaris. Salvia pratensis. Tragopogon orientalis L., Amoria montana (L.) Sojak, Achillea millifolium, Knautia arvensis (L.) Coult, Leucanthemum vulgare reduce their phytocenotic role.
Species saturation is 55 species per 1 m2.
Stage IV of mixed herbs-shore brome-ryegrass (since 1982) was identified on the basis of materials by V.D. Sobakinskikh (2000) and our data.
The main species composition of the meadow steppes and the reference qualities of the communities are preserved.
While maintaining the species diversity of grasses participating in herbage, Bromopsis riparia and Stipa pennata remain abundant. The dominant species is Arrhenatherum elatius, which is found in all the census areas and overlaps the forbs aspect.
There are 4 types of sedges in herbage. Carex humilis is found singly.
There are numerous legumes (13 species) with scattered and single abundance. Only Vicia tenuifolia is abundant in places.
In areas with a periodic mowing regime, the spread of Calamagrostis epigeios is restrained, because. its seeds ripen late, the growth of its shoots is slowed down by haymaking.
Species saturation remained at the same level.
Changed the coefficient of moisture L.G. Ramensky due to an increase in the abundance of meadow elements, meadowing occurred.
Chapter 6
steppes with successions in other reserves of the European forest-steppe
The chapter considers the experience of protection of the following reserves: "Stone steppe" of the Voronezh region (Kazantseva, 2002); "Gapichey mountains" of the Lipetsk region (Danilov, 2005); "Volga forest-steppe", Penza region(Kudryavtsev, 2002); "Mikhailovskaya virgin lands" in Sumy region, in the branch of the Ukrainian steppe nature reserve (Tkachenko, 2005); "Cossack Steppe" of the Central Black Earth Reserve (Neshataev, 2006).
A comparison of the succession of the steppe phytocenoses of the Streltsy steppe with the successions in the forest-steppe reserves showed that under an absolutely reserved regime, trees and shrubs overgrow in place of the meadow steppes, in some cases at a faster pace (for 65 years in the Stone Steppe, the overgrowth of trees and shrubs was 60% ). With the development of succession in the unmowed areas of the above protected areas, there was a decrease in species saturation. Decreased spread of weeds. Ryegrass (on hayfields) and nettle (on unmowed plots) communities appeared on the Mikhailovskaya virgin lands, just as in the Streletskaya steppe.
In all the considered reserves, with successions directed towards the development of tree and shrub communities, overgrowing occurs due to similar species (Cerasus fruticosa, Chamaecytisus ruthenicus, Prunus spinosa, Malus sylvestris, Pyrus piraster, Quercus robur, Acer platanoides, Acer tataricum, Ulmus glabra, Ulmus minus).
In the meadow-steppe phytocenoses under the haymaking regime, it is noted that the species saturation and cenotic diversity in these regimes is greater.
The study of the duration of succession stages in the Streletskaya steppe and other reserves of the forest-steppe zone makes it possible to pay attention to the duration of the stages when plant communities change. In all the reserves under the reserved regime, after 40-50 years in the steppe phytocenoses, the emergence of tree and shrub communities, light forests was observed. At the same time, in some cases this happened through the shrub phase (“Galichya Gora”, “Privolzhskaya forest-steppe”), in others, the simultaneous spread of trees and shrubs (“Streletskaya and Kamennaya steppes”).
The successions of meadow-steppe phytocenoses of the European forest-steppe confirm the need for haymaking to preserve these communities.
2. The cause of changes in phytocenoses in the areas of the Streletskaya steppe is an exogenous anthropogenic factor that has been acting for a long time. Due to the action of an absolutely reserved regime (removal of grazing, haymaking, burning), the influence of endogenous factors is observed (accumulation of rags and bedding, changes in temperature, changes in moisture, shading). Successions in plots with a periodical mowing regime arose as a result of replacing annual mowing with hay rotation, removing grazing, changing the timing and methods of mowing.
