PROFESSOR OLGA V. SMIRNOVA’S SYSTEM OF VIEWS IN FOREST ECOSYSTEM ECOLOGY

Original Russian Text © 2019 O. I. Evstigneev, V. N. Korotkov published in Forest Science Issues Vol. 2, No. 4, pp. 1–36.

© 2022                                             O. I. Evstigneev ^{1, 3 *}, V. N. Korotkov²                

¹State Nature Biosphere Reserve “Bryanskii Les”,
Nerussa Station, Bryansk Oblast 242180, Russia

²Yu. A. Israel Institute of Global Climate and Ecology, 20B
Glebovskaya st., Moscow 107258, Russia

³Center for Forest Ecology and Productivity of the RAS,
Profsoyuznaya st. 84/32 bldg. 14, Moscow, 117997, Russia

*E-mail: quercus_eo@mail.ru

Received: 16.09.2019

Revised: 07.10.2019

Accepted: 07.10.2019

 

Professor Olga V. Smirnova, Doctor of Biological Sciences, is a prominent scientist in the field of plant demography, population biology, and forest ecosystem ecology. Professor Olga V. Smirnova’s edifice is based on ideas about the leading role of plant and animal populations in the organization of the biogeocenotic cover. In this case, it is implied that a continuous generational turnover in edificator (keystone species) populations is necessary to maintain the species and structural diversity in communities and ensure their sustainability. This system of views was influenced by Professor Alexey A. Uranov. The development of these ideas was consistent and gradual. First, Professor Olga V. Smirnova studied the biology of different plant species life forms. Examining their individual development, with identification of ontogenetic stages, is necessary for demographic research. She then developed the theory of coenopopulations as supraorganismal systems, which can self-sustain under different conditions. Finally, she developed the doctrine of biogeocenosis as a system of interacting populations and created the concept of anthropogenic transformation of the forest cover in the Holocene. Her contributions helped researchers to understand the mechanisms of the formation of modern zonality that are due to human activity.

Key words: plant biological age, plant population strategy, coenopopulation, edificator, forest ecosystem ecology, modern zonality, historical ecology

October 9, 2019 marks the anniversary of Professor Olga Vsevolodovna Smirnova, Doctor of Biological Sciences, a prominent scientist in the field of plant demography, population biology and forest ecosystem ecology (Fig. 1).

Professor Olga V. Smirnova was born to the family of intellectual workers in 1939. Her mother, Nina Nikolaevna, was a French to Russian translator, and her father, Vsevolod Mikhailovich, was an engineer. Her maternal grandfather, Nikolai A. Zhukov, graduated from the Moscow Higher Vocational School of Commerce (now Financial Academy) and worked as an economist with the People’s Commissariat for Foreign Affairs. Her paternal grandfather, Mikhail I. Smirnov, graduated from the Moscow Archaeological Institute in Nizhny Novgorod. He was an outstanding professional who specialized in local history and founded the Pereslavl-Zalessky Historical, Architectural and Art Museum-Reserve in 1919. Olga V. Smirnova spent her childhood in Gagarinsky Lane in the very heart of Moscow (Zhukova, 2006).

She showed interest in biology in her high school years, attending a young naturalists hobby group at the All-Russian Society for the Protection of Nature (VOOP) headed by a renowned biologist Pyotr P. Smolin. In 1963, Olga V. Smirnova graduated from the Chair in Geobotany at M. V. Lomonosov Moscow State University. In 1968, she defended her Candidate of Sciences thesis under supervision of Professor Alexey A. Uranov that was devoted to the topic “Life cycles, number and age composition of populations of the main components of oak grass cover”. In 1983, she defended her Doctor of Sciences thesis devoted to the topic “Behavior of species and functional organization of grass cover of deciduous forests (a case study of plain broad-leaved forests in the European part of the USSR and linden forests of Siberia)”. From 1966 to 1992, Olga V. Smirnova had been employed with the Problem-Centered Biology Laboratory (PBL) at the V. I. Lenin Moscow State Pedagogical Institute (MGPI) (Fig. 2, 3). In 1987, she published the book Grass Cover Structure of Broad-Leaved Forests that resulted from her Candidate’s thesis and Doctor’s thesis. Since September 1, 1992, Professor Olga V. Smirnova has been employed as Principal Researcher at the Center of Forest Ecology and Productivity of the Russian Academy of Sciences (CEPF RAS). From 1993 to 2008, she taught at Pushchino State University (PuschGENI) at the Department of System Ecology, founded and headed by Professor Alexander S. Komarov (Fig. 4, 5). In 1994, a collective monograph Eastern European Broadleaf Forests, and in 2004, a two-volume book Eastern European Forests: History in Holocene and Contemporaneity were published under her editorship. In 2017, Springer published a revised version of this book, European Russian Forests: Their Current State and Features of Their History as requested by the Plant and Vegetation editorial board. The fundamentals of Professor Olga V. Smirnova’s research, starting with the young naturalists hobby group and to the present day, are her expeditionary studies she conducts every year (Fig. 6–8).

In 2015, Professor Olga V. Smirnova founded an international research journal, Russian Journal of Ecosystem Ecology. The journal covers the functioning and dynamics of ecosystems, the organization of biogeocenotic cover, and other issues of ecology. Professor Olga V. Smirnova supervised 25 successfully defended Candidate’s theses (Sugorkina, 1989; Evstigneev, 1990; Argunova, 1993; Istomina, 1993; Korotkov, 1993; Nedoseko, 1993; Chumachenko, 1993; Kiseleva, 1994; Shanijazova, 1994; Barinova, 1997; Ripa, 1997; Samohina, 1997; Sarycheva, 2000; Bobrovskaja, 2001; Braslavskaja, 2001; Turubanova, 2002; Bobrovskij, 2004; Shestakova, 2005; Bogdanova, 2006; Romanovskij, 2006; Lugovaja, 2008; Popov, 2008; Aleinikov, 2010; Zaprudina, 2012; Kharitonenkov, 2012); five of her students were awarded the degree of Doctors of Sciences (Chumachenko, 2006; Argunova, 2010; Evstigneev, 2010; Bobrovskij, 2013; Nedoseko, 2018). On December 2, 1994, she was awarded the title of Full Professor in Botany. To date, Professor Olga V. Smirnova has published over 300 works. All her publications, including an extensive reference list, are available online at http://istina.msu.ru/profile/sov1933/.

Professor Olga V. Smirnova’s views in forest biogeocenology / forest ecosystem ecology are based on ideas about interacting populations of living beings, which were shaped under the influence of her mentor, Professor Alexey A. Uranov (Shorina et al., 2014). Within the framework of said system of views, Professor Olga V. Smirnova made a significant contribution to enhancing the concepts of plant biological age and plant population strategy by developing the doctrine of coenopopopulations and biogeocenosis as a system of interacting populations, as well as to ideas about modern zonality as an anthropogenic phenomenon.

 

The Concept of Plant Biological Age

Classifying plant populations into ontogenetic (age) groups constitutes the basis of population demographics studies. The works of Tikhon A. Rabotnov (1950) and his followers, including Professor Olga V. Smirnova, substantiated and developed an approach to age differentiation in individual plants based on studying the ontogeny of living organisms from birth to death. The method provides for the allocation of stages in individual plant development, or ontogenetic states that reflect the biological age of an individual plant. Professor Olga V. Smirnova studied the ontogeny of over 30 plant species growing in Eastern European forests and linden trees of Western Siberia (Table 1). She presented early descriptions of the ontogeny in three collective monographs edited by Professor Alexey A. Uranov, Ontogenesis and Age Composition of Flowering Plant Populations (1967), Morphogenesis of Flowering Plants and The Structure of Their Populations (1968) and Age Composition of Flowering Plant Populations Due to Their Ontogenesis (1974).

