{"id":4156,"date":"2021-03-30T09:54:34","date_gmt":"2021-03-30T06:54:34","guid":{"rendered":"https:\/\/jfsi.ru\/?p=4156"},"modified":"2021-04-05T11:35:25","modified_gmt":"2021-04-05T08:35:25","slug":"4-1-2021-ruchinskayagornov","status":"publish","type":"post","link":"https:\/\/jfsi.ru\/en\/4-1-2021-ruchinskayagornov\/","title":{"rendered":"INFLUENCE OF SINGLE TREES ON FLORISTIC DIVERSITY OF STEPPE MEADOWS AND POPULATION STRUCTURE OF SOME RARE PLANT SPECIES"},"content":{"rendered":"

\"\"<\/a>Original Russian Text \u00a9 2020 E.V. Ruchinskaya, A.V. Gornov published in Forest Science Issues Vol. 3, No. 4, pp. 1-25<\/a><\/span><\/p>\n

E.V. Ruchinskaya*<\/sup>, A.V. Gornov<\/strong><\/span><\/p>\n

Center for Forest Ecology and Productivity of the Russian Academy of Sciences <\/em><\/span><\/p>\n

Profsoyuznaya st. 84\/32 bldg. 14<\/em>,<\/em> Moscow 117997<\/em>,<\/em> Russian Federation<\/em><\/span><\/p>\n

\u00a0<\/em><\/span><\/p>\n

*<\/sup>E-mail: elena.ruchinskaya@mail.com<\/span><\/p>\n

Received 01.11.2020<\/span><\/p>\n

Accepted 14.12.2020<\/span><\/p>\n

In the zone of broad-leaved forests of the European Russia, steppe meadows have been preserved showing rich floristic composition and making a significant contribution to the biological diversity of the territories. Bryansk oblast is one of the forest regions in Russia where such meadows are found. Here, steppe meadows with high floristic diversity and a large number of rare plant species have survived. Trees from surrounding forest areas are constantly encroaching on these meadows. Most of the young trees die from regular grass fires and economic activity. However, some individuals survive and reach a generative state, becoming relatively resistant to ground fires. The influence of single trees on the floristic diversity of steppe meadows was studied at two levels of living system organization \u2013 coenotic and population levels. Polydominant steppe meadows and polydominant steppe meadows with single generative trees were studied at the coenotic level; and coenopopulations of Iris aphylla<\/em>, Anemone sylvestris<\/em>, and Anthericum ramosum<\/em> were studied at the population level. Collecting the material, we used different methods: geobotanical, demographic, and measurements of environmental factors (illumination, slope steepness, and the frequency of grass fires). Polydominant steppe meadows were found to be were preserved in the middle part of steep slopes unsuitable for haymaking and grazing and subjected to infrequent grass fires. These communities have high floristic diversity and stable coenopopulations of model species. Ontogenetic spectra of Anemone sylvestris<\/em>, Anthericum ramosum<\/em>, and Iris aphylla<\/em> are of the complete left-hand type with the maximum number of individuals. Single trees (Quercus robur<\/em>, Tilia cordata<\/em>) have controversial influence on the vegetation of polydominant steppe meadows. On the one hand, with the introduction of trees, species diversity of communities increases. This is due to the fact that trees offer resting places and shelter for birds that spread plant diaspores. On the other hand, mature trees shade the herb cover. This leads to cover reduction and occurrence of steppe and dry meadow species, as well as affects their population structure. The ontogenetic spectrum of Anemone sylvestris<\/em> is still complete, whereas that of Iris aphylla<\/em> becomes incomplete, and the spectrum of Anthericum ramosum<\/em> becomes unfinished.<\/span><\/p>\n

Key words<\/strong>: steppe meadow, single trees, floristic diversity, coenopopulations, ontogenetic spectrum, state of coenopopulations, Anemone sylvestris, Anthericum ramosum, Iris aphylla<\/em><\/span><\/p>\n

Steppe meadows have survived in the zone of broad-leaved forests of the European Russia (Bulohov, 1977, 2001; Bosek, 1980; Skvorcov, 1982; Averinova, 2010; Semenishhenkov, 2010, 2012; Evstigneev et al., 2011; Panasenko et al., 2013, 2015, etc.). These communities, in general, have rich floristic composition and make a significant contribution to the biological diversity of the territories. However, due to human economic activity and grass fires, such coenoses are in danger of extinction (Zelenaja …, 2012; Evstigneev et al., 2018a; Ruchinskaya, 2019). Woody plants from the surrounding forest areas are constantly encroaching on the steppe meadows preserved in the zone of broad-leaved forests. Most of the young trees die from regular grass fires and economic activities such as haymaking, grazing, etc. However, some individuals survive and go into a generative state (Evstigneev et al., 2018a). Mature trees are relatively resistant to ground fires: their renewal buds are located high and the thick crust of the trunk protects the cambium (Serebrjakov, 1962). Single trees affect the growing conditions of other plants in the meadows. It is known that in phytogeneous fields of trees, illumination, air temperature and humidity, soil temperature and humidity, the amount of precipitation penetrating through the crown, quality of litter, concentration of nutrients and other soil characteristics change significantly (Uranov, 1965; Samojlov, 1983; Nikonov et al., 2002; Ipatov, 2007; Zhuravleva et al., 2012; Orlova et al., 2016). In addition, single trees attract animals of different ecological groups, e.g. soil invertebrates, mouse-like rodents, birds, etc. (Manning et al., 2006; Prevedello et al., 2018). On the one hand, these influence the growing conditions of plants, and on the other hand, they participate in the creation of both intracoenotic and intercoenotic flows of diaspores. Therefore, the objective of this work is to consider the influence of single trees on the floristic composition and the state of coenopopulations of some rare plant species in the steppe meadows.<\/span><\/p>\n

