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116.Журнал Сибирского федерального университета. Сер. Биология №1 2009

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Журнал Сибирского федерального университета
2009
Journal of Siberian Federal University
2 (1)
Биология (Спецвыпуск «Изменения климата и экосистемы»)
Biology (Special Issue «Climate Change and Ecosystems»)
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Editorial Advisory Board
Chairman:
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Vladimir V. Shaidurov
Veniamin S. Sokolov
CONTENTS / СОДЕРЖАНИЕ
В.В. Зуев
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Tatiana A. Blyakharchuk
Western Siberia, a Review of Holocene Climatic Changes
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Editor-in-Chief:
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Editorial board for Biology:
Sergey I. Bartsev
Alexander Y. Bolsunovsky
Tatiana G. Volova
Eugene S. Vysotski
Nikolai A. Gaevsky
Egor S. Zadereev
Valentina A. Kratasyuk
Elena N. Muratova
J. Woodland Hastings
Frank D. Salisbury
Malcolm K. Hughes
Ernst-Detlef Schulze
Akira Osawa
Takayoshi Koike
Marc D?Alarcao
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Journal of Siberian Federal University. Biology 1 (2009 2) 4-12
~~~
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Western Siberia, a Review of Holocene Climatic Changes
Tatiana A. Blyakharchuk*
Institute of Monitoring of Climatic and Ecological Systems SB RAS
10/3 Akademichesky, Tomsk, 634055 Russia 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
This article generalizes published palaeoecological and palaeoclimatic data based on pollen and other
proxy data for Holocene of Western Siberia and describes the most significant palaeoreconstructions
and hypothesis in explanation of past climatic changes in Western Siberia, although it does not pretend
on overall review of all palaeopalynological data available for this region.
Key words: Siberia, pollen analyses, palaeovegetation, palaeoclimate.
Introduction
Pollen analyses of peat and lake sediments
accompanied by radiocarbon dates and,
were possible, by macrofossil data are strong
instrument for study of past changes of vegetation
and landscapes. Taking into account a close
connection of types of vegetation with climatic
conditions palaeopalynological data can be used
for qualitative and quantitative reconstructions
of changes of palaeoclimate, which allow to
observe long time climatic oscillations in the
past long before instrumental era of observations
of climatic parameters. To the end of 20-th
century quite a lot of pollen diagrams supplied by
radiocarbon dates were published for territory of
Western Siberia. Generalizing paleopalynological
data, macrofossil and some other proxy data
(macrofossil data and information about
change of lake levels) researchers performed
reconstructions of palaeoclimates of Western
Sibiria in the Late Glacial and the Holocene
time. Each generalized palaeoclimatic curve is
*
1
based on well proved detailed research of one or
another natural archive of proxy data. Comparing
of these curves allocated on the same time scale
can provide information about local and regional
fluctuations of climate, among which fluctuations
which are similar in different curves and proxies
reflect more global and regional events in climatic
changes and those presented only in individual
curves reflect change of local climates.
Short geographic description of the area
The complicated character of Holocene
climatic changes in Western Siberia is a result
of the continental position of this extensive
low plain (52-72 degrees N, 60-90 degrees E).
The climate of Western Siberia is temperatecontinental and is controlled by the alternation
of Atlantic storms and Asiatic anticyclones,
with the influence of the Arctic air mass from
the north and the subtropical air mass from
Central Asia (Trifonova, 1982). Clear latitudinal
vegetation zones in Western Siberia range from
Corresponding author E-mail address: tarun5@rambler.ru
© Siberian Federal University. All rights reserved
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Tatiana A. Blyakharchuk. Western Siberia, a Review of Holocene Climatic Changes
tundra in the north to boreal forest (taiga) to birch
forests and steppe in the south, with transitional
subzones at ecotones.
Patterns of the Holocene climatic changes
in Western Siberia and hypothesis
connected with them
On the basis of available palynological
data from the whole of Western Siberia, plantmacrofossil investigations in the Yenisey River
region, and investigations of lake-level changes
in the forest-steppe and steppe zones, the
palaeoclimatic reconstructions were performed
for the entire Holocene by different researches
during second half of 20-th centuary. With use of
different methods of qualitative and quantitative
reconstructions of paleoclimate the results
have been presented as palaeoclimatic curves
(Fig. 1.) In available publications each researcher
presented well established palaeoreconstructions
of past climates, but in many cases there were
not clear contradictions in results of different
researcher. The aim of this article is to compare
different palaeoclimatic curves and to analyze
possible reasons of similarities or dissimilarities
of them. One of the first qualitative palaeoclimatic
curve was constructed by A.V. Shnytnikov on the
base of available historical data for Euroasia and
Northern America (Fig. 1. C) (Shnytnikov, 1951).
This curve reflects mostly fluctuations of humidity
of climate. Zonal method of palaeoclimatic
reconstructions based on close connection of
different types of vegetation (and their pollen
specters) with climatic zones and on special
meaning attributed to indicator types of pollen
was used by S. S. Savina and N.A. Khotinskiy
(1984) as well as by V.S. Volkova and T.P. Levina
(1985) for paleoreconstruction of average annual
temperatures and annual amount of atmospheric
plecipitations (Fig. 1, A, B).
Information-statistic method of Klimanov
based on pollen taxa of tree species and
common composition of pollen specters in each
pollen sample from palynological diagrams
(Klimanov, 1976; Blyakharchuk, 1989; Glebov
et al.., 1996; Volkova and Klimanov, 1988)
produced another type of palaeoclimatic curves
an examples of which for south-eastern part
of Western Siberia is presented in Fig. 1. F, G.
(Blyakharchuk, 1989). One can mention more
complicated appearance of these palaeoclimatic
curves, which can be result of influence of more
uncertainties involved in mathematic analyses
of pollen data. Similar approach to pollen data,
but with use of another mathematical method
such as discriminant analyses and equations
of regression was used by G. Yu. Zubareva
(1987) for pollen data located slightly to the
east from Western Siberia in Minusinskaya
steppe (Fig. 1, E). There is evident correlation
between curves of precipitations constructed
by these two methods for two sites located in
forest-steppe zone (Fig. 1, E and F). This is
evidence both of similar mathematical approach
to pollen data and of regional (or global) nature
of the reflected climatic changes. As additional
reassurance of the last conclusion we can present
palaeoecological curves constructed by method
of ecological scales (Blyakharchuk, 1989) on
the base of macrofossil composition of peat
from mire located in floodplain of Ob? River
in subtaiga zone of Western Siberia (Fig. 1, H),
which demonstrates similar wet interval 2000
years ago, dry interval between 1 and 2 thousand
years ago and dry interval between 2 and 3
thousand years ago. If similar climatic events
have been fi xed by few independent methods, it
increases considerably their probability.
Among other methods it is need to mention
mathematical method of analogues (Burashnikova
et al.., 1982) and method of ecology-geographic
analyses of carpological materials (Koshkarova,
1989). Very interesting palaeoclimatic curves
were constructed by L.A. Orlova, (Orlova, 1986;
#5#
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Tatiana A. Blyakharchuk. Western Siberia, a Review of Holocene Climatic Changes
Fig. 1. Correlation on one time scale of palaeoclimatic curves for Holecene of Western Siberia constructed
with use of different qualitative and quantitative methods based on pollen data, macrofossil data and historical
data and: where: A ? palaeoclimatic curves constructed by N.A.Khotinskiy (1977) on the base of pollen data.
B ? palaeoclimatic curves constructed by V.S.Volkova and T.P. Levina (1985) based on the pollen data. C ? curve
of humidity of climate constructed by A.V.Shnytnikov (1951) on the base of historical data for all northern
hemisphere. D ? palaeoclimatic curves constructed by T.P.Levina and L.A.Orlova (1993) for south of Western
Siberia on the base of pollen data (in deviations from modern level). E ? palaeoclimatic curve of annual amount of
precipitation constructed by G. Yu. Zubareva (1987) for Minusinskaya steppe with use of discriminant analyses of
pollen data. F and G ? palaeoclimatic curves of average annual temperatures and annual amount of precipotation
(in deviations from modern level) constructed by T.A.Blyakharchuk (1989) with use of information-statistic
method of Klimanov (1976) on the base of pollen data from south-eastern Siberia, were F ? reconstructions for
subtaiga zone, and G ? reconstructions for middle and southern taiga zone. H ? curve of change of moisture on
the mire in subtaiga zone constructed by T.A. Blyakharchuk (1989) with use of method of ecological scales of
moisture of soils on the base of macrofossil composition of peat deposits (1989) (in steps of ecological scale)
Levina and Orlova, 1993) for forest-steppe zone
of Western Siberia (Fig. 1, D) on the base of
palaeopalynological and radiocarbon data in
which special attention have been paid to such
pollen types as trees pollen, pollen of Artemisia,
Chenopodiaceae and Poaceae in combination
with Betula nana pollen. As a limitation of most
palaeoreconstructions one can mention the use of
rather scattered palynological data for so vast and
diverse an area as Western Siberia and insufficient
attention to complex local conditions in affecting
pollen spectra.
According to most published data the
Holocene climatic changes in Western Siberia
consisted of alternation of cool and warm periods
in combination with changes in humidity and
continentality. These climatic oscillations were
superimposed on the main trend of gradual
warming in the first half of the Holocene and
gradual cooling in the second half. This picture
is in accordance with hypothesis of maximum
of summer insolation, which took place at the
beginning of Holocene at about 10 000 years ago.
At the same time the climate of the Holocene
fluctuated between stages of relative stability and
periods of rapid changes (Khotinskiy, 1984). The
warmest climatic conditions of the Holocene,
so called climatic optimum, took place in the
middle Holocene at about 5500-6000 yr BP in
central part of Western Siberia according to
#6#
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Tatiana A. Blyakharchuk. Western Siberia, a Review of Holocene Climatic Changes
V.S.Volkova and T.P. Levina (1983) (Fig. 1, B).
According to N.A. Khotinskiy (1984) the model
of Holocene climatic changes in Western Siberia
differs from that of Europe by the stronger Boreal
thermal maximum, which was accompanied
by an increase in precipitation (Fig. 1, A).
Summarizing this information V.S.Volkova and
V.A. Klimanov (1988) concluded, that in Western
Siberia Boreal (BO) and Subboreal (SB) thermal
maxima took place at 8500 and 3500 yr BP. At
the same time one can mention, that the Boreal
climatic maximum was more clear expressed
on the north of Western Siberia, but in the
middle part of Western Siberia only Atlantic
(AT) climatic optimum was pronounced, and
in the south in forest-steppe zone the picture
was not so contrast, but more complicated with
numerous displacements of southern boundary
of forest zone to the north and to the south.
This can be result of different parameters used
for determination of conception of «Holocene
optimum». In the north Holocene optimum was
fixed by northward shifting of forest zone caused
by warming of climate. In the middle part of
Western Siberia (in taiga zone) the Holocene
optimum was marked by maximal spreading
of Abies sibirica the most thermophylous and
methophyllous tree species among Siberian tree
species (Shumilova, 1962), that was connected
with warming and increased humidity of climate.
In the south the limiting factor for spreading of
forests is humidity of climate, which fluctuated
in Western Siberia during Holocene many times
(Fig. 1, D). Reflecting of these situation we can
observe in the picture of correlation of different
palaeoclimatic curves for Western Siberia
(Fig. 1). The next important peculiarity of West
Siberian climate was the cooling and increase of
continentality in the second half of the Holocene.
It is so-called Siberian continental type of past
climatic change. These fluctuations were caused
by the rearrangement of atmospheric circulation
above northern Eurasia and by change in the
Arctic ice (Khotinskiy, 1984).
Other researchers (Levina and Orlova, 1993)
in reconstructing the development of vegetation
and climate in the south of Western Siberia,
distinguished 10 uniform cycles with a duration
of 1000 years, each of which includes 4 intervals
of 250-300 years. Each cold period continues for
500-600 years; with the same duration for each
warm period (Fig. 1, D). The cycle started by
warm and dry interval followed by warm and wet
interval, then by cold and wet and then by cold and
dry intervals. The dry periods were more clearly
expressed in the first half of the Holocene and the
wet periods in the second half after 6650 yr BP.
Finer fluctuations of climate were superimposed
on these cycles.
According to point of view of V.A.
Klimanov and A.A. Sirin (1997) the Holocene
climatic oscillations in Western Siberia were
cyclic, but have a quasi periodic character. Under
these conditions in the north the precipitation
increased during warm periods and decreased
during cold periods. In the south the picture was
opposite, and in the middle latitudes the maxima
of precipitation occurred between extremes of
warming and cooling. This hypothesis finds
conformation in correlation of different types
of palaeoclimatic curves (Figire 1). Southern
type of climatic oscillations is clear expressed
in climatic curves constructed by T.P. Levina
and L.A. Orlova (1993) for forest-steppe zone
of Western Siberia. Climatic curves constructed
by method of Klimanow for forest zone of
south-eastern part of Western Siberia are more
close to northern type of climatic fluctuations
(Fig. 1. F, G). But at some time intervals, such
as in the middle Atlantic period (AT-7) and in
Boreal period (BO-9) it changed to southern type
of fluctuations, that is clear evidence of shifting
of southern boundary of forest zone in Western
Siberia during these time intervals.
#7#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Tatiana A. Blyakharchuk. Western Siberia, a Review of Holocene Climatic Changes
During the Holocene in Western Siberia
the main changes in climate and landscape as
described by different methods were probably
caused by global factors. Bellow is generalized
picture of climatic and landscape changes in
Western Siberia during the Holocene.
Compilation of palaeoclimatic
reconstructions according
to periods of Holocene
Preboreal period (PB). If to accept
chronostratigraphic division of the Holocene
according to J. Mangerud et al., (1974), then the
first Holocene warming in Western Siberia took
place in the first half of the Preboreal (PB). From
this time severe continental climatic conditions
with widespread permafrost changed to a milder
climate, tundra-steppe vegetation of periglacial
areas was replaced by birch forest-tundra
with shrub under storey, and the thawing and
lowering of permafrost started to spread quickly
(Blyakharchuk and Sulerzhitsky, 1999). The
further warming of climate promoted the spread
of spruce along the river valleys and lowlands.
Climatic conditions became much warmer than
in late-glacial time but did not reach the present
level.
In the second half of the Preboreal (PB)
the climatic conditions became drier as a result
of further warming. That was favorable for
the spread of birch forest-steppe in the area
of modern forest zone (Volkova and Levina,
1985; Koshkarov, 1998; Blyakharchuk and
Sulerzhitsky, 1999). At the same time in foreststeppe zone humidity increased during second
half of preboreal. Some authors mention the
cooling of climate during this time (Khotinskiy,
1977), but the spread of such a thermophylous
tree like Abies sibirica Ledeb. (Blyakharchuk
and Sulerzhitsky, 1999) and thermophylous water
plants (P?yavchenko, 1983; Koshkarov, 1998) does
not agree with this conclusion. About 9500 yr BP
the forest plant communities existed in a zone of
modern tundra on Cape Karginsky (Firsov et al.,
1974). Just as at the beginning of the Holocene
in the Yenisei River area, part of the forest zone
of Western Siberia show changes from foreststeppe to birch forest and then to dark coniferous
forest of the modern taiga (Koshkarova,1989;
Koshkarov,1998). By the opinion of these authors
the decreasing continentality of the climate and
the increasing moisture caused these change in
forest-steppe zone.
Boreal period (BO). The next remarkable
warming of climate took place in the Boreal
period 8300-8500 yr BP. Precipitation increased
(Khotinskiy,1984), promoting the widespread
expansion of spruce throughout the vast
territory of Western Siberia. Some scientists
consider that the climate of Western Siberia in
Boreal time was cooler than today and that the
steppe zone was shifted to the south and situated
out of Western Siberia (Volkova and Levina,
1985). Other investigators show that average
temperatures in January and annual precipitation
were higher than modern north of 60 degrees N,
and the opposite to the south (Velichko et al.,
1997; Savina and Khotinskiy, 1984). At the same
time the average July temperatures were lower
than modern north of 60 degrees N. According
to V.L. Koshkarova (1989) and A.D. Koshkarov
(1998) during the Boreal warming the climate
was warmer and drier than modern in the
southern taiga zone in the Yenisey River area.
The amount of precipitation was 40-50 mm less,
and latitudinal zones were shifted to the north by
4-5 degrees from the modern position. Forests
expanded in the tundra zone (Levkovskaya et
al., 1970). In the modern taiga zone the larchspruce forests spread together with birch foreststeppe (Blyakharchuk and Sulerzhitsky, 1999).
Accordingly palaeoclimatic curve of southern
type (Fig. 1, D) reflects mostly drier climate
during BO with short increase of humidity
#8#
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Tatiana A. Blyakharchuk. Western Siberia, a Review of Holocene Climatic Changes
in the middle of BO and more wet climate in
taiga zone (Fig. 1, A, and G). The process of
podsolization was interrupted by formation of
steppe soils in the zone of the modern southern
taiga (Koshkarov, 1998). The type of latitudinal
zonation in the Boreal was rather different from
modern.
Atlantic period (AT). The climatic optimum
of the Holocene is best expressed in Western
Siberia at 5500-6000 yr BP. Most investigators
consider that during this time the climate was
warmer and wetter than modern in the forest
zone. It caused the new expansion of forests
into the tundra zone on Cape Karginsky (Firsov
et al.., 1974; Levkovskaya et al.., 1970) and the
spread of lime, oak, and elm in the western part of
the forest zone of Western Siberia (Volkova and
Levina, 1982). At the opinion of N.A. Khotinskiy
(1984) the southern boundary of the forest zone
in Western Siberia was stable in the second half
of the Holocene. But later there were found quite
a lot of evidences, that the southern boundary
of the forest zone was considerably shifted to
the north in the Atlantic climatic optimum, as
indicated by the formation of a second humus
horizon in the relic soils of the modern southern
taiga (Orlova and Panychev, 1989). In the steppe
zone the great decrease in the size of Lake Chany
as a result of drying was dated as 5530 yr BP
(Orlova, 1986), and steppe communities spread
into the forest-steppe zone. Peat accumulation
started on some dried lakes in the forest-steppe
zone at this time. Moreover it was proved, that
there were numerous less profound movements
of southern boundary of forest zone in Western
Siberia caused by climatic fluctuations (Levina
and Orlova, 1993). The deviations of climatic
parameters from modern values in the Holocene
optimum changed in accordance with latitude.
Thus in the forest zone the annual temperatures
exceeded modern by 1-1.5 degrees, and the
precipitation were 25-50 mm greater. These
conditions promoted widespread Abies forests in
Western Siberia, which expanded to the north of
the modern position by 200-300 km (Glebov et
al., 1996; Blyakharchuk and Sulerzhitsky, 1999;
Koshkarov, 1998). According to other scientists
(Volkova and Levina, 1985) the climate of the
Holocene optimum was 3.5 degrees warmer
than modern, and latitudinal zones were shifted
500-600 km north of their modern positions.
The average January temperature in the modern
forest-tundra zone was increased 5-6 degrees
(Burashnicova et al., 1982; Khotinskiy, 1984).
Another opinion is that the expansion of forests
to the north was caused not by warming of the
climate, but by the increase of continentality
(Danilov, 1989). The author considers, that in
more continental climate due to more hot summers
the permafrost in soil thawed deeper and it was
favourable for grows of trees. According to
A.A.Velichko (1989), in the Holocene optimum
the increase of average global temperature
at high latitudes was maximal in winter (+4
degrees), whereas in the south the positive
deviations decreased, and at 40-50 degrees N
mostly cooling took place. Precipitation increase
was the same in high and low latitudes. In the
middle areas (steppe zone of Western Siberia)
precipitation was less than modern. At the same
time palaeoclimatic curves constructed by
T.P.Levina and L.A Orlova (1993) do not show
pronounced warming in forest-steppe zone, but
rather more stable steppe conditions compare
with previous and next Holocene periods (Fig. 1,
D). On the contrary, palaeoclimatic curves from
forest zone demonstrate more warm climate
with cooling in the middle AT and other smaller
fluctuations of temperature and humidity (Fig. 1,
A, B and G).
Subboreal period (SA). The climate of
the Subboreal period was unstable, but mostly
cooler than that in the Atlantic in the forest
and forest-tundra and forest-steppe zones. The
#9#
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Tatiana A. Blyakharchuk. Western Siberia, a Review of Holocene Climatic Changes
northern boundary of the forest zone retreated
to the south by almost 2 degrees. Most scientists
consider 4500 yr BP as a time of dry cooling in
the late Holocene (Blyakharchuk, 1989; Volkova
and Levina, 1985; Koshkarov, 1989; Khotinskiy,
1984). But in the opinion of F.Z. Glebov et
al., (1996) the dry warming of the Subboreal
allowed cotton-grass peat to form on the mires
of Western Siberia. In any case it was a dry
period, as indicated by the renewed regression
of lakes in the steppe and forest-steppe (Orlova,
1986). After the dry period the climate became
wet and cool, causing the afforestation of foreststeppe and partly of the steppe zone. The cooling
at 4300 BP stimulated the freezing of some wet
mires in the middle taiga zone and formation of
palsa bogs in the northern taiga and forest-tundra
(Blyakharchuk and Sulerzhitsky, 1999; Peteet et
al., 1998). Relatively thermophyllous Abies forests
retreated to the south to the position of the modern
southern taiga. The widespread expansion of
siberian pine ( Pinus sibirica) started. In central
areas of Western Siberia in the second half of the
Holocene the continentality increased. This fact
supports the model of Siberian continental type of
climatic changes (Khotinskiy,1984). Fluctuations
of climate to more cool and dry conditions are
expressed in palaeoclimatic curves for forest zone
(Fig. 1, A, B, F and G, H) and to more unstable
conditions in curve for forest-steppe (Fig. 1, D).
Subatlantic period (SB). During subatlantic
time the cold phases took place at 2800, 1700
and 500 yr BP (Levina and Orlova, 1993). The
following dry events at 2500-3000, 1500, and
600-800 yr BP were marked by afforestation
of mires in the southern taiga zone and by
regressions of lakes in the forest-steppe and steppe
(Blyakharchuk, 1989; Orlova, 1986; Levina and
Orlova, 1993). At the opinion of N. A. Khotinskiy
(1989) the century oscillations of climate during
last millennium had the tendency that warming in
humid areas coincided with stages of aridification
in the steppe zone, and cooling in humid areas
coincided with wetter conditions in arid zones.
Conclusion
Correlation of palaeoclimatic curves of
temperature and humidity (which have been
constructed with use of different methods on
the base of pollen, macrofossil and radiocarbon
data) demonstrated complicated pattern of
climatic changes in Western Siberia during the
Holocene. Complexity is expressed both in space
and in time scales. But never the less there are
common features in different palaeoclimatic
curves, which are caused by global, possibly,
solar-orbital driving forcing of climatic changes
in Western Siberia. On this pattern a smaller
oscillations are superimposed, possibly induced
by changes of local climates. The last were caused
by the rearrangement of atmospheric circulation
above Western Siberia and by peculiarities of
local environment in each study site. For more
correct modeling of past and future climatic
changes in such vast and diverse area like
Western Siberia, which could include not only
time palaeoreconstructions, but also space-time
models, there is obvious need in an accumulation
of more sites supplied by detailed palaeoecological
data of high scientific level accepted on modern
palaeogeographic research. And in each separate
case a special attention should be paid to complex
local conditions in affecting pollen and other
palaeo proxy data so to avoid mix of local and
regional signals. Correlation of different proxies
on one time scale is a good instrument for this
aim.
Acknowledgments
The author thanks Professor Dr Herbert
E.Wright (University of Minnesota, USA) for
advices and linguistic improvements of this entry.
The work was performed under support of grants
of RFBR ? 05-05-64266 and ? 06-05-65127.
# 10 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Tatiana A. Blyakharchuk. Western Siberia, a Review of Holocene Climatic Changes
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 13-17
~~~
??? 595.768.13: 632.76
???????????? ?????? ? ????????? ????????
Acanthoscelides obtectus Sav. (Coleoptera, Bruchidae)
?? ???????? ??????? ?????? ????????
?.?. ???????, ?.?. ?????????, ?.?. ?????????*
??????? ??????????????? ???????????,
634050 ??????, ?????, ??. ?????? 36 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
??????????? ???????????? ????????, ??? ?????? ???????????? ????? ???????
???????????????? ?? ??? ???????? ??????. ????? ?? ??????? ????????, ?????????
??????????? ? ???????? ????? ??????, ???????? ?????????, ????? ??????? ???????? ??????
????????? ???????? (Acanthoscelides obtectus Sav.). ???????? ??????????? ? ??????? ?? ????
??????? ?????? ????????, ? ? ???????? ???????? ?? 90 % ????? ?????????? ??????????.
???????? ???????? ???????? ? ??????????????: ? ????? ????????? ??????????????
??????? ????, ??? ? ????? ????????.
???????? ?????: ??????, ????????? ????????, ?????????????? ?????.
????????
???????? ????????? ? XXI ?. ????? ???????????? ?? ????? ?????????, ? ??? ????? ?? ??????????? ???????????? ???????????????????? ??????? ? ???????????????
? ????????????? ???????? ??????????
?????. ??????????? ??????????????? ??
????? ? ?????? ??????? ? ??????????????
??? ??????????? ?? ??? ?????????????? ?
???????????? ?????????? ???-?????????
??????? ? ????? ????????????? ??????,
??????? ?? ??????. ??? ???? ????????
??????????? ? ???, ??? ? ????????? ?????
90 % ???????? ???????????? ?????????
??????? ?????????? ???? ?? 20 ????? ????????. ? ?????????? ???????????? ?????????????? ???????? ?????? ???? ??????
*
1
??????? ??????????? ????? ??? ???????
??????? (???????, 2001).
????????? ??????? ? ???????? ????????
???????????? ??????? ? ?? ??????? ??????????????? ???????????? ?????? ?? ??????
???????? ? ???????????? ??????????. ? ?????
? ??????????? ?????????????? ??????? ???????? ?????? ???????????? ??????? (??????,
???), ? ????????? ?? ??????????? ?? ?????,
?????????? ????????? ?????????? ???????????? ?? ???????????? ? ??????????? ???????? ????? (??????? ? ??., 1989).
?????? ???????????? ? Phaseolus vulgaris
(L.) Savi. ? ?????????????? ??????? ????????,
????????????? ??????? ?? ?????? ???????????? ??????? ???????? ???????????? ????????????????? ??????? ????????? ?????????
Corresponding author E-mail address: quality_tsu@mail.ru
© Siberian Federal University. All rights reserved
# 13 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ? ?. ?????????? ???????????? ?????? ? ????????? ???????? Acanthoscelides obtectus Sav?
????????. ?????? ??????? ?? ??????? ???????
? ???????? ????????. ??????????? ? ???????????? ?????? ?? ???????? ????????????????
??? ?????? ???????. ?????? ????????????? ?
???????? ????? ? ?????, ?????? ???????????
?????????, ??????????? ? ????????? ???????????? ???? (?????) ???????? ???????????????? ???????????? (???), ??????????????? ?
??????????? ?? ???????????? ?? ??? ??????
???? ???????? ??????. ? ????????? ???? ???
???????? ????????? ? ?????????? ??????????,
? ????? ? ??? ?????? ?????? ?????? ????????????? ??????, ???????????? ??????????????,
??????? ???????????? ? ???????? ????????????? ? ?????????? ? ????????.
???????? ??????????????? ?????????
????? ?????? ?????? ????????? ? ???????? ??????????? ???????? ? ?????????????
????????? ???????????????????? ???????
?????, ? ????? ? ??????? ??????, ???????????? ????????? ???????? ? ??????????
?????????? («??????? ??????»). ????????????? ???? ?????? ???????????? ? ?????
??????????? ????? 30 ??????. ???????? ?????????? ????????? ????? ??????, ??????????? ?? ??????? ???????????? ??????? ??????, ?? ????????? ???????????? ????? ??????
? ?????? ?????. ????? ???????? ?????? ?????? ???????? ???????????? ?????, ?????????
?????????????? ??????, ????? ??? ????????,
?????????, ??????? ? ??. ??? ??????????????? ?????????? ???????????? ? ??????????????. ??? ??????????? ??????? ??????
?????? ???????????? ??????? ?????? ??????
??????????? ? ???????? ????????.
???????? ???????, ????????? ??????????? ? ???????? ????? ? ????? ??????, ? ????????? ? ??????? ???????.
???????? ??????? ????????? ?????? ? ????????? ???????? (Acanthoscelides obtectus Sav.),
????????????? ???????????? ?????????, ??????????? ? ????????? ???????? (Bruchidae).
??????? ????????? ???????? ???????? ??????????? ???????; ? ????? XIX ?. ????????? ????? ? ????????????? ?????????? ?? ???????.
???????? ?????? ???????? ????? ???????????????? ? 1914 ?., ??? ???? ??????? ? ????????
????? ?? ??? ????? ??????????? ?????????.
? ????????? ????? ????????? ??????? ??????? ????? ??????? ?????? ? ??????????? ?
????? ???????? ?????? (??????????? ????,
????????????? ????). ????????? ???????? ????????????? ? ???????? ??????????? ? ????????? (?????????, 1998), ?? ????? ?????? ?????
????????????? ?????? ????????? ??????. ?
????? ? ??????? ??????????????? ?????? ??
?????? ??????????? ????? ?????? ? ? ??????
??????????? ??????????? ? ????? ?????????
???????? (????????, ???????, 2004, 2005),
???? ?? ???????? 80-? ??. ?? ?. ???? ?????????
??????????? ?? ????????? ?? ??????????? ???????? 52є ?.?. (??????? ? ??., 1989).
????? ???????????, ??? ??????? ???????????? ?????? ????????? ????? ???????? ??
????? ???????? (????????, 1999), ?? ???????? ???????? ???????? ? ??????? ?????????????? ????? (Ciepelewska et al, 1997), ??
??????? ???????????? ??????? ?????? ????????? (????????, ???????, 2004). ? ?????,
?????????? ? ????????????? ????????????
????????? ???????? ? ????? ?????? ? ??? ?
?????????????? ??????? ?????? ????.
????? ????? ?????? ???? ???????? ????? ???????? ????????? ???????? ? ???????
??????????? ????? ?????? ??????, ?????????????? ?? ??? ??????? ???????. ? ????????? ????????? ?????????? ?????????????
?????? ???????????? ????? ???? ??????????
?????? ?????? ? ????????? ????????.
??????? ? ?????? ????????????
?????????????? ?????? ????????? ????????? ?? ???? ????????????? ?? ???????? ????????????? ?????? ? ???????? ? ?????????.
# 14 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ? ?. ?????????? ???????????? ?????? ? ????????? ???????? Acanthoscelides obtectus Sav?
???? ???????? ????????? ?????????????
??????? ? ????????? ??? ??????????????? ?
???????? (?. ?????????? ????????????? ???.)
??????? ?????? ?? ???????? ??????? ?????????? ?????? ????????????? ????. ????????
???????? ?????, ??????? 35?50 ??. ??????
????????-??????????, ?? 2?5 ?? ?????????.
???? ??????????????, ??????????????, ? ??????????? ????????? ? ??????? ????????,
? ???????????? ?????. ???? ?????-????????
????? ? ?????????? ??????????????. ??????
??????? ???????????? ???? ? ???????? ? ??????? ? ???? ?????. ?????? ???????, ??????????? ?????, ??????-??????? ? ??????????
????????, ?????? ????? ????????? ??????????. ???? ???????? ?????????? ???????????
? ????????????? ????????????, ?????????????? ? ??????.
???? ??????????? ? 1989 ?. ? ???????????????? ? ???????-????????? ????????.
?? ?????? ?????, ? ???????? ??? ???????
??????? ???? ?????????? ?????????? ???????????? ? ???????? ????????? ??????????
?????. ????? 1000 ????? 390?600 ?. ?????????? ????? 22?25 %. ????????????? ?????? ? 88?94 ???. ??????????? ????? ????????? 34,1 ?/??.
???? ????????? ???????? ????????????? ??????? ?? ??????????? ???????
??????? ???????. ???? ???????????. ???????? ???????? ?????, ??????? 40?52 ??.
?????? ???????? ???????, ?????. ???? ??????????????, ? ?????????? ??????? ?? ????????????? ??? ?????????? ?????, ???
????????????? ???? ? ???????, ? ??????????? ???????? ???????, ?????? 11,7?12,8
??. ???? ???????, ???????? ???????????,
?? ??????? ?????????????? ????????. ?????? ?????????????, ?????, ?????? ?????.
????? 1000 ????? 290?320 ?. ???????????
????? ????????? 90 ?/??. ???? ???????? ?
??????????.
? 2002-2007 ??. ???? ?????????? ????? ????????? ?????? ?????? ? ????????????? ????????? ???????????? ???????.
??? ?????????? ???????????? ?????????
?????????? ???????? ????????? ????????
?????? ????? ??????????? ? ????????????
?????? ????????? ? ???????? ????????????? ? ??????? ??????? ?? ??????????????
?????????. ???? ????? ????????????? ?
?????? ?????? ???????? ? ??????????? ??
?????? ??????????. ???????? ????????, ??????????????? ?.?. ????????? (1975), ?
?????? ?????? ?????? ??? ??????? ?????
???????? ?? 100 ?????. ?????? ?????????
??????? ?? ?????, ?????????? ???????????? ????????.
?????? ????????????? ? ???????? ?
???????? ?????? ? ?????????? ?? ????????
? ??????????. ?????? ????????? ? ?????????
???????? ??? ??????????? 25 ?? ? ???????????? (??? ? 10 ????) ???????????? ???????? ??
??????? ?????????.
??????????? ?????????? ?? ?????????
????????? ???????? ? ???? ???????? ????? ???????? ?????????, ???????? ????????
? ??????? ?????? ?? ????????????, ???????
??????????? ?????. ??????????? ???????
?????, ???????????? ?????????, ? ??????????? ????????? ?????? ????????? (???????,
??????? ? ?????) ? ??????.
?????????? ? ??????????
??????????? ????????????, ??????????? ?? ???? ????? ???, ???????? ??????????? ????????? ??????????????? ??????
???????????? ? ????? ??????? ??????? ???????.
?????? ?????????????????? ?????????
???????? ??????????, ??? ?? ??? ???????
??????? ???? ????????? ???????????? ?????????? ? ??????? ????????? 20 ???. ???????????????? ????????? ????? ????????
# 15 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ? ?. ?????????? ???????????? ?????? ? ????????? ???????? Acanthoscelides obtectus Sav?
?????. ???????? ???????? ????????? ??????? ? ????? ????????? ? ???????? (????????
?????????? ? ????????????????? ??????).
???????????? ??????? ????????? ????????
??? ??????????? ??????????? ?? ?????? ??
?????, ?? ? ? ?????? ???????? ?????. ??????????? ? ????????? ? ????????? ???????, ??????? ?????????? ????? ??????????? ? ???????? ????????? ????????. ???? ?
????? ???????? ?????? ????????? ???? ?
??????? ?????????????? ??????? ??? ????????? (? ? ??????? ????, ? ?????? ???????? ?
??????????, ?? ????), ?? ? ?????? ? ??????
???? (? ?????? ???????? ? ?????????? ? ??
????).
?? ??????? ?????? ?????? ????? ????????? ???????? ???? ???????? ? ?????? ????????? ???????? ????????. ? ??????????????
??????? ???????? ??????????????? ??????
????????? (??????????????? ???????), ?????? ????? ???????????? ????? ?? ?????????
20-22 %.
? ???????? ???????? ????? ??? ??????????? 25 ?? ??????????? 2?3 ?????????
????????? ????????, ? ????? 5?7 ???????
???????? ??????????? ???????? ???????
????????? ????????? ?????????. ? ?????,
??????? ????? ????? (?? 90 %) ?? ?????? ???????? ???????????? ?????????. ? ?????????
??????? ??????????? ?? 20?25 ???????????. ????????? ????? ?????????? ? ???????
?? ???? ??????? ???????? ? ? ???? ???????,
??????? ??? ?????.
???????????? ?????????? ???????, ????????????? ? ????? ??????, ????? ????????
?? 28, ?????? ? ??????? ????? ??????? ?
????? ?????? ?? ????????? 10-15.
???? ???????? ???????? ???????? ???????? ? ??????? ??????????? ????? ??????
?? ??????? ??????? ???????? (????. 1). ??????
????????? ????? ???????? ???????????? ??????? ??????, ??? ????? ?????????. ?? ????
?????????, ??? ??????? ? ????????????????
????????????? ???????? ????? (??? ? ?????
????????? ??????, ??? ? ????? ????????).
????? ????, ?????? ????? ????????? ????????? ????? ? ?????????? ?????? ??????????
????????? ????????? ? ????.
???????? ????????????? ????????????
????????? ????????, ?????????????? ??
?????????? ?????????????????, ????????
?????? ???????????? ????? ? ?????? ????????????. ???????? ??????? ? ????????????
?????????? ?????? ??????????????? ?????
????? ? ?? ?????????????? ? ??????? 2?3
????? ??? ??????????? ????? 15 ?? ??? ???????? ? ??????? ?? ???? ??? ?????? ????????????? ??????????? (2?5??). ??? ????????
? ???????? ????????? ??????????? ????????????? ???????????? ????????? ??????? ??
???? 8-10 %. ? ????? ?????? ???? ??? ???????????? ????? ??? ????? 15 ?? ? ???????
????? ??????????? ??????????? ?????? ?????? ????????? ????????, ?????? ????????
???????? ? ?????? ?????????? ????????? ??
????????.
??????? 1. ??????? ??????????? ????? ????????? ?????? ?????? ???????????? ????????? ?????????
??????????
??????????? ????????? ? 1
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? ??? ?????:
? ???????
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Lim
X + mx
lim
X + mx
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3-27
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0-18
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3,0 ±0,6
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10,7 ±0,6
# 16 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ? ?. ?????????? ???????????? ?????? ? ????????? ???????? Acanthoscelides obtectus Sav?
?????? ??????????
???????? ?.?. (1999) ????????????? ??????????? ?????? ?????? ?? ?????????? ? ???????? ?????????????? ????: ???????. ???. ? ????. ????. ????. ?????????, 22 ?.
??????? ?.?., ????? ?.?., ???????? ?.?. (1989) ???????? ? ????????????? ??????? ??????? ???????. ???????: ???????, 280 ?.
??????? A.A. (2001) ?????????? ??????? ???????? ???????? (???????-???????????? ??????). M.: ???????, ? 208 ?.
???????? ?. ?., ??????? ?. ?. (2004) ????????? ???????? ? ???? ?????? ? ??? // ????????
?????????. ? 8: 26-29.
???????? ?. ?., ??????? ?. ?. (2005) ????????? ???????? // ?????? ? ???????? ????????.
? 12: 34-35.
??????? ???????? ?????????? ???????????????????? ????????. (1975) /??? ???. ?-?? ?.-?.
???? ?.?. ????????. ?.: ?????, 239 ?.
????????? ?.?. (1998) ??????????? ???????? ????????? ???????? // ???????? ????????????
????????????????? ????????. ?. 6. ? 2: 139-142.
Ciepelewska D., Kostyra H., Nietupski M., Fornai L., Gudaczewski W. (1997) Wplyw modyfikowania
okrywy nasiennej na porazenie nasion fasoli przez strakowca fasolowego (Acanthoscelides obtectus
Sav.) // Post. ochr. rosl. V. 37. ? 1: 202-209.
The Resistance of Haricot (Phaseolus Vulgaris) for Bean Weevil ?
Acanthoscelides Obtectus Sav (Coleoptera, Bruchidae)
on the North Border of Crop?s Distribution
Andrey S. Babenko, Svetlana I. Mikhailova
and Irina V. Nikolaeva
Tomsk State University,
36 Lenina, Tomsk, 634050 Russia
Long-term experiments carried out have shown the ability of haricot being cultivated in the south of the
forest zone of Western Siberia. The main factors decreasing ripening and quality of beans and haricot
seeds are pests. The most dangerous pest for haricot is Bean Weevil (Acanthoscelides obtectus). The
pest may be in the beans at different stages of its development; during storage about 90 % of seeds are
damaged by this pest. The are some differences in seeds between haricots of two sorts: vegetable sort
«Zhuravushka» are damaged more significant in compare with seed sort «Bijchanka».
Key words: haricot, Bean Weevil, seed damage.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 18-29
~~~
??? 551.89
???????????? ???????? ??????? ??????
????????? ?????? ??????
?.?. ?????????*
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???,
634055 ??????, ?????, ??. ????????????? 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
? ????????????? ? ????????????? ????? ????? (????????, ????????????, ??????????) ??????????
???? ????? ????? ?????? ?????? ????????? ?????? ???? ?????????????????? ?????????, ???????
???????? ?? ?? ???????? ????????????? ?????????, ??????? ???????? ????? ??????????
? ????????????? ????, ??????????? ?????? ???????????????? ?????? ? ??????????? ???
? ??????? ????????????? ???????. ???????? ???? ?????? ?????? ? ???????, ??????????? ?
??????? ???????? ??????????, ????????? ??????? ???????? ??? ????? ????????? ???????,
??????? ???? ??????? ? ???????????? ?????????? ? ????????????? ????.
???????? ?????: ??????????? ?????, ?????, ?????????????, ??????????.
????????
???????? ?????????????? ???????? ????? ?????????? ??? ???????????? ??????????
(???????) ? ?????????? (??????????) ????????. ???????? ???????? ??? ???? ??????????? ??????? ?????????? ?????????, ???????
? ???????? ?????????? ??????? ??? ????????????? ?????????, ??????????? ? ?????????????? ????????, ? ? ????? ?????????? ???
???????. ?? ?????????? ???????? ????????????? ??????? ??????? ??????????? ??????,
????????? ????? ??? ? ????, ?????????????,
??? ???? ????????, ? ??? ? ???????? ? ?????? ??????. ????? ???????, ????? ??????
?????????, ??? ??? ? ????????? ??????????
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????? ???????????? ????? ? ?????, ? ?????
???????, ? ?????????? ?????????? ? ? ???*
1
???, ?.?. ????????? ? ?.?. ?????? ? ???????? ???????? ???? ?????????????? ????????????? ????? ?????????????? ???????????
(?????????, 1946). ????? ??????????, ??? ?????????? ?????????????? ??? ????????????
???????????????? ?????????????? ???????
????????? ??????? ? ?????????????? ???????
??????? (?????????? ????????????? ???????
? ?????????? ?????, ???????????? ??? ????????? ??????? ????? ???????).
? ?????? ??????? ????? (??????????????
? ???????????) ????????????? ??????? ????? ????????????? ????????? ??????? ??????
?????. ????? ?????????? ?????????? ?????????? ? ??????????? ???????? ?????? ????,
?????????? ?????? ? ??????? ?????. ????? ?
?????? ?? ???????? ??????? ????? ????. ? ???
