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Biosystems
Diversitv
Biosystems
Diversity
ISSN 2519-8513 (Print) ISSN 2520-2529 (Online) Biosyst. Divers., 26(4), 286-291 doi: 10.15421/011843
Comparison of commonly used ecological scales with the Belgard Plant Ecomorph System
B. Baranovski, N. Roschina, L. Karmyzova, I. Ivanko
Oles Honchar Dnipro National University, Dnipro, Ukraine
Article info
Received 08.10.2018 Received in revised form
14.11.2018 Accepted 18.11.2018
Oles Honchar Dnipro National University, Gagarin Av., 72, Dnipro, 49010, Ukraine. Tel.: +38-095-779-99-94. E-mail:
Baranovski, B., Roschina, N., Karmyzova, L., & Ivanko, I. (2018). Comparison of commonly used ecological scales with the Belgard Plant Ecomorph System. Biosystems Diversity, 26(4), 286-291. doi:1015421/011843
There are several ecological scales developed both for phytoindication of ecological factors and plant ecomorphs. Among them, the scales of Ellenberg and Tsyganov are the most commonly used. L. G. Ramensky and P. S. Pogrebnyak had developed a phytoindication method; they also were founders of first ecological scale of plant species in relation to various environmental factors. One of first ecomorph systems was developed by Alexander Lyutsianovich Belgard. In 1947, Belgard presented a tabular ecomorph system in his doctoral dissertation, and later in monograph "Forest vegetation of the South-East of Ukraine". In the system he used abbreviated Latin names applying terminology proposed in the late 19th century by Dekandol, Warmin and other authors. He considered ecomorphs as adaptations of plants to environmental conditions in forests of the steppe zone of Ukraine where forest cenoses are exposed to processes of steppization, prairification, swamping, salinization, and thus clarification of relationships between forest, meadow, steppe, marsh and weed plant species was essential. Therefore, development and introduction of cenomorph terms as "adaptation of plant species to phytocenosis as a whole" were an absolutely new contribution to the concept of ecomorph system. In environmental factor scales of Ellenberg and other authors, environment characteristics based on phytoindication were underlined; in the Belgard Plant Ecomorph System, ecomorphs reflect ability of plant species to grow within certain ranges of a given factor. These approaches are quite comparable, and ecomorphs of the Belgard system correspond to certain grades of the Ellenberg and Tsyganov scales. The Belgard ecomorph system has been applied in a number of fundamental and applied works on plant ecology and phytocenology. It is convenient for characterizing ecological features of plant species growing in the steppe zone with a wide range of environment factors such as lighting, humidity, and soil richness. Other authors have expanded and supplemented the Belgard Plant Ecomorph System based on its strategy. A number of ecomorphs was introduced; they reflect intermediate or extreme gradations of factors. A new cenomorph - silvomargoant - has been proposed by the authors of this paper.
Keywords: ecological scales; environmental factors; habitats; heliomorphs; hygromorphs; trophomorphs; halomorphs; cenomorphs; ecomorphic analysis
History of phytoindication scales and ecomorph systems
A large number of environmental scales currently exist. On the one hand, they are intended for phytoindication of ecological factors, and on the other, as a system of plant ecomorphs (Diduh & Pljuta, 1994). One of first ecomorph systems was developed by Alexander Lutsianovich Belgard on the basis of concepts of D. I. Mendeleev, V. V. Dokuchaev, V. N. Sukachev. Together with his teacher G. N. Vysotsky, Belgard developed a new branch of science "Steppe Forestry". The Belgard Eco-morph System was presented by the author in his doctoral dissertation "Forests of South-Eastern Ukraine" in 1947, and then in his monograph "Forest Vegetation of South-Eastern Ukraine" (1950). In development of the ecomorph system he applied known terminology and abbreviated Latin plant names (Belgard, 1950). For nearly 70 years, this ecomorph system has been successfully applied in survey and characteristics of cenoses of both specific biotopes and landscapes. The Belgard Ecomorph System was developed in a number of fundamental works on plant ecology and phytocenology. It can be quite wide ranging: in assessment of phytocenotic structure, in ecological characteristics of flora on suprace-notic levels, as well as in assessing the state of the environment with the main physical and chemical parameters.