3. The direction of succession of meadow-steppe phytocenoses is reflected in the change in species richness. Absolutely protected areas of the Streletskaya steppe almost halved the species saturation (from 110 to 49 plant species per 100 m2 and from 44 to 24 plant species per 1 m2). Species saturation in areas with periodic mowing has not changed since the establishment of the reserve and is 87 plant species per 100 m 2 and 57 plant species per 1 m2.
4. In absolutely protected areas, the process of succession goes in the direction of the disappearance of annuals and juveniles, the loss of perennial rosette plants, the decrease in steppe plants and the increase in meadow and forest plants. Modern phytocenoses in areas with periodic mowing have retained their main species composition. Under the regime of periodic mowing, the abundance of steppe plants also somewhat decreased and the abundance of meadow species increased. The composition of cereals has changed due to the wide distribution of rhizomatous cereals in phytocenoses.
5. While maintaining the indicators of biological productivity of terrestrial phytomass in absolutely protected areas (9 t/ha), the percentage of economic and botanical groups of plants has changed. The weight participation of cereals has now increased to 70%. The biological productivity of terrestrial phytomass in the mown areas was 4.48 t/ha. Changes have taken place in the ratio of the main economic and botanical groups. At present, the participation of grasses (52.5%) prevails over forbs, although the predominance of forbs over grasses was noted during the organization of the reserve.
1) Fescue-forb.
3) Veynikovskaya.
1) Fescue-forb.
3) Forb-coastal rump.
8. Comparison of the succession process of meadow-steppe phytocenoses in the Central Chernozem Reserve with successions in other reserves of the European forest-steppe shows that under an absolutely reserved regime, light forest appears in steppe phytocenoses either through the shrub phase (Galichya Gora, Privolzhskaya forest-steppe), or with the simultaneous spread of trees and shrubs (Streletskaya and Kamennaya steppes). The most optimal for the conservation of steppe phytocenoses is the mode of periodic mowing, which preserves the features of the reference northern forb steppes.
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3. Avanesova A.A. Influence of Anthropogenic Factors on the Species Composition of the Steppe Communities of the Streltsy Area of the Central Chernozem
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4. Avanesova A.A. Changes in the steppe phytocenoses of the CCR under the influence of the anthropogenic factor /A.A. Avanesova// Education in decision environmental issues: materials of the international scientific conference. September 18-21. - Kursk, 2001.- S-91-92.
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9. Avanesova A.A. Experience in the protection of the meadow steppe in the Central Chernozem Reserve. prof. V.V. Alekhina /A.A. Avanesova, V.D. Sobakinsky // Kulikovo field. Historical landscape. Nature. Archeology. Story. Volume 1. -Tula, 2003, - S. 169-186.
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11. Avanesova A.A. Morphological features of Peoria tenuifolia (Paeoniceae) on the Streltsy site of the Central Chernozem Biosphere Reserve named after Professor V.V. October 14-18, 2002, - St. Petersburg, 2002, - S. 11-12.
12.Avanesova A. Influence of absolutely reserved mode on steppe phytocenosis of Central-Chernozem reserve / A.Avanesova // Importance of science education in the light of social and economic changes in the central and east European countries: The materials of the IV IOSTE symposium for Central and East European Countries.- Kursk, KSU, 2003.- P. 254-256.
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Avanesova Anna Alexandrovna
Successions of steppe phytocenoses of the European forest-steppe
(on the example of the Central Chernozem Biosphere Reserve named after V.V. Alekhin)
License for publishing activities ID No. 062248 dated 12.11.2001 Signed for printing 13.09.06 Format 60x84/16 Offset paper. Offset printing Circulation 100 copies. Order No. 12
Publishing House of the Kursk State University 305000, Kursk, st. Radishcheva, 33
Printed in the laboratory of information and methodological support of KSU
Chapter 1
1.1. Characteristics of virgin upland steppe phytocenoses of the Streletskaya steppe.
1.2. Study of the flora of the Streletskaya steppe.
1.3. Studies of productivity and species saturation of steppe phytocenoses.
1.4. Successional changes in steppe phytocenoses under different protection regimes.
1.5. The study of biomorphological features of plants of the Streletskaya steppe.
Chapter 2. Area, materials and research methods.
