Bibliography
Voronov A.G. Geobotany. Proc. Allowance for high fur boots and ped. in-comrade. Ed. 2nd. M.: Higher. school, 1973. 384 p.
Ipatov V.S., Kirikova L.A. Phytocenology: Textbook. St. Petersburg: Publishing House of Leningrad State University, 1997. 316 p.
Stepanovskikh A.S. General ecology: Textbook for universities. M.: UNITI, 2001. 510 p.
Sukachev V.N. Fundamentals of forest typology and biogeocenology. Fav. tr. L.: Nauka, 1972. T. 3. 543 p.
Program and methodology of biogeocenological research / Study of forest biogeocenoses / M.: Nauka, 1974. S. 281-317.
Tsvetkov V.F. Forest biogeocenosis. Arkhangelsk, 2003. 2nd ed. 267 p.
Questions
1. Identification of the boundaries of biogeocenosis.
2. Biogeocenosis-forming role of phytocenosis in the landscape.
3. Definition of the concept "phytocenosis".
4. The main signs of phytocenosis.
5. The minimum size of the area for detecting phytocenosis.
6. On the boundaries of phytocenosis. The concept of the vegetation continuum.
7. Differences between the concepts of "phytocenosis", "association" and "plant community".
To manage biogeocenotic processes, one must know the patterns to which they are subject. These patterns are studied by a number of sciences: meteorology, climatology, geology, soil science, hydrology, various departments of botany and zoology, microbiology, etc. Biogeocenology, on the other hand, synthesizes the results of the listed sciences from a certain angle, focusing on the interactions of the components of biogeocenoses with each other and revealing general patterns that govern these interactions. This field of knowledge studies biogeocenosis as a whole, explores its inherent processes.
1. Identification of the boundaries of biogeocenosis
It is known that only when the exact boundaries of the biogeocenosis are established, it is possible to analyze it. And the more specifically and precisely the biogeocenosis is limited in space, the more objectively it is possible to quantitatively characterize the processes and phenomena occurring in it.
The difficulty of establishing these boundaries accurately is well known, "... drawing boundaries between them is often conditional, to a certain extent subjective in nature ...". V.N. Sukachev wrote about this: "...Different biogeocenoses, of course, have different vertical thicknesses, for example, forest, steppe, desert, etc. However, as a rule, we can assume that the upper limit of the biogeocenosis is located several meters above the vegetation cover, the lower part lies several meters below the soil surface ... "(Fundamentals of forest biogeocenology, 1964: 32.
E. M. Lavrenko (1962) understands the boundaries of biogeocenoses in approximately the same way when he singles out a part of the biosphere, which he called " phytosphere ".
The question arises, what criteria are most accessible when distinguishing biogeocenoses in nature?
1. You need to start with the analysis of the terrain. Although the relief, as you know, is not part of the biogeocenosis components, it is a very important factor in its existence. For the first orientation in forest biogeocenoses during their isolation and differentiation in nature, it can play a very important role. The description of biogeocenosis begins with it.
2. Within a homogeneous relief, the most indicative sign of the homogeneity of a biogeocenosis is the homogeneity of the soil and vegetation cover. Of these two indicators, it is especially suitable for the identification of biogeocenoses homogeneity vegetation cover due to its visibility. Therefore, when distinguishing biogeocenoses in nature, it is advisable to use phytocenosis. The boundaries of each biogeocenosis separately are determined by the boundaries of the phytocenosis.
Visibility is a very important detail, but the main thing is that among the components of the biogeocenosis, it is the phytocenosis that plays the decisive - biogeocenosis-forming role.
2. Biogeocenosis-forming role of phytocenosis in the landscape
Phytocenosis is the main component, a nodal subsystem of biogeocenosis in all respects, in which the main processes of formation and transformation of what is the basis of life on the planet - organic matter take place. It determines the spatial boundaries of the biogeocenosis, its structure and appearance, internal climate, composition, abundance and distribution of animals, microorganisms, features and intensity of the material and energy exchange of the entire biogeocenosis system.
Phytocenoses serve:
1) main receivers and solar energy transformers,
2) the main suppliers of products in the biogeocenosis,
3) their structure objectively reflects all the processes occurring in the biogeocenosis,
4) at the same time, they are easily accessible for study directly in nature,
5) for them, for several decades, effective field research methods and methods of office processing of actual materials have been developed and are being developed.
Based on the foregoing, a detailed study of phytocenosis is obligatory part any biogeocenological research. Vegetation always begins the study of any natural system. In our lectures, the main attention will be paid to phytocenosis and methods of its study. Moreover, many of the patterns inherent in phytocenosis also apply to zoocenosis and microorganisms.
In a general form, the study of phytocenoses is reduced to solving the following major issues:
- Determination of the role of phytocenosis in the accumulation of organic matter and energy and the transformation of matter and energy into common system biogeocenosis (a key issue!).
- Determination of the role of phytocenosis in the dynamics of biogeocenosis.
- Determination of the nature and degree of impact of phytocenosis on other components of biogeocenosis.
- Accordingly, the determination of the influence of other components of the biogeocenosis on the properties, features and efficiency of the “work” of the phytocenosis.
- Determination of the nature and degree of impact of phytocenosis on neighboring biogeocenoses.
- Determination of the form, methods and means of direct and indirect impact on phytocenosis by human economic activity in order to increase the biological productivity of biogeocenosis and enhance its other useful properties.
3. Definition of the concept of "phytocenosis"
The first definition of a plant community was given by G.F. Morozov (1904) for the forest, and then distributed by V.N. Sukachev (1908) to all plant communities. The term "phytocenosis" was applied in 1915 by I.K. Pachosky for "pure thickets" (formed by one plant species), and for all communities - V.N. Sukachev (1917) and G. Gams (Gams, 1918).
Phytocenosis, like biogeocenosis, should be understood as a geographical phenomenon, namely, each terrestrial phytocenosis occupies a certain territory with its own hydrological regime, microrelief, microclimate, soils.
V. N. Sukachev (1956) very successfully defines phytocenosis: "... Phytocenosis, or a plant community, is a collection of plants growing together in a homogeneous area, characterized by a certain composition, structure, composition and relationships of plants both with each other and with environmental conditions. The nature of these relationships is determined, on the one hand, by the vital, otherwise, ecological properties of plants, on the other hand, by the properties of the habitat, i.e., the nature of the climate, soil and the influence of humans and animals ... ".
The structure and state of the phytocenosis well reflects both competition and mutual assistance of plants.
EXAMPLE. Broad-leaved-linden with liana vegetation hazel forb phytocenosis in the middle part of the slope north-north-east. exposure (altitude marks 250-300 m a.s.l.; average slope steepness is 15-20).
The tree stand consists of 2 tiers. It is characterized by a very high density of crowns - 0.97. The trees are closed by crowns and twilight reigns under their canopy. The first tier is formed by Mongolian oak, Amur and Manchurian lindens, Manchurian walnut, single old trees of black birch and small-leaved maple.
Most of the lindens are represented by slender full-wood trees with smooth trunks. Average taxation indices of trees of the 1st tier: Dm 18-20 cm, Hm - 17-18 m. In the well-defined second tier, linden predominates - mostly younger than in the upper tier, and stunted trees, small-leaved maple. Insignificant admixture of mountain elm, Amur maakia, heart-leaved hornbeam; dimorphant, false-sibold maple and small-carp are rare. In addition, the crackling, Maksimovich's spindle tree "penetrate" into it, the bulk of the individuals of which are concentrated in the undergrowth.
The further existence of any forest is ensured by the renewal of the species of the parent stand. Undergrowth in the amount of 8.6 thousand specimens/ha is represented by all types of forest stand. Small-leaved maple dominates in its species composition, in self-sowing there are single specimens of a dark coniferous species - whole-leaved fir (Abies holophylla). ).
The undergrowth is dense, it is dominated by Manchurian hazel, mock orange, eleutherococcus are common, currant Maksimovich is less common in large bushes, early-flowering honeysuckle, single large-winged and few-flowered euonymus, green maple. Rarely, viburnum Bureinskaya grows in groups and, as a rule, with it - single shoots of Amur barberry.
Due to the high density of trees and shrubs, the herbage is sparse. In addition to the spring forest poppy, it is dominated by tuberous cornflower, sedges: Ussuriyskaya, returned, long-nosed, Bunge's chickweed, Daurian bedstraw, ferns. As in the forest stand, tiers can be distinguished in the herbage. The upper one, up to 1 m high, is formed by large-grass species growing everywhere: mountain peony, black cohosh, Daurian and Amur angelica, pointed raven, red-flowered sapling, two-row lily; single Volzhanka Asiatic and powerful stalk. Sometimes small densely covered micro-groups form grasses of medium size (forbs), up to 0.5 m high - hairy smilacin, wintering horsetail, deaf nettle, lily of the valley, and small herbs, not more than 0.25 m high: Ussuri skullcap, Franchet buttercup, two-leaved mullet, rooting trigonotis, musk adoxa, meringia lateral, Colin's violets, Siebold's hoof, jeffersonia dubious, different types corydalis.
In addition to plants forming tiers, in the described phytocenosis, so-called extra-tiered plants can also be distinguished, for example, creepers of actinidia, magnolia vine, and grapes.
All types of herbs can be divided into groups according to seasonal development (some are spring ephemeroids (anemones, corydalis, adonis, lloydia, etc.), go through a development cycle within a month and are at rest in June. In others (double row lily, sparkling lychnis, powerful stalk, etc.) the culmination of development occurs in July, the third (plectranthus, desmodium, saussurea, aconites) bloom and remain green in September), by origin (taiga forests, small-leaved, nemoral, meadow, etc.), by abundance (some of them are found in such a significant amount that they form a continuous cover, others are rare, and others are single).
Thus, six above-ground tiers can be distinguished in this forest: two arboreal, one shrub (with undergrowth) and three herbaceous.
Having dug a trench in such a forest, one can also observe underground layering (though less pronounced than aboveground): the roots and rhizomes of grasses are located in shallower soil horizons, the roots of shrubs and trees - in deeper ones. Due to the underground layering, plants use different layers of soil to obtain moisture and nutrients.
Thus, phytocenosis characterized by:
1. a certain species composition of the plants that form it,
2. a certain abundance of them,
3. certain structure and
4. confinement to a certain habitat.
4. The main signs of phytocenosis
Significant signs of phytocenosis - phytocenotic relations (relationships between plants) and the presence phytocenotic environment.
The creation of a phytoenvironment is the first sign of a phytocenosis in time of appearance, because. the influence of plant organisms on the environment can already be where there is no influence of plants on each other. PHYTOCENOTIC ENVIRONMENT begins to form even at a time when individual plants that have appeared on a previously devoid or not having a coherent vegetation cover of the territory grow scattered, without forming a continuous cover.