Other authors have expanded and supplemented the Belgard Plant Ecomorph System based on its strategy. A number of ecomorphs reflecting intermediate or extreme gradations of factors have been introduced into the system. A new cenomorph - silvomargoant - has been proposed by the authors of this paper. There is a practical need for expanding the Belgard Plant Ecomorph System and further introduction of additional
ecomorphs. The objective of this work was characterization of the peculiarities of the Belgard Plant Ecomorph System (including its supplemented version) and its advantages in comparison with the scales of Ellenberg and Tsyganov.
Ramensky was founder of the phytoindication method that is based on the use of species composition of vegetation. He was also originator of the first ecological scale of plant species in relation to various environmental factors (Ramenskiy, 1929). According to Tsyganov (1983), "works of Pogrebnyak and Ramensky are sources of European score-and-scale methods in ecological analysis. The first scientific work of Pogrebnyak including justification of his methodology was published in 1927. The first scientific work of Ramensky where he justified and applied in detail an ordinary coordination method was published in 1929, i.e. much earlier than similar works of Ellenberg and other European researchers." Pogrebnyak (1955) proposed an ecological scale of tree species, Ramensky (1956) developed detailed ecological scales of soil humidity, richness and salinity of meadow plant communities.
There exist a large number of ecological scales (Table 1), both phy-toindications of ecological factors and plant ecomorphs. Many scales were developed by their authors based on characteristics of plant communities and biotopes surveyed. Among them, the most commonly used scales are: the Ellenberg (1950, 1974) scale, the Tsyganov (1975) scale, the Landolt (1977) scale. The Ellenberg scale is the most commonly used (Gilhaus et al., 2017; Ford et al., 2018; Goedecke et al., 2018; Hancock et al., 2018). A value set of environmental factors for vascular plants in Central Europe is defined in this scale (Ellenberg 1979, 1988; Ellenberg et al., 1991). It is widely used both in Europe and
adjacent territories (Kalusova et al., 2016; Berg et al., 2017; Britton et al., 2017; Ceplova et al., 2017; Chmura et al., 2017; Hulber et al., 2017; Muir, 2017; Pruchniewicz, 2017; Santini et al., 2017; Johansen et al., 2018; Roeling et al., 2018). Its modern version was supplemented and adapted by many foreign authors (Douda et al., 2016; Dyderski et al., 2016; Ewald & Ziche, 2016; Koch et al., 2016; Van Dobben & de Vies, 2016; Elst et al., 2017; Mitchell et al., 2017; Vitasovic Kosic et al., 2017; Kill? et al., 2018; Kosanic et al., 2018).
The scales are divided into point and amplitude scales. The point scale indicates the ecological range of plant species by a particular environmental factor. The amplitude scale determines the coordinate of plant species on the axis of environmental factor.
Table 1
General characteristics of ecological scales (by Diduh & Pljuta, 1994)
Author Hd fH Tr Rc Total score in the scale Nt Gm Ar Tm Om Kn Cr Lc Dg
Ellenberg 12 - 3 9 9 - - 9 - 9 - 9
Landolt 5 4 2 5 5 5 5 5 - 5 - 5 -
Zoiomi 11 - - 6 - - - 7 - 2 - - -
Frank & Klotz 12 2 3 9 9 - - 9 - 9 - 9 -
Zazhitskii* 6 - 3 6 5 5 5 5 - 5 - 5 -
Tsyganov* 23 11 19 13 11 - - 17 15 15 15 9 -
Ramensky* 120 20 30 - - - 10 - - - - - 10
Tsatsenkin* 120 - 30 10
Notes: * - scales wherein factor amplitude is displayed; signs of the factors: Hd - soil moisture, W - soil moisture changes, Tr - generalized salt regime, Rc - acidity, Nt - content of mineral nitrogen, Gm - content of humus, Ar -soil aeration, Tm - thermoclimate, Om - ombroclimate, Kn - climate continen-tality; Cr - cryoclimate; Lc - light intensity in cenosis; Dg - pasture digression.