These works show that defining the ontogenetic (age) state is incomparably more important for demographic research than the analysis of numerical age. This is due to two reasons: 1) different individual plants of the same species often reach the same ontogenetic state in different numerical terms; yet, since they are at the same development stage, they play the same part in the population and in the community; 2) the time individual plants of different species and life forms take to go through the same ontogenetic states may vary. All this means that it may be more logical to associate comparative assessment of significance of plants in coenosis with not numerical age but development stage, that is, the ontogenetic state.

Table 1. Ontogeny of plants studied by Professor Olga V. Smirnova

 

The Concept of Plant Population Strategy

Based on the system of ideas about plant coenotypes proposed by Leonty G. Ramensky (1935) and the concept of plant strategies developed by J. Grime (1979), Professor Olga V. Smirnova substantiated a new approach to studying plant population strategies (population behavior) (Smirnova, 1980, 1987; Smirnova, Chistyakova, 1980). The essential provisions of this approach are as follows.

1. The following properties are considered as integral, phytocoenotically significant plant population strategies: competitive, phytocoenotically tolerant, and reactive. Competitive strategy (violent, competitive power) means the ability of species to create and control the environment in a community, as well as suppress other living organisms due to the great vitality and highly intensive environment use. Phytocoenotically tolerant strategy (patient, resistance, endurance in an extremely unfavorable phytocoenotic environment) means the ability of species to survive for a long time in the area occupied by other living organisms, due to the maximum lowered vitality. Reactive strategy (explerent, dynamism, pioneering, ruderality) means the tendency of a species to the fastest possible development of released resources in the community due to vigorous vegetative growth and significant reproductive effort.

According to said definitions, a plant population strategy means the ability of species to dominate or occupy a subordinate position in a community, which resulted from a long evolution in preagricultural climax coenoses undisturbed by humans. The described population strategies reflect the phytocoenotic potencies of a species. The real-life ranking of a species in a particular coenosis makes up its phytocoenotic position. Phytocoenotic potencies and phytocoenotic positions may be expected to overlap completely in climax communities in the preagricultural age. Real-life position of a species in modern communities differ significantly from its role in climax coenoses, since the communities structure has been fundamentally transformed by humans.

2. Integral properties (competitive, phytocoenotically tolerant, reactive) are inherent in every species but expressed to various degrees. Species that are predominantly competitive are violent, species that are predominantly tolerant are patient, and species that are predominantly ruderal are explerent. Aside from groups of species characterized by these three strategy types (behavior), following J. Grime (1979), Professor Olga V. Smirnova identified the groups of species that occupy an intermediate position.
3. Investigation of a plant strategy is based on investigation of biological properties of a species. This assumes a differentiated approach to studying biological (behavior) and ecosystem properties of a species. Knowing the ecosystem properties, one could reveal the requirements of a species to environment resources, whereas studying biological properties could help to define the method and nature of how to use said resources. In other words, plant ecosystem properties determine the species composition in a community, whereas biological properties determine the predominant or subordinate role of the species in the community. This approach to identifying types of plant behavior differs significantly from the methods of studying the coenotypes proposed by Leonty G. Ramensky (1935) and plant strategies developed by J. Grime (1979). Thus, Ramensky’s patient plants (or Grime’s stress-tolerant plants) are allocated based on ecosystem properties of species, while violent (competitive) plants and explerent (reactive) plants are allocated based on biological properties.
4. The history of studying phytocoenotic potencies in plants shows that it is not possible to single out any standalone, independent feature that would determine the type of plant behavior in its entirety. At the same time, each type of plant behavior is characterized by a set of particular (differential) properties, which are specific manifestations of competitive, tolerant and reactive strategy.
5. It may be advisable to analyze the types of behavior in plants of relative life forms occupying the same space-time niche and belonging to the same trophic level, i. e. the same synusia (Smirnova, 1987). This is determined by the fact that the types of a single synusia are characterized by similar impact on the environment and play a similar role in the community. Moreover, the biological originality of a given species is most fully manifested if we investigate the entire historically formed set of species at the same time. In temperate forests, synusias of trees, shrubs, summer broadleaf herbs, and early spring ephemeroids are usually considered as such (Eastern European …, 1994).

Based on these provisions and a detailed study of the biological plant properties, Professor Olga V. Smirnova developed a classification of plant species by strategy (behavior) type in the synusias of spring ephemeroids and summer broadleaf herbs (Smirnova, 1987). This approach was successfully implemented when studying the types of strategy of trees and shrubs in Eastern European forests (Smirnova, Chistyakova, 1980; Evstigneev, 2004, 2010; Evstigneev, Didenko, 2004). Below is an example of a plant classification by type of behavior in the synusia of summer broadleaf herbs (Smirnova, 1987).

Type I. Competitive species (violent).

Group 1 — vegetatively mobile: Aegopodium podagraria L., Convallaria majalis L., Carex pilosa Scop., Mercurialis perennis L.

Type II. Tolerant species (patient).

Group 1 — vegetatively slightly mobile: Asarum europaeum L., Carex digitata L., C. rhizina Blytt ex Lindbl., Paris quadrifolia L., Polygonatum multiflorum (L.) All., Pulmonaria obscura Dumort., Viola mirabilis L.

Group 2 — vegetatively immobile: Brachypodium sylvaticum (Huds.) Beauv., Bromopsis benekenii (Lange) Holub, Carex sylvatica Huds., Campanula latifolia L., C. rapunculoides L., C. trachelium L., Dactylis glomerata L., Festuca gigantea (L.) Vill., F. sylvatica L., Geum urbanum L., Melica nutans L., Lathyrus vernus (L.) Bernh., Poa nemoralis L., Ranunculus cassubicus L., Scrophularia nodosa L., Scutellaria altissima L.

Type III. Reactive species (explerent).

Subtype 1: competitive reactive species. Group 1 — vegetatively mobile: Ajuga genevensis L., A. reptans L., Galeobdolon luteum Huds., Milium effusum L., Viola odorata L. Group 2 — vegetatively immobile: Lamium maculatum (L.) L.

Subtype 2: Actually reactive. Group 1. Vegetatively mobile: Galium odoratum (L.) Scop., Glechoma hederacea L., Stachys sylvatica L., Stellaria holostea L., Urtica dioica L. Group 2: vegetatively immobile: Alliaria petiolata (Bieb.) Cavara & Grande, Chaerophyllum temulum L., Geranium robertianum L., Torilis japonica (Houtt.) DC.

Professor Olga V. Smirnova showed that defining phytocoenotic potencies in plants makes it possible to understand some features in the organization of climax coenoses which differed in maximum species diversity (Smirnova, Chistyakova, 1980; Smirnova, 1983, 1987). Competitive plant species constituted a stable basis for every synusia, for they were predominant in number and biomass, involved the largest portion of matter and energy in the community, significantly changed the coenotic environment, and executed the function of edificators. Tolerant plant species, having an extremely low vitality level, used resources that could not be occupied by competitively powerful plants. Reactive plant species “roamed around” from one disturbance to another and “patched the holes” that occasionally occurred in the community in areas where individuals died in populations of edificators. Species with different strategy (behavior) types act as complementary formations, thanks to which the community resources are used most efficiently.