Already at the first stages of vegetation development, microclimate conditions change, with dead plants some chemicals are introduced into the soil or soil, while others are extracted by living plants, the nature of the microrelief changes (for example, trails of dusty and sandy particles are formed near the stems of plants), in a word, there is transformation of the environment by plants. In the future, through the interaction of plants, the phytocenosis changes the environment more and more and creates its own phytoenvironment. At the same time, the environmental conditions in different parts of the phytocenosis (on the surface of the soil, on the trunks and crowns of trees, at different heights above the soil surface, etc.) are not the same.
The presence of phytocenotic relationships is the most significant feature of phytocenosis, but INTERACTIONS BETWEEN PLANTS begins somewhat later than the impact of plants on their habitat. It can take place only at a certain density of vegetation cover. However, it is very difficult to notice this moment when the interaction between plants begins, since it does not always involve direct contact between organisms.
Therefore, various stages of development of the vegetation cover should be attributed to phytocenoses, except for the very first moments of plant settlement in a territory devoid of vegetation.
Another issue is the degree of expression of the phytocenotic environment and the degree of expression of relations between plants in phytocenoses. In some deserts, in the polar tundra, the vegetation cover is so sparse that it cannot be considered formed from phytocenoses. With such a sparse cover, it is very difficult to determine the degree of influence of plant root systems on each other and it is almost impossible, using existing research methods, to determine the degree of influence on the environment and on other plant organisms of microscopic plants - algae and bacteria, which can be very significant. Under such environmental conditions, it is likely that each section of a long-term and, therefore, adapted to the environment vegetation cover should be divided into phytocenoses.
Since phytocenosis is not any collection of plant species, but only a completely natural combination, formed as a result of a long historical process and being in connection with the external conditions of existence, then V. V. Alekhin and other representatives of the Moscow phytocenological school believe that “the ability to restore” or “the ability to relative restoration” is put forward as one of the mandatory features of the community (Prozorovsky. 1956).
From this point of view, cultivated vegetation, plant groups that settle in areas devoid of vegetation, as well as all those natural combinations of plants that are not restored after their disturbance or destruction, cannot be considered phytocenoses.
So, it would be impossible to consider as phytocenoses not only secondary birch or aspen forests that appear on the site of the primary ones after they are cut down, but also the primary northern temperate forests that live in areas with shallow groundwater, since these forests do not regenerate after cutting down or burning out. , and the areas of cutting areas and burnt areas become swampy. Shrub communities (forest forests, sea buckthorn forests), which have replaced tall-stemmed phytocenoses (oak forests, cedar-broad-leaved forests), cannot be considered either.
It is unlikely that such a point of view can be recognized as correct. Indeed, in cultural communities, and in primary and secondary forests, and in pioneer plant groups (perhaps, with the exception of the very initial stages of their existence), there are those signs that constitute the essential features of a phytocenosis: the creation of a phytoenvironment and the presence of phytocenotic relationships.
There are two types of relationships between plants in the existing phytocenosis:
1) competition with each other because of the means of subsistence or the struggle for existence in the broad sense, as Charles Darwin understood it. On the one hand, this weakens the plants, but on the other hand, it forms the basis natural selection- the most important factor in speciation and, consequently, the process of evolution.
EXAMPLE. Differentiation and self-thinning of the stand with age or deterioration of growing conditions - the strongest survive. From tens of thousands of seedlings and self-sowing, less than 1% remains by the age of ripeness.
In the process of natural selection, the composition of the phytocenosis included such species that are interconnected with each other or depend on each other. It includes not only flowering, gymnosperms, ferns, club mosses, horsetails, mosses, but also lower plant organisms: fungi, algae, bacteria, lichens.
What has been said about the role of natural selection in the formation of communities is true only for undisturbed, established natural phytocenoses. In phytocenoses developing on areas previously devoid of vegetation, at the first stages of development, there is no direct influence of individual plants on each other, and therefore the relationship between individual species has not yet been expressed.
2) mutual assistance- plants in a phytocenosis have a beneficial effect on each other: shade-loving herbs live under the canopy of trees that cannot grow or grow poorly in open places; tree trunks and branches of shrubs serve as a mechanical support for vines, on which, in turn, epiphytes not associated with the soil settle.
5. The minimum size of the area for detecting phytocenosis
How to determine the minimum size of the area on which phytocenosis can be distinguished?
Obviously, the smallest territory for detecting a phytocenosis should be of such size that all the signs of the phytocenosis itself (species composition, structure, etc.) can appear, as well as all the main features of the soil, microclimate, surface microrelief, in a word, features of the phytoenvironment.
It goes without saying that for different phytocenoses the size of this smallest territory is not the same: the simpler the structure of the phytocenosis, the less its impact on the habitat, the smaller the size of the territory.
In the temperate zone, they are smaller for grasslands compared to forests. For temperate forests, they are smaller compared to tropical forests.
6. On the boundaries of phytocenosis. Concept of land cover continuum
There can be sharp boundaries between phytocenoses, but more often the transitions are gradual, imperceptible. This causes difficulties in isolating phytocenoses. The gradual transition from one type of phytocenosis to another is a consequence of a gradual change in the environmental conditions of the habitat. If the changing values of any factor (for example, moisture conditions, salinity, etc.) are plotted on a graph, then combinations of plant species that gradually replace each other will also correspond to them. Based on this, L.G. Ramensky developed the doctrine of vegetation continuity(Moscow school of geobotany), or, as it is often called, the doctrine of continuum. As L.G. points out Ramensky (1910, 1925, 1937, 1938), "... the ability of plants to form various combinations is downright inexhaustible ...", i.e. the number of associations is unlimited.
Many scientists, following this doctrine, do not recognize the reality of the existence of phytocenoses. According to L.G. Ramensky, to some extent conditional and depends not only on the relationship of this phytocenosis with neighboring ones, but also on the "target setting, work." In other words, the boundaries between two phytocenoses may be absent and may be drawn differently in different cases or by different researchers.
At present, the doctrine of the continuity of vegetation cover has become widespread, especially in the works of American scientists: H. Gleason (Gleason, 1939), D. Curtis (Curtis, 1955, 1958), R. Whittaker (Whittaker, 1953, 1956, 1960) , F. Igler (Egler, 1951, 1954), Polish researcher V. Matuszkiewicz (Matuszkievicz, 1948). The main idea of this doctrine is the impossibility of putting all the variety of combinations of plants with many different transitional groupings into a limited number of associations. Proponents of this concept, as a rule, consider phytocenoses (associations, plant communities) to be conditional, abstract categories that do not exist in nature, although, as some of them believe, they are necessary from a practical or theoretical point of view.
In this regard, the question of the existence of sharp or gradual boundaries between phytocenoses becomes essential. L.G. Ramensky, T.A. Rabotnov (1967) and other supporters of the continuum theory believe that the rule is gradual, unclear boundaries, and the exception is sharp ones.
Supporters of another doctrine (Leningrad school) - about the discreteness of the vegetation cover, for example, V.N. Sukachev, G. Durie, in contrast to the view of L.G. Ramensky believe that, as a rule, the boundaries between associations are sharp, but sometimes smooth transitions are possible.
In fact, both theories have the right to exist. The nature of the boundaries between phytocenoses reflects the influence of edificatory plants on the environment. Smooth transitions from one cenosis to another are more often observed where the influence of edificators does not change the environment so much, for example, in meadows, and abrupt transitions where one powerful edificator is replaced by another (for example, at the boundaries of forest plots formed by various tree species).
As pointed out by B.A. Bykov (1957), edificators by their influence largely determine the sharpness of the boundaries even with a gradual change in habitat conditions. Therefore, "... the boundaries of plant associations are indisputably outlined in nature more sharply than the boundaries of habitats ..." (Nitsenko, 1948).
7. Differences between the concepts of "phytocenosis", "association" and "plant community"
Between the terms "phytocenosis" and "plant community" many scientists for a long time put an equal sign, using them as synonyms. V.N. Sukachev repeatedly pointed out that the term "phytocenosis" (plant community) can be applied both to specific areas of vegetation cover and to designate taxonomic units of various ranks: associations, formations, types of vegetation, etc.
Recently (Voronov, 1973), the term "phytocenosis" has been applied only to specific areas of the vegetation cover that correspond to the above definition by V.N. Sukachev ..
Association called the typological unit of phytocenoses. The same meaning is put into this term by S.M. Razumovsky (1981). In his interpretation, an association, an elementary unit of vegetation, unites areas with the same species of each tier and the same succession trend. Drawing an analogy, we can say that phytocenosis and association relate to each other in the same way as a specific plant and the species of this plant.
EXAMPLE. Driving along the Vladivostok-Ussuriysk highway, we can say that almost all forest vegetation on this stretch is represented by secondary oak forests. The areas of these forests are very similar to each other. The stand is oak with a single admixture of Dahurian birch. The undergrowth is fragmentary, represented by hazel and lespedecia. The ground cover contains forbs and sedges. Nevertheless, despite the very high similarity, it is impossible to find even two sites with exactly the same phytocenoses.
In other words, all these similar phytocenoses are of the same type, and together they represent one association, or one type of forest - sedge-forb oak forest.
Association is the first step in the system of taxonomic units of vegetation cover of various ranks. The following steps are: group of associations, formation, group of formations, class of formations, type of vegetation; There are also intermediate categories.
Such an abundance of hierarchical levels caused the need for a special term that could be applied to a taxonomic category of any rank. G. Duriez (Du-Rietz, 1936) suggested "plant community" as such a term. This concept is broader than "phytocenosis". It does not have a specific volume. So, V.V. Alekhine (1950) writes that "... plant community... these are plant associations of both a larger and a smaller volume, both a forest in general and a coniferous forest, and subdivisions of the latter, as well as this particular section of it ... ".
Thus, a plant community refers to any plant "collective" where plants are in certain relationships, without indicating its dimension and nature of organization. In this case, the plant community can be considered as a phenomenon of joint social life of plants. In such a broad sense, this term was used by V.V. Alekhin (1935, 1950, 1951)
Phytocenoses belonging to the same association are scattered. Usually they border on phytocenoses belonging to other associations. On the ground, combinations of phytocenoses are usually found, often they are quite noticeably different from each other. This is due to differences in other natural components, and as a result, different biogeocenoses, or so-called natural territorial complexes, are formed. It is with such complexes that one usually has to deal with when using plant resources.
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, Kursk State University
Supervisor:
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 Chernozem 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 forest 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 to protect meadow steppes, confirm the assessment of an 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 questions of biomorphological features of plants of the Streltsy steppe are considered (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 trial 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 Microsoft Excel 2002, Biostat programs.