Chapter 3
3.1. Species composition and productivity in absolutely protected and adjacent mowing phytocenoses in the western part of the Streletskaya steppe.
3.2. Species composition and productivity in absolutely protected and adjacent mowing phytocenoses in the central part of the Streletskaya steppe.
Chapter 4
4.1. The direction of vegetation change in areas with an absolutely protected regime. 4.2. Expansion of trees and shrubs on the Streletskaya steppe.
4.3. Stages of succession of steppe phytocenoses in areas with an absolutely reserved regime.
Chapter 5. Successions of steppe phytocenoses under the influence of haymaking.
5.2. Stages of succession of steppe phytocenoses in areas with periodic mowing.
Chapter 6
Introduction Thesis in biology, on the topic "Successions of steppe phytocenoses of the European forest-steppe"
Relevance of the topic. The study of succession is not only of deep theoretical interest, but also of practical importance. It is also important to study the successions of the vegetation cover, which occur as a result of the changes that a person is currently producing with his economic activity (Kamyshev, 1964).
Central Chernozem State Biosphere Reserve named after V.I. prof. V.V. Alekhin (Kursk region) is the basis for studying changes in virgin steppe phytocenoses under the influence of certain protection regimes. The meadow steppes of the Central Chernozem Reserve are unique systems that have almost disappeared throughout Eurasia due to plowing and human economic activity (Alekhin, 1936). The processing of biological material accumulated in the Central Black Earth State Biosphere Reserve is of great scientific importance in connection with the compliance with the UNESCO program "Man and the Biosphere" (MAB) and the Convention on the Biological Diversity of Wildlife Species adopted in 1992.
The successions of meadow-steppe phytocenoses under various protection regimes have not been sufficiently studied. It is necessary to know the direction of these changes in order to correct the processes occurring in them and optimize the steppe nature management (Lavrenko, 1959).
A large number of studies of vegetation cover have been carried out in the forest-steppe zone. This is due to its economic importance, as well as the fact that the study of steppe vegetation leads to the identification of phytocenological and floristic problems (Kultiasov, 1981). One of the problems is the "steppe issue", which reveals the reasons for the lack of forests of the steppes (Komarov, 1951).
Recently, great changes have been observed in steppe phytocenoses and the spread of trees and shrubs in areas with a reserved regime.
Data on the state of meadow-steppe phytocenoses, obtained in different years, are the material for identifying succession stages.
Purpose and objectives of the study. The purpose of this study is to study the succession of steppe phytocenoses of the European forest-steppe (on the example of the Central Chernozem Biosphere Reserve named after V.V. Alekhin).
To achieve this goal, the following tasks were set:
1. To identify the features of the current state of steppe phytocenoses with a periodic mowing regime (RPK) and an absolutely protected regime (RAS), paying special attention to the distribution of trees and shrubs.
2. Determine the directions of succession of steppe phytocenoses with RAS and RPC.
3. Identify stages of succession of steppe phytocenoses during RAS and RPK.
4. Compare ongoing successions of the vegetation cover of the Central Chernozem Reserve with changes in similar meadow-steppe communities of the European forest-steppe.
Scientific novelty of the work. For the first time, the stage of light forests in phytocenoses with an absolutely reserved regime was described, and the stages of vegetation succession were identified under an absolutely reserved regime of protection and a regime of periodic mowing. The dynamics in the expansion of trees and shrubs in all unmowed areas of the Streltsy section of the Central Chernozem Reserve has been established. Meadow formation was established in areas with the PKK regime and afforestation in areas with RAS in the steppe phytocenoses of the Streletskaya steppe.
Basic provisions for defense.
1. Analysis of changes in the species composition, species saturation, productivity and moisture indices according to L.G. Ramensky indicates the meadowing of the steppe communities of the Streletskaya steppe during the period of the reserve's existence.