???????? ??????????? ???? ??????????? ?
Corresponding author E-mail address: bor@ggf.tsu.ru
© Siberian Federal University. All rights reserved
# 18 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ???????????? ???????? ??????? ?????? ????????? ?????? ??????
????????? ???????, ??????????? ? ??????,
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??????? ??? ??????????? ??????. ??? ???????? ?????????? ???????? ??????? ????.
? ???? ?????? ???? ?????????, ?????, ?????????? ? ??????. ? ???????? ????? ????? ??????
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?.?. ?????? ?????? ????? ? ???, ??? ?????
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????? ????????????? ??????? ????? 20 ???
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????? ???????? ? ?????????, ?? ???? ????
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??? ? ????????? ????????? (32190 +/-260 BP
(Beta 137035 UM993)). ???????? ????????????
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?? ?????????? ?? ????????? ???? ?????????
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? ?????? ?????? ??? ???? ???. ????. ???? ???
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3 ?? 5,5 ???. ?? ? ???, ?.?. ???? ? 2,5-4 ????
?????? ???????????? ???????? ????? ??? ??
?????? ????? ???. ? ??????????, ??? ???????? ?????? ????? ? ????????? ????? ?? 2000
?? 1720 ? ??????? ??????? ????????? ??????????? ? 4-5 ???. ???? ???????????? ???????
??????????? ?????? ???????? ????????????
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# 23 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ???????????? ???????? ??????? ?????? ????????? ?????? ??????
???. 2. ???????? ????? ???????? ??????-????????? ???????????? ? ??????? ??????????????
? ? ??????? ????????? ? 1650 ? ? ? ?????????.
? ??????? ????????????? ??????-??????????
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???????? ? ???????? ???????, ??????????
???? (???????, 1947; ???????, 1972; ???????,
1959; Benson, 1978; Bobek, 1963; Butzer, 1961;
Gilbert, 1890; Morrison, 1965,1970; Thomson
et al., 1986). ??????? ??????? ????????????
????????????? ?????????, ????? ??????????
?????? ??? ??? ????? ??? ??? ??????????
???????? ?????? ????????? ?????????????????? ???????????. ????????? ???????
????, ??????? ? ????????? ???????? XIX
????????, ??????????, ? ?????, ?????????
??????????? ??????? ????????????????, ?
?????????, ?????????? ??????????????? ??????????? ????, ??????????? ?? ?????????
???????? ??????????????? ????????: ?.?. ?????????, ?.?. ?????-?????????, ?.?. ????????, ?.?. ????????, ??????? ???? ??????
?????????????? ?????????????? ?????????
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?? ?? ?????????. ????? ???????? ???????
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? ?.?. ???????, ???????? ?.?. ???????, ??????? ?.?. ???????. ?????????????? ??????????? ???? ???????? ???????? ??????????
? ?????? ?.?. ????????? (1948), ?.?. ?????????? (1981), ?.?. ????????? (1971).
????????? ??????? ???? ???????????? ????? ?????????? ??????? ?????????
?????????. ????????????? ? ????????? ?????????? ??? ???????, ???????????????
??????????? ???????, ??????? ??????? ?
??? ?? ?????. ???????? ?????????? ?????
?????? ? ????????? ????? ?????? ?????????? ???? ?????????? ???????????????? ?????? ? ???????????. ????????? ??????????? ???????? ??? (?????, ???????, ?????,
# 24 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ???????????? ???????? ??????? ?????? ????????? ?????? ??????
???, ??????, ??????? ? ??.) ? ???????? ?
???????????? ??????? ???????????? ????? ? ??????, ?????????????? ??????????
? ????????. ?? ?????????? ???????? ???????? ?????? ?????? (????????, 1981; ?????????, 1972; ???????, 1959) ????????????? ??
???? ?? ??????? (??????? ???????????) ??????????. ?? ?????? ????? ??????????????,
?????????? ?????????? ???? ?????????????,
?? ?????? ?????? ? ?? ???????? ?? ????????? ?????? ??? ???????? ? ???????? ??????? (????????, 1981). ?????????? ????????
??????????????, ?? ?????? ?.?. ??????????
(1972), ?????? ???????? ???????? ? ?????????? ??????? ????? 12000 ??. ?? ? ????????? ???????? ? ?????? ??????? ?????-?????,
??????-???????, ????-???????, ?????,
?????-??????, ??????????, ?????, ???????
? ?????? ? ????? ?????????? ??????????
????????? ? ?????? ?? ??????.
???????? ?????????? ? ?????? ?????????? ????????? ??????? ???? ??????????
????? ???????????? ???????. ??? ?? ?????????? ??? ??????????? ?? ?? ?????? ????????
????????? ??????? ???????? ??????????, ??
???????? ??????? ????????? ????? ??????????? ????????? ??? ????????? ??????? «?????????? ? ?????? ???? ? ??????? ???????»
?????? ?????? ?? ?????????. ????????, ???
???????? ????????? ??????? ?? ????? ????????????? ? ???????? ?????? ??????????.
?????????? ??????????? ??????? ?????? ??
????? ?????????? ?????????? ???? ????????? ?, ??? ?????????, ???????? ??????????
????????????? ??????????. ?????????, ?
???? ???????, ??????? ????????? ? ???????
??????? ?????????? ????????.
? ??????-??????? ??????????????????
?????? ????????? ??????? ???? ??????????? ??? ?????? ???????? ?????? ???????
??????????-?????????????? ????????????,
??????? ????????????? ??????? ??????
???????? ? ???????? ?????????????? (??????????????, 1982; ????????, 1981).
? ??????? ?????????????? ???????????
????? ????-??-???, ????-???, ??????-???,
?????-??? ? ??????-??? ???????????? ? ?????? ?????? (???. 3) ? ???????????? ????????
?????? 1260-1265 ?. ????? ????????????? ?????? ????????? ???????, ??????? ?????????
???????? ????????? 23158 ??. ??, ????????????
??????????????? ????????? ???????????
????????? ?? 30 ?? 60-70 ?, ????????????
?? ??????? ???????-?????????? ? ????????????????? ????????????. ? ???????????
??????? ????????? ???????? ??????????? ??
????????? ??????????????? ?????? ???????
????????????. ? ?????, ??? ???????? ??????,
??????????????? ? ?????? ??????? ???????????, ???????? ????????? ? ????? ???????????
??????. ??????? ? ???????? ??????? ???????
????????? ??????????? ?????????? ????????? ?????????? ???? ??????????????? ? ??????????? ?????????? ???????????? ? ?????????.
???????????? ???????????? ????????
?????????????? ??????? ? ??????? ??????
??????? ?? ??????? 1170-1180 ?, ??? ????
??????? ????????? ?????????? 14362 ??.??.
??????????????????????? ??????? ??????????? ??????????? ? ???? ???????? ?? ?.
??????? ? ?? ??????? ??. ??????-???. ???
???????????? ????????-?????????, ?????
??????????, ???????, ?????????? ? ????????? ?????????? ? ???????????. ? ?????????
? ?????? ???? ??????? ???????? ??????????
????????. ???????? ??????????????????????? ??????? ???? ????? 15-30 ? (???. 4).
???????? ???????? ??????? ?????????
???????? ?????????????? ?? ??????????????????????? ???????? ?? ????????????? ?????
????? ??????????-?????????????? ???????
??????: ????? ?????????? ??????????? ????? ? ???????? ???????? ?????, ????????????
?????, ????????? ????, ????, ?????? ? ????.
# 25 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 3. ??????? ????????? ??????? ???? ? ??????????? ???????? ? ? ???????? ???????????? ????????
??????????????
???. 4. ????????? ??????? ????????-???????? ??????? ?????????????? ? ??????? ??????? ??????.
???????? ????????? ??????-????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ???????????? ???????? ??????? ?????? ????????? ?????? ??????
???. 5. ?????? ????????? ??????? ????? ??????-??? ?? ????????? 400 ???. ???.( ?? ?.?. ?????????, 1981
? ????????????)
??????? ????????? ??????????? ???????????
??? ????????????? ?????????, ???????????
??????? ??????????? ?? ????? ?????? ? ?????
???????. ?? ??????????? ??????? ????????? ???? ? ?? ???????????, ? ????? ?? ??????????? ? ???????? ??????? ??????? ?????????? ??????????????? (????????????) ?????
??????? ???????? ????????? ??????? ??????????????????????? ??????? ????????????
(???. 5). ??????? ??????? ???????? ??? ??????
?? ??????? 1140-1180 ? (IV ??????? ? 10-150 ?),
1120 ? (III ??????? ? 90 ?) ? 1090 ? (II ??????? ?
60 ?). ? ?????? ???????? ???????? ????? ????? ????????? ?????????. ? ?????????, ?? ????
??????????????? ?????????? ??????? ???????
?? ?????? ??. ????-??-???.
?????? ????? ????? ??????????????
???? ????????????? ???? ???????? ???????????? ?????????? ?????. ??? ???????????? ?
???????? ?????????? ????????, ?? ?????????????? ?????? ????????????? ???????.
??? ???? ????????? ?????-??????? ? ?????
????????, ???????????? ????????????? ????????? ?????, ???????????, ???? ? ??????????? ???????????? ???????. ???????? ?????
?????????????? ??????? ?????????????
????? ????? ????????? ????????? ? ???????????????, ???????????????? ? ??????
????????. ?? ?????? ??????? (??????????????, 1982), ? ??????????? ?????? ???? ?
?????? ?????? ????? ?? ?????????? ???????
???????? ?????????????? ???????-?????????
??????-????????????? ?????? ?????????
?? ?????? ?? ???????? ??????, ? ????????
??????? ??? ?????????? ?????????? ?????
??????? ??????? ????????.
??????????
??? ????????????? ???? ?????????????? ???? ?????????? ? ?????????????
??? ???? ??????? ? ????????? ??????????.
?? ???????? ????? ??????? ?????????????
???????, ?????????????? ? ???????? ? ??????? ? ??????? ??????????????. ??? ????? ???????? ????? ??????????? ??? ???? ??????????
? ??????? ??????????? ????????? ??????????????? ???. ????????????? ? ??????????????? ??????? ?????????? ? ????? ???????
??????????, ??? ?????????????? ?????????,
??????? ? ?????????? ??????? ??????.
???????????? ????????? ??? ?????????? ????????? ???? (????? 08-05-00037-?)
# 27 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ???????????? ???????? ??????? ?????? ????????? ?????? ??????
?????? ??????????
????????????? ??????????? ???????? ?????????? (1982) /??? ???. ?.?. ??????????,
?.?. ????????. ?.: ?????, 260 ?.
????????? ?. ?. ????????? ????? ?????????? ????? ???? ? ?????????? ????????? (1946) //
???. ?? ????. ???. ?????. ? ??????. 2: 3-17.
???????? ?.?. (1981) ???????? ?????????? ???? (????????????, ?????????????, ??????????) //??. ????????-??????????? ?????.-????. ????. ????. ?.: ?????, ???. 27, 196 ?.
??????? ?.?. (1948) ????????? ??????? ???? ? ???????? ???????? ? ????????????? ??
?????????? //??. 2-?? ????????. ?????. ??????, ?.: ??????????, ?.1, ?. 367-378.
???????? ?.?., ??????? ?.?., ?????? ?. ?. (1971). ?????????????? ????????????? ??????? ?????????? ???????? //???. ???, ?. 103, ???. 5. ?.397-411.
??????? ?.?. (1947) ??????? ??????????? ???? ? ?? ??????? ?????????, ?????????
???????? //???. ?? ????. ???. ????. 5: 17-36
??????? ?.?. (1972) ???????????????? ? ??????? ?????????? ???? ? ???????????. ?.:
?????, 335 ?.
????????? ?.?. (1972) ???????????? ? ????????????? ??????????? ???????? ??????????. ?.: ?????, 293 ?.
??????? ?.?. (1959) ??????????? ????. ?.: ??????????, 456 ?.
Benson L.V. (1978). Fluctuation in the level of pluvial Lake Lahontan during the last 40000 years.
Quaternary Research 9, 300-318.
Bobek H. (1963). Nature and implications of Quaternary climatic changes in Iran ? Symposium
on change of climate. Rome, Oct.1961. UNESCO-WMO,. pp. 403-413.
Butzer K.W. (1961). Climatic change in arid regions since the Pleistocene. ? Arid Zone Res., ?17,
pp. 31-56.
Gilbert G.K. 1890. Lake Bonnevile. ? US. Geol. Surv. Monogr., vol.1, monogr., 438 p.
Morrison R.B. 1965. Lake Bonnevile, Quaternary stratigraphy of estern Iordan valley, south of
Salf Lake Sity, Utah. ? Geol. Surv. Profess. pap., ? 337, 80 p.
Morrison R.B. 1970. Conflicting pluvial-lake evidence on climatic changes between 14 and 9
millennia ago, with particular reference to lakes Lahontan, Bonnevile, and Searles. AMQUA Abstracts
1: 97-98.
Thomson R.S., Benson L.V., Hattori E.M. 1986. A Revised chronology for the Last Pleistocene
lake cycle in the central Lahontan basin. Quaternary Research 25, pp. 1-9.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????. ???????????? ???????? ??????? ?????? ????????? ?????? ??????
Quaternary Evolution of the Altay Mountains lakes Systems
Pavel S. Borodavko
Institute of Monitoring of Climatic and Ecological Systems SB RAS,
10/3 Akademichesky, Tomsk, 634055 Russia
Global cooling in the Early Pleistocene caused extensive continental glaciation in the northern
hemisphere including the arid areas of Central Asia. The reduction of temperatures (particularly
summer temperatures) reduced evaporation and strengthened the importance of precipitation. The
simultaneity of «lakes periods» (pluvials) and stages of glaciation is established experience confirmed
by investigations in the west of North America and Russia. In the Russian and Mongolian Altay new
evidence for similar conditions is found.
Key words: Altay mountains, pluvial lakes, neopleistocene, glaciation.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 30-37
~~~
??? 630*56:582.47:551.583.4(235.222)
???????????? ????????????? ?????????? ???? ???????
? ??????? ????? ??????? ????? ?????????? ??????
(??????????? ?????)
?.?. ???????*
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???
634055 ??????, ?????, ??. ????????????? 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
? ?????????? ???????????? ????????? ?????????? ??????????? ???????? ???????? ?????,
??????????? ? ??? ? ??????? ??????? ???? ? ??????? ????? ??????? ????? ????? ?????
?????????? ??????. ???????? ????????????? ?????? ??????????, ??????? ?? ????????????,
???????? ?????????????? ?????????????? ? ????? ? ???????????? ? ???????? ????????????
??????. ????????, ??? ?? ??????? ???? ???? ? ??????????? ????? ?????????? ???????
???????? ?? ????????? ??????????? ? ?????????. ????????????? ??????????? ????????
??????????? ? ????????????? ???, ???? ? ????, ????????????? ? ? ???????? ???. ?
??????? ????? ??????? ????? ??????? ??????????? ? ???????????? ????? ?? ?????????????
????????? ????????. ??? ????????? ????????? ???????????????? ???????, ??????????
??????? ? ??????????? ?????????? ??? ???????? ?? ??????? ???? ???? ? ?????-?????? ????.
? ??????????? ?????????????? ????? ????????????? ??????????? ?? ??????? ????????????
????. ???? ? ??? ?????? ?? ????? ??????? ?? ????????????? ??????????? ??? ? ????,
?????????? ????????????? ??????????? ??????????? ? ??? ? ???????? ???????. ??????????
?????????? ??????????????? ? ?????????????? ???????? ??????????? ? ????????? ?
????? ???????????? ?????? ?????????, ? ????????? ?????????? ??????? ????? ??????? ?????
?????????? ??????.
???????? ?????: ????????????????, ??????????????????, ???? ?????????, ???????????
?????????, ??? ?????????, ?????? ?????.
????????
??????? ????????????? ???????? ?? ???????? ??????????? ???????? ???????? ???????? ??????? ?? ???????? ?? ???????????????, ??? ????? ?????????? ???????? ??????
????????????? ?????? ? ????? ? ?????????????????? ?????????????? ??????? ? ???????
????????????? ??????? (??????? ? ??., 1996).
*
1
???? ??????? ????????, ?????????????? ?
???????? ? ???????? ? ??????? ????????,
????????????, ?????? ?????, ????????????
???????????? ??????????? ??????? ???????
???? (Fritts, 1976; Briffa et. al., 1990; ??????,
1986; ???????, ??????, 2000). ??????? ??????????? ?????? ??????? ??????? ????????
?? ????????? ???? ???????????? ?????? ??-
Corresponding author E-mail address: bochar74@mail.ru
© Siberian Federal University. All rights reserved
# 30 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ????????????? ?????????? ???? ??????? ? ??????? ????? ??????? ??????
?????, ? ????????? ??? ???????? ????????
???????????? ?????????, ??????????? ?????
???????? ? ????????????? ?????????, ???????????? ? ?????????? ????? (Luckman,
1990).
?? ?????? ?????? ???????????? ????????? ?????? ????????? ?????????????? ???????
?? ??????? ???? ???????? ??? ?? ??????????
????????????? ? ????????? ?????????????
????????, ?? ??????????? ????????? ???????????? ?? ??????????? ???????? ? ???????????
?? ???????? ?????????.
?????? ?????? ???????? ?????? ??????????? ???????????? ????????? ? ????????
???????????? ?????????? ???????????? ????? ??? ??????? ????????? ????????, ???????? ?? ???? ????????, ??????? ?????????
? ???????????? ???????? ?????????????,
?????? ????????? ? ???????????? ???????????? ??????? ?????? ????????? ?? ?????????? ????????????? ?????????.
????????? ? ??????
???????? ??????????? ????? ???????? ?????, ??????????? ? ??? ??????????? ?? ???????????? ??????????? ????? ????? ?????????? ??????, ? ?????????? ??????? ?????
??????? ????? ? ?? ??????? ??????? ????.
??? ?????? ???????????? ?????????? ????????? ??????? ? ?????? ?????? (?? 2500 ?) ?
???????? ?????????? ???????????, ???????? ?? ?????????? ?????? 800-1000 ? (???????,
1996). ????????? ????? ?????? ? ?. ??????
(2506 ?). ???????? ??????? ??????????? ??????
?? ????? ?????? ??????????, ???? ?????? ??????????? ????????? ???? (????????, 1981).
?? ????????????????? ?????????????
????? ????????? ? ??????????-?????????
??????????-?????? ????????? ????????????? ? ???????????? ????? (??????, ?????,
1967). ??????? ??????? ???? ????????? ?????????? ?????? 2100-2150 ? ? ????????????
????????????? ????????? ????????????. ?
??????? ????? ??????? ????? ?????????????? ???????????? ???????? ? ???????????????????? ????. ? ?????? ????? ??????? ?????
???????-????????????? ???? ??????? ??????
??????????????. ?? ??????????????? ??????? ??? ???????????? ?????? ? ???????-??????
????. ? ??????????? ?????? ??? ??????? ????
??????? ?????????????????? ????? ??????,
????? ???????? ??????????? (???????, 1999;
???????? ? ??. 2000; ??????? ? ??., 2002).
??? ?????????????????????? ????????
???????? ??????????? ???????? ???????? ??
??????? ??????? ????????????? ?????????????? ?????? ???????? ?????, ??????????? ?
??? ? ???? ??????? ????????, ?????????? ??
?????????? ??????? (2000-2100 ?). ? ??????? ????? ??????? ????? ??? ??????? ??????????? ???????? ?????????????? ??????? ? ????
??????? ????????, ?????????? ? ?????????
?????????? ????? (1550-1850).
????????? ????? ? ????? ??????? ? ???????? ?????? ????????, ? ?????????? 10-20 ??????? ?????? ?????? ? ?????? ??????? ???????. ?????? ???????? ????? ?????????? ?
??????? ?????????????? ????????? LINTAB
? ??????? ???????? TSAP. ?????????? ????? ??????? ?????????????? ????????????????? ????? ??????????? ????????????
??? ?????? ????????? COFECHA (Holmes,
1992?). ?????????????? ????????? ?????
??????????????????? ? ??????????? ? ?????????? ??? ?????? ????????? CRONOL
(Holmes, 1992b). ??? ?????????????? ????????
???????? ? ?????? «Statistica for Windows».
??? ???????????????????? ??????? ?????????? ?????????? ?????????????? ???? ??????
???????????? ??????? ?? 1936-1993 ??.
?????????? ? ??????????
?? ??????? ?? ???? ?????????????
???????? ?? ??????? ??????? ???? ??? ??????
# 31 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ????????????? ?????????? ???? ??????? ? ??????? ????? ??????? ??????
?? ????? ???? ????????? ??????????????
???? ??????????? ???????? ???????? ?????,
??????????? ? ???. ?????????? ???? ???????? ??????? ??????????????? ?? ??????? ???
?????? ?????????, ??? ? ????? ?????????????? ????????? ??????? ????. ???????????
?????????? ?????????????? ????? ?? ?????,
??????????? ? ??? ? ?????? ?????????????
???????? 0,53, 0,62 ? 0,46 ??????????????. ???
??? ????????? ?????????? ?? ? ??????????
?????????? ??? ??????? ??????? ???? ????? ????? ?????????? ??????. ????????????
?????????? ?????????? ????????? ??? ???????? ????? 113 ???, ??? ? 108 ???, ??????????? ? 65 ???. ????????? ?????????????? ?????????????? ?????????? ????????? ?????
????????? ? ????. 1.
?????????? ???????? ????????????????
? ???????????? ?????????? ? ?????????????? ? ?????????? ?????? ??????????? ?
???????????. ??? ????????? ????? ??????????, ??? ???????? ??????????? ???????? ?????, ??????????? ? ??? ?????????? ????????? (???. 1). ??????????? ?????????? ?????
???????????? ???? ????? ? 0,48. ???????????? ???????????? ?????????? ????????????
???????? ? ??????? ??????? ??? ??????????
? ???????? ??????????????? ? ????????????
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???????? ? ??????????? ??????????.
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?? ??????? ??????? ????????????? ?????????? ?????????? ?????????? ? ????????????
? ???????? ?????????????? ??????? (???????????).
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??? ????????? 0,42, ? ???????????? ???? ?
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?????????? ? ???????? ??? (r = ?0,31). ? ????? ????? ????? ?????????? ?????????????
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?? ????: 0,39, 0,34 ? 0,31 ?????????????? ?
???????????? ????????????? ??????????? ?
???????? ??? (r = ?0,45). ? ????? ? ??????????? ???? ?????????? ????????????? ?????
???????? ? ????????????? ???????? 0,3 ? 0,27
??????????????. ??????????? ?????? ??????????? ???????? ??? ? ??????????????? ????????????? ?????????? ?? ?????????????.
??? ??????? ????? ??????? ????? ????? ???? ????????? ?????????? ????????????????? ?????????? ?? ???????? ?????, ??????????? ? ??? ? ???? ??????? ????????.
????????? ?????????????? ??????????????
?????????? ?????????? ????????? ? ????. 2.
?????????????? ?????? ??????????
?????????? ????? ? ?????? ?????????????
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??????? ?????. ??????? ??????????? ??????????? ?????????? 0,57. ?????????? ?????????????? ????? ?????????? ? ????????
????? ?????????????? ? ?????? ????????
??????????? ??????? ????? ??????? ?????
? ?????? ???????? ??????? ?????????????.
??????????? ???????????? 0,6; ?? ?????, ???????????? ?.?. ???????? (1986) ???
?????? ??????? ?????????? ?? ?????, ???????????? ????? ???????. ??????? ??????????? ?????????? ????? ???????????????
??????? ?????????? ?????? ?????????? ??
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? ???????????? (1600?1650 ?) ????????? ??????? ? ????????, ?????????????? ? ??????
???????? ??????????? ??????? ????? ??????? ????? (1650?1850 ?). ?????????? ????????
# 32 #
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1550 1575 1600 1625 1650 1675 1700 1725 1750 1775 1800 1825 1850 1875 1900 1925 1950 1975 2000
????
???. 2. ?????????? ?????????? ??????????? ???????? ?????, ??????????? ? ??? ??????? ????? ???????
?????, ?????????? 7-?????? ?????????? ???????
???????????????? ????? ?????????? ?????????? ?? ???????????.
?? ????????? ?????????? ??????? ?????????? ?????????? ?????????? ????? ????? ??? ???? ?????????? (???. 2). ?????????? ????????????????? ?????????? ?? ?????
????????? 437 ???, ?? ??????????? ? 391 ???,
?? ??? ? 260 ???. ??????????? ??????????? ?????????? ?????????? ?? ????? ? +0,53.
??????????? ?????????? ? ??????????? ???????????????? ? 0,16. ??????????? ??????????? ?????????? ?????????? ?? ??? ? +0,48.
??????????? ?????????? ?????????? ? 0,16,
??????????? ???????????????? ? 0,17. ? ??????????? ? 0,16 ? 0,21 ??????????????.
??????????? ?????????? ??????????
????? ?????????? ??????? ???????????? ?
???????????? ???????? ???????? ???????? ?
?????? ????????????? ????? ????????????
????????? ???????. ???????? ?? ?????????
???????????????? ?????????? ?????????? ?
??????????? ???????? ?????????? ?????,
??????????? ???????????? ????? ??????????,
??? ???????????? ??????????? ???????????
???????? ???????? ? ????????? ??????????
??????? ????? ??????? ????? ????? ???????
?? ?????-?? ????? ????????. ? ???????????
??????????? ???????? ?????, ???????????
? ??? ????? ????? ??????? ?????????? ?
???????? ?????. ?? ?????????????? ? ?????????, ? ????? ? ??????????????? ???????
????????? ?????, ???, ??????, ??????? ??????? ????????????? ? ??????????????? ????????? ?????, ??????????? ? ??? ?? ???? ?
?? ?? ??????????? ??????? ???????? ? ???????? ????????? ?????????? (??????? ? ??.,
1998; Vorobjev et ?l., 1998).
??? ????????? ?????????? ??????????
? ???????????? ? ???????? ? ??? ?? ????????
?????????? ????? ??????????? ???????? ?
??????? ?????????? ?????????????? ???????
?????????? ? ???????????. ?????????????
??????????? ?? ??????? ???????????? ????
0,45. ? ????? ????? ????????????? ??????????? ?????????????? ?? ??????? ???????????? ??? ? ???? ? 0,26 ? 0,32 ??????????????,
# 34 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ????????????? ?????????? ???? ??????? ? ??????? ????? ??????? ??????
? ??? ?????? ? ???????????? ??? ? 0,44. ??????? ????????????? ??????????? ????????
????? ? ??? ? ???????? ??????? (r = ?0,28) ?
(r = ?0,33) ??????????????. ??????????? ?????? ??????????? ???????? ???????? ??????????? ? ????????? ?????????? ??????? ?????
??????? ????? ? ???????? ?????????????? ??????? ?? ???????????.
?? ????????? ?????????????? ?????
?????????, ??? ???????? ??????????? ???????? ???????? ?????, ??????????? ? ??? ??
??????? ??????? ????????????? ? ???????????? ??????????? ??????. ? ????? ? ??????????? ??????????? ????? ????????????? ?????
?? ???????? ????????????? ? ??? ?? ???? ?
???????????? ????????, ????????????? ? ?
???????? ???. ? ????????? ?? ??????????
??????? ????? ??????? ????? ??????????? ????????? ????????? ??????? ???????????????
????? ?? ????????? ??????????? ? ???????.
?????????? ??????????? ??????????? ?????
?? ?? ??????? ???????????? ???????-??????
???????, ? ?????? ? ???????????? ????. ?????????????? ????? ??????????? ???????
???????? ?? ?????? ? ????? ? ???.
??????? ??????? ??????????????? ?????
? ???????????? ???????????, ?? ?????? ??????, ??????????? ??????? ?????????????
??????? ???????? ??????? ?????????????.
??????? ?????? ????? ? ??????? ???? ????????? ????????????? ????? ????? ? ??????????? ? ???????????? ???. ?? ??????????????
?????? ?????? ??????? ??????? ??????? ????,
??????????????? ??????, ??????? ??, ????????? ????? ?????? ????????? ???????? ????? ?
???? ? ??? ??????? ??? ????? ?????????? ?
???????? ?????? ???? ? ????. ?????????????
????? ???????? ????? ? ??????????? ? ???????????? ????????, ??????, ????? ?????????
«??????????? ???????? ? ??????? ???????», ?.
?. ???????? ???????? ????? ? ????? ???????
???? ?? ??????? ???? ??? ?????????.
????????? ??????? ???????????????
????? ? ?????-?????? ????? ???????????
????? ?????????????? ????????? ?????????????, ??? ? ???????????? ???????????. ???
????????? ? ?????????? ??????????? ?????????? ?? ???? ??????????? ???? ?? ???????
????, ????????? ????????? ???????, ????????? ??????? ?????????? ?????? ???????,
?????????? ????????????????? ?????????????? ???????. ??? ??? ?????? ????? ???????????? ??????? ??????????? ?????? ??????? ?? ?????????? ??????? ????????. ??????
????? ??????, ? ????????? ??? ????? ? ???,
???????? ????????????? ????? ???. ??? ??????? ? ???, ??? ? ????????? ????????????
?????????? ??? ??????? ???? ?????? ????? ?
??????? ???? ???? ??????? ?????????, ??? ?
????????????? ?????????????. ????????,
?????????? ????? ????????????? ????? ???????? ??????????? ? ????????? ??????????
??????????? ?????? ? ???????????? ????,
? ? ???? ???????????. ???? ? ???? ????????
???????????? ????????? ??????????? ??????????? ???????? ?????? ?? ??????????? ?????? ??????? ?????? ? ????, ?????????? ?
??????????? ??????? ???? ???. ??????????? ????????????? ??????????? ????? ? ??? ?
???????? ???????, ?? ??? ??????, ??????????
????????. ????????? ????? ????, ??????????
?????????? ? ????????? ????????? ???????
? ???? ??????? ????? ? ???????? ??? ??? ?
????? ???? ????? ??????? ???? ????????, ????????? ????????????????? ??????????????
???????.
??????
??? ????????? ?????????? ?? ?????, ??????????? ? ???, ?????????????? ?? ??????? ??????? ???? ? ????? ????? ??????????
??????, ??????????, ??? ???????? ?????????? ????????? ?????? ?? ????? ?????, ??????? ??????????? ???????? ????? ? ???????-
# 35 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ????????????? ?????????? ???? ??????? ? ??????? ????? ??????? ??????
???? ?? ????????? ??????????? ? ???????
?????????.
? ?????????? ??????? ????? ??????? ????? ??????? ???????? ????? ? ?????? ????????
?????????????????? ????? ?????????? ??????? ????????????. ????????? ???? ???????????? ???????? ???. ?????? ????????????
???????????????? ?????????? ????? ?? ??????? ??????? ???? ????????? ??????? ? ????????, ?????????????? ? ???????? ??????????
??????? ????? ??????? ?????.
??????? ?????????? ? ???????????? ??????????? ????? ??????????????? ????? ??
??????? ???? ??????????????? ? ?????????????? ???????? ????????????? ????????, ?????? ????? ??????????? ? ???????, ? ????????
?????? ??????????.
? ??????? ????? ??????? ????? ????? ??????????? ???????? ??????????? ? ???????????? ????? ? ?????????? ??????????????
?????? ? ????????? ???????? ? ????? ????????, ??? ??????? ?????????? ???????????????? ???????, ??????????? ???????
? ???????, ????????? ???????????? ??????????????? ????? ???????? ??????????????
????? ?????????? ??????.
?????? ??????????
???????? ?.?. (1981) ??????? ?????????????? ???????? ??????? ????? ? ????????. ?.:
?????, 132 ?.
??????? ?.?., ?????? ?.?, ???????? ?.?. (1998) ????????????????????? ?????????????
???????? ????? ????? ?????????? ? ??????????? ????????? ?? ????????? ???????? (?????) //
???????? ??????? ?????, ?????????, ?. 11-12.
??????? ?.?. (1999) ?????????? ????????? ???????-????????????? ?????????? ?? ????????? ???????? (??????????? ?????) //???????????? ??????????? ?????? ????????? ??????,
??????????, ?.18-19.
??????? ?.?., ???????? ?.?., ???????? ?.?. (2002) ??????????? ????????????? ???????? ???????? ?????????? ?? ????????? ??????? ????? ?????????? ?? ????????? ????????
???????????? ????? //???????????? ??????? ???????, ???. 2 /????????-????????? ??????
?????????????? ????????? ????, ??????????, ?. 177-181.
??????? ?.?., ?????? ?.?. (2000) ???? ? ????????? ???????? ????? ???????. ???????????: ?????, 232 ?.
??????? ?.?., ?????? ?.?., ?????? ?.?. (1996) ??????????????????? ???????????? ?
?????-????????? ??????????. ???????????: ?????, 246 ?.
???????? ?.?., ???????? ?.?., ????? ?.?., ??????? ?.?., ?????????? ?.?., ??? ?.?., ??????? ?.?., ?????? ?.?, ??????????? ?.?. (2000) ??????????? ?????????????? ????????????
?????????? ????? //???????? ???????????? ????????, ???. 6. ???????????: ???-?? ?? ???,
?. 21-22.
?????? ?.?., ????? ?.?. (1967) ???? ???????? ? ????????????? ????? ??????? ?????. ?.:
?????, 224 ?.
??????? ?.?. ?????. (1996) ???????-?????????????? ?????? ??????????????????. ???????: ???-?? ???. ??-??, 240 ?.
?????? ?.?. (1986) ???????????????? ??????? ??????? ???? ?? ?????. ?.: ?????, 136 ?.
Luckman B. H. (1990) Mountain areas and global change: a view from the Canadian Rockies.
Mountain research and development, Vol. 10, No. 2. University of Western Press, Ontario, p. 183-195.
# 36 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????????? ????????????? ?????????? ???? ??????? ? ??????? ????? ??????? ??????
Briffa K., Cook E. (1990) Method of response function analysis. Methods of dendrochronology.
Dordrecht-Boston-London, p. 240-247.
Fritts H.S. (1976) Tree rings and climate. London: Acad. Press, 245 p.
Holmes R.L. (1992?) Program COFECHA: Version 3. Tuscon: Laboratory of tree-ring research,
University of Arizona.
Holmes R.L. (1992b) Program CRONOL. ? Tuscon: Laboratory of tree-ring research, University
of Arizona.
Vorobjev V., Savchuk D., Dutova N., Bocharov A. (1998) Dendrochronological study of
interrelationships between Siberian stone pine and Siberian larch in subalpine forests of Altai
mountains. Forest ecosystems and land use in the mountain areas. Seoul, p. 556-561.
Climatogenetic Radial Growth of Conifers in the Upper Forest Belt
of the Seminsky Range (the Central Altai Mountains)
Anatoly Yu. Bocharov
Institute of Monitoring of Climatic and Ecological Systems SB RAS
10/3 Academichesky, Tomsk, 634055 Russia
Tree-ring width chronologies were built for Siberian stone pine, Siberian larch and Siberian spruce
in the forest limit and the upper forest belt of the Seminsky Range. The chronologies were compared,
their synchrony and statistics were estimated, and correlation with temperatures and precipitations
were received. In the forest limit the growth responses to temperature and precipitations are similar
in pine and large. Tree ring width correlates positively with May, June and July temperatures and
negatively with May precipitations. In the upper forest belt the responses of the trees are different
because of degree of density, crown closure, and phytocoenological conditions are increased from the
forest limit to the upper forest belt. The ring width and June temperatures are positively correlated
in larch. In pine and spruce the correlation is positive with May and July temperatures and negative
with August precipitation. The results are evidence of paramount importance of temperature and
precipitation in forest trees of Seminsky Range.
Key words: dendrochronology, dendroclimatology, Siberian stone pine, Siberian larch, Siberian
spruce, Altai Mountains.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 38-53
~~~
??? 581.526:574.45
????????????? ?????????????? ????????????
? ????????? ????? ??????????? ??????? ???????? ??????
?.?. ??????????*
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???,
634055 ??????, ?????, ??. ????????????? 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
?????????? ?????? ??????????? (1999-2007 ??.) ????????? ??????? ????????, ????????
???????? ????????? ? ?????????, ?????? ????????? ????????? (NPP) ???????? ?????????????
????????????? ? ?????????? ????, ????????????? ?? ?????????? ??????????? ???????
???????? ?????? (??????? ???????). ????????, ??? ?????? ????????? ? NPP ??????????
?????? ????? ? ??? ???? ???? ?? ????????? ? ????????????. ???????? ??????? ?????????
? ????????? ???????????? ??????? ???????? ????????????? ???????, ??????????
????????????? ? ????????????????? ?????????. ???????? ??????????? ????? ????????
???????? ????, ???????????? ? ???? ? ?????? ????????? ?????????? ???????? ?????????.
???????? ?????: ???????? ??????????, ????????????? ??????????????, ???????? ????????
?????????.
????????
???????? ?????????? ?????? ???????????? ???? ? ?????????? ??????????? ????????
? ????????. ?????????, ??? ?????? ????????
?????????? ????????? ? ?????????????
?????????? ????????, ?? ???? ? ??? ??????????? ??????? ??????????? ???????? ????????? ? ???? ?????????????? ? ????????
?????????. ???????? ?????????? ????????
????? 3-5 % ?? ??????????? ????, ?? ???
???? ? ??? ????????????? ???????????? ?????? ???????? ? ???? ???????? ???????, ??????? ?????????? ?? 120 ?? 455 ??C (Gorham,
1991; ??????????, 1994). ?? ?????? Botch et
al. (1995), ?????? ???????? ? ??????? ??????
*
1
????? 215 ??C. ??????? ???????? ?????????
???????? ?????? ????? 42 % ?? ??????? ????? ??????, ??? ???? ? ?????????? ????????
?????? ?????????? ????? 36 % ?? ??????
???? ?????????? ???????? ?????? (??????????, 1994; ??????? ? ??., 1994; ?????????
? ??., 1998). ?? ?????? ???????, ???????????????? ?????? ???????? ?? 55 ?? (???????,
2007) ?? 70 ?? (Sheng et al., 2004).
??????????? ???????? ? ???????? ??????????? ? ??????????? ??? ? ???? ????????????? ???????? (????????????? ??????????????, NPP ? net primary production) ? ???? ??
???????? ??????? ????????, ??????? ????????????? ?????????? ??????????? ???????-
Corresponding author E-mail address: golovatskaya@imces.ru
© Siberian Federal University. All rights reserved
# 38 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
??? ?????????? ???????. ????? ?????????
????, ????????? ? ???????????? ????????,
???????? ?????????? ???????? ???? ?? ??????? ????. ?? ?????? ???????, ???????
?????????????? ???????? ??????????? ???????? ???????? ????? 16-29 ??/?2 ? ??? (??????????, 1994), ??? ?????????? 7-30 % ??
????????? ????????? ?????????????? ?????
(??????????, 1994; ?????????, 1985).
???????? ????????????? ?????????????? ????????????? ?????? ????????? ???
????????? ??????? ?????????? ?????. ?
60-??70-? ??. ???????? ???? ? ?????? ????????????? ????????? ??? ???????????????
??????????????? ???????? ?????????,
???????????????? ? ????????????? ?????????????? ? ????????? ???????? ???????,
??????????? ??? ???????? ?????. ????????? ? ???????? ??????????, ????? ???????
???????? ???????? ??????? ?????? ???,
??????, ????????? ? ??. (Backeus, 1990;
Bartsch, Moore, 1985; Dyck, Shay, 1999; Grigal
et al., 1985; Vasander, 1982; Moore et al., 2002
? ??.). ?? ?????????? ??????? ???? ??????????? ???????????? ? ??????????, ???????,
???????? ??????, ??????? ??????? (?????????, 1967; ????????, ??????, 1976; ??????,
???????, 1975; ??????? ? ??., 1994; ?????,
2003; ??????????-????????, 2001; ?????????, 1967; ?????????, 2001, 2007; ?????????? ?
??., 1978; ????? ? ??., 1984; ????????, 1986 ?
??.). ? ?????????? ???????? ??????? ??????????? ???????? ?? ???????????? ???????????????? ???????? ?????????, ???????????
???????? ??????????, ??????????????? ????????????? ?????????????? ???????????.
????????? ?????? ?? ????????????? ?????????????? ???????? ????????? ???????? ?
?????????? ?.?. ????????? (1993). ????????
?????????????? ??????? ?? ???????? ???????
????????????? ??????????, ????????????? ?
???????????????? ???????. ?????? ?????-
???, ???????? ?? ????????????? ??????????????, ???????? ??????????????? ?????
???????? ??????????, ??? ??? ? ???????????
?? ??????? ?????? ?? ??????? ???????????
??????, ???????-??????, ???????-???????
??? ??????? ??????????. ???????? ?????
??? ???????? ????????????? ?????????????? ????????? ???????? ???????, ????????? ?
????????? ?????????? ?????, ? ?? ????? ???
????????? ?????????, ???????????? ?? 40 ??
90 % (Aerts et al., 1992; Titlyanova et al., 1999),
??????????? ?? ???????.
? ????????? ???? ?????????? ???????????? ???????????? ?????????????? ??????????? ???????? ?? ???????????? ??????, ??????????? ?????????? ?????????????? ???????????
?????, ????????? ????????? ???????????????? ???????? ????????? ? ?????????
?????-??????????? ???????? (??????? ? ??.,
1993; ??????????? ??????, 2000; ?????, 2001;
??????????-????????, 2001; ?????????, 2001
? ??.). ????????? ????????? ?????????????
?????????????? ?????????????? ????????
????????? ??????? ?????????? ????? (?????????, 1985; ??????, ????????, 1977; ???????? ? ??., 1994; ????????? ? ??., 1995, 2000; ?????, 2003 ? ??.), ?????? ???????? ?????????
???????? ????????? ??-???????? ?????????? ??????, ???????? ? ?????? ???????????
??????????, ??? ? ?????????? ????? ????????? ??????? ??????? ????????? ?????????????? ??????????? ???????? ????????? ???
????????? ??????? ??? ??? ?????????????
?????????????.
??????? ? ??????
???????????? ????????????? ?????????????? ???????????? ? ????????? ????????
????????????? (???) ??????????? ? 1999 ??
2007 ??. ??????? ???????????? ??????????? ?? ?????????? ?????????? «?????????»
????????? ??????????? ????????????? ?
# 39 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
????????????? ?????? ?? ??? (??????????
?????, ??????? ??????? 56o58`?? 82o36`??).
????????? ????? ???????????? ?????????? ?? ??????-????????? ??????? ????????
???????????? ?????? ?????????? ? ?????????? ??? ????-?????? ? ??????????????? ???????????? ?????? ???????? ??????????? ????????????? ????. ???????????? ???????????
?? ???????????? ??????????? ???????????
??????? ?. ????, ??????? ?????????? ????????? ???????? ???? ???????????? ????????
