River valleys as the object of ecological and geobotanical research
The river valleys are characterized by high diversity of ecosystems, their originality, complex structure and dynamics. Changes in vertical displacements from the plain to channel in various parts of сatena, which can be seen as an intricate complex integrating coenoses formation, geomorphological...
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irk-123456789-1779152021-02-18T01:26:47Z River valleys as the object of ecological and geobotanical research Дідух, Я.П. Чусова, О.О. Ольшевська, І.А. Поліщук, Ю.В. Геоботаніка, екологія, охорона рослинного світу The river valleys are characterized by high diversity of ecosystems, their originality, complex structure and dynamics. Changes in vertical displacements from the plain to channel in various parts of сatena, which can be seen as an intricate complex integrating coenoses formation, geomorphological and pedogenic processes, are very important. Plant communities serve as indicators of territorial and temporal changes. Combinations of the communities within the ecological series influenced by environmental factors are characterized as ‘ecomers’, which reflect the trends of ecosystem changes, i.e. ȕ-coenodiversity. Quantitative assessment of these plant communities is possible based on the synphytoindication methodology. There were selected three model river valleys, namely, those of the Sluch, Dniester, and Krasna rivers. Comparison of these rivers is of a special interest because they have different dimensions and flow in different natural areas. The topological differentiation in ecological series of the river valleys served for comparison of the ‘ecomers’, i.e. for assessment of ȕ-coenodiversity differentiation towards changes of leading ecological factors. Долини річок характеризуються високою різноманітністю екосистем, їх оригінальністю, складною структурованістю і динамічністю. Важливе значення у формуванні закономірностей територіального розподілу цих екосистем мають вертикальні переміщення від плакору до русла на різних ділянках катени, яку можна розглядати як складний комплекс, де інтегруються геоморфологічні, грунтотвірні процеси та ценозоутворення. Індикаторами цих територіально-часових змін виступають відповідні рослинні угруповання, поєднання яких у межах екологічного ряду під впливом екологічних чинників трактується як екомери, що відображає градієнт змін, тобто beta-різноманіття. Кількісна оцінка цих рослинних угруповань можлива на основі методики синфітоіндикації. Для роботи було обрано 3 модельні річки (Случ, Дністер і Красна), які мають різну розмірність і протікають у різних природних зонах. На їх прикладі проведено порівняння екомерів, тобто оцінено диференціацію beta-ценорізноманіття щодо зміни провідних екофакторів. Долины рек характеризуются высоким разнообразием экосистем, их оригинальностью, сложной структурированностью и динамичностью. Важное значение в формировании закономерностей территориального распределения этих экосистем имеют вертикальные перемещения от плакора до русла на разных участках катены, которую можно рассматривать как сложный комплекс, где интегрируются геоморфологические, почвообразующие и процессы ценозообразования. Индикаторами этих территориально-временных изменений выступают соответствующие растительные сообщества, сочетание которых в пределах экологического ряда под влиянием экологических факторов трактуется как экомеры, что отражает градиент изменений, то есть ȕ-ценоразнообразие. Количественная оценка этих растительных сообществ возможна на основе методики синфитоиндикации. Для работы были выбраны три модельные реки (Случ, Днестр и Красная), которые имеют разную размерность и протекают в разных природных зонах. На их примере проведено сравнение экомеров, т. е. дана оценка дифференциации ȕ-ценоразнообразия по отношению к изменению ведущих экофакторов. 2015 Article River valleys as the object of ecological and geobotanical research / Я.П. Дідух, О.О. Чусова, І.А. Ольшевська, Ю.В. Поліщук // Український ботанічний журнал. — 2015. — Т. 72, № 5. — С. 415-430. — Бібліогр.: 30 назв. — англ. 0372-4123 DOI: http://dx.doi.org/10.15407/ukrbotj72.05.415 http://dspace.nbuv.gov.ua/handle/123456789/177915 en Український ботанічний журнал Інститут ботаніки ім. М.Г. Холодного НАН України |
| institution |
Digital Library of Periodicals of National Academy of Sciences of Ukraine |
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DSpace DC |
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| topic |
Геоботаніка, екологія, охорона рослинного світу Геоботаніка, екологія, охорона рослинного світу |
| spellingShingle |
Геоботаніка, екологія, охорона рослинного світу Геоботаніка, екологія, охорона рослинного світу Дідух, Я.П. Чусова, О.О. Ольшевська, І.А. Поліщук, Ю.В. River valleys as the object of ecological and geobotanical research Український ботанічний журнал |
| description |
The river valleys are characterized by high diversity of ecosystems, their originality, complex structure and
dynamics. Changes in vertical displacements from the plain to channel in various parts of сatena, which
can be seen as an intricate complex integrating coenoses formation, geomorphological and pedogenic
processes, are very important. Plant communities serve as indicators of territorial and temporal changes.
Combinations of the communities within the ecological series influenced by environmental factors
are characterized as ‘ecomers’, which reflect the trends of ecosystem changes, i.e. ȕ-coenodiversity.
Quantitative assessment of these plant communities is possible based on the synphytoindication
methodology. There were selected three model river valleys, namely, those of the Sluch, Dniester, and
Krasna rivers. Comparison of these rivers is of a special interest because they have different dimensions
and flow in different natural areas. The topological differentiation in ecological series of the river valleys
served for comparison of the ‘ecomers’, i.e. for assessment of ȕ-coenodiversity differentiation towards
changes of leading ecological factors. |
| format |
Article |
| author |
Дідух, Я.П. Чусова, О.О. Ольшевська, І.А. Поліщук, Ю.В. |
| author_facet |
Дідух, Я.П. Чусова, О.О. Ольшевська, І.А. Поліщук, Ю.В. |
| author_sort |
Дідух, Я.П. |
| title |
River valleys as the object of ecological and geobotanical research |
| title_short |
River valleys as the object of ecological and geobotanical research |
| title_full |
River valleys as the object of ecological and geobotanical research |
| title_fullStr |
River valleys as the object of ecological and geobotanical research |
| title_full_unstemmed |
River valleys as the object of ecological and geobotanical research |
| title_sort |
river valleys as the object of ecological and geobotanical research |
| publisher |
Інститут ботаніки ім. М.Г. Холодного НАН України |
| publishDate |
2015 |
| topic_facet |
Геоботаніка, екологія, охорона рослинного світу |
| url |
http://dspace.nbuv.gov.ua/handle/123456789/177915 |
| citation_txt |
River valleys as the object of ecological and geobotanical research / Я.П. Дідух, О.О. Чусова, І.А. Ольшевська, Ю.В. Поліщук // Український ботанічний журнал. — 2015. — Т. 72, № 5. — С. 415-430. — Бібліогр.: 30 назв. — англ. |
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Український ботанічний журнал |
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415ISSN 0372-4123. Укр. ботан. журн., 2015, 72(5)
Introduction
The biosphere depends on the water regime which
determines distribution of ecosystems and their
diversity. Rivers play important role and the way they
function depends on many factors such as climate,
topography, geological structure, water resources,
vegetation, etc. On the other hand, rivers are difficult
paragenetic systems which affect the nature of
topography, landscape, soil, and biotic components of
the ecosystems. These functions of the river are defined
by paragenetic structure of the river valley that is formed
by a complex interaction of zonal-regional location,
climate impact, watercourse power, altitude, exposure
and slope of the banks, geological structure, landscape
structure, riverbed nature, presence of terraces, degree
of floodplain inundation, vegetation, hydrological
regime, etc. The movement of physical, chemical, and
biotic components of the energy flows occurs both in
a horizontal direction, from the source of the river to
its mouth, and vertically, from the plain to the channel.