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 the penetration of 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 herbage show that grasses 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 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 successional 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 along the Far field of the Cossack steppe, the steppe territories are also overgrown 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 course of development of vegetation on unmowed plots 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; Dohman, 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 herbs grow (^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.
During 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 preserves species diversity and basic 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); "Privolzhskaya 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" 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 reserves considered, 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, by 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, mowing, 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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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
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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 Black Earth 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 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).
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 forest 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. An assessment 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, mowing, 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 ecological 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. Alekhin 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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1The phenotypic and soil confinement of German elecampane (Inula germanica L.) on the territory of the Kabardino-Balkarian Republic was studied. It has been established that German elecampane occurs in areas of typical steppes (north and north-east of the republic) on carbonate chernozems. The main limiting factor in the distribution of the species is the carbonate content of the soil. The confinement of the species to a certain phytocenotic environment has not been revealed. However, the highest occurrence of the species is confined to habitats with a lower representation of turf and rhizomatous grasses. In the ontogenetic structure of Inula germanica L. cenopopulations, the largest number of individuals (65–70%) falls on middle-aged generative plants. The low proportion and absence of juvenile and immature individuals indicate the weak competitiveness of elecampane germanica. Vegetative and seed reproduction of Inula germanica L. as a result of the growth of turf grasses and steppe shrubs are reduced, resulting in competitive displacement and natural aging of the population. The ability to exist in unfavorable conditions due to ecological specialization and low competitiveness testify to the ecological patentity of the species Inula germanica L. load. The conservation of the species Inula germanica L. will be facilitated by the creation of optimal conditions for vegetative and seed renewal in natural habitats (chopping and incorporation of sod into the soil, loosening the soil); formation of reserve populations in culture; expansion of work on the reintroduction of the species into habitats corresponding to it in terms of biotopic parameters.
elecampane Germanic
phytocenosis
carbonate chernozems
competitiveness
environmental susceptibility.
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German elecampane (Inula germanica L.) is a Euro-West Asian species of the forest-steppe and steppe zone, found in Central Europe, Central Asia, Western Siberia and in the Caucasus. It is a typical representative of forb-fescue-feather grass steppes on the southern differences of ordinary chernozems and on southern chernozems (Azov-Black Sea steppes, Donetsk-Middle Don steppes). It grows in the steppe, forest-steppe areas, on fescue and grass-forb-feather grass, feather grass and forb-grass steppes, on forest-steppe and steppe slopes, along edges, between shrubs and clearings, along carbonate outcrops between shrubs, without forming numerous populations. Elecampane is a herbaceous long-rhizome perennial. The life form according to Raunkier is a hemicryptophyte. In relation to the temperature and water regime, this species is a steppe euxerophyte, and in relation to calcium, it is an obligate calciophil.
Flavonoids, tannins, essential oil were found in the grass and leaves of elecampane germanica. In folk medicine, Germanic elecampane is used for inflammatory diseases of the oral mucosa, lymphatic system; in veterinary practice it is recommended for diseases of the stomach and intestines, bronchitis and bronchopneumonia. Elecampane Germanic is a honey and ornamental plant. Thus, Inula germanica L. can be classified as a valuable multi-purpose herbaceous plant species.
Unfortunately, as a result of the expansion of arable land, the area of typical steppe plots in the North Caucasus is annually decreasing, which requires the adoption of measures to preserve them, at least in the conditions of specially protected areas.
In connection with the foregoing, it is important to study the distribution area and ecological and phytocenotic characteristics of Germanic elecampane in steppe phytocenoses. The object of the study was the cenopopulations of Inula germanica L. in the steppe phytocenoses of the Kabardino-Balkarian Republic, located in the Central part of the North Caucasus.
The purpose of the study is to study the phytocenotic and soil confinement of Inula germanica L. on the territory of the Kabardino-Balkarian Republic.
Research methods
Geobotanical descriptions were carried out during the flowering period of German elecampane (July-August) in the course of field studies on test plots with an area of 100 m2 in the places of soil sampling (Prokhladnensky district - the vicinity of the village of Altud, the village of Chernigovskoye, Tersky district - the vicinity of .p. Planovskoe, s.p. Tambovskoe, s.p. New Khamidie, s.p. Krasnoarmeyskoye).
The abundance of species in phytocenoses was assessed using the Brown-Blanque scale, and the similarity in species composition was assessed using the Jaccard coefficient KJ. The ontogenetic structure of cenopopulations was determined as the ratio of individuals of different ontogenetic states in them. The type of cenopopulation was determined according to the delta-omega classification.
Soil sampling was carried out using the envelope method from the upper layer (0–20 cm). Agrochemical analysis of soils included the determination of hygroscopic moisture, humus (according to Tyurin), pHwater, mobile phosphorus (according to Machigin), exchangeable potassium (according to Machigin), calcium (by effervescence from a 10% HCl solution), cation exchange capacity (complexometric determination of magnesium express method TsINAO).
Results and discussion
The steppe zone is located in the northeastern part of Kabardino-Balkaria (200-600 m above sea level) on an inclined, slightly undulating Kabardian plain. Due to the active use in agricultural production, steppe phytocenoses have been preserved here in small areas.
The climate of the steppe zone is temperate continental warm, in some areas it is dry hot with a pronounced period of summer desiccation. The average annual air temperature is +11.6 ºС, and the sum of temperatures for the period of active vegetation is 3000-3600 ºС. The average annual rainfall is 533 mm, in summer - 201 mm. Winters are mild with an average temperature of minus 1 ºС and frequent thaws.
The soils of the study area are southern calcareous chernozems (village Altud, vil. Chernigovskoye), southern carbonate residual-meadow chernozems (village Novoe Khamidie), ordinary calcareous chernozems (village Tambovskoye, vil. Krasnoarmeiskoye, s.p. Planovskoye) - low-humus, medium loamy. Soil reaction is weakly alkaline, due to the high carbonate content of the parent rock. Soil-forming rocks are characterized by a high absorption capacity - 25-45 mmol (equiv.) / 100 g of soil. The content of mobile phosphorus is average and increased (village settlement Chernigovskoye, village village Tambovskoye, village village Krasnoarmeiskoye), exchangeable potassium - average (village village Altud), increased (village village Tambovskoe village Planovskoye settlement of Novoe Khamidie), high (village of Krasnoarmeiskoye), calcium carbonate - high (village of Altud, settlement of new Khamidie) and medium (Table 1).
Table 1
Agrochemical analysis of soils in the study areas
|
Humidity hygroscopic, % |
Absorption capacity, mmol (equiv.) / 100 g |
Average content |
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|
Р205, mg/kg |
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|
No. 1 - with. p. Altud |
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No. 2 - p. n. Chernihiv |
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No. 3 - p. Tambovskoe settlement |
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No. 4 - p. Planovskoye |
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No. 5 - p. New Hamidiye |
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No. 6 - p. p. Krasnoarmeiskoye |
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The species composition of the studied phytocenoses differs significantly, as evidenced by the KJ values, which are in the range of 0.10-0.23 (Tables 2-3).
table 2
The number of common species (c) and the number of the total species list (d) for phytocenoses of the foothill steppes of the Kabardino-Balkarian Republic
|
Indicators and district numbers |
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Note: the number of species in each phytocenosis is marked diagonally.
Table 3
Jaccard similarity coefficients calculated for phytocenoses of foothill steppes of the Kabardino-Balkarian Republic
|
Indicators and district numbers |
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In plant communities of steppe phytocenoses, meadow-steppe (creeping wheatgrass, sickle-shaped alfalfa, narrow-leaved peas, common agrimony, etc.), steppe (common bearded vulture, pig fingered, plowed and field clover, Austrian flax, etc.), meadow (shoot-forming bent grass) , common reed, multicolored briar, meadow clover, St. The wide representation of ruderal species will be explained by the influence of anthropogenically created agrophytocenoses.
Phytocenosis s.p. Altud herb-grass. The subdominants are common chicory, wild carrot, steppe sage, St. among the cereals dominated by pig fingered, Valisian fescue, gray foxtail, comb-footed, feather grass, awnless and field fire. Legumes are represented by clovers (strawberry, meadow, creeping), alfalfa (yellow and hop-like), Caucasian bird's-foot. In the phytocenosis there are many individuals of Germanic elecampane, but the coverage is not large.
Phytocenosis in the vicinity of the village Chernihiv cereal-forb. The role of subdominants is played by the common bearded vulture, field brome, green foxtail, comb-footed, narrow-leaved bluegrass, field brome, creeping wheatgrass. Legumes are represented by clovers (meadow, field, plowed), crescent alfalfa, multi-colored elm, and herbs - sprawling cornflower, common yarrow, common chicory, prostrate woodruff, common bruise. In the phytocenosis, Germanic elecampane is rare and has a small projective cover.
In the grass-forb community, s.p. Tambovskoye is dominated by field bonfire, pig finger, couch grass, narrow-leaved and common bluegrass, green foxtail. Legumes are represented by crescent alfalfa, meadow clover, multi-colored elm. Multi-species herbs: common wormwood, steppe euphorbia, green strawberry, tall sorrel, horse sorrel, tall mullein, common yarrow, hooked thistle, annual falacroloma. The number of individuals of Germanic elecampane is large, but the coverage is insignificant.
In the forb-couch grass-pig phytocenosis (environment of the village of Novoe Khamidie), the subdominants of the community are creeping couch grass and pig fingered couch grass. Multi-species forbs: common yarrow, prostrate woodruff, shaggy woolly, Palestinian pterotheca. Of the legumes, meadow and field clover, medicinal sweet clover, and narrow-leaved peas were noted. In the phytocenosis, Germanic elecampane is rare and has a small projective cover.
Phytocenoses in the vicinity of s.p. Planovskoye and s.p. Krasnoarmeyskoe grass-forb with dominance of couch grass and narrow-leaved bluegrass. Among the legumes, meadow clover, sickle-shaped alfalfa, multi-colored elm, and medicinal sweet clover dominate. Herbs are represented by St. John's wort, common chicory, field bindweed, green strawberry, steppe euphorbia, real bedstraw, common yarrow, purple mullein, etc. German elecampane has a slight coverage, but individuals are sparse.
In the steppe plant communities of all study areas, xeromesophytes predominate. The largest proportion of typical xerophytes was noted in the vicinity of the settlement. New Khamidie, Chernihiv and Altud, and mesophytes - in the village. Tambovskoe and s.p. Planovskoye (figure).