MATERIALS AND METHODS<\/strong><\/span><\/p>\n

The research was carried out in the south-east of Bryansk oblast within the Melovitskie Slopes natural monument (Figure 1). The site is located in Komarichsky-Sevsky physiographic region. It consists of elevated loess plains with ravines, gullies, slopes and outcrops of carbonate rocks on the western offsets of the Central Russian Upland. Botanically and geographically, the territory belongs to the Eastern European Province of the European broad-leaved forest region (Rastitel’nost’<\/em>…, 1980). Komarichsky-Sevsky district has temperate continental climate. The mean annual temperature is 5.4\u00a0\u03bf<\/sup>\u0421. The duration of the warm season with above-freezing temperatures is 228 days; the growing season with the temperature above + 5\u00b0C is 188 days. Mean annual precipitation is 613 mm; the mean precipitation in the warm season is 342 mm (Prirodnoe …, 1975).<\/span><\/p>\n

The research was carried out at two levels of living system organization, i.e. at the coenotic and population level. Polydominant steppe meadows and polydominant steppe meadows with single generative trees were studied at the coenotic level. At the population level, the objects of the study were coenopopulations of model plant species Iris aphylla <\/em>L., Anemone sylvestris <\/em>L., and Anthericum ramosum <\/em>L. Iris aphylla<\/em> is a short rhizome rosette spring-flowering summer-green plant (Figure 2, A<\/em>). It is a geophyte. Anemone sylvestris<\/em> is a perennial herbaceous spring-flowering summer-green short rhizome plant (Figure 2, B<\/em>). It is a hemicryptophyte and geophyte. Anthericum ramosum<\/em> is a perennial herbaceous summer-green summer-flowering short rhizome plant (Figure 3). It is a hemicryptophyte and geophyte. These species were chosen due to the fact that they are rare, endangered and listed in Red Books of many regions (Krasnaja<\/em>…, 2002, 2004, 2015, 2016, etc.). Moreover, Iris aphylla<\/em> is listed in the Red Book of Russia (2008).<\/span><\/p>\n

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Figure 1.<\/strong> Location of the Melovitskie Slopes natural monument. The green line represents the boundaries of the research object. Background satellite imagery by Microsoft Bing Maps<\/span><\/p><\/div>\n

The following research methods were used: geobotanical, demographic, statistical, and measurements of habitat factors. Relev\u00e9s were made on plots of 100 m2<\/sup> in 11-fold repetition for each community type. A complete floristic list was compiled at each plot. Species participation was evaluated as scores according to the cover-abundance scale proposed by J. Braun-Blanquet (Mirkin et al., 1989). Species richness and species density were used to assess the species diversity of communities. Species richness is the total number of species in a community, which is obtained on the basis of 11 relev\u00e9s. Species density is the average number of species per unit area. The names of vascular plants were given according to The Plant List international database (http:\/\/www.theplantlist.org\/<\/a>). For relev\u00e9s analysis of communities, ordination was carried out using Detrended Correspondence Analysis (DCA). This method works successfully with heterogeneous data of relev\u00e9s (Dzhongman et al., 1999). PC-ORD software was used for calculations. Demographic research was based on ontogenesis periodization proposed by T. A. Rabotnov (1950) and supplemented by A.A. Uranov (1975) and other scientists (Cenopopuljacii …, 1988). Ontogenesis is divided into stages that show morphological and functional differences. Individuals belonging to the same ontogenetic state are grouped together: j<\/em> \u2013 juvenile, im<\/em> \u2013 immature, v<\/em> \u2013 virginile, g1<\/sub><\/em> \u2013 young generative, g2<\/sub><\/em> \u2013 mature generative, g3<\/sub><\/em> \u2013 old generative, ss<\/em> \u2013 sub-senile, s<\/em> \u2013 senile. Ontogenetic states of the model species were determined on the basis of publications (Evstigneev et al., 2018; Ruchinskaya, 2019). The state of coenopopulations was estimated using the number, density, and type of the ontogenetic spectrum. Number is the number of individuals in the study area (Chernova, Bylova, 2007). Population density is the average number of individuals per unit area (Odum, 1986; Cenopopuljacii …, 1988). The type of the ontogenetic spectrum was named according to the classification proposed earlier (Zaugol’nova, 1994b). In the meadows and under the trees, the illuminance was measured hourly with a light meter on a cloudless June day from 10 a.m. to 6 p.m. Illuminance in lux was converted to a percentage of the total illuminance, which was measured in the open space. The slope steepness was measured using a Nikon Forestry Pro rangefinder. The frequency of grass fires was determined by the age of shoots of formation in shrubs (Frangula alnus<\/em> Mill., Corylus avellana<\/em> L.). These shoots emerge from dormant buds located in the basal part of the shrub. The former aboveground shoots were destroyed due to fire damage (Figure 4).<\/span><\/p>\n

RESULT AND DISCUSSION<\/strong><\/span><\/p>\n

Polydominant steppe meadows<\/em><\/strong> were preserved in the middle part of steep slopes hardly suitable for haymaking and grazing (Figure 5). Grass fires mainly occur once every two years. They limit the introduction of woody plants as young tree species are most vulnerable. For instance, seedlings and juvenile oak plants often die during grass fires (Komarov, 1951). As a result, polydominant communities with high species diversity are formed (Table 1; Suppl. materials). These coenoses are unique since they include species that are characteristic of steppe communities: Ajuga genevensis <\/em>L., Anemone sylvestris<\/em>, Aster amellus <\/em>L., Astragalus cicer <\/em>L., Campanula sibirica <\/em>L., Prunus cerasus <\/em>L., Galium tinctorium <\/em>L., G. verum <\/em>L. etc. The ecological-coenotic structure is dominated by plants of the dry meadow group, which also includes the above-named steppe plants. Moist meadow (Festuca pratensis<\/em> Huds., Hypericum maculatum<\/em> Crantz, Succisa pratensis<\/em> Moench, Thalictrum lucidum<\/em> L. \u0438 \u0434\u0440.), nemoral forest-edge (Brachypodium pinnatum<\/em> (L.) Beauv., Peucedanum cervaria <\/em>(L.) Cusson ex Lapeyr, Laserpitium latifolium<\/em> L., Lathyrus pisiformis<\/em> L., L. sylvestris<\/em> L., Pyrethrum corymbosum<\/em> (L.) Scop.), and nitrophilous forest-edge (Rubus caesius<\/em> L. and Valeriana officinalis<\/em> L.) plants are often found as well. Small participation is typical of forest species, i. e. nemoral \u2013 Convallaria majalis<\/em> L., Corylus avellana<\/em>, Quercus robur<\/em> L., Viola mirabilis<\/em> L., boreal \u2013 Frangula alnus<\/em>, and piny \u2013 Pteridium aquilinum<\/em>, Solidago virgaurea<\/em> L., and Viola collina<\/em> Besser. Diaspores of forest and forest-edge species are brought here by animals and wind from the neighbouring pine forest. Moist meadow and nitrophilous species are brought from floodplain communities adjacent to the slope.<\/span><\/p>\n