River valleys are the subject of study in many disciplines
such as geography, landscape ecology, and geobotany.
Ranking of the configuration of river basins reflects
particular features of their organization. The lower rank
rivers are known to be more sensitive to seasonal climate
change as well as other phenomena and appear to be
more uniform than rivers of higher levels. In the basins
of lower rank rivers, the volume of flow depends on af-
forestation areas, waterlogging, soil cover, and rainfall,
whereas in the basins of higher rank ones these depen-
dencies entirely disappear and are not present within
the levels 6 and 7 (Hrodzynskyi, 2005).
In addition to zonal-regional characteristics, river
valleys also vary from the source to the mouth and are
divided into the upper, middle and lower reaches, each
with its own characteristics.
http://dx.doi.org/10.15407/ukrbotj72.05.415
YA.P. DIDUKH, O.O. CHUSOVA, I.A. OLSHEVSKA, YU.V. POLISHCHUK
M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine
2, Tereshchenkivska str., Kyiv, 01004, Ukraine
ya.didukh@gmail.com,
chusovaolia@mail.ru
RIVER VALLEYS AS THE OBJECT OF ECOLOGICAL AND GEOBOTANICAL RESEARCH
Didukh Ya.P., Chusova O.O., Olshevska I.A, Polishchuk Yu.V. River valleys as the object of ecological and
geobotanical research. — Ukr. Bot. J. — 2015. — 72(5): 415—430.
The river valleys are characterized by high diversity of ecosystems, their originality, complex structure and
dynamics. Changes in vertical displacements from the plain to channel in various parts of сatena, which
can be seen as an intricate complex integrating coenoses formation, geomorphological and pedogenic
processes, are very important. Plant communities serve as indicators of territorial and temporal changes.
Combinations of the communities within the ecological series influenced by environmental factors
are characterized as ‘ecomers’, which reflect the trends of ecosystem changes, i.e. -coenodiversity.
Quantitative assessment of these plant communities is possible based on the synphytoindication
methodology. There were selected three model river valleys, namely, those of the Sluch, Dniester, and
Krasna rivers. Comparison of these rivers is of a special interest because they have different dimensions
and flow in different natural areas. The topological differentiation in ecological series of the river valleys
served for comparison of the ‘ecomers’, i.e. for assessment of -coenodiversity differentiation towards
changes of leading ecological factors.
K e y w o r d s: river valley, -coenodiversity, catena, ecomer, plant communities
© YA.P. DIDUKH, O.O. CHUSOVA, I.A. OLSHEVSKA,
YU.V. POLISHCHUK, 2015
416 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
The patterns of territorial distribution, the structure
of valleys, and the form of landscape components are
indicators of the processes they were formed by, which
can be described within the framework of fractal theory,
such as sinusoidal tortuosity of channel, arborescence
of hierarchical combination of tributaries, character
of circuses and ravines systems on shores, linearity of
alluvial sediment, shape and location of the oxbow
lakes.
Apart from the patterns that reflect the nature
of regional changes of river valleys and horizontal
movement of the material-energy reserve, change in
vertical displacements from the plain to channel in
various parts of the сatena including water parting,
slopes, terraces, floodplain, and riverbed, is also
essential.
The functioning of the catena depends heavily on
geomorphological factors associated with relief, namely
changes in exposure, steepness, and, consequently, heat,
light, humidity, and evaporation. Significant altitudinal
differentiation of relief in some catenas may result in
vertical change in the altitudinal zones of vegetation
and soils. In this case, the shape of slopes is important.
Convex slopes indicate the presence of active deep ero-
sion, while concave slopes indicate the reduction of soil
erosion and reduction in the rate of vertical movements
(Devis, 1962). Hydrological regime is greatly affected
by the geological structure, which causes the internal
pressure and outflow of ground water to the surface.
Peculiarities of relief lead to changes in the distribution
of sediments, and, consequently, formation of soil and
its wealth. In the eluvial zone, the internal ecogenetic
nature of successions defines the processes of natural
accumulation of matter and organics, leading to autog-
enous pedogenesis. These processes slow down as slope
steepness of the river valley increases. This transit zone
is dominated by the processes of denudation and sur-
face erosion, which cause soil depletion. In turn, soil
depletion halts and even completely stops certain stages
of the vegetation succession. Geological sediments and
organics-rich soils tend to accumulate at the foothills
of the slopes. In some places, colluvial sediments in the
form of talus, landslides, and marl accumulate under
the rocky outcrops. Alluvial fans of shallow sedimen-
tary rocks are observed at the base of the slopes and oc-
casionally can be found even in the floodplain.
All this determines complex nature of the ecosystems
differentiation in the catena. The catena can be regarded
as an intricate complex which integrates formation of
coenoses, geomorphological and pedogenic processes.
Geomorphological processes affect the kinematics of
relief, alluvial, transitive, and accumulative processes.
The formation of coenoses within catena is determined
by successions, which, in contrast to plakor, do not
reach the final stages, but rather transition to the stable
state due to the action of a specific external limiting
factor on a particular succession unit, which should
be interpreted as ecoclimacteric. Nodal stages (links)
of plant communities form through these processes of
certain direction and speed. The order in which these
stages are located in the river valleys are considered as
series (raws). Based on the role of a limiting factor or
a group of thereof, lithosere, hydrosere, psammosere,
and autogenous series can be recognized. Therefore,
the catena allows for the fullest expression of natural
spatial and temporal properties of ecosystems, which
characterize diversity of region’s ecosystems and
its dynamics (Didukh, 2008). In this context, the
vegetation cover is an important indicator of landscape-
territorial and temporal changes. Floodplain exposes
an entirely different pattern of differentiation and
functioning, the one determined by the direct action
of the water environment and can be considered as a
terraqueous component of landscape (Lavryk (2013)
includes riverbed to it). Unlike mesorelief of slopes,
geomorphological differentiation of the floodplain is
not as significant and is caused by microrelief, where
a variegated diversity of habitats is determined by the
layer of groundwater.
In addition to the changes of microrelief proper and
depth of groundwater, seasonal changes in water regime
are of importance as well. Together with other factors,
they determine the type of plant cover. Based on the
duration of flooding, short-term, middling streamside,
and long-term flooding communities are recognized.
Certain indicator species are present in each of these
communities (Ramenskii, 1939; Mirkin 1974; Kurkin,
1976). To assess the variability of damping, there are de-
veloped appropriate integral scales (Ramenskii, 1939;
Didukh, 2012).