Distribution of herbaceous plant species in relation to moisture, %: K - xerophytes, KM - xeromesophytes, MK - mesoxerophytes, M - mesophytes
Based on the data of agrochemical analysis of the soil and the occurrence of elecampane in various steppe phytocenoses, it can be argued that the main limiting factor in the distribution of the species is the carbonate content of the soil. Germanic elecampane, as a stenotopic species, has a narrow ecological amplitude, which makes it possible to use it in haloindication (carbonate salinization of soils). The confinement of the species to a certain phytocenotic environment has not been established. However, the highest occurrence of the species is confined to habitats with a lower representation of turf and rhizomatous grasses.
In the studied phytocenoses, elecampane rarely forms clumps and is mainly represented by solitary sparse individuals. In the ontogenetic structure of Inula germanica L. cenopopulations, the largest number of individuals (65-70%) falls on middle-aged generative plants. The studied cenopopulations are normal, full-membered (areas No. 1,3,4,6) and incomplete (No. 2, No. 5). According to the "delta-omega" classification, cenopopulations of area No. 1 are classified as maturing, Nos. 3,4,6 - as mature, Nos. 2.5 - as aging (Table 4).
Table 4
Integral characteristics of the structure of cenopopulations of Inula germanica L.
The low proportion and, in some cases, the absence of juvenile and immature individuals indicate a weak competitiveness of elecampane germanica. Vegetative and seed propagation of Inula germanica L. as a result of the growth of turf grasses and steppe shrubs are reduced, as a result of which competitive displacement and natural aging of the population are observed. The ability to exist in unfavorable conditions due to ecological specialization (on saline, dry or stony substrates) and low competitiveness testify to the ecological patience (stress tolerance) of the species Inula germanica L.
German elecampane (Inula germanica L.) on the territory of the Kabardino-Balkarian Republic has a narrow distribution area - areas of typical steppes on carbonate chernozems in the north and northeast of the republic. The species is confined to a greater extent to forb-grass phytocenoses with a low proportion of turf grasses.
The limiting factors for the resumption of elecampane cenopopulations in the steppe zone of the Kabardino-Balkarian Republic are pasture degradation of vegetation cover, erosion processes on slopes, destruction of habitats (plowing land), grazing, and increased recreational pressure. Therefore, Inula germanica L. is preserved only in slightly disturbed communities.
The conservation of the species Inula germanica L. will be facilitated by the creation of optimal conditions for vegetative and seed renewal in natural habitats (chopping and incorporation of sod into the soil, loosening the soil); formation of reserve populations in culture; expansion of work on the reintroduction of the species into habitats corresponding to it in terms of biotopic parameters.
Bibliographic link
Tamakhina A.Ya., Gadieva A.A., Lokyaeva Zh.R. ECOLOGICAL AND PHYTOCENOTIC CHARACTERISTICS OF GERMAN NECKEL (INULA GERMANICA L.) IN STEPPE PHYTOCENOSES OF THE KABARDINO-BALKARIAN REPUBLIC // Modern Problems of Science and Education. - 2016. - No. 4.;URL: http://science-education.ru/ru/article/view?id=24980 (date of access: 02/01/2020). We bring to your attention the journals published by the publishing house "Academy of Natural History"
Phytocenosis, which are found in nature, are complex and multi-species. The number of species in them can reach 100 or more. Very rarely there are simple phytocenoses, which consist of one or two species, as a rule, of lower plants. Phytocenoses formed by one species are called colonies by botanists. The floristic composition determines the features and appearance of the phytocenosis. The study of the floristic composition is the basis of phytocenotic research.
The number of species included in the phytocenosis is called species floristic richness.
Species floristic richness is full list plant species that form a phytocenosis.
The number of species in a given phytocenosis that grow per unit area (1 m2 or 100 m2), is called the species saturation of the phytocenosis.
Phytocenosis, which belong to different associations, differ very sharply in species richness. So, in conditions of strong salinity or significant dryness of the soil, phytocenosis can grow, consisting of one species of higher plants. they are called pure thickets. For example, pure thickets of sarsazan (Halocnemum strobilaceum) on salty surfaces, or pure mangroves (Rizophora mangle) in the tidal belt of the seas and oceans. Such phytocenoses do not belong to colonies, because next to one species of higher plants, many species of lower plants are included in their composition. Tropical forests are an example of phytocenoses with an increased number of species of higher plants. For example, in the forests of Sri Lanka alone there are more than 1,500 tree species, and in the forests of the Amazon basin - about 2,500. In addition to species diversity, tropical forests are also characterized by a high saturation of species. Thus, in the tropical forests of West Africa, up to 100 species of higher plants can grow on an area of 100 m2, not counting epiphytes. Sometimes, in unfavorable, at first glance, conditions, phytocenoses are formed that are very diverse in species composition. Studies carried out in the ephemeral desert near Tashkent found more than 5,000 plant specimens belonging to 47 species per 1 m2 of area. The species composition and, especially, the species saturation of higher plants in the phytocenoses of Ukraine is small. For example, in dark spruce and beech phytocenoses on an area of 100 m2, no more than 10 species of flowering plants can be found, and up to 100 species in the meadows of the steppe zone.
When studying phytocenosis, as a rule, all types of plants are included in the list of plants, but with a single examination, the species composition cannot be determined. Some species have a very short development period, and spend most of the year as seeds or underground organs. But the more fully the species composition of the phytocenosis is studied, the more data we get to analyze the relationships between plants, to determine the state of the group itself and abiotic factors. So, for example, the presence in the meadows of Bilousu compressed or yarn (Nardus stricta) indicates soil degradation, and cinquefoil erect (Potentilla erecta)- to reduce the wealth of the soil. The presence in the phytocenosis of representatives of the hazel genera (Corylus) elder (Sambucus) bird cherry (Padus) indicates the richness of the soil. The presence of certain species may also indicate the direction of change in groupings. If in a spruce forest there is a hoofed hoof and an anemone oak forest in the grass cover, this means that earlier there was an oak forest type phytocenosis in this area.
The species saturation of phytocenoses increases from arctic deserts and tundra to broad-leaved forests and the northern steppe, then again decreases to a minimum in deserts, and again grows in savannahs, laurel-type forests and tropical rainforests. When determining species saturation, one should correctly choose the unit of area on which it is determined. In the northern steppe (where herbaceous plants predominate), this figure will be too high even for 1 m2 - up to 80 species, and in the humid tropical forest, the species saturation per 1 m2 will be too low and will not objectively characterize the plant groups existing there. With an increase in the size of registration areas in the tropics, the species saturation index will increase. Species saturation of phytocenosis is determined by:
By eye assessment in points;
Numerical method after determining the number of individuals of each plant species in the trial plots or in the accounting plots.
An example of an eye scale for assessing species saturation is a fairly common phytocenosis. four-point scale BUT. Drude, which uses special characters:
Soc (socials) - plants close with their upper parts and form a kind of background;
Cop3 (copiosae) - plants are very common;
Cop2 - plants are common;
Cop1 - plants are quite common;
Sp (sparsae) - plants are found in small quantities, occasionally, scattered;
Sol (solitariae) - there are single plants.
Modern researchers most often use the Drude scale, as well as other visual scales, only for route studies, when it is necessary to make at least an approximate description of existing phytocenoses or make a general description of vegetation. Eye scales give a scale-free, very approximate assessment of the species richness of phytocenoses. Numerical methods of direct accounting are more objective. An assessment of the species saturation of a phytocenosis is carried out by counting the number of individuals per unit area, their mass or projective cover. For this purpose, in the most characteristic (this is a rather subjective moment of these methods) phytocenoses, trial plots are laid, on which accounting work is carried out. Sample plots can be placed evenly or by random placement in the entire population of phytocenoses that are being studied.
The species richness of plant communities depends on many factors: the climatic zone, the species richness of the flora of the region, environmental conditions, the degree of human influence, and others. Species richness and species saturation of a phytocenosis depend on the lifetime of a given phytocenosis. The older the phytocenosis, the higher the university, ceteris paribus, will be the number of species included in its composition.
Species that are part of phytocenoses can be united in groups on various grounds:
According to ecological and floristic features;
For business purposes;
According to the systematic composition;
By geographic distribution;
origin and the like.
The simplest groupings - according to economic purpose and geographical distribution. By economic purpose herbaceous plants are most often divided into four groups:
legumes;
Forbs.
Representatives of these groups differ in the peculiarities of the fodder use of plants. The division of plants according to economic purpose depends on the characteristics of economic activity in the region. For example, for the Caspian steppe, the following grouping was proposed:
Dry hodgepodges;
Juicy hodgepodges;
legumes;
Spring ephemera;
Other herbs and dwarf shrubs and shrubs;
Poisonous plants.
This grouping by economic feature is important for the development of steppe phytocenoses.
Species that are part of the phytocenosis and have similar ranges are combined into groups that reflect the geographical elements of the flora.
In the phytocenosis, there may be an element of the steppe flora of Europe - pontic, European broad-leaved forests - immoral, dark coniferous forests - boreal. Such an analysis of the flora allows a better understanding of the origin and development of phytocenoses.
To understand the structure and essence of phytocenosis, it is necessary to know the quantitative participation of species in phytocenosis.
A dominant species is a species that is represented in a grouping by a significant number of specimens.
Assektator - these are species that are not dominant, the number of which in the phytocenosis is insignificant.
Dominant species act, as a rule, as elements of a phytocenosis that form the environment, especially when they are strong edificators. Dominants and assektator often have different requirements for certain environmental conditions but, at the same time, grow on the same area. So, other, shade-tolerant tree species grow in the first tier of tree species, under the canopy of which even more shade-tolerant plants have found their place (Fig. 12.4). Dominant species can be found on all plant layers. So, in a four-tiered forest (scotch pine - brittle buckthorn - blueberries - green mosses) there are four dominants that dominate the corresponding tier. The quantitative participation of a species in a phytocenosis determines its role and the degree of influence on other living organisms and on inanimate nature.
Rice. 12.4. View layout in phytocenosis, which differ in their requirements for environmental conditions
Tundra vegetation. The tundra is distributed mainly in the northern hemisphere. It occupies large spaces on the northern outskirts of the continents of Eurasia, North America, and is also found on the Antarctic islands of the southern hemisphere. In the USSR, the area occupied by tundra is 14.7% of the entire territory of the country.
The ecological conditions of the tundra are extremely peculiar and cause the appearance of a certain adaptability of plants.
In the conditions of the arctic and subarctic climate, a characteristic feature of the tundra is the absence of woody vegetation. Of the existing environmental factors, the thermal and chemical features of tundra soils, which explain the causes of its treelessness, are of the greatest importance. Many hypotheses have been put forward regarding the treelessness of the tundra. One of the main reasons should be considered the phenomenon of “physiological dryness”, which is created in supercooled thawed soil, when the roots of trees cannot “use” soil water due to low temperatures (B.N. Gorodkov). According to some scientists, the environmental conditions for the germination of tree seeds are deteriorating in the tundra due to climate change (V. B. Sochava).