The Theory of Coenopopulations

The team at the Problem-Centered Biology Laboratory at the Moscow State Pedagogical Institute where Professor Olga V. Smirnova was employed at the time, published four outstanding books on plant demography, Plant Coenopopulations (Basic Concepts and Structure) (1976), Plant Coenopopulations (Development and Relationships) (1977), Dynamics of Plant Coenopopulations (1985), and Plant Coenopopulations (Essays on Population Biology) (1988). The books are based on ideas of plant biological age. These monographs present the concept apparatus and propose a system of methods in plant population biology. Having summarized her long-term studies at the Problem-Centered Biology Laboratory and Chair in Botany at the Moscow State Pedagogical Institute, Professor Olga V. Smirnova, in collaboration with Lyudmila B. Zaugol’nova, developed the ideas of characteristic ontogenetic spectrum (COS) and elementary demographic unit (EDU).

COS is a full-membered ontogenetic spectrum with a certain ratio of ontogenetic group number, which allows for a continuous generational turnover. This spectrum is due to the plant biological properties: 1) total duration of ontogeny and individual age states; 2) rate of development in individuals having various ontogenetic states; 3) methods of population self-sustainment; 4) intensity and frequency of inspermation and elimination; 5) ability to create a soil reserve of seeds or other vegetative rudiments; 6) area of resource consumption by individuals at different stages of ontogeny (Zaugol’nova, 1994; Zaugol’nova, Smirnova, 1978; Smirnova, 1987; Zaugol’nova et al., 1992; Eastern European …, 1994, 2004). At first, in the framework of studies conducted by said researchers, COS was considered synonymous with “basic ontogenetic spectrum”. However, the authors later limited the concept of the basic ontogenetic spectrum with the modal one, obtained by averaging data on several coenopopulations belonging to one community variant (Smirnova et al., 1993; Zaugol’nova, 1994).

 

 

COS reflects the dynamically stable (definitive) coenopopulation state to which it returns from deviations caused by external influences. The real ontogenetic spectrum is most consistent with the COS in undisturbed (climax) communities. In human-transformed coenoses, the ontogenetic spectrum of a population generally deviates from COS to varying degrees (Coenopopopulations…, 1976; Smirnova et al., 1987, 1989, 1990, 1991, 1992; Eastern European …, 1994).

Professor Olga V. Smirnova showed that plants have three types of characteristic ontogenetic spectra (Smirnova, 1987; Eastern European …, 1994). The first type is left-sided spectrum, with the maximum falling on pregenerative individuals. It is found in trees, monocarpic and oligocarpic tap-root herbs, bulbous, tuber-bulbous, and tuberous geophytes. These plants actively propagate by seed and/or deep rejuvenated vegetative rudiments. The second type is a centered spectrum: the largest number of individuals are located on middle-aged generative plants. It is characteristic of tap-rooted, long- and short-rhizomed herbs, sod grasses, and semi-shrubs. They have a weakly expressed aging period, propagate by seed or have a mixed propagation type, their vegetative reproduction is not accompanied by deep rejuvenation. The third type is a bimodal spectrum with two maxima: one in young individuals, and the other in mature or old generative individuals. This type is described in dense and loose sod grasses, tap-rooted and short-rhizomed herbs, in semi-shrubs. These plants have a significant life expectancy with a well-defined period of aging, their active propagation by seed is combined with vegetative reproduction with no deep rejuvenation.

EDU is a population unit, which is a set of individuals of different ages of the same species, necessary and sufficient to ensure sustainable generational turnover in the minimum allowable area. Important EDU characteristics include: 1) minimum number of individuals which allows for a continuous generational turnover; 2) minimum space necessary for a steady flow of generations; 3) lifetime of one generation (Smirnova et al., 1989; Zaugol’nova et al., 1993). EDU of different types are arranged in continuous series by values of each of the listed features (Table 2).

Table 2. Some parameters of elementary demographic units (EDU) in plants growing in broad-leaved forests (Smirnova et al., 1992)

The idea of EDU allowed Professor Olga V. Smirnova to propose a deeper definition of an important concept in forest ecology, namely the edificator (Smirnova, 1998; Smirnova, Toropova, 2008). This category includes species with the largest EDU and population mosaics existing for a long time. They include the largest portion of matter and energy in the cycles of generational turnover. Edificators belong to powerful environment converters. Populations of edificators with spontaneous development may transform a habitat to the greatest extent: they can change the hydrology, temperature, and lighting regime of a community, create micro- and mesorelief, and transform the soil cover. The intrinsic heterogeneity of the edificator EDU habitat allows for the coexistence of ecologically and biologically diverse species with smaller EDUs, and also maintains a high biodiversity level. The concept of edificator is synonymous with those of keystone species and ecosystem engineer. In the forest zone, edificators include species of various trophic groups and systematic positions: for example, large trees, needle- and leaf-eating insects, and tree-destroying fungi; river beavers join them in floodplain communities.

The Concept of Biogeocenosis as a System of Interacting Populations

Professor Olga V. Smirnova has been actively developing an idea of the structure and dynamics of undisturbed (climax) forest biogeocoenoses that existed in the preagricultural age with no human intervention (East European …, 1994, 2004; Smirnova et al., 1988, 1989, 1990; Smirnova, 1998, 2000; Smirnova, Toropova, 2008). This understanding is based on a population view of the community and biogeocenotic cover. According to the concept of biogeocenosis as a system of interacting populations, the forest cover should be considered a hierarchy of population units in species of different trophic groups. The population life of edificators unites this multiscale mosaic into communities. Population mosaics of keystone species create an environment suitable for sustainable life of populations of many subordinate species and determine the maximum species diversity in communities.

Phytogenic mosaicism in undisturbed forests results from the population life of edificator trees. In forests, the population life of trees creates a mosaic of lighting, water and soil regimes. This mosaic results from gaps in tree canopy that occur due to aging and death of one or several trees growing nearby. The death of a tree and associated soil perturbation determine the development of wind-soil complexes. At the same time, it builds a specific dumping microrelief, including hills, depressions, and coarse woody debris (Bobrovskij, 2004, 2013). Heterogeneous gap-like environment and wind-soil complexes, created as a result of generation flows in edificator tree populations, determines the presence of the maximum possible set of subordinate species of plants, animals, fungi, and representatives of other kingdoms in undisturbed forests. Having studied said mosaicism, Professor Olga V. Smirnova and her students created a new system of understanding forest ecology, the gap paradigm (Korotkov, 1991, 1993; Smirnova, 1998; Assessment …, 2000).

Professor Olga V. Smirnova convincingly shows that mosaicism caused by vitality of animal phytophages is as characteristic a feature of forest landscapes as phytogenic one (Smirnova et al., 1993; Eastern European …, 1994, 2004). Zoogenic mosaicism in preagricultural forests resulted from population life of animal edificators. In undisturbed European forests, these animals included: 1) large herd ungulates (bison, auroch, tarpan, etc.); 2) leaf- and needle-eating insects; 3) beavers. Large herd ungulates that destroyed young trees, shrubs, and grasses, as well as compacted and cherished the soil, created zoogenic glades with meadow, forest margin, and meadow-steppe flora. By destroying leaves and needles, insects increase the illumination on the grass cover surface and the temperature of the air and soil, enrich the soil with nitrogen and other minerals, and also contribute to an increase in number of light-loving and nitrophilic types of herbs. Beavers, by building dams on streams and small rivers, create ponds and lowland swamps, increase species diversity and the number of related plant and animal species. By destroying trees and shrubs in the coastal strip, beavers form glades with light-loving flora and fauna.