In general, the floodplain is a highly dynamic sys-
tem. Floodplain ecosystems are the least stable, because
their resistant stability increases and plastic stability
decreases in the direction from riverbed to slopes and
eluvial plains and vice versa, with plastic stability be-
ing the highest in the ecosystems of hydrogenous type
(Didukh, 2014). Accumulation of mechanical deposits
and organic sediments (the highest biomass productivi-
ty leads to accumulation of highest energy reserves) and
constant turnover of these alluvial deposits are charac-
417ISSN 0372-4123. Укр. ботан. журн., 2015, 72(5)
teristic for the floodplain. Therefore, the floodplain can
be considered as an area of perturbation of paragenetic
configuration of landscape, which is characterized by
high degree of seasonal fluctuation, turbulence, and
even catastrophic processes that are of great interest for
the research of ecosystem dynamics. Unique vegetation
classes (Вidentetea tripartite, Nano-Juncetea, Salicetea
purpureae, Alnetea glutinosae, Phragmito-Magnocarice-
tea) form in the floodplain. Furthermore, floodplains
allow for the incursion of invasive species and often
have the highest concentration of transformer species
(Didukh, 2014).
The third component of the river valley is the chan-
nel proper. Its functions are determined by such factors
as width, depth, water flow capacity, speed and nature of
water movement, seasonal changes in its extremes, bot-
tom sediment substrate, the presence of rapids, flatwa-
ters, pits, chemical composition of water (in particular
the degree of eutrophication), to name a few. This type
of habitats is distinguished by being the most dynamic.
Rapid recovery (plasticity) is the key to their stability.
Specific helophyte communities (Lemnetea, Potame-
tea, Zosteretea), algal and zoological complexes, such
as plankton and benthos, are important components of
these habitats. Indicator scales and indices developed for
the these communities and complexes are used in the as-
sessment of water (Rozenberg, 1984; Didukh, 2012).
It can be concluded that the river valleys are
characterized by high diversity, originality, complex
structure, and dynamics of ecosystems. Plant
communities serve as indicators of territorial and
temporal changes. Quantitative assessment of
these plant communities is possible based on the
synphytoindication methodology (Didukh, 2012).
The success of such research requires logical ordering
of distribution of territorial and temporal components
of ecosystems reflecting these patterns. Methodology-
wise, it is important to shape researcher’s understanding
of the river valley as a complex functional system.
Topological differentiation and temporal dynamics
of the river valley ecosystems
Differentiation of ecosystems can be analyzed from
three angles:
1. typological ( -diversity), reflected in classification of
syntaxa;
2. topological ( -diversity), reflected in the way coenoses
are connected in the landscape (phytocoenomers);
3. territorial ( -diversity), the way coenoses are located
in any given territory, and estimate the size of their
areas (phytocoenokhors) (Didukh, 1995).
Phytocoenomers are understood as order of a set
of communities that are linked to each other within
a certain ecological series, thus reflecting a gradient
of changes occurring throughout these communities,
namely -coenodiversity. When coenotic and ecological
indicators are analyzed, in other words, when changes
at the level of habitats are assessed, phytocoenomers are
understood as ecomers. The terms ‘phytocoenomer’ and
‘phytocoenokhor’ were introduced by V.B. Sochava
(Sochava, 1972; 1979) in the framework of mapping of
vegetation, once it was determined that hierarchy of the
syntaxonomic units does not correlate and cannot be
used for mapping of vegetation at any scale. In Western
Europe this is reflected in Symphytosociology (Tuxen,
1979) and subsequently in the dynamic-catenal phyto-
sociology (Rivas-Martinez, 2005).
Logics behind a legend to a map differs from the logics
behind classification of vegetation. The former depends
primarily on the size of the plots occupied by coenoses
and the way they are linked, rather than on the extent of
taxonomic differences. When similar types of coenoses
are merged into one, their rank is elevated to the
highest syntaxanomic category they have in common.
If, however, coenoses are very different, they cannot be
presented as one system, since their properties expressed
in khoric units are lost. Hence, there is a need to find
other traits or factors that can be used as a basis to form a
unified system. In this case, instead of transcending the
boundaries of the coenotic (syntaxonomic) categories
proper, other characteristics, such as ecological series,
stages of succession are used. In contrast to the well-
developed and fine-tuned syntaxonomic classification,
research into underpinning such systems is in its
infancy. Novel mathematical methodology that can
be used to evaluate the ranks of phytocoenomers and
phytocoenokhors, to determine their scope, degree of
differentiation, subordination, and, first and foremost,
to determine principles upon which a sound system can
be formed, is urgently needed.
Ecomers can be viewed as analogous to stratigraph-
ic columns in geology, with suites being equivalent to
the link of plant communities, facies, and series being
equivalent to eco-topological series of plant communi-
ties. Research on spatial heterogeneity of plant cover at
the level of landscape brings to light yet another prob-
lem, namely, which elementary units (links) should be
used. This problem is further complicated by the con-
418 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
tinuous mode of changes, variation in extent of differ-
ence between adjacent departments, and presence of
different stages of succession.
Rivas-Martinez (2005) proposed to call series of
plant communities the sigmeta. Depending on the
structure, distribution groups and factors that deter-
mine this distribution, he proposed to allocate geosig-
meta, permasigmeta and permageosigmeta and have
even developed principles for giving them Latin names
according to the syntaxon, which represented a particu-
lar link. The development of these ideas finds support
in mapping complex combination of plant communities
(Pedrotti, 2013).
Thus the lowest part of an ecomer is a link. All links
of various stages of succession combined together are
understood as series. Serial communities are the ones
which are at present in the intermediate stages of devel-
opment, relative to the stable (climax) state. Dynami-
cally, each plant association belongs to one or another
of initial, intermediate, mature or deviant stages out of
a vegetation series (Rivas-Martinez, 1987; 2011). Com-
bination of links which change each other in a limited
area and are caused by the limiting activity of a single
or several mutually dependent factors, e.g., humidity,
salinity, drainage, are considered as ecological (eco-
topological) series (Isachenko, 1969; Didukh, 1995).
M.A. Holubets (2005) defines ecological series as a
naturally occurring or imaginary arrangement of living
systems according to the gradient of environment or
according to selected structural and functional indica-
tors of a system that is being studied, provided that the
indicators reflect the gradual changes of the ecological
gradient. The combination of regularly occurring di-
verse communities, which are distributed based on the
particulars of micro-relief forms, occupy limited areas,
and do not form elements of landscape units is called
a ‘complex’. Regularly located communities (or com-
plexes) that reflect the territorial distribution of com-
munities caused by changes in mesorelief are consid-
ered as mesocombinations. Totality of heterogeneous
communities, complexes or mesocombinations which
reflect change in ecological conditions of macrorelief
are known as macrocombinations (Isachenko, 1969).
Since the actual slopes of the river valley, floodplain,
and channel are interconnected as spatial paragenetic
parts of a landscape, they are combined at the level of
the macrocombination. However, because they differ
in structure, influence of limiting factors, the nature
of dynamic processes, they are assigned to separate
mezocombinations, which may be of different size
and structure. The banks can be steep, high, slopes —
precipitous, floodplains — absent, and vice versa, a
wide floodplain with oxbows, low banks, and slight
terraces. The same applies to the relation between these
components of the river valley and the channel.
Distribution of communities across the catena of
slopes and on the floodplain, direction and speed of
successions are different. Changes in habitats of the
slopes are defined by the changes in mesorelief and are
considered as mesocombination determined by the in-
fluence of a single system-forming ecofactor, which al-
lows to maintain the state of dynamic equilibrium with-
in the given system. Elementary mesocombination is a
combination of coenoses represented by phytokhor of
landscape level (100 × 100, 200 × 200 m), and mapped
on the scale of 1:100000 (Yurtsev, 1988). Mesocombi-
nation is formed by a set of eco-topological series of a
given landscape, which changes depending on altitude,
steepness of slopes, and geological rock (Belikovich,
2000). In a different landscape, certain series, or links
of some series can vanish altogether.