The flora of the tundra is distinguished by a certain originality. Its species composition is poor and includes no more than 500 species of higher plants. In the process of adaptation to the conditions of existence in the tundra, certain phytocenoses were formed. A particularly important role in them is played by green mosses and lichens, as well as perennial plants adapted for development under conditions of a short growing season.
The herbaceous vegetation of the tundra is undersized (5-15 cm), forms many shoots, due to which it often takes on a semi-oval shape in the form of “pillows” (for example, grains, saxifrage plants). Due to the abundance of light in summer time tundra plants have large and bright flowers (polar poppy, forget-me-not, tar, etc.).
Shrub plants with a characteristic woody stem and dull leathery leaves with a wax coating and pubescence (blueberries, cranberries, bearberries, dreadas, etc.) are also common in the tundra.
Of the shrubs in the tundra, only dwarf birch and various types of willows are common. These undersized shrubs have small pubescent leaves, lying trunks, often hidden in the moss cover. Of the coniferous shrubs near the southern border of the tundra, juniper is found, and in Eastern Siberia - elfin cedar.
The tundras distributed on three continents do not remain homogeneous. Within their limits from north to south, one can outline a regular change of certain tundra types(formations).
Arctic tundra is common on the coast of the Northern Arctic Ocean. Its vegetation cover is characterized by sparseness (the coverage area is not more than 60%). Of the shrubs, only dread is found here. Herbaceous species are represented by sedge, cotton grass, polar poppy, etc. The moss cover is formed by polytrichus and green mosses. Lichens are characterized by scale forms. Areas not occupied by vegetation are stony placers, areas complicated by stone polygons and polygons.
The moss-lichen tundra is characterized by a complex vegetation cover. On clay soils, a moss cover of green mosses is developed. Willows, blueberries, and dreadas are represented in the upper tier of this formation; from herbs - sedges, arctic bluegrass.
Lichen associations are common on sandy soils. The following geographical regularity can be outlined in their distribution. In the western Arctic (up to the Yenisei River), in the presence of deep snow cover, reindeer moss tundras are common (lichen - reindeer moss predominates), which are valuable pastures for reindeer herding. In the eastern part of the Taimyr Peninsula, fruticose lichens (allectoria, cetraria) are common in winters with little snow.
Moss-lichen tundras are characterized by significant waterlogging (up to 30%). The swamps are predominantly low-lying moss and sedge with a characteristic hilly relief (Fig. 64).
Shrub tundra - a formation that replaces the moss-lichen tundra to the south. In the western sector of the Arctic
shrubs are represented by dwarf birch (dwarf birch). East of the river Lena is dominated by various types of willows and alders. Wide development of plants-shrubs is characteristic. Very large areas (up to 50%) are occupied by swamps.
From the south, the tundra is limited by the forest tundra. In this subzone, an alternation of light forest and tundra areas is noted. So, in the European and North American forest-tundra, birch and spruce woodlands are common, in Asian - larch. In the forest-tundra, the trees are far apart, their height is no more than 6-8 m, have thin curved trunks. The ground cover is dominated by lichens, green mosses and grasses (Fig. 65).
The forest tundra should be considered as a transition zone to the forest zone. Its border is very tortuous. On flat interfluves, the tundra moves south. On the contrary, along the river valleys and mountain slopes of southern exposure, the forest moves northward. The sparse forest-tundra stand is a consequence of unfavorable climatic conditions associated with low rainfall.
It should be noted that the boundary between tundra and forest does not remain constant and is subject to certain dynamics. Such facts as the overgrowth of the tundra of the Far Northeast and North America with larch forest, the overgrowth of patchy and polygonal areas of the Arctic tundra indicate that the forest boundary tends to shift to the north.
Taiga vegetation.This type of vegetation is widespread in the temperate zone of the northern hemisphere - Eurasia and North America. Within the USSR, the taiga occupies more than 11 million hectares. km 2.
The vegetation of the taiga type is mesophilic and is represented by such life forms as coniferous trees, shrubs, herbs, etc. This type of phytocenosis is distinguished by a complex structure and great diversity. In the future, the characteristics of coniferous forests will be given on the example of the taiga of the USSR.
The classification of forests is carried out taking into account forest-forming species, i.e. those species of trees that predominate in the composition of the forest. On this basis, coniferous forests are divided into dark coniferous(spruce, fir, cedar) and light coniferous(pine, larch).
The distribution of coniferous forests is largely determined by the area of distribution of their forest-forming species (see Fig. 62). At the same time, in order to make the characteristics of forests more specific and show all the diversity, they use the concept of forest types. Forest types are distinguished on the basis of common physiognomic features of phytocenosis, its floristic composition and habitat conditions. In a taxonomic sense, the type of forest approaches the concept of an association.
Consider a brief description of the main formations of coniferous forests.
Spruce forests. This formation is most common among the dark coniferous forests of the European part of the USSR and Western Siberia. The forest-forming species is spruce. In the USSR, spruce has up to 10 species, of which the most common spruce (European part of the USSR) and Siberian spruce (north-east of the European part of the USSR and Western Siberia).
Spruce is one of the shade-tolerant species, so spruce forests are dense and shaded. The undergrowth in the spruce forest is not widely developed. In the undergrowth there are shrubs: buckthorn, honeysuckle, juniper. The herbaceous cover is characterized by shrubs (lingonberries and blueberries), fern, club moss, oxalis, wintergreen, etc. Green mosses and bearded lichen are abundant in the spruce forest.
Spruce forests form a large number of associations. Currently, to identify them, as well as to determine the patterns of distribution, it is used ecological series method, developed by acad. V. N. Sukachev. Ecological series make it possible to reveal the sequence of arrangement of plant associations in connection with changes in environmental conditions (Fig. 66).
If we consider the change in spruce forest types depending on environmental conditions, the following regularity can be outlined. Optimal growing conditions (point O) corresponding association spruce forest. With an increase in dryness and a decrease in soil fertility (a number of BUT) association replaces spruce lingonberry. Grows in dry conditions spruce-lichen. In wet areas with signs of stagnant
swamps (row AT) develops spruce-blueberry. A typical spruce forest association is spruce forest, which grows in low places of excessive moisture. In swamp conditions, it is replaced by sphagnum spruce forest- an association where spruce is strongly oppressed, and sphagnum moss forms a continuous cover (the final member of the series AT). Under conditions of flowing humidification (series D) an association is formed along the stream valleys marsh-grass spruce forest(stream) with developed undergrowth. Also of note is the association oak spruce forest, which is confined to the most fertile soils of the forest zone (a number WITH). Broad-leaved species (oak, linden) usually form an admixture to spruce, hazel in the undergrowth, and a cover is also developed, where broad-leaved oak-grass predominates.
The above associations of the spruce forest do not exhaust the diversity. The considered ecological series show how the composition of associations changes with a change in ecological conditions and position in the relief (Fig. 67). Thus, ecological series can be considered as one of the methods for studying plant associations with their frequent changes in space.
A certain regularity is outlined in the distribution of dark coniferous forests on the territory of the USSR. Within the European part of the USSR, spruce forests predominate, occupying wide and flat watersheds. In the West Siberian Lowland, forests of this type gravitate toward river valleys as more drained areas. In southern Siberia, they are found in the mountains, where the participation of fir and cedar (Tien Shan) is increasing.
Fir forests are close to spruce forests in their characteristics. Their predominant associations belong to the “green moss” group.
Fir forests have a more limited distribution. They avoid both dry and wetlands. Often there are mixed fir forests with an admixture of Siberian spruce and Siberian pine (West Siberian lowland, Urals, Altai, Sayan mountains, etc.).
Larch forests occupy a large area in the USSR. The area of distribution of larch falls on Eastern Siberia (to the north of 48 ° N), where Dahurian larch is common. This breed, having a superficial root system, is well adapted to growing in conditions of a sharply continental climate on swampy soils with close occurrence of permafrost. Siberian larch is common in the mountains of southern Siberia.
A characteristic feature of larch forests is that they form pure stands. Since the density of crowns in the forest is small, the larch forest has a park-like appearance. In the upper tier larch reaches 30-35 m height. The ground cover is dominated by upland grasses and shrubs (reed grass, cranberries), and in some places sphagnum mosses. Larch forests have a large supply of valuable industrial wood, and are also valuable hunting grounds.
Pine forests. This is the most widespread type of coniferous forest in the temperate zone. The main forest-forming species is Scotch pine. Other types of pine (Crimean, Pitsunda, etc.) have a very limited range.
Compared to other tree species, pine has a large ecological range. It is most widespread on sandy plains and river terraces. At the same time, it forms peculiar associations in peat bogs. Pine forests penetrate the forest-steppe zone and are found in the mountains.
Pine forests, like larch forests, often have pure stands of one or two tiers. The grass cover in them is poor and is mainly characterized by the predominance of shrubs (heather, blueberries, lingonberries, etc.). In pine forests of dry habitats, a cover of lichens or green mosses is developed (Fig. 68, 1).
The main associations of a pine forest and the patterns of their change can be expressed by a system of ecological series similar to spruce forests, which include associations of certain ecological conditions (lichen pine forest, green moss pine forest, sphagnum pine forest, etc.).
In terms of the composition of their associations, cedar forests are close to pine forests. Their distribution area is confined to the northeast of the European part of the USSR and Western Siberia. Cedar forests, like pine forests, are found in sphagnum bogs. In the mountains of Siberia, cedar forests of a mixed type - cedar-larch (Altai, Sayan, etc.).
Geographic patterns of distribution of coniferous (taiga) forests in the northern hemisphere are quite complex. Distributed in the temperate zone, coniferous forests in their composition change markedly both from north to south and from west to east.
Currently, geobotanists divide taiga-type forests into three groups of formations: northern taiga, middle and southern taiga, which are characterized by the predominance of certain forest types. Geographically, they represent wide bands (subzones), which are clearly visible on the plains (Russian Plain, West Siberian Lowland). Thus, the northern taiga of the European part of the USSR, located south of the forest-tundra, is distinguished by the predominance of species of Siberian origin (spruce, cedar, larch). Here the predominant type of forest is green moss spruce forest. The middle taiga is characterized by spruce (spruce-blueberry) and spruce-fir forests; southern taiga - spruce forests with an admixture of broad-leaved species (oak, elm, linden, maple). The northern and middle taiga are very heavily swamped, and sphagnum pine forests are common in raised bogs.
Taiga in the Asian part of the USSR retains the same subdivision. Its distinctive feature is the large swampiness of forests (up to 50%).