???: ???????-?????????????-????????? ????????? ? ??????? ???; ????? ?? ????????? ?
?????????? ?????????? ? ?????? ???; ???????? ???????-????????? ????. ??? ?????????
?????? ?????????? ?? ??????????? ??????? ??????????????? ?????? ??????? ???????? ??? ? ??????????????? ??????? (??????? ? ??., 2003). ???????? ???????? ??????
? ?????? ???????????? ?????????? ?? 1 ?? 3 ?
??? ???????? ?? 3 ?? 5 ???. ??? (??????? ?
??., 2000).
??????? ??? ? ??????? ????????? ??????????????? ??????? ??????? 20-30 ?. ??????????? ??????????? ???????? ????????? ? ????????????? ??????? ??????? ?? 50
??, ??????????? ????? 50 % ???????????.
?????????????? ???????? ???? ???????????
???????-?????????????-????????? ??????????. ????????? ???? ??????? ?? ????? (Pinus
sylvestris) ? ?????????? ?????????? Pinus
sibirica ? Betula pubescens. ??????? ?????? ????????? 18 ?, ??????? ??????? ???????
????? 20 ??. ??????????? ???????? ?????????? ????? 90 %. ?????????????? ???? ??????
?????, ?????????? ????????? ?????? 50 ??,
??????????? ???????? (?.?.) ? 90 %. ??????????? ???????? Ledum palustre, Chamaedaphne
calyculata, ???????? ??????? ????????
(Vaccinium vitis-idea) ? ?????? (Oxycoccus
microcarpus). ? ???????? ??????? ??????????? Carex globularis, Eriophorum vaginatum,
Rubus chamaemorus. ????? ?.?. ?????????
????? 15 %. ??????????? ???????? ???????
(96 %) ????????? ????????? ??? (Sphagnum
angustifolium). ?? ???????????? ??????????
??????????? ????????? ?????????? ???????
????, ?? ??????? ???????? ? ????????????
???????????? ?????????. ???????? ??????
???????? ???? ????? ???????? 1 ?. ? ????????? ?????? ? ???? ????????? ????????? ?????
??????? ??????? ??????????, ????? ???? ?????????? ???? ??????: ????????-?????????
? ????????-????????. ??????? ???? ??????????? ???????-????????? ??????.
?????? ???. ??????????? ??????? ????
????????? ??-?? ???????? ?????????? ??????? ??????? ??????? ??????? ????? 30 ??,
????????? ?? 3 ?. ?????????????? ??????? ????
????? ????????? ? ???????-?????????????????????? ?????????? ? ?????? ??????. ?????????? ????????? ???? ??????????? Pinus
sylvestris f. Litwinowii. ??????? ?????? ????????? 2-3 ?, ??????? ??????? ??????? 3 ??. ?.?.
?????????? ????? 30 %. ?????????????? ????
?????? ??????? ?? ???????????????, ?????
??????????? ???????? ? 60-70 %. ???? ?????? Ledum palustre, Chamaedaphne calyculata,
Andromeda polifolia ? Vaccinium uliginosum. ??
???????? ????? ?????? ?????? ????????????
(Oxycoccus microcarpus). ???????? ???? ????? ?.?. ????? 5 % ? ??????????? ?????????
Eriophorum vaginatum, Rubus chamaemorus
? Drosera rotundifolia. ? ??????? ??????? ??
?????????? ?????????? Sphagnum fuscum
(95 %), ????? ????, ?? ??????????? ?????????? ??????????? Sph. angustifolium ? Sph.
magellanicum. ???????? ?????? ??????? ????
????????? ???????? 3 ? ? ????? ?????????
??????? ??? ????????. ? ????????? ?????? ?
???? ????????? ?????, ??????? ???????????
?????? (1.5 ?) ???? ????????? ????? ????
?????: ???????????? ?? ??????? ????????
?????????? ? ?????? ? ? ??????. ?? ????????
# 40 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
????????? ? ????????? ??????? ????????????? ?????? ????????? ??????????? ?????????????????? ?????.
???????-?????????
????
???????? ??????????? ????? ????????? ???????.
??????????? ??????????? ??????????????? ???????????????? ??????? ?? 20 ??.
?????????????? ???????? ???? ????????? ?
?????????????-???????-????????? ??????????. ?????????????? ???? (Chamaedaphne
calyculata, Andromeda polifolia, Oxycoccus
microcarpus) ??????? 10 ?? ?????? ?????, ????????? ?? ????????????? ????????? ????????????. ??????????? ???????? ??????????????? ????? 20 %. ? ???????? ????? (?.?. 40 %)
???????? ??????? Eriophorum vaginatum
? Carex rostrata. ??????? ???? ??????????? ?????????? ?????? ????????? ????
(Sphagnum fuscum, Sph. angustifolium ? Sph.
magellanicum), ??????????? ???????????.
??????? ???????? ??????? ?????? ?????
100 %. ???????? ?????? ???????-?????????
???? ????????? ???????? 2,5 ?. ? ?????????
???? ?????? Carex limosa, C. chordorrhisa,
C. diandra, C. dioica, Rumex aquaticus, Cicuta
virosa, Calamagrostis neglecta, Galium palustre,
Oxicoccus palustris ? ??. ??????? ?????? ?????????, ??????????? ???????? 40-60 %. ?
??????? ??????? ?????????? ??????? ???:
Tomenthypnum nitens, Aulacomnium palustre,
Helodium blandowii, Drepanocladus aduncus,
Brachythecium mildeanum, ???? ???????????
Sphagnum warnstorfii, Brium pseudotriquetrum
? ??????. ???????? ?????? ????????? ? ????????? ???? ???????? ???????. ???????
???????? ???????? ?????? ? ???????????
???????? ????? ?????? ????? 3-4 ?, ???????????? ??????? ???????? ? ????????????
????? ????????? ????????? ? ? 5,5 ?. ???????? ?????? ?? ??????? ????? ??????? ?????
??????????? ?????????? ???????? ? ?? ??????? 2-3 ? ?????? ??????? ???????? ????????? ? ???????-????????? ?????? ?????.
????????????? ?????????????? ??????????????? ????? ???????? ???????????, ????????? ?? ??????? ????? ???????? ?? ??????-
?????? ????????? ?????? ????? ?????????
????? ????????? ? ????????-????????? ????
(2 ?). ??????? ???? ?????? (1 ?) ?????????
????????? ?????? ? ????????-???????????
??????? ?????? ??????? ??????????.
????? ????, ??????????? ????????????
????????????? ?????????????? ?? ?????????
?????? ??????, ???????? ????? 400 ??, ????????????? ?? ?????? ???????????? ???????
?. ?????? ? ???????????? ?. ?????????.
????????-???????? ?????????. ???
??????????? ????? ????????? ??????? ???????? ??????? ?????????? ??????? ? Betula nana,
? ?????? ?????? ??????????? ????? ? ?????
(?????? 2-6 ?, ??????? ??????? 2-7 ??), ????? ????, ??????????? ?????? ?????? ??????:
Salix cinerea, S. rosmarinifolia, S. pentandra.
??????????? ???? ??????????? ?????????
??? ???????-???????? ????? ???????? ??-
?????? ?????? ???????, ? ????? ??????? ???
???????? ?????????? ???????? (??????, ????????, 1977). ????? ????? ???????? ???????? ? ???? ????????? ???????: ????????? ? ????? ???????????? ???????? ????????
? ????????? ? ??????? ???????????? ???????? ????????. ????????? ?????????????? ??
????????????????? ????????? (??????? ????? ????, ????? ?????? ???????????, ???????
?????) ? ??????????????????? ?????????
(???????????????? ????? ?????? ? ???? ???????, ????????????? ?????? ??????????? ?
????? ????? ????????), ? ????????? ???????
?? ???????? ?????????????????? ???? ????? ? ???????? ????? ? ???????? ?????? ????
? ??????????? ? ?????.
????? ???????? ????????????? ? ??????
????????????? ?? ???????? (????? ???? ? ?????? ???????). ????????? ????????? ??????-
# 41 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
???? ??????? ??????? (??? ????? ??????????
?????), ????????? ? ??????? ?????????. ???
??????????? ??????? ????????? ?????????
???????????? ???????? ???????? ? ????????
???????? 50?50 ?? ? 3-5 ????????????. ???
??????????? ?????? ????????? ?????????
????? ???????? ???????? 10?10 ?? ? ???????? ?? 20 ??. ??? ?????? ???????????????
???????? ????? ???? ? ????????????. ????
????????? ? ??????????? ???????? 50*50 ??,
????? ???? ??? ??????? ??????????? ? ?????,
? ????? ??? ??? ???????? ???????. ?????
????, ? ?????? ?????? ? ?????? ????? ??????
? ?????? ? ???????? ????? ???? ? ????????????, ?? ??? ?? ?????????? ????? ? ?????????
(? ???????? ??????? ?????? ?????????? ???
????). ???????????? ???????? ????????? ??
???????? ? ???????????? ? ?????????, ??????? ??????????????? ? ???????????????
?????????????. ???????? ???????? ???????????? ?????????? ? ??? ?? ????????. ???????? ???????? ???????? ???????? ????????
???????????? ?? ???????????? ???????? ??????? ?????? ???????? ? ???????? ?????, ???????? ? ??????? ????.
?? ?????? ?????? ?? ??????????? ??????? ????????? ?????????????? ??????
????????? ????????? ????????????? ???????
??? ????? ????????? ???? ????? ???????? ??
??????? ??????? ?? ???.
??????????? ????????? ?????????? ??
?????????????????? ????????????????: ???????????? ???????, ??????????? ???????, ???????????? ???????????????? ???????????
(???), ???????? ?????? ???????? ??? (???).
?????????? ? ??????????
???????? ???????
???????? ??????? ???????? ??????
????????, ??????????? ??????? ?? ????????? ????????????? ???????. ? ??????
???????????? ?????? ?? ??????????? ???-
???. 1 ?????????????? ??????????? ??????? ?
?????????? ??????? ?? ??????? ?????? ?? 19992007 ??.
???? ? ?????????? ??????? ?? ?????????
???????????? ??????, ????????????? ? 30
?? ????????? ??????? ?????????? (WMO ID
293280, 57 °05'N, 81 °55'E). ??????????????
(1988-2007 ??.) ????????????? ???????????
????? 0,53 °?, ?????????????????? ?????????? (STD) ? 0,56 °?. ??????? ??????????? ?????????? ?? ????? 0,35 °? ? 2006 ?? 1,74°? ?
2007. ??????? ??????? ?????????? ??????? ?????????? (488±117) ?? ? ???. ??????????? ??????? ??? ??????????? ? ??????? ?????????
?? ???. 1. ??????? ????????????????? ??????????? ??????? 193 ???. ??????? ??????????? ?????????????? ??????? (???-????????)
????? (13,6±0,7 °?). ????? 58 % ??????? ???????? ? ??????? ?????????????? ??????? (???
(285±91) ??).
?????? ???????????? ???????????????
??????? ????????? ?????????. ????????
???????? ???? ???? ? 2005-2006 ??. ???????
??????????? ?????? 2006 ?. (????? 31,9 °? )
???? ?? 14,2 °? ???? ?????. ???? 2001-2002
? 2006-2007 ??. ???? ???????????? ???????. ??????? 2006, ?????? 2007 ? ?????? 2002
???? ?? 8,57, 7,6 ? 8,4 °? ?????? ?????. ???????? ???? 2005-2006 ??. ???????????????
?????? ??????????? ???????. ????????????
??????? ???????? ??????? ?????????? 56 ??,
??? ?? 19 ?? ???? ????????????? ?????.
# 42 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
???????????? ???????? ?????????? ???????
???? ????? ?? ????? ??????, ??? ??????, ???? ???????? ???????????? ???????? ???????
??????. ?????????? ?????????? ??????? ??????????? ????? 2000-2001 ??. ??????? ???????? ??????? ????????? 83 ??, ????? ??????
??????? ????????? 165 ?? (88 ?? ???? ?????).
????????????? ??????? (???-????????)
? 1999-2007 ??. ???? ??????, ?? ????????? ?
20-?????? ???????. ???? ? ???????? 2006 ?.
???? ???????????? ??????? (?? 3,81 ? 3,35 °?
???? ?????), ?? ??????????? ??????? ??
2,63 °? ???? ?????. ?????? ???????????
2001 ? 2003 ??. ???? ?? 0,86 ? 0,9 °? ??????
?????. ???? ???????????? ?????????? ? ??
?????????? ??????? ? ??????? ??????????????
???????. ???????? ??????????? ???? 1999 ?.
(158 ??) ? 2006 ?. (189 ??). ????? 2007 ?. ??????????? ???????????? ??????? ??????????
??????? ? 416 ??, ??? ? ??????? ???? ????,
??? 20-?????? ????????????? ?????.
????? ???????? ?????????? (????
10 °?) ? ???????????????? ??????????? ?????????? (???) ????????????? ?????- ? ??????????????????? ?????????????? ???????
??????????. ? ?????????? ?????? ????????????? ??????? 1999-2007 ??. ???? ? ??????? ???????????? ??????? (??????? ?? 1999-2007 ??.
????? ???????? ?????????? 1907 °?) ? ???????????? ???????????? (??? 1,2). ???????????? ????? ???????? ?????????? ??????????? ? 2001 ?. (2063 °?), ??????????? ?
2007 ?. (1698 °?). ???????? ?????????? ???
????????????? ?????? 1999 ?. (??? 0,54), ?
???????? ??????? ? 2007 ?. (??? 1,9).
??????? ???????? ??? ?? ???????????? ?????? ??????????? ????????????? ??
???????? ???? ? ??????????? ???????? ??????????? ???? (????. 1). ?? ???. 2 ????????
???????? ???????? ?????? ???????? ??? ??
??????????? ???????? ???????????. ?????-
???? ????????? ?????? ?????? ?????????????? ??????? ???????????? ?? ??????? ????,
??????????? ? ?? ???????? ????. ?????????
?????? ??????????????? ???????? ????????
???????? ???.
?????? ????????
????? ?? ???????? ??????????? ????????????? ?????????????? ???????? ????? ?????????. ???????????? ??????? ????????? ????????, ??? ???????????? ?????? ??????????
??? ?????????? ??????, ????? ????????????
????????????? ????????????? ???????? ????????? ??????????????? ??????? ???, ?????
??????? ?????? ??? ? ???????? ???? (???. 3).
?????????? ????????? ???????????? ?????????? ????????????? ???????. ? ???????
?????? ????????? ?? ????? ?????????? ??
?????? ???????????? ???. ?????? ???????????? ?????? ????????? ?????????? ??????????? (???. 3); ???, ?????????? ???? ? ???????????? ??????? ???????? ? ??????? ????
?????? (1 %), ????? ??? ?? ???????? ???? ? ??
????????? ?????? ????? ???? ? ?????? ????????? ??????????? ? ?????????? ???????? 8
? 4 % ??????????????. ??????????? ??????
?????????? ? ???????????? ??????? ???????? ???? ???????????? ?????????? ? ??????
????????? ??????? ??????? ? ??????? ????????????. ????????? ?????? ????????, ??????????????? ??????? ??????????? ???????????? (Betula nana) ?, ?????????????, ????????
????? ? ?????? ????????? ?????? ?????? ????????????. ?????? ???????????? ????? ?
?????? ????????? ?? ???? ??????????? ???
?????? ????? ???? ? ???????????? (48-62 %).
??????? ??????? ? ?????? ????????? ?????
??????? ????? ???? (?? 10 % ?? ????????? ?????? ?? 36 % ?? ???????? ????) ? ??????????? ??????? ? ???????? ???? (21-27 %).
?????? ????????? ???????????? ?? ????????????????? ?????? ???????? (???????
# 43 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
??????? 1. ????????????????? ?????? ???????? ??? ?? 1999-2007 ??. ?? ??????????? ????????
???????????, ??
??????? ???????? ???
??????? ???
?????? ???
???????? ????
????????? ??????
22 (7-38)
10 (-1-19)
5 (-1-11)
6 (2-9)
??????????. ????????????? ???????? ???????? ??????? ???? ???? ???????????, ? ??????? ???????
??????????? ? ???????????? ??????? ????????.
???. 2. ????????? ??? ?????? ???????? ???. 1 ? ??????? ???, 2 ? ?????? ???, 3 ? ???????? ????, 4 ?
????????? ??????
???. 3. ????????? ??????? ????????? ???????????? ? ????????? ??? (??????? ?? 1999-2007 ??.)
????? ???? ? ????, ?????? ????????????) ? ??????????????????? (???????????????? ????? ?????? ? ???? ???????, ?????????????
?????? ??????????? ? ????? ????? ????????).
?????????? ????????????????? ?????????
????? ??????? ?? ????????? ??????????????,
?? ?????????? ????? ?????????????? ? ??
???? ??? ???????????? ????? ?????? ??????? ????? ???? (52-79 %). ????? ??????????
?????? ?????????? ??????? ????????????
(22-29 %), ?? ??????????? ???????? ????. ??
???????? ???? ? ????????? ?????? ? ???????????? ??????? ???? ? ???????????? ???????
????????????? ? ?? ????? ? ?????? ????????????????? ????????? (17 ? 22 % ??????????????) (????. 2).
??????? ????????, ??? ????????????
????? ? ????? ?????? ???????? ?? ??????????? ??? ?????? ?????????, ??????? ?????????? ?? 63 % ?? ?????? ?????? ???????? ??
# 44 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
??????? 2. ?????? ????????????? ????????, ??????? ???????? ?? 1999-2007 ??., ?/?2 (??????? ???????? ±
??. ????.)
?????
9±7
4±6
????????
????
80±18
?????? ????????????
119±35
107±49
18±13
112±31
???
285±78
370±88
369±75
220±72
????? ????????????????? ?????????
413±63
481±97
466±82
423±60
?????? ????????????
231±84
154±44
12±6
368±84
????? ???? ? ????????????
669±276
584±313
556±122
1411±514
??????? ??? ?????? ???
?????????
??????
91±48
????? ??????????????????? ?????????
900±296
738±394
563±137
1779±633
????? ?????????
1313±287
1220±362
1035±157
2202±516
????
189±114
164±75
232±101
184±121
??????? ????
4075±724
3895±639
2460±536
3633±363
????? ?????????
4239±710
4059±606
2692±527
3794±433
????? ????????
5577±916
5279±692
3727±584
5996±1082
????????? ?????? ?? 77 % ?? ?????? ????
????????????? ?????? (????. 2). ????????
????? ????????? (????? 90 %) ????????????
??????? ??????. ?????????? ???????? ?????
??????? ?? ????????? ???????? ??????? ?????????? ???????? ??????. ???, ???????????
?????? ??????????? ?? ???????? ????, ???
??????? ???? ???????? ?????? ?????????.
?????? ???????? ????? ?? ???????, ??????
???? ? ????????? ?????? ????? ???????????? ????????. ????????? ??????? ???? ????????????? ?????? ??????????? ?????? ?????????, ?? ? ?????????? ??? ?????? ???????
? ???????? ????????? ?? ????? ????????
? ???????????? ?????, ??????? ??????????
????????? ????????? ??????????????.
?????? ????? ?? ???????, ?????? ???? ?
????????? ?????? ??????????? ?? ?????????? (????. 2); ??-????????, ??? ??????? ?? ??????? ??????????????, ??? ??? ??? ??????????
??????????????? ???????? ?????? ?????????
??????????????? ?????, ??????? ? ????????
???????? ??????????? ?????, ??????????
??????????? ?????????????? ????????????? ?
????? ?? ?? ?????. ????? ??? ?? ???????? ????
??????????? ??????????? ???????????, ?
??????????? ???????? ???? ?????????? ?????
50 %, ??? ???? ???????? ??????????? ?????.
?????? ????????? ????????? ?? ???????????? ?????? ?????????? ?? 480 ?? 650 ?/?2,
??? ? ??????? ????????????? ??????, ?????????? ??? ????????? ???????????? ?????.
?????? ?????? ? ???? 0-20 ?? ?? ???????????
?????? ????? ??????????? ? ???????? ??????
??? ????? ???? ? ????????? ??????? (Moore et
al., 2002). ???? ?????? ? ????? ????????????? ???????, ?????????? ????? ? ??? ???????????????? ?????. ???????? ????????????
?????? ?????? ????? ????????? ?? ???????
????????? ????? ?????????? ?? 1900 ?? 17100
?/?2 ? ??????????? ?? ???? ????? (?????????,
1967; ????????? ? ??., 1995; ?????, 2003; ????????, ??????, 1976). ?? ?????? ?.?. ????????? ? ?????. (1994), ????? ????? ????????????? ???????? ? ??????? ???? ????????????
????? ???????? ?????? ???????? ?? 2300 ??
3100 ?/?2 ? ??????????? ?? ????????????.
Moore et al., (2002) ?????????, ??? ?????
????????? ???????? ?????????? ????????
??????????? ? ??????????? ?????? ???????? ??? ?? ?????? Mer Bleue (??????): ???????? = 10,44*????106, r2 = 0,75. ????????
# 45 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
??????? ?????????? ???????? ?? ????? ?????
??????? ??????????? ?? ?????? ???????? ???:
???????? ??????? = 2,78??? + 77, r2 = 0,53, ?
???????? ????????? ???? ?????? ?? ????? ??????????? ????? ? ??????? ???????? ???. ?
????? ???????????? ???????? ???????????
????????? ?????????? ???????? ?? ?????? ???????? ???: ????????? = -18,78??? +
910,68, r2 = 0,60.
?????? ??????? ???????? ??????? (??????????? ? ?????????? ???????) ?? ??????
???????? ????????, ??? ?????????? ???????
? ??????????? ??????? ?? ?????? ???????????? ???? ? ?????????? ????????, ??? ??? ???
?? ??????????? ???????? ???????? ????????
???????? ????????????? ???????, ???????, ?
???? ???????, ???????????? ??????? ???????? ???.
????? ???????, ?????? ?????? ???????
???????? ???? ??????????? ?????? ???????,
??????????? ?????????????? ????????????? ??????? ???????? ?? ?????? ????????????? ? ??????????? ?? ??????? ?????????????
???????.
???????? ??????? ????- ? ?????????
???????????? ???????? ????????? ????????, ??? ???????????? ????????? ????????? ??????????????? ????????? ??????, ?
??????????? ? ???????? ????. ??????? ????????? ?? ???????? ???? ????????? ????????? ? ???????, ?? ?????? ???? ? ? ????????,
?? ????????? ?????? ? ? ????. ? ???????? ??????????? ???????? ???????? ?????????, ??
??????????? ??????? ????, ???????? ?????
?????????? ?? ????????? ??????, ??? ??? ?????????????? ????? ???????????? ? ????????
???????????? ????????????? ? ???????.
???? ???? ????????? ??????????? ???????? ??????? ????????? ????, ???????????? ?
???? (???.4). ???? ???? ? ???????? ?????????
?????????? ?? 1 % ?? ??????? ? ?????? ????
?? 12 % ?? ???????? ????. ???? ???????????? ??????????? ???? ? ????????? 55 ? 63 %
?? ??????? ???? ? ????????? ??????, ?????
???????? ???????????? ? ????? ??????? ????????? ?? ???????? ???? ????? ????? 9 %.
????? ?????? ???? ???? ? ???????? ?????????, ??????? ?????????? ?? 33 % ?? ?????????
?????? ?? 85 % ?? ???????? ????. ????? ???????, ?? ???????????? ?????? ???????? ???????? ????????? ????????????, ???????
???????, ????????? ??????? ?????? ????, ?
?? ????????? ? ????????? ????????????.
???????? ????????? ? ????????? ????
?????????? ? ??????????? ?? ???????? ??????? ???? ????????????. ????????, ????? ????????? ?????? ????? ? ??? ? ??????????
????? ??????????? ???????? ?????????????
????? ????????, ????????? ???? ??????????
???????????, ????????? ? ??????? ??????????? ???????, ????, ????????, ??????????
????????. ????????? ??????? ????????? ?
???????? ???????????? ????????? ?????????,
?? ???? ??? ???????? ?????? ??????????????
?????? ????? ???????????????? ????????????? (??????????????? ??????? ????????????????????????), ?????? ???????? ??????????,
?????? ???????, ??????????????? ????????????? ? ?????????? ??????? ????????? ??? ?
????????? ????, ??? ? ?? ???? ??????.
?????????????? ?????? ???????, ???
???????? ????????? ?? ???????????? ???
?? ??????? ?? ???????? ???????, ? ?? ?????
??? ???????? ????????? ?????????? ??????
??????? ?? ??????????? ??????? (r=0,58) ?
?????????? ??????? (r=0,61). ????? ?????????
??????, ??????????? ??? ????????? ???????????? ???????? ????????? (????, ????,
????????????), ???????, ??? ??????? ??????????? ?????????????? ?? ???????? ???? ?????
??????? ?? ??????????? ??????? (r=0,59), ? ??
?????? ???? ? ?? ?????????? ??????? (r=0,45) ?
??? (r=0,53). ??????????? ??????? ?????????
# 46 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
???. 4. ???????? ???????? ????????? ? ????? ???????????? ? ????????? ??? (??????? ?? 1999-2007 ??.)
??????? ?? ??????? ???? ?? ????????? ?????? (r=0,71), ? ????? ?? ??????? ???? ? ???????????? ?????????? ?????? ?????? ??????????
??????? (r=0,51 ? r=0,47 ??????????????).
????? ???? ???????, ??? ?????? ????
? ???????? ????????? ?????? ?????? ???????
???????? ???, ?????? ?????????? ??????????????? ??????? ????????, ??? ??????? ???
??????????? ? ???????? ?? ????????? ??????,
?????? ?????????? ??????? ?????????? ???
???????? ???????????? (r=-0,70). ????????
??????? ??? ?? ??????? ???? ? ?? ???????
???? (r =0,69).
????????? ??????? ????? ?? ??????? ?????????????? ??????? ???????? ?????????
????????: ?????? ? ?????? ???? ????? ???????????? ????????, ????? ??? ?????? ??????
??????????????? ??????? ?????????? ?????
(???. 4). ???????? ???????? ????? ?????????? ???, ??? ?????? ???????????? ? ?????,
???????? ? ??????? ??????, ?? ???? ?? ???????? ???????????, ????? ????, ? ??? ??????????? ????? ????????, ????????? ? ??????
?????? ??? ????????? ????? ? ?????.
??????????? ?????? ???????, ??? ?????????? ????? ??????? ?? ???????? ?????????
???? ? ???????????? (????. 3). ?? ???? ??????????? ????????????? (?? ??????????? ???????? ????) ??????????? ??????? ?????????????
????? ????? ????????? ????????? ? ??????.
? ??????? ?????????????? ?????? ????????
????? ? ????????? ????????? ? ???????????,
?? ???? ??? ???????????? ??????? ?????????
???????? ??????????? ?????? ?????????.
?? ?????????? ???????? ????????? ???? ? ???????????? ? ????????? ????????? ?? ???????
???? 33 %, ?????? ? 43 % ? ????? 90 % ?? ???????? ????. ??? ?????????? ?????? ??? ???????? ??????? ???? ? ?????????? ???? 18 %.
?????? ????????? ?????????
?????? ????????? ????????? ??????
??????????? ?????????? ???????? ? ???? ????????????? ????????. ?????????? ???????????? ???????, ???, ???????? ?? ????????????
???????? ? ??????? ????????????? ???????, ?
??????? ???????????? ???????? ?????????? ????? ??????? ???????? NPP ???????-
# 47 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
??????? 3. ??????????? ???????? ????????? ?? ???????? ?????????
???????????
????????? ? ?????
?????????
??????? ???
0,43
?????? ???
???????? ????
????????? ??????
??????????? ??????????
?????????
????????? ? ?????????
? ????????? ????
????????????
-0,73
0,53
-0,88
-0,75
-0,82
-0,95
-0,99
-
-0,86
-0,95
-0,71
?????????????-???????? ????? ?? ???????,
?????? ???? ? ???????? ???? ? 558, 587, 571
?/?2 ? ??? ??????????????. NPP ?? ?????????
?????? ? 1,8 ??? ???? ?? ????????? ? ???????????? ??????? (1056 ?/?2 ? ???) (???. 5).
???????? ????? ? ????????? ?? ??????????? ???????????? ???????? ??? ??????
????? ???? ? ???????????? (47-57 %) ? ????????? ??? (23-31 %). ?? ??????? ? ?????? ????
????? ?????? ???? ???????????? ? 18 ? 17 %
??????????????. ?? ???????? ???? ????? ???????? ????? ????? ??????????? ?????????????? (16 %), ? ???? ???????????? ?????????
?? 7 %. ??????????? ?????????? ???????????? ?????? ????????? ?? ????????? ??????,
??? ???????? ????? ? NPP ?????? ???????????
(31 %), ???? ???? ? ?????? ????? ? ?????????? 26 %. ??????? ????????, ??? ??????? ?????? ?????????? ?????? ??????????????? ???????????? ? ????????? ??? ?????? ?????? ??
?????????? ??????????????? (??????), ??????? ??????? ?????????? ????? 40 %.
???????? ?????????????? ?? ??????????? ???? ??????????????? ?????? ?????????
?? ???? ??? ? 2002 ?. ? ?????????? ? 2003 ?.,
?????????? ?????????? ????????? ??????, ??
??????? ???????? ????????? ??????????? ?
2005-2006 ??. (???. 6).
?????? ??????? ???????? ??????? ??
NPP ??????????? ??????????? ????????,
??? ???????? ?????????? ??????? ???????? ??????? (???) ????????? ??????? ????
(r= -0.84), ???????? ???? (r= -0.53) ? ??????-
???? ?????? (r= -0.61). ?????? ??????? ??????????? ? ??????? ?? NPP ?????????????, ???
????????? ?? ??????? ???? ? ???????? ????
??????? ?? ??????????? ??????? ? ???????
?????????????? ??????? (r= 0,64 ? r= 0,69, ??????????????). ?? ?????? ???? ? ?????????
?????? ??????????? ?????? ??????????????
????, ? ?? ????? ??? ?????????? ??????? ?????
??????? ??????? ?? ?????????????? ???????
???? (r= -0,88) ? ?????????? ?????? (r=-0,61).
?????????????? ?????? ????? ????? NPP ?
??????? ???????? ??? ???????, ??? ??? ????????? ??????? ?? ?????????????? ???????
???? (r= -0,74).
????? ????????? ??????, ???????????
??? ????????? ????, ???? ? ????????????
????????, ??? ??????? ???????? ??? ?????? ?? ????????? ????????? ???????. ???,
????????, ?? ??????? ???? ???????? ????????????? ??????????? ????????? ???? ??
?????? ???????? ??? (r=0,62) ? ?????????????
??? ????????? ???? (r= -0,70). ???????? ??????? ??????????? ??? ??????? ????, ??? ?????
????? ??? ? ?????????? ???? ????? ????????????? ???????? (r= -0,56), ????? ??? ??? ????
??? ????? ???????????? (r= 0,70). ?? ????????
???? ??????? ???????? ??? ????????? ???????
?? ????????? ???? ? ???????????? (r= 0,46 ?
r= 0,75 ??????????????). ??? ?????????? ?????? ??????????? ????? ????? ??????? ????????
??? ? ?????????? ???? (r= -0,65).
??? ????????? ????? ????? ????????
???????? ????, ???????????? ? ???? ? NPP
# 48 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
???.5. ?????? ????????? ????????? ???????????? ? ????????? ??? (??????? ?? 1999-2007 ??.)
???.6. ???????? ?????? ????????? ????????? ???????????? ? ????????? ???
??? ???????? ????????????? ??????, ?????????? ???????? ????????? ? ????. 4.
?????????? ??????????? ??? ????????
???????? ? NPP ????????????? ??????????
????? ??????? ???????? ??? ??????????
? ????????? ????? (Moore, 2002; Backeus,
1990; Grigal et al., 1985; Reader, Stewart, 1972;
Szumigalski, Bayley, 1997; Thormann, Bayley,
1997; Vasander, 1982). ?????????? ??????????? ????????? ????????? ????????? ????????
??????????? NPP, ??????? ?????? ?????????.
?????????????? ?????? ?? ???????
???????? ? NPP ??? ?????????? ????????
??????? ? ?????? (Grigal et al., 1985; Reader,
Stewart, 1972; Szumigalski, Bayley, 1997;
Thormann, Bayley, 1997; Bartsch, Moore, 1985;
Bartsch, Schwinzter, 1994; Dyck, Shay, 1999;
Vitt, 1990; Wallen, 1986; Wallen, Malmer, 1992)
??????????????? ? ??????? ????????? ????????; ??? ??????????? ???, ??? ???????????
???????? ?????????? ??? ???????? ???????????? ??????? ???????????? (?? 10 ?? 50 ????? ?????????? ???????? (??????, ????????,
1977; Moore, 2002), ?????????? ??????? ???????????? ???????????? ??????????, ?????
???????? ? NPP ??????? ??????? ?? ??????????????? ? ?????????????????? ???????. ?????? ????????? ????????? ????? ? ????????
?????? ? ??????????? ?? ???? ?????? ?????
????? ?????????? ? ??????? ???????? ? ??
210 ?? 780 ?/?2/??? (?????????, 1967; ?????,
2003; ????????, ??????, 1976; ??????, ???????, 1975). ??????? ??????? ?? ????????????
??????? ???????? ?????? ????????? ?? 133 ??
464 ?/?2/??? (?????????, 1967, 1985; ??????,
????????, 1977; ?????????, 1967).
# 49 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
??????? 4. ????????????? ????????? ??? ????? ?????? ????????? ????????? ? ????????? ??????
??????? ??????????????
NPP ????
NPP ????????????
NPP ??????
NPP
NPP
NPP
NPP
????????? ?????????
Log10NPP=0.87log10(????????? ????) +0.25
Log10NPP=0.43log10(????????? ????????????)+1.17
Log10NPP=0.74log10(????????? ??????)+0.44
Log10NPP=6.26log10(NPP????)
Log10NPP=0.26log10(NPP ??????) +2.10
Log10NPP=0.62log10 (?????????) +0.85
Log10NPP=0.35log10(???????? ??????) +1.78
??????????
??????????? ???????????? ????????????? ?????????????? ???????????? ?
????????? ???????? ??? ????????, ??? ?????? ???????? ? ??????????? ???????????? ??? ??????????? ????????? ?? ??????? ????????? ??????? (???????? ????),
?????????????????? ???????? ????????
???????????? ?????????, ? ???????????
????? (???????? ????). ?????? ????????
?????????? ?????? ??????????? ? ????????
???????? ???????? ????. ?????? ????????? ?? ????????? ?????? ????? ? ??? ????
???? ?? ????????? ? ????????????. ????????????? ?????????????? ???????????
???????? ????????? ? ????? ???????????? ????? ????? ? ???????????? ??????????? ???????? ????????????. ??????? ???
???????? ????????????? ??????????????
?????? ???????? ????????? ????????????
????????? ????????????? ??????? ? ???? ?
??? ????????? ????? ????????. ?????????
?????????????? ? ??????? ?????? ????????
???????????? ???????? ??????? ?? ???????????????? ?????? ?????, ????????? ???????? ??????????????? ???? ? ??????? ??????????? ?????????.
? ???????? ??????????? ?????????? ??????? ????- ? ????????? ??????? ??????????
n
27
27
27
27
27
27
27
R2
0,96
0,35
0,37
0,99
0,31
0,39
0,39
??????? ?? ???????? ??????? ??????????????,
????????? ?????????????, ?????????? ???????, ??????????? ??????? ? ?????? ????????
???.
???????? NPP ?????? ?????????????
????????????? ?????? ????? ??????? ????????, ????? ??? NPP ?????????? ??????
? 2 ???? ????. ??????? ????? ?????????
?????????????? ??????? ???????? ? ?????. ??????? ??????????? ????? ???????? ???????? ???? ???????????? ? ???? ?
?????? ????????? ?????????? ????????
?????????, ??????? ????????? ?????????
???????? ??????????? NPP ???????? ?????????.
? ????? ?????????? ? ?????????? ???????????? ?????? ?? ??????? ???????? ? ????????? ???????? ????????????? ????????????? ?????????? ????? ?????? ??? ????????
????????? ?? ?????????? ??? ??????, ??? ?
??????, ??? ? ??????. ?????? ???????????
?????????? ????????? ??????? ??????? ???????? ? ???????????????? ??????? ?? ?????????
???????? ????????????? ?????????????? ???????? ?????????, ??????? ??????? ?????????
?????????????? ? ??????? ???????? ??? ????????? ??????? ??? ??? ????????????? ????????????? ? ????? ????????????? ?????????????? ? ?????????? ???????? ? ????????.
?????? ????????? ??? ????????? ?????????? ????????? ???? (08-05-00426a, 0805-92501).
# 50 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
?????? ??????????
????????? ?.?. (1967) ?????????????? ? ????????????? ?????????? ? ??????? ???????
?????? ????????? //???????????? ???????, ?.3, ???. 4:567-589.
????????? ?.?. (1993) ????????????? ?????????????? ????????? ???????? ???????. ?.:
?????, 295 ?.
???????? ?.?., ?????? ?.?. (1976) ?????? ? ???????? ??????? ????????????? ? ???????????????????. ??????????, 59-82.
??????????? ?????? (????????? ???????, ????????? ? ????????????????) /??? ???. ?.?.
???????? (2000). ?????: ????, 136 ?.
?????????? ?.?. (1994) ?????????????????? ??????????? ????? ? ?? ???????????? ?????????????. ?.: ?????, 5-37.
?????? ?.?., ??????? ?.?. (1975) ? ????????????? ?????????????? ???????? ?????,
??????????????????? ? ????????????????????? ????????? //???. ????., ?. 60, ? 9: 13361347.
????? ?. ?., ???????? ?. ?., ???????? ?. ?. (1984) ??????????-?????????????? ??????????? ? ???????? ???????? ????????? ???????. ?., 128 ?.
??????? ?.?., ???????? ?.?. (2007) ????????????????? ??????????? ??????????????? ?
????????????? ????????????? ???????? ? ???????????? ??????????? ???????? ?????? //????????? ??????? ?????????????? ???????? ?????????? «????????? ???????? ?????? ? ????
????????: ??????? ? ?????????», ?. 95-97.
??????? ?.?., ???????? ?.?., ?????????? ?.?. (1994) ?????? ???????? ? ??????????? ?????
//??????? ? ??????????? ????? ? ????? ?????? /??? ???. ?.?. ????????? ? ?.?. ??????. ??????????, ?. 128-139.
???????? ?.?., ??????? ?.?., ????? ?.?., ??????????-???????? ?.?., ????????? ?.?.
(1994) ????????????? ?????????????? ? ????? ????? ?????? ????????? //????????? ????????????? ??????, 3:253-267.
??????? ?.?., ?????????? ?.?., ?????????? ?.?., ???????? ?.?. (2003) ??????????????????????? ??????? «?????????». ????????? ??????? ?????????. ?????, ????, 88 ?.
?????????? ?.?., ????????? ?.?., ????????? ?.?. (1978) ???????? ????????????? ???????? ? ???????? ????????????????. ?.: ?????, 176 ?.
???????? ?.?. (1986) ????????????-?????????????? ???????? ? ??????????? ????????
????? ????????? //????????, N 3:35-41.
????? ?.?. (2001) ???????? ??????? ????????? ? ????????? ????? ???????? ????? //????????? ???????? ?????? ? ???? ????????: ??????? ? ?????????. ???????????,
? 94-96.
????? ?.?. (2003) ???????? ?????????? ??????? ???? ???????? ??????: ????????? ?
?????????: ???????. ???. ? ???? ????. ????, 23 ?.
??????? ?.?., ???????? ?.?., ?????????? ?.?. (2000) ?????? ?????????????? ?????? ???
?????? ???? ???????? ??????. Krylovia, ?.2, 1:38-43.
??????????-???????? ?.?. (2001) ???????? ??????? ? ????????? ?????????????? ?????
????? ????? //????????? ???????? ?????? ? ???? ????????: ??????? ? ?????????. ???????????, ?. 106-107.
# 51 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
??????? ?.?., ????????? ?.?., ??????? ?.?., ??????? ?.?., ??????????-???????? ?.?.,
??????? ?.?., ??????? ?.?., ??????? ?.?. (1993) ??????? ?????? ?? ????? ??? ???????? ?????? //???, ?. 330. ? 3:388-390.
????????? ?.?. (1967) ? ?????????????? ????? ???????? ?????? //???????????? ???????,
67, ?.3,4: 523-532.
????????? ?.?. (1985) ???????? ??????, ?? ????????? ? ????????????? ????????. ?.: ?????, 152 ?.
????????? ?.?. (2001) ??? ?? ????? ? ????????? ?????? //????????? ???????? ?????? ?
???? ????????: ??????? ? ?????????. ???????????, ?. 136-139.
????????? ?.?. (2007) ?????? ????????? ????????? ???????? ? ???????? ????????? //
????????? ????????????? ??????, 5: 763-770.
????????? ?.?., ??????? ?.?., ????????? ?.?., ?????? ?.?., ??????? ?.?., ??????????
?.?. (1998) ?????? ? ?????? ????????????? ???????? ? ?????? ?????? //????????????, 1:51-59.
????????? ?.?., ????? ?.?., ??????????-???????? ?.?. (1995) ????????????? ?????????????? ????? ?????? ?????????. ?????? ?????? ?.?. ??????, ?????, ?. 59-63.
????????? ?.?., ????? ?.?., ??????????-???????? ?.?. (2000) ??????? ???????? ???????? //????????? ????????????? ??????, 5:653-658.
?????? ?.?., ???????? ?.?. (1977) ?????? ? ???????? ?????????? ?????????? ?????????, ???????????: ?????, ????????? ?????????, 219 ?.
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and P in a dry heathland ecosystem. Biogeochemistry 15:175?190
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in subarctic peatlands. Can J Bot 63:1241?1248
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of forested bogs in northern Minnesota. Can J Bot 63:2416?2424
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production and CO2 exchange in an ombrotrophic bog. J Ecol 90:25?36
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Beilman D.W. and Dubinin P. (2004). A high?resolution GIS?based inventory of the west Siberian peat
carbon pool. Global Biogeochem. Cycles, 18: GB3004
# 52 #
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?.?. ??????????. ????????????? ?????????????? ???????????? ? ????????? ??????