Professor Olga V. Smirnova shows that strong anthropogenic impact, destroying the population mosaic, breaks the cycles of generational turnover in keystone species. As a result, the development of communities becomes unidirectional, or succession, before the natural mosaic is restored. Understanding biogeocenosis as a system of interacting populations and a quantitative assessment of population parameters of principal coenosis builders make it possible to reconstruct the potential structure of biogeocenotic cover in an area, quantify the degree of disturbance in communities and their complexes, as well as to streamline existing succession systems.

The Concept of Anthropogenic Transformation of the Forest Cover in the Holocene

Despite the large number of works devoted to anthropogenic transformation of the forest cover in the Holocene, the paradigm of climate migration still prevails in domestic science. Summarizing historical and paleontological data, Professor Olga V. Smirnova proposed a new, “anthropic” system of views that defines human impact as the essential factor in biogeocoenotic cover transformation in the Holocene (Smirnova, Bobrovskij, 2000; Smirnova et al., 2001a, 2001b, 2006, 2013; Turubanova, 2002; Smirnova, Turubanova, 2003; Haritonenkov, 2012; Smirnova, Toropova, 2016; Kalyakin et al., 2016; Smirnova, Toropova, 2016; Smirnova et al., 2018). This new paradigm can be described as follows.

Written sources indicate that, over the recent 1–2 millennia, the diversity of life on Earth has been rapidly decreasing due to anthropogenic transformation. However, paleontological studies show that significant transformations on the Russian Plain and throughout Northern Eurasia date back much earlier. The first man-made environmental crisis in said area occurred 22–18 thousand years ago. It was caused by extermination of crucial edificators of late Pleistocene — that is, mammoths, woolly rhinoceros, giant deer, and other animals. They used to edify the composition and structure of plants and animals at that time. Herbs, primarily grasses, growing on meadow-steppe glades and forest margins were their primary source of nutrition. Meadow-steppe communities alternated with small clusters of trees. At the same time, palynology studies show that coniferous and broad-leaved tree species used to inhabit the entire Russian Plain in the Pleistocene. The soils of cryogenic savannah with a layer of permafrost underneath are chemically similar to modern chernozem soils. Their productivity throughout Northern Eurasia was so great that it allowed for sustainable existence of huge herds of giant phytophages and their retinue. Paleozoologists call this feature “the paradox of prehistoric pastures”. It was in cryogenic savannahs of the late Pleistocene (Upper or Final Paleolithic) that the sites of mammoth hunters with highly-developed farming and culture were found.

Over the recent 10.000–7.000 years, there has been a general climate warming, which coincided with gradual destruction of keystone species of giant and large animals of the mammoth complex as a result of hunting. The degradation of the mammoth complex, which began in the late Pleistocene, led to woody plants strengthening their role. The pastures the mammoths and their companions used to graze were then inhabited by trees. The first trees appeared by volatile seeds and a rapid generational turnover: that were birch, willow, aspen, and pine trees. They were followed by dark coniferous (spruce, fir) and broad-leaved (oak, linden, maple, ash, beech, hornbeam, etc.) trees. The near extinction of the giants and largest phytophages in the mammoth complex, combined with warming, marked the beginning of development of a forest belt in the early Holocene at the site of former Pleistocene cryogenic savannahs.

At the beginning of the Middle Holocene (7.000–2.500 years ago), the forest belt was almost completely developed on the Russian plain with broad-leaved and dark coniferous trees being predominant species; it occupied the space from the northern to southern seas. Within the forest belt, due to transforming activity of bisons, aurochs, tarpans, the saiga and other animals, zoogenic glades with meadow and steppe plants occurred constantly. Beavers built settlements with wetlands along small watercourses. As a result, the biogeocenotic cover of the Middle Holocene was a set of forest-meadow-bog complexes, created and regulated by keystone species, that is, large herd ungulates, beavers, and trees.

Since the mid-Middle Holocene occurred the production economy as the most powerful factor of biogeocenotic cover impact (agriculture, cattle breeding, smelting). The osteological material of this time includes a greatly reduced ratio of bones of wild ungulates (bison, auroch, tarpan, etc.) and increased ratio of livestock bones, whereas pollen of cultivated grasses appeared in the spore-pollen spectra. The production economy changed the biogeocenotic cover structure fundamentally. First of all, large herd ungulates and beavers disappeared not only due to hunting, but also due to the radical transformation of their habitats under the influence of slash-and-burn agriculture, logging and other harvesting trades. With the destruction of keystone animal species, the ratio of natural meadow-steppe ecosystems decreased, and that of forest ecosystems increased. As a result, the life of light-loving tree species (primarily oak and pine), as well as all light-loving plant species of other life forms, and many animal species that had previously inhabited zoogenic glades, became possible only in anthropogenic habitats. For example, pioneer tree species regenerated mainly on abandoned arable land.

Relatively “independent” from humans remained the ecosystems of “shadow” coniferous/broad-leaved forests; their spontaneous development is possible even now under a natural reserve regime. However, only part of the region’s natural flora and fauna can sustainably exist in these communities.  Ecosystems representing the “fragments” of mid-Holocene forest-meadow-bog complexes have been preserved in the modern forest cover of the Russian Plain only as a small number of refugiums, untouched by strong anthropogenic transformation of recent centuries.

It is from the late mid-Holocene that it becomes fundamentally impossible to restore the potential (former) biogeocenotic cover in a spontaneous mode, since, on the one hand, the populations of keystone animal species (large herd ungulates and beavers) have greatly decreased in number, and, on the other hand, humans have become the most powerful environment transforming force. Human activity began to determine the existence of certain subordinate plant and animal species.

By the late mid-Holocene, forest burning for the slash-and-burn agriculture cycle pushed the southern border of the forest belt to the north. The spread of nomadic cattle breeding in the south of the Russian Plain resulted in the formation of steppe and semi-desert-steppe zones. These events were a major step towards modern zonality and probably had a significant impact on changes in the macroclimate of Eurasia in its entirety. They maybe were a reason for the growing climate instability in the second half of the Holocene.

From Iron Age to early Middle Ages (2.500–500 years ago), the northern borders of the ranges of broad-leaved tree species significantly retreated to the south, mainly due to slash-and-burn agriculture, which marked the beginning of the modern taiga — a forest strip where said tree species do not exist. At the same time, specific pyrogenic forests with predominance of Pinus sylvestris developed on the sandy soils of the forest belt. Slash-and-burn, and then cross-bed and arable agriculture, forest grazing, gathering litter and coarse woody debris as well as other kinds of forest use resulted in soil cover degradation in large areas. Forest burning on the northern border was the reason why the tundra zone developed from the forest tundra and northern taiga in the late Holocene.

In general, the production economy of the mid- and late Holocene divided the unified forest-meadow-bog complex into two groups: 1) ecosystems capable of supporting themselves with spontaneous development (“shadow” forests), which formed the forest belt itself; 2) ecosystems that require constant anthropogenic impact for life (floodplain and land meadows, meadow steppes, forests of pioneer tree species). At the same time, there was a final step in formation of anthropogenic zonality — under human impact, the unified forest belt on the Russian Plain divided into coniferous, coniferous/broad-leaved and broad-leaved forests.