In the floodplain, phytocoenosis changes are
determined by microrelief. It is within microrelief
that fluctuations in groundwater and type of alluvial
deposits are observed, and, consequently, processes of
soil formation, which determine the type of succession,
transition from peat-swamp, silty-marsh to meadow
alluvial type. This natural change of communities
from coastal vegetation to riverine sediment manes
and shafts within a microrelief are considered as
complexes. However, great variety, large areas of similar
sites (links) and complexes in the floodplain allow to
consider the totality of these elements on the level of
mesocombination. Patterns of change of phytocoenoses
(biotopes) within the riverbed are tied to the depth of
water, speed of its movement, and type of the bottom
substrate. Transition from hydrotops proper to floating
(Lemnetea), rooted (Potametea) and coastal hydro-
aerial communities (Phragmito-Magnocaricetea)
forms ecological series, which can be joined into
mesocombinations.
Therefore, the cross-section of the river valley can
be viewed as an ecosystem of landscape level, which is
characterized by a conjunction of three types of meso-
combinations: valley slopes are a mesocombination of
pedo-lithogenic type, floodplain is the one of pedo-
hydrogenous type, and channel — of hydrogenous type
proper; each of these combines individual links (specif-
ic plant communities or habitats) and includes relevant
complexes, series (Fig. 1). The plakor portion is consid-
419ISSN 0372-4123. Укр. ботан. журн., 2015, 72(5)
ered as a separate mesocombination or even a complex
of thereof, and is a subject of a separate analysis. The
totality of mesocombinations within given landscape is
treated as a macrocombination.
Mesocombination changes, loss of individual series,
which are observed from the upper river to its mouth
characterize regional, ecokhoral aspects which, albeit
important, are not going to be considered here. The
phenomenon of a shift in phytokhorion boundaries
along river banks due to the presence, on the southern
slopes, of communities of a vastly different type from
the ones characteristic for the given region, is well-
known. Particularly relevant is the question whether
the river valley can be interpreted as a barrier (bound-
ary) between the ecokhors of regional level (zoning), or
whether it is a link within the zoning units. When large
rivers are considered, their differentiating role is not
questioned, whereas the role of middle rivers is up for
discussion. In traditional geo-botanical zoning, when
khorions were delimited based on the characteristics of
plakor vegetation, boundaries between khorions were
generally tied to rivers. However, eco-geo-botanical
zoning of Central-Podilskyi geobotanical district, con-
ducted by our group based on the analysis of distribu-
tion of all types of communities when plakor vegetation
is violated or destroyed, has led us to question how ap-
propriate it is to establish such boundaries strictly based
on the plakor areas (Didukh, Vasheniak, 2013).
River valleys are interesting from the standpoint of
dynamics, kinetics of ecosystems, which explore pat-
terns underpinning changes in state of certain points
(elements) in time. Kinematic approaches have already
made their way into geomorphology (Devdariani, 1964);
however, they are relevant to ecology as well. Environ-
Fig.1. The cross-section of the river valley
Symbols: — 1; — 2; — 3; — 4; — 5; — 6; — 7; — 8; — 9
Legend: 1 — Quercus robur; 2 — Prunus spinosa; 3 — Vicia angustifolia; 4 — Poa angustifolia; 5 — Koeleria cristata; 6 — Festuca
pratensis; 7 —Salvia pratensis; 8 — Alopecurus pratensis; 9 — Phragmites australis
MACROCOMBINATION MACROCOMBINATION
mesocombination
mesocombination mesocombination
complex
link
mesocombination
420 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
mental kinematics is clearly a promising direction of
future research, and a lot of valuable data can be gained
from research on the river valley ecosystem dynamics.
Of particular interest, from this standpoint, is research
on positional-dynamic and paragenetic aspects, which
reflect the type, speed, and magnitude of change in
matter-energy processes (Hrodzynskyi, 1993), as well
as direction, speed, degree of stability, risk of loss, ap-
pearance of some and disappearance of other elements,
all of which are the basis for predicting future changes.
Furthermore, successional development of vegetation
has long been known to be complicated by the large-
scale fluctuating changes, reflected in the fact that, in
different years, vegetation of the same plot was recorded
as belonging to different formations (Shennikov, 1924;
Rabotnov, 1974).
The problem of the dynamics of ecosystems is par-
ticularly relevant due to the spread of the invasive trans-
former species, which change the structure of natural
coenoses. The fact that river valleys serve as not only
migration routes for the hydrophytes proper, which are
waterborne, but also for species that, at first glance, are
not directly connected to water, has been recorded in
botanical literature long ago (Shennikov, 1924). Type
of mezorelief provides far more diverse conditions than
the plakor areas. When zonal climatic conditions are
similar due to drainage, angle of slope (steepness), ex-
posure, and baseline erosion, indicators of the thermal
regime are higher and those of humidity are lower than
on plakor. This, in turn, leads to a significant increase
in intensity of evaporation and affects the hydrothermal
regime. That said, excessively humid habitats of flood-
plains are the coldest. Observed elevated extent of plant
cover differentiation occurs because of this high gradi-
ent of changes in hydrothermal regime. At the same
time, species distributed along the border of their range
are of narrower ecological amplitude relative to zonal
species, and, under such diverse ecological-coenotic
conditions, are able to find plots suitable for their distri-
bution. The river valleys are highways spreading south-
ern, more xerophytic species up north and northern,
boreal species down south (Didukh, 2008).
This is of particular importance for the climate
change research. We associate distribution of invasive
species along the river valleys with the indirect climate
changes. Temperature rise has been proven to affect
ecosystems of high, 'cold' latitudes the most. Of all
ecosystems of our temperate latitudes, the coldest and,
thus, the most vulnerable are hydrophilic ones, and
the toughest ones are the dry, lithophilic ecosystems.
Furthermore, the hydrophilic ecosystems of floodplains
transformed the most through reclamation, creation of
reservoirs, dams, and straightening of channels, which
all cause eutrophication. Together, all of these factors
determine the type of transformation of the floodplain
ecosystems.
Therefore, research on the patterns of spatial-
temporal changes and functioning of the river
ecosystems is of utmost importance.
Methods of ecomer assessment
Field research of these aspects involves establishing a
system of ecoprofiles and visual assessment of changes in
plant cover based on the traits of the dominant species,
which reflect the lowest level of syntaxa (variants). The
geobotanical descriptions are provided for each site. If a
given plot is small, several plots, with similar plant cover,
relief, geology, and soil are described. Depending on the
type of limit of important ecological factors and their
interrelation (orographic features, height and steepness,
slope exposure, outcrops of the geological rocks and
their location, the impact of anthropogenic factors, type
of soil, among others), each specific ecoprofile reflects
this distribution, which is determined by the interaction
of various factors. The task to be completed is to describe
sufficient number of ecoprofiles to adequately represent
all types of combinations. The next step is to process the
data collected during the field research, and to create a
database of geobotanical descriptions and to calculate
ecofactor indicators based on phytoindication scales.