Belgard presented his ecomorphic system of vascular plant species in relation to the main environmental factors with the purpose of classifying ecological characteristics of forest communities in the steppe zone of Ukraine (Belgard, 1950). In 1947, A. L. Belgard presented a tabular ecomorphic system in his doctoral dissertation, and then in his monograph "Forest vegetation of the South-East ofUkraine" (1950). The author developed an ecomorphic system using terminology proposed in the late 19th century by Decandol (Dekandol, 1956), Warming (Warming, 1903), and by other authors. The Tabular Belgard Plant Ecomorph System was compiled for the main environmental factors: light intensity (heliomorphs), temperature (termomorphs), humidity (hygromorphs), soil richness (trophomorphs). The author designated names of ecomorphs with abbreviated Latin names of environmental factors. Tabulated eco-morphic analysis of plant species within a community or system of supra-cenotic level (ecological certification of species) gives insight into the ecological structure of flora (Table 2).
Table 2
Fragment of table on ecological characteristics of plant species of genus Equisetum in the steppe zone of Ukraine according to Belgard's Plant Ecomorph System
№ Species within genus Heliomorph Trophomorph Hygramorph Cenomorph
1. Equisetum arvense L. ScHe MsTr HgMs RuSilPr
2. E. fluviatile L. ScHe MsTr HelHg AqPal
3. E. hyemale L. ScHe MsTr HgMs PrSil
4. E. palustre L. ScHe MsTr MsHg PrPal
5. E. pratense Ehrh. ScHe MsTr HgMs Pr
6. E. ramosissimum Desf. ScHe MsOgTr Ms PrPs
7. E. sylvaticum L. Sc MsTr HgMs Sil
8. E. telmateia Ehrh. HeSc MsTr MsHg SilPal
9. E. variegatum Schlech. ex Weber et Mohr. He MsTr MsHg Pr
Note: plant species names are given in accordance with the nomenclature of Mosyakin & Fedoronchuk (1999)
Alexander Grossheim, famous botanist, author of Angiosperm Phylogeny Group Classification (1946), emphasized the following: "The advantage of the ecological analysis method proposed by A. L. Belgard resides in the fact that it covers all the most important environmental factors affecting the existence of plant species in a phytocenosis.
Usually, a more vivid impression is obtained on application of the ecomorphic method in a phytocenosis. According to A. L. Belgard, the characteristic of cenose is multisided in this case. In application of the author's formulae and graphs, interrelations existing in nature have many-sided coverage, due to which maximum approximation to the truth is obtained in comparison with the use of other, usually one-sided methods".
Correspondence of the Belgard Plant Ecomorph System to other environmental scales
Ecomorphs are considered by Belgard (1950) as adaptations, adjustments to environmental conditions (Belgard, 1950). Belgard's scheme "was made specifically for forest survey in the steppe zone of Ukraine, where forest cenoses are often exposed to processes of steppization, prairification, swamping, salinization, spread of weeds, and where it is important to find out the relationships between forest, meadow, steppe, marsh and weed species" (Belgard, 1950).
Choice of ecological scale largely depends on geographical area, biotopes studied and survey specificity. Environmental characteristics associated with phytoindication form the basis of scales of primary factor regimes developed by Ellenberg and other authors. But in the Belgard system, one or another ecomorph reflects the ability of plant species to grow in certain ranges of factors' influence, i.e. it serves as an ecological characteristic of the species. These approaches are quite comparable, and ecomorphs of the Belgard system correspond to certain gradations of the Ellenberg and Tsyganov scales (Table 3-6). Differences in these systems concerning the meaning of soil richness are based on specificity of geographical areas for which the scales have been developed. Authors apply concept of nitrogen richness to the forest zone when soils are characterized by low level of mineralization and a large amount of slowly biodegradable organic matter. For the steppe zone, the concept of soil richness reflects its total nutrient content because soils with greater mineralization and decomposition of organic matter are formed within this zone. And the presence of highly mineralized soils is reflected by the cenomorph of high salinity (alkotrophs).
In his ecomorph system, A. L. Belgard developed and first applied the term "cenomorph". Cenomorph shows the confinement of a plant species to a particular phytocenosis: Sil (Silvaticus) - sylvant (forest species), St (Stepposus) - stepant (steppe species), Pr (Pratensis) - pra-tant (meadow species), Pal (Paludosus) - paludant (marsh species), Aq (Aqantus) - aquant (aquatic species), Ps (Psammophyton) - psammo-phant (species of sandy ecosystems), Pt (Petrophyton) - petrophyte (pet-rant) (species of stony ecosystems), Ru (Ruderatus) - ruderant (weed species), H (Halophyton) - halophyte (species of saline soil), Cu (Cultus) -culturant (cultivated species), etc. (Table 7). Belgard also applied intermediate values to cenomorphs. For example, a complex coenomorph SilPr (forest-meadow) shows the species belonging to different phyto-cenoses, i.e. its cenotic amplitude.