 CONCLUSION

Professor Olga V. Smirnova’s edifice is based on ideas about the leading role of plant and animal populations in the organization of the biogeocenotic cover. In this case, it is implied that a continuous generational turnover in edificator (keystone species) populations is necessary to maintain the species and structural diversity in communities and ensure their sustainability. This system of views was influenced by Professor Alexey A. Uranov. The development of these ideas was consistent and gradual. First, Professor Olga V. Smirnova studied the biology of different plant species life forms. Examining their individual development, with identification of ontogenetic stages, is necessary for demographic research. She then developed the theory of coenopopulations as supraorganismal systems, which can self-sustain under different conditions. Finally, she developed the doctrine of biogeocenosis as a system of interacting populations and created the concept of anthropogenic transformation of the forest cover in the Holocene. Her contributions helped researchers to understand the mechanisms of the formation of modern zonality that are due to human activity.

We would like to congratulate Professor Olga V. Smirnova on her anniversary! We wish her a long life full of success in her new creative endeavors, vigor, energy, and good health! Her colleagues and students, as well as Forest Science Issues editorial board, join the congratulation.

 ACKNOWLEDGMENTS

The authors thank Oleg G. Barinov, Maria A. Barinova, Maxim V. Bobrovskij, Natalia N. Bogomolova, Alexey V. Gornov, Elena S. Esipova, Ekaterina L. Zheleznaya, Ekaterina A. Kobozeva, Svetlana I. Ripa, Mikhail S. Romanov, Andrey M. Romanovskij, Elena P. Sarycheva, Nikolay S. Smirnov and Larisa G. Khanina for their help in preparing this article.

REFERENCES

Aleinikov A. A., Sostojanie populjacii i sredopreobrazujushhaja dejatel’nost’ bobra evropejskogo na territorii zapovednika “Brjanskij les” i ego ohrannoj zony: avtoref. dis. … kand. biol. nauk (Population status and environmental activity of the European beaver on the territory of the reserve “Bryansk forest” and its protection zone. Extended abstract of Candidate’s thesis), Tol’jatti: In-t jekologii Volzhskogo bassejna RAN, 2010, 22 p.

Argunova M. V., Ekologicheskoe obrazovanie v interesah ustojchivogo razvitija kak nadpredmetnoe napravlenie modernizacii shkol’nogo obrazovanija: avtoref. dis. … d-ra. ped. nauk (Environmental education for sustainable development as a non-objective direction of modernization of school education. Extended abstract of Doctor’s thesis), Moscow: Mosk. gos. obl. un-t, 2010, 47 p.

Argunova M. V., Populjacionnaja organizacija dubovo-grabovyh lesov Zapadnoj Ukrainy i optimizacija ih struktury: avtoref. dis. … kand. biol. nauk (Population organization of oak-hornbeam forests of Western Ukraine and optimization of their structure, Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1993, 16 p.

Barinova M. A., Vlijanie vodohranilishha na sinuzii zelenyh mhov doliny Giljuja: avtoref. dis. … kand. biol. nauk (The reservoir effect on sinusoi green mosses of the valley Gilya. Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1997, 16 p.

Bobrovskaja N. E., Formirovanie struktury kron listvennyh i hvojnyh derev’ev v ontogeneze: avtoref. dis. … kand. biol. nauk (Formation of crown structure of deciduous and coniferous trees in ontogenesis, Extended abstract of Candidate’s thesis), Moscow: MGPU, 2001, 19 p.

Bobrovskij M. V., Bioticheskie i antropogennye faktory dolgovremennoj dinamiki lesnyh pochv Evropejskoj Rossii: avtoref. dis. … d-ra biol. nauk (Biotic and anthropogenic factors of long-term dynamics of forest soils in European Russia. Extended abstract of Doctor’s thesis), Vladimir: im. A. G. i N. G. Stoletovyh, 2013, 48 p.

Bobrovskij M. V., Raznoobrazie rastitel’nosti i pochv zapovednika “Kaluzhskie zaseki” i ego svjaz’ s tradicionnym prirodopol’zovaniem: avtoref. dis. … kand. biol. nauk (Diversity of vegetation and soils of the reserve “Kaluga Zaseki” and its connection with traditional nature management. Extended abstract of Candidate’s thesis), Moscow: MGPU, 2004, 23 p.

Bogdanova N. G., Formirovanie travjanogo pokrova hvojno-shirokolistvennyh i shirokolistvennyh lesov v hode vosstanovitel’nyh sukcessij v Nerusso-Desnjanskom poles’e: avtoref. dis. … kand. biol. nauk (The formation of grass cover coniferous and deciduous forests during the restoration successions in Nerussa-Desna Polesye), Moscow: MGPU, 2006, 21 p.

Braslavskaja T. Ju., Biologicheskoe raznoobrazie i dinamika rastitel’nosti v pojme maloj reki Juzhnogo Nechernozem’ja (na primere r. Nerussa, Brjanskaja obl.): avtoref. dis. … kand. biol. nauk (Biological diversity and vegetation dynamics in the floodplain of the small river of the southern non-Chernozem region (by the example of the Neruss river, Bryansk region), Extended abstract of Candidate’s thesis), Moscow: MGPU, 2001, 21 p.

Cenopopuljacii rastenij (ocherki populjacionnoj biologii) (Plant coenopopulations (essays on population biology)), Moscow: Nauka, 1988, 184 p.

Cenopopuljacii rastenij (osnovnye ponjatija i struktura) (Plant coenopopulations (basic concepts and structure)), Moscow: Nauka, 1976, 217 p.

Cenopopuljacii rastenij (razvitie i vzaimootnoshenija) (Plant coenopopulations (development and relationships)), Moscow: Nauka, 1977, 135 p.

Chumachenko S. I., Biojekologicheskaja model’ raznovozrastnogo lesnogo cenoza: avtoref. dis. … kand. biol. nauk (Bioecological model of uneven-aged forest cenosis. Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1993, 16 p.

Chumachenko S. I., Imitacionnoe modelirovanie mnogovidovyh raznovozrastnyh lesnyh nasazhdenij: avtoref. dis. … d-ra biol. nauk (Simulation modeling of multi-species multi-age forest stands. Extended abstract of Doctor’s thesis), Moscow: Mosk. gos. u-t lesa, 2006, 32 p.

Dinamika cenopopuljacij rastenij (Dynamics of plant coenopopulations), Moscow: Nauka, 1985, 208 p.

European Russian Forests. Their Current State and Features of Their History, Smirnova O., Bobrovskij M., Khanina L. (eds.). Dordrecht: Springer Nature, 2017, 566 p.

Evstigneev O. I., Didenko E. G., Populjacionnye strategii vidov kustarnikov (Population strategies of shrub species), Vostochnoevropejskie lesa: istorija v golocene i sovremennost’ (Eastern European forests: history in Holocene and contemporaneity), Vol. 2, Moscow: Nauka, 2004, pp. 205–224.

Evstigneev O. I., Fitocenotipy i otnoshenie listvennyh derev’ev k svetu: avtoref. dis. … kand. biol. nauk (Fitocenotype and the relation of deciduous trees to light. Extended abstract of Candidate’s thesis), Moscow: MGPI, 1990, 17 p.

Evstigneev O. I., Mehanizmy podderzhanija biologicheskogo raznoobrazija lesnyh biogeocenozov: avtoref. dis. … d-ra biol. nauk (Mechanisms of maintenance of biological diversity of forest biogeocenoses. Extended abstract of Doctor’s thesis), Nizhnij Novgorod: NNGU, 2010, 48 p.

Evstigneev O. I., Populjacionnye strategii vidov derev’ev (Population strategies of tree species), Vostochnoevropejskie lesa: istorija v golocene i sovremennost’, Moscow: Nauka, 2004, Vol. 1, pp. 176–205.