Such calculations will allow to correct the position of
the individual links relative to changes of indicators of
ecofactors in order to construct ecological series.
In theory, these series can be constructed based
on indicators of all 12 ecofactors. However, that is
not necessary, and such series are constructed with
respect to changes in key ecofactors or a group of
thereof, or based on the corresponding structure of
the coenoses (e.g. lithogenous, sylvatic, hydrogenous,
psammogenous) (S m rghi an, 2003). Ecological
series should be distinguished from dynamic ones,
which reflect successional development. Consequently,
their links are represented by time-serial communities.
At the same time, syntaxonomic vegetation
classification is constructed. Indicators evaluating
similarity characteristics (floristical, ecological)
between syntaxa, type of displacement of dominant
species or individual groups are calculated as well.
The next step is the selection of ecomers
(mesocombinations). The essence of this step lies
421ISSN 0372-4123. Укр. ботан. журн., 2015, 72(5)
in construction of one or two conditional series
based on actually existing profiles, ecological series,
and classification data. Conditional series, in turn,
characterize patterns in link distribution for a given
region. Such series (mesocombinations) differ from
ecological series in that the latter reflect how the links
are ordered (the order of links) relative to changes in
the dominant factor or several, mutually dependent
dominant factors, such as change of steppe, forest,
psammophyte vegetation.
Mesocombination reflects the sequence of links
typical for this particular region. For example, the
upper part of the slope of the bank can be composed of
quaternary loess rocks, on top of which develop steppe
communities (Festucion valesiacae, Fragario viridis-
Trifolion montani), whereas on the slope of a bank,
where the soil is washed away, steppe communities are
represented either with certain differences or entirely
different syntaxa. Furthermore, in the middle of the
slope it is possible that the outcrops of tertiary limestone
or chalk can occur in the middle part of a slope, and
Devonian or Silurian sediments might be present
in the bottom part of the slope. These links can be
represented with Stipo pulcherrimae-Festucion pallentis,
Alysso-Sedetalia, and, on the cliffs, with Asplenietea
trichomanis communities. At the base of the slopes,
where collin deposits of rocks accumulate, communities
of the class Thlaspietea rotundifolii, shrubs (cl. Rhamno-
Prunetea), or the edge coenoses (cl. Trifolio-Geranietea)
form. Further down, where alluvial sandy sediments
occur, Agrostion vinealis coenoses are present, which
transition into meadow communities on the terraces
and floodplains. In other cases, different pattern of
alternation of coenoses may occur, e. g., typical nemoral
forest coenoses (Carpinus-Quercus) are replaced by the
thermophilic forests (Quercion petraea), but on the
«foreheads», open edge coenoses (Trifolio-Geranietea)
are observed. These latter coenoses transition into the
meadow—steppe coenoses (Cirsio-Brachypodion),
which, in turn, are replaced with steppe shrubs (Prunion
fruticosae, Berberidion vulgaris). Forests of the Alnion
incanae alliance are common at the bottom of the slope.
Thus, on one hand, each mesocombination
combines elements, links of various ecological series,
and, on the other hand, includes elements that could
be absent from any given actually existing profile due to
various reasons.
Research results
Let us consider this approach on the example of specific
valleys, namely, those of the Sluch, Dniester, and
Krasna rivers. Comparison of these rivers is of a special
interest because they have different dimensions and flow
in different natural areas. However, we did not intend
to provide an exhaustive description or comparison of
the ecomers of these rivers, which would require more
extensive data collection. In the present study, we were
merely trying to illustrate how the methodology of this
approach could be practically applied.
The Sluch river takes its source in the Forest-Steppe
zone of Podillya Upland near the village Chervony
Sluch of Teofipol District, Khmelnitsky Region, flows
through Polissya lowlands, and falls into the Horyn river.
Based on the peculiarities of physical and geographical
characteristics, the river is divided into upstream,
middle part, and downstream. The upper part of the
river stretches from the source in Khmelnitsky Region
to the town of Myropil (Romaniv District, Zhytomyr
Region), where the banks of Sluch are composed of
loess rocks. The middle part of the river cuts through
the Ukrainian Crystalline Massif from Myropil to
Sosnove village (Bereznovsky District, Rivne Region),
and the downstream part stretches from Sosnove to the
confluence into the Horyn river in Rivne Region. The
Sluch is a stream of the fourth order, with length of 451
km (Hrodzynskyi, 2007). Width of its valley is from 0.2
to 0.8 km in the upper part, to 5 km downstream. Width
of the channel is from 5 to 50 m, with a maximum of
110 m. The slope of the river is 0.4 m / km.
Studies were conducted in the upstream and in the
middle part of the river. Eco-coenotic profiles were
established in the environs of the town of Staryi Ostropol
(Starokostiantyniv District, Khmelnitsky Region). The
South-Eastern slope of the left bank of the Sluch river
is 20 m high. Its upper part is occupied by coenoses of
Artemisio marschalliani-Elytrigion intermediae alliance
(dominated by Elytrigia intermedia (Host) Nevski,
Bromopsis inermis (Leyss.) Holub, and occasional
Chamaecytisus ruthenicus (Fisch. ex Wol.)Klask.).
Coenoses of Festucion valesiacae alliance with dominant
species Festuca valesiaca Schleich. ex Gaudin, Medicago
falcata L., and some Carex humilis Leyss. are present in
the middle part of the slope, and bottom part of the slope
is occupied by the Koelerio-Corynephoretea coenoses,
with Sedum acre L., Euphorbia cyparissias L. Melica
transsilvanica Schur being the dominant species. This
422 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
series combined meadow-steppe vegetation in the upper
part, and psammo-pteridophytic in the bottom part.
Another profile was established near the village
Mala Tokarivka (Romaniv District, Zhytomyr Region)
on the left bank of Sluch, whose height reached 15
m, where the conditions are more mezophytic than
in the previous site. Upland area was represented by
the meadow coenoses of Cynosurion cristati, the well-
heated and well-drained upper part of the slope — with
Cariceto praecocis-Thymetum marschalliani, the middle
part of the slope (with the 20° steepness) — by Artemisio
marschalliani-Elytrigion intermediae (with dominant
Elytrigia intermedia), and the bottom part — by
Arrhenatherion (with dominant species Briza media L.).
Geobotanical descriptions of these plots were used
to develop syntaxa classification and to perform their
comparison based on the indicators of 12 ecofactors
according to the method of synphytoindication.
Dniester is one of the longest rivers in Ukraine, with
length of 1362 km. It originates in the north-eastern
slopes of the Ukrainian Carpathians, flows through the
Pre-Carpathian basin, Lviv basin, southern Podillya
Plate, slopes of the Ukrainian Crystalline Massif, Black
Sea lowlands, and falls into the Black Sea. Based on the
peculiarities of orographic characteristics, Dniester is
divided into upper (Carpathian), middle (Podolian),
and lower (Black Sea plain) parts. The middle part of
the river is the most interesting and unique. Here, the
valley narrows and deepens, acquiring the canyon-like
shape, with steep slopes (30—70°), cliffs (over 80°) up to
250 m high, and winding meanders.