In the northern taiga of the West Siberian Lowland (the southern border coincides with the latitudinal segment of the Ob River), cedar, spruce-larch, and pine forests are common. East of the river Yenisei in the area of permafrost development, larch forests predominate.
The middle taiga in Western Siberia is characterized by the dominance of dark coniferous fir-spruce forests (“urmans”) and cedar forests. In Central Siberia and Yakutia, they are replaced by larch-pine forests (Dahurian larch).
The southern taiga is distinguished by the predominance of pine and birch forests, and in the lower areas - spruce-fir (Western Siberia), cedar-fir (Central Siberia) and larch woodlands (Transbaikalia). A special variety is represented by marshy larch forests "Mari", widespread in Central and Eastern Siberia and the southern regions of Transbaikalia. The forests consist of Dahurian larch, have an undergrowth of yernik (birch) and a continuous cover of sphagnum. This type of forest is confined to river valleys, where peat-bog soils are formed.
Coniferous forests of Western Europe do not form marked subzones and grow only in the mountains (Alps, Pyrenees, Carpathians, etc.). In addition to ordinary pine and spruce, there are European larch and fir, which form a special forest belt.
Coniferous forests in North America occupy large areas (Labrador, Alaska, mountains of the Pacific coast, Atlantic plains). Unlike European coniferous forests of the American type, they amaze with a wide variety of species of pine, spruce, fir, and larch. There are also specific species in the forests, especially on the western coast of the mainland, such as douglas fir, thuja, in the Sierra Nevada mountains - giant sequoia(mammoth tree). These tree species are distinguished by their gigantic height (up to 80-100 m). The main reason for the species richness of the American taiga is the favorable conditions for the migration of species during the Ice Age.
Summer green deciduous forests. Deciduous forests in temperate latitudes are spreading in a maritime climate. In Eurasia, these forests are typical of Western Europe, the south of the Russian Plain, the Caucasus and the Carpathians. Further to the east they are replaced by coniferous forests. The range of deciduous forests is represented in the Far East of the USSR, in eastern China and on the Japanese islands. Deciduous forests are found in North and South America (Patagonia).
Deciduous forests are divided into broad-leaved and small-leaved.
In broad-leaved forest-forming species, are oak, beech, linden, maple, elm and ash. These forests have a well-developed crown, and the upper layer in them is usually composed of a single forest-forming species. Since the forests are shady, the undergrowth and grass cover are weakly expressed. Herbal species are dominated by ephemeroids, which develop intensively in spring and autumn. Typical formations of deciduous forests are as follows.
Beech forests are most widespread in Western Europe. Near the northern border of their range, they are distributed on the plains, in southern Europe - in the mountains, where they form a forest belt. Within the USSR, beech forests are found in the west of Ukraine, in Moldavia, as well as in the Carpathians, Crimea and the Caucasus, where they form a special belt.
The beech forests of Europe are of the same type. The forest-forming species in them is beech. Due to the large shading, undergrowth and summer grasses are usually absent. In the mountains, the companions of beech are fir, yew.
The beech forests of North America (eastern United States, Canada) are different from European ones. The forests have a wide variety of species, but American beech and sugar maple dominate. Creepers from wild grapes are characteristic.
Oak forests are the more common type of broadleaf forest. In the USSR, oak forests are widespread in the European part, where they form a subzone of deciduous forests. In oak forests, the main forest-forming species is the pedunculate oak, which is mixed with maple, ash, linden, and elm. Oak forests are multi-tiered. They have an undergrowth of hazel, euonymus. Plants are represented in the grass cover - oak broad grass (snot, lungwort, hoof etc.), the aspect of ephemeroids with under-snow development is also characteristic in spring (see Fig. 68, 1, 2). Oak forests have a wider range than beech forests and are found in other subzones, for example, forest-steppe, where they form ravine forests along the gullies. Forest-forming species in the oak forests of Western Europe are holm and downy oak, which are combined with evergreen rhododendron and yew (Ireland). In North America, oak forests are common in the continental regions of the west bordering the prairies. In contrast to European ones, there is a large participation of broad-leaved species here: several species of oak, maple, walnut, plane tree, etc. Lianas are also characteristic of oak forests.
small-leaved forests (birch, aspen, alder) along with conifers and broad-leaved trees, they are especially widespread. These forests are originally secondary, which have become widespread after cutting down broad-leaved and coniferous forests. However, examples of primary birch forests are known in the West Siberian forest-steppe. (pegs) and in Kamchatka.
There is also a mixed type of forests - coniferous-broad-leaved, in which small-leaved species occupy a significant place. These forests are predominantly distributed on the border of coniferous and broad-leaved forests. Depending on the change in environmental conditions, there is an alternation of various types of coniferous and broad-leaved forests: oak forests are confined to the slopes of the southern exposure of river valleys, pine forests are common on terraces, and spruce forests are common on flattened watersheds. Mixed forests are especially typical of the European part of the USSR, where they form an entire subzone.
Steppe vegetation.Steppes are a herbaceous type of xerophytic vegetation with a dense herbage that develops in a temperate climate with a deficit of summer precipitation. The steppes occupy the largest expanses in the USSR, stretching in a wide strip in the European and Asian parts. Steppes (Pasht) found within the Danube lowland. In North America, the steppes are called prairies.
Vast spaces of the steppes (pampas) in subtropical parts of South America, Africa and Australia.
The main features of the steppe vegetation will be considered by us on the example of the steppes of the USSR. Steppe vegetation in terms of physiognomic features differs sharply from other herbaceous types (for example, meadows, swamps), since it has xerophytic traits. To endure summer droughts, plants have developed adaptations such as a wax coating on the leaves, their pubescence, and in some cases, the reduction of the leaf blade. All terrestrial vegetative parts of plants have a dull green hue, which creates a certain background of the steppe.
The steppe vegetation is characterized by the predominance of special floristic groups. Widespread development is being turf grasses, having narrow folded leaves and turf deeply seated in the soil (feather grass, fescue, thin-legged). Also presented sedge, legumes, forbs, ephemera.
Most hallmark steppe is her dynamism. The flora of the steppe consists of plants that do not coincide in their phenological phases. Therefore, the appearance of the steppe, its color background changes in different periods. So, in early spring, the appearance of the steppe is determined by the flowering of yellow tulips, blue hyacinths, golden goose onions and white crocuses. The silvery color of stepsh in May is due to feather grass. In June, the flowering of tumbleweed plants is noted. The golden-green background of the steppe in July is associated with the flowering of feather grass - "tyrsa". In August, wormwood, steppe asters begin to bloom.
For the entire flowering period in the steppe, there are up to 12 colorful phases (Streletskaya steppe). The noted change of aspects in the steppe should be considered as the adaptation of vegetation to certain ecological conditions.
V. V. Alekhin divided the steppes of the USSR into types: northern (meadow) steppes and southern steppes located on the border with semi-deserts.
The northern steppes are characterized by the development of colorful forbs. The participation of feather grasses is insignificant (common and narrow-leaved feather grass). Of the cereals, loose soddy ones predominate (bonfire, wild oats, bent grass, etc.). The maximum species saturation is noted when the number of species per 1 m 2 reaches 80, and their number - up to 2000. Steppe phytocenoses have a complex layering.
The southern steppes are distinguished by the predominance of feather grasses, forming an aspect: feathery feather grass (fescue) and feather grass - "tyrsa". There is very little forbs in the grass cover. Only the phase of spring ephemeroids (tulips) is sharply expressed, and the number of tumbleweed plants is increased. In the southern steppes, the grass cover is very sparse and, in general, is characterized by low species saturation compared to the northern steppes: by 1 m 2 area accounts for no more than 12 species.
The northern and southern steppes in the European part of the USSR have been completely plowed up. The surviving virgin areas are declared reserves.
The Siberian steppes have many features in common with the European ones. However, in the conditions of a peculiar ridged relief (Baraba steppe), they are combined with grassy swamps and solonchaks. In the steppes of Western Siberia, feather grasses are predominant. Near the border of the birch forest-steppe, there are forest, marsh species, as well as saltwort plants.
The North American prairies are similar in floristic composition to the European steppes. Three types of cereals are dominant in the American prairies: feather grass, wheatgrass and gram (the latter is not found in the European steppes). A feature of the prairies is the distribution of deep-rooted species, in which the roots go to a depth of 1.6-2 m. At the same time, cereals are distinguished by a large height (80-120 cm).
The steppe types of North America are extremely diverse. The tall grass prairies of the Great Plains are close to the steppes of the "northern" type. Grass formations are dominated by such grasses as bearded man, Indian grass, feather grass, wheatgrass. In spring, blooming forbs are abundantly presented.
In the drier conditions of the Prairie Plateau, short-grass steppes are common, where densely tufted grasses (gram grass, endemic bison grass) predominate.
The "Pampas" of South America occupy vast plains in the south and east of the mainland (Argentina, Uruguay). The vegetation has all the features of xerophytes and is characterized by distinct changing aspects. The main elements of the steppes are perennial grasses of the genus feather grass, bearded vulture, and millet. Variety of herbs.
Desert vegetation. The desert type of vegetation is characterized by the predominance of shrubs and shrubs. The factors determining the development of the desert are essentially climatic, since they are formed in an arid climate with a high moisture deficit in extratropical and tropical regions of the globe (deserts of the USSR, Central Asia, Sahara, Colorado, South America, Australia, etc.) - Unfavorable in Deserts also have edaphic conditions: soils are depleted in humus and saline, groundwater is located at great depths. Desert vegetation, compared with steppe vegetation, is characterized by a sharp increase in drought-resistant species. In the process of adaptation of plants to adverse climatic conditions, a number of life forms. Among them, the most adapted to the transfer of dryness plants - xerophytes, having a developed taproot and superficial lateral roots, a rigid stem, reduced leaves. The most typical representatives are semi-shrubby plants, such as camel's thorn, black and white saxaul, etc. (Fig. 69).
Another predominant life form is ephemeroids. These tuberous and bulbous perennials are finishing
vegetation for 1-2 months before the onset of the dry period (bulbous bluegrass, tulips, onions). At this time, the deserts are covered with a continuous carpet of flowering plants.
A number of desert plants have the ability to accumulate water reserves in the hairs covering the leaves (for example, in the kokpek shrub) or in the tissues of the leaves and stem (plants succulents). The latter are very characteristic of many deserts of the globe. Their main representatives are cacti, spurges, etc. The amount of water that they accumulate can be 96% of their weight. These life forms form a complex complex with saltwort plants.
By the nature of environmental conditions and, first of all, by the mode of precipitation, the nature of the substrate, the deserts of the globe can be divided into a number of types: clayey, sandy, stony, solonchak. We will consider their main features on the example of the deserts of the USSR.