2. In phytocenoses with PKK, the moisture index changed (from 51.9 to 52.7 on the moisture scale of L.G. Ramensky). Currently, phytocenoses with PKK are located on the border between meadow steppes and dry, fresh meadows. Phytocenoses with RAS are characterized as fresh meadows (level 56 on the moisture scale of L.G. Ramensky) with the expansion of forest species, trees and shrubs.
3. Evaluation of the succession of phytocenoses with RAS and RPC in the Streletskaya steppe and other protected areas showed that the identified changes are typical for protected meadow-steppe communities throughout the European forest-steppe.
Practical significance. The materials presented in the dissertation summarize the accumulated experience in the protection of meadow steppes in the Central Chernozem Reserve, allow you to choose rational methods for the protection of meadow steppes, confirm the assessment of the absolutely reserved regime of N.S. Kamyshev (1965), as a regime leading to the loss of meadow-steppe phytocenoses and the emergence of light forests.
Approbation of work. The main provisions of the dissertation were presented and discussed at scientific conferences "Phytocenoses of the northern forest-steppe and their protection" (Kursk, 2001), "Flora and vegetation of the Central Chernozem region" (Kursk, 2002,2003,2004), "Anthropogenic impact on flora and vegetation" (Lipetsk , 2001), "Studying and protecting the nature of the forest-steppe" (Kursk, 2002), II International Conference on anatomy and morphology of plants (St. Petersburg, 2002), "Historical landscape. Nature. Archeology. History "(Tula-Kulikovo field, 2002).
Workload. The dissertation of 166 pages consists of an introduction, 6 chapters, a conclusion, conclusions and 29 pages of an appendix; contains 120 pages of main text, 22 figures, 9 tables. The list of references includes 217 titles, 9 of which are in foreign languages.
Conclusion Dissertation on the topic "Botany", Avanesova, Anna Aleksandrovna
1. Meadow-steppe phytocenoses of the Streletskaya steppe of the Central Chernozem Reserve have been studied since the beginning of the century; at different stages of the study, complete lists of phytocenoses were compiled, which makes it possible to trace the direction of successional processes and identify the stages of succession in absolutely protected areas and periodically mowed phytocenoses of the Streletskaya steppe.
2. The cause of changes in phycocenoses in the areas of the Streletskaya steppe is an exogenous anthropogenic factor that has been acting for a long time. Due to the action of an absolutely reserved regime (removal of grazing, haymaking, burning), the influence of endogenous factors is observed (accumulation of rags and bedding, changes in temperature, changes in moisture, shading). Successions in plots with a periodical mowing regime arose as a result of replacing annual mowing with hay rotation, removing grazing, changing the timing and methods of mowing.
3. The direction of succession of meadow-steppe phytocenoses is reflected in the change in species richness. Absolutely protected areas of the Streltsy steppe almost halved the species saturation (from 110 to 49 plant species per 100 m and from 44 to 24 plant species per 1 m). Species saturation in areas with periodic mowing has not changed since the establishment of the reserve is 87 plant species per 100 m and 57 plant species per 1 m2.
4. In absolutely protected areas, the process of succession goes in the direction of the disappearance of annuals and juveniles, the loss of perennial rosette plants, the decrease in steppe plants and the increase in meadow and forest plants. Modern phytocenoses in areas with periodic mowing have retained their main species composition. Under the regime of periodic mowing, the abundance of steppe plants also slightly decreased and meadow plants increased. The composition of cereals has changed due to the wide distribution of rhizomatous cereals and ryegrass in phytocenoses.
5. While maintaining the indicators of biological productivity of terrestrial phytomass in absolutely protected areas (9 t/ha), the percentage of economic and botanical groups of plants has changed. The weight participation of cereals has now increased to 70%. The biological productivity of terrestrial phytomass in the mown areas was J 4.48 t/ha. Changes have taken place in the ratio of the main economic and botanical groups. At present, the participation of grasses (52.5%) prevails over forbs, although the predominance of forbs over grasses was noted during the organization of the reserve.