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gradient in central Alberta in relation to environmental factors. Ecoscience 4:385?393
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gradient in southern boreal Alberta, Canada. Ecoscience 4: 374?383
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comparison between a North American and a Scandinavian peat bog. Acta Societatis Botanicorum
Pol 61:75?87
Biological Productivity of Oligotrophic and Eutrophic Mires
in the Southern Taiga of Western Siberia
Evgeniya A. Golovatskaya
Institute of Monitoring of Climatic and Ecological Systems,
Siberian Branch of the Russian Academy of Science
10/3 Akademichesky, Tomsk, 634055 Russia
The results of long-term (1999-2007) investigations of Phytomass (above and below-ground living
biomass), dynamics of phytomass and mortmass and net primary productivity of oligotrophic and
eutrophic mire in southern taiga in Western Siberia (Tomskaya oblast) are presented. The storages of
phytomass on different biogeocenoses of oligotrophic bog have similar values. Reserves of phytomass
of eutrophic peatland are 2 times higher in comparision to oligotrophic bogs. Net primary production
of different oligotrophic biogeocenosis has close values. NPP at eutrophic peatland is 1.7 times higher
in comparision to oligotrophic bogs. The major part of production is provided by mosses and plant
roots. The qualitative composition of phytomass depends on vegetation of the studied phytocenosis.
The dynamics of phytomass and NPP depends on precipitation, air temperature, and water table level.
The correlation between phytomass of herbs, shrubs and mosses and NPP peatland ecosystem are
revealed.
Key words: Peatland, net primary production, dynamics of phytomass.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 54-79
~~~
??? 634.0.416.1.: 582.475.4
?????????????? ? ???????? ??????? ? 5-??????? ?????
???????? ? ????????? ????:
???????????????? ? ????????????? ?????????????
?.?. ??????????*, ?.?. ?????
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???
634055 ??????, ?????, ??. ????????????? 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
???????? ????????????? ?????? ????????? ? ???????? ??????????? ? ??????? ??????? ?
5-??????? ????? ???????? ? ????????? ????. ??? ?????? ????????????????? ??????????????
??????? ? ????? ??????????? ?????????? ? ???????????? ??????, ??? ????? ?????????? ?
????? ????????????? ???? ?????? ? ???????. ? ???? ????????? ????????? ???? ????? ???????
?? ?????? ??????: (1) ????????????-?????????????? P. pumila, (2) ?????????-????????????
P. sibirica ? P. cembra, (3) ??????????? ?? ????????? ????? P. koraiensis ? P. parviflora,
(4) ?????????????? P. armandii. ???????? ????? ?????? ?? ????????? ??????????? ??????
?? ???? ??????? ? ???????? ? ?? ???????????? ??????. ?? ????????????????? ???????????
??????????? ? ??????? ?? ?????????? ?????? ????????????? ??????? ???? ????????
??? ???? ?????: P. sibirica + P. cembra; P. koraiensis + P. armandii; P. parviflora + P. pumila.
??? ????????? ????????? ???????? ? ?????? ???????????????? ???????? ?????????????
??????????? ????????? Cembrae. ?????????????? ??????? ?????? ??????? ? 12 ???????????
?????????????? ???????? ???? ?????????? ????? Pinus ?? ?????? Cembrae (P. sibirica, P. pumila
? P. koraiensis). ? ???? ???? ????? ? ??????? ????? ????? ? ??????? ?????????????? ???????
??????????? ???? (????????) ? ???? (????????) ???????????? ?????. ? ??????? ????? ?????
? ??????? ?????????????? ??????? ??????????? ??? ????? ? ??? ???????????? ??????. ?
?????? ?????? ????????????? ????????? ?????, ?? ??????? ??? ??????? ????? ???????????
?????? ?????. ????? ??????????? ???????? ?????, ?? ??????? ? ?????? ???????? ??????????
?????????????? ??????? ??????????? ???????? ?????. ???????????, ????????? ????? ?
??????????? ??????? ?? ?????? ?? ???????????? ???????. ??????? ????? ?????????????
? ?????????? ????? ????????? ??????. ???????? ? ????????? ???????? ????? ?????? ?
????????? ?? ????????? ????????? ??????.
???????? ?????: ????????? ? ???????? ??????????? ? ????????????? ???????, 5-???????
????? ???????? ? ????????? ????, ????????, ?????????.
??? ??? ?????????? ???? Pinus ?? ??????? Haploxilon (P. sibirica Du Tour, P. pumila
(Pall.) Regel. ? P. koraiensis Sieb. & Zucc.) ??
?????? J. R. Shaw (1914) ? R. Pilger (1926)
*
1
?????? ? ?????????-?????????? P. cembra L.
? ?????????????????? P. albicaulis Engelm.
???????????? ? ?????? Cembrae. ?????????????? ???? ?????? ????? ???????? ?????????-
Corresponding author E-mail address: gorosh@imces.ru
© Siberian Federal University. All rights reserved
# 54 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
?? ?????? ?????????? ????????, ?????? ??
??? ??? ??????? ? ?????????? ???? ?????
??????. ???????, ?????? P. sibirica ? ?????
??????? ? ???? P. cembra ???????? ???????
«????????» ?? ? ?????????? ?????? ?? ?????? Strobi. ? ????????? ???? ?????? ????? ???
???????? ????????????. ???, ? ????????????
???????? ?????? W. B. Critchfield (1986) «?????» P. pumila ?? ????????? P. parviflora Sieb.
et Zucc., ? P. albicaulis ? ?? ?????????????
P. flexilis James.
?????????? «???????????????» ????? ????? ??????????? ?????? Cembrae ???
????? ???????? ????????? ?????????. ???,
???????????????? ???????? ???????????, ??? ???? ?? ???????? ???? P. pumila ?
P. koraiensis ????? ?????? ? ??????????? ?????????? ?? P. sibirica (??????????, 1974),
?? ?? ????????? ????????? ?????, ????????,
P. sibirica ? P. pumila ????????? ? ??????, ?
P. koraiensis ? ? ??????? ???? (??????????,
??????????, 1974). ?? ????? ?????????????
? ??????????, ?????????? ???????????? ???????? ????????????? ???????????. ??????
???????????? ????????????? ??????? ??????? ??????????? ?????? ???????? P. pumila
? P. koraiensis ? ?????????? ???????????? ??
??? ????????? P. sibirica (?????????? ? ??.,
1990; Shurkhal et al., 1992; Goncharenko et al.,
1992). ???????????? ???????????? ??? ???????????? ???? ??????????????? ??????? ?
???????????? ??????????? P. pumila ?? ?????
??????? ? ???? ????????? ???? ?????? ?????
(Krutovskii et al., 1994).
?????????? ??????, ???????? ?????????, ????????????? ?????? ? ???????
??? ??? ???? ???????? ?? ???????? ??????? ? ??????? ???? «??????????» ????????, ?????????????? ??????? ? ??????????
???????? ??????? ????? ??????????. ? ????
????????? ?? ??? ??????? ?????????? ???????? ?????????, ??????????????? ????????-
??? ? ?????????????? ???????. ??? ????????
????????? ?? ?????? ??? ????????? ????????? ??????????????? ????? ???????????
??????, ?? ? ??? ??????? ???????????? ????? ????????????? ??????????. ??????????
???? Pinus ?? ?????? Cembrae ? ?????????
?????? ??? ???????????? ?????? ????. ???????? ?? ????? ??????? ??????? ??? ???????????? ??? ??? ???????? ???????? ????
????????? ?????????????? ?????????? ?????:
?????????????-????????????? (P. pumila),
?????????-???????????? (P. sibirica) ? ??????????? (P. koraiensis). ??? ???? ?????? ???
???????? ???????? ????????, ? ??? ?????? ? ?
????? ?? ???? ????????? ??????? ????????? ?????? ????????. ??? ??? ???? ????????
???????? ?????? ? ?? ??????? ?? ????? ?????? ???? ???????????? ????????????? ???????. ?? ???????? ? ??????? ? ????????? ??????
????? ??????. ?????????? ?? ???????????
????????????, ?????? ?????????, ??? ????????????? ? ???????????? ??????????. ? ????
????? ???????? ????????????? ?????? ??????????? ??????? ??? ?????????? ????????
????????? ???????? ? ???????????? ???????? ?????, ????????????? ?? ????????????
? ??????????????.
Pinus, ? ??????? ?? ????????? ?????
Pinaceae, ????? ????? ????????????? ????????? ?????? (Lanner, 1976). ? ?????????????
????? ??????? ????????????? ?????? ??????
???????????? ??????????? (??? ????????
????????), ? ? ????????? ????? ?????? ? ??????????? ???????????. ? ?????????? ????
???????? ?????? ????????????? ? ????????????????, ??? ???????, ? ????? ??????????????????: ?????????????, ???????????
??????, ??????????? ?????????? ??????,
?????. ???????? ????? ????? ???????? ??
?????? (???????????????) ??? ?? ??????????
(???????????????) ???????????? ???????.
???????? ???????????? ?????? ???? ?????:
# 55 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
???????? ? ??????. ???????? ?????? ??????????? ?? ???????? ?????, ???????????? ?
?????????????? ????. ?????? ?????? ??????????? ?? ????????? (??????) ????? ???
??????? ??????? ?????.
????????? ????????? ?????? ??????????? ??????? ?? ?????????????? ??? ?????????? Pinus. ?????? ??? Shaw (1914) ???????,
??? ???? ???????, ??????????, ??????????????, ?????? ?????? Diploxylon ??????????
?? ??????? Haploxylon ??????? ????? ?????????? ???????????? ?????????? ? ???????????????. C.C. Doak (1935) ?? ????????
????????? ???????, ??? ????- ? ??????????????? ???? ????????? ????? ???????????
? ????? ????????. ?????? ?????????????
????????? ????????? ????????? ?????? ???????? ??????? ?????, ??? ???????? ?????
????????? ???-???? ???? (Stone, Stone, 1943).
???? Haploxylon ?????? ????? ?????? ????
???????? ?????, ? ???? Diploxylon ? ?? ??????? (P. rigida Mill.).
???? Pinus ?????????????? ?? ?????
????????? ????????? ?? ??????????????
?? ??????????? ?????. ???? ??????????????? ????? ? ??????????? ????????? ?????
???????????? ??????? ?????????? ? ??????
?? ??. ??? ??????????? ???? ???????? ????????????????. ????? ???????????? ??
??? ???????????? ??????? ????????? 6-7 ??.,
????????, ? P. caribaea Morelet (Chudnoff,
Geary, 1973). ??????????? ???????????????
????? ???????????? ????????? ????. ??? P.
sylvestris L. (Henry, 1846 ? ??.), P. monticola
Dougl. (Owens, Molder, 1977), P. resinosa
Ait. (Sucoff, 1971), P. lambertiana Dougl. ? P.
ponderosa Laws. (Sacher, 1954). ? ?? ?? ?????,
???? ????? ????? ?????????? ????? ??????? ???? ???????????????: P. contorta Dougl.
(Van den Berg, Lanner,1971; Owens, Molder,
1975), P. banksiana Lamb. (Curtis, Popham,
1972), P. strobus L. (Owston, 1968).
?????? ? ?????????????? ????????????
??????? ?????? ?? ????????????? ?????????????? ????????? ????????? ??????. ? ?????
?? ???? ????????? ? ????? ??? ?????????
????? ????????, ??? ????????????? ??????
???????????? ???????, ??????????? ????????? ????? ???????? ??????? ? ??????
??????. ??????????? ????????? ?????????
?????? ? ?? ??????? ?????????????, ? ?? ??????????????? ?????????????? ?????? ????
??????? ?????? ????????????? ??? ???????????? ????????????? ????? ? ????? ?? ????????? ? ???????? ??????????? ???????.
????????? ???????????? ????? ???????? ????????? ???? ???????????? ??? ??????? ????????, ???????? ????? ?????. ?????
???, ????????, ???????????? ???????? ?????
????? ???????????? ?????? ?????? ??????????. ? ???????????????? ?????? ? ??????????
? ??????????? ???????? ???? ??????????????
???????????, ??? ?? ?????, ??? ????????????? ????????? ???????? ???????? ?????? ?????????? ? ?????????? ????????? ????????
(?????????, 1987 ? ??.). ? ?????????? Pinus ??
?????????? ?????? ?????????? ??? ??????????????? ????????? ??? ???? ???????????:
????????????? (?????????? ? ??????????),
?????????? (????? ? ?????????? ????????
????????) ? ?????? (??????????? ? ?????????????). ???????? ? ????????? ???? ?????
???????????? ????????????: ??? ?????????
??????????? ???? ?? ??????? ? ????? ????????? (Mirov, 1967 ? ??.). ??? ???????????????
???????? ????????????? ?????????? ???????
??????. ??????? ????????????, ??? ?? ????????????? (??????????) ????? ???????????
?????? ?????????? ????????, ????????????
????????????? (?????????????) ????????,
????????, ?????????? ? ??????? ?????.
?? ????? ?????? Cembrae ?????????? ?????? ???????????? ?????? ?????? ? P. sibirica
(??????, 1971; ?????????, 1972). ???????? ??-
# 56 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
??????? ??????????? ???????????? ? ??????
???????? ? ??????? ? ???????? ?????? ?????
?????? ??????????? ?????? ???? (????????)
?????, ? ? ??????? ?????, ??? ???????, ???
(???????? ? ??????), ??????? ?????? ??????????? ?? ????????? ? ???????? (??????????, 1994). ? ???? ?????? ?????????? ?????
Cembrae ????????? ? ???????? ??????? ??????????? ?????????? ?? ???????. ?? P. pumila
??????? ?????? ????? ????? ???????? ? ?????? ????????? ? ??????? ???????????? ??
???????? ?????? ???????? ????? ????????
??????? (???????????, 1960; Hanawa, Sasaki,
1970). ???????? ? ???????? ?????? ? ??????
???????? P. koraiensis ?????? ?? ?????????.
???? ????????? ?????? ? ??????? ?????????
? ???????? ??????????? ?????? ? 5-???????
????? ???????? ? ????????? ????, ? ?????
???????? ????????????? ?????? ????????????
? ?????????????? ??????? ? 12 ???????????
????????????? ???????? ???? ?????????? ????? Pinus ?? ?????? Cembrae.
???????? ? ????????
??? ???????????? ??????????? ?????? ?
??????? ?? ???????????? ????? ????????????? ????? 5-??????? ?????:
(1) ???? ???????????, P. cembra L., ??
??????? ????? ??????? ????? ???? ? ??????;
(2) ???? ?????????, P. sibirica Du Tour, ???????????????? ? ????????-???????????????
??????? ??????? ???? ? ? ??????? ????? ???
????? ??????;
(3) ???????? ???????, P. pumila (Pall.)
Regel, ??????????????? ?????????????? ?
????????????? ??? ? ???????? ??????? ?
???????? ? ????????? ???? ?? ??????? ??
??????? ? ?? ???????? ???? ?? ???????????
????? ?. ?????;
(4) ???? ?????????, P. koraiensis Sieb.
& Zucc., ??????????? ??? ?? ??????????,
????????? ? ????????;
(5) ????? ??????????????, P. parviflora
Sieb. & Zucc, ???????? ???????, ???????????
?? ?. ???? ?? ?. ???????? ? ????? ??????? ?
??????-???????????????? ?????;
(6) ????? ???????, P. armandii Franchet,
??? ?? ???????????? ? ?????? ?????,
????????????????
?
??????
?????????????????????? ????? ???????????????
? ???????????? ????? ?? ??????? ?? 1500 ??
3500 ? ??? ??????? ????.
P. cembra, P. sibirica, P. pumila ? P.
koraiensis ??????????? ? ????????? Cembrae,
? P. parviflora ? P. armandii ? ? ?????????
Strobi (Price et al., 1998). ?????? P. cembra ???????? ?? ????????? (???????? ????, ?????
? ?????????, ???????????); ????????????? ? ???????? ?????, ???????? ??????????,
?????? 1800 ? ??? ??????? ???? (????????
???????? ???? ????????? ?????????, ? ??????? ????????????? P. cembra). ?????? P.
sibirica ??????? ? ?????-??????????? ????????????? ??????????: ???-?????? ???????????????? ???????, ????-?????? ??????????, ??????? ????? ?????, 50-70 ?? ?? ?????
??????? ??????. ?????? P. pumila ???????
? ??????????????? ????? ??? ??????: ???????? ???????, ??????-??????? ??., 900 ? ???
??????? ???? (?????? ????? ???????????
????? ????????? ?????). ?????? P. koraiensis
??????? ? ??????-????????? ?????, ????????? ??????????, ???????? ????? ??. ??????????, ?????? 700 ? ??? ??????? ???? (??????? ????? ????? ???????-????????????????
?????). ?????? P. parviflora ??????????? ?
????? «??????» (?. ???????); ????????????? ? ???????? ?????????? ??. ????? ? ?????? ?????, ?????? ??? ??????? ???? 1900 ?
(??????? ????? ??????-?????????????????
? ???????? ??????, ??????? ????? ????????
????? ????????? ?????). ?????? P. armandii
??????? ? ????? ???????? ???????? ?? ??????: ??????????? ?????, ????????? ??????,
# 57 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
1992; 1994). ??????? ? ????????? ??????
???????? ?????? ?????? ?? ???????? ? ???-
??. ???????, ?. ??????????, ?????? 2600 ?
??? ??????? ???? (?????? ????? ????? ??????? ?????; ??????? ????? ????????? ?????????, ? ??????? ????????????? ???? ???). ? ???
???????, ????? ?? ???? ???????? ?????? (P.
armandii, P. sibirica, P. pumila ? P. koraiensis),
????????????????? ???????? ???? ?? ?????
????. ?????? P. cembra ? P. parviflora ????
????????????????? ????? (? ?????? ????????).
??? ?????? ???????????????? ? ?????????? ???????? (6 ??????? ??? ???????????
?? 0 ?? +5°?). ????? ??????????? ?? ??????
?????? ??? ? ???????? ????? ?? ??????? ?????????? «????» ????????? ??????????? ????????????? ? ????????????? ?????? ?? ???
(30 ?? ? ??? ?? ??????, ???-?????? ???????????????? ???????, ??????? ?? ???????
????????? ???????? ??????. ????? ????????
P. pumila ????????????? ??? ??? ???????? ????? ??????: ??????????????, ?????????????
? ???????????????, ?????? ????????? ???????????? ???????? ???????? ?? ??????????
?? ???????????. ??? ??????? P. koraiensis
?????? ?? ?????????????: ??????? ????? ?
??????-????? ??????. ???????? ???????? ?
???????? ??? ??????? ???? ????????????
???????????. ??????? ???????? ??? ?????? ?
?????? ???????, ?? ?? ???? ??????? ?????????????? ?????? ?????, ?????????? ? ???????????? ???? ?????????? ????????? ????????
??? ????? ? ?? ??????? ?????????? ????????? ???????? (?? ???????? ???? G1 ?? ???????? ???? G3 ?? ??????????? ????? ??????????
????). ????? ?????-????? ??????, ??????????.
?????? ???? ?????????? ??????? ??????,
??????? ??????????? ??? ??????????? ??
? ???????? ????????? ????????? (????? ??
???? ?????????????, ?????? ????????). ??????? ??????????? ?? ?????????, ?????????, ?????? ?? ???????? ? ??????????, ?????????? ? ?????? ??????? ?? ???????????.
? ??????? ???? ????????????? ????????
? ?????? ??? ?? ????? ???????? ????????? ?
10-??????? ?????????? ?? ???????????? ?
?????? 25 ??????? ??????? ????. ?? ????????? ??????? ?????? ??????? ?????????????:
???????? ????????? ? ???? ????? ???????????????, ?????????? ?? ??? ? ????????????
?????, ? ????? ??????????? ????????? ????????? (???????? ??? ??????) ? ????? ??????????? ????? ???????.
???????? ?? ?????? ??????? ??? ??????
? ???????????? ??????????? ???? ??????????
????? (????. 1). ????????? ? ???????? ???????
P. sibirica ? ???? ?????????????? ????????
(?????????? ????? ? ????? ??????? ?????)
???? ???????? ??????? ????? (??????????,
?????????). ??????? ???????? ? ???? ???????? ???????? ??? ??????? ?? ?????????? ???????? ? ???????? ????????????????? ???????
(?? ?????????????? ? ?? ??????????).
? P. sibirica ??????? ?????? ????????????? ? ??????? ????? ????? ? ??????????????? ???????? ??????? ??????, ? ???????
?????????? ? ??????? ????? ????? ? ?????????? ?????? ?????? (????????, ??????????,
1991). ??????????????? ??????????????
???? ???????????, ??? ? ?????? ???????? ???
??????? ?????????????? ????????????? ????????? ?? ????????????? ?? ???????? ?????
????????, ? ?????????? ?????????? ? ????
????????? ??????? ? ??????? ?????? (??????????, 1994). ??????? ? ????????? ?????? ?????????????? ??? ???? ????????: ?????
?? ???????? (1) ? ???????? (2) ?????? ?????
? ???????????? ?????????? ?? ??????????
????????? 12-15 ??? ?????????????? ???????.
? ??????? ???????? ??????? ?? 5 ??????? ?
5 ??????? ??????. ?? ??? ??????????? ?????????????? 10 ????????? ???????? ???????: ????? ???????????? ???????, ????? ???
# 58 #
???
Pinus sibirica
Pinus pumila
50°45?
?.?.
56°55?
?.?.
6-7. ????
8. ??????
54°00?
?.?.
5. ??????
136°25?
?.?.
121°5?
?.?.
119°55?
?.?.
103°40?
?.?.
51°30?
?.?.
3-4.
????????
77°41?
?.?.
109°05?
?.?
64°40?
?.?.
1. ?????????
2. ??????? 15°15?
?????
?.?.
??????
??????
???????
??? ?????????????,
??????
600 ????? ????? ??.
??????, ?????? ?.
???????
1600
????????????? ????
?????????
?????? ????? ?????
???????????? ?????
1100 ??????-??????
?????????? ??????,
???????? ?????
600 ????? ?????
????????? ??????,
?????? ???????
??????? ?. ??????
800 ???????? ??????????, ?????? ?.
????????, ??????
(40°) ???????? ?????
1300 ???????? ?????
????????? ??????,
?????? ?. ?????????
60 ??????? ???????????
??? ????? ?
????????
??????
???
???????
????, ?
??????? ????? ?????
?????????????
?????
?????? ????? ?????
?????????????
?????
?????? ????? ?????
???????????? ?????
??????? ????? ?????
???????????? ?????
???????? ?????
???????? ???????
?????
????????? ????,
???????? ????
?????????????? ????????? ? ??????
??????? 1. ?????????????? ???????? ????????????
1600
600
1000
1500
1500
1400
800
6?
3?
1?
-
10?
6?
4?
8?
1?
1?
10?
4?
4?
1?
1?
IV
-
V
IV
IV
III
V
?????
?????
?????????? ?????? ???????? 10°?
????
????????????
? ????????? ??
????????? ????????
?????????????
?????????????
? ????????? ??
????????? ????????
?????????????
? ????????? ??
????????? ????????
???????????????
? ????????? ??
????????? ????????
????????????
?????????????????????????
??? ????
?????????????? ??????????
180
4
0,5
1,5
200
150
3
2
150
20
120
23
14
200
240
170
???????, ??????,
???
?
-
-
-
-
-
26
36
24
???????
??????,
??
??????????????
????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Pinus koraiensis
Pinus pumila
139°13?
?.?.
137°10?
?.?.
131°35?
?.?.
51°17?
?.?.
11. ?????? 50°30?
?.?.
12. ??????- 49°05?
?????
?.?.
9-10.
?????????????
??????????? ????. 1.
???????????
??????????????????????????? ????
??????????????????????????? ????
?????? ????? ?????
???????????? ?????
500
???-?????? ??.
????? ??????,
??????? ?????????
?????
??????? ? ????? ?.
????
????????? ??. ????,
??????? ????????
?????
200
250
??????
????????????
??????? ?. ????
100
2000
1900
1800
1800
6?
4?
6?
4?
8?
2?
10?
II
III
IV
V?
?????????????
?????????????
?????????????
????????????????????? ?
????????? ??
????????? ????????
240
150
200
200
30
17
22
4
48
20
41
-
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
? ???? ??????? ?? ???, ? ????? ????? ? ???????????? ???????? ???????: ??????????????, ?????, ?????????? ? ???????????
???????, ????????? ?????. ? ?????? ????????
???????? ???????? ?? ????? ??????????????? ?? ?????? ?? ???? ??????? (????????, ??????????, 1989; Vorobjev et al., 1994).
??? ???????? ?????????????? ??????
???????????? ????????? ??????? ????????. ????????? ????? ????????? ??????? ??
?????????? ? ?????????. ?????????? ????? ???????????? ????? ??? ? ?????????????
? ??? ????????? ??????? ?????????? ?????.
?????? ????? ? ??? ?????????? ? ??????????? ?? ???????? ????????, ???????????.
????????? ????? ???? ???????????, ?.?. ????????? ???????? ????????, ? ???????????, ?.?. ???????? ???????? ?????????. ???????? ????????? ????? ????????????? ???
?? ?????????? ?????? (? ??????? ??????????
????), ??? ? ?? ?????? (? ??????? ??????????
??????????). ????????????? ? ????????? ?????? ???????? ???? ???????? ??? ????????
???? ???????? ????????: (1) 5-??????? ??????????? ?????? ??? ???????????; (2) ???????? ?????. ?? 2-?????? (??????????????) ?????? ??? ???????? ????? ???? ?????????????
? ??????????? ???????????, ???? ?? ?????????????, ???????? ?????????? ??????? ???
??????????????. ????????? ?????? (?? ?????????? kryptos ??????, ???????) ??? ? ????
????? ????????? ? ??? ???????? ?????????
???? ???????????? ???????? ???????? ? ????????? ????? (Borthwick, 1899). ?????????
????? ? Pinaceae, ?????????????, ????????????. ????? ???????????? ??? ????? ??????
?? ?????? ??????? ???????????? ??????, ??
? ????? (Bannan, 1942). ???????? ???? ????????? ?????? ??????????? ????????????
??????? ?? ????????? ????? (Owens, Molder,
1974). ?????? «????????????» ????? ??????
???????? ?? ????????????? ???????????-
?????????. ??????? ?? ???????????? ??? ?
????? ??????.
??????? «?????????? ?????» ? «?????? ?????» ????????????? ????? ??????????.
?????????????? ????? ???????, ??????? ???????? ??????????? ? ?????????????? ??????????? ?????????????????????? ????????
(Lanner, 1976; ??????????, 1994). ?????, ?????????? ???????? ????????? ?? ??????????
?????? ?????????????? ???????, ? ????????
?????. ?? ????? ???????? ?? ?????? (???????????????) ???? ?????????? (???????????????) ???????????? ???????. ???????????? ?????? ?????????? ?? ???? ?????????:
??????? ????? ???????????? (?????? ??????
? ?????? ? ??????? ? ?????? ??????) ? ??????? ????? ????????? ????? ??????????
(???????? ? ??????? ? ????????). ???????????? ?????? ??????? ?? ??? ???? ?? ?????????????: ???????? ??????, ? ??????? ????????? ????? ? ?????????? ?????? ?????????
???????? ??????? ?????, ? ?????? ??????, ?
??????? ????????? ????? ? ?????????? ?????? ?????????? ?? ?????????? ?????? ?????????????? ???????. ?????? ?????? ? ?????????? ??????????? ???? ????????: ??? ??
????? ?? ????????? ??????????????, ??? ???
?????????? ?? ??????? ??????? ?????, ?????
???? ???????????? ???????? ????????????
?????, ?????????? ???????? ????? ?????????? ????.
????????????? ???????? ????? ??????
? ????????? ?????????? ??? ?????? ?????????????? ??????? (????? ???????? ?????????? ?????). ? ???????? ??? ???????? ??????? ??? ??????. ??????? ?????????? ????? ?
???? ? ????? ???????? ?????????? ??????????
???????? ????? ???? ??? ? = 0,95.
??????????
????????? ? ???????? ??????????? ?????? ? ???????. ????? ????????? ????????
# 61 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
??????? ????? ??????? ? ???????? ? ???????????? ??????? ?????. ??? ?????? ????????????????? ?????????????? ??????? ?
????? ??????????? ?????????? ? ???????????? ??????, ??? ????? ?????????? ? ????? ????????????? ????, ?????? ???????? ?? ?????
?????????, ??? ???????, ??????????? ??????, ??? ?? ????? ??????. ? 1-?????? ???????
????? ???????? ? ???????????? ???? ???????
?? ??????? ??????????? ?????. ??????? ????? ???????????? ?????????? ????????????
2-?????? ??????? (???. 1). ? ???? ?????????
????????? ???? ????? ??????? ?? ??????
?????? ? ???????????? ? ?????????? ??????, ??? ????????? ??????? ????? ?? ??????:
(1) ????????????-?????????????? P. pumila,
(2) ?????????-???????????? P. sibirica ? P.
cembra, (3) ??????????? ?? ????????? ?????
P. koraiensis ? P. parviflora, (4) ?????????????? P. armandii.
????????? ? ???????? ??????????? ?????? ????????????? ??? ???????? ????????:
????????????????? ??????????? ????? ?????????? (???? ??? ??? ????????????? ???????)
? ??????? ?? ??? ???????? ???????? «???????» ???? ? ????????????. ???????? ?????
?????????? ?????? ?? ????? ?????????
?????????? ???????????, ?????? ?? ???? ??
??? ?? ????????? ???????-?????? ?????? ??????????? ?? ????????????????? ?????????????? ??????? ? ????? ??????????? ?????????? ? ???????????? ?????? ?????. ???,
????????????? ?????????? ???? ????????
??????????? ?????? ?????????? ??? ?????
?? ????? ????????????? ????????? ???, ?
???? ? ??????????????? ??? ?????????????
?????????? ?????? ???????? ???????????
?????? ???????? ???????? ????? ????? ????? ?? 30 ?? 70є ?.?. ??????? ???????????? ??
?????????? ?????? ????? ?? ??????? ? ????????: ?? ? ???????????? ?????????? ????? ?
?????????????? P. Pumila, ? ??????????????
P. armandii. ? ?? ?? ?????, ????? ?????????
????? ???? ???????????? ? ??? ????, ??????? ?? ????????? ? ???????? ???????????
?????? (????. 2):
(1) «???????????????» ?????? ? P. sibirica
+ P. cembra. ???? ???????? ??????????? ?????? ?????? 1-??????, ??????????? ?? ??? ?????.
(2) «??????????» ?????? ? P. koraiensis +
P. armandii. ???? ???????? ??????????? ?????? ?????? 1-??????, ??????????? ?? ??? ???? ??????.
(3) «?????????» ?????? ? P. parviflora +
P. pumila. ???? ???????? ???????????
?????? ????????????? ??? ??????????????? 2-??????, ??????????? ?? ???
?????? ??? ???????????.
????? ?? ???????? ????? ????? ??????
????????. ?????? ?? ???????? ????? ????? ??
??????????? ?????????????? ????????? ????
??????????? ? ????????? ????? ? ????????
(????. 3). ?????????? ?? ??????? ?????????????? ?????? ????? ?????? ???????????. ??
???????? ?????? ????? ???????? ????????
?????????? 15-25 ??. ??? ????????????? ?
????????? ???????: ?????????????, ???????????, ????????????, ??????????? ???????????. ?? ???? ??????? ????? ???????? ???????
?????? ??????????? ??????? ?? ???? ???????.
?????????? ????????, ?.?. ???????? ?????????????, ???????? ?? 40 ?? 70 % ???????.
???? ????????????? ??? ???????????? ??????? ?? ???????? ?????, ?? ??????? ????? ???????????? ????? ??????? ??????, ??? ???????
????? ??????????????. ???? ?? ????????????
??? ??????? ?????? ??????? ??????, ?? ?? ???
???? ?????????????? ???????? 50-75 % ??
?????? ????? ???????? ???????????. ? ?????????? ????? ??????? ?????? ?????????????
??? ???? ???????????, ??? ???? ???????????
??????????. ???????????? ???????? ? 40-90,
??????????? ??????????? ? ? 10-30 % ???????.
# 62 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
P. pumila
P. sibirica
P. cembra
P. koraiensis
P. parviflora
P. armandii
???
????
????
??????
????????
???. 1. ????? ????? ??????? ? ???? ? 2-?????? ??????? ????????? ????? Pinus ?? ?????? Strobus
??????? 2. ????????? ??????????? ?????? ? ????????????? ????? Pinus ?? ?????? Strobus
????
P. cembra
P. sibirica
P. pumila
P. parviflora
P. koraiensis
P. armandii
??? ??????????? ??????, ????????????? ? %
1-?????? ??????????
1-?????? ??????????
2-?????? ??????????
?????, ???????????
?????, ???????????
?????, ???????????
???????????
???????
???????
85
15
0
90
10
0
30
10
60
0
0
100
0
100
0
0
100
0
?? ??????? ????? ?????? ???????
?????????, ? ?????? ???????, ???????????
?????? ??? P. pumila ???????????? ?????
????????????? ? ????????????? ????? ??????. ??? ??????? ?????????? ? ???? ????????, ?? ???????? ? ?????? ???????. ? ??????????????? ??????? ????? ????????????
????? ??? ???????? ? 42 % ???????, ? ?
???????????????? ??????? ??????? ? ????
???? ???????? ? 12 % ???????. ?? ??????? ?
??????????????? ??? ????????????? ?????
????. ?? ???????? ??????? ?? ????? ????????? ?? 1 ?? 12 ??., ????????? ? ??????? 4-8
??. ???? ? ?????? ????, ??? ?????????????
???????????? ??????? ?????? ?? ?? ????
???????, ?? ??????? ????? ? 2-6 ??? ????????? ????? ??????? (??????????) ?????????????, ? ????? ????? ????????????? ?
P. pumila ??????????? ? 6-10 ??? ??????, ???
? ???? ?????? ?????. ? ??????????? ???????? ?????? ????? ????????????? ????????????? ??????. ?? ????????? ?????? ?? ????
???????? ?? ?????? ?????? ?? ???????????? ?????? 10 ??? ?????. ?? ???? ?? ??????
? ??????? 5 % ?????????? ????????????? ?
???? ???? ????? ?????? ????? ?????? ??????? ??????.
?? ????????? ????????? ????? ??????
??? ???????????? ????????. ?????????????
???????? ???????????? ?????? ????????? ?????????????? ????????, ??? ?? ??????????????? ? ????????????? ????????. ???????
??? ???????? ?????? ? ??????????? ?????.
? ????????? ???????? ????????? ????????????? ???? ?????????? ??????? ??????? ? ????????? ?? ?????.
# 63 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
??????? 3. ?????????????? ??????? ?? ???????? ????? ?????
????????
???? ? ???????
Pinus pumila
4
5
8
Pinus sibirica
1
2
3
Pinus koraiensis
10
11
12
?????? ?? ???????? ?????
???? ??????? ???????,
%
????? ??????, ??
50
72
64
56
60
48
42
50
60
1,9 ??*
1,75 ?
2,1?
2,2 ?
2,3 ?
2,05 ??
1,07 ?
2,16 ?
1,15 ?
-
-
-
5,11 ?
2,2 ?
1,81 ?
-
-
9,9 ?
9,3 ?
12,2 ?
6,94 ?
12,2 ?
12,8 ?
7,4 ?
11,41
??
6,8 ?
9,3 ?
6,48 ?
6,6 ?
5,14 ?
7,5 ?
0,8 ?
0,91 ?
0,79 ?
0,53 ?
0,4 ?
0,2 ??
0,23 ??
0,11 ?
0,29 ?
0,2 ??
24,7 ?
22,2 ?
14,7 ?
17,4 ??
14,9 ?
????? ????????????, ??. 6,84 ?? 4,4 ? 6,59 ?? 8,2 ?? 8,5 ?? 12,7 ?
????? ??????????????,
??.
12,1 ? 14,9 ?? 15,7 ?? 14,3 ?? 17,3 ? 13,2 ??
????? ????? ????????
????????, ??
19,8 ?? 20,1 ?? 23,2 ? 24,7 ? 27,03 ? 26,1 ?
???? ??????????????, %
61
74
68
58
64
51
4,9 ?
6,2 ??
11,5 ?
6,3 ??
15,81
??
12,7 ??
16,4 ?
22,1 ?
18,9 ??
70
72
67
????? ?????????????
?????????????, ??.
????? ????????????, ??.
????? ??????????????,
??.
5,9 ?
7,7 ?
6,3 ?
7,32 ?
????? ??????????
?????????????, ??.
0,78 ?
0,6 ?
0,7 ?
0,83 ?
????? ???????????
????????????, ??.
0,14 ?? 0,09 ? 0,15 ?? 0,19 ??
????? ????? ????????
????????, ??
16,8 ? 17,7 ?? 19,4 ?? 20,4 ?
?????????? ??????? ??????
??????????. ???????? ??????????? ???????? ????? ? ????? ????????? ? ?? ???????????? ? ????. 1
* ? ??????? ?????????? ????? ? ???? ? ????? ???????? ?????????? ?????????? ???????? ????? ???? ???
? = 0,95.
????? ?? ???????? ????? ?????. ????????????? ? ????????? ????????? ??????. ? ??????? ? ?????????????????? ????????? ?????? ? ???? ????? ??????? ????? ?????? (???. 2).
??????????? ???? ???????????? ??????? ??
???? ????????????? ?????? ???????? ? ????
???? ?????. ? ???????? ????? ??????????
?????? ???? (????????) ?????????? ?????,
?????? ? ??? ?????????? ????? ?????????????
??????? ???????????????????? ?????????
??????? ?????. ?????????? ????????? ???????? ??????????? ???????????, ????????? ???????? ??????????. ??????? ????????? ??????? ??????????? ?????? ? ????????
?????? ???, ??? ????????? ????????????? ???
??? ??????, ????????? ? ?????????? ????????
?????????? ? ??????? ?????? ??????????????
???????.
??? ????????? ???? ??????????? ??????????? ?? ????????????? ????????? ?????
?????????????. ???????? ????? ????????
?? ???????????? ??????? ????????????? ?
??????? ??????? ??????. ? ???? ??? ??????????? ???? ????? ??????, ?????? ????????????? ???????? ????? ????????? ?????
????????. P. pumila ?????? ?????????? ?? ???.
???, ???? ??????? ? ??????? ? ?. sibirica ?????????? 88-93%, ? P. koraiensis ? 90-97%, ? ?
P. pumila ? 44-87%. ? ???????? ???? ???????? ?????????? ??????? ????????????? ????-
# 64 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
???. 2. ??????? ?????? P. koraiensis (?), P. sibirica (?) ? P. pumila (?) ? ????????? ??????? ????? ? ?????????
?????: 1 ? ???????????? ?????, 2 ? ??????????? ?????????? ?? ?????? ?????? (?????), 3 ? ???????????
?????????? ?? ?????? ??????, 4 ? ??????????? ?? ?????? ??????, 5 ? ?????????? ?? ?????? ??????,
6 ? ?????????? ????????? ??????? ??????, 7 ? 1-?????? ????? (?????), 8 ? ???? ?? ????????? ?????,
9 ? ??????????? ?????????? ?? ???????? ?????? (?????), 10 ? ??????????? ?????????? ?? ???????? ??????,
11 ? ?????????? ??????????? ?? ???????? ??????, 12 ? ?????????? ?? ???????? ??????, 13 ? ?????????????
??????????? ?? ???????? ??????, 14 ? ?????????? ????????? ????????? ??????
?????? ? ???????? ? ???????????? ????????.
??????????? ???????? ??????? ?????? ? P.
pumila, ? ??? ????????? ???????????????
?????? ? ???? ????????? ????? ? ??????????? ?????, ??? ? ????????? (??????) ????????.
??????? ? ???????? ???????????? ??
????????????? ??????? ?????? ????? ??????
? ?????? ??? ?????????. ? ??????????? ????? ???? ???????? ??????? ?????? (8-10 %)
?? ????? ??????? ?????? ?? ??????? ??????.
? ????????? ???????? ?????????? ???????
?????? ? ??????? ????????. P. pumila ??????????????? ??????????? ??????? ?????????????? ??????? ??????. ???? ? ???????? ????-
???? ? ???? ????????? ? ??????????? ?????
?????????? ???????????????? ??????? ????
????? ???????? ???????? ??????? ?????
??????? ?????????. ? ??????????????? ?
????????????? ???????????????? ????????
????????????? ??????? ?????? ?????????? ????? 15 %.
? ????? ??????????? ????? ???????
????? ??????????? ??????????? ???????????
??????? ??????, ? ?????? ????? ? ??????????
????? ??????? ???????????. ? P. pumila ????????? ??????? ??????? ??? ????, ? ???????
????? ?? ??????????? ??????????? ???????
??????. ???? ? ?????????? ????????????????
? ???????????? ???????????????? ????????
# 65 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
?????? ????? ?? ?????????? ??????? ???????
???? ???????, ??? ???????????. ? ?????????
???????? ??????????? ??????? ??????? ??????? ?????????.
?????????? ???????? ????? ??????
? ????????? ?? ??????? ??? ???????, ?????
???????? ????? ????? ??????? ?????????
(????. 4). ?? ?????? ????? ???????? ??????????? ????????? ???? ????? ??????. ???????? ????? ????????? ???????? ??????,
?????? ? ??????????? ???????????????????
?????????????? ??????? ????? ????????
???????? ??????????. ??? ??????????????
???????? ????? ???????? ? P. pumila. ? ??????????? ????? ??? ??????????? ?? ??? ????
??-?? ??????? ???????? ?????????????? ????????, ? ??????????? ???????? ????? ????????
???????? ?????????. ???? ??????? ?????? ?
???????? ?????????? ???????? ? P. pumila ???????????, ? P. sibirica ? ????????? ????, ? ?
P. koraiensis ? ??????????? ????, ??? ? ????
?????? ?????. ???????? ????? ?????????
?????????????. ???????????? ????? ????????? ?????? ????????? ???????????? ??????
????? ???????? ????????. ?????? ????????
????? ????????? ???????? ????? ????, ?.?. ?
??????????? ???????? ???????? (P. sibirica, P.
koraiensis) ? ??????????????????? ?????????????? ?????? (P. pumila) ??????????? ?? ??????
????? ?????????, ?? ? ????? ??????????.
????? ???? ?? ???????? ?????? ? P. pumila
??????????? ??????, ??? ? ???? ?????? ?????. ????????????? ???????????? ?? ?????
???????? ? ?????????? ???????? ?????????
???????????? ?? ????? ?????? ? ?????? ????? ?????????. ?? ????? ???? ?? ?????? ?????? ???????? ????? ?????? ??????? ??????:
P. sibirica ?????????? ???????? P. koraiensis ?
?????????? ??????????? P. pumila. ????????
????? ????????? ?????????????. ???? ??
?????? ?????? ? ???? ???? ????? ??????????? ??????, ??? ?? ????????. ?????????????
????????? ?????? ?? ????? ???????? ???????? ???????? ? P. pumila, ? ???????? ? ? P.
koraiensis.
????? ?? ???????? ????? ?????. ????????? ????????? ??????. ? ???? ???? ????? ?
????????? ????????? ?????? ?????????????
?????????? ?????????, ? ????????? ?????
?????? ???????? ?????????? ????????? ?
?????????????, ??????????????, ???????????? ????????????? ? ??????? (????. 5).