On the cross-section profile of the banks, three
parts can be clearly distinguished. Upper part, usually
a gentle slope composed of Quaternary loam, clay, and
Miocene deposits; middle part, with cliffs composed
of solid rocks of middle Baden (gypsum, limestone),
chalk (limestone, sandstone), Silurian and Devonian
(limestone, dolomite, mudstone and marl) deposits; and
lower part, where products of these rocks accumulate.
This three-tiered profile of the Dniester Canyon plays
an important role in the development of exogenous
processes in the valley. The upper and middle parts
of slopes are affected by such processes as erosion,
sheet flood, in the middle part water denudation and
landslides occur, and at the bottom part accumulative
processes take place (talus, colluvial deposits and
alluvial fans are formed).
Dniester valley is presented by the complex of
heterochronous terraced levels. Of the 7—11 currently
recognized levels of the terraces (Herenchuk, 1980),
only terraces of the levels 1—5 are present within the
canyon. Fourth Middle Pleistocene terrace is located at
the height of 70—60 m, third — at the height of 50—
45 m. Narrow bottom of the valley is formed by Late
Pleistocene terraces, the second one at the height of
35—25 m, and the first one at the height of 15—10 m.
Terraces are expressed fragmentarily on convex, sloping
areas of meanders.
Floodplain is rather narrow and clearly separated
from the terraces by steep ledges. It can be divided into
high (7—5 m) and low (3.5—0.5 m) parts.
Gray-forest and black-forest soils have formed in
loess-loam and loess areas; sod-carbonate soils (ren-
dzinas) formed on carbonate rock weathering products.
Rendzinas are typically washed away on steep slopes.
Certain peculiarties of the climatic characteristics of the
Dniester Canyon can be attributed to the Orographic
heterogeneity surfaces. A specific microclimate forms
on the slopes and near foot of the slope. This micro-
climate causes the effect of Transdnistrian «warm Po-
dillya». The effect is caused by the canyon-like valley
that serves as a barrier that protects from the wind and
retains heat accumulated by the slopes from solar radia-
tion. Consequently, elevated temperatures are observed
within the canyon. Winter period here is 20—25 days
shorter, and the average temperature of air 1.5—2.0° С
higher than that of the uplands. This difference in tem-
perature has a marked effect on vegetation of plants
(Herenchuk, 1980; Denysyk, 2007).
Warm and mild climate, diversity of geological and
geomorphological structures, and edaphic conditions
contributed to formation of nemoral forests with
elements of Mediterranean flora, Central and Eastern
European steppe coenoses, and petroglyphic vegetation
within the canyon, which includes a large number of
endemic, relict and rare species. Accordingly, diverse
plant communities, which form certain ecological
series, are present in the valley. Steppe-petroglyphic
ecological series occurs on the southern slopes: Festuca
valesiaca and Bothriochloa ischaemum (L.) Keng are
dominant in the upper part of the slope near plakor
Teucrium chamaedrys L., Festuca ovina L. — in places
of quaternary rocks denudation on convex slopes
Seseli hippomarathrum Jacq, Teucrium pannonicum
A.Kern. are dominant on petrous limestone screes
Teucrium chamaedrys, Melica transsilvanica — further
down, on the colluvial deposits. Another Steppe series
is represented by the following change of dominants:
Poa angustifolia L. on plakor (upland) Festuca
valesiaca, Bothriochloa ischaemum dominate dry areas
423ISSN 0372-4123. Укр. ботан. журн., 2015, 72(5)
near plakor Stipa capillata L. is dominant on convex
slopes Elytrigia intermedia — in areas with loess
deposits Poa versicolor Besser is a dominant species
on the outcrops of limestone with intense denudation
Teucrium chamaedrys, Melica transsilvanica — on
the colluvial deposits Agrostis tenuis Sibth., Briza
media — on alluvial sediments. Presence of the Central
European Steppes, forming a following ecological series
of changing of dominants on the northern slopes, is a
peculiar feature of this part of the valley: Brachypodium
pinnatum (L.) P. Beauv. is a dominant species on
the even areas Carex humilis (Sesleria heufleriana
Schur) — in the upper part of the convex slopes
Seseli hippomaranthrum, Teucrium pannonicum — in the
middle part and on carbonate cliffs Inula ensifolia
L., Anthericum ramosum L. — further down, in areas
with intense denudation and flushing carbonates
Melica transsilvanica, Teucrium chamaedrys, Bupleurum
falcatum L. — in the bottom part, on colluvial and diluvial
deposits. Shrub series on dry slopes is characterized by
Prunus spinosa L. growing on the top Lembotropis
nigricans (L.) Griseb. is a dominant species further
down Caragana frutex (L.) K. Koch — in the dry
conditions of middle part Swida sanguinea (L.) Opiz,
Ulmus minor Mill. — in the bottom part. Forest series is
represented with Fraxinus excelsior L., Carpinus betulus
L., and Quercus robur L. on flat plakor areas Carpinus
betulus — on slopes near the top Carpinus betulus,
Quercus robur, and Stellaria holostea L. Quercus
petraea (Matt.) Liebl., Cornus mas L., and Aegonychon
purpurea-coeruleum Holub. — on the convex southern
slopes Fraxinus excelsior, Aegopodium podagraria
L. — in the bottom part, on accumulative soils. These
examples do not exhaust the list of series of the middle
Dniester, which are presented by various combinations
and require further study.
The Krasna river is a left feeder of the Seversky
Donets and flows within the southern part of Central
Russian Upland (Steppe zone) from north to south. The
length of the river is 124 km, the drop height of riverbed
is 0.5 m/km. River valley is quite wide and its length
is 3.5 km, while the width of the floodplain is 200—
250 m. The river is fed predominantly by meltwater.
Geomorphologically, the area is located 150—200 m
above the sea level and is composed of cretaceous rocks,
which are well developed here. Carbon rocks form the
base of this territory, with Upper Cretaceous deposits
further up, and, finally, the Quaternary loess. Relief is
dominated by the denudation forms. The right bank
of the river is fairly high (up to 60 m), steep (30—35°),
sometimes scarp, frequently dissected by ravines mostly
in the west-east direction. The ravines are fairly long for
the most part, and assume the shape of a «circus» along
the banks of the Krasna river. Outcrops of Cretaceous
rocks, sand, and loess deposits are observed here. The
left bank of Krasna is gently sloping (3—7°). Three
wide terraces can be traced within the right bank:
floodplain, sandy terrace, and a terrace above the sandy
terrace proper. The latter terrace transitions into plakor
(Ekomerezha, 2013).
In regard to climate, the investigated area is
characterized by average rainfall of 450—500 mm.
The average annual temperature is 7—8º C, average
January temperature is from –8 to –6º C, average
July temperature is from 21 to 22º C, indicating
subcontinental (subarid) climate with deficit of moisture
and drought in August and September.
Medium-humic blackearth predominate in the soil
mantle. Rendzinas (sod-calcareous soils) or chalk
outcrops occupy the slopes of the right bank (Hrytsenko,
2004).
The vegetation of the valley is represented by
deciduous ravine forests, with plakor occupied by sites
of forb-fescue-feather-grass steppe. Chalk outcrop
vegetation is present on the steep right bank, along with
forb-herb-grass meadows which form a patchy complex
of combinations in the catena and are characterized by
high gradient of changes of various ecological factors.