The systems of Ellenberg and Tsyganov were developed for phyto-indication of forest plant communities within the forest zone. And Bel-gard's system of ecomorphs was developed for survey of forest plant communities in the steppe zone. On the one hand, it meets the requirements of these ecosystems, and on the other hand, it is more suitable for phytoindication of different types of plant communities (forest, steppe, meadow, marsh, aquatic, etc.). The Belgard Plant Ecomorph System has a wider range of environmental factors in forest phytocenoses of the steppe zone. It is particularly convenient and can be successfully applied for data processing in ecological analysis both of individual plant communities and flora in large areas (supracenotic level) having considerable species richness. For a more convenient analysis, Matveev suggested a numerical equivalent of ecomorphs (Matveev, 2006).
Application and development of Belgard's Plant Ecomorph System
The Belgard Plant Ecomorph System was applied in several fundamental and applied scientific works on plant ecology and phytocenology (Tarasov, 1981, 2012; Matveev, 1995, 2006; Baranovsky, 2000; Eko-
flora Ukrajiny, 2000; Brygadyrenko, 2016; Baranovsky et al., 2017). Bel-gaid's scale is applied to characterize the environmental features of plant species in the steppe zone with a wide influence range of environmental factors such as moisture, soil richness, mineralization.
In a multi-volume edition "Ekoflora Ukrajiny" (2000-2010), Tsy-ganov's scheme was underpinned by the ecomorph system with numerical gradations of environmental factors, although Belgard's cenomorph
system was used in section "Cenotop" of the monograph (Belgard, 1950). The Belgard Plant Ecomorph System is applied in tabular form (Table 2), so it is convenient for computer data processing in performing ecological analysis of flora of large areas with significant species richness (Baranovsky et al., 2017). The letter designation of ecomorphs allows one to calculate quantitative relationships between ecomorphs using the software Microsoft Excel and even Microsoft Word (using "replacement" option).
Table 3
Light intensity factor according to Belgard's Plant Ecomorph System in comparison with other ecological scales
Complemented Belgard scale (**) (point) Ellenberg scale (point) Tsyganov scale (amplitude)
letter fcological. t optimum, points designation (by Matveev) ecomorphs relation to environment factor points relation to environment factor identifica-tion relation to environment factor points relation to environment factor
USc** 0-1 (0.5) ultra- sciophytes ultra-shade species 1 Extreme shade-loving plants (growing at light intensity up to 1%, rarely at light intensity more than 30%) S ultra-shadow 9 of particularly shady forests
Sc 1 sciophytes obligate shade-loving plant spe cies 2 3 From strongly shade-loving to shade-loving (between 1 and 3 units) Shade-loving plants (grow in light intensity up to 5%, but can grow in lighter places) + s thicket-shadow shady-forest 8 7 of shady forests/particularly shady forests of shady forests
HeSc 2 helio-sciophytes facultative shade-loving plants 4 5 From shade-loving to shade-tolerant (between 3 and 5 units) Shade-tolerant plants (in most cases grow at light intensity more than 10%, as an exception at total illuminance) + M thick light forest light forest 6 5 of light forests/shady forests of light forests
ScHe 3 scio-heliophytes facultative sun-loving plants 6 7 From shade-tolerant to light-loving (between 5 and 7 units, rarely grow at light intensity less than 20%) Sun-loving plants (in most cases grow in total light, but can grow in shade - up to 30%) + g sparse forest shrubby 4 3 of semi-open spaces/light forests of semi-open spaces
He 4 heliophytes obligate sun-loving plants 8 From sun-loving plants to extremely sun-loving (exceptionally the plants can grow at light intensity up to 40%) + forest meadow (sublight) 2 of open spaces/semi-open spaces
UHe** 5 ultra-heliophytes ultra sun-loving species 9 Extremely sun-loving plants (growing only in lighted places, in open areas, at light intensity no less than 50%) G outside-forest (sun) 1 of open spaces
Note: ** - amendments of Matveev (2006). Table 4
Temperature factor according to Belgard's Plant Ecomorph System in comparison with other ecological scales
Complemented Belgard scale (**) (point)
Ellenberg scale (point)
Tsyganov scale (amplitude)