?????? ? ????? ?????????????????? ??? ?????? ???? ???????? ???????? ????????????? ?? ???????????? ??????? ??????? P. sibirica
? P. koraiensis. ? P. pumila ????? ???????????
?????????????? ????????????, ?????????
????????????? ? ????????????? ????? ?????? ????? ??????????? ??????????? ? ?????????????. ???????? ?? ????? ??????? ???????????????, ????????????? ????????????
???????? ???????? ?????????? ?????????
??????. ?? ????????????? ?? ????? ???????????? ??????????? ?? ???????????????????
?????????????? ???????. ???, ??? ????????
??????? ? ?????????????? ? ???????????
??????????????? ??????????, ?? ??????????
??????????? ??? ??????????? ?????? ?????
?????????????? ? ????????? ??????????????? ? ???????????? ??????????????? ??????????. ???? ? ?????????? ? ??????? ????????
??????????? ????????????? ?????????????
?? ????? ? ??????? ???? 50 % ???????. ????? ????????????? ?? ????? ??????? ?????????
?? 2 ?? 26 ??., ? ? ??????? ?????????? 4-10 ??.
?? ?????? ?????? ?? ???????????? ?? ????????. ? 10-??????? ???????? 70-80 % ????????????? ????????????? ??????.
??????? ????? ???????? ???????? ????????? ?????? (80-90 %) ? ???? ???? ????? ?????????? ???????????. ? ?????????? ?????
?????? ????????? ??????????? ??????????
????????????, ??????????? ??????????? ?
?????. ? P. sibirica ? P. pumila ? ??????????
# 66 #
5,7 ??
0,3 ?
5
????? ????????? ??????, ??
????? ??????? ??????, ??
????? ??????? ??????, %
10,1 ?
5?
50
9,9 ?
4,6 ?
46
5
0,2 ?
3,8 ?
52
5,8 ?
11,2 ??
4
0,3 ?
6,8 ?
41
3,2 ?
7,9 ??
4
0,3 ?
6,8 ?
7,1 ?
5
13
7,1 ?
41
3,1 ?
7,5 ?
5
0,3 ?
6,2 ?
6,5 ?
5
2,5 ?
45,4 ??
47,9 ??
5
35
2,2 ?
6,4 ?
6
0,2 ?
2,9 ??
3,1 ?
8
3,2 ?
34,7 ?
37,9 ?
6
-
-
-
4
0,1 ?
2,3 ?
2,4 ?
10
3,4 ?
29,1 ?
32,6 ?
7
Pinus pumila
???? ? ???????
40
3,4 ?
8,6 ??
4
0,4 ?
9,6 ?
10 ?
5
3,1 ?
54,3 ?
57,4 ??
8
* ? ??????? ?????????? ????? ? ???? ? ????? ???????? ?????????? ?????????? ???????? ????? ???? ??? ? = 0,95
??? ???? ? ??. ???????? ??????????? ???????? ????? ? ????? ????????? ? ?? ???????????? ? ????. 1
????? ???? ?????????
??????, ??
????? ???? ??????? ??????,
??
????? ???? ??????? ??????,
% ?? ????? ???? ?????????
??????
17
4?
2,5 ?
6,1 ??
5,6 ?
5,3 ?
52,5 ??
4
50 ?
27,9 ?
38,7 ??
47,7 ??
3
41,6 ??
33,5 ?
44,1 ??*
12
2
6 ??
1
Pinus sibirica
????? ???????? ????????
?? ???????? ??????, ??.
????? ???????? ????????
?? ???????? ??????, ??.
????? ???????? ????????
?? ?????? ??????, ??.
????? ???????? ????????
?? ?????? ??????, %
????? ????????? ??????, ??
????????
??????? 4. ?????????????? ???????? ????????? ?????? ?? ?????? ?? ???????? ????? ?????
36
2,9 ?
8,1 ??
6
0,6 ?
8,9 ?
9,5 ?
11
7,4 ??
57,7 ?
65,1 ?
9
62
6,6 ?
10,7 ??
17
1,5 ?
7,5 ??
9 ??
31
10,9 ??
24,7 ?
35,6 ?
10
69
9?
13,1 ?
27
5,8 ?
15,6 ?
21,4 ?
29
19,5 ?
46,8 ??
66,3 ?
11
51,??
12
72
8,1 ?
11,2 ??
26
3,1 ?
9?
12,1 ?
25
12,8 ??
38,4 ??
Pinus koraiensis
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
0,40?
2,00?
2,05?
2,11?
0,43?
1,94?
0,62??
0,55?
0,51?
25,0?
35,0?
0,98?
-
2
Pinus sibirica
-
1
2,23?
0,42?
2,12?
0,43?
1,0?
38,0?
-
3
1,91?
2,50?
1,50?
3,0?
0,80
40,0?
4,64?*
4
2,20?
2,0?
1,75?
2,5?
0,70??
38,0??
2,48?
5
1,84?
0,65?
1,20??
0,9?
1,39?
31,0??
0,23?
1,32?
0,70?
0,59?
1,0?
1,32?
25,6?
0,6?
7
Pinus pumila
6
2,11?
2,90?
1,63?
3,5?
0,90?
46,0?
2,22?
8
* ? ??????? ?????????? ????? ? ???? ? ????? ???????? ?????????? ?????????? ???????? ????? ???? ??? ? = 0,95
????? ???????????
????????????, ??.
????? ?????, ??.
?????????? ??????? ??????
????? ?????????? ??????
?????, ??.
????? ???????????
????????????, ??.
????? ?????, ??.
????? ?????????????
?????? ?????, ??.
????? ????????????, ??
?????? ?? ???????? ????? ?????
????????
???? ? ???????
??????? 5. ?????????????? ???????? ????????? ?????? ?? ?????? ?? ???????? ????? ?????
3,16?
1,58?
2,68??
1,72??
0,82?
52,0??
0?
9
3,60?
0,08?
3,0?
0,22?
0,14?
21,3?
-
10
3,5?
0,02?
3,20?
0,32?
0,28?
43,0??
-
11
Pinus koraiensis
3,1?
0,01?
2,90?
0,27?
0,18?
35,0?
-
12
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
????? ????????? ?????? ?????? ??????? ????
???????????. ???????? ????? ????????? ??
????? ???????? ?????????????. ???? ? ?????????????? ??????? P. sibirica ??????????
????????????? ????????? ?????? ??-?? ???????? ???????? ?????????????? ????????. ??
??????? P. koraiensis ?????????? ???????????? ??????????? ?????. ?? ????? ???? 1438 % ???????. ?? ?????? 10 ??? ????? ? ????
????? ? ???????? ?????? 70-80%, ? ???? ?????? 5-7 % ?????????? ?????????????.
?? ????? ??????????? ????????????
??? ????????? ???? ????? ???????????. ? P.
koraiensis ?? ????? ????, ? ?? ??????? ? ??????? ? ??????????? ??? ?????. ? P. sibirica ??????????? ???????????? ?????????? ??????,
? ??????? 0,5 ??. ?? ?????. ???????? ?????
????????? ?? ????? ???????? ? ????? ???-
? P. Koraiensis, ??????- ? ??????????? ??????? ? ? P. Sibirica, ? ???????????? ?????? ?????????? ????? ?????? ?????????? ?????.
????? ?? ???????? ????? ?????. ????????? ??????? ??????. ? ???? ????? ?? ??????
?????? ??????????? ??? ???? ???????? ????????: ???????????, ???????????? ? ??????????? ??????????? (????. 6). ??? ??????? ????
???? ????????????? ????? ??? ?????????????
?????? ? ????????? ??????????????????. ?????? ????? ??????? ?????? ? P. sibirica ??????????? ???? (60-75 % ???????), ??? ????????. ?
P. koraiensis ? P. pumila ??????? ????? ??????
??????? (?????????????? 65-78 % ? 70-97 %)
?? ???????? ??????? ?????? ????? ????????
????????. ??????????? ?????? ??????? P.
sibirica ????? ????????? ????????????, ????
??????????? ? 1-2 ??????????? ???????????. ??
???????? ????? ?????????????. P. pumila
??????????????? ???????? ?????????? ??????????? ???????????? ?? ???????? ??????
(? 2-7 ??? ??????, ??? ? P. Sibirica, ? ? 4-10
??? ??????, ??? ? P. koraiensis). ?? ??????? ?
??????? ????? ???? ??????? ??????? ??????, ??? ?? ??????? ??? ?????, ?? ??? ?????
?????????? ??????. ? P. pumila ?? ????? ???????? ??????? ???????????? ???????? ????? ?????????. ??????? ? ??????? ? ??????
???????????????????? ?????????????? ??????? (??????????????? ?????????, ??????- ?
???????????? ???????????????) ????? ??????????? ????? ??????????? ????????????.
???????????? ????? ????? ?? ??????
?????????? ??? P. koraiensis. ? ????? ???? ??
???????? ????????? ?????? ? 6, 7 ? ???? 8
???????. ????? ????? ?? ??????? P. sibirica
??????????? ??????. ? ????? ???????????
????? ???????? ????? ????????? ?? ?????
???????? ??????????? ???????????. ????????, ??????? P. pumila ?????? ??????????? ??
????? ????? ?? ???????. ????????? ??????????????? ?????? ? ???? ????????? ??????
?????? ??????? P. koraiensis ???? ????????????
??????????? ????, ? ???? ????????????? ? ??
????? ??? ????, ??? ? P. sibirica. ???????? ????? ????????? ?????????????. ? ???????? ???????? ???? ????????????? ????????? ????????????? ?? ???? ???????? ???? ????????????. P.
pumila ????? ?????????? ?? ???? ?????? ?????
?????????? ????? ????????????. ????????,
???? ??????????? ???????????? ? ????????
????????????? ??????????? ????, ??? ? ??????????? ?????. ???????? ????? ?????????
????? ??????. ?????????????, ????? ? ????
???????????? ??????????? ?????????? ? ??????????? ??????????????????? ??????????????
???????. ?? ??????????? ????? ?????????????? ? ???????????? ????????????? ??????? P. pumila ?????????? ????? ??????? ?? ???
??????. ???????? ????????? ?????????????
?????????? ??? ??????? ?? ??????????????????? ????????: ???????????????? ??????????
? ???????????????? ??????- ? ?????????????.
? ??????????????? ? ????? ??????????? ???????? ????? ????????????? ?? ?????? ??????
???????? (??? ? P. sibirica).
# 69 #
1
2
Pinus sibirica
61
19
24
????????????
??????????? ???????????
26
68
24
8
1,7?
0,6?
0,2?
90
55
5
4
60
28
12
1,5?
0,7?
0,3?
85
60
9
5
16
71
13
0,5?
2,3?
0,4?
46
100
38
6
9
88
3
0,33?
3,1?
0,1?
11
100
11
7
Pinus pumila
38
26
16
1,8?
0,8?
0,5?
86
70
16
8
* ? ??????? ?????????? ????? ? ???? ? ????? ???????? ?????????? ?????????? ???????? ????? ???? ??? ? = 0,95
24
13
13
57
???????????
63
1,6?
0,8?
3,7?
95
74
95
3
1,3?
1,4?
????? ???????? ????????, % ?? ?????? ??????????
??????????? ???????????
1,0?
?????? ?????
0,7?
3,5?
3,0?*
???????????
66
86
85
90
85
88
????? ???????? ????????, ??. ?? ?????
??????????? ???????????
?????? ?????
???????????
????????????? ???????? ????????, % ?? ????? ???????
????????
???? ? ???????
??????? 6. ?????????????? ???????? ??????? ?????? ?? ?????? ?? ???????? ????? ?????
21
50
29
1,5?
3,7?
2,2??
69
87
46
9
30
4
66
3,3??
0,39?
7,2?
97
33
100
10
20
2
78
4,0?
0,3?
15,2?
100
35
100
11
Pinus koraiensis
22
2
76
2,8?
0,2?
9,6?
100
22
100
12
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
????? ?? ???????? ????? ?????. ????????????? ?????? ????????? ????????? ? ??????? ???????. ? ???? ???? ????? ????????
????? ???????? ???????? ???????? ??????????? (80-90 % ?? ?????? ????? ????????
????????). ??????? ????? ???????????? ??????????? ?? ???????? ??????. ???? ???????
???????? ?????? ? P. koraiensis (22-26 %). ?
P. sibirica ??? ??????????? ?????? (7-12 %), ?
? P. pumila ? ???????? ???? (2 % ? ???????????????? ?????????? ??????? ? ????? 1 % ?
???? ?????????).
???????????? ???????? ????? ? P.
koraiensis (0,7-1,4 % ?? ?????? ????? ???????? ????????), ??????, ????? ???????? ??
??????????? ?? ?????? ?????? (52-72 %). ?
P. sibirica ???? ????????????? ???????????
???? (3,6-4,4 %), ? ??? ???????? ???????
???????????? ????? ????? ????????????? ????????. ???????? ???????? ?????????
????????????? ?????????? ??? P. pumila (4,611,0 % ?? ?????? ????? ???????? ????????).
????? ????????? P. pumila ??????? ???????
???????? ?? ????? ????????, ? ????? ?? ???????????? ????????????? ?? ???????? ??????.
?????????? ???? ????????????? ??????????
??? ?????????? ???????????????? ???????,
?????? 72 % ?? ?? ?????? ?????????? ?????????? ?? ?????? ?????. ? ???????????????
? ????? ??????????? ???????? ???? ???????
?????? ?? ????????? 7 %, ? ?? ???????? ?????? ??????????? ????????????? ???????????
? ??? ????????????? ????? (67-84 %).
???????????? ???? ??????????? ???????????? (?? ?????? ????? ???????? ????????) ?????????? ??? P. koraiensis ? 5,9-9,3 %.
????? ??? ??? ?????? (92-95 %) ??????????? ?? ?????? ??????. ? P. sibirica ???? ???????????? ????????? ?????? (4,2-5,4 %), ? ??
?????????????? ? ??????? ?????? ????????
?? ??? ???? (65-79 %). P. pumila ? ?? ???? ????????? ?????????? ??????? ?????????????.
???? ??????????? ???????????? ????????? ??
2,8-3,5 % ? ??????- ? ????????????? ????????
?? 5,0-7,2 % ? ??????????? ? ???????????????? ?????????? ????????. ??????? ?????
??????????? ???????????? ?????????? ??
???????? ??????. ???? ??????? ????????? ??
5-10 % ? ???????????????, ??????- ? ????????????? ???????? ?? 33 % ? ????????????????
?????????? ???????. ????? ?????????? 2,08,6 % ?? ?????? ????? ???????? ????????
?? ??????? ??????. ??? ??????????? ???? ?
???? ????????? ??????. ???? ????? ? ???
?? ???? 4,4 %. ??? P. pumila ??? ????????
????????, ????????, ??????????? (??????????????? ????????? ??????). ???????????
???? ????? (2,0 %) ???????? ? ????????????? ???????.
??????????
????? ???????, ?????? ???????????? ?????????????? ????????? ????????? ?????????
????????? ???????? ? ?????? ???????????????? ???????? ????????????? ???????????
????????? Cembrae, ? ????????? ??? ?????????? ?????????? ? ?? ??????????????, ?????????????, ? ????????????? ?? ?????????. ?
?????, ??????????? ?????????? ?????? ????????????? ???????? ??????? ???????????? ????????? ???????????? ?????? ??????
????????????????? ?????: ??? ?????? ??? ???
???? ??????? ??? ????????? ????, ??? ?????? ?? ?????? ? ??? ??????????? ? ?????? ? ?
????????????
????????-??????????????
????????? ??????. ??? ??????????? ? ?????????? ????????? ??????????? ? ??????????
????????? ????????, ?? ??????????? ?????????? ?? ???????? ?? ???????? ?????
????????. ???????? ?????????, ??????????????? ???????? ??????????? ??????, ?? ? ?????? ???? ???????? ????? ???????. ?????????? ????? ? ??? ????? ??? ???????? ?????.
?? ???? ?????????? ? ?????????? Pinus ? ???
# 71 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
??????? ??????, ????? ?????? ???????????
????? ??????? ? ???????? ??????? ?? ?????
??????????? ????????. ??? P. parviflora ??
?????? ? ????????-?????? ???????????? ???????? 2-?????? ???? ???????? ???????????
??????, ??????, ?????? ?????????. ??? ???????????????? ?????????? ? ??? «????? ???????» P. pumila, ??????????? ???????? ?????, ?????????? ??????-????????? ??????
? ?? ????? ??????????????? ??????????????
????????, ??? ????????????? ????????, ?????????????? ?????????? ??????????? ????.
??????????? ??????? ??????? ????????????
????????? ??????????? ??????, ??????????? ? ????? ????, ???? ??????????????? ? ???
????????????? ?????????, ? ???????????? ?
????????? ????? ??? ???????? ????????. ??????? ? P. pumila ????????????? ????? ??????
? «?????????» ?????????? ??????????? ?????? ?????????? ???? ?? ??????????? ?????
????????.
???????? ? ??????? ????????? ?????? ?
?????? ????????, ??????? ????????, ??? ???
????????? ?????????? ???? ????? ?????
?????? ? ????????? ? ???????? ?????????
??????. ? ??????? ????? ????? ?? ???????
?? ????????????? ????????????? ??????, ? ?
??????? ? ?? ????: ????????? (?????????) ?
??????????????? (???????). ? ???? ???? ?????
?????? ????? ?????????? ?? ?????? ? ??????????? ???????? ????????? ?? ????? ?????????, ????? ??? ? ????. ???????? ?????????
????????? ?????? ? Pinus ?????? ??????????????? ??? ????????????? ???????? ????????????? ?????. ???, ?.?. ????????? (1971)
??????????, ??? ?? ?????? ????????????????? ????? ?????? ???? ????? ??????????????? ???????? ??????, ?????? ??? ??? ???????? ?????????? ?????????. ?? R.M. Lanner
(1976), ?????????? ??????????? ?????? ??????? ?????????? ??? ?????, ?????????????? ?
??????? ? ??????? ????????????? ????????,
???????? ? ?????? ?????. ? ????? ????????
????????? ???????????????? ??????????????
????? ???????? ??????? ?? ????????? ??????
?????? ??????? ???????????? ??????????? ????????????? ????????????? ???????? ?????.
???????? ????????? ????????? ?????? ? ????
????????? ????? ? ????????? ??? ???????????? ???????? ?? ?????????????. ???????
??????? ?????? ????????? ????????????, ???
?????????????? ????? ??????, ?????? ?????,
???????????? ? ????? ?????? ???????, ???
???, ??????? ?????????? ??? ???????? ??????? ????.
? ????? Pinus, ??????? ? ?????????
????? ???????????? ? ????? ???????? ?????????? ????? ??? ? ???????????, ?????? ?????? ?????? ?? ??????? ? ???????? (Tepper,
1963; Lanner, 1970; Sirikul, Kanninen, 1990).
??????? ????? ????????? ?????? ? ?????????? ???? ???????????? ????????? ????????
? ???????? ????, ??? ??????? ????????????
?????? ?? ??????????????? ??????? ????????? ??? (Lanner, 1976). ? ???? ???? ????????? ????? ?????? ????? ??????????? ???????? ????????? ?? ????? ? ????? ????????
????????. ???????? ????? ?? ???????????
???? ???????????? ??????? ?????????? ? ???
????????????? ??????????? ???? P. strobus
?? ??????? ?????? Strobi (Paul, 1957; Owston,
1968). ??? ????? ?????????????? ? ???????????? ??????? ????????????? ????????.
????????? ????? ?????????????? ? ??? ????????? ????? ????? ?????. ???????????
????????????? ? ?????? ??????? ?????? ?????????? ??? ?????????????-??????????????
P. pumila, ???????????? ? ??? ????????????
P. koraiensis. ?????????? P. sibirica ????????
????????????? ?????????. ???? ?????????????? ?????????? ????????? ? ??????? ?????? ??????????? ??????????? ? ???? ?????.
???? ????? ? ????? ?????????????? ???????
???????, ?? ???? ????????? ??????. ???? ? P.
# 72 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
koraiensis ?????? ????? ?????? ???????? ????? ? ???? ???????. ? ???? ?????? ????? ???
?????????? ?????? ?????? ?? ?????????????. ???????? ????? ???? ? ???? ???? ?????
????????????? ?? ???????? ??????.
???????? ??????? ??????? ?????? ? ??????????? ?????????? ?????? ??? P. koraiensis.
? P. sibirica ???????????? ?? ?????? ?????? ?????? ????, ? ?? ???? ????????????. ? P. pumila
?????? ????? ??????????? ?? ????????? ?
???????????? ????????? ????????. ????????
?????? ???????? ????? ?????? ?? ???????
???? ????? ???????????? ??????? ? ?????????. ? P. koraiensis ??? ?????????????? ?????
????????????? ?? ???? ???????, ? P. sibirica ?
? ???????? ?? ???? ???????, ? ? P. pumila ? ?
???????? ?? ???? ????????? ??????.
????? ???????, ? ??????????? ????????????????? ? ??????????????????? ?????????????? ??????? ???? ????????? ?????????????? ????????????? ??????? ?? ????
???????????? ???????. ??? ???????? ?????? ??????? ?? ??????? ??????? ???????????? ? ?????????: ? P. pumila ??? ?????
????????? ????????? ?? ???????? ??????, ?
? P. koraiensis ? ????? ????????? ?????????
????? ????? ????????????? ????????. ??????? ????????: ? ??????? ???????????????
P. koraiensis ?????? ???????? ???? ???????
?????????????, ? ? ??????? ???????????????
P. pumila ? ?? ????????????? ????????. ??????? ? ?????? ?????? ???????????? ?????
???????????, ? ?? ?????? ? ?? ???????????
?????????? ??????? ????? ????????.
???????? ????? ????????? ?? ?????????
??????? ??????? ? P. sibirica ? P. koraiensis ???????? ? ??????? ?????? ? ????????? ?????????????? ????????, ??? ? ????????. ???? ? P.
koraiensis ?????????? ???????????? ????????????? ???????? ????? ???? ?? ?????????
????? ?????????? ???????? ? ??????? ????????????, ?? ??? P. sibirica ????? ???????-
??? ???? ????????????, ??? ? ?????? ???????????? ??? ??? ??????????? ? ?????????????
????????? ???????. ????? ????, ???????????
? ??????? ???????????, ?????????? ? ???????? ????????????? ??? P. sibirica ????????
??????-?????????? ????? (??????????, 1994)
? ????? ??????? ????? ???????? ??????
(??????????, ?????????, 1992), ??????????,
??? ???? ? ??????? ?? ????????????? ???????? ?????? ?????? ????????? ????????? ?????? ? ????? ???? ?? ????????? ????????????.
?????????, ?? ?????? ??????, ??????? ? ???,
??? ? ????????????? ???????? ? ????????
?????? P. sibirica ?? ??? ?? ??????. ?? ????
?????????? ?????? ??? ???????? ????????
? ???????? ??????????????? ?????? ???????- ? ????????????? ?????. ? P. pumila ??????? ???????? ???????? ?? ????????? ??????
?? ??????????????. ??? ?????? ???????????,
?.?. ???? ?????????? ??????????? ? ??????????? ??????????? ?????? ?????????????
??????????? ?????????? ???????? (????????, 1958; ??????????, 1975; ????????????,
1995). ????????, ???????????? ????????????? ???????? ????? ????????? ?????? ? ??????????. ?? ????????, ??? ??????? ?? ?????
???????? ???????????? ??????? ?????? ??????. ??????? ??????????????? P. pumila ??????????? ????? ????????? ? ?????????????
?????????. ???? ? ???????? ??? ?????????
??????? ????????, ??????? ???????????? ?
????????? ??????, ?????? ????????? ?? ?????
????????? ? ????? ????????????????, ?????????? ???????????? ??? P. sibirica, ?? ???????????? ??????? ?????????????, ?????? ??????????? ??? P. koraiensis. ??????? ????????
??????????????? P. pumila ? ????? ???????
????????????? ?????????, ??-????????, ???????? ??????????? ????? ?????, ??????????? ???? ? ??????? ???????? ? ?????? ??????? ????????, ? ??????? ????????????? ?
????????????? ????????? ? ???????? ???-
# 73 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
??????????. ?????? ???????????, ??? ???????????? ????????? ????????? ?????? ???????? ???????????? ????????????? ? ????
??????? ? ???????? ?????? P. pumila. ??? ????
??????????? ?????????? ?????? ??????????????: ??? ?????? ????????????????? ? ??????????????????? ?????????????? ???????,
??? ????????? ?????? ????? ? ???????????
??? ??????????? ?????. ??? ???? ?????????? ????????????? ??????? ?????? ? ?????
????????? ?? ??? ???????? ????????????????
????? ??????? ????????????? ?????????????
????????. ???????????? ??????? ????????????? ???????, ????? ???????????? ?????
? ?? ???? ?? ?????? ????? ????????? ?????
??????????? ?????????? ? ???? ?? ???????????????? ????????????? ? ????????????????
?????????? ???????. ??-????????, ??? ? ????????? ???? ???????? ????????? ? ??????????? ????????????? ? ????????? ????????????? ????.
????????????? ? ????? ????????????
????????? ???????? ????? ?????? ? ????????? ?? ????????????? ????????????? ? ??
???????????? ?? ???????. ??????????? ???????????? ????? ???????? ?????, ?????????
???????? ?????????? ??????????????? ?????????? ? ????????????????? ? ????????
????? ???????? ???????????? ???????????
(Powell, 74; Owens, Molder, 1973). ???????
?????? ???????????, ??? ???????????? ???????? ?????????? ??? ????????????? ???????.
? ????????????? ????????? ??? ???????????
??? ??? ????????? ??????? ? ??????? ??????? ????? ????????. ?? ??????? ???? ????????????? ??????? ????, ? ?? ????????? ? ??????????? ???????????? ??????????? ????,
??? ?? ???????. ?? ?? ????? ??????????? ?
??? ????????? ??????? ??????? ???? ?????:
????? ? ???????? ???? ????????????? ?????????? ? ??????????? ??????? ?????? ? ???? P.
koraiensis ? P. sibirica ? P. pumila. ???? ???-
???? ??????????? ????? ?????? ? ? ????
?????????, ?? ??????? P. pumila ????????
????????????. ????????? ?? ??. ????? ? ????
????????????? ????????????? ? ???????????
??????? ?????? ? ???? ?? ????????????????
?????????? ? ???????????????? ????????????? ???????. ?????????? ???????? ????
????? ?????? ???????. ????????? ????? ? ???
????????? ?????, ?????????????? ??????????? ???????????????? ???????? ? ?????? ???
??????????? ??? ????????? ??????? ????????. ??????????? ????? ??????????? ?????????? ? ??????? ????????????? ????????.
??????? ? ????????????? ???????????????
????????, ????????????? ????????? ????????????? ? ??????????? ???????? ????? ???????.
??????????? ??????? ? ?????? ?? ??????????? ??????? ??? ?????????? ?? ???????????? ??????? ????????? ?????. ??? ?????
??????? ???????? ? ? ?????????? ????????
????????? ????????? ????????, ???????? ??????????? (???????, 1964 ? ??.) ? ????????,
????????? ? ??????????? ?????????????
(Busgen, Munch, 1931? ??.). ? ????????? ???????? ????????????????? ?? ????????? ????? ???????????? ???????????? ?????????
?????? ?????????, ??????? ?? ??????? ??????
?????????? ?????? ????? ???? ????????????
????????? ?????, ????????? ?? ???????
? ????????? (????????, 1963; Harmer, 1990;
?????????, ?????????, 1992 ? ??.). P. sibirica
??????????????? ?????? ????????? ????????? ????? ? ???????? 200-300 ??? (??????????, 1992; ??????????, ?????????, 2000).
??? ?????????? ????????? ? ?????????????? ??-?? ?????? ??????????? ???????????
??????????? ????????? ??????????? (??????????, 1990 ? ??.) ?, ??? ?????????, ????????
?????? ???????????? ??? ??????? ????. ???
??????????????? ??????????? ??????????
???????????? ???????? ? ??????? ??????-
# 74 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
????? (???????????? ? ??????????????) ??
???? ?????? ?? ????????????????-??????????
???????? (??????????, 1956 ? ??.). ? ?????
???????? ???????-?????? ???????????? ???????? ????????? ????? ???? ?? ????????. ??
?????????, ??? ?????? ??????? ???????????? ????? ?? ???????????? ???????????
???????? ????? ????????????? ? ?? ???? ??
??????? P. koraiensis ??????????? ? 3-4 ????
??????, ??? ? P. sibirica.
??? ???? ??????? ????????? ????? ?
??????????? ???????????? ???????????
?????? (Bannan, 1942 ? ??.). ??????? ?????
???????????????? ?????? ?????????????
?????????? ? P. sibirica ?? ??????? ???????
??????????????? ? ????? (??????????, 1998).
???? ? ??????????? ????? ????????? ?? ????????? ????? ????? ????? ???????? ???????? ?????? ?? ???????? ??????, ?? ? P. pumila
??????????? ???????????? ???????????
?????? ?????????? ??????????? ? ?????????,
???????? ????????? ????????? ?????????
????????? ???? (??????????, 1975; ????????????, 1995 ? ??.). ????????, ??? ????????? ????? ???????? ????? ????????????? ??
??????? P. pumila ??????? ?????? ? ???? ???????????? ????.
?????????, ??? ??? ????????, ???????
????????? ? ????????? ????????????? ?
P. pumila (?? ?????, ???? ?? ?????? ????? ?????????, ???????????? ?? ??????), ?? ????????
???????? ?? ???? ?????? ????????????: ??
????? ????? ? ?????? ????, ?? ?????? ??????
?????? ????????, ? ?????? ?????? ????? ????????? ? ? ?????? ????????. ??-????????,
??? ?????? ????????? ? ????????? ????? ????????? ?? ????? ???????????, ?????? ????????
????????? ????????????? ??????. ? ??????
?????? ?????? ?????? ??????? ????? ????
???????????. ?? ????, ? ?????????, ??????????????? ???????????? ???????? ?? ??????
????????? ????????????? ? ?? ????????? ??
???????? ??????. ? ??????? ?????????????
???????? ??? (??? ? ??? ????????? ??????)
????????????? ?? ???????? ??????, ? ? ????????????? (??????????????? ?????????
??????), ??????? ???????, ?? ?????? ??????.
? ? ??? ? ? ?????? ?????? ???????????? ???????? ?? ???????????? ?????? ???? ??????? ?????: ????? ????????????? ? ?????????????. ? ???? ???????? P. pumila ?? ?????
???????????? ??????? ????? ? ? ??????????
????????? ??? Pinus ????? ? ? ????????????? ????? ????????? ??????. ??????? ? ?????
????? ????????????? ? ????? ????????? ??
???? ?????????, ??????????????? ???????????. ???? ? ??? ????????, ??? ????????????? ???????????? ?????? ? ?????????????,
??????? ????????, ? ??????? ??? ?????????? ???????????. ????????????? ???????????? ???????? ?? ?????? ?? ????????? ??
??????, ?? ? ?? ?????? «?????????»: ?? ?????????? ?? ?????? ?????? ???????????? ?
?????? 10 ??? ?? ?????. ????????, ??? ???
???????????? ???????? ? ????????????? ??????????????? ? ?????????????????? ??????
??? ??????????? ??????????? ??????.
????? ???????, ???????? ????????? ????????? ?????? ???????????? ???????? ????????????? ???? ?????????? ????? Pinus ??
?????? Cembrae, ? ???????????? ????????
????? ?????? ? ????????? ????????????? ??????? ???????? ?????????????? ????????? ????????? ??? ????????? ????????
? ????????????? ? ???? ????????. ????????? ????????????? ????????? ????????? ?????? ? ??????????? ??????????????????? ?????????????? ??????? ? ???? P.
koraiensis ? P. sibirica ? P. pumila ????????
???????? ?????????????? ?????????????
???????????? ???????, ???????? ???????
??????, ?????????? ????????? ? ????????? ?????????????? ????????, ??????????
????? ?????????????. ???????? ????????
# 75 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
????????? ????????? ?? ??????? P. pumila
???????????????? ??????? ? ????????????
?? ????????????? ? ??????????? ????? ???
???????????? ????????? ????? ???????.
?????? ????????? ??? ????????? ?? ??? (?????? ? 6.3.1.16; 7.10.1.3, ?????????????? ?????? ? 53), ?????????? ??? (?????? 16.10 ? ???? (??????? ? 07-04-00593)
?????? ??????????
????????? ?. ?. (1971) ???? ????? ???????? ??????? ? ????????? ????? ????? ?? ???????? // ???. ??. ?????. ???? ?? ????. ???. 78. ?. 52-57.
???????? ?. ?., ?????????? ?.?. (1989) ???????? ???????????????? ???????? ????????
???????? «????????» Pinus sibirica (Pinaceae) // ???. ????. 74: 554-557.
???????? ?. ?., ?????????? ?. ?. (1991) ???????????? ????????? ????? ????? ??????????
// ???????????.. ? 1. ?.42-48.
????????? ?. ?., ????????? ?. ?. (1992) ??????????????? ??????????? ??????? ???????
????? ????????????? // ???????????. ? 3. ?. 61-68.
?????????? ?. ?. (1992) ??????????? ????????????? ? ????????????? ???????????? ? ????? ??????? ??????? ????? ?????????? ? ?????????? ???????? // ???????? ???. ???. ????. ? 3. ?. 368-377.
?????????? ?. ?. (1994) ? ??????????????? ????????? ? ???????? ??????? Pinus sibirica
(Pinaceae) // ???. ????. 79: 63-71.
?????????? ?. ?. (1998) ? ????????? ????? ???????? ? ????? ?????????? ?? ??????? ??????? ??????????????? ? ????? // ???????? ??????? ?????: ???. ????. ????????. ????. ?????????. ?. 19-20.
?????????? ?. ?., ????????? ?. ?. (1992) ? ??????????? ????????????? ? ?????????????
???????????? ? ????? ?????????? // ?????????. 23: 268-276.
?????????? ?. ?., ????????? ?. ?. (2000) ????????? ????? ????? ?????????? (Pinus
sibirica) ?? ???????????? ????? ?????????? //???. ???. ????. 2: 110-122.
?????????? ?. ?. (1956) ???????? ???? ???????? ???????. ?.-?., 261 ?.
?????????? ?. ?., ??????? ?. ?., ??????? ?. ?. (1990) ?????????? ???????????? ?????????????? ???????? ????? ??????? // ????????. ?. 26. ? 4. ?. 694-707.
?????????? ?. ?., ?????????? ?. ?. (1974) ?????? Cembra ???? Pinus, ?? ????? ? ????? ??
???????????????? ?????? // ???. ????. ?. 59. ? 5. ?. 630-664.
?????????? ?. ?. (1974) ??????????? ???????? ?????? ????????? ???? ? ????? Cembrae
???? Pinus // ???. ????. ?. 59. ? 10. ?. 1501-1505.
??????? ?. ?. (1964) ???????? ?????? ?????. ?.-?., 88 ?.
???????? ?. ?. (1958) ????????? ??????????? ????? ? ???????????? ???????? ????? ?
?????? ????????? ???????? ? ????? ?????? //??????? ????????? ????????? ??????? ?? ??
????. ???. 1. ?. 96-103.
?????? ?. ?. (1971) ? ??????????? ????? ?????????? //???????????. ? 4. ?. 27-36.
???????????? ?. ?. (1960) ? ???????? ????????? ???????? (Pinus pumila Rgl.) ?? ????????
// ????. ????. ?????? ?????. ????. ?????. ? 3. ?. 136-141.
?????????? ?. ?. (1975) ???????? ??????? ?????? ?????????? ????????? ???????????.
?., 203 ?.
# 76 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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????? ????? ? ???????-????????? ???????. ??????????, 288 ?.
????????? ?.?. (1987) ??????? ?????????????. ?.: ?????. 440 ?.
???????? ?. ?.(1963) ??????? ?????? // ???????????? ???. ?., ?. 217-229.
???????????? ?. ?. (1995) ???????? ????????? ???????? (Pinus pumila (Pallas) Regel) ??
????????. ???????????, 227 ?.
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?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?????? ?. ??????????, ????????? ?. ?????. ?????????????? ? ???????? ??????? ? 5-??????? ??????
Shoot Morphological Structure and Development
in 5-Needle Pines of Northern and Eastern Asia:
Phylogenetic and Climatic Interpretation
Sergey N. Goroshkevich and Alexandr G.Popov
Institute of Monitoring of Climatic and Ecological Systems,
Siberian Branch of the Russian Academy of Science
10/3 Akademichesky, Tomsk, 634055 Russia
Comparative analysis of juvenile and adult shoot structure in 5-needle pine species of Northern and
eastern Asia are realized. The more the duration of vegetation period and sum of effective temperatures
in natural habitat, the later shoot growth in seedlings begins and terminates. On this point, examined
species are neatly divided into 4 groups: (1) subalpine-subarktic P.pumila, (2) boreal- mountain taiga
P.sibirica and P.cembra, (3) nemoral species of monsoon forest (P. koraiensis and P.parviflora),
(4) subtropical P. armandii. Differences between species by juvenile shoots were not related with
climate in its natural range. There are 3 pairs of species were distinguished by duration of juvenile
morphogenesis and presence of the mature morphostructures on juvenile shoots: Pinus sibirica + P.
cembra, P. koraiensis + P. armandii, P.parviflora + P.pumila. The presented dates adds argument in
favor of polyphyletic hypothesis of modern subsection Cembrae origin. The morphological structure
of adult shoots in 12 contrasting climatic ecotypes of three Russian Pinus species from Cembrae
group (P. sibirica, P. pumila ? P. koraiensis) was examined. In all species, male shoots produce one
(winter) bud and one (spring) elementary shoot during the growing season. Female shoots were found
to initiate two buds and two elementary shoots during the growing season. At the beginning of the
season a temporal bud is initiated giving rise to the summer shoot, which is formed during the season
without any period of dormancy. Then a winter bud is formed producing the first (spring) elementary
shoot in the beginning of the next season. There are short shoots, dormant buds, and lateral long
shoots on each elementary shoot. Differences in annual shoot structure between species and ecotypes
were showed and discussed.
Key words: structure and development of juvenile and adult shoot, 5-needle pines of Northern and
Eastern Asia, evolution, adaptation.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 80-89
~~~
??? 581.132. 1. 541.144: 582.475.4
????????? ? ??????? ???????????????? ????????
????? ?????????? ? ????? ???????????? ?????
?.?. ????????*, ?.?. ??????
???????? ??????????? ?????????????
? ????????????? ?????? ?? ??? ,
634055 ??????, ?????, ??. ????????????? 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
??????????? ???????????????? ?????????, ?????????? ????????????????? ????????? ?
?????????????? ?????????? ???????????? ?????????? ???? ????? ?????????? (Pinus sibirica
Du Tour), ??????????????? ? ??????????? ????? ??????? ????? ?? ??????? 1570, 1710, 1900 ?
2000 ? ??? ??????? ????. ? ???????? ? ???? ???????? ?????????? ????? ? ???????????????
???????????, ? ????? ????? ????. ?? ??????? 2000 ? 1900 ? ????????? ??????? ??????? ?
??????? ????. ???????? ?????????? ?????????? ??????????? ? ???????????? ?? ?????????
???????, ???, ????????, ??????? ? ???????????? ?? ?????????? ??? ?????? ??????????? ?
??????????? ? ????????????? ????????? ????????. ? ???? ????????, ?????????????? ? ???????
????? ??????? ??????? ????????? ?????????? ????????-???????????? ??????? ?? ??????
??????????? 2 ? ?????????? ?????????????? ????????????????????, ? ????? ??????????
????????????? ?????????. ???????? ????????????? ????????????????? ??????????
?? ??????? ??????? ????, ????????, ???????? ?????????? ????????????? ???????????,
??????????? ????? ?????????? ????????? ?????????? ????????????????? ? ???????????
??????????? ? ???????????.
???????? ?????: Pinus sibirica, ???????????, ?????????? ????, ????????????????? ????????,
?????????????? ?????????? ????????????.
????????
???? ?????????, ????????? ??????? ????????????? ????????????, ???????? ???????? ?????, ??????? ????? ?? ???????????????? ?????????????? ????????? ???????? ??
??????? ??????? ???? ? ??????????? ?????
??????? ?????. ??????? ??????????? ?????
??????????????? ?????????? ??????????????? ????????: ????????? ????????????
???????? ??2 ? ?2, ?????????? ??? ????? ?
?????????????, ??????? ?????????? ?????*
1
?????, ??????????? ?????????????? ??????? ? ??. ????? ??????????, ?????? ???????
???????? ???????? ???????? ? ??????? ?????
???, ?? ????????? ? ?????????? ?????????
????? ??? ? ??? ????, ????? ?? 60 ???? ??????????? ????????????????? ??????????????
??????? (??????, 1997; ???????????, 1998).
????? ????, ???? ????? ?????? ??????????????? ????????????? ????????????????
????????????? ??????: ????????? ????????
(?????? ? ?????????) ? ??????? ??????????, ?
Corresponding author E-mail address: zotik.05@mail.ru
© Siberian Federal University. All rights reserved
# 80 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
?????????????? ? ???????????. ??????????
?????????????? ???????? ? ??????? ?????????? ?? ???? ??????????? ?????????? ?, ???
????????? ?????, ?????????? ????? ?????????? ?????? (??????????, 1985). ?????????
???? ???????, ??????????, ????????? ???????????? ??????? ?? ???????????? ? ????????????????? ???????????? ??????????. ???
???????????? ??????? ?????????? ???????
????????????? ????????????, ????????? ? ???????, ???????????, ??????????? ? ????????? ??????, ??? ???????????? ????????????
?????????? ????????? ?? ?????? ??? ??????????????? ? ??????????? ??????????, ??? ?
???????? ??????????????? ???????.
? ???????? ????????? ????????????????? ????????? ???????? ? ???????? ????????
????? ???????? ???????????, ??????? ????????????? ? ??????? ????????????, ? ?????? ???????, ??????????????? ????????????
????????? ???????? ???????? ???????? ????????? ?????????????? ?????????, ????????????? ???????? ? ???????????? ???? ?????????? ???????? ? ?????????????? ????????.
??????? ????????????????? ??????????????
????? ??????????????? ??? ????????? ????????? ?????? ????????? ????????????? ?????????. ????? ????, ??? ???????? ??????????
???????????? ????? ???? ?????????? ????????? ???????????? ?????????? ? ????? ? ??????????? ????????? ?????????? ????????
?? ????????? ?????????.
????? ????????? ?????? ????? ????????? ????????? ???? ?? ?????? ??????????,
? ????? ??????????? ????????? ? ?????????????? ?????????? ???????????? ????? ??????????, ??????????????? ? ???????? ???????????.
????????
??????? ????????????. ????????????
????????? ? ??????? ???? ??? ? ???????????