Table 1.<\/strong> Characteristics of communities on the steppe slopes. Melovitskie Slopes natural monument<\/span><\/p>\n

\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n
Indicators<\/span><\/td>\nCommunities<\/span><\/td>\n<\/tr>\n
1<\/em><\/span><\/td>\n2<\/em><\/span><\/td>\n<\/tr>\n
Slope angle<\/span><\/td>\n<\/tr>\n
Slope angle, M \u00b1 \u03c3<\/em><\/span><\/td>\n37 \u00b1 2.4<\/span><\/td>\n31 \u00b1 2.4<\/span><\/td>\n<\/tr>\n
Slope angle range<\/span><\/td>\n33\u201341<\/span><\/td>\n28\u201337<\/span><\/td>\n<\/tr>\n
Number of measurements<\/span><\/td>\n21<\/span><\/td>\n33<\/span><\/td>\n<\/tr>\n
Fires<\/span><\/td>\n<\/tr>\n
Fire frequency, M \u00b1 \u03c3<\/em><\/span><\/td>\n2.3 \u00b1 1.2<\/span><\/td>\n2.2 \u00b1 1.0<\/span><\/td>\n<\/tr>\n
Number of measurements<\/span><\/td>\n52<\/span><\/td>\n33<\/span><\/td>\n<\/tr>\n
Characteristics of the diversity of vascular plant species<\/span><\/td>\n<\/tr>\n
Average number of species per 100 m2<\/sup>, M \u00b1 mM<\/sub><\/em><\/span><\/td>\n51 \u00b1 1.2<\/span><\/td>\n59 \u00b1 1.2<\/span><\/td>\n<\/tr>\n
Range of number of species per 100 m2<\/sup><\/span><\/td>\n44\u201356<\/span><\/td>\n52\u201366<\/span><\/td>\n<\/tr>\n
Number of species on 11 plots of 100 m2<\/sup> each<\/span><\/td>\n98<\/span><\/td>\n107<\/span><\/td>\n<\/tr>\n
Number and proportion (%) of species of different ecological-coenotic groups<\/span><\/td>\n<\/tr>\n
Dry meadow<\/span><\/td>\n77 (78.6)<\/span><\/td>\n79 (73.8)<\/span><\/td>\n<\/tr>\n
Moist meadow<\/span><\/td>\n5 (5.1)<\/span><\/td>\n7 (6.5)<\/span><\/td>\n<\/tr>\n
Nemoral forest<\/span><\/td>\n4 (4.1)<\/span><\/td>\n8 (7.5)<\/span><\/td>\n<\/tr>\n
Nemoral forest-edge<\/span><\/td>\n6 (6.1)<\/span><\/td>\n6 (5.6)<\/span><\/td>\n<\/tr>\n
Piny<\/span><\/td>\n3 (3.1)<\/span><\/td>\n3 (2.8)<\/span><\/td>\n<\/tr>\n
Boreal forest<\/span><\/td>\n1 (1.0)<\/span><\/td>\n1 (1.0)<\/span><\/td>\n<\/tr>\n
Nitrophilous forest-edge<\/span><\/td>\n2 (2.0)<\/span><\/td>\n3 (2.8)<\/span><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n

Note<\/em>. M<\/em> is the arithmetic mean, and \u03c3<\/em> is the standard deviation. Communities: 1<\/em> \u2013 polydominant steppe meadows, 2<\/em> \u2013 polydominant steppe meadows with single generative trees<\/span><\/p>\n

Iris aphylla <\/em>is one of the predominant species in the herb layer of polydominant steppe meadows of the Melovitskie Slopes. Population density of Iris aphylla<\/em> is 82 plants per 1 m2<\/sup>. The ontogenetic spectrum is complete single-peak with a maximum at v<\/em> and g\u043d<\/sub> <\/em>plants (Figure 6, 1a<\/em>). Iris aphylla<\/em> is well adapted to high illuminance of open spaces due to the structure of its leaves: they are flattened laterally and vertically oriented (Evstigneev et al., 2018b). Seed renewal of Iris aphylla<\/em> is facilitated by the activity of animals such as ants and mouse-like rodents that inhabit the slopes and create disturbances. These microsites are characterized by sparse herb cover, loosened substrate, increased aeration and soil temperature, and significant microbiological activity (Zrjanin, 2003; Dauber, Wolters, 2000; Kostrakiewicz, 2004, etc.). For example, a population locus consisting of 10 juvenile plants was found in a 0.03 m2<\/sup> earth ejections of the mouse-like rodent. The spread of Iris aphylla<\/em> diaspores is facilitated by ants (Figure 7). R.E. Levina (1957) states that fresh seeds attract these animals with sweet, sticky liquid that is contained in the shell. Our observations showed that ants also spread dry seeds (Evstigneev et al., 2018b).<\/span><\/p>\n