Grime J. P., Plant strategies and vegetation processes, N. Y., 1979, 222 p.

Istomina I. I., Kvazisenil’nost’ i ee rol’ v zhizni drevesnyh rastenij: avtoref. dis. … kand. biol. nauk (Quasi-senility and its role in the life of woody plants. Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1993, 16 p.

Kharitonenkov M. A., Rol’ antropogennogo faktora v formirovanii rastitel’nogo pokrova juga Zapadno-Sibirskoj ravniny v jepohu tradicionnogo prirodopol’zovanija (s pozdnego paleolita do konca XIX v.): avtoref. dis. … kand. biol. nauk (The role of anthropogenic factor in the formation of vegetation cover of the South of the West Siberian plain in the era of traditional nature management (from the late Paleolithic to the end of the XIX century), Extended abstract of Candidate’s thesis), Moscow: Mosk. gos. obl. un-t, 2012, 25 p.

Kalyakin V. N., Turubanova S. A., Smirnova O. V., The origin and development of the East European taiga in late Cenozoic, Russian Journal of Ecosystem Ecology, 2016, Vol. 1, No 1, pp. 1–26, DOI: 10.21685/2500-0578-2016-1-2.

Kiseleva L. L., Jekologo-floristicheskij analiz jekotonnyh soobshhestv central’noj lesostepi: avtoref. dis. … kand. biol. nauk (Ecological and floristic analysis of ecotonic communities of the Central forest-steppe. Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1994, 16 p.

Korotkov V. N., Demutacionnye processy v ostrovnyh lesnyh massivah (na primere GIZL “Gorki Leninskie” i Kanevskogo zapovednika): avtoref. dis. … kand. biol. nauk (Demutation processes in island forests (on the example GISL Gorki Leninskie and Kanev nature reserve). Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1993, 16 p.

Korotkov V. N., Novaja paradigma v lesnoj jekologii (New paradigm in forest ecology), Biologicheskie nauki, 1991, No 8, pp. 7–20.

Lugovaja D. L., Rol’ jekotopicheskih i antropogennyh faktorov v formirovanii vidovogo i strukturnogo raznoobrazija juzhnotaezhnyh lesov (vostok Kostromskoj oblasti): avtoref. dis. … kand. biol. nauk (The role of ecotopic and anthropogenic factors in the formation of species and structural diversity of South taiga forests (East of Kostroma region), Extended abstract of Candidate’s thesis), Moscow: MGPU, 2008, 28 p.

Nedoseko O. I., Ontomorfogenez Salix pentandra L., Salix caprea L., Salix cinerea L.: avtoref. dis. … kand. biol. nauk (The ontomorphogenesis Salix pentandra L., Salix caprea L., Salix cinerea L. Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1993, 16 p.

Nedoseko O. I., Stanovlenie zhiznennyh form i arhitektoniki kron boreal’nyh vidov iv podrodov Salix i Vetrix Dumort. v ontogeneze: avtoref. dis. … d-ra biol. nauk (Formation of life forms and architectonics of crowns of boreal species of willows of subgenera Salix and Vetrix Dumort. in ontogenesis. Extended abstract of Doctor’s thesis), Moscow: MSHA im. K. A. Timirjazeva, 2018, 43 p.

Ocenka i sohranenija bioraznoobrazija lesnogo pokrova v zapovednikah Evropejskoj Rossii (Assessment and conservation of forest cover biodiversity in reserves of European Russia), Moscow: Nauchnyj mir, 2000, 196 p.

Ontogenez i vozrastnoj sostav populjacij cvetkovyh rastenij (Ontogenesis and age composition of flowering plant populations), Moscow: Nauka, 1967, 156 p.

Popov S. Ju., Struktura i dinamika rastitel’nosti Kerzhenskogo zapovednika: avtoref. dis. … kand. biol. nauk (Structure and dynamics of vegetation of Kerzhensky reserve. Extended abstract of Candidate’s thesis), Moscow: MGPU, 2008, 20 p.

Rabotnov T. A., Zhiznennyj cikl mnogoletnih travjanistyh rastenij v lugovyh cenozah (Life cycle of perennial herbaceous plants in meadow cenoses), Trudy BIN ANSSSR, Ser. 3, Geobotanika, 1950, Vol. 6, pp. 7–204.

Ramenskij L. G., O principial’nyh ustanovkah, osnovnyh ponjatijah i terminah proizvodstvennoj tipologii zemel’, geobotaniki i jekologii (On fundamental principles, basic concepts and terms of production typology of land, geobotany and ecology), Sovetskaja botanika, 1935, No 4, pp. 25–41.

Ripa S. I., Populjacionno-cenoticheskij analiz gornyh bukovyh i smeshannyh lesov Ukrainskih Karpat: avtoref. dis. … kand. biol. nauk (Population-cenotic analysis of mountain beech and mixed forests of the Ukrainian Carpathians. Extended abstract of Candidate’s thesis), Moscow: MGPU im. V. I. Lenina, 1997, 16 p.

Romanovskij A. M., Osobennosti ontogeneza i fitocenoticheekaja rol’ eli evropejskoj v lesah Nerusso-Desnjanskogo poles’ja: avtoref. dis. … kand. biol. nauk (Features of ontogenesis and phytocenotic role of spruce in the forests Nerussa-Desna Polesye. Extended abstract of Candidate’s thesis), Moscow: MGPU, 2006, 18 p.

Samohina T. Ju., Struktura i spontannaja dinamika hvojno-shirokolistvennyh lesov Srednego Urala: avtoref. dis. … kand. biol. nauk (Structure and spontaneous dynamics of coniferous broad-leaved forests of the Middle Urals. Extended abstract of Candidate’s thesis), M.: MGPU im. V. I. Lenina, 1997, 16 p.

Sarycheva E. P., Strukturnoe i vidovoe raznoobrazie chernool’hovyh lesov centra evropejskoj Rossii (na primere zapovednikov “Brjanskij les” i “Voroninskij”): avtoref. dis. … kand. biol. nauk (Structural and species diversity of black alder forests in the center of European Russia (on the example of reserves “Bryansk forest” and “Voroninsky”) Extended abstract of Candidate’s thesis), M.: MGPU, 2000, 16 p.

Shanijazova Z. P., Populjacionnaja biologija jefemerov Karakalpakii: avtoref. dis. … kand. biol. nauk (Population biology of the ephemera of Karakalpakstan. Extended abstract of Candidate’s thesis), Moscow: MGPU, 1994, 16 p.

Shestakova A. A., Jekologo-cenoticheskie i floristicheskie osobennosti organizacii briobioty na territorii Nizhegorodskoj oblasti: avtoref. dis. … kand. biol. nauk (Ecological-cenotic and floristic features of the organization of mosses in the Nizhny Novgorod region. Extended abstract of Candidate’s thesis), Nizhnij Novgorod: NNGU im. N. I. Lobachevskogo, 2005, 28 p.

Shorina N. I., Kurchenko E. I., Grigor’eva N. M., Aleksej Aleksandrovich Uranov (1901–1974) (Alexei Alexandrovich Uranov, 1901–1974), Samarskaja Luka: problemy regional’noj i global’noj jekologii, 2014, Vol. 23, No 1, pp. 93–129.

Smirnova O. V., Bakun E. Ju., Turubanova S. A., Predstavlenie o potencial’nom i vosstanovlennom rastitel’nom pokrove lesnogo pojasa Vostochnoj Evropy (Understanding of the potential and restored vegetation cover of the forest belt of Eastern Europe), Lesovedenie, 2006, No 1, pp. 22–33.