letter designation ecological optimum, points (by Matveev) ecomorphs relation to environment factor scores relation to environment factor identification relation to environment factor scores relation to environment factor
cryophytic species of the polar zone cold climate - K hyper-cryothermic 1st 1 of very harsh winters (average temperature ofthe coldest month < -32)
UOgT** 1 ultra-oligo-thermophytes 1 Arctic and highmountain (alpine and + hyper-cryothermic 2nd 2 of very harsh winters/harsh winters
nival) species L percryothermic 1st 3 of severe winters (average t of the coldest month -24 to -32)
cryophytic species of taiga and tundra 2 from cold to cool (1-3 units) + percryothermic 2nd 4 of harsh winters/moderately harsh winters
OgT 2 oligo-thermophytes 3 cool climate (subalpine heights) M cryothermic 1st 5 of moderately harsh winters (average t of the coldest month -16 to -24)
4 cool to moderate (4-5 units) + cryothermic 2nd 6 of moderately harsh winters/moderate winters
moderately cryophytic species of broadleaf forest zone 5 temperate (warm-temperate) climate N subcryothermic 1st 7 of moderate winters (average t of the coldest month -8 to -16)
MsT 3 meso-thermophytes 6 from moderately warm to warm (5-7 units) + subcryothermic 2nd 8 of moderate winters/mild winters
7 warm climate O hemi-cryothermic 1st 9 of mild winters (average t of the coldest month 0 to -8)
+ hemi-ayothermic 2nd 10 of mild winters/warm winters
MgT 4 mega- heat-loving plants of 8 from warm to extremely warm, sub- P acryothermic 11 of warm winters (average t of the coldest month 0 to +8)
thermophytes steppes and Mediterranean + sub-thermophilic 1st 12 of warm winters/very warm winters
deserts (7-9 units) Q sub-thermophilic 2nd 13 of very warm winters (average t of the coldest month +8 to +16)
UMgT** 5 ultra-mega- heat-loving plants of the tropical zone 9 extremely warm, + thermophilic 1st 14 of very warm winters/unpronounced winters
thermophytes Mediterranean R thermophilic 2nd 15 of unpronounced winters (average t of the coldest month is above + 16)
Note: ** - amendments of Matveev (2006).
Table 5
Moisture factor according to Belgard's Plant Ecomorph System in comparison with other ecological scales
Complemented Belgard scale (* *, ***) (point) Ellenberg scale (point) Tsyganov scale (amplitude)
letter designation ecological optimum, points (by Matveev) ecomorphs relation to environment factor p° int s relation to environment factor identification relation to environment factor points relation to environment factor
very dry habitats (plants growing on dry soils, often on slob habitats) D dry desert 1 desert
UX** ultra- species of very dry 1 + middle desert 2 desertic/semidesertic
_ xerophytes habitats d semidesertic 3 semidesertic
+ desert-steppe 4 semi-desertic/dry steppe
X 0-1 (0.5) xerophytes species of dry habitats 2 from very dry to dry s sub-steppe 5 dry steppe
(between 1 and 3 degrees) + dry steppe 6 dry steppe/medium steppe
MsX 1 meso-xerophytes species of preferably dry places occurring also 3 on fresh soils dry habitats (plants occur on dry soils more frequently than on fresh soils) S middle-steppe 7 middle-steppe
species on fresh soils occur also on dry habitats + fresh-steppe 8 fresh-steppe/meadow-steppe
XMs 1-2(1.5) xero- 4 from dry to fresh habitats C moist-steppe 9 meadow-steppe
mesophytes (between 3 and 5 units) + sub-forest-meadow 10 meadow-steppe/dry-forest-meadow
inhabitants of fresh soils fresh habitats (medium-moist) c dry-forest-meadow 11 dry-forest-meadow
Ms 2 mesophytes 5 + fresh-forest-meadow 12 dry-forest-meadow/moist-forest-meadow
HgMs 2-3 (2.5) hygro- species of fresh soils oc-mesophytes curring also in wet habitats 6 from fresh to moist habitats (between 5 and 7 units) f moist-forest-meadow 13 moist-forest-meadow
MsHg 3 meso-hygrophytes species of wet soils occurring also in fresh 7 moist habitats (well-saturated with water but not wet) + not quite wet-forest-meadow 14 moist-forest-meadow/wet-forest-meadow
habitats F wet-forest-meadow 15 wet-forest-meadow
Hg 4 hygrophytes species of wet soils 8 from moist to wet habitats (between 5 and 7 units) + watery-forest-meadow 16 wet-forest-meadow/swampy-forest-meadow
p swampy-forest-meadow 17 swampy-forest-meadow
UHg** ultra- hygrophytes species of watery habitats 9 wet habitats (mainly oxygen-poor soils) + sub-swampy 18 swampy-forest-meadow/swampy
5 P swampy 19 swampy
HelHg* ** helo-hygrophytes species of temporarily flooded wetlands 10 temporarily flooded habitats (plants of intermittent water bodies) + wetland 20 swampy/shore