Anthericum ramosum<\/em> is a codominant species in the herb layer of steppe meadows. Population density of Anthericum ramosum<\/em> is 56 plants per 1 m2<\/sup>. Ontogenetic spectrum is complete, left-hand type, single-peak with a maximum at v<\/em> and g\u043d<\/sub><\/em> plants (Figure 6, 2\u0430<\/em>). The formation of the maximum at v <\/em>and g\u043d<\/sub><\/em> plants is determined by the following: 1) short duration of j <\/em>and im <\/em>states; 2) recruitment of plants resting from flowering; 3) recruitment of v<\/em> with plants of vegetative origin formed as a result of disintegration of g2 <\/sub><\/em>plants.<\/span><\/p>\n

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Figure 2.<\/strong> Model plants species in steppe meadows of the Melovitskie Slopes natural monument: \u0410\u00a0\u2013 Iris aphylla, B \u2013 Anemone sylvestris. Photo by A.V. Gornov<\/span><\/p><\/div>\n

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Figure 3.<\/strong> Model plants species in steppe meadows of the Melovitskie Slopes natural monument: \u0410,\u00a0B \u2013 Anthericum ramosum. Photo by E.V. Ruchinskaya<\/span><\/p><\/div>\n

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Figure 4. Coppice shoots of burnt sour cherry (Prunus cerasus). A \u2013 general appearance of the shrub, B \u2013 base of the shrub. 1 <\/em>\u2013 stump of a burnt perennial shoot, 2 <\/em>\u2013 dead burnt biennial coppice shoot, 3<\/em> \u2013 live annual shoot that woke up from a dormant bud after a ground fire in the spring (from: Ruchinskaya, 2019)<\/span><\/p><\/div>\n

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Figure 5.<\/strong> Polydominant steppe meadows in the territory of the Melovitskie Slopes natural monument. Photo by A.V. Gornov<\/span><\/p><\/div>\n

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Figure 6.<\/strong> Ontogenetic spectrum of coenopopulations of model plant species in steppe meadows. 1 \u2013 Iris aphylla, 2 \u2013 Anthericum ramosum, 3 \u2013 Anemone sylvestris. The X axis shows ontogenetic states, and the Y axis \u2013 the proportion of individual plants, %. Circled is population density (the number of plants per 1 m2) is shown. Communities: \u0430 \u2013 polydominant steppe meadows, b \u2013 polydominant steppe meadows with single generative trees. Ontogenetic states of trees: j \u2013 juvenile, im \u2013 immature, v \u2013 virginile, g\u043d \u2013 temporarily not flowering generative plant, g1 \u2013 young generative, g2 \u2013 mature generative, g3 \u2013 old generative, ss \u2013 subsenile, s \u2013 senile<\/span><\/p><\/div>\n

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Figure 7.<\/strong> Dispersal of fresh Iris aphylla seeds by the red forest ant (Formica rufa). Photo by E.V. Ruchinskaya<\/span><\/p><\/div>\n

Anemone sylvestris<\/em> can be both an assectator and a codominant in the herb layer of steppe meadows. Population density of A. sylvestris<\/em> is 75 plants per 1 m2<\/sup>. Ontogenetic spectrum is complete left-hand type with a maximum at im<\/em> plants (Figure 6, 3a<\/em>), the density of which is 27 trees per 1 m2<\/sup>. No A. sylvestris <\/em>individuals of seed origin was found in the studied community. Therefore, the spectrum can be called vegetative and complete. High population density and the complete ontogenetic spectrum are determined by the biology of A. sylvestris<\/em>. Large number of plants of the pregenerative state is due to the ability of A. sylvestris<\/em> to vegetative reproduction with deep rejuvenation (Figure 8). Vegetative individuals develop from buds that appear on horizontal adventitious roots (Starostenkova, 1986; Barykina, Potapova, 1994). The beginning of the growth season in early spring, before the herb layer rises, contributes to the accumulation of a sufficient amount of macronutrients required for the formation of generative organs in plans. It is worth noting that the absence of seed individuals is evidence of adverse conditions for the coenopopulation. Apparently, this is due to the spread of fire on the slopes, which destroys young seed individuals of A. sylvestris<\/em>.<\/span><\/p>\n

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Figure 8.<\/strong> Vegetative renewal of Anemone sylvestris: A \u2013 generative individual with a root shoot and adventitious buds on the root (from: Barykina, Potapova, 1994, as supplemented), B \u2013 juvenile and virginile individuals of root shoot origin (from: Gornov et al., 2013). j \u2013 juvenile individual, g \u2013 generative individual, v \u2013 virginal individual, r m p \u2013 root of the parent plant, rh \u2013 rhizome, a b \u2013 adventitious bud<\/span><\/p><\/div>\n