????? ??????? ????? ? ?????? ??????????
??????. ????? ????? ? ???????????? ???????? ????? ?????????? (Pinus sibirica Du
Tour) 60-80-??????? ???????? ?? ????????????????????? ???????????? ????? ? ????????,
?????????????? ?? ????????? ???????.
??? ???? ???????? ???????? ??????? ??????? ?? 4-5 ??????? ?????? ?? ??????? ?????
????? ? ??????? ???????? ?????? ???????
???????. ????? ???? ???? ?????????? ??
??????? ????????, ????????????? ?? ???????: 1570, 1670, 1900 ? 2000 ?????? ???
??????? ????.
??????????? ????????????????? ?????????. ??? ??????????? ????????? ?????
??????? ????????????? ????????? ??????????? 96 %-??? ???????? ???????. ???????
????????? ????? ???????, ????? ????????
?????????? ????????? ????????? ????????
?? ????????? 0,6 ??. ?????????? ????????? ?????????? ?????????????????????
(Spectrophotometer UV-1601PC Shimadzu) ??
???????? ?????????? ????????? ??? 665,
649 ? 440 ?? (????, 1971). ???? ??????????, ????????? ? ??????????????? ????????
(???), ???????????? ? ?????? ???????????
????????? a/ ????????? b, ??????? ??? ???
1,2 (????? ? ??., 1988).
??????????? ?????????????? ?????????? ????????????. ??????????? ??????????? ?? ???? ?? ???????? (West, Wiskich, 1968),
???????????????? ??? ??????? ????????
(???????? ? ??., 1987). ???? ????????? ? ????? ????????? ?????????? ???????: 0.05 ?
????-HCl-????? ?H 8,0, 0,4 ? ????????, 1 ??
????, 1 ?? ?g?l2, 0,5 % ????? ????????.
?????????????? ?????????? ????????????
?????????? ????????????????????? ? ?????????????? ???????????? ????? (??????????, ????????, 2003). ??????????? ?????
?????????: 100 ??? ????, ?? 7,4, 10 ???
NaCl, 10 ??? MgCl2, 3 ??? K3Fe(CN)6, 4 ???
# 81 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
???, 10 ??? KH2PO4. ??????? ???????? ???
????????? ?????? ????? ?????? ?????????????? 150 ??/?2. ???????? ??????????????
???????????????????? ????????? ?? ?????
??????????????? ???????, ??????? ?????????? ??????? ????? (??????????, ????????,
2003).
?????? ?????????? ????????????? ?
??????? ???????????? ???????? Excel ?
Statistica. ? ???????? ? ?? ???????? ????????? ??????? ?????????????? ???????? ??
??????? ????????????? ???????????? ? ??
??????????? ??????.
??????????
? ??????????? ?????? ?????????????
????? ?????????? ????????? ???????? ????????????? ????????????? ???????? ??????.
????? ? ??????????? ???? ??????????? ??
10?25 %. ???????????? ?????????? ???????? ????? ????? ????: ???, ?? ?????? 2000 ?
??? ???????? ????????? 25 % ?? ????????? ?
??????? 1570 ? ??? ??????? ???? (????. 1).
?????????? ??????????????? ??????????, ??? ??????????????? ?????? ???. 1, ?
???? ????? ???? ???????? ?????????? ?? ????????? ??????? ?????????? ???????????
? ???????????? ? ??????? ?? 1 ? ????? ????? ????. ?????????? ?????????? ?????????
???? ????? ?????? ?? ??????? 1900 ? 2000
? ? ?????? ????, ???, ????????, ??????? ? ????? ???????? ??????? ???????? ???????????
????????? ? ????? ?? ??????? ??????? ???????? ?? ????????? ? ???????. ????????
???????? ?????????? ??? ?????????? a, ???
? ?????????? b, ?????????????? ? 2 ????, ?
???????????? ? ? 1,5. ?????????? ???????
????? ???????????????? ??????????? ????? ?? ????????? ????????? ???????? ? ?????
????. ? ???????? ???????? ????? ????????
? ??????? ??????? ?????????? ?? ????????? 1,5. ????????? ??????????/??????-
????? ???? ??????? ???? ?? ??????? 2000
? ???????? 1900 ?, ??? ?? 1710 ? 1570 ? ???
??????? ????, ??? ??????????????? ? ??????
????????? ???? ???????????? ? ??????????
?????, ???????? ?? ?? ?????? ??????????
(????. 2). ??????? ??????????????? ?????????? (???) ? ?????????-???????? ??????????
?????????? (???) ????? ???????????, ???
?? ????????? ???????, ??? ? ?? ?????? ?????. ?? ??????? ??????? ???????? ????????
??? ? ??? ???? ??????????? ? ?????? ?????????????? ???????, ????? ??? ??????????, ??
??? ? ?? ????????? ????????, ??????????? ??
?????? ??????? (????. 2).
?????????? ???????????? ??????? ?????????????????? ???????????? ????? ? ?????????????? ???????????????????? ???????
????? ???????????? ????????? ? ????????????? ????????? ?????????????? ????????? ? ???????? ? ??????????? ?? ?????? ?????????????. ??? ????? ?? ????. 3, ????????
???????? ?????????????????? ????????????
? ?????????????? ???????????????????? ?
??????????? ?????? ????????????? ????????
??????????? ? ??????? ????? ?????????????? ??????. ???????? ??????? ?????????????? ??????????? ???????? ???????????,
?????????? ?? ????????, ??????????????
?? ?????? 1710 ?. ?? ??????? 1900 ? 2000 ?
?????????? ??????? ????? ? ??????????????
???????????????????? ? ???????????? ????
???????? ?? 20 ?? 50 % ? ????? ?? ????????? ? ??????? ???????? ????????? (1710
? 1570 ?). ???????, ??? ? ???????? ? ????
????????? ?? ?????? ???????? ???? ????????????? ?????????, ?? ????? ?? ??????????, ?
??? ??????????????? ?????????? ?????????
?/2? (????????? ???????? ??????????????
???????????????????? ? ???????? ??????? ?????) ?? ??????? ??????? ????????
(????. 3). ????? ???????, ?????????? ?????
?????????? ????????? ?? ?????????? ????-
# 82 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
??????? 1. ???????????????? ?????????? ?????????? ???? ????? ??????????, ??????????????? ??
????????? ??????? ?????????? ?????? ??????? ?????
?????? ???
??????? ????, ?
?????, ??
1570
58,3±0,6
???????
???????????
???????, ??2
0,44±0,02
???????
???????????, ??2
?????
1 ???????, ??
171,4±3,2
35,9±0,9
1710
55,2±0,8
0,39±0,02
160,5±3,6
33,2±0,6
1900
49,7±0,7
0,37±0,02
151,9±2,6
30,5±1,7
2000
45,2±0,5
0,34±0,01
129,7±2,3
26,5±0,3
????????? a
0,9
0,7
0,5
0,3
23.6
13.7
2.8
22.8
11.9
?????????? ?????????,
??/? ????? ?????
????????? b
0,45
0,35
0,25
0,15
23.6
13.7
2.8
22.8
11.9
????? ????????????
0,35
0,3
0,25
0,2
0,15
23.6
1570 ?
13.7
1710 ?
2.8
1900 ?
22.8
11.9
????
2000 ?
???. 1. ???????? ?????????? ????????????????? ????????? ? ?????????? ???? ????? ?????????? (Pinus
sibirica Du Tour) ?? ?????? ??????? ????????????? ? ???????? ??????? ?????, ??/? ????? ?????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
??????? 2. ?????????????? ??????????? ????????? ?????????? ???? ????? ??????????, ???????????????
?? ????????? ??????? ?????????? ?????? ? ?????? ?????, ????????????? ????????
???? ??????
????
??????????
?????? ??? ??????? ????, ?
1570
1710
1900
2000
???
1.07.
0,77±0,03
0,75±0,02
0,35±0,02
0,43±0,01
26.07.
0,70±0,02
0,84±0,03
0,54±0,03
0,55±0,01
9.09.
0,83±0,02
0,88±0,05
0,57±0,02
0,60±0,04
???
1.07.
0,23±0,01
0,31±0,02
0,12±0,01
0,15±0,01
26.07.
0,31±0,01
0,39 ±0,01
0,26±0,01
0,24±0,01
9.09.
0,50±0,01
0,390,01
0,29±0,01
0,23±0,02
????????? ?/
1.07.
1,84±0,03
2,12±0,07
1,98±0,06
1,97±0,08
????????? b
26.07.
2,15±0,05
2,11±0,09
2,25±0,09
2,16±0,04
2,31±0,05
2,05±0,12
9.09.
2,50±0,01
2,18±0,09
??????????/
1.07.
4,30±0,07
5,59±0,04
2,92±0,07
3,48±0,11
???????????
26.07.
4,61±0,08
4,98±0,08
3,79±0,12
3,95±0,02
9.09.
4,02±0,06
4,68±0,12
3,62±0,11
3,67±0,10
??????????: ??? ? ?????? ???????????????? ????????? ????????????; ??? ? ?????? ????????????????? ?????????? ?????????? ???????????.
??????? 3. ??????????? ?????????????? ?????????? ???????????? ?????????? ???? ????? ?????????? ??
?????? ????????????? ???????? ? ?????? ?????????? ?????? ??????? ?????
?????? ???
??????? ????, ?
1570
1710
1900
2000
????
??????
????
30.06
30.07
30.06
30.07
30.06
30.07
30.06
30.07
???????? ??????? ?????,
??? ????????????/
?? ?????????? ???
484,1±8,5
456,8±11,1
546,4±9,7
596,3±19,5
384,5±8,3
335,8±7,8
323,0±10,5
363,3±3,3
?? ??????????? II ? ???????????? ?????????? ???? ????? ?? ????????? ?????????.
? ?????? ????? ???????????? ?????????????? ?????????? ???????????? ??????????? ?????? ??????? ??????? ???????????
??????, ??? ?? ????????? ???????. ??????
?????????? ???????? ????????????? ????????? ?????????????? ???????, ??? ? ?????????? ?????????, ???? ???????? ? ?????
??????? ? ???? ????????, ?????????????? ??
?????? 2000 ? (????. 3).
????? ????, ? ???????? ?????????? ?????????? ??????? ??????????? ??????????
?????????????? ??????? ?? ?????? ?? II ??
??????? ????? ??????????? (???. 2). ??????-
???????? ??????????????
????????????????????, ???
??????? / ?? ?????????? ???
426,2±10,2
428,5±6,4
437,1±8,5
560,5±18,1
285,3±11,8
256,6±12,4
257,6±7,2
242,3±11,8
? / 2?
0,88
0,94
0,88
0,94
0,74
0,74
0,80
0,75
??? ????????? ??????????? ??????? ? 20-22
?? 2 0?, ??? ????? ?????????? ? ?????? ?????,
???????? ???????? ?????????? ????????????? ????????? ?????????????? ??????? ????? ??? ? 2 ???? ?? ?????? ??? 2000, ??? ? 1710
? ??? ??????? ????. ??? ???? ?????? ??????????? ?????????????? ????? ??????????
?????????? ?????????? ?????????? ? ?????????? ????????????????????, ?? ?????? 2000
? ??? ????????? ????????? ? 0,82 ?? 0,71, ??
?????? 1710 ? ? ? 0,96 ?? 0,87.
??????????
?????? ?????????? ?????? ??????????????? ? ???????? ? ???????? ? ???? ? ?????
# 84 #
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???????? ??????? ????? ??? ????,
??? ?? ??? ?/?? ?? ?
?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
500
400
300
200
100
0
1710
2000
?????? ??? ??????? ????, ?
???????? ??????? ????? ??? +
22°?
???????? ??????? ????? ??? + 2°?
???????? ???? ??? + 22°?
???. 2. ??????? ??????? ???????? ??????????? ??????? ?? ?????????????? ?????????? ????????????
?????????? ???? ????? ??????????, ??????????????? ? ?????? ?????: ?? ? ?????????; ?? ????????????;
? ? ??????; ???? ? ????????????? ????????????????????
?????????? ??? ???????? ????????????????
????????, ??? ? ??? ?????????????? ??????????. ?? ??????? 2000 ? 1900 ? ?????????
??????? ??????? ? ??????? ????. ??????? ????????, ??? ?????????? ?????????????????
????????? ? ?????????? ???? ????? ?? ????
??????? ???????? ? ?????? ???????? ????
??????????? ???? ?? ????????? ? ?????????, ??????????????? ? ????????? ???????? ?????? ?. ?????? (????????? ? ??., 2000).
?? ???????? ?????????????????? ????????
? ????????, ?????????????? ? ?????, ????????? ??????? ???????? ????????, ? ?????? ???????, ????? ?? ??? ????????????, ??,
???? ?? ????? ???????????, ?????????????,
??????????, ???????? ????? ?? ??????????.
???????? ????????? ??????????? ??????? ?
?????? ????? (???????????, 1998) ? ??????
???????????? ??????? ?????????????? ??????????? ???? ?? ?????? 1600 ? ??????????
?????????????? 14 0?, ?? ?????? 2000 ? ???
???????????? ? 9 0?. ????????????? ???????-
???? ??????? ?? ??????? ??????? ????????????? ????? ????? 5 0?. ?????????? ????????
?????????? ??????????, ????????, ?????
????????? ????????????? ???????????? ??
??????????, ? ????? ?????????? ?????????????? ????????? ??-?? ???????? ?????????
?????? ? ?????? ??????????????? ????????.
????? ????, ?????????? ???????? ??????????? ????????? ? ???? ????????????? ????????
(???. 1) ????? ???? ?????????? ??????????
??????? ????????? ? ?????.
?? ???????????? ??????, ???????? ????????????? ??????? (???????, ???????????
??????, ???????????), ??? ???????, ?????
?????????? ?????????? ??????????? ?????????? ????????? ? ??????? ?? ???????
???? ????? (?????? ? ??., 1989). ? ???????? ?
???? ?????????? ?????????? ??????????? ?
??????? ??? ??????????? ????, ??????, ???????? (Tranguillini, 1979; James et al., 1994).
???????????? ?????????? ???????????? ? ??????????? ??????? ???????? ??? ? ????????
# 85 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
(Maslova, Popova, 1993), ??? ? ? ????????? ?
??????????? ?????? ????????????? ????????
(Todaria, 1988). ???, ??-????????, ??????????? ?????????????????? ? ??????? ????????????? ??????? ????????????? ? ?????.
?? ?????? ?.?. ???????? ? ?.?. ?????????
(2003), ???????? ????????? ??????????? ??????????? ????? ? ?? ?????????? ??????????
????? ???????? ?????????, ????? ?????????
?????, ????????????? ???????????? ? ???????? ???????? ???????? ?????. ?????????
???? ???????????? ? ?????????? ?????????
???? ??????? ? ???????? ???????? ? ??????????????? ???????????????, ???????????
??????? ? ????? ?????????? ?????????? ??????? ?????????, ?????????? ??? ????? ? ?????????????.
???????? ?????????? ????????? ?
?????????? ???? ????? ?????????? (???. 2) ?,
??? ????????? ?????, ????????? ??????????
?????????????? ?????????? ????????????
(????. 3) ? ??????????? ? ????? ????, ????????, ??????? ? ???????? ?????? ? ?????????
???????????????? ???????? ???? ????????
????. ? ????????????? ???????? ????????????? ? ????????? ?????????????? ??????? ????????, ?????? ?????, ????????? ????????????
???????????? ?????????????? ???????????
? ??????? ????????????? ? ?????? ??????
???????????????? ????????. ??????????? ?????????? ????????????????? ????????? ??
???????? ????????? ? ??? ????????? ? ?????
???????????? ? ???????? ?????? ???? ???????? ??????? ???????? (?????????, ????????????, 1981).
????????????????? ???????? ? ????????
? ???????? ??????????? ??????????? ??????? ??????? ?? ????????? ?????????????
??????????. ??????? (Todaria, 1988), ???????
???????????? ?????? ?? ???????????? ?????????, ????????, ??? ??? ??????????? ?????
??????????? ???????? ???????? ???????????
???? ????????? ????????????? ???????????
?? ????????????? ????????? ???????. ??????? ?????? ??? ? ?????????, ??? ? ? ????????? ??????????????? ???????????? ????????? ????????, ?????????? ?? ??????????
?????????? (Tranguillini, 1979; Todaria, 1988).
?? ???? ????? ???? ???????? (Pisek, Winkler,
1958), ??? ????????????? ?????????? ????????? ???????? ???? ?? ???????? ??????? ????
??????????? ????, ??? ? ????????, ????????
? ????????? ????????. ????? ?? ?????? ????? ????????, ?? ?????? ???????, ????? ????
?????????? ?????????? ?????????? ?? ??????? ???????.
??????????? ???? ???????????? ????????, ??? ?? ??????? ??????? ???????? ???????????? ? ??????????? ?????????? ???????
????????? ? ???????? ??? ? ??? ?????????
?????????????? ?????????? ???????????? ?
???? ?????. ??-????????, ???????? ????????
?????? ?????????? ???????????? ??? ??????????? ????????? ????????????? ????????
??????? ????? ???????? ?????? ??????????
????????? ? ?????? ????????-????????????
???? ?? ?????? ?? ?? II (??????????? ??????? ??????????? II). ?????????? ?????????????????? ???????????? ? ?????????????? ???????????????????? ???? ?????????
???????????? ? ????????? ? ??????????? ?????????? ? ?????????? ??????????? ???????
??????????? II ? ?????????? ?????? ????????????? ?????. ???????? ?? ??????????????? ????????????? ? ?????? ????????? ?????
??????????????? ?????????? ????????????
? ????????? ?????????? ???????? ? ???????? ??????????. ??? ??????? ? ??????????? ???????? ?????????? ? ??????? ?, ????????,
?????? ?????????????? ??????????? ???????
??????????? ???????????.
?.?. ????????? ? ?????????? (Tselniker,
Chetverikov, 1988) ??????? ????? ? ???, ??? ?
???? ???????? ??? ? ?????, ??????????????
# 86 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
? ?????????? ???????, ?????????? ?? ?? I ?
?? II ???? ????? ??????? ? ???????????? ??????? ? ?????. ???????????? ?????????????? ?????????? ???????????? ???? ?????
???????????? ??? ?????? ???????????? ???????? ?????????? ??????????? ??????????
????????? ?? II ? ????????????? ????????? ????? ???????????????? ?????????? a
? ???????????????? ?????????? ?????????
(Ottander et al., 1995). ??????????? ????????????? (Tausz et al., 1995), ???????? ??????????? ???? ??? ?? ?????????????? (????????
?????????? ?????????, ?????????? % ??????????? ?????????, ????????? ?????????? ??????????????), ???????, ??? ???????
?????????? ??????????? ???????? ????. ??
????? ????????? ? ????????????? ???????
?????????? ???????????????? ???????? ??
????????????????? ??????? ? ?????. ???????
??????? ?????????? ?????????? ?? ???????
??????? ??????? ?? ????????, ???????? ? ????????????????? ????????. ???? ????????,
??? ???? ?????????? ????????????? ?? ??????????? ??????????? ????????????? ??????? ????????????????? ? ??-? ?????????
?? ????????? ? ??????, ??????????????? ?
???????, ?? ???????? ?????? ????????????
?????????????? (Rau, Hofmann, 1993).
?? ?????? ?.?. ?????????? (1977), ?????????? ???????????? ? ??????????????
???????????????????? ???????? ? ??????????? ?? ????????????? ????? ? ???????????. ??? ?? ??????????? ??? ???? ????????? ????????????? ?????????????? ?????
??????? ?????????????? ????????????????????. ??????????? ???????????????????? ???????? ?????????, ? ??? ???????
?????? ? ???????? ??????? ????????????
?? ??????????? ????????. ?????????????
?????????? ?????? ?? ??????????????
?????????? ???????????? ????? ?????????? ? ???? ????? ??????, ????? ????????,
??? ??????? ??????? ????????????? ?????
???? ?????? ? ???????????? ???? ??????????????? ????????? ??????????? ?????
?????????????? ??????????? ????????????
?????????? ?????????????? ???????????, ?
????????? ????????? ?????????????? ????????????????????.
????? ???????, ? ?????? ???????? ?????
? ??????????? ?????? ????????????? ?????
?????????? ??????????? ??????? ???????????????? ????????, ????????? ??????????
????????????????? ????????? ? ??????????? ?????????????? ?????????? ????????????
?? ?????? ??????????? II.
?????? ????????? ??? ????????? ????????? ?? ??? 6.3.1.16 «????????? ???????????? ? ??????????? ?????????? ?????: ???????????? ? ?????????????? ???????», ??????????????? ??????? ?? ? ??? ??? ? 53 ? ?????? ???? ? 07-04-00593.
?????? ??????????
???????????? ?.?. (1996) ????????? ?????????????????? ???????? ? ?????????????? ????????? ???? ??????????? ?????????? ?????? //?????????? ????????. 43. ? 3: 391-398.
????????? ?.?., ?????? ?.?., ???????? ?.?., ????????? ?.?., ??????????? ?.?., ????????
?.?. (2000) ?????????? ??????? ???????-?????????????? ????????????? ??? ?????????????? ???????????? ?????????????????? ???????? ???????? //?????? ????????? ? ??????. 13. ?2:
539-542.
?????????? ?.?., ???????? ?.?. (2003) ??????? ????????? ?? ???????????. ?.: ????????,
256 ?.
# 87 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
????????? ?.?. (1977) ???????-??????????????? ??????? ???????? ??????????? //XXXVII
????????????? ??????. ?.: ?????, 56 ?.
???????? ?.?., ??????? ?.?., ???????????? ?.?., ?????? ?.?., ???????? ?.?. (1987) ???????????? ? ?????????????? ???????? ???????????? ?????????? ?????, ?????????? ?? ?????
? ? ??????? //?????????? ????????. 34. ???.3: 445-451.
?????? ?.?. (1997) ??????? ?????????? ?????. ???????????: ???-?? ??????????????
???. ??-??. 177 ?.
?????? ?.?., ??????? ?.?., ?????? ?.?. (1989) ??????????? ??????????? ???????? ???????? ????????? ????????-?????????????? ??? //???????-??????????????? ???????????? ??????????? ? ??????? ????????. ?.: ?????, ?.115?140.
??????? ?.?., ????????? ?.?. (2003) ???????? ???? ??????????? ?????????? ?????????
????? ???????? ?????????? ?????? ??? ????????? ? ???????????? ???????? //??????????
????????. 50. ? 1: 34-42.
????? ?.?., ?????????? ?.?, ?????????? ?.?., ??????? ?.?. (1988) ????????? ?????????
?????? ??????????? ??? ?????????? ???????????????? ????????? ???????? //?????????? ?
????????????? ??????? /??? ???. ?.?. ????????????. ?.: ?????, ?. 29-39.
?????????? ?.?. (1985) ???????????? ????? ?????? ????? ??????. ???????????: ?????,
123 ?.
??????????? ?.?. (1998) ?????? ???????????? ??????? ????? ? ????. ?????: ???-?? ???,
201 ?.
????????? ?.?., ???????????? ?.?. (1981) ???????? ???? ????? ? ??? //???????????????????? ?????? ????????? ?????????????? ??????? ????? ???????????? ??????. ?.:
?????, ?. 108-119.
???? ?.?. (1971) ??????????? ??????????? ? ???????????? ? ?????????? ??????? ??????? //????????????? ?????? ? ??????????. ?.: ?????, ?. 154?170.
James J.C., Grace J., Hoad S.P. (1994).Growth and photosynthesis of Pinus sylvestris at its
altitudinal limit in Scotland. Journal of Ecology. 82: 297-306.
Maslova T.G., Popova I.A. (1993) Adaptive properties of the plant pigment systems. Photosynthetica.
29 (2): 195-203.
Ottander C., Campbel D.,Oguist G. (1995) Seasonal changes in photosystem II organisation and
pigment composition in Pinus sylvestris. Planta. 197: 176-183.
Pisek A., Winkler E. (1958) Assimilationsvermoegen und Respiration der Fichte (Picea excelsa
Link) in verschiedener Hoehenlage und der Zirbe (Pinus cembra L.) an der alpinen Waldgrenze. Planta.
51: 518-543.
Rau W., Hofmann H. (1993) Sensitivitдt gegen UV-B Strahlung von Pflanzen verschiedener Arten
der gleichen Gattung aus unterschiedlichen Hohenstufen der Alpen. Wetter und Leben. 45, ? 4: 5-16.
Tausz M., Muller M., Bermadinger-Stabentheiher E., Grill D. (1995) Physiologische Methoden als
Erganzung zur Stre?diagnose an Fichten. Osterr Forstztg. 106, ? 11: S. 38-49.
Todaria N.P. (1988) Ecophysiology of Mountain Plants: 1 Photosynthesis. Acta Physiol. Plant. 10,
? 3: 199-226.
Tranquillini W. (1979) Physiological ecology of the alpine timberline: tree existence at high
altitudes with special references to the European Alps. New York: Springer-Verlag., 137p.
# 88 #
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?.?. ????????, ?.?. ??????. ????????? ? ??????? ???????????????? ???????? ????? ???????????
Tselniker Y.L.,Chetverikov A.G. (1988) Dynamics of chlorophull content and amounts of reaction
centres of photosystems 1 and 2 in Pinus silvestris L. and Picea abies Karst. needles during a year.
Photosyntetica. 22. ? 4: 483-490.
West K.R., Wiskich I.T. (1968) Photosyntetic control by isolated pea chloroplast. Biochem. J. 109.
? 2: 527-540.
Structure and Function of Siberian Stone Pine Assimilation
Apparatus in Central Altai Mounteens
Albina P. Zotikova and Olga.G. Bender
Institute for Monitoring of Climatic and Ecological Systems,
Siberian Branch of Russian Academy of Science
10/3 Academichesky, Tomsk, 634055 Russia
Needle morphology, photosynthetic pigment contents and chloroplast functional activity were
measured in the four Siberian Stone pine stands: the subalpine (1570 and 1700 m asl) and the timberline
(1900 and 2000 m asl). At the high altitudes the length, assimilation surfaces and mass needles were
decreased. At the timberline the needles had necrotic and chlorotic areas. Chlorophyll (a+b) and
carotenoid contents, chloroplast functional activity at the level of photosystem II: the Hill reaction
rate and noncyclic photophosphorylation were decreased with increasing elevation. Probably,
reduced pigment contents and low activity of photosynthetic process were induced by low temperature
and photooxidation. Due to morphological and physiological adaptation Siberian Stone pine trees
supported growth and reproduction ability.
Key words: Pinus sibirica, altitude, needle morphology, pigment contents, chloroplast functional
activity.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 90-102
~~~
??? 582.35/99+551.435.34+551.324.22(235.222)
??????? ???????????? ???????? ?? ??????? ???????
??????? ????????? (????-??????? ??????,
??????????? ?????)
?.?. ???????*, ?.?. ?????, ?.?. ??????????
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???
634055 ??????, ?????, ??. ????????????? 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
?? ??????? ??????? ??????? ????????? ? ???????????? ???????? ????-???????? ??????
??????? ??????? ?????? ?????????? ???????? ? ???? ?? ?????? ?????? ?? ?????????. ?????????
???????? ?? ????????????? ???????????? ????? ????? ??????? ?????.
???????? ?????: ??????? ??????, ??????? ????????????, ?????????? ????????, ???, ??????????? ??????.
????????
???????? ???????? ??????? ???????? ??
?????????????? ??? ???? ???????????, ???
??????????? ??????? ??????? ?????????????
??????????? ?.?. ???????? (1964), ??????????? ?????, ????????? «??? ??????????
????? ???????? ????? ????? ???????, ???????? ????????????? ? ??????? ??????? ??
???????? ????????????? ????? ?? ?????? ??
??-??? ???? ??? ???????????? ?? ????????».
???????????? ?????????? ????????
?????, ?????????? ? ???????? XIX ?. (??????????, 1901; ??????, 1949; ???????, 1966;
????????, ??????, 1999; ????????, 2001;
????????, ???????, 2003 ? ??.), ??????? ?
????, ??? ? ??????????? ?????? ???????????
?????????? ??????? ? ??????? ??????????
????? ??? ???? ???????????? ????????????
??????????. ???, ?????????? ??????? ??*
1
???: ?????????, ???????????? ? ??., ??? ????????? ? ?????????? ??????? ????? ??? ??
2 ??.
?????? ?? ???????? ???????????? ???????? ?? ???????????, ?????????????? ???
????, ? ??????? ?????????? ???????????????. ??? ?????????????? ????????? ?????
?????????? ?? ?????? (??????????? ?????
??????-???, Cooper 1931, 1939), ? ???????????
?????? (Palmer, Miller, 1961), ?? ??????? ?????????? ?????? (???????????, 1960) ????????
???? ?????????, ??????? ???????, ????????
???????? ??? ???????????? ?????????????? ???? ?????????? ????????. ?? ?????, ??
?????? ?????? ???????, ?????????? ?? 28 ??
83. ? ?????? ?. ???? ( Ludi, 1945) ??? ?????
???????? ????????? ??????? (???????????
?????) ?????? ????? ?????? ??????????
???????? (141 ??? ? ?????? ?? 83 ????? ?
Corresponding author E-mail address: timoshokee@mail.ru
© Siberian Federal University. All rights reserved
# 90 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
34 ????????) ? ???????????????? ????, ? ????? ??????? ???????? Racomitrium canescens,
Polytrichum juniperinum, P. piliferum.
? ????? ????? ?????? ????? ???????????? ?? ?????????? ??????? ????????. ?.?.
???????? (1996) ??? ??????????, ?????????????? ??? ???? ? ?????-???????? ?????? ???????, ??????? ?????? 40 ????????? ?????,
???????????? ? ?????? 3-5 ??? ????? ?????????? ???????. ? ????? ??????? ????????????
???????? ?? ??????? ??????? ?? ?? ?????????????. ??????????? ????????? ??????? ????? ????????, ? ????????, ? ??????????????? ???????? ????? (??????-??????? ??????,
??????????? ?????) ???????? ? ????? ??????? (???????, ????????, 2002; ???????
? ??., 2003; ?????, 2006; ??????? ? ??., 2008;
Timoshok et al., 2003; Timoshok, Skorokhodov,
2006). ?? ????-??????? ?????? ????? ???????????? ????? ?? ???????????.
?????????????? ??????????
????-??????? ?????? ??????? ????? ?
???????? ???????????, ??????????? ? ???
??????? ?????????. ?????? ?????? ??????????? (????? ? ????????). ??????????
??????? ? ??????????, ?????????? ? ??????? ?????????, ?? ??????? 1700-1800 ? ???
??. ????, ??????????? (100-150 ??), ?? ??????? 2000-2500 ? ??? ??. ?. ????????? ???? ?
250-300 ?? (????????????????? ??????????
?, 1962). ? ??????, ???????? ???????? ?????
?????? ?????????? ??????? ??????????? ?????? ? ?? ????????? ?????? ?.?. ????????
(1961), ????? 1000 ??. ??????????????????
????????????, ??????????? ?.?. ??????????
(1987) ? ???????????? ????? ?? ?????? 2500 ?
??? ??. ?., ????????, ??? ?? ????? ??????????
??????? ?????? ?????? ????? ??????? ?????????????????????. ???????? ???? ??? ??????????? ? ???????? ?????????? ???????????,
??????? ? ??????? ??? ???? ? ???????? ????,
? ????, ????? ????????? 38-40 є?. ?????????????? ??? ?????????????? ? ????? (????????, 1987). ? ??????? ????? ???????? ?????
(2100-2500 ? ??? ??. ?.) ????????? ??????????? ?????????? ???????????????-????????
??????. ? ?????? 2500 ? ???????? ??????????? ???????? ??????????? ?????????? ???????????? ?????????? ?????? ? ???????????
??????? ??????????, ???????? ???????, ????? ??????? ? ???????? ? ?????????? ??????????? ? ??????????? ?????. ?????? ???? ??
???????. ?? ?????? ??????? ???????? ?????
???????????? ????????? ???????? ???????????? ??????????? (Juniperus pseudosabina,
Lonicera hispida, Betula rotundifolia), ?????????? ??????????????????? ?????????
? ??????????? ???????????-???????? ??????. ? ???????????? ??????? ????????????? ????? (???? 2700 ? ??? ??. ?., ????????,
1987) ?????? ?????????????? ???????????
?????????? (?????????? ??????????? ????,
??????????? ???????????? ?????????????,
??????????-????????? ??????). ?????? ??????? ???????????? ?????????????? ?????????? ????? ???????? ?? ??????? ??????? ?
?????? ? ?? ?????? ???????? ? ??????????
???????.
??? ???????? ???? ??????????, ? ??????????????? ???? ? ?????? ?. ????? ????????????: ???????????? ?????? ? ????????
?????? ?? ???????? (?obresia myosuroides,
?. simpliciuscula), ???? (Carex ensifolia, C.
atrofusca) ? ??????? ?????? ?? Drepanocladus
aduncus ? Sanionia uncinata; ??????? ?? (Salix
pseudopentandra, S. sajanensis) ? ???????????
????????? ???????? ???????? ?? Astragalus
austrosibiricus, Leymus dasystachis, ?obresia
myosuroides, Potentilla multifida, Oxytropis
alpina, Leontopodium ochroleucum, Saussurea
schanginiana; ?????????????????? ?????????????? ?? ?????????? ? ???????????? ??????? ? ??????? ??????? ???????? ? Carex
# 91 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
altaica, C. atrofusca, ?. media, C. mel?nantha,
Hordeum brevisubulatum, Elymus schrenkianus,
Calamagrostis macilenta, Alopecurus alpinus,
Poa attenuata, Festuca rubra, Kobresia
simpliciuscula, Oxytropis deflexa, Scrophularia
incisa, Pedicularis abrotanifolia, Pedicularis
tristis, Galium ruthenicum, G. mollugo,
Dracocephalum origanoides, Saxifraga hirculus,
Artemisia dolosa, Matricaria ambigua. ?? ?????? ?????? ?? ?????? ?????????? ???????
? ???????? ??????? ??????? ??????? ??????????? ??????? ?????????? ? ???????? ??????? ? ???????????? ????? (Artemisia dolosa,
A. dracunculus, A. ?haeolepis, Seseli libanotis, S.
condensatum, Potentilla nudicaulis, P. bifurca,
Veronica porphyriana, Youngia tenuifolia ?
??). ? ??????? ???????????? ???????????
(Juniperus pseudosabina, Betula rotundifolia).
?? ????????? ?????? ?????? ?? ??????? ????????? ??? ? ?????????? ???????? ?????
???????????. ? ????????????? ???? ???????
??????????? ???, ???????? ????????? ?
?????????? Saxifraga hirculus, S. oppositifolia.
?? ?????? ??????? ? ?????????? ???????, ??
????????? ????????? ? ???????? ??????????? ??????? ????????????? ?????????? ? ???????? ?? (Salix pseudopentandra, S. sajanensis,
S. glauca ? ??.), ??????? ?????????????
(Betula rotundifolia), ?????????????? ?????????? ? ?????????????? (Juniperus sibirica ?
J. pseudosabina), ????? ??????? ???????????
????????? ???????????.
???????? ?????? ????????? ?????????? ? ???????????, ???????? ???????????
? ??????? ????????? ????? ????????? ??????????? ????-???????? ??????, ? ??????????????? ???????? ????? (???. 1).
???????? ???????? ????? ??????? ???????? ? 1898 ?., ????? ??? ?????? ?????????
?????? ?. ????? ????????? ????? ???????
???????????? ????????? ???????? ?????????????? ???????????? ?.?. ??????????.
???????, ? 1939 ?., ?????????, ???????????
??????????? ???????? ???????????? ?.?.
????????, ????????, ??? ?? ????????? ??
??????? ?????????? ?.?. ??????????? ?????? (41 ???) ?????? ???????? ?? 1100 ? ??
??????? ????????? 26 ? ? ???. ?? ??????
?.?. ????????, ?.?. ???? ? B.C. ????????
(1967), ? 1963 ?. ???????? ?????????? ???????
??????????? ???????????: ? 22,5 (?? 1951 ?.) ??
10 ? ? ???. ?????????? ????????? ?????????,
???????????? ????????, ? ???? ?????? ??????
????????. ??? ??????? ?????? ???????? ??????? ?????????, ??????????? ?.?. ???????? (2001), ? 1850 ?? 2000 ??. ?????? ????????
?? 2710 ?, ?? 102 ???? (1898-2000 ??.) ? ?? 1910
?. ????????????? ???????? ??? ?????????? ?????????? ????? 18,7 ? ? ???. ??? ????? ???????????? ???????? ??????????? ???????
???????????? ?????????? ?????? ??????????? ????? ????? 70 ? ?? 1850-2000 ??. ????? ?????????? ??????? ??????? ?? ????????? 150
??? ????????? 10,3 % ??? 1,94 ??2.
? ????????? ????? ????????? ??????? ????????? ??????????? ????? ??? ?? 2
??. ??????? ?????? ????? ??????? ??????
???????? ? ??????????? ??????? ??????
????? ? ?????????? ?????? ??????? ? ???????????????? ?????? ???????? XIX ?.,
????? ??????-????????? ??????????. ???
??????? ??????????? ??????????????? ???????? ???????????????? ??????????? ?????????. ??????? ?????? ????? ???????
??????, ? ??????? ???????????? ??????????? ???????? ? ???????????????? ???????
? ??????? ??????????? ??????? ? ???? (??
6 ? ? ????????); ???????? ?????? ???????
?????? (25-30°) ?? ??????? ????????????:
????????? ???????, ?????????? ?????? ?
???????????, ?? ??????? ????????? ?????????; ??????? ? ??????????? (????? 1 ??
????? ? 300-400 ? ??????) ???????? ????????? ????, ?????????? ??????? ?????
# 92 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
???. 1. ??????????????? ??????? ????? ? ?????? ????????? (???? ?.?. ???????)
??????????? ???????? ?? ????? ??????. ?
?????? ????????????? ?????? ??????? ???
?????????? ?????? ??????????? ??????.
??????????????? ??? ????? ??????? ???????? (??????? ? ??., 1967). ? 2000 ?. ?????
????? ??????? ????????? ?? ?????? 2484,5 ?
(????????, 2001).
???????? ???????????? ????????????? ???????? ????????????? ???????? ?.?. ????????? (1960), ???? ? ?.?. ????????? (1974).
????????? ???????? ?????????? ???????? ?????????, ? ????????, ?? ?????? ?.?.
?????????? (1995), ??????????????? ???? ? ??
????????? ? ????????
?????????? ? ?? ??????????
?? ??????? ??????? ??????? ?????????
????????? ???????? ??????????????? ???????????? ???????? ??????????? ???? ? ??????-
?? ??????? ??????? ??????? ????????? ???? ???????? 126 ????? ??????????
???????? ?? 71 ????, 32 ????????; 18 ?????
??????????????? ???? ?? 11 ?????, 10 ????????. ????? ????, ?? ????????? ? ???????? ????????? ???????? ????????? ?? ?????
Cladonia, Peltigera, Stereocaulon.
?????? ????? ?????????? ????????
??????????, ??? ??? ????? ????????? ???????????? ?????????? ?????????? ? 107
????? (93,8 %). ? ?????? ?????????? ???????? ????????? ?????? ???? ?????. ?????
??? ? ???????????????????, ?????????????? ?????? ???????? Woodsiaceae (Woodsia
ilvensis), Athyriaceae (Athyrium distentifolium,
Cystopteris fragilis, Gymnocarpium dryopteris).
????? ????????? ????? ???????????? ????-
???????????? ??????????, ?????????????
?? ?????? ???????? ?? ???????, ??????? ??
????? ?????? ????? ? ????? ???????, ?????????? ???????-???????? ?????. ??? ??????? ????? ??????? ???????? ???? ?????????? ?????????????? ??????, ??????? ?????
200 ?????? ???????? ?????????? ???????? ?
50 ??????? ????. ?????? ???????? ?????????
????? ? ????????????? ????????? ????????
? ??????????????? ?????????? ??????????
(????????, 1950) ?? ?????? ??????? ?????????
??????? ?????, ??? ?????????? ??? ???????
????????? ???????? ????????????. ??? ?????????? ?????????????? ??????? ??????????
?.?. ????????, ?.?. ????????? (2003, 2004).
# 93 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
????????? ?????????????? ?? ?????????
Huperziaceae ? Huperzia selago. ???????????? ???????? ?? ??????? ??????? ???????
????????? ???????????? ?????? ????????
???????? ? Pinaceae (Larix sibirica, Picea
obovata) ? ??????????? ? Cupressaceae
(Juniperus sibirica).
? ??????????????? ??????? ?????????
???????? ? ??????? ?????????? (? ???????
?????? ? ?????), ????????? ?????? ?????? ?????????: Asteraceae (23 ????, 17,9 %), Salicaceae
(19 ?????, 15,5 %), Poaceae (13 ?????, 10,2 %),
Caryophyllaceae (10 ?????, 7,0 %).
????????? ????????????? ? Asteraceae
?????????? ?????????? ??????? (13 ?????) ? ??????? ?????????????: Antennaria
dioica, Artemisia dolosa, Aster alpinus, Crepis
crocea, C. karelinii, C. multicaulis, C. nana,
C. polytricha, Erigeron altaicus, E. eriocalix,
E. politus, Leontopodium ochroleucum, Matricaria
ambiqua,Pyrethrum
pulchrum,
Solidago
dahurica, Taraxacum altaicum, T. officinale, T.
pingue, Tephroseris heterophylla, T. praticola,
T. pricei, Trommsdorfia maculat?, Waldheimia
tridactylites.
?????? ????? ???????? ????????? ?????? ? Salicaceae ? 2 ?????? ? 19 ??????. ?????
?????????????? ????? ????????? ???????????
???? ???? ???: Salix bebbiana, S. berberifolia,
S. coesia, S. coesia subsp. tschujensis,
S. divaricata, S. glauca, S. hastata, S. nummularia,
S. pseudopentandra, S. pyrolifolia, S. rectijulis,
S. recurvigemmis, S. reticulata, S. sajanensis,
S. saposhnikovii, S. turczaninovii, S. vestita.
?? ??????? ????? ????????? ???????? ? Poaceae, ?????????? 5 ????? ? 13 ?????: Calamagrostis pavlovii,
Calamagrostis
pseudophragmites,
Elymus
schrenkianus, E. transbaicalensis, Festuca
altaica, F. kryloviana, Poa alpina, P. altaica,
P. attenuata, P. glauca, P. mariae, P. stepposa,
Trisetum mongolicum.
?? ????????? ????? ????????? ?????????? ? Caryophyllaceae, ?????????? 6 ????? ? 10 ?????: Cerastium lithospermifolium,
C. pusillum, Dianthus versicolor, Gastrolychnis
apetala, Minuartia verna, Silene chamarensis,
S. graminifolia, S. jenisseensis, Stellaria
dichotoma, S. peduncularis.