Smirnova O. V., Bobrovskij M. V., Vozdejstvie proizvodjashhego hozjajstva na sostav i strukturu lesnogo pokrova (The impact of the producing economy on the composition and structure of forest cover), Ocenka i sohranenija bioraznoobrazija lesnogo pokrova v zapovednikah Evropejskoj Rossii (Assessment and conservation of forest cover biodiversity in reserves of European Russia), Moscow: Nauchnyj mir, 2000, pp. 22–26.

Smirnova O. V., Chistjakova A. A., Analiz fitocenoticheskih potencij nekotoryh drevesnyh vidov shirokolistvennyh lesov Evropejskoj chasti SSSR (Analysis of phytocenotic potencies of some woody species of broad-leaved forests of the European part of the USSR), Zhurn. obshh. biol., 1980, Vol. 41, No 3, pp. 350–362.

Smirnova O. V., Chistjakova A. A., Drobysheva T. I., Cenopopuljacionnyj analiz i prognozy razvitija dubovo-grabovyh lesov Ukrainy (Cenopopulation analysis and forecasts of development of oak-hornbeam forests in Ukraine), Zhurn. obshh. biol., 1987, Vol. 48, No 2, pp. 200–212.

Smirnova O. V., Chistjakova A. A., Popadjuk R. V., Evstigneev O. I., Korotkov V. N., Mitrofanova M. V., Ponomarenko E. V., Populjacionnaja organizacija rastitel’nogo pokrova lesnyh territorij (na primere shirokolistvennyh lesov evropejskoj chasti SSSR) (Population organization of vegetation cover of forest areas (on the example of broad-leaved forests of the European part of the USSR)), Pushhino: ONTI Nauchnyj centr biol. issledovanij AN SSSR, 1990, 92 p.

Smirnova O. V., Chistjakova A. A., Popadjuk R. V., Populjacionnye mehanizmy dinamiki lesnyh cenozov (Population mechanisms of the dynamics of forest cenoses), Biologicheskie nauki, 1989, No 11, pp. 48–58.

Smirnova O. V., Chistjakova A. A., Ripa S. I., Lysyh N. I., Populjacionnaja organizacija bukovyh lesov Zakarpat’ja (Population organization of beech forests in Transcarpathia), Bjul. MOIP. Otd. biol., 1989, Vol. 94, No 5, pp. 78–91.

Smirnova O. V., Geraskina A. P., Aleinikov A. A., The concept “complementarity” as the basis for model and nature reconstruction of potential biota in the current climate, Russian Journal of Ecosystem Ecology, 2018, Vol. 3, No 3, pp. 1–21, DOI: 10.21685/2500-0578-2018-3-1.

Smirnova O. V., Kaljakin V. N., Turubanova S. A., Bobrovskij M. V., Sovremennaja zonal’nost’ Vostochnoj Evropy kak rezul’tat preobrazovanija pozdneplejstocenovogo kompleksa kljuchevyh vidov (Modern zoning of Eastern Europe as a result of transformation of the late Pleistocene complex of key species), Mamont i ego okruzhenie: 200 let izuchenija (Mammoth and its environment: 200 years of study), Moscow: Geos, 2001a, pp. 200–208.

Smirnova O. V., Lugovaja D. L., Prokazina T. S., Model’naja rekonstrukcija vosstanovlennogo lesnogo pokrova taezhnyh lesov (Model reconstruction of the restored forest cover of taiga forests), Uspehi sovrem. biol., 2013, Vol. 133, No 2, pp. 164–177.

Smirnova O. V., Popadjuk R. V., Chistjakova A. A., Populjacionnye metody opredelenija minimal’noj ploshhadi lesnogo cenoza (Population methods for determining the minimum area of forest cenosis), Bot. zhurn., 1988, Vol. 73, No 10, pp. 1423–1433.

Smirnova O. V., Popadjuk R. V., Janickaja T. O., Korotkov V. N., Puti vosstanovlenija populjacionnoj struktury i vidovogo raznoobrazija v lesnyh demutacionnyh kompleksah (Ways of restoration of population structure and species diversity in forest demutation complexes), Biologicheskie nauki, 1992, No 5, pp. 7–25.

Smirnova O. V., Populjacionnaja organizacija biocenoticheskogo pokrova lesnyh landshaftov (Population organization of biocenotic cover of forest landscapes), Uspehi sovrem. biol., 1998, Vol. 118, No 2, pp. 148–165.

Smirnova O. V., Populjacionnaja organizacija biogeocenoticheskogo pokrova lesnyh territorij (Population organization of biogeocenosis cover of forest territories), Ocenka i sohranenija bioraznoobrazija lesnogo pokrova v zapovednikah Evropejskoj Rossii (Assessment and conservation of forest cover biodiversity in reserves of European Russia), Moscow: Nauchnyj mir, 2000, pp. 14–22.

Smirnova O. V., Povedenie vidov i funkcional’naja organizacija travjanogo pokrova shirokolistvennyh lesov (na primere ravninnyh shirokolistvennyh lesov Evropejskoj chasti SSSR i lipnjakov Sibiri): dis. … d-ra biol. nauk (Behavior of species and functional organization of grass cover of broad-leaved forests (on the example of flat broad-leaved forests of the European part of the USSR and Linden forests of Siberia). Doctor’s thesis), Leningrad: LGU im. A. A. Zhdanova, 1983, 685 p.

Smirnova O. V., Povedenie vidov i funkcional’naja organizacija travjanogo pokrova shirokolistvennyh lesov Evropejskoj chasti SSSR (Behavior of species and functional organization of grass cover of deciduous forests of the European part of the USSR), Bjul. MOIP. Otd. biol., 1980, Vol. 85, No 5, pp. 53–67.

Smirnova O. V., Struktura travjanogo pokrova shirokolistvennyh lesov (Grass cover structure of broad-leaved forests), Moscow: Nauka, 1987, 208 p.

Smirnova O. V., Toropova N. A., Potencial’naja rastitel’nost’ i potencial’nyj jekosistemnyj pokrov (Potential vegetation and potential ecosystem cover), Uspehi sovrem. biol., 2016, Vol. 136. No 2, pp. 199–211.

Smirnova O. V., Toropova N. A., Potential ecosystem cover — a new approach to conservation biology, Russian Journal of Ecosystem Ecology, 2016, Vol. 1, No 1, pp. 1–16, DOI: 10.21685/2500-0578-2016-1-1.

Smirnova O. V., Toropova N. A., Sukcessija i klimaks kak jekosistemnyj process (Potential vegetation and potential ecosystem cover), Uspehi sovrem. biol., 2008, Vol. 128, No 2, pp. 129–144.

Smirnova O. V., Turubanova S. A., Bobrovskij M. V., Korotkov V. N., Hanina L. G., Rekonstrukcija istorii biocenoticheskogo pokrova Vostochnoj Evropy i problema podderzhanija biologicheskogo raznoobrazija (Reconstruction of the history of the biocenotic cover of Eastern Europe and the problem of maintaining biological diversity), Uspehi sovrem. biol., 2001b, No 2, pp. 144–159.

Smirnova O. V., Turubanova S. A., Formirovanie i razvitie Vostochnoevropejskih shirokolistvennyh lesov v golocene (Formation and development of Eastern European broadleaf forests in the Holocene), Bjul. MOIP. Otd. biol., 2003, Vol. 108, No 2, pp. 32–40.