Polydominant steppe meadows with single generative trees. <\/em><\/strong>On the slopes, there are single Quercus robur<\/em> and Tilia cordata <\/em>generative trees (Figure 9), which survived fire during their virginile stage. Mature oaks and linden trees are relatively resistant to ground fires: their renewal buds are located high, and the thick crust of the trunk protects the cambium (Serebrjakov, 1962). The steepness of the slope and the frequency of grass fires are similar to polydominant steppe meadows. Ordination of relev\u00e9s divided polydominant steppe meadows and polydominant steppe meadows with single trees into distinct groups (Figure 10). The communities differ in the maximum values of species richness and species density (Table 1; Suppl. materials). High species diversity is due to several factors. First, in the past, the communities were not subjected to active grazing or haymaking, as they are also located on steep parts of the slopes. Second, single trees offer resting places and shelter for many animals, including birds. These are known to disperse seeds of meadow and forest plants (Manning et al., 2006; Prevedello et al., 2018). As a result, communities with free-standing trees have higher species richness than polydominant steppe meadows. The ecological-coenotic structure of the community is dominated by dry meadow and steppe plants. Under the crowns of single trees, the illuminance is reduced to 60% of the total. Shading reduces the cover of light-loving dry meadow and steppe plants. However, the number of species in this group increases. Allium oleraceum<\/em> L., Artemisia absinthium<\/em> L., Carex montana<\/em> L., Cirsium decussatum <\/em>Janka, Fallopia convolvulus<\/em> (L.) \u00c1. L\u00f6ve, Filipendula vulgaris<\/em> Moench, Hypericum perforatum<\/em> L., Silene vulgaris <\/em>(Moench) Garcke, S. viscaria <\/em>(L.) Jess., Stachys officinalis <\/em>(L.) Trevis., Veronica spuria<\/em> L. etc. appear. In addition, the species composition of other ecological-coenotic groups also expands: Carex hirta<\/em> L., C. lachenalii<\/em> Schkuhr, Galeopsis bifida<\/em> Boenn are found in the moist meadow group; the group of nemoral plants is supplemented by Euonymus europaeus <\/em>L., Lathyrus niger<\/em> (L.) Bernh. and Pyrus communis<\/em> L., and the group of forest-edge nitrophilous group \u2013 by Galium aparine<\/em> L. Apparently, this is due to the activity of animals, primarily birds, that use single trees like resting places and brought diaspores of these plants. It is known that birds actively disperse seeds of many plant species (Levina, 1957; Cramp, 1998, etc.). Thus, thanks to single trees, the polydominant composition of the community with maximum species diversity is maintained. However, shading has an adverse effect on the state of coenopopulations of model plant species.<\/span><\/p>\n

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Figure 9.<\/strong> Polydominant steppe meadows with single trees: \u0410 \u2013 general appearance of the slope, B \u2013 crown of Tilia cordata from above. Photo by A.Yu. Sitnikov<\/span><\/p><\/div>\n

\u00a0<\/em><\/span><\/p>\n

Iris aphylla<\/em> loses positions in the herb layer of polydominant steppe meadows with single generative trees. Population density is 50 plants per 1 m2<\/sup>. This is almost twice as low as in polydominant steppe meadows. A decrease in density is explained by the fact that due to the small amount of light, very few fruiting individuals able to produce viable seeds are formed in Iris aphylla<\/em>. This leads to a four-fold drop in the number of young seed plants in the coenopopulation. Established plants feature an increased area of the leaf blade (Table 2). This adaptation allows the plants to get more of the scattered light. However, due to the lack of light, most individuals of Iris aphylla<\/em> develop only to the v<\/em>-ontogenetic state, and then die. As a result, an incomplete ontogenetic spectrum is formed with a maximum at v<\/em> plants (Figure 6, 1<\/em>b<\/em>). If, over time, the number of trees on the slope increases, and they form a close-canopy area of the forest, the coenopopulation of Iris aphylla<\/em> will disappear.<\/span><\/p>\n

T<\/strong>able 2.<\/strong> Length and width of the leaves of Iris aphylla<\/em> in the light (1) and in the shade (2)<\/span><\/p>\n

\n\n\n\n\n\n\n
Measurement<\/span><\/td>\nN<\/em><\/span><\/td>\nM \u00b1 mM<\/sub><\/em><\/span><\/td>\n\u03c3<\/em><\/span><\/td>\nU<\/em><\/span><\/td>\n<\/tr>\n
1<\/span><\/td>\n2<\/span><\/td>\n1<\/span><\/td>\n2<\/span><\/td>\n1<\/span><\/td>\n2<\/span><\/td>\n1<\/span><\/td>\n2<\/span><\/td>\n<\/tr>\n
Leaf length, cm<\/span><\/td>\n22<\/span><\/td>\n26<\/span><\/td>\n31.4 \u00b1 2.22<\/span><\/td>\n60.6 \u00b1 1.70<\/span><\/td>\n10.4<\/span><\/td>\n8.7<\/span><\/td>\n6 <\/strong>(p<\/em> = 0.000000)<\/span><\/td>\n<\/tr>\n
Leaf width, cm<\/span><\/td>\n22<\/span><\/td>\n26<\/span><\/td>\n1.6 \u00b1 0.09<\/span><\/td>\n2.3 \u00b1 0.08<\/span><\/td>\n0.4<\/span><\/td>\n0.4<\/span><\/td>\n56 <\/strong>(p<\/em> = 0.000002)<\/span><\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n

Note<\/em>. Communities: 1 \u2013 polydominant steppe meadows, 2 \u2013 polydominant steppe meadows with single generative trees. N<\/em> \u2013 number of measurements, M <\/em>\u2013 arithmetic mean, mM<\/sub><\/em> \u2013 error of the arithmetic mean, \u03c3<\/em> \u2013 standard deviation, U <\/em>\u2013 Mann\u2013Whitney test values, p <\/em>\u2013 probability of error. Significant differences in the U<\/em> test are shown in bold.<\/span><\/p>\n

Anthericum ramosum<\/em> shows low cover in the herb layer of polydominant steppe meadows with single generative trees. Under the tree canopy, population density of A. ramosum<\/em> decreases sharply and only reaches 7 plants per 1 m2<\/sup>. This is eight times lower than in polydominant steppe meadows. Here, an unfinished left-hand type ontogenetic spectrum is formed with a maximum at v<\/em> and g<\/em>\u043d<\/sub><\/em> individuals (Figure 6, 2<\/em>b<\/em>). There are no ss<\/em> or s<\/em> individuals in the coenopopulation, which is probably due to the death of plants already in the g3<\/sub><\/em> state. In addition, the decrease in the population density is explained by the fact that due to a small amount of light, the mortality of j<\/em> and im<\/em> individuals increases. The group of v<\/em> and g<\/em>\u043d<\/sub><\/em> plants is replenished by single temporarily not flowering generative plants and particles formed as a result of disintegration of g2<\/sub><\/em> plants. For the same reason, the number of g3<\/sub><\/em> individuals, which are represented by branching and non-branching particles, increases slightly.<\/span><\/p>\n