6. The study of the succession of meadow-steppe phytocenoses of the Streletskaya steppe showed that currently phytocenoses with an absolutely reserved regime are characterized as fresh meadows (index 56 according to the environmental scale of L.G. Ramensky) and undergo expansion of trees and shrubs (from 5 to 20% of the area unmowed areas). An analysis of the descriptions on ecological scales shows that steppe phytocenoses with periodic mowing have changed the moisture index from 51.9 to 52.7 and are currently on the border between meadow steppes and dry, fresh meadows.
7. For the first time, stages of succession of meadow-steppe phytocenoses were identified and their time limits were determined. In areas with an absolutely reserved regime of the Streletskaya steppe, 4 stages of successions are described:
1) Fescue-forb.
2) Shore-kostretsovo-feather grass.
3) Veynikovskaya.
4) Development of woodlands with a predominance of ryegrass association.
In areas with periodic mowing, the following stages of succession are distinguished:
1) Fescue-forb.
2) Fescue-coastal brome-forb.
3) Forb-shore-streak.
4) Forbs-shore-brim-ryegrass.
8. Comparison of the euccession process of meadow-steppe phytocenoses in the Central Chernozem Reserve with successions in other reserves of the European forest-steppe shows that under an absolutely reserved regime, light forest appears in steppe phytocenoses either through the shrub phase (Galichya Gora, Privolzhskaya forest-steppe), or with the simultaneous spread of trees and shrubs (Streletskaya and Kamennaya steppes). The most optimal for the conservation of steppe phytocenoses is the mode of periodic mowing, which preserves the features of the reference northern forb steppes.
Suggestions for using the results of the study
The materials presented in the dissertation, summarizing many years of experience in the protection of meadow steppes in the Central Chernozem Reserve, provide environmental institutions (reserves, State Committees for Environmental Protection) with information on the direction of vegetation changes, allowing them to choose rational ways to protect meadow steppes and herbaceous communities. Succession data can be used to test the hypotheses of vegetation dynamics.
Conclusion
One of the most important tasks facing the reserves is the study of biocenoses in all relationships and monitoring their dynamics in various environmental conditions. In this regard, observations of successions in the vegetation cover are of particular interest. Productivity and species composition are the main indicators of communities that provide information characterizing biological systems. Based on these indicators, it is possible to determine the position of associations in the hierarchy of these systems, analyze their economic and ecological significance, make florogenetic, ecological and phytocenotic analyzes, and identify the stages of community succession during long-term observations.
The currently available protection regimes in the Streletskaya steppe ensure the safety of its phytocenoses to varying degrees.
The regimes of periodic mowing, in contrast to the reserved regime, preserve the species diversity of phytocenoses of the Streletskaya steppe, which have the characteristic features of northern, meadow, colorfully mixed grass steppes. This mode creates conditions that are closer to those conditions in which the phytocenoses of the Streletskaya steppe were in the historical past.
The protected regime has led to great changes in phytocenoses. They are currently dominated by large quantities of cereals, their abundance and share in the mass of cuttings of phytocenoses is great. The occurrence and abundance of herbs decreased. Trees and shrubs appeared, the crown projection of which reaches 20% in some places.
The reserve regime did not exist before the creation of the reserve, its creation is part of the experiment laid down by V.V. Alekhin. At present, forest elements are expanding in areas with this regime. These areas reflect the phytocenotic relationship between the forest and the steppe. Studies of succession in the Streletsky section of the Central Chernozem Reserve, as well as data on changes in steppe phytocenoses in other reserves of the European forest-steppe, confirm the “cyclic” theory of the relationship between forest and steppe, expressed by G.E. Grosset. Forest and steppe in the forest-steppe belt exist on an equal footing, and between them there is only a temporary replacement of each other in different areas. But the main inhibitory factor in the advance of the forest on the steppe should be considered anthropogenic, and not changes in growth conditions. This point of view was expressed by V.I. Taliev. Keller (1931), Semenova-Tyan-Shanskaya (1966), Smirnova (2000) studies are devoted to this problem.
Experience of V.V. Alekhine on regime measures in the Central Chernozem Reserve is of unique importance and worthy of comprehensive study and coverage, its results should be taken into account when creating new reserves and natural monuments in the forest-steppe zone.
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