????? ????????, ??????? ????????
??????? ????? (4-7) ?????, ????? ???????? ???????? ? Cyperaceae (6,3 %, 2 ????)
? 7 ??????: Carex aterrima, C. capillaris,
C. eleusinoides, C. media, C. melanocephala,
C. sempervirens ? Eriophorum altaicum. ?
???????? ????????? ? ???????????? ????????????? ????????? ?????????? ? Juncaceae
(3,1 %, 2 ????, 4 ????): Juncus arcticus,
J. castaneus ? Luzula sibirica, L. spicata. ??
?????? ???? ???????? ?????????: ???????????? ? Crassulaceae (Orostachis spinosa,
Rhodiola algida, Rh. coccinea, Rh. rosea), ??????????? ? Rosaceae (Dryas oxyodontha,
Pentaphylloides fruticosa, Potentilla nivea,
Sibbaldia
procumbens),????????????
?
Scrophulariaceae (Euphrasia altaica, Lagotis
integrifolia, Pedicularis anthemifolia, Veronica
porphyriana).
??????????? ?? ????? ????????? ???????? ????- ? ??????????? ?????????: Fabaceae
(Lupinaster eximius, Oxytropis alpina, O. deflexa),
Betulaceae (Betula pendula, B. rotundifolia),
Brassicaceae (Draba cana, D. fladnizensis),
Grossulariaceae
(Ribes
atropurpureum,
R. graveolens), Pyrolaceae (Orthilia obtusata,
Pyrola rotundifolia), Ericaceae (Ledum palustre,
Vaccinium uliginosum), Gentianaceae (Ciminalis
grandiflora, Comastoma tenellum), Caprifoliaceae
(Lonicera altaica, L. hispida), Ranunculaceae
(Aconitum glandulosum), Papaveraceae (Papaver
pseudocanescens), Empetraceae (Empetrum
nigrum), Onagraceae (Chamaenerion latifolium),
Primulaceae (Primula nivalis), Boraginaceae
(Myosotis austrosibirica), Lamiaceae (Thymus
# 94 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
proximus),
Campanulaceae
(Campanula
rotundifolia).
????? ????? ?? ?????????? ????? ?? ?????? ????? ? ??? Salix ? 19 ??????, ?? ??????
????? ??? ???? ? Carex ? Poa (?? 6 ?????).
????????????? ?????? ???????, ???
????? ????????, ???????????? ?? ?????????????? ???????? ???????? ?????????,
??????????? ???????? ? ???? ???????? ????????????? ?? ?????? ??????????? ? 81 ???
(64,3 %). ????? ??? ??????? ????????? ???????? ?????????? ?????????? (50,6 %), ???
???????? ????? Festuca altaica, F. kryloviana,
Poa alpin?, Trisetum mongolicum, ?????
Carex sempervirens, C. melanocephala, ????? Luzula sibirica, L. spicata, ??? S. rectijulis,
S. reticulata, S. turczaninovii, S. vestita, ??????? Rhodiola coccinea, Rh. rosea, Betula
rotundifolia, Stellaria peduncularis, Lagotis
integrifolis ? ??. ???????????? ???? ????
?????? ?????????? ??????????????? (21 %)
P. glauca, S. recurvigemmis, S. sajanensis,
Oxyria digyna, Gastrolychnis apetala, Saxifraga
cernua, S. oppositifolia, Lonicera hispida, Crepis
nana, ?. karelinii, Waldheimia tridactylites
? ??.; ??????? ???? ? ??????????????
(14,8 %) Chamaenerion latifolium, Matricaria
ambiqua, Lonicera altaica ? ??. ? ??????????????? (13,6 %) Carex eleusinoides, Eriophorum
altaicum, Juncus arcticus, J. castaneus, Salix
saposhnikovii, S. hastata, S. nummularia,
Saxifraga hirculus, Primula nivalis ? ??.
??????????? ???? ??????? ?????, ??????????? ? ?????? ????????????? ???????:
?????????? (17 ?????, 13,5 %), ????????? (17
?????, 13,5 %) ? ?????????? (11 ?????, 8.7 %).
?????????, ???? ?????????? ?????????????, ????? ???????????? ?????????? ??????????? Poa stepposa, Stellaria dichotoma, Aster
alpinus ? ??.; ???????????????? Artemisia
dolosa, Pyrethrum pulchrum, Orostachis spinosa
? ??.; ??????????????? Larix sibirica, Poa
attenuata, Dianthus versicolor, Trommsdorfia
maculat?; ???????????????? Populus laurifolia
? Oxytropis deflexa. ?????????, ??? ? ???? ???????????? ???????? ??????????, ?? ???
?????? ????????????? ??????, ??????? ?
????????, ??? ????? ???????????????? ??
???????, ?????? ????????, ????????, ?? ?????????? ??????????? ??????. ????? ???
?????????? ????????, ? ??? ????? ??????
???? Betula pendula, Salix bebbiana, Pyrola
rotundifolia, Orthilia obtusata, Calamagrostis
pavlovii, Gymnocarpium dryopteris ? ??. ? ????????????? Poa altaica, Draba cana, ???????? ??????????? Cystopteris fragilis, Woodsia
ilvensis. ????????? ? ???????? ?????????
??????????? ?????????????, ??? ?????
????????? ?? ????? ?????, ??????? ???,
???????, ?? ??????? ???????, ???????????? ?????????? ??????????? Calamagrostis
pseudophragmites, Carex aterrima, C. capillaris,
C. media, Elymus transbaicalensis, Ledum
palustre, Salix coesia ? ??????????????? Picea
obovata, Salix pseudopentandra, S. pyrolifolia,
Vaccinium uliginosum.
??????????????? ??? ?? ??????? ??????? ??????? ????????? ???????????? ????????????? ????? ?? ???????? Bryaceae (Bryum
caespiticium, B. argenteum, B. lonchocaulon),
Ditrichaceae (Ceratodon purpureus), Grimmiaceae
(Racomitrium canescens), Mniaceae (Pohlia
filum, P. cruda, P. nutans, P. wahlenbergii),
Polytrichaceae
(Pogonatum
urnigerum,
Polytrichum juniperinum, P. piliferum), Pottiaceae
(Syntrichia ruralis) ? ???????????? ????? ??
???????? Amblystegiaceae (Campylium stellatum),
Bartramiaceae (Philonotis seriata, Ph. fontana),
Brachytheciaceae (Brachythecium salebrosum),
Scorpidiaceae (Sanionia uncinata). ????? ????
?? ??????? ????? ???? ? ?????? ?????????.
??????? ???????, ??? ??????? ???? ????, ???????????????? ?? ????? ?? ????? ?????????
??????? ???????: Brachythecium salebrosum,
# 95 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
Ceratodon purpureus, Pohlia wahlenbergii,
Pogonatum urnigerum, Polytrichum juniperinum,
P. piliferum, Racomitrium canescens, Sanionia
uncinata ? ??. ?? ????? ???????????? ? ???????????? ???? ???? Bryum criophilum, Philonotis
seriata, Pohlia nutans. ????? ??????????? ???,
????????? ??????????????? ?? ?????? ???????, Philonotis seriata, Racomitrium canescens;
??????????????? ?? ??????????? ? Bryum
lonchocaulon, ?? ??????????? ???? ????, ?????????????? ? ??????????? ?????????, ???????????? ?? ?????????? ?????, ?????, ?????
?????? ? ??.
????? ?????????? ???????? ? ???? ??
??????? ??????? ??????? ????????? ???????? ????????? ?? ????? Cladonia (C.
fimbriata ? ??.), Peltigera (P. polydactyla ? ??.)
Stereocaulon (S. vesuvianum, S. paschale).
??? ???????????? ?????????? ????? ????? ?.?. ????????? ? ???? 1973 ?. ?? ???????
????? ???????, ? 7 ? ?? ?????? ???? ????, ????
??????? ?????? ?????????: Chamaenerion
latifolium, Papaver, Oxyria digina, Matricaria
ambiqua, Stellaria peduncularis ? ??. ?????
??????? ????????? ???????? ??? ?????????
???????? ????? ???????? ?????????? ????????? ????, ?? ??????? ??? ? ??????????.
? ????? ??, ? ??? ??? ????? 30 ? ???????? ??
???? ???????? (????????, 1996).
????, ????? 30 ??? ????? ?? ??????????,
?? ????????? ? ??????? ???????, ????????? ???????????????? ??????????, ????
???????? ?????????? (??????? ??????????)
????? ????????? ???????? Chamaenerion
latifolium, Saxifraga oppositifolia, Waldheimia
tridactylites, ? ????? ?????????? ???? Bryum
cryophilum, Pohlia nutans, P. wahlenbergii,
Pogonatum urnigerum.
? ???????? ??????? ? ??????? ?????????? (50-350 ? ?? ????? ??????? ?????????)
???????? ??????????, ??????? ???????, ??
???????? ? ????????? ???????? ??? ???????
???????. ????? ????? ?????????? ????????
? ?????? ?????????? ?????????? ?? 8 ?? 27
(???. 2). ?????????? ????? ????? ? 27 ????????? ???????? ? 6 ????? ???? ? ???????? ??
?????????? 200-250 ? ?? ????? ???????.
??????? ???????, ????? ???????? ?????????? ? ????????? ?? ?????? ????????:
????????????? Artemisia dolosa, Crepis nana,
Waldheimia tridactylites; ?????????? Cerastium
lithospermifolium, C. pusillum, Gastrolychnis
apetala, Stellaria peduncularis, Minuartia verna;
????????????? Draba cana, Draba fladnizensis;
???????? Elymus transbaicalensis, Trisetum
mongolicum;
?????????????
Saxifraga
oppositifolia; ????????? Oxyria digyna; ?????????? Luzula spicata, L. sibirica; ?????? Salix
divaricata, S. glauca, S. hastata, S. sajanensis.
???? ?????? ????????????? ????? ???????????
????????, ???????? ????????? Calamagrostis
pseudophragmites, Carex aterrima.
???? ????? ? ???? ?????????? ??????????? Chamaenerion latifolium, ???????????
???????? ???????? ?? ????????? ????????
?????????? ????? 1 %. ?? ????????? ??? ????????? ?????????????? ? ?????????? ????????. ?? ????????? ???????? ?????????
?????????? ????????? ??????????????? ??????????????? ? ???????????? ?????? ?????????? Waldheimia tridactylites, Cerastium pusillum,
Gastrolychnis apetala, Draba fladnizensis. ??
?????????? ???????????? ??????????
???????? ???????? ???????: ??????????
Stellaria peduncularis, Trisetum mongolicum,
?????????????? Cerastium lithospermifolium,
???? ???????????? Elymus transbaicalensis.
?????????? ???????????? ????? ????????
??????????, ???????????? ? ????????????
????? ????? ? ??????? ???????????? ? ???????, ?? ??????????????? ??? ????? ??????- ?
????????? Crepis nana, Saxifraga oppositifolia,
Calamagrostis pseudophragmites, Carex aterrima,
Oxyria digyna, Draba cana, Minuartia verna,
# 96 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
70
60
50
40
30
20
10
0
58
46
41
47
56
52
29
?
80
090
0
?
95
010
00
?
75
0
?
70
0-
50
50
?
60
06
50
?
50
05
40
04
35
0
?
?
30
0-
?
25
0
0
50
10
01
30
-5
4 6 45
13 16
12 8 10
20
0-
3
51
31 3 3
27
?
?????????? ?????
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
?????????? ?? ???? ???????
???. 2. ?????????? ????? ?????????? ???????? ? ?????? ?????????? (? ??????????? ?? ?????????? ??
???????) ?? ??????? ??????? ??????? ?????????
Luzula spicata, L. sibirica. ? ??????????, ??
????????? ???????? ??? ??????? ??????? ?????????? ? ?????? ????????? ?????????????
??? (Salix glauca, S. divaricata, S. sajanensis, S.
hastata, S. recurvigemmis) ?? ?????? ????????????? ?????.
????? ?????????? 8 ????? ????: Bryum
caespiticium, B. criophilum, B. argenteum,
?eratodon purpureus, Pohlia cruda, P. nutans,
P. wahlenbergii, Pogonatum urnigerum. ??? ???
???????????? ???, ???????????? ?? ?????????? ??????????? ?????????, ?????????????? ?????? ?????? ???? ??????????????
? ?????????. ?????????? ???? ??? ????????? (1-4 ?? ? ????????). ??????? ? ??????
?????????? ???? ??????, ??? ???????????
?? ?????? ?????????? ???????????? ? ?????????? ?????? ?? ???????? ????????, ??
??????? ???????????? ????????? ? ?????????? ??????????. ????????, Bryum argenteum
? Pohlia cruda ? ????????? ? Luzula sibirica;
?eratodon purpureus ? Pogonatum urnigerum ?
? Luzula spicata ? Chamaenerion latifolium.
?? ????????? ????????? ??????? ????????
Stereocaulon vesuvianum.
?? ??????? ??????? ??????? ?????????
?? ?????? ?????? ?????????, ??? ? ?? ???????
???????? ???????? ???????? ????? (????????????? ??????) (??????? ? ??., 2002, 2003;
Timoshok et al., 2003), ??????????? ?????????
??????????? ????????. ??? ??? ????? ?????
??? ????????? ? ????? ?????? ?????????????
?? ???????. ?? ????????? ???????? ?????????? ?????????? ? ?????????? ???????????? ?? ???????????????, ? ????????, ????????
?????????.
???????? ??????? ??????? ????????????
???????? ?????????????, ??????? ? 400-450 ?
?? ???? ???? (???. 2). ???????? ???????? ?????,
? ????????, ??????? ?????? (????? ? ??????? 5-10°). ??? ??????? ???????????????
??????? ???????? ??????????, ?? ???????
??????? ? ??????????? ??????? ? ??????????????, ???????? ? ????????????? ??????????. ??? ?????? ????? ???????? ?????????
????, ??????? ????????? ??????????? ??????
???????? ?. ?????, ? ???????????? ?? ???
???????? ????????? ?????????.
?? ???? ???????? ?? ??????? ????? ????? ?????????? ???????? ? ?????? ?????????? ????????? ?? 31 ?? 58, ???? ? ?? 4 ?? 8 ?
??????????? ?? ??????????????????? ???????
???????? ?????????. ????? ??????????????
? ????????????? ???????? ?? ??????? ?????
????????, ??? ? ????? ??????? ? ???????
??????????.
?? ??????? ??????? ?????, ?? ???????? ? ??????????????? ?????????? (?????,
# 97 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
????????), ?? ???????? ??????????? ???????? ? ????????????????, ??????? ?????
??????? ????????? ??????? (???????? ??????? ? 3?8 ?), ???????? ?????????????? ???????, ???????? ?????? ?? ???????, ????????
?????????? ???????? (???? Chamaenerion
latifolium, Waldheimia tridactilites) ? ???
(Polytrichum piliferum, Pogonatum urnigerum,
Sanionia uncinata) (???. 3).
?? ???????????????? ????????? ??????????
???????????
Chamaenerion
latifolium, ??????????? ???????? ????????
? ??????? ?????????? 3-5 %, ?? ?????????? ?????????? ???????? ? 5-7 %, ? ?????????????????, ???????????? ? ??????????
????????? ???????, ????? ????????? 40 %.
????????? ? ??? ? ??????? ???????????????? ?????????, ??????? ???????, ??????????????? ????? Stellaria peduncularis, Poa
glauca, Trisetum mongolicum. ????????? ???????????????? ???????? ? ???? ???? Salix,
??? ???????????? ???-??? ????? ?????? ????? ?? ??????? 30-40 ?? ?? ???? ??????????
??????????. ????? ?????, ??????????? ?
??????? ????????? ????????????????, ??
??????? ????? ? ????????? ????????? ????????? ??????????????? ? ????????????
?????? ???? Erigeron altaicum, Antennaria
dioica, Leonthopodium ochroleucum, Festuca
kryloviana, ??????????? Cystopteris fragilis,
??????????? Empetrum nigrum, ??????????
Ribes graveolens. ????? ?? ??????? ???????? ????????? ??? ????? ??????? ?????
??????????? ????????? (Larix sibirica), ?????? ??????? ?? ????????? 50-60 ??. ??
????? ????????? ?? ??????? ??????????
?????????? ????? ???? ?? ?????? ??????????? Betula rotundifolia, Lonicera hispida,
Carex eleusinoides. ? ?????, ????? ???????? ??????? ? ????????, ? ???????? ?
??????????? ??????, ???????? ??????????? ???????? Pentaphylloides fruticosa,
Ribes atropurpureum, Athyrium distentifolium
? Gymnocarpium dryopteris, Calamagrostis
pavlovii ? ???????? Ledum palustre. ?? ?????
????? ?????? ??????? ???????? Stellaria
dichotoma.
??? ?????? ?? ???????, ??????? ? 600650 ?, ?? ?????? ????????? ????????? ??????? ????????? ???????????? ???????????????? ?????????? ????? ?????????????? ?
???????? ?? ???????? ???????.
? ???????? ??????? ?????, ?? ??????????????? ???????? ??????? ? ?????? ?????? ????????? ???, ? ?? ??????? ????????? ??????????????? (???. 4) Eriophorum
altaicum, Juncus arcticus, J. castaneus, Luzula
spicata, Carex eleusinoides, ???????? Carex
capillaris, ? ????? ???????????? ??? ?? ????????? ???????????? Polytrichum juniperinum,
P. piliferum, Pogonatum urnigerum. ?????????
? ???? ??????? Ceratodon purpureus ? ???
??????????????? ????????????? ? ??????????? ??????????? Philonotis fontana, Ph.
seri?ta.
? ?????????????? ? ????? ?????????
????????? ??????????? ????????????? ???????????????? ? ????????????? ?? Salix
sajanensis, S. divaricata, S. hastata, S. glauca,
Salix berberifolia, S. turczaninovii ?? ??????
????????????? ????? (???. 5). ?? ??????????? ?????, ??????????????? ??????????????? Stellaria peduncularis, Saxifraga sibirica,
Poa glauca, Trisetum mongolicum, ? ?? ?? ????
???????????? ???? (Pogonatum urnigerum,
Polytrichum juniperinum, P. piliferum) ????????? ? Brachythecium salebrosum.
?? ?????????? ??????????? ?????????
????????? ??????? ???????? ???????? ?????????? ??????? ?????? ?, ?????? ?????, ?????
???? ?????? (Poa glauca, P. altaica, P. alpina,
P. attenuata, P. mariae, P. stepposa) ? ????????
(Festuca altaica, F. kryloviana), ???????????
? ?????? ????????????? ???????. ? ??????
# 98 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
???. 3. ?????? ???????????????? ?? ???????
??????? ??????? ????????? (???? ?.?. ?????)
???. 4. ???????????????? ? ???????? ??????????
?? ??????? ??????? ??????? ????????? (????
?.?. ?????)
???. 5. ???????????????? ? ???????? ?? ?? ??????? ??????? ??????? ????????? (???? ?.?. ?????)
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
??????? ?? ??????????? Trisetum mongolicum,
Calamagrostis pavlovii, C. pseudophragmites,
Elymus schrenkianus, E. transbaicalensis,
Antennaria dioica, Leontopodium ochroleucum ?
??. ????? ???????? ??????? ??? ???????????
????????? ???????????? ??? ?? ????? Bryum
(B. caespiticium), Pohlia (P. cruda, P. nutans) ?
Ceratodon purpureus. ?? ????????????????
???????? ??-???????? ?????? Chamaenerion
latifolium, ??????????? ? ????? ?????????????? ????? ???????????, ????????? ?? ??????? ?????.
?? ???????? ???????? ??????, ??
?????????-????????? ???????? ? ??????????? ????? ?????????? ??????????? ????????? ????? ??? ????????? ????????? ??????????????? ?????? ??????? ??????????
Poa stepposa, Artemisia dolosa ? ??????????
Orostachis spinosa, Thymus proximus, Dianthus
versicilor, ? ????? ? ???????? ???????????????????? Rhodiola algida, Rh. ?occinea,
Dryas oxyodontha. ?? ???????? ??????????
???????? ???????????? ????????? Empetrum
nigrum, ??????????? ???????? ????????
???????????? ???????.
??????? ????? ???????? ???????????
????????? ??????????? ? ?????????? ???????????, ??????? ? 350-400 ?, ??? (Picea
obovata) ? ? 450-600 ? ?? ???????????? ?????
???????, ?????? ??????? ???????? ?????? ???????? (Betula pendula) ? ? 550-600 ? ?? ????.
??????
1. ? ????????? ??????? ????? ???????
????????? ????????? 126 ????? ??????????
????????, 18 ????? ????, ????????? ?? ?????
Cladonia, Peltigera, Stereocaulon.
2. ? ??????? ????? ????????? ??????????? ????????? ???????? (93,8 %). ???????? ??????????? ???????? Asteraceae ?
17,9 %, Salicaceae ? 15,5 %, Poaceae 10,2% ?
Caryophyllaceae ? 7,0 %.
3. ????? ????????, ???????????? ?? ?????????????? ???????? ???????? ?????????,
??????????? ???? ???????? ?????????????
?? ?????? ??????????? ? 64,3 %. ????? ?????
?? ?????? ????????????? ????? ???????????
????: ?????????? ? ????????? ? ?? 13,5 %,
?????????? ? 8,7 %.
4. ? ??????????? ?? ??????????? ?? ????? ??????? ? ??????????????????? ???????
???????? ????????? ?????????? ????? ?????????? ???????? ? ???????? ??????? ? ??????? ?????????? ??????? ????? ??????????
?? 3 ?? 27, ? ????? ????????? ? ?? 31 ?? 58.
?????????? ????? ??????????????? ???? ????????? ?????????????, ?? 4 ?? 9.
5. ??????? ?????? ????????, ?????????? ???? ?? ??????? ??????? ??????? ?????????, ??????????, ? ????????, ???????
????? ????????????? ? ???? ??????????
?????? ? ??????????????? ????????? ? ?????? ?. ?????.
?????? ??????????
????????????????? ?????????? ?? ?????-????????? ?????????? ??????? (1962), ?: ???????????????, ?. 84.
???????? ?.?. (1974) ??????????????? ??? ????? ? ????. ???????????: ?????, 319 ?.
??????? ?.?., ??? ?.?., ??????? B.C. (1967) ? ??????????? ?????????? ???????? ?. ?????
// ??????????? ?????. ???. 5. ?????: ???-?? ???????? ????????????. ?. 163-176.
??????????? ?.?. (1960) ??????????????? ?????????????? ???????? ???????? ??????? ?
?? ???? ? ??????????????? ???????? ?????? ?????????. ???. ????. ?. 45, ? 9: 1249-1258.
????? ?.?. (2006) ????? ??????? ????? ???????? ???????????? ?????: ???????. ???. ?
????. ????. ????. ?????, 19 ?.
# 100 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
??????? ?.?., ???????? ?.?. (2003) ????? ???? ??????? ????? ??????????? ??????. ?.:
Scientific Press LTD, ?.1, 608 ?.
??????? ?.?., ???????? ?.?. (2004) ????? ???? ??????? ????? ??????????? ??????. ?.:
Scientific Press LTD, ?.2, 960 ?.
???????? ?.?. (1960) ???????????? ?????? ?????. ???????????: ?????. ???. ???., 449 ?.
???????? ?.?. (1987) ?????????????? ????????? ????? ????-???????? ?????? //??????????????? ???????????? ? ???????? ? ??????? ??????. ???????????: ?????. ???. ???., ?. 30-37.
???????? ?.?. (2001) ????????? ?????? ? ????????? ????????? ??????????? ???????
????? ?? ????????? 150 ??? //????????? ???????????????? ????????????. ?., ???. 90, ?.
117-125.
???????? ?.?., ?????? ?.?. (1999) ???????? ???????? ????? ? ???????????? ????
?????? ??????????? ??????? //????????? ???????????????? ????????????. ?., ???. 87, ?. 119123.
???????? ?.?., ??????? ?.?. (2003) ??????????? ?????????? ????? ?? ?????? XXI ????
//????????? ???????????????? ????????????. ?., ???. 95, ?. 93-102.
???????? ?.?. (1950) ??????? ???????? ??????? ????????? ??? ????? ????????????? //
???????? ????????. ?.: ?.: ???-?? ?? ????, ???. 1, ?. 465-483.
??????? B.C. (1966) ????????? ????????? ? ??????????? ?????????? ????-????????
?????? //??????????? ?????. ?????: ???-?? ???????? ????????????, ???. 5, ?. 149-161.
???????? ?.?. (1996) ??????????? ????????????? ????? ?????-???????? ?????? ???????
(?????????????, ???????????, ?????????). ???????, 287 ?.
??????? ?.?. (1961) ????????????? ?????????????? ?????????? ??????? ? ??????????? ????? //???????? ???. ?. 93. ???.6: 74-88.
?????????? ?.?. (1949) ?? ?????. ?: ????????????, 579 ?.
??????? ?.?., ???????? ?.?. (2002) ??????????? ???????????? ?????????????? ?? ??????? ??????? ? ??????????? ????? //????????????? ??????? ? ????????? ????? ? ????????
? ??????????? ??????. ???????????, 2002, ?. 398-401.
??????? ?.?., ????? ?.?., ??????? ?.?., ?????????? ?.?. (2003) ??????????? ?????????
????? ? ?????????????? ?? ???????? ?????????? ????? ?????????? ????? ??????-????????
?????? (??????????? ?????) //???????????? ???????????? ? ????????? ??????: ????????? XI
?????? ???????? ????????????? ????????. ?. 1. ???????, ?. 406-407.
??????? ?.?., ????? ?.?., ?????????? ?.?. (2008) ???????????? ?????????????? ??????? ????? ???????? ???????? ??????-???????? ?????? (??????????? ?????) ??? ????? ????????????? ??????????? //????????? IV ?????????????? ?????????? «???????? ? ???????????? ?????????? ?????». ?????, ?. 144-146.
?????? ?.?. (1949) ?????? ?????????? ?????. ?.: ?????????, 376 ?.
????????? ?.?. (1995) ?????????? ???????? ?????? ? ???????????? ??????????. ???., 990 ?.
???????? ?.?. (1964) ???????? ? ???????????. ?.: ???-?? ???, 380 ?.
?ooper W. S. (1931). A third expedition to Glacier Bay, Alaska. Ecology, vol. 12. ? 1: 61-65.
Cooper W. S. (1939) A fourth expedition to Glacier Bay, Alaska. Ecology. vol. 20. ? 2: 59-130.
Ludi W. (1945) Besiedlung und Vegetationsentwicklung fur den jungen Seitenmorenen des
Grossen Aletschgletschers. Bericht uber das Geobot. Forschungsinst. Rubel in Zurich. S. 32-59.
# 101 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ?????? ??????? ???????????? ???????? ?? ??????? ??????? ??????? ??????????
Palmer W.H., Miller A.K. (1961) Botanical evidence for the recession of a glacier // Oikos, vol.
12, ? 1: 75-86.
Timoshok E.E., Narozhnyi Yu. K., Dirks M.N., Berezov A.A. (2003) Expirience in Combined
Glaciological and Botanical Studies on the Primary Successions on Young Morains in the Central
Altai. Russian Journal of Ecology, vol. 34, ? 2: 91-97.
Timoshok E.E., Skorokhodov S.N. (2006) Species, coenotic and ecosystem diversity in headwater
basin Aktru (Severo-Chuisky Range, Centrai Altai). Global Changes in Mountain Regions. M.F. Price,
ed. Duncow, Sapiens Publ., ?. 174-176.
Species Diversity of Plants on New Moraines
of the Sophysky Glacier (Juzhno-Chuysky Ridge, Central Altai)
Elena E. Timoshok, Marina N. Dirks
and Sergey N. Skorokhodov
Institute of Monitoring of Climatic and Ecological Systems SD RAS
10/3 Academichesky, Tomsk, 634055 Russia
On new moraines of the Sophysky Glacier in cryoarid conditions of Juzhno-Chuysky ridge species
composition of vascular plants and mosses was revealed in different stages of colonization. Information
about ecological peculiarities of species of new moraines flora is given.
Key words: new moraines, species diversity, vascular plants, mosses, Juzhno-Chuysky ridge
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 103-116
~~~
??? 504.7.05: 551.582 (571.1/5)
??????????? ?????? ???????????????
? ????????????????? ?????????.
II. ????????????? ????????? ? ??????????? ???????????
?????? ?????????
?.?. ??????*
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???,
634055 ??????, ?????, ????????????? 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
???????? ????????????? ????????? ??? ???????????? ????????? ?????????????,
??????????????? ? ????????????? ????????, ??????????? ??????????? ??????
??????????????? ? ????????????????? ????????? ??????????? ?? ?????? ???????????
??????? ??????????? ????????? ????, ??????????? ???????? ? ???????????? ?????????
? ???????????? ????????-????????????? ???????.
???????? ???? ????????-???????????? ?????? ?????????. ?????????? ??????? ???????
????????? ???????? ?? ??????????????? ????????. ??????? ????????? ????????????
????????????????? ????? ??????????????? ? ????????????????? ????????? ???
????????????? ??????????. ??????????? ????????? ?????????????? ?????? ???????????
??????????? ????? ????????????????? ????????? ? ???????????? ??????? ????????????????????? ???????. ???????? ???????? ????????? ????????? ? ????????? ????????,
??????????? ???????????? ????????? ????????? ????????? ?????? ????????? ? ????
????????????? ???????????.
???????? ?????: ?????, ?????????????, ???????????? ?????????, ????????????
?????????, ??????????? ???????????, ??????????????? ?????????.
????????
?? ??????????? ??????? ?????? ????????????????? ????????? ???????, ??? ??????????????????? ??????????? ????????
? ?????????? ???????? ??????????????????????????? ????????? ???????? (????????) ????????????? ??????? ? ????????
???????? ?????? (??????, 2008). ????????????? ????????? ??????? ????????? ???*
1
?????? ?????????????? ???????? ? ????????? ? ???????? ??????????????????
?????????. ?????????????? ????????????
??????????-??????????????
???????????
?????????? ?????? (?????? ? ??., 1995; ??????, 2003) ? ????????????? ????? (???, 1991)
??????? ? ????????? ????????????, ???????????????? ? ?????????????????? ?????????
? ??????????? ???????? ? ?????????????
Corresponding author E-mail address: shishlov@imces.ru
© Siberian Federal University. All rights reserved
# 103 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ??????????? ?????? ??????????????? ? ????????????????? ??????????
?????????????????????? ????????? ? ????????? ?????????.
? ????????? ?????? ???????????????
?????????????? ?????????????????? ????????? ??? ???????? ??????????????
????????? ? ????????? ??????? ????????? ? ????????????? ?????????????????
????????, ??????????? ??????????????
?????????????, ??????????? ???????????
????????????????? ????????? ??? ????????????? ????????? ? ???????????? ??????? ????????-????????????? ???????. ?
?????????? ??????? ??????????? ???????
??????????? ????????? ????????? (???????? ??????????, ?????????? ??????? ??????????? ???), ??????????????? ?????????
(????????? ???????????, ????????) ? ???????????? ??????????? ?? ??????????????????
? ???????-????????????? ?????????. ?????? ???????? ??????? ????????????????
?????????? ???????? ? ????????????????
???????? ????????-???????????? ??????.
???????? ???? ????????-???????????? ?????? ?????????. ?????????? ??????? ???????
????????? ???????? ?? ??????????????? ????????. ????????? ????????? ?????? ????????????????????, ??????????? ????? ? ?????
? ??????????????????? ???????? ? ???????
???????????? ? ????????? ???????? ?????.
??? ???????????, ? ????? ??????? ??????????????? ?????????????? ??????????? ??
??????????? ? ????????????? ??????? (???????, ???????, 2006) ? ??????? ???????????????? ????????.
? ?????? ????? ???????????? ?????? ?
??????????? ???????????? ????????? ????????????????? ??????? ????????? ? ?? ??????????? ??????? ????????-????????????
????????? ???????????? ????????????????? ????? ????????????????? ?????????
??? ????????????? ?????????, ??????????????? ???????????? ????????? ? ???????-
???????? ????????-????????????? ????????
? ???????????? ??????????? ????????????
???????????? ????????-????????????? ????????? ? ????????. ??????????? ???????? ???????? ?????? ????????? ????????? ? ????????? ????????, ??????????? ????????????
?????????, ??????? ????????? ?????????
?????? ????????? ? ???? ????????????? ??????????? ? ???????????? ??????? ????????????????????? ???????. ?????????? ??????
? ??????????? ????????? ????????? ??????????? ?????????? ???????, ?????????? ?????
??????????? ????? ???????? ????????? ?
????????-????????????? ????????.
?????????????????? ?????????
????????????? ?????????????? ????????? ? ?????? ????????? ???????, ?????????? ? ??????????? ??????????????? ????????? ? ???????? ???????????????????
??????? ? ????????????? ?????????????????????? ?????????, ????????? ??????? ??????????? ??????? ????????????????? (??????, 2003). ?????????????? ?????????????
???????? ? ????????? ? ???????? ?????????????????? ????????? ???????? ??????????????? ??????? ? ????????? ??????????????? ???????????? ??????????????????
?????????. ????????? ??????? (???????????
?????, ??????????? ????, ?????? ???????)
????????? ???????? ????????? ????????,
??????????????? ? ???????????????? ????????? ? ???????????? ??????? ??? ??????? ??????? ??? ???????? ?????????????????
???????. ?? ???? ? ????????? ????????????? ??????? ????????????? ???????????????
???????? ??????? ????????? ????????.
???????? ? ???????????? ???? ? ????,
???????? ? ???????? ???????? ????? (?????
????????????? ????????????? ? ???? ? ???????????????? ? ?????) ???????? ???????????? ????????? ??????????? ???????????.
# 104 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ??????????? ?????? ??????????????? ? ????????????????? ??????????
????? ?????? ??????????? ??????????? ??????, ????????? ??????? ????? ?????????? ????????? ???????? ??????????? ?? ?????????, ?
????? ?????????? ? ????? ?????? ????. ????
? ??? ????? ??????????? ???????. ???????????? ??????????? ???????? ???????? ??, ???
??????????? ?????? ??????????? ? ????????
???????????? ???????? ?????????????? ????????. ??????? ? ???????? ?????? ??? ??????? ?????????? ??????? ????????????
?????????? ?????. ?????????? ???????? ???????? ?????? ?? ????? ???????????? ???????? ??????? (????????? ??????? ???????????)
? ???????????? ?????????????? ???????????? ??????? ??????????, ?? ??????? ???????
????????????? ???????????????? ???????.
????????????? ??????, ?????????? ? ??????
???, ??????????? ?????? ?????????????
?????, ??????? ????? ? ???????? ???? ???????
??????? ??? ???????????? ????????????? ?????????????????? ???????? ?? ????? ???????
? ???????? ??????????? ??????????? ? ???????? ?????????????? ????? ????? ??? ??? ??????? ????? ??????????? ???????. ?????????
???????????????-??????????? ???????????
??????? ???????? ???????? ? ??????????????
??????? ????????? ????????? ??? ? ????????????? ????? ??????????????? ? ?????????????????? ?????????, ??????? ?????? ??
????????????? ?????????. ??????????????
??????? ??? ?????????? ???????????? ???????????, ??????????? ???? ?????????????
???, ????????????? ????????? ?????????????? ??????????? ???????? ?????????? (??????????? ????? ? ???????? ??????), ??? ???????? ?????? ?????? ??????????, ????????
???????? ?????, ????????? ????????????????
????????? ?????????. ? ?????? ?????? ?????
????? ?????? ??????? ?????????????? ???????????? ???????? ?? ???????? ????????????????? ???????? ????????????? ???. ??????
???????? ??????????? ?????????????????
????????????????? ??? ? ?????? ??????????
???????????? ??????????????????? ????????
?????????? ??????????? ?????????? ? ???????? ? ?????????????? ??????? ????????????? ????????????? ? ?????????????????
??????? ??? ??????? ??????????????? ??????????? ?? ??????????? ???????????.
????????????????? ??????? ?? ???????? ????? ? ??????? ??????????? ????????? ??????? (?????, ?????????????, ????), ?.?.
??????? ???? ? ??? ???? ??????, ??? ????????? ??? ???????? ??????? (?????????, 1978).
???????? ?????? ????? ?? ??????????? ???????????? ?????????????? ?? ???? ????????????? ???????, ??????? ?? 20-40 % ????,
??? ?? ????. ???? ?????? ????????????
?????? ?????????? ??????????? ? ????????
????? ?? ????????????? ???????. ?? ??????
???????? ??????? ???????? ????????? ???????
????????? ??????? ??????????? ????????????? ???????? ??????? ????? ?? ????????? (3040 %). ?????? ??????????? ??????? ?????????? ???? ??????????? ???? ? ?????? ????????
???????? ????? ?????????? ????? ???????.
???????? ???????????? ???????? ???????????????? ??????? ?? ??????? ??????????.
???, ????????????? ??????????? ? ???????
????????? ???????????? ????? ??????????
?????????? ?? 1-1,5 °C. ?????????? ??????? ???????????? ?? ??????????? ??????? ?
?????? ??????????? ?? ?????????? 2-5 ??, ?
????????????? ? 35-60 ??. ??????? ????????
?????? ?????????? ???? ?????????? ????????????? (??????? ?????? ???????????
2170 ??2, ????????????? ??????????? ? ????
????????????? 8-10 ?), ?????????? ? ??????
????????????????? ?????? ????? ? ?????????, ?????????? ???????????? ? ???????
?????????????? ???????? (?????????, 1978).
?????? ??????? ?????????? ?????????????????? ? ???????????????? ??????????? ???????? ????????? ????????????? ?? ??????????,
# 105 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ??????????? ?????? ??????????????? ? ????????????????? ??????????
????????????????? ??????? ??????????????
??? ???????????? ??????????, ??????????,
?????????, ????????????, ???????????,
?????????? ?????????.
???????-????????????? ????????? ? ??????????????? ???????? ?? ????? ??????? ???
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???????? ??????? ??????? ?? ????? ??? (6 ?????), ????? (5 ?????), ?????? (3 ?????) (????????, ?????????, 1997). ?????? ????????????
????????? (Shishlov, 2001) ???????? ??????????
????????-???????????? ????? ???????? ? ????????? ??????????? ? ???????? ??????? ??????????? ????????? ????????? ? ???????????
(?) ???????? ??????? (???. 1). ??????? ?????????? ???????????? ??????? ?????? ????? ???
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??? ? ????. ?????? ?? ????? ??????? ??????? ?
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?? ???? (???.2), ??????? ????????? ??????????
???????? ??????? ????????????? ????????
?? ????????? ????? ???? ?? ?????? ?????????
? ?????????? ???????????? ???????????????
????? (??????, 1999). ?? ????? ???????? ????????? ?????????????? ??????????? ????
????? ?? ?????? ????????????, ????????? ?
??????? ??????????? ??????. ??? ????????
??????? (-S) ????? ????????????? ???????
(+S) ?????????? ? ???????????? ??????? ????????.
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???=???+?(??)+
??(?,??)+???(??1,??2,??3).
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? ????????????? ??????? (???????, ???????, 2006). ? ???, ???? ????? ??? ?????????
# 106 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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??3 ? ????????????? ???????, ?- ???????????, ? ? ????????? ??????? ????, ? ? ??????????, ? ? ??????,
?? ? ????????? ?????????, ?? ? ????????????? ????, ? ? ?????, ?, ??? ? ?????????? ???????,
? ? ?????, ?? ? ?????? ?????????, ?? ? ????????????? ?????????, ? ?? ? ????? ???????????????
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# 107 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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????????????? ????????: ???????, ?????,
????????? ???????? ? ???????? ??????????
????????? ???????????????? ???????????
(???????????, 1981). ?????????????? ? ????????? ????????? ??????? ?????????? ????? ????????? ??? ?????????? ?????????? ??????????????.
??????????? ???????? ???????????????? ????????????? ????????-????????????? ?
???????????????? ????????? (?????? 1998;
??????, 2002), ??????? ????????? ????????? ????? ??????????? ????????? ??????????? ????????? ? ?????????????? ????????? ?
?????????? ????????????? ??????? (???. 3) ?
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????????? ???????? ????????????? ????????? ???????????? (???): ??????????? ????,
???? (????????, ???????, 1978). ? ??????????
??????? ??????? ????????????? ?????????
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???????? ????????, ???????? ?? ?????????? ???????????, ??????????? ???????????
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????????? ???????? ?????????? ????????????? ??????. ??????? ??????????? ?? ?????????? ????????????? ???????? (????, ???????? ????????? ? ?????, ???????????????)
???????????? ????? ????????? ???????????????????? ????????. ??????????? ???? ????????? ???????? (???????????? ?????????
??????????????, ???????????? ???????????
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(???????????? ??????????? ??????? ?? ???????? ? ?????, ?????????? ?????????? ?????,
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???????? ?????????? ????? ????????????????????? ?????? (Shishlov, 2001) ??????? ????????? ??????????, ????? ???????,
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# 108 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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??????) ? ????????? ???? ????????, ????????????? ??????? ???????????? ? ???? ????????. ???????? ?????????? ???????? ??
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???? ?? ???????????? ???????.
?? ????? ????? (???. 3) ???????? ???????? ??????????? ????????? ????? ??????????: ???????? ??????? (1), ???????? ?????? (2)
? ????? (3) ? ??????-«?????????» ???????
???????: ??????? (4), ????????? (5), ??????????? ?????????: ?????? (6) ? ?????????
(7), ?? ???????????? ????????? (???????
????????? ?????) ? ??????????? ?????????? (9), ????????????????? ??????????
(10) ? ????????? ?????????? (11), ???????
??????? ?????? ??????? ??????????? (??)
? ????? ????? (12) ??????????????????????
?? (????????? ?????, ??????, ????, ????????). ???????? ?????????? ???????????
????? ????? ??????????? ? ??????? ????????? ????????? ?? ??????? ???????????
?????????. ????????????????? ??????? (13),
?????????? ?????? (?) ?????????? ????????,
??????? ????????????? ??????? (??), ? ?????
?????? ?? ???????????????? ? ????????? ?????????????? ?? (??????, 2002).
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?????????? (??? ???????? ????? ?????????)
??????????? ??????? ?? ??????? ????????????? ???????? (????????, 1983; ???, 1992).
# 109 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ??????????? ?????? ??????????????? ? ????????????????? ??????????
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? ?????????? ???? ?? N? ????? ?????????? ?
???????? ??????????? ? ??????????? ??????? ???????? ???? N?? ?????. N??=(0,15ч0,2)N?
(???, 1992).
?????? ?????????? ???????? ??????? ?
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(????????? ??????, ???? ?????????) ???????? ???????????? ??????????? ???????. ?
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N??<0.02N??. ???? ???????? ? ????????????????? ????????? ?????? ??????? ??????. ??