\"\"

Figure 10.<\/strong> Results of DCA-ordination of relev\u00e9s of steppe plant communities in the axes of the greatest variation in floristic composition. Communities: 1 \u2013 polydominant steppe meadows, 2 \u2013 polydominant steppe meadows with single generative trees<\/span><\/p><\/div>\n

The projective cover of Anemone sylvestris<\/em> is significantly less than in polydominant steppe meadows. Population density decreases six-fold \u2013 there are only 12 individuals per 1 m2<\/sup>. In the shade, most anemones do not form flower stalks. Therefore, the generative fraction is represented by single fruiting individuals with predominant g1<\/sub><\/em> plants. They are mainly found on the periphery of the crowns, where there is lateral illumination. As a result, the density of generative plants is ten times less than in the former community. This leads to a significant reduction in the replenishment of the coenopopulation with young plants, since the number of g2<\/sub><\/em> individuals that produce the largest number of root shoots decreases. Despite this, the ontogenetic spectrum of Anemone sylvestris<\/em> is still complete and left-hand type. However, it shows an extremely low participation of generative individuals and a shift of the maximum to v<\/em> plants (Figure 6, 3b<\/em>). The latter is due to the relatively long duration of the v<\/em>-ontogenetic state and the insignificant replenishment of the coenopopulation by j<\/em> and im<\/em> plants.<\/span><\/p>\n

CONCLUSION<\/strong><\/span><\/p>\n

The maximum species diversity of polydominant steppe meadows is maintained on steep slopes unsuitable for ploughing up, where haymaking and grazing are difficult and fires are infrequent. This contributes to the formation of stable coenopopulations of model species. Their ontogenetic spectra belong to one type \u2013 complete left-hand type, with the maximum accounted for by young individuals. The mechanism of formation of this spectrum is species-specific. Thus, individuals of Iris aphylla<\/em> and Anthericum ramosum<\/em> are characterized by vegetative reproduction with shallow rejuvenation of particles and frequent breaks in flowering. In addition, the left-hand type structure is provided by high seed productivity, and in case of Anemone sylvestris<\/em> \u2013 by active vegetative reproduction, when plants are deeply rejuvenated. Single trees (Quercus robur<\/em>, Tilia cordata<\/em>) have controversial influence on the vegetation of polydominant steppe meadows. On the one hand, with the introduction of trees, species diversity of communities increases. This is due to the fact that trees offer resting places and shelter for birds that spread plant diaspores. On the other hand, mature trees shade the herb cover. This leads to reduced cover and occurrence of steppe and dry meadow species, as well as affects their population structure. The number of individuals of all ontogenetic states is significantly reduced. The ontogenetic spectrum of Anemone sylvestris<\/em> remains complete, whereas in Iris aphylla<\/em> it becomes incomplete, and in Anthericum ramosum<\/em> it becomes unfinished. If, over time, the number of trees on the slope increases, and they form a close-canopy area of the forest, coenopopulations of the model species will gradually disappear from the community.<\/span><\/p>\n

ACKNOWLEDGEMENTS<\/strong><\/span><\/p>\n

The work was carried out as part of the state assignment of the CEPF RAS “Methodological approaches to the assessment of structural organization and functioning of forest ecosystems” (state registration number AAAA-A18-118052400130-7). The authors express their gratitude to Ye.A. Gavrilyuk, senior research officer of the CEPF RAS, for his help in drawing up the map.<\/span><\/p>\n

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Reviewer:<\/strong> candidate of Biological Sciences, Associate Professor Panasenko N.N.<\/span><\/p>\n

Supplementary materials<\/em><\/span><\/p>\n

Species composition of the Melovitskie Slopes natural monument communities<\/span><\/p>\n