The nature of the chalk outcrops (the presence or
absence of soil cover, the mobility of the substrate,
chemical and physical properties of the chalk) is one of
the leading factors of formation of ecological series in
the valley of the Krasna river.
Forest vegetation in the floodplain areas is represented
by communities with dominance of Alnus glutinosa (L.)
Gaertn., Salix alba L., and Populus nigra L., whereas
communities with Fagetalia sylvaticae (Scillo sibericae-
Quercion roboris) are present on the slopes of ravines and
areas adjacent to plakor. Shrub steppe, with dominant
species Caragana frutex and Prunus stepposa Kotov,
can often be encountered on the slopes of the valley
and ravines. Communities of Amygdalus nana L. and
Spiraea hypericifolia L. are more rare. In more humid
conditions, at the bottom, thickets of Acer tataricum L.,
Rhamnus cathartica L., Padus avium Mill. occur.
Steppe series form on rendzinas and on the blackearth
soils with underlying loess. Gentle slopes of the upper
part are occupied by forb-grass steppe of Fragario
viridis-Trifolion montani alliance. Festucion valesiacae
steppe coenoses are common in drier areas adjacent
424 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
to plakor. In places with good drainage, communities
of Artemisio marschalliani-Elytrigion intermediae
alliance are formed, and, further down, typical steppe
communities Stipion lessingianae are present.
In the middle reaches of the Krasna river, the right
bank is very steep and is characterized by the most
flushing regime. Chalk outcrops are observed here, and
the ravine system is represented by narrow ravines with
steep slopes. From upland to foot of the slope, the fol-
lowing series is usually formed: Stipion lessingianae
Centaureo carbonatae-Koelerion talievii Artemisio
hololeucae-Hyssopion cretacei Festucion valesiacae
Artemisio marschalliani-Elytrigion intermediae.
The steeper are the slopes, the bigger is the area
occupied by communities with dominance of Artemisia
hololeuca M. Bieb. ex Besser (predominantly on the
slopes with eastern exposure), and by communities
with Hyssopus cretaceus Dubj. as a dominant species.
The latter communities favor slopes with eastern and
southern exposure, composed of denser chalk. On more
gentle slopes, less affected by erosion and land-loss,
the communities of tomillares are either fragmented
or do not form at all. Under such conditions, lower
alpine communities Centaureo carbonatae-Koelerion
talievii with dominant species Thymus cretaceus Klokov
& Des.-Shost. and Pimpinella tragium Vill. occur
on the loose chalk substrate. Centaureo carbonatae-
Koelerion talievii communities alternate with xerophyte
communities, such as Festucion valesiacae or Artemisio
marschalliani-Elytrigion intermediae. At the bottom of
the slopes, in the area with talus material accumulation,
xerophytic communities with considerable share of
Krascheninnikovia ceratoides (L.) Gueldenst. are formed.
These coenoses transition into Artemisio marschalliani-
Elytrigion intermediate grass communities, followed by
meadow-steppe communities of the floodplains.
Thus the following series of coenoses is formed: Sti-
pa capillata + Festuca valesiaca S. lessingiana L. +
F. valesiaca Artemisia hololeuca Thymus cretaceus
+ Pimpinella tragium Bromopsis inermis + Elytrigia in-
termedia Krascheninnikovia ceratoides. Сoenoses of
tomillares are not present when this series occupies gen-
tle slopes. Shrub communities with dominance of Cara-
gana frutex are often formed on plakor and upper parts
of the steppe slopes of the northern, northeastern, and
eastern exposures. The latter species is adapted to steppe
conditions and can withstand considerable grazing and
mowing. Therefore, its communities often develop
Fig. 2. Patterns of distribution of the mean values of leading ecological factors in river valley catena: 1 — Sluch; 2 — Dniester;
3 — Krasna
1
2
3
425ISSN 0372-4123. Укр. ботан. журн., 2015, 72(5)
where grass communities would have otherwise thrived.
An example of ecological series involving Caragana frutex
is as follows: Stipa capillata + Festuca valesiaca Festu-
ca valesiaca + Poa angustifolia Caragana frutex + Poa
angustifolia + Elytrigia intermedia Bromopsis inermis
+ Elytrigia intermedia Thymus cretaceus + Pimpinella
titanophyla, and further down Ceratoides papposa.
Comparative analysis of ecomers on river's slopes.
The examples of changing links in ecological series of
the valleys of three model rivers served for comparison
of ecomers i.e. for assessment of -coenodiversity
differentiation towards the changes of leading ecological
factors.
Ecological factors by the nature of the mean value of
bias can be divided into two groups: the first in which
the highest rates associated with the moisture regime
(soil humidity, variability of moisture, aeration, mineral
nitrogen's content in soil, humidity) are characteristic
for the Sluch valley, and the second, with the chemi-
cal properties of soil (salt regime, acidity, carbonate
content and continental climate), typical of the Krasna
valley (Fig. 2). Indicators of the Dniester valley occupy
an intermediate position, except termal climate and
crioclimate, which are the highest ones here, leading to
the effect of «warm Podillya». Thus, the mean values in
general depend on regional location of the valleys and
in the system of global ecological space according to cli-
mate indicators occupy the optimal position (50—60 %
of the scale), except for humidity of the Krasna which
is at around 45 %.
By the indications of soil humidity (Hd) ecomers
of the Krasna (6.5—9.3 points) and Sluch (9.2—11.4
points) river valleys are characterized by the widest
gradient among groups of grassy slopes and of the
Dniester valley — by the narrowest one (8.0—9.2
points). Although given forest communities, the picture
changes dramatically and gradient of ecomers of Sluch
is the narrowest and the widest is on valley Krasna
(Fig. 3, a). The stabilizing role of forests is to increase
the ecosystem humidity by two points approximately
as it can be seen on these data. This is also confirmed
by indicators of soil aeration (Ae), which is closely
correlated with the previous one (Fig. 3, c). Important
role in the accumulation and nitrogen mineralization
play the forests, where the figures differ significantly
from those of herbaceous communities (Fig.3, h).
Herewith, among grass habitats the wettest indicators
Hd are for the Sluch, accordingly, the driest ones are for
the Dniester and Krasna valleys, while the driest forests
are characteristic for the Dniester valley. With regard to
the variability of damping, these figures for all valleys
have similar amplitudes (2 points); however, they are
naturally reduced from the Sluch to Dniester and to the
Krasna valley (Fig. 3, b).
Indicators of chemical properties of soil vary in
the opposite direction. The lowest and the narrowest
parameters Rc, Sl and Ca are characteristic for the
Sluch, the larger and wider ones — for the Dniester
and the highest gradient — for the river Krasna, where
the biotopes from loess and arid slopes to tomillares
are represented. The indicators Rc, Sl, Ca of these
coenoceses does not overlap with other and differ
significantly, indicating a different type of conditions of
their existence (Fig. 3, d—f).
Estimating of ecomers by the climatic indicators is
important. The warmest biotopes (Tm, Cr) are charac-
teristic for the canyonlike Dniester valley, not for the
open steppe of the Krasna river valley (Fig. 3). As for
those environmental parameters that are associated
with precipitation (Om, Kn), a clear zonal pattern of
their changes is observed (Fig. 3, a, i). Forests increase
these figures in the first case and reduce in the second,
although not essential, wherein range of amplitudes of
all four climatic factors increases from the forest zone
to the steppe.