The deserts of Central Asia occupy more than 2 million hectares. km 2(10% of the area). According to climatic conditions, they can be divided into northern and southern.
Northern deserts are formed in a temperate continental climate with a uniform distribution of precipitation throughout the year. These are predominantly clay deserts (Ustyurt, Bet-Pak-Dala, etc.).
A common feature of their vegetation cover is the predominance of xerophyte shrubs in combination with saltwort. Depending on the soil, a number of typical phytocenoses are formed. The most common are sagebrush deserts (clayey), which are characterized by the predominance of various types of sagebrush and saltwort. The vegetation cover is monotonous and very sparse (the cover does not exceed 40%).
In areas with highly saline soil, saltwort deserts become widespread, represented by a group of associations dominated by a low semi-shrub kokpek and a pillow-shaped semi-shrub biyurgun.
These types of deserts do not occupy large areas. In conditions of complex microrelief, depending on the degree of soil salinity, they form complexes. For example, on the bottom of the basins and saucer-shaped depressions, the kokpek desert is widespread, on their slopes - saltwort, on elevated areas - sagebrush deserts (Fig. 70). This complexity of the vegetation cover is reflected in the legends of geobotanical maps.
Climatically, the southern deserts are quite different in terms of temperature and a pronounced spring-autumn maximum of precipitation. This explains their significant difference and a wide variety of types.
Clay deserts of the southern variant are formed on piedmont loess plains (Kopet-Dag, Pamir-Alay, Tien Shan). Their vegetation cover is mesophytic in nature. In the spring, a continuous sod of grass is formed, which can be compared to a meadow. The most typical plants are bulbous ephemera: sedge, bulbous bluegrass, which form a dense, albeit low (20 cm) sod. The projective cover reaches 80-100%. In spring, the ephemeral desert is used as pasture. In the summer, all ephemera die off and the soil surface dries out.
Sandy deserts (Karakum, Kyzylkum, Muyunkum, Balkhash sands, etc.) are distinguished by the greatest variety of plant associations. There is a wide variety of life forms here, including ephemera, shrubs and trees.
This is largely due to the properties of sands, in which a favorable water regime is created (permeability, poor capillarity, the ability to condense moisture). In the sandy desert at a depth of 100-150 cm there is a constant moisture horizon, which in spring is supplemented by a "hanging horizon". Among the unfavorable environmental factors on the sands should be attributed to their mobility. Plants as an adaptation for growing on moving sands form deep adventitious roots.
The distribution of plant associations depends on the degree of sand fixation. Tall grass dominates on mobile dunes and dune chains Celine(up to 1 m), juzgun, sand acacia- low trees (up to 6 w) with a characteristic "weeping" shape (Fig. 71).
In places of development of ridge and hilly sands, gnarled bushes are widespread. white saxaul(up to 2-3 m). Its distinctive feature is the summer shedding of branches. From other shrubs, juzgun and tree-like saltworts are found. Under the canopy of white saxaul, a grass cover of ephemera and ephemeroids develops.
Saline deserts are usually confined to river terraces (the Amu Darya and Ili rivers, etc.), sea coasts, and deep depressions where highly saline soils are developed. They are characterized by a peculiar saltwort (halophytic) vegetation, among which succulents predominate. The most typical plants are: sarsazan - semi-shrub with a fleshy stem (sparse thickets of sarsazan occupy large areas on the coast of the Caspian Sea), succulent saltwort(tamarixes) - plants whose leaves are covered with a salt crust, and sulfur wormwood.
Associations are widespread on weakly saline soils. black saxaul- leafless tree-like saltwort, reaching a height of 5-9 m. Dense thickets of black saxaul are widespread in the Balkhash region, in the lower reaches of the Amu Darya, etc. Saxaul is a good fuel, which is slightly inferior in calorific value to some varieties of coal.
The desert type of vegetation on the globe occupies significant areas. In Eurasia, outside the USSR, there are vast deserts that form in a subtropical and tropical climate (Tar, Registan, Rub al-Khali, Syrian, Gobi desert, Alashan, etc.). These are predominantly upland deserts, which are characterized by the development of thorny cushion-shaped shrubs (astragalus), succulents, and wormwood.
Large areas (up to 40%) are occupied by the desert in Africa (Sahara, Kalahari, etc.). The vegetation of the Sahara approaches to some extent the deserts of Central Asia. Shrubs (camel thorn, acacia) and grasses (related to seline) are common on sandy areas. Significant areas are also occupied by a rocky desert ("gammada"), which is characterized by lichens that form a continuous crust on stones, and rare semi-shrubs. The deserts of South Africa are characterized by the dominance of succulents, which are distinguished by an abundance of species: aloes, spurges (Karoo desert).
Grasses predominate in the sandy Kalahari desert, and acacia dominates from woody forms.
The Namib Desert (Orange River basin) is distinguished by its greatest originality, where special forms of succulents dominate, which look like stones - "stone plants". A very peculiar endemic relict plant velvichia(obviously a representative of the Mesozoic flora). The life expectancy of Velvichia is more than 100 years. Its woody stem is up to 1.5 in diameter m rises to a height of no more than 20 cm(Fig. 72).
The deserts of North America are peculiar: along with the indicated types (saltwort, black wormwood, etc.)
Spread creosote shrub desert (Colorado Plateau, California coast). Mexican upland deserts at 1000-2250 m above sea level are the center of education cactus flora. Up to 500 species of cacti, prickly pear, agave, tree-like yuccas are represented in this area (Fig. 73). Giant cacti in the desert are combined with thorny bushes (mimosa, creosote bush, etc.). The vegetation cover of the Mexican desert is very sparse.
The deserts of South America and Central Australia are saline plains and plateaus abounding in sands and salt marshes. The vegetation background in them is formed by halophytic shrubs, succulents and herbs. Specifically Australian are bushy deserts with a predominance of acacia and shrub eucalyptus. Extensive sandy expanses are occupied by the formation of hard and prickly cereals (spinifex) growing on loose sands and stones (Western Australia).
Subtropical shrub-wood vegetation.Humiditynye subtropical forests are typical for areas of subtropical climate. This type of forest is common in East Asia, Central and South America and other areas and is represented by evergreen trees and shrubs. Dominant in hardwood forests are laurel, plane tree, oak, boxwood. Ferns and mosses are plentiful and often grow as epiphytes. In the forests of Florida, Chile and Patagonia, evergreen beech predominates.
A variation of this formation is evergreen hardwood forests and shrubs, typical for countries
Mediterranean, as well as California, southwest Australia, South Africa (Cape), South America (Patagonia). The life forms of hardwood forests are very peculiar. Plants have xerophilic adaptations: hard leaves covered with resinous secretions, twig-like stems. The dominant species in the forests of the Mediterranean are stone and cork oak. The forests have an undergrowth of evergreen shrubs such as myrtle and heather.
In Southern Europe, North Africa and the countries of Asia Minor, evergreen forests begin to be dominated by groves of pine pine and Lebanese cedar.
The hard-leaved forests of the subtropical zone are everywhere combined with thickets of various tall shrubs, which in the Mediterranean are called maquis. The maquis formation is characterized by strawberry tree, myrtle, tree heather. In thickets of low evergreen shrubs called gariga, shrub oak, thyme, rosemary, gorse, and others dominate (south of France). In North Africa and southern Spain, the gariga is represented by a dwarf palm. The hard-leaved eucalyptus forests of southern Australia are very peculiar with evergreen undergrowth and thickets of shrubs (scrub) from various types of acacias, shrub eucalyptus, etc.
Humid tropical (rain) forests. This type of forest formations is spreading in the equatorial climate zone. Forests occupy vast areas in Africa (the Congo and Niger basins), Central and South America (the Amazon basin), and Southeast Asia. Tropical rainforests stand out sharply from all other forest formations with a large variety of species. The trees are dominated tree ferns, ficus different types palm trees(coconut, oilseed, wine) rubber plants(Hevea brazilian).
The structure of tropical forests is the most complex. The number of tiers in them reaches 4-5. Tall trees (up to 60 m and above), with characteristic board-shaped roots. The shading of the forest is maximum and the surface of the soil reaches only 1/150 of the incoming light. It should also be noted that there is also a smaller variety of grass cover species, where mainly spore plants predominate: ferns, club mosses. Another characteristic feature of tropical forests is the abundance vines and epiphytes(Fig. 74). The spread of such life forms is explained by the great shading of the tropical forest. The most common: liana palms (up to 300 m), creepers from the philodendron family, pepper, vanilla, etc. Epiphytes are herbaceous species from the fern and orchid families, as well as mosses and algae.
A very characteristic formation of humid tropical forests is mangrove vegetation, common in the tidal zone of bays and lagoons of the northern and eastern coasts of South America, West Africa, Hindustan, etc. Mangrove vegetation is a thicket of evergreen shrubs. The participation of trees in them is small. In terms of species, this formation is extremely monotonous (rhizophora predominates, some types of palms). This is due to the specifics of environmental conditions, since tree crowns rise from the water during high tide, and at low tide, trunks, stilted and respiratory roots are exposed. Such a root system is a kind of adaptation for the transfer of oxygen during the flooding of the coast. Mangrove plants also have features characteristic of halophyte plants (Fig. 75).
In the climate region of the equatorial monsoons, a special type of deciduous (winter-green) tropical rain forests develops (Indo-China, Hindustan, Sunda Islands). The forests are similar to tropical ones, but during the period of drought, the trees shed their leaves. These forests are different in the nature of the stand. As part of mixed forests, forest-forming species are valuable tree species (sandalwood, rosewood), bamboo, palm trees. There are many flowering shrubs and herbs in the forests. Lianas and epiphytes are also characteristic, which lose their foliage during dry periods.
With increasing dryness in the tropical zone, winter-green tropical rainforests are replaced by dry xerophilous forests and thorn bushes. This formation occupies large areas in Africa (Rhodesia, Angola, Somalia),
Argentina, northern Australia. Xerophilic forests are short and sparse. They are dominated by leafless trees and shrubs such as acacia, palm trees, intertwined with lianas. The undergrowth is dominated by thorny bushes. The xerophilous forests of Brazil are peculiar "caatinga" with many specific species: cacti, including tree-like ones, trees from the bombas family, having swollen barrel-shaped trunks several meters in diameter (Fig. 76). The distribution of these formations can be seen on the world vegetation map.
This is a woody-herbaceous type of vegetation, which is transitional from tropical forests to tropical deserts. Savannahs are widespread in southern and central Africa (basins of the Niger and Upper Nile rivers), South America (Brazil, the Orinoco river basin) and Australia.