HgHel* ** hygro-helophytes species of semi-aquatic habitats a water near the shoreline 21 shore
Hel*** 6 (Hd) ** hydrophytes helophytes species of shallow-water habitats 11 shallow-water environment (amphibious plants) + shallow-water 22 shore/aquatic
Pl*** pleistophytes species floating on the water's surface 12 underwater environment A water 23 water
Hy*** hydatophytes underwater plant, mostly totally immersed in water (underwater plants)
Notes: ** - amendments of Matveev (2006); *** - amendments of Baranovsky (2000, 2017). Table 6
Trophicity factor according to Belgard's Plant Ecomorph System in comparison with other ecological scales
Complemented Belgard scale (**) (point) Ellenberg scale (point) Tsyganov scale (amplitude)
letter designnation points of ecological optimum (by Matveev) ecomorphs relation to environment factor points relation to environment factor desig nation relation to environment factor points relation to environment factor
1 2 3 4 5 6 7 8 9 10
UOg Tr** 0-1 (0.5) ultra-oligotrophs species growing on oligotrophic soils 1 extremely nitrogen-poor habitats j nitrogen-free soil 1 anitrophilous
OgTr oligotrophs species growing on 2 from extremely poor to poor soils + intermediate between j and k 2 subnitrophilous 1st
nutrient-poor soils (between 1 and 3 units) k soils very poor in 3 subanitrophilous
1 nitrogen 2nd
MsOgTr meso-oligotrophs species that grow on poor soils, but can also occur on soils medium in fertility 3 nitrogen-poor habitats (plants occur on nitrogen-rich soils only as an exception) + intermediate between k and l 4 heminitrophilous 1st
OgMsTr oligo-mesotrophs plants growing on soils medium in fertility but can occur on poor soils 4 plants growing on soils rich in nitrogen from poor to moderately (between 3 and 5 units) l soils poor in nitrogen 5 heminitrophilous 2nd
MsTr 2 mesotrophs species growing on soils medium in fertility 5 soils moderately rich in nitrogen (moderately rich) + intermediate between l and m 6 subnitrophilous 1st
MgMs Tr mega-mesothophs plants growing on soils moderate in fertility, but can occur on rich soils 6 plants growing on soils from moderately to high rich in nitrogen (between 5 and 7 units) m soils sufficiently rich in nitrogen 7 subnitrophilous 2nd
MsMg Tr 3 meso-megathophs species that grow on rich soils, but can also occur on soils medium in fertility 7 nitrogen-rich habitats (plants occur on nitrogen-poor soils only as an exception) + intermediate between m and n 8 nitrophilous 1st
MgTr - megatrophs species tending to prefer soil high in fertility 8 from rich to very rich in nitrogen soils (between 7 and 9 units) n nitrogen-rich soils 9 nitrophilous 2nd
1 2 3 4 5 6 7 8 9 10
UMgTr - ultra-megatrophs species tending to prefer the highest-fertility soil - - + intermediate between n and o 10 nitrophilous 3rd
Nitr - nilrophilic group species that grow on soil enriched in nitrogen 9 extremely nitrogen-rich habitats (nitrogen-contaminated) extremely nitrogen-rich soils 11 nitrophilous 4th
AlkTr (not cur- 4 (HMg Tr) ** species tending to prefer saline soils low in fertility not provided because the lack of analogues in Belgard system not provided because not provided because
rently developed) 5 (Hal) ** alkotrophs the lack of analogues in Belgard system the lack of analogues in Belgard system
Ac (not currently developed) - acidophilic group species that grow in more acidic environment not provided because the lack of analogues in Belgard system not provided because the lack of analogues in not provided because the lack of analogues
Ca - calciphilous group species that grow on soil enriched in lime Belgard system in Belgard system
Spr - saprophytes heterotrophic species - - -
Par - biophytes heterotrophic species - - -
S/Par - semi-bicphytes myxotrophic species - - -
Note: ** - amendments of Matveev (2006). Table 7
Cenomorphs according to Belgard's Plant Ecomorph System in comparison with other ecological scales
Letter Points of ecological designation optimum (by Matveev) Ecomorphs Relation to environment factor
Aq - aquant aquatic species
Pal - paludant swamp species
Pr - pratant meadow species
Sil - sylvant forest species
SMn*** - sylvomargoant forest margin species
St - stepant steppe species
Ps - psammophyte species of sandy soils
Pt - petrophyte species of stony biotopes
Ru - ruderant ruderal species
Hal - halophyte species of saline soils
Cul* - culturant cultural species
Notes: "-" is not provided by this system; * - amendments of Tarasov (1981, 2005); *** - amendments of Baranovsky (2000, 2017).