The next stage of analysis of the biotopes in river val-
leys was to assess their similarities-differences by com-
plex of ecological performance. For this purpose, a den-
drogram was built by Ward method that reflects Link-
age Distance. Dendrogram at D>30 is divided into two
large clusters according to the habitat specifics, not to
the river valley grounds. The «A» cluster can be called a
«humid» and the «B» cluster an «arid» (Fig. 4). The first
in its turn is divided into two groups at level D>10: A
1
—
nemoral deciduous forests of river basins, A
2
— grassy
meadows communities (A
21
, A
22
), which at the highest
level (D>5) are adjacent to xerophytic forests Querceta-
lia pubescenti-petraeae and rock communities Asplenion
rutae-murariae (A
23
). This strange combination can be
explained by the fact that petrophytic rock communities
were not available elsewhere except the Dniester valley.
Instead, A
21
and A
22
clusters include strictly speaking
biotopes of the Sluch valley. The «arid» «B» cluster has
a lower degree of differentiation and at the level (D>5)
is divided into three parts: B
1
— steppe biotopes, B
2
—
426 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
Fig.3. Distribution of xerophytic steppe communities depending on ecological scales: a — soil humidity; b — variability of
damping; c — aeration of soil; d — nitrogen content; e — acidity; f — total salt regime
a b
c d
e f
427ISSN 0372-4123. Укр. ботан. журн., 2015, 72(5)
Continuation fig.3. Distribution of xerophytic steppe communities depending on ecological scales: g — carbonate content in
soil; h — thermal climate; i — cryo-climate; j — humidity; k — continental climate; l — light in community
Legend: 1. Carpinion betuli; 2. Querco - Fagetea, Quercetalia robori-petraeae; 3. Agrostion vinealis; 4. Brizeto mediae - Anthoxanthetum
(Cynosion cristati); 5. Artemisio marschalliani-Elytrigion intermediate; 6. Festucion valesiacae; 7. Thymo pulegioides-Sedion
sexangulare; 8. Asplenietea trichomanis, Androsacetalia vandellii (with dominant species Asplenium trichomonas); 9. Carpinion
betuli; 10. Qurcion petraeae; 11. Prunion fruticosae; 12. Festucion valesiacae; 13. Artemisio marschalliani-Elytrigion intermediate;
14. Allyso-Sedion (with dominant species Sedum acre); 15. Allyso-Sedion; 16. Galio campanulatae-Poion versicolor; 17. Asplenion
rutae-murariae; 18. Scillo sibericae-Quercion roboris; 19. Prunion fruticosae; 20. Fragario viridis-Trifolion montani; 21. Artemisio-
Kochion; 22. Artemisio marschalliani-Elytrigion intermediae; 23. Festucion valesiacae; 24. Stipion lessingianae; 25. Centaureo
carbonatae-Koelerion talievii; 26. Artemisio hololeucae-Hyssopion cretaceae
g h
i j
k l
428 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
desert steppes Artemisio-Kochion, B
3
— tomillares Ar-
temisio hololeucae-Hyssopion cretaceae and petrophytic
steppes Centaureo carbonatae-Koelerion talievii.
In turn, B1 is divided into two clusters at the level
D>4: B
11
— steppes and petrophytic communities of
the Dniester river and B
12
— steppe communities of
the Krasna river, adjacent to loess outcrops biotopes
Artemisio marschalliani-Elytrigion intermediae (13) of
the Dniester valley, which are also represented on the
slopes of the Krasna valley (22). At the same time, we
note that communities of this union are also present in
the Sluch valley (5) although they are quite different
from the south ones.
Conclusions
Analysis of ecomers of the river valleys indicates that
each valley includes diverse, enough differentiated by
the ecological conditions habitats, which analogues
may occur in different areas and different by structure
and size river valleys. This confirms the high diversity of
river valleys as important ecological corridors providing
biodiversity conservation. At the same time, the
technique permits to conduct quantitative comparative
analysis both of individual habitats and ecomers of the
valleys entirely, to assess their topological structure that
reflects change in -coenodiversity. Such studies are
important for developing specific habitat conservation
measures and forecasting their possible changes.
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430 ISSN 0372-4123. Ukr. Bot. J., 2015, 72(5)
Дідух Я.П., Чусова О.О., Ольшевська І.А.,
Поліщук Ю.В. Річкові долини як об’єкт еколого-
геоботанічних досліджень. — Укр. ботан. журн. — 2015. —
72(5): 415—430.
Інститут ботаніки імені М.Г. Холодного НАН України
вул. Терещенківська, 2, м. Київ, 01004, Україна
Долини річок характеризуються високою різноманіт-
ністю екосистем, їх оригінальністю, складною
структурованістю і динамічністю. Важливе значення
у формуванні закономірностей територіального роз-
поділу цих екосистем мають вертикальні переміщення
від плакору до русла на різних ділянках катени,
яку можна розглядати як складний комплекс,
де інтегруються геоморфологічні, ґрунтотвірні
процеси та ценозоутворення. Індикаторами цих
територіально-часових змін виступають відповідні
рослинні угруповання, поєднання яких у межах
екологічного ряду під впливом екологічних чинників
трактується як екомери, що відображає градієнт
змін, тобто -різноманіття. Кількісна оцінка цих
рослинних угруповань можлива на основі методики
синфітоіндикації. Для роботи було обрано три модельні
річки (Случ, Дністер і Красна), які мають різну
розмірність і протікають у різних природних зонах. На
їхньому прикладі проведено порівняння екомерів, тобто
оцінено диференціацію —ценорізноманіття щодо
зміни провідних екофакторів.
К л ю ч о в і с л о в а: річкова долина, -різноманіття,
катена, екомери, рослинні угруповання.
Дидух Я.П., Чусова О.А., Ольшевская И.А.,
Полищук Ю.В. Речные долины как объект эколого-
геоботанических исследований. — Укр. ботан. журн. —
2015. — 72(5): 415—430.
Институт ботаники имени Н.Г. Холодного НАН Украины
ул. Терещенковская, 2, г. Киев, 01004, Украина
Долины рек характеризуются высоким разнообразием
экосистем, их оригинальностью, сложной структуриро-
ванностью и динамичностью. Важное значение в
формировании закономерностей территориального
распределения этих экосистем имеют вертикальные
перемещения от плакора до русла на разных участках
катены, которую можно рассматривать как сложный
комплекс, где интегрируются геоморфологические,
почвообразующие и процессы ценозообразования.
Индикаторами этих территориально-временных
изменений выступают соответствующие растительные
сообщества, сочетание которых в пределах
экологического ряда под влиянием экологических
факторов трактуется как экомеры, что отражает
градиент изменений, то есть -ценоразнообразие.
Количественная оценка этих растительных сообществ
возможна на основе методики синфитоиндикации.
Для работы были выбраны три модельные реки (Случ,
Днестр и Красная), которые имеют разную размерность
и протекают в разных природных зонах. На их примере
проведено сравнение экомеров, т. е. дана оценка
дифференциации -ценоразнообразия по отношению к
изменению ведущих экофакторов.
К л ю ч е в ы е с л о в а: речная долина, -разнообразие,
катена, экомеры, растительные сообщества.
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