In the harsh dryness of tropical climates, evergreen trees in the savannas have stiff, hairy leaves that shed during the dry season. Another feature is the umbrella shape of the crown as an adaptation from strong winds. Trees in the savannah are scattered in small groups. The main representatives are umbellate acacia and baobab - giant tree, reaching a height of up to 25 m and having a diameter of up to 9 m(can reach the age of 5000 years) (Fig. 77). In the Australian savannas, eucalyptus trees predominate, in the savannahs of South America - "llanos" - varieties of palms. The herbaceous cover of savannahs is dominated by tall xerophytic hard-stalked grasses (such as bearded vulture).
Swamp vegetation. Bogs develop in various climatic zones - from equatorial to subarctic. They are especially characteristic of the temperate forest zone.
Swamps are characterized hygrophilic plants growing in conditions of excessive moisture and experiencing the influence of "physiological dryness". When plants die, peat accumulates.
The modern classification of swamps is based on the allocation types(formations) according to the following features: 1) position in the relief, 2) conditions of moisture and nutrition, 3) prevailing plant associations.
The most common type are lowland swamps. They form on the bottoms of the valleys of streams, ravines, gullies and are characteristic of all natural zones. Humidification of swamps is associated with the close occurrence of mineralized groundwater. Green mosses, various sedges and grasses dominate in the grass cover of lowland swamps. Birch, alder and willow appear on older swamps. This type of bogs is characterized by low peatiness (the thickness of the peat layer does not exceed 1-1.5 m).
A variety of grass and hypnum-grass bogs are tall-grass bogs with thickets of reeds, reeds, cattails, widespread in floodplains and lake basins in the forest, forest-steppe and steppe subzones of the European part of the USSR, Siberia and the Far East.
Raised bogs have specific features. They form on flat watersheds and occupy vast expanses of the forest zone (for example, the West Siberian and Pechora lowlands, Polissya). Humidification of raised bogs is associated exclusively with atmospheric precipitation and in conditions of a flattened relief.
becomes excessively stagnant. The surface of the swamp acquires a convex profile due to the uneven growth of peat. The excess of the central part of the swamp over the periphery reaches 3-4 m. During the development of the swamp, a complex microrelief is formed on its surface, represented by a combination of small ridges and depressions (see Fig. 82).
The ecological conditions of the raised bog are very peculiar. Plants adapt to growing in an environment depleted in mineral nutrition, especially nitrogenous compounds. Common in upland bogs sphagnum mosses and accompanying marsh species: cotton grass, wild rosemary, cassandra, heather, cranberry and others. The peculiarity of their life forms is that they have adventitious roots and a moving growth point. The vegetation of the raised bog (Fig. 78) is also adapted to the transfer of "physiological dryness".
Shrub plants of the raised bog have narrow leathery leaves with a wax coating, woody stem. Sedges, cotton grass, etc., which have narrow hard leaves, also acquired a specific appearance. Of the tree species on the raised bog, they are most often found pine and Birch, less often cedar and larch. The trees in the swamp are very oppressed and stunted.
The named plant dominants for the raised bog (trees, shrubs, mosses, etc.) form certain associations depending on the environmental conditions. In their distribution, the following regularity is outlined: in the central part of the raised bog, pine-sphagnum association, towards the periphery of the flywheel in the places of development of the ridge-hollow complex, it is replaced by pine-shrub, on the edge of the swamp
is gaining ground pine-cotton grass association (the nature of the distribution of associations in the raised bog and their location in the plan, see Fig. 79).
Raised bogs are of great practical importance. Their peat layer often reaches a thickness of 6-10 m. Peat is used as a fuel, fertilizer and for the production of a number of chemicals.
Raised bogs are common in North America (mainly in the northeast). Here, the swamps are similar in nature to European ones, but larch predominates among tree species. In South America, sphagnum bogs are found in the Andes and Tierra del Fuego.
Transitional swamps are mixed in nature, they form on terraces or concave slopes of interfluves. The vegetation cover of such marshes combines the features of lowland and raised marshes and is characterized by a predominant distribution sphagnum-cotton grass-sedge associations.
The process of development of swamp massifs is very complicated. With the death and accumulation of plant residues in the swamp, its water regime is essentially associated, since the growth of the peat layer reduces the inflow of groundwater. This, in turn, entails a change in ecology and the corresponding swamp associations. The above types of swamps can be considered as certain stages this development. Low-lying sedge-grass bogs of soil moisture are replaced by transitional sphagnum-sedge bogs of mixed nutrition. Raised bogs of exclusively atmospheric nutrition represent the final stage of the development of the bog, at which the tendency of peat growth in height is manifested.
The planned scheme gives the most general idea of the development of swamps. At the same time, the causes of swamping are extremely diverse. During the development of spruce and pine formations, swamping is facilitated by the development of a moss cover of cuckoo flax and sphagnum. Often, forest fires and clearings are swamped. And, finally, swamps appear on the site of lakes and river oxbow lakes, which are inhabited by wetland vegetation. Let's take a look at this process in general terms.
Waterlogging of a lake is most often overgrowth. With gentle shores and a gradual increase in depth, aquatic vegetation is located in the form belts in the following sequence: 1) in the deepest places, a belt of green algae stands out (at a depth of over 6 m); 2) a belt of flowering plants submerged in water (hornwort, pondweeds); 3) a belt of broad-leaved pondweeds, which have wide floating leaves and inflorescences on the surface of the water (at a depth of 4- 5 m); 4) belt of water lilies at a depth of 2-3 m(water lily); 5) belt of reeds up to 2 m(bulrush, reed, horsetail); 6) a belt of large sedges at a depth of up to 0.7 m; 7) belt of small sedges (Fig. 80).
Each of the marked belts is not durable and is replaced by the neighboring one, which is less deep. This is due to the fact that the deposited plant residues contribute to the shallowing of the reservoir. This is how associations are consistently shifted from the periphery to the center. Its last stage is the transformation of the lake into sedge swamp.
Of all the listed swamp associations, the most common on the globe are reed swamps with a predominance of tall reed beds (up to 6-10 m). In African tropical swamps along the banks of rivers and lakes thickets of papyrus, reeds and reeds stretch, in Indian - thickets of bamboo.
Meadow vegetation.Meadows are a herbaceous type of vegetation of a mesophilic nature, growing in conditions of moderate moisture in various natural zones of the globe (tundra, forest zone, tropical, etc.).
By location, meadows can be subdivided into floodplain (flood) and watershed (upland). The floodplain meadows are of the greatest interest as a type of plant formations. Their vegetation is formed on fertile soils under the influence of a long flood regime and the deposition of loose river sediments.
Floodplain meadows are characterized by a number of floristic features: the predominance loose sod grasses(meadow timothy, fescue, bluegrass, awnless bonfire), legumes(clover, alfalfa, mouse peas) and forbs(meadow geranium, meadow cornflower, common daisy and a number of others). The herbage of the meadows is distinguished by high merits and is a valuable hayfield!
On the floodplains of large rivers, which are distinguished by a complex microrelief and different moisture, a variety of environmental conditions are created. When crossing the floodplain from the riverbed towards the slope, a complex ecological line.
The most arid conditions are created in the elevated near-channel part of the floodplain, composed of sandy alluvium. Here, in the sparse grass cover, long-rhizome grasses (couch grass, awnless bonfire) predominate, and some steppe species are also found.
The central part of the floodplain is composed of clayey alluvium and has a high standing groundwater. Loose sod grasses, legumes and flowering forbs dominate here, having a dense herbage.
The nature of the meadow changes dramatically in the terraced part of the floodplain. This lower section of the floodplain at the foot of the slope is the most humid due to the stagnation of hollow waters and the outflow of springs. The terraced floodplain is often forested. Alder groves with blackcurrant undergrowth are the primary forest type.
Sod grasses (for example, pike) and sedges predominate in wet meadows.
In the forest zone, upland meadows are also distinguished, which are formed on watersheds. Unlike floodplains, dry valleys are of secondary origin, as they arise on the site of cut down or burnt forests. Therefore, in their herbage, in addition to purely meadow mesophytes, there are elements of forest forbs and a well-developed moss cover. The productivity of such meadows is low.
In the steppe zone, watershed meadows are formed in depressions (estuaries, pods, floods, etc.).
The vegetation of meadows is influenced by the type of vegetation (steppe, desert, etc.) in which it is formed. So, on the floodplain of the river. Oka, crossing the subzone of deciduous forests, there are steppe species, on the floodplain of the river. Teberdy - subalpine species. On the floodplains of the steppe rivers Sala and Manych, saltwort plants, characteristic of semi-deserts, predominate.
Alpine vegetation is a special type of alpine low-grass meadows, which are located above the upper border of the forest. The altitudinal position of alpine meadows varies depending on the geographical latitude of the mountains, the exposure of the slopes, and the degree of continentality of the climate. For example, in the Alps and in the Western Caucasus, alpine meadows are located at altitudes of 2200-3000 m above sea level.
Alpine vegetation is similar to tundra in a number of ways: the predominance of perennial shrubs, cushion plants, etc. All this is an adaptation of alpine vegetation to the conditions of a short growing season with sharp temperature contrasts during the day. In conditions of high intensity of illumination, alpine plants acquire a stunted squat shape and bright colors of flowers.
Alpine vegetation is divided into two altitudinal belts: the lower one is subalpine, the upper one is alpine meadows.
Subalpine meadows are most widespread in the mountains of Central Asia and the Caucasus. Their vegetation is distinguished by extraordinary diversity and brightness. The grass cover is dominated by cereals and flowering forbs (geraniums, scabiosa, bluebells, alpine poppies, forget-me-nots, anemones, asters and many others). This type of herbaceous phytocenosis is characterized by complex layering and high species richness. Subalpine meadows of the Western Caucasus are characterized by the development tall grass, reaching up to 2 m and more (bellflower, cow parsnip, elecampane, catchment area). There are also thickets of Caucasian rhododendron, juniper, willows.
Alpine meadows replace subalpine ones at altitudes above 3000 m. In the grass cover of the Tien Shan meadows, mesophytes dominate. An important element of it are turf sedge(kobresia) with a very large and bright flower. The size and brightness of the color of the flowers are a characteristic feature of the alpine vegetation. Flowering forbs are similar to the subalpine belt, but in appearance the vegetation of alpine meadows is distinguished by a low herbage. The leaves of the herbs are arranged in the form of a rosette, often without a stem (stars, forget-me-nots, buttercups, alpine poppies and many others). There are many bulbous plants in the alpine meadow: tulips, hyacinths, mountain lilies and others. The Alpine meadows of the Alps are characterized by an abundance of species of tundra flora (for example, dryad), there are endemic species, such as edelweiss, primrose.
- Source-
Bogomolov, L.A. General geography / L.A. Bogomolov [and d.b.]. – M.: Nedra, 1971.- 232 p.
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