Like any other scientific concepts, Belgaid Plant Ecomorph System can be developed and expanded. Other authors have expanded and supplemented the Belgard Plant Ecomorph System based on its strategy. Several ecomorphs have been later introduced; they reflect intermediate or extreme gradations of factors for terrestrial ecosystems (Tarasov, 1981, 2012; Matveev, 1995, 2006), as well as for aquatic ecosystems (Baranovsky, 2000). Multi-year analysis of ecological features of plant species in their native environment (Baranovsky, 2000, 2008; Belgard, 1950; Matveev, 1995, 2006; Tarasov, 1981, 2012) generated the need to introduce a new cenomorph - sylvomargoant (species of forest margins, from Latin words: Margo - edge, boundary, border, Margino - edged, framing (Dvoreckij, 1976), Margo - forest margin, ad margines silvarum - at margins of deciduous forests (Kirpichnikov & Sabinkova, 1977). The "Sylvomargoant" cenomorph applies to species that can grow on forest margins or forest clearings, but not in the shady areas of forest plant communities (Baranovsky, 2017; Baranovsky et al., 2017).
Vascular plant species of forest clearings present a special ecomorph due to the specificity of natural conditions in this biotope. This is especially true of forest clearings of the steppe zone where the lack of moisture as a limiting factor is more significant and manifests itself in more distinct gradations. Vegetation of forest margins and clearings had previously been allocated in a separate group (Rastenija lesnih poljan i opuschek, 1986). Previously, in ecomorphic analysis the authors attributed these plants to steppe or forest species (Baranovsky, 2000; Matveev, 2006; Tarasov, 2012). However, the majority of these species classified as stepants are not typical representatives of steppe biotopes, and species classified as sylvants almost never occurred in the undergrowth.
Conclusion
The Belgard Plant Ecomorph System has its own characteristics, but it corresponds to the more widespread systems of Ellenberg and Tsyganov. It includes fewer gradations of environmental factors, so it is
more applicable for phytoindication of different plant associations, especially for large areas. Its advantage is that the letter designation of eco-morphs gives the possibility to tabulate calculation of ratio between plant species of different habitats. The Belgard Plant Ecomorph System is especially suitable for characterizing ecological features of higher plant species in biotopes of the steppe zone with a wide range of such factors as moisture and mineralization. The Belgard scale covers specifically a wide range of influence of environmental factors. In his ecomorph system, A. L. Belgard first used the term "cenomorph" which indicates the confi-nedness of a species to a particular phytocenosis. Multi-year studies and analysis of reported and archival scientific materials allow us to propose new ecomorphs for extreme values of factors and separate a new ceno-morph - sylvomargoant - in the context of development of the Belgard Plant Ecomorph System. Botanists and ecologists of Oles Honchar Dnipro National University are successors of Belgard's scholarly traditions. Currently, "ecological passports" have been developed for almost 2,000 vascular plant species of the steppe zone of Ukraine. In this paper, we set ourselves the task of acquainting our colleagues and revealing as fully as possible the essence of Belgard's Plant Ecomorph System. Application of the methodology of this system, as well as the ecomorph system itself, can be useful and applicable by our foreign colleagues.
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