Smirnova O. V., Voznjak R. R., Evstigneev O. I., Korotkov V. N., Nosach N. Ja., Popadjuk R. V., Samojlenko V. K., Toropova N. A., Populjacionnaja diagnostika i prognozy razvitija zapovednyh lesnyh massivov (na primere Kanevskogo zapovednika) (Population diagnostics and forecasts of development of protected forests (on the example of Kanev reserve)), Bot. zhurn., 1991, Vol. 76, No 6, pp. 860–871.

Smirnova O. V., Zaugol’nova L. A., Popadjuk R. V., Populjacionnaja koncepcija v biocenologii (Population concept in biotsenology), Zhurn. obshh. biol. 1993, Vol. 54, No 4, pp. 438–448.

Smirnova O. V., Zhiznennye cikly, chislennost’ i vozrastnoj sostav populjacij osnovnyh komponentov travjanogo pokrova dubrav: dis. … kand. biol. nauk (Life cycles, number and age composition of populations of the main components of oak grass cover. Candidate’s thesis), Moscow: MGPI im. V. I. Lenina, 1968, 285 p.

Smirnova Ol’ga Vsevolodovna, Intellektual’naya Sistema Tematicheskogo Issledovaniya Naukometricheskih dannyh (Intelligent System case Studies Scientometric data), available at: http://istina.msu.ru/profile/sov1933/ (September 14, 2019).

Sugorkina N. S., Ontogenez i osobennosti populjacionnoj biologii vidov roda geran’: avtoref. dis. … kand. biol. nauk (Ontogenesis and peculiarities of population biology of geranium species. Extended abstract of Candidate’s thesis), Moscow: MGPI, 1989, 17 p.

Turubanova S. A., Jekologicheskij scenarij istorii formirovanija zhivogo pokrova Evropejskoj Rossii i sopredel’nyh territorij na osnove rekonstrukcii arealov kljuchevyh vidov zhivotnyh i rastenij: avtoref. dis. … kand. biol. nauk (Ecological scenario of the history of the formation of the living cover of European Russia and adjacent territories on the basis of the reconstruction of the habitats of key species of animals and plants. Extended abstract of Candidate’s thesis), Moscow: In-t biol. Komi nauchn. centra Ural’skogo otd. RAN, 2002, 23 p.

Voprosy morfogeneza cvetkovyh rastenij i stroenija ih populjacij (Morphogenesis of flowering plants and the structure of their populations), Moscow: Nauka, 1968, 234 p.

Vostochnoevropejskie lesa: istorija v golocene i sovremennost’ (Eastern European forests: history in Holocene and contemporaneity), Moscow: Nauka, 2004a, Vol. 1, 479 p.

Vostochnoevropejskie lesa: istorija v golocene i sovremennost’ (Eastern European forests: history in Holocene and contemporaneity), Moscow: Nauka, 2004b, Vol. 2, 575 p.

Vostochnoevropejskie shirokolistvennye lesa (Eastern European broadleaf forests). Moscow: Nauka, 1994, 364 p.

Vozrastnoj sostav populjacij cvetkovyh rastenij v svjazi s ih ontogenezom (Age composition of flowering plant populations due to their ontogenesis), Moscow: MGPI im. V. I. Lenina, 1974, 216 p.

Zaprudina M. V., Mikromozaichnaja organizacija travjano-kustarnichkovogo i mohovogo pokrova srednetaezhnyh temnohvojnyh lesov Urala: avtoref. dis. … kand. biol. nauk (Micromosaic organization of grass-shrub and moss cover of middle taiga dark coniferous forests of the Urals. Extended abstract of Candidate’s thesis), Moscow: MGPU, M., 2012, 23 p.

Zaugol’nova L. A., Zhukova L. A., Popadjuk R. V., Smirnova O. V., Kriticheskoe sostojanie cenopopuljacij rastenij (The critical state of plant coenopopulations), Problemy ustojchivosti biologicheskih system, Moscow: Nauka, 1992, pp. 51–59.

Zaugol’nova L. B., Smirnova O. V., Komarov A. S., Hanina L. G., Monitoring fitopopuljacij (Monitoring of phytopopulation), Uspehi sovrem. biol. 1993, Vol. 113, No 4, pp. 402–414.

Zaugol’nova L. B., Smirnova O. V., Vozrastnaja struktura cenopopuljacij mnogoletnih rastenij i ee dinamika (Age structure of perennial plant coenopopulations and its dynamics), Zhurn. obshh. biol., 1978, Vol. 39, No 6, pp. 849–858.

Zaugol’nova L. B., Struktura cenopopuljacij semennyh rastenij i problemy ih monitoringa: dis. … d-ra biol. nauk (Structure of seed plant coenopopulations and problems of their monitoring. Doctor’s thesis), SPb: SPb gos. un-t, 1994, 70 p.

Zaugol’nova L. B., Svjaz’ vozrastnogo spektra cenopopuljacij s biologicheskimi svojstvami vida (Relation of age spectrum of coenopopulations with biological properties of a species), In: Vozrastnoj sostav populjacij cvetkovyh rastenij v svjazi s ih ontogenezom (Age composition of flowering plant populations due to their ontogenesis), Moscow: MGPI im. V. I. Lenina, 1974, pp. 38–55.

Zhukova L. A., Mir prinadlezhit optimistam (The world belongs to the optimists), Polivariantnost’ razvitija organizmov, populjacij i soobshhestv, Joshkar-Ola: Mar. gos. un-t, 2006, pp. 229–278.

СИСТЕМА ВЗГЛЯДОВ О. В. СМИРНОВОЙ В ЛЕСНОЙ БИОГЕОЦЕНОЛОГИИ

О. И. Евстигнеев^{1, 3*}, В. Н. Коротков²

 ¹Заповедник «Брянский лес», Россия, 242180, Брянская обл., ст. Нерусса

²Институт глобального климата и экологии имени академика Ю. А. Израэля, Россия, 107258, Москва, ул. Глебовская, 20Б

³Центр по проблемам экологии и продуктивности лесов РАН, Россия, 117997 Москва, ул. Профсоюзная, 84/32, стр. 14

*E-mail: quercus_eo@mail.ru

Поступила в редакцию: 16.09.2019

После рецензирования: 07.10.2019

Принята к печати: 07.10.2019

О. В. Смирнова — профессор, доктор биологических наук, крупный ученый в области демографии растений, популяционной биологии и лесной биогеоценологии. Система взглядов О. В. Смирновой построена на представлениях о ведущей роли популяций растений и животных в организации биогеоценотического покрова. При этом подразумевается, что непрерывный оборот поколений в популяциях эдификаторов (ключевых видов) — необходимое условие для поддержания видового и структурного разнообразия сообществ, а также для обеспечения их устойчивости. Эта система взглядов сформировалась под влиянием идей профессора А. А. Уранова. Развитие этих представлений было последовательным и постепенным. Сначала О. В. Смирнова изучает биологию видов растений разных жизненных форм, их индивидуальное развитие, в котором выделяет онтогенетические состояния, необходимые для демографических исследований, затем — разрабатывает теорию ценопопуляций как надорганизменных систем, способных к самоподдержанию в различных условиях, и, наконец, — развивает учение популяционной организации биогеоценозов и создает концепцию антропогенной трансформации лесного покрова в голоцене. Все это помогает понять механизмы формирования современной зональности, которые обусловлены деятельностью человека.

Ключевые слова: биологический возраст растений, популяционная стратегия растений, ценопопуляция, эдификатор, лесная биогеоценология, современная зональность, историческая экология

Reviewer: Candidate of Biological Sciences A. V. Gornov