\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n\n
Plant name<\/span><\/td>\nPO<\/span><\/td>\nECG<\/span><\/td>\n<\/tr>\n
Communities<\/span><\/td>\n<\/tr>\n
1<\/span><\/td>\n2<\/span><\/td>\n<\/tr>\n<\/thead>\n
Achillea millefolium<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Agrimonia eupatoria<\/em> L.<\/span><\/td>\nV (+)<\/span><\/td>\nIV(+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Ajuga genevensis<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Allium oleraceum<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Anemone sylvestris<\/em> L.<\/span><\/td>\nV (1)<\/span><\/td>\nII (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Anthericum ramosum <\/em>L.<\/span><\/td>\nV (1)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Anthyllis vulneraria<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\n\u00a0\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Artemisia absinthium<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Artemisia vulgaris<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Asparagus officinalis<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Aster amellus <\/em>L.<\/span><\/td>\nV (3)<\/span><\/td>\nII (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Astragalus cicer<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Astragalus glycyphyllos <\/em>L.<\/span><\/td>\nI (+)<\/span><\/td>\nIV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Brachypodium pinnatum<\/em> (L.) Beauv.<\/span><\/td>\nII (2)<\/span><\/td>\nV (3)<\/span><\/td>\nNm-FE<\/span><\/td>\n<\/tr>\n
Bromus inermis<\/em> Leyss.<\/span><\/td>\nV (3)<\/span><\/td>\nV (4)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Calamagrostis epigejos<\/em> (L.) Roth<\/span><\/td>\nIV (+)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Campanula bononiensis<\/em> L.<\/span><\/td>\nV (+)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Campanula rapunculoides<\/em> L.<\/span><\/td>\nII (+)<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Campanula sibirica<\/em> L.<\/span><\/td>\nII (+)<\/span><\/td>\n\u2013\u00a0<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Carex hirta<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Carex lachenalii<\/em> Schkuhr<\/span><\/td>\n\u2013<\/span><\/td>\nIV (+)<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Carex montana<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Carex praecox<\/em> Schreb.<\/span><\/td>\nIII (+)<\/span><\/td>\nIV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Centaurea jacea <\/em>L.<\/span><\/td>\nIII (+)<\/span><\/td>\n\u2013\u00a0<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Centaurea phrygia<\/em> subsp. pseudophrygia<\/em> (C.A. Mey.) Gugler<\/span><\/td>\nIII (1)<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Chamaecytisus ruthenicus<\/em> (Fisch. ex Woloszcz.) Klaskova<\/span><\/td>\nV (+)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Cichorium intybus<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013\u00a0<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Cirsium decussatum<\/em> Janka<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Cirsium pannonicum<\/em> (L. fil.) Link<\/span><\/td>\nIV (1)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Convallaria majalis <\/em>L.<\/span><\/td>\nIV (1)<\/span><\/td>\nIV (1)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Convolvulus arvensis<\/em> L.<\/span><\/td>\nIII (+)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Corylus avellana<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nV (+)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Dactylis glomerata<\/em> L.<\/span><\/td>\nIII (+)<\/span><\/td>\nV (+)<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Elymus repens<\/em> (L.) Gould<\/span><\/td>\nII (+)<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Equisetum arvense<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Erigeron annuus<\/em> (L.) Desf.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Euonymus europaeus<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nIII (+)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Euphorbia esula<\/em> L.<\/span><\/td>\nII (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Euphorbia semivillosa<\/em> (Prokh.) Krylov<\/span><\/td>\nV (2)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Fallopia convolvulus<\/em> (L.) \u00c1. L\u00f6ve<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Festuca pratensis <\/em>Huds.<\/span><\/td>\nII (+)<\/span><\/td>\n\u2013\u00a0<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Filipendula vulgaris<\/em> Moench<\/span><\/td>\n\u2013<\/span><\/td>\nIV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Fragaria viridis<\/em> Weston<\/span><\/td>\nIII (+)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Frangula alnus<\/em> Mill.<\/span><\/td>\nIV (+)<\/span><\/td>\nII (+)<\/span><\/td>\nBr-Fo<\/span><\/td>\n<\/tr>\n
Galatella linosyris<\/em> (L.) Rchb.f.<\/span><\/td>\nII (+)<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Galeopsis bifida<\/em> Boenn.<\/span><\/td>\n\u2013<\/span><\/td>\nIII (+)<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Galium aparine<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nIV (+)<\/span><\/td>\nNt-FE<\/span><\/td>\n<\/tr>\n
Galium boreale <\/em>L.<\/span><\/td>\nV (+)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Galium mollugo<\/em> L.<\/span><\/td>\nV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Galium tinctorium <\/em>L.<\/span><\/td>\nV (1)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Galium verum<\/em> L.<\/span><\/td>\nV (+)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Genista tinctoria<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Geranium sanguineum<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nIV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Hypericum perforatum<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Inula hirta<\/em> L.<\/span><\/td>\nII (1)<\/span><\/td>\nIV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Inula salicina<\/em> L.<\/span><\/td>\nV (1)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Iris aphylla<\/em> L.<\/span><\/td>\nV (3)<\/span><\/td>\nV (2)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Knautia arvensis<\/em> (L.) Coult.<\/span><\/td>\nIII (+)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Lactuca serriola<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Laserpitium latifolium <\/em>L.<\/span><\/td>\nIV (1)<\/span><\/td>\nV (2)<\/span><\/td>\nNm-FE<\/span><\/td>\n<\/tr>\n
Lathyrus niger<\/em> (L.) Bernh.<\/span><\/td>\n\u2013<\/span><\/td>\nV (1)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Lathyrus pisiformis<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\n\u00a0\u2013<\/span><\/td>\nNm-FE<\/span><\/td>\n<\/tr>\n
Lathyrus sylvestris<\/em> L.<\/span><\/td>\nII (+)<\/span><\/td>\nIV (1)<\/span><\/td>\nNm-FE<\/span><\/td>\n<\/tr>\n
Lavatera thuringiaca<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Leucanthemum vulgare <\/em>(Vaill.) Lam.<\/span><\/td>\nIV (1)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Linum flavum<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Lithospermum officinale<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nIV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Medicago falcata<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Myosotis ramosissima<\/em> Rochel<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Nepeta nuda<\/em> L.<\/span><\/td>\nI (1)<\/span><\/td>\nIV (2)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Origanum vulgare <\/em>L.<\/span><\/td>\nIV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Peucedanum alsaticum<\/em> L.<\/span><\/td>\nV (1)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Peucedanum cervaria<\/em> (L.) Cusson ex Lapeyr.<\/span><\/td>\nIV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nNm-FE<\/span><\/td>\n<\/tr>\n
Peucedanum oreoselinum<\/em> (L.) Moench<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013\u00a0<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Phlomoides tuberosa<\/em> (L.) Moench<\/span><\/td>\nII (+)<\/span><\/td>\nII (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Pilosella piloselloides<\/em> subsp. bauhinii<\/em> (Schult.) S.Br\u00e4ut. & Greuter<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Plantago lanceolata<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Plantago media<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Poa angustifolia<\/em> L.<\/span><\/td>\nV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Poa trivialis<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nIV (+)<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Podospermum purpureum<\/em> (L.) W.D.J. Koch & Ziz<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Polygala comosa<\/em> Schkuhr<\/span><\/td>\nIII (+)<\/span><\/td>\n\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Polygonatum odoratum<\/em> (Mill.) Druce<\/span><\/td>\nIII (+)<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Prunus cerasus<\/em> L.<\/span><\/td>\nIV (1)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Pteridium aquilinum <\/em>(L.) Kuhn<\/span><\/td>\nIV (2)<\/span><\/td>\nV (2)<\/span><\/td>\nPn<\/span><\/td>\n<\/tr>\n
Pyrethrum corymbosum<\/em> (L.) Scop.<\/span><\/td>\nV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nNm-FE<\/span><\/td>\n<\/tr>\n
Pyrus communis<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Quercus robur<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nV (+)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Ranunculus polyanthemos<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Rubus caesius<\/em> L.<\/span><\/td>\nII (+)<\/span><\/td>\nIII (1)<\/span><\/td>\nNt-FE<\/span><\/td>\n<\/tr>\n
Salvia pratensis<\/em> L.<\/span><\/td>\nV (3)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Salvia verticillata<\/em> L.<\/span><\/td>\nII (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Securigera varia<\/em> (L.) Lassen<\/span><\/td>\nIV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Sedum maximum <\/em>(L.) Suter<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Sedum telephium<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Serratula tinctoria<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nNm-FE<\/span><\/td>\n<\/tr>\n
Seseli libanotis <\/em>(L.) W.D.J. Koch<\/span><\/td>\nI (2)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Silene latifolia <\/em>Poir.<\/span><\/td>\nI (+)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Silene nutans<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Silene viscaria<\/em> (L.) Jess.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Silene vulgaris<\/em> (Moench) Garcke<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Solidago virgaurea<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nIII (+)<\/span><\/td>\nPn<\/span><\/td>\n<\/tr>\n
Stachys officinalis<\/em> (L.) Trevis.<\/span><\/td>\n\u2013<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Stachys recta<\/em> L.<\/span><\/td>\nV (3)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Succisa pratensis<\/em> Moench<\/span><\/td>\nI (+)<\/span><\/td>\n\u00a0\u2013<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Taraxacum officinale<\/em> Wigg.<\/span><\/td>\nII (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Thalictrum lucidum<\/em> L.<\/span><\/td>\nIII (+)<\/span><\/td>\n\u00a0\u2013<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Thalictrum minus <\/em>L.<\/span><\/td>\nIII (1)<\/span><\/td>\nIII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Tilia cordata<\/em> Mill.<\/span><\/td>\n\u2013<\/span><\/td>\nIV (+)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Tragopogon dubius<\/em> Scop.<\/span><\/td>\nI (+)<\/span><\/td>\n\u00a0\u2013<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Trifolium alpestre<\/em> L.<\/span><\/td>\nV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Trifolium montanum <\/em>L.<\/span><\/td>\nV (+)<\/span><\/td>\nII (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Valeriana officinalis<\/em> L.<\/span><\/td>\nIII (+)<\/span><\/td>\nII (+)<\/span><\/td>\nNt-FE<\/span><\/td>\n<\/tr>\n
Verbascum lychnitis<\/em> L.<\/span><\/td>\nII (+)<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Verbascum nigrum<\/em> L.<\/span><\/td>\nV (+)<\/span><\/td>\nIV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Veronica austriaca<\/em> subsp. teucrium<\/em> (L.) D.A. Webb<\/span><\/td>\nV (1)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Veronica chamaedrys<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Veronica spuria<\/em> L.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Vicia tenuifolia<\/em> Roth<\/span><\/td>\nV (2)<\/span><\/td>\nV (1)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Vicia tetrasperma<\/em> (L.) Schreb.<\/span><\/td>\n\u2013<\/span><\/td>\nI (+)<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Vincetoxicum hirundinaria <\/em>Medik.<\/span><\/td>\nV (1)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Viola canina<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\n\u2013\u00a0<\/span><\/td>\nM-Md<\/span><\/td>\n<\/tr>\n
Viola collina<\/em> Besser.<\/span><\/td>\nIII (+)<\/span><\/td>\nIV (+)<\/span><\/td>\nPn<\/span><\/td>\n<\/tr>\n
Viola hirta<\/em> L.<\/span><\/td>\nIV (+)<\/span><\/td>\nV (+)<\/span><\/td>\nD-Md<\/span><\/td>\n<\/tr>\n
Viola mirabilis<\/em> L.<\/span><\/td>\nI (+)<\/span><\/td>\nIII (+)<\/span><\/td>\nNm-Fo<\/span><\/td>\n<\/tr>\n
Number of species<\/span><\/td>\n98<\/span><\/td>\n107<\/span><\/td>\n<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<\/div>\n

Note.<\/em> Communities: 1<\/em> \u2013 polydominant steppe meadows, 2<\/em> \u2013 polydominant steppe meadows with single generative trees PO \u2013 average occurrence points, Arabic numerals and \u00ab+\u00bb \u2013 points of cover-abundance scale proposed by J. Braun-Blanquet. ECG \u2013 ecological-coenotic groups: D-Md \u2013 dry meadow, M-Md \u2013 moist meadow, Pn \u2013 piny (boreal forest-edge), Nm-FE \u2013 nemoral forest edge, Nm-Fo \u2013 nemoral forest, Nt-FE \u2013 nitrophilous forest-edge, Br-Fo \u2013 boreal forest.<\/span><\/p>\n","protected":false},"excerpt":{"rendered":"

Original Russian Text \u00a9 2020 E.V. Ruchinskaya, A.V. Gornov published in Forest Science Issues Vol. 3, No. 4, pp. 1-25 E.V. Ruchinskaya*, A.V. Gornov Center for Forest Ecology and Productivity of the Russian Academy...<\/p>\n","protected":false},"author":1,"featured_media":0,"comment_status":"closed","ping_status":"closed","sticky":false,"template":"","format":"standard","meta":[],"categories":[20],"tags":[],"_links":{"self":[{"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/posts\/4156"}],"collection":[{"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/posts"}],"about":[{"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/types\/post"}],"author":[{"embeddable":true,"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/users\/1"}],"replies":[{"embeddable":true,"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/comments?post=4156"}],"version-history":[{"count":4,"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/posts\/4156\/revisions"}],"predecessor-version":[{"id":4236,"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/posts\/4156\/revisions\/4236"}],"wp:attachment":[{"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/media?parent=4156"}],"wp:term":[{"taxonomy":"category","embeddable":true,"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/categories?post=4156"},{"taxonomy":"post_tag","embeddable":true,"href":"https:\/\/jfsi.ru\/en\/wp-json\/wp\/v2\/tags?post=4156"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}