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# 110 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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1999) ???????? ??????? ??????????? ??????????????????? ????????? ????????? ? ?????? ???????? ??????????? ??????????????
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(??1, ??2, ??k) ? ???????????? ??? ??? ???????? ????????? ????????? ? ?????????????
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????????
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????????????????;
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?????;
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???? ????????????? ??????.
???????? ????????? ???????? ????????????? ?????????? ???????????? ????
(Oke, 1982):
? ??????????? ??????? ??????? ?? ?????????? ??????? ?????;
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???? ? ??????;
? ?????????? ?????? ????? ?? ????????????? ????;
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# 111 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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??????? (?j). ????? ????, ????????? ???????? ?j ?????? ?? ???????????? ??????? ???????????? ??????? ? ????????????? ???????
(???? ???????????? ?????). ?????????????
????????? ???? (??) ? ??????????? ????,
??????? ?????, ???????? ????????? ?????????????? ??????????? ??????? (??????
???? ???????????? ??????). ????????? ???????????? (???? ?????????, ???? ????????????? ??????????????????) ???????????????
?????? ?? ????????????? ??????? ???????.
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?????? ???????? ? ????????? ??????? ????????? ????? ? ?????????? ???????? ??????? (??????, 2005). ??? ????????????
???????????? ?? ?????? ???????? ?????? ?????????? ???????? ?????? ?????????? (????,
2001). ???????? ?????, ????????????? ? ????
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# 112 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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- ???? ????????????? ????????????????? (????????-???. ????
???????????) ? ??????????????? ????????? ???????
???????????? ??????? ???????????? ???????
? ????????????? ???????
???. 4. ????? ?????????????? ?????? ??????????? ??????????? ????????????????? ????????? ?
???????????? ??????? ????????-????????????? ???????
?????????????? ? ??????? ??????????? ??????? ??????? ????, ?????????????? ???????,
???????????? ???????, ???? ??????? ???????
(????????) ????????? ? ?????????? ????????????. ???? ??????? ??????????? ??????????? ??????? ?? ?????? ??????????? ?? ?.
?????????? ?????? ???????? ???????????? ??????? ?????????? ????????? ? ????. 1.
?????????? ? 1914-1950 ??. ?????? ????????
????????. ??????? ?????????? ???????????
? ???????? ???????? (????????, ?????????).
?? ??? ?????? (????, ???????, ???????) ??-
??????? ???????? ???????????? ???????
??????????? ????? ????? ? 1951-1968 ??. ????????? ?????????? ?? ??????????? ????? (?
1978 ?.) ??????????????? ????????? ????????
???????? ????????? ? ??????? ? ??????????
????????????? ????????????. ???? ???????
??????? ????????? ? ?????????? ????????????? ???????????? ??????????? ? ????????? ??????? ?????? (? ????????? ??????? ????????? ? ? > 1,3 °? ?????????? ?? 0,1 ?? 0,52,
? ????????? ??????? ????????? ? ? > 0 °? ?????????? ?? 0,06 ?? 0,37, ? ????????? ? ? 0,07
# 113 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ??????????? ?????? ??????????????? ? ????????????????? ??????????
??????? 1. ?????? ???????? ???????????? ??????? ??????? ???????????, °?
?????
1881-1913
1914-1950
1951-1968
1969-1977
1978-1994
1995-2001
2002-2008
????????
-1.6ч2.4
-1.7ч1.7
-1.8ч2.1
-3.5ч0.9
-0.7ч2.2
-0.9ч2.6
-0.1ч2.6
?????????????
????????
-3.8ч-0.2
-2.8ч0.3
-3.9ч1.0
-4.9ч0.7
-3.3ч0.6
-2.9ч1.5
-1.9ч1.6
-10.8ч-4.2
-8.3ч-2.9
-8.7ч-3.6
-9.0ч-5.3
-8.5ч-3.7
-8.3ч-2.9
-7.6ч-4.1
????
???????
?????????
-1.5ч2.4
-1.5ч2.3
-1.0ч3.1
-2.0ч2.0
0ч3.6
0.3ч3.1
1.3ч4.0
-1.6 ч+2.8
-0.6ч2.1
-0.1ч3.5
-1.0ч3.0
0.3ч4.3
1.1ч4.3
1.1ч4.9
-3.5ч0.2
-4.2.ч-0.4
-2.3ч1.0
-2.3ч1.1
-2.2ч1.6
-
-3.3ч0.6
????????
-4.1ч0.5
-3.8ч-0.3
-2.8ч-0.7
-4.4ч-0.6
-3.0ч0.6
-2.0ч0.5
-2.8ч1.4
?????????
-9.9ч-5.2
-8.2ч-5.1
-9.7ч ? 5.2
-10.0ч-5.2
-8.5ч ? 3.9
-8.2 ч-3.4
-7.5ч ? 4.2
???????
-2.5.ч-0.2
-2.7ч0.3
-1.8ч0.6
-1.6ч0.4
-0.5ч1.9
0ч2.3
0.3ч2.9
??????
-4.1ч-2.1
-3.8ч-0.9
-3.7ч-0.1
-3.8ч-0.6
-2.6 ч0.7
-1.2ч 0.8
-2.0 ч1.7
?????
-
-7.6 ч-4.2
-7.7ч ? 5.5
-8.5 ч-5.0
-7.7ч ? 4.0
-6.7ч ? 4.6
-6.9 ч-4.0
?? 0,32). ??????????? ???????? ???????? ??????? ?????????? ? ???????? ???????.
???????? ????????? ????? ??????????
??????? ??????, ????????? ???????????
???, ???????. ? ???????? ???????? ??????????? ???? ??????? ??????????? ???????????
??? ??????? ?????????????? ??? ??????????
???????? ???????????? ??????? ???????????
(?????????, ????????, ?????). ??????????? ??????????? ???????? ???????? ????????? ? ???????????? ? ??????????? ???????
??????????? ????????????????? (?), ?? ??????????? ??????? ????? ??????????????? ????????? (??) ? ???????? ????? ? ??????????
(?) ?????. ??????????? ????????? ????? ??
???????, ??? ??? ??????????? ??????? ????????????? ???????? ????????? ? ?????? ???????????? (??????? ????????????? ?????????
? ????????? ? 0,33, ?????????? ? 0,23, ????????? ? 0,17, ?????? ? 0,28, ??????? ? 0,43, ????????? ? 0,43, ????????? ? 0,36). ?????????? ?
????????? ?????? ????? ???????? ????????.
??????????? ????????? ?????????? ? ??????
???????? ???????????? ???????? ? ???????
?????????? ? ????? ???????? ?????? (????,
??????, ???????, ???????).
?????? ? ????????????? ??????????? ??????????????? ????????????? ???? ??????????? ????????? ??????????? ??? ????????
???????? ????????? ??????? ?. ????? ????????, ??? ???????? ???????????? ?????????,
??????????
??? ?????? ????? ?????? ????????? ????????? ??????? ???? ? ????????? ????????, ???
??????????? ?????? ????????????? ????????. ???????? ????? ???????? (?) ???????
????????? ? ??????????? ????????????? ?
????????? ????? ? ????????????? ??????.
??????????? ?????????????? ????????? ???????? ????????????? ?????????
? ???????? ? ????????? ??????? ????????????????????? ???????. ??? ??????????????
??????? ??????????? ?????????? ???????????? ???????????? ???????? ???????????
?????????????? ????????? ? ???????????
????, ????????? ?????????? ? ?????? ?????????????????? ?????????. ??????? ??????? ???????? ??????-???????????? ?????
????????-????????????? ??????? ???? ??????????? ??????????? ????????? ?????????
(?????????????, ???????????????) ? ????????????? ????????, ?????? ?? ??????? ??
??? ????????????? ?????????, ???????????
# 114 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ??????????? ?????? ??????????????? ? ????????????????? ??????????
??????????? ?????? ??????????????? ? ????????????????? ????????? ? ????????????
??????? ????????-????????????? ???????.
??? ??????? ?????????????? ?????????? ??
??????????? ??????????? ????? ?????? ???????? ???????? ????????? ? ?????????? ????????? ????????? ?? ???????????? ???????
????????????? ????????? ??????????? ???
?????????????? ????????? ????????? ?????-
???????? ????????? ?? ?????? ????????? ???????? ??????? ??????? ????????? ????????.
?????? ??????????? ????????????? ??????????? ?? ????? ????????????? ?????????
??????????? ???????? ???? ???????????, ??????????? ??? ?????????? ??????????????
?????? ????????????? ??????????? ??? ?
??????????? ??????????? ????? ?????????
(??? ?????).
?????? ????????? ??? ?????????? ????????? ??????? 10 ????????? ??????????
??? ?16.
?????? ??????????
??? ?.?. (1992) ????????????? ????????????? ??????? ?????. ???????????: ?? «?????»,
????????? ???????????? ?????, 200 ?.
???????? ?.?. (1983) ????????????? ?????? ???????????? ????????????? ???????? ?????.
???????????: ?????, ????????? ?????????, 254 ?.
????????? ?.?. (1978) ?????? ? ??????????? ??????????? ?????? ?????. ???????????:
?????, ????????? ?????????, 214 ?.
???????? ?.?., ????????? ?.?. (1997) ???????????? ? ??????????, ? 1, ?. 81-89.
??????? ?.?., ??????? ?.?., ?????? ?.?. (2000) ???????? ??????? ? ?????????????
???????????? ????????-????????????? ????????? /?.?. ??????? (???.) //???????????? ?????????? ?????????. ?. IV. ????????-????????????? ?????????. ?????, ??? «?????», ?.
200-252.
??????? ?.?., ??????? ?.? (2006) ?????????? ? ????????????? ????????-?????????????
????????? ? ?????? //?????? ????????? ? ??????. ?. 19. ? 9, ?. 1-14.
???? ?.?., ???????? ?.?., ?????? ?.?. ? ??. (2001) ???????? ??????? ???????? ?????? ?
?? ??????????????? ???????? /?.?. ?????? (???.). ????: ???? ? ?°, 584 ?.
??? ?.?. (1982) ??????? ???????????? ???? /???. ? ????. ??? ???. ?.?. ??????. ?.: ???????????????, 360 ?.
??????????? ?.?. (1981) ?????????????? ???????? ??????????. ?.: ?????, 231 ?.
?????? ?.?. ? ??. (1999) ???????????? ??????. ????????????? ????????????, ??????????
???????? ? ??????????????? ????????. ?.: ??? «??????», 704 ?.
???????? ?.?., ??????? ?.?. (1978) ???????????? ????????????? ?????????. ?.: ?????,
352 ?.
?????? ?.?. (1998) ?????? ??????? ????????????? ???????????? ?? ??????????? ?????
? ???????????? //?????? ?????? ?.?.??????: ????????? II ??????????????? ?????????????
???????????. ?????:???, ?.121-122.
?????? ?.?. (1999) ??????? ????????????? ???????? ?? ????????????? ?????????? ? ????????? ??????? ? ???????? ?????? //?????? ????????? ????????? ?? ????????????????????? ???????????: ?????? ????????. ?????: ???-?? ??? ?? ???, ? 86-87.
# 115 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ??????????? ?????? ??????????????? ? ????????????????? ??????????
?????? ?.?. (2002) ?????? ???????????? ????????? ? ?????? ????????????? ????????????? ????????? ?????????????? ? ???????????? ????????? ????? // ????????? II ?????????????? ?????????? «???????? ? ???????????? ?????????? ?????». ?????: ???-?? ?? ???,
?. 28-32.
?????? ?.?. (2003) ?????? ???????????? ???????????? ????????????? ????????? ? ?????? ? ?? ??????????? // ????? ????????? ????????? ?? ???????-?????????????? ???????????. ?????, ?. 119-123.
?????? ?.?. (2005) ?????? ??????????? ????????????? ????????????? ?????????? ?????? // ?????? ????????? ????????? ?? ???????-?????????????? ???????????. ?????, ?. 254257.
Shishlov V.I. (2001) Analysis of ecological problems in antropoecosystems // Proceedings KORUS
The 5th Korea-Russia International Symposium on Science and Technology. Tomsk. KORUS 2001. v.
2, p.80-83.
The Organization of Processes Cycles Forming a Climate
and Environment.
II. Landscape Transformation and Reorganization of Process Cycles
Viktor I. Shishlov
Institute of Monitoring of Climatic and Ecological Systems, SB RAS,
10/3 Akademichesky, Tomsk, 634055 Russia
Problems of landscape transformation under influence of climatic, hydrological, and anthropogenesis
factors combination and reorganization of the environmental and climate formation cycles have been
investigated. It has been done by means of the joint analysis of the associated changes of environments,
geosphere components and intersystem relations in regional natural and climatic system.
Chains of cause-effect relations of processes are revealed, and influence functions of factor
combinations on the hydrological processes are defined. The formation concept of the modified forming
cycle of environment and climate processes under transformation of landscapes has been developed.
The structure of conceptual model of reorganization of the cycle organization of climate processes and
formation of local natural and environmental conditions has been developed. The means of description
of the natural processes and the factor combination, the organization of interconnections allow the
analysis of changes in the transformation of the components.
Key words: cycles, transformation, intersystem relations, ecosystem changes, reorganization,
hydrological changes.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 1 (2009 2) 117-132
~~~
??? 551
??????? ????????????? ???????? ?? ?????????? ???? ?????
? ??????????? ? ???????? ? ????????? ?????????? ?????
?? ??? ???????? ??????
?.?. ?????????*, ?.?. ????????
???????? ??????????? ?????????????
? ????????????? ?????? ?? ???
634055 ??????, ?????, ??. ?????????????, 10/3 1
Received 3.03.2009, received in revised form 10.03.2009, accepted 17.03.2009
??????????? ??????? ??????????? ? ??????? ?? ???????????? ???????? ????? ???????
?????? ????? ?????????? (Pinus sibirica Du Tour) ? ??????????? ????????? (Larix sibirica
Ledeb.) ? ???????? ? ????????? ?????????? ?????. ????????, ??? ??? ??? ???? ???????????
??????????? ?? ??????? ??????????? ????? ?????? ?? ??????????? ? ?????? ?? ??????????
?????????????? ???????, ??? ???????????????? ??????????? ????????????? ????????? ?????
???????? ????. ???????????? ????????? ???? ??????? ? ??????????? ??????????? ???????
?? ?????????????? ????????? ???????? ??????? ?? ?????????? ???? ???????? ? «?????» ?
«??????» ????, ?????? ? ???????? ????? ??????? ???? ?????????? ??????? ? ???????? ??????
??????, ? ?? ?????? ????? ??????????? ??? ??????? ??????????? ? ????????? ???????. ?
????????????? ????? ???? ??????? ?? ????????? ????? ?? ?????????????? ?????? ???????
????, ????????? ? ???????????? ???? ?????????????? «???????» ????? ???? ??????????
???. ???? ??????? ???????? ?????????????? ???????? ??????? ?? ??????? ? ???????
?????????????? ??????, ? ? ???????????? ???????? ????? ????????????? ??????? ?????????
?????? ???????????.
???????? ?????: ???? ?????????, ??????????? ?????????, ?????????? ????, ???????????,
??????, ????????? ?????.
????????
?????? ????????????? ???????? ?????????? ???????????? ??????? ?? ?????????
???????, ??? ?????????? ? ?????????? ????????? ????????? ??????????? ? ????????? ????????? ??????? ????????????????? ?
?????? ????????????? ????? ? ?????????
??????? ??? (Jones, Briffa, 1992; ??????????,
1992; ???????, 1997; ?????????, ????????,
2001; ? ??.). ? ?????? ???? ????????? ?????
*
1
?????????? ?????????? ???????? ?????????,
?????????? ?? ????? ?????????? ????
??????? ??????? ? ????????? ??????? ???
????? ???????, ??? ???? ?????????????? ????????? ??????????? ? ??????? ????? ??????? ????????? ????????? ? ????????????????? ????????. ???????? ?????? ????????
??? ??????????? ???? ????????? ????????
????????? ????????, ??????? ????????? ??????????? ????? ???????? ???????? ???????-
Corresponding author E-mail address: velisevich@imces.ru
© Siberian Federal University. All rights reserved
# 117 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
???? ??????????, ??????????????? ? ???????? ???????.
? ??????? ?????????? ???????? ???????????? ??????????? ???????? ?? ???????
???????? ???????? ?? ?????????? ????
???????, ?????????? ? ???? ???????????
???????????? ? ?????????????? ? ???????????? ??????? ? ????? ?????????? ??????????
?????????? ??? ????????????? ???????. ?
????????? ??????????? ?????????, ???????????? ?????????? ????????????????? ?
??????????? ???????????? ??? ????-??????,
???????????, ??? ??????????? ???? ? ????
?????????? ?? ?????????? ??????? ? ???????????? ??????? (??????, 1986; ??????????, ???????, 1999; ?????????, ??????????,
1999; ? ??.) ? ?? ???????? ??????? ??????????????? (????????, ??????, 1975; ???????
? ??., 1996; ??????, 1999). ? ????????? ?????
?????????? ?? ???? ???????? ????????????
?????? ?????? ? ????? ? ???????? ???????
??????: ???????? ????????????? ??????????? ? ???????????? ???? ? ??????? ???????
(Vaganov et al., 1998; ???????? ? ??., 2002).
? ????? ????? ????????????? ??????
?? ??? ?????? ???????????? ???? ????????,
??? ?? ??????????? ????????. ???? ????????
? ????????? ??????? ??????? ?? ????????????
??????? ? ?????????? ? ??? ????????????????? ????????? ????? ? ????????. ? ?????
???????? ? ???????? ??????? ????????????
???? ????? ???????????? ????? ????????????, ?????????? ??????????? ??????? ????????????? ??? ?? ??????? ????????? ?????.
?? ??????? ????? ???? ???????? ??????? ??????????? ????????? ???????? ???????????
???????? ? ???????? ??????????? ? ???????????? ????????????? (????????, ???????,
1999; ????????, 2003; ????????, 2003), ? ????? ??????? ???????? ???????? ?? ??????
?????? ??????? ? ??????????? ?? ?????????
?????????? ???? (??????, ??????????, 1976;
?????????, ??????????, 2000; ? ??.). ??? ??
?????, ? ??????????? ??????????? ???????
????????????, ????????????? ??????????
???????????? ??????????? ????? ?? ????????? ???????????? ???????????????? ??????
???? ?????????? ?????????????. ???????????? ?????????? ????? ???????? ???????????? ???????? ???????? ? ????????? ????????, ???????????? ???? ????????.
??? ??????????? ????? (?????? Strobus),
? ??????? ??????????? ? ???? ?????????,
??????????? ?????? ???????? ???????? ????????????? ???????? ? ?????? ??????????
?????? ? ???????? ???????. ??????, ? ??????? ?? ????? ????????? ?????????????, ????
??? ? ???????????? ????????????? ????????
?????????, ?????? ???????????????? ?? ????????? ?????????? ???????? ?????? (???
? ??., 2004). ??????????? ????? ??????????
???????????? ? ???????? ?????????? ????????? ????? ?????? ??? ?????????? ???????? ??? ????????????. ???????????????
????????????? ?????????? ???????? ??????
(Blyakharchuk, Sulerzhiysky, 1999; Crawford et
al., 2003; ????????, ???????, 1999), ? ?????
????????????? ????????? ???????? ??????????????? ???? ? ?????? ? ???? ?? ????????
?????? ??????? ??????????, ?? ?????? ???????????? ???????????? ?????????? ????????????. ????? ????, ????, ??? ????????????
???????????? ??? ????????? ????? ???????????? ?????? ???????? ? ????? ?????? ???????????, ? ??? ????? ? ??? ????? ????????,
??? ???????? ???????????? ???????? ???????? ????????????? ????????? ?????.
???????????? ??????? ???????? ? ??????????? ??????????? ?????????, ???????????? ?? ???????????? ???????, ? ?? ?????
?????????? ? ??? ?????, ????? ????? ????????????? ? ??????????????? ????????????.
?????? ??????????? ?? ??????? ?? ????????
??????? ??? ????????? ???????????????
# 118 #
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?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
?????? ???? ?????????? ????????, ??? ? ???????? ????????? ????????????? ???????
????????? ??????????? ?????, ? ?? ?? ????????? (????????, ??????, 1975; ????????,
???????, 2004).
?????? ????????? ?????? ? ????????
????????????? ?????? ??????? ???????????
? ??????? ?? ???????????? ???????? ?????
????????? ?????? ????????????? ????? ?????????? ? ??????????? ??????????? ?????????
? ???????? ? ????????? ?????????? ????? ?
???????? ????? ??????? ????? ???????? ??????.
????????? ? ??????
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????? ???????? ??????, ? ?????????? ???
??? ? ??????. ?????? ?? ??? ??????????
? ???????? ????? ?????????? ??? ? ????,
?? ?????? ?????? ?????? ????? (N 56є11?, ?
84є25? ? N 56є12?, ? 84є24?). ????? ??? ???????? ????? ???????? ????? ?????????? ??? ??????? ???????????? ????????????? ???????
?????? 2,2 ?? ? ??????? ????? ??????? 11є.
??????? ????? ????? ??????? ? ?????? ??????? ??????? ?????????? ????? 40 ?. ???????? ?????? ????????????? (??????? ????????), ??????????????? ????????????????
????? ????????????? ???????? ?? ?????????
????????? ????? (????. 1).
??? ??????? ??????? ??????? ????????? ?? ?????? ?????? ?????. ? ??????? ?????
??????? (?? 1), ??????????????? ?? ??????
??????????? ??????, ? ????????? ???????????? ???????? ????? ?????? ? ??????????
??????? ???? ??????????? ???? ?????????,
?????????? ???? ??????? ?????????? ???????? ???????? ?????. ??????????? ?????
?????????? ? ???????? ??????? ?????????
????? ? ??????? ????? ?????? (?? 2) ??????? ????????????? ??????? ??? ????? ????
??????? ?????, ? ????? ?????????? ? ???
?????. ?????? ?????????? ???????????????? ??????????? ????? ??? ???? ???????? ?
?? ????????????? ? ?????? ????? ?????????,
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30 %. ??????? ??????? ? 3 ? ????????? ???????????? ? ?????? ????? ??????, ????????
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???????? ????????? ???????-???????? (??
4), ??? ?????????? ???????? ?????. ?????
????? ??????? ????? ???? ????????? ????????? ?????????? ?????? ?? ??????????? ????????? ???, ?????? ??? ????? ??????, ??? ???
??????? ?? ?????? ?????????????? ?????.
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???????? ??????? ???????-??????????????
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????????? ????? ??????? ??????????
??????? ??????? ? 5-7. ???????-????????
????? ?? ????????? ???? (?? 5) ????????????? ?? ?????? ??????????-??????? ?????? ?
?????????? ????????? ???????????. ??????????? ??????? ???????-?????????? ? ??????? ????? ????? ?????? ????? (?? 6) ??????????? ?????????? ??????? ???????? ?????
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????????????? ?????? (?????????, 1974; ???
2004). ???????? ??????? (?? 7) ????????????
????? ????????? ?????????????? ?????????????? ????? ? ???????? ????????, ??????????????? ????????????? ??????. ?? ??????????????? ??? ??????????? ?????????? ?
????? ???????? ?????????????? ????????
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(?? 8) ? ?????? ????????? ? ??????? ??????,
??????????? ? ??????????? ????? ????????? ?????? ?????. ?? ???????????? ??????,
???????? ???????? ?????? ????? ??????????
?? ????? 3,5 ? (????????, 1960). ? ????????
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? ??????????????? ????? ???????? ????? ???????? ????????????? ?? ???????.
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85є06? ? N 56є25?, ? 85є07?) ? ?????????? (N
57є01?, ? 85є55? ? N 56є57?, ? 85є43?), ??? ????
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??????? (155-200 ? ??? ??. ????) ?? ???????????? ??. ????? ? ????, ? ???????? ??????????
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??????, ??????, ??????.
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(????????????? ?, 1997; ??????? ? ??.,
1999), ?????????????? ???????????? ??????? ? ????????????? ????????????? ?? ??????????? ????????????? ?????? (?????????
? ??., 1956). ??? ??????? ??????????? ?????
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1986). ??? ???????? ?????????????? ??????
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187
17,2
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6
214
12,3
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7
308
12,8
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118
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16.5
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19.8
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20.4
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90
13.9
33.3
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152
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22
99
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87
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?
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?? ???? (?????????? 6), ???, ????????, ??????? ? ????????? ????????????? ?????????????? ??? ? ???? ????????.
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(r = -0,51*), ???? (r = -0,22), ???? (r = -0,63*) ?
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? ????? ? ??????? ????? ?????????????? ??????
(?????????? 10) ???????????? ?????? ?? ?????????? ???? ???????? ??????????? ? ??????-
?????? ?????????????. ? ????????? ?????
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(????. 3). ??? ?? ?????, ???????? ?????????
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# 124 #
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??????? ???????????. ????????????? ????? ?
????????????? ??????? ??????????? ?????? ? ?????????? ??????? ??????? ? 4, ???
????????? ???????? ?? ? ? 6-8 (?? ??????????? ???????????).
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? ???? ??????? ?????????? ? ??????? ????????????????. ??????????? ?????????????
????? ??????????? ???? ???????????, ??????
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???????????? ? ? 3 ? 5, ? ??????????? ? ?
4. ?????? ?????????????? ??????? ????????
??? ?????????? ?????????? (??????? ??????? ? 1 ?? 5, ? ? 6 ?? 8) ? ??????????? ?? ?????????? ??????? (???. 2). ??? ???? ????? ?????
??????? ?? ?????? ?????? ? ??????????? ??????????? ??????? ???????? ? 1-5 ?????????? ?? r = +0,63, ? ? 6-8 ?? r = -0,32. ?????
????, ?????????? ????????????? ????? ?????
?????????? 6-8-? ??????? ???????? ?? ??????? ???????????? ?????? ??????? (r = +0,20).
??? ???? ?????????? ??????????? ??????????? ??? ??????? ???????? ??????????? ?????:
??????? ? 70-? ??. ?? ?. ?? ?????? ??????????
?????? ??????? ???????? ????? ???? ?????,
?? ? ? 1975 ?. ??????? ??????????? ??????????
?????????????? (???. 2).
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???????????, ??? ????? ???????????? ???????? ????? ????????? ?????????????, ??? ?
? ?????????????? ???????????? ?????????????? ???????, ???? ????????? ????????????
«????????-????????» ???????? ?????????????
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?? ??????? ????????? ?????.
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# 125 #
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250
240
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220
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180
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1880 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010
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??? ?????? ????????????, ?????????????
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?? ??????, ?????????? ? ???? (????????, 1980),
???????? ?????? ?????? ????????? ????? ??
?????? ???????? ?? ?????????? ????? ?????????????? ?????? ????? ? ?????-????????
?????????????? ?????? (???????, 1969).
?? ????? ????? ???????? ??????? ????????. ??? ????????, ??????? ????????? ?
???????????? ?????????? ???????, ?? ???????? ??????? ??????? ? ????????????? ?????? ?????????? ????, ?????? ???? ??????
??????????? ?????? ? ??????????? ?? ?????????? ?????????????? ??????. ???????????
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(???. 4) (???????, 1986).
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? ???? ???????? ?????? ????? ????????????
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?????? ??????????? (????????? ? ??., 1993).
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??????? ??? ?? ??????????? ???????? ?????, ????? ? ?????????????? ?????? ???????? ??????? ???? ?????????????? ??? ?????
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????????????? ?????.
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? ????? ?????? ????. ?? ?????? ?????? ????? ????????? ????? ????? ??? ??? ???????
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??????????? ? ?????????????? ? ?????????
??????????? ?? ??????, ????????? ? ??????????? ?????????? ????? (?????, 1961; ?
??.). ??????? ??? ????????????? ???? ????????? ??????????? ????????????? ??????????????? ????????????, ?????????? ? ?????? ??????? ????????? ??????? ???????? ??
??????????? ? ????? ???.
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?????????, ??? ?????????? ????????????
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(???????? ? ??., 1999). ?????????? ??? ????????? ?? ???????? ??????????? ?????????
?????? ?????????????????? ?????????? ?
???????? ????? ???????????????? ???????, ??
# 127 #
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??????????, 1994) ? ??????????? ????????? (???????, 1986)
???? ??????? ? ???????? ?????? ???? (?????,
1961). ???????? ????? (????????, 1961), ???
????????????? ??????? ??? ????? ??????????? ???????????? ? ??? ??????????????, ???
???????? ???????? ????? ??????? ?? ??????
????? ? ???????? ???????. ?????? ? ???????????? ????? ?????? ???? ???????????? ?????????? ????? ????? ?????????? ????????
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??????????? ??? ???????? ?????? ????? ? ?????????????? ???? ?????. ? ???????? ?????????? ??????? ? ????????????????? ??????????
?????????, ??????? ?? ????????? ???????? ?
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? ?????? ?? ????. ? ???????? ???????????
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?? ?????? ? ????? ?????? ????.
?????????? ???? ????? ???????? ?????
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?????? ? ????? ?????????????? ???????????.
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????? ???? ??????????? ??????? ???????????? ????? ????. ? ????????????? ????? ????
??????? ?? ????????? ????? ??????????????
????? ????? ?????????? ???. ???????? ?????
????????? ???????? (???. 4) ????????? ??????
??????? ?????????? ?? ?????? ?????? ????
??????????? ? ?????? ????? ???? (?????????, 1972; ??????????, 1994). ??????? ? ?????
??????????? ???????????, ??????????? ???
???????????? ??????? ??????, ?? ?????????
????? ???????? ????? ???? ?????????? ???.
????????? ????? ???????? ????? ????? ????
? ?????? ?? ??????????? ? ?????????????
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???? ????????? ?????? ???????. ???? ????
?????????? ?? ??????????? ???????????, ?
??????? ???? ??????? ?????????? ???? ?????
????????? ????? ????.
??? ??????????? ??????? ??????????????? ????? ???????? ? ?? ??????? ?? ????????? ????????????? ?????????? ????????
???????? ????? ????????????? ? ??????????????? ????????? ????????????? ???????,
?????? ??????????? ????????? ???????? ?????????????? ???????? ????????? ? ?????????
???????, ? ??????????? ????????? ????????
???????????? ???????? ? ? ????????? ?????????? ? ??????? ?????????????? ??????. ?
????? ??? ????????? ? ????? ??????????????
???????????????? (Lloyd, Bunn, 2007). ?????
????, ?? ????????????, ??? ????????????? ??????? ????????, ?????????????? ? ????????
?????????????? ?????????? ???????? ??????,
?? ?????????? ??????????? ???????, ??????? ?
?????????? ??????????? ?????????? ?????
????, ?????????? ? ????? ?? ??????????????? ??????????? ??? ????????.
??????????
????????? ?????????? ???????????
??????????, ??? ???? ? ??????????? ??????????? ??????????? ?? ??????? ??????????? ????? ?????? ?? ??????????? ? ??????
?? ?????????? ?????????????? ???????, ???
???????????????? ??????????? ????????????? ????????? ????? ???????? ????. ???????? ???????????? ????????? ???? ???????
? ??????????? ??????????? ????? ????????
?? ?????????????? ????????? ???????? ??????? ?? ?????????? ???? ???????? ? «?????» ?
«??????» ????, ?????? ? ???????? ????? ??????? ???? ?????????? ??????? ? ????????
?????? ??????, ? ?? ?????? ????? ???????????
??? ??????? ??????????? ? ????????? ???????. ? ???????? ?????????? ??????? ?????????? ????????? ????? ?????? ?? ?????????? ????? ??????? ???????????? ???? (???,
????). ? ???????? ??????????? ??????????
????????????? ??????? ????????? ????
?????? ????, ?????????????? ???????????
????????????? ?????? ????????? ???, ???
??????? ????????????? ???????? ???????
???????? ?? ??????????. ? ????????????? ????? ???? ??????? ?? ????????? ????? ?? ?????????????? ?????? ??????? ????, ?????????
? ???????????? ???? ?????????????? «???????» ????? ???? ?????????? ???. ??????????? ????????? ???????? ??????????????
???????? ????????? ? ????????? ???????, ?
??????????? ????????? ???????? ???????????? ???????? ? ? ????????? ?????????? ? ??????? ?????????????? ??????.
?????? ??????????
????????? ?.?., ???????? ?.?. (2001) ? ???????????? ????????????? ??????? ??????
? ?????? ? ????? ?? ???????? //??????? ????????? ??????. ???. 24 / ??? ???. ?.?. ??????.
?????: ???. ???. ??-?, ?. 56-59.
??????? ?.?. (1986) ???????? ???? ??????????? ????????? ? ??????? //????? ?????.
???????????. ??. 39, ?. 2. ??????????, ?. 24-38.
???????? ?.?. (2003) ??????? ???????? ???????? ?? ??????? ????? ???????????? ? ??????? ????? ?????? //????????? XI ?????? ???????? ????????????? ????????. ???????????, 1727 ??????? 2003 ?., ?. 15-16.
??? ?.?., ???????? ?.?., ????? ?.?. (2004) ????? ???????? ?????????. ?????: ???, 160 ?.
??????? ?.?., ?????? ?.?., ?????? ?.?. (1996) ??????????????????? ???????????? ?
?????-????????? ??????????. ???????????: ?????, 246 ?.
# 129 #
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?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
???????? ?.?., ??????? ?.?. (1999) ?????????? ??????? ????? ???????????? ?? ???????
? ?????????. ?????? ? ???????????? ???? ? ????? ????? ??????????? ??????????????????. ????????? ?????????. ?.: ????, ?. 99-101
???????? ?.?., ??????? ?.?., ???????? ?.?., ???????? ?.?. (2002) ??????????????????? ?????? ??????????? ???????? ????? ?????????? (Pinus sibirica) ? ?????-?????????? ???????? ?. ????? (??????????? ?????) //???????? ?????????????? ?????????? ? ????????????
????????? ??????? ? ????????? ????? ? ??????? ???????? ??????. ???????????: ???-?? ???
?? ???, ???. 1: 71-78.
?????? ?.?., ?????????? ?.?. (1976) ???????? ?????? ???????? ????? ? ????? ? ?????????????????? ???????????? ? ????????? ????? ???????? ????? //??????????? 4: 56-62.
?????????? ?.?. (1994) ? ??????????????? ????????? ? ???????? ??????? Pinus sibirica
(Pinaceae) //???????????? ??????. ?. 79, N5: 63-71.
????? ?.?. (1961) ??????????? ????????? ?????? ? ???????? ???????. ???????????? ?
????????? ????????????. ?.: ???-?? ?? ????, 210 ?.
??????? ?.?., ???????? ?.?., ???????? ?.?. (1999) ???????????? ???????????? ?? ???????????????? ???????? ????. ?????: ???-?? ?? ???, 39 ?.
??????? ?.?. (1969) ???????? ?????? ????????. ?.: ?????, 312 ?.
????????????? ???? ?????? (1997) ?.: ????? ????????? ???????? ??. ?.?. ?????????,
236 ?.
?????????? ?.?. (1992) ?????????? ??????. ???.: ?????, 359 ?.
??????? ?.?. (1997) ?????????? ????????? ??????? ? ??? ????????? ???????????. ???????????: ?????????, 102 ?.
????????? ?.?., ?????????? ?.?. (2000) ?????? ?????????? ??????? ? ????? ?????
???????-????????? ??????? (????????? ? ???????????????-????????? ????????) / ???. ???.:
?.?.??????, ?.?.????????. ??-? ????????? ?? ???. ???????????: ???-?? ?? ???, 216 ?.
?????? ?.?. (1999) ??????? ??????? ?? ???????? ??????????? ???????? ??????? ? ?????????????? ??????? ??????? //??????????? 6: 14-21
????????? ?.?. (1972) ????????????? ?????? ??????????? ????? ??????????. ???????????:
?????, 272 ?.
???????? ?.?., ??????? ?.?. (2004) ??????? ????????????? ???????? ? ????????????
?????? ?????????? ???? ??????????? ?????? ?? ?????????? ??????? ??????????? ? ?????
//??????????? 6: 3-13.
?????????? ?.?., ??????? ?.?. (1999) ????????????????????? ?????????????? ??????????? ????????? (Larix sibirica Ldb.) ?? ??????? ??????? ???? ? ?????? ????? //????????? ????????????? ?????? 2: 145-152.
????????? ?.?., ?????????? ?.?. (1999) ???????????? ????????????? ???????? ??????????? ???????? ??????????? ? ?????? ????? //??????????? 6: 22-32.
???????? ?.?. (1963) ?????? ???????? ????? ?????????? ??? ? ????. ???????????? ???? ?
?????? ??????. ?.: ???-?? ?? ????, ?. 65-95.
???????? ?.?., ??????? ?.?. (1999) ?????????? ????????????? ???????? ????? ????????
?????? //?????? ? ???????????? ???? ? ????? ????? ??????????? ??????????????????. ????????? ???????????. ?.: ????, ?. 79 ? 82.
# 130 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
???????? ?.?., ?????? ?.?. (1975) ??????? ???????????? ?????? ?? ?????????? ???????
???????? ? ????????? ???????? ?? ????????????? //???????? 6: 30-35.
????????? ?.?. (1963) ?????? ?????????????. ?.: ???-?? ?? ????, 192 ?.
????????? ?.?., ???????? ?.?., ??????? ?.?., ??????? ?.?. (1956) ????????????? ??????
???????? ?????? ?? ????????????? ???????. ?.: ??????????, 472 ?.
???????? ?.?. (2003) ???????????? ????????? ?????? ????????? Pinus sylvestris ????????? ???????? ?????????????. ????????? XI ?????? ???????? ????????????? ????????. ???????????, 17-27 ??????? 2003 ?., ?. 92-93.
???????? ?.?. (1980) ??????????? ????? ?????? ??????? ? ?????????? ???????? ??????
???? ????????????? //??????????? 1: 89-93.
???????? ?.?. (1961) ???? ??????????? ? ???????? ?????????????????? ?????. ?.: ???-??
?? ????, 159 ?.
????????? ?.?., ??????? ?.?., ??????? ?.?. (1993) ???? ? ????????? ??2 ? ?????? ????????. ?.: ?????, 256 ?.
?????? ?.?. (1986) ???????????????? ??????? ??????? ???? ?? ?????. ?.: ?????, 136 ?.
???????? ?.?., ???????? ?.?., ??????? ?.?., ???????? ?.?., ???????? ?.?. (1999) ???????
????????????? ??????? ??????????? ??????? ?? ??????? ????? //??????????? 5: 41-49.
Blyakharchuk T.A., Sulerzhiysky L.D. (1999) Holocene vegetational and climatic changes in the
forest zone of western Siberia according to pollen records from the extrazonal palsa bog Bugristoye.
Holocene. V.9: 621-628.
Crawford R.M.M., Jeffree C.E., Rees W.G. (2003) Paludification and forest retreat in northern
oceania environments. Annals of botany. V. 91: 213-226.
Jones P.D., Briffa K.R. (1992) Global surface air temperature variations during the twentieth
century: Part 1. Spatial, temporal and seasonal details. Holocene. V. 2, N 2: 165-179.
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variability. Environ. Res. Lett. V. 2.
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summer air temperature at high latitudes in the northern hemisphere for the last 1.5 ka: a comparative
analysis of the data on annual tree rings and ice cores. Doklady Earth Sciences, 359(2): 267?270.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
Effects of Climatic Factors on Radial Growth
of Siberian Stone Pine and Siberian Larch in Sites With
Different Soil Humidity in the South of Western Siberia
Svetlana N. Velisevich and Oleg V. Khutornoy
Institute of Monitoring of Climatic and Ecological Systems of SB RAS
10/3 Akademicheskiy, Tomsk, 634055 Russia
The effects of temperature and precipitations on the formation of annual rings in stem wood of Siberian
stone pine (Pinus sibirica Du Tour) and Siberian larch (Larix sibirica Ledeb.) in sites with different
soil humidity are discussed. It is shown that these two species essentially differ on reaction of radial
growth to temperature and precipitations during the growth period that is presumably caused by
features of a seasonal cycle of needle development. The formation of a xylem year layer at deciduous
larch depends on a favorable combination of weather conditions during two periods ? «growth»
and «activity» of needles. Damp and moderate-warm weather is necessary for young needles during
growth period but its activity is more productive at high temperature and moderate precipitations. At
evergreen Siberian stone pine the growth of xylem on the initial stage is not limited by young needles
growth due to «activity» of alive needle developed during previous years. The xylem growth of trees of
the unwaterlogged sites depends on precipitations during growth season, and in waterlogged sites the
slight positive influence is caused by summer temperature.
Keywords: Siberian stone pine, Siberian larch, radial increment, temperature, precipitation, soil
humidity.
???????? ??????? ?????????
?????? ???????????.
???????? ?????: ???? ?????????, ??????????? ?????????, ?????????? ????, ???????????,
??????, ????????? ?????.
????????
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???????, ??? ?????????? ? ?????????? ????????? ????????? ??????????? ? ????????? ????????? ??????? ????????????????? ?
?????? ????????????? ????? ? ?????????
??????? ??? (Jones, Briffa, 1992; ??????????,
1992; ???????, 1997; ?????????, ????????,
2001; ? ??.). ? ?????? ???? ????????? ?????
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?????????? ?? ????? ?????????? ????
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??? ??????????? ???? ????????? ????????
????????? ????????, ??????? ????????? ??????????? ????? ???????? ???????? ???????-
Corresponding author E-mail address: velisevich@imces.ru
© Siberian Federal University. All rights reserved
# 117 #
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?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
???? ??????????, ??????????????? ? ???????? ???????.
? ??????? ?????????? ???????? ???????????? ??????????? ???????? ?? ???????
???????? ???????? ?? ?????????? ????
???????, ?????????? ? ???? ???????????
???????????? ? ?????????????? ? ???????????? ??????? ? ????? ?????????? ??????????
?????????? ??? ????????????? ???????. ?
????????? ??????????? ?????????, ???????????? ?????????? ????????????????? ?
??????????? ???????????? ??? ????-??????,
???????????, ??? ??????????? ???? ? ????
?????????? ?? ?????????? ??????? ? ???????????? ??????? (??????, 1986; ??????????, ???????, 1999; ?????????, ??????????,
1999; ? ??.) ? ?? ???????? ??????? ??????????????? (????????, ??????, 1975; ???????
? ??., 1996; ??????, 1999). ? ????????? ?????
?????????? ?? ???? ???????? ????????????
?????? ?????? ? ????? ? ???????? ???????
??????: ???????? ????????????? ??????????? ? ???????????? ???? ? ??????? ???????
(Vaganov et al., 1998; ???????? ? ??., 2002).
? ????? ????? ????????????? ??????
?? ??? ?????? ???????????? ???? ????????,
??? ?? ??????????? ????????. ???? ????????
? ????????? ??????? ??????? ?? ????????????
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???????? ? ???????? ??????? ????????????
???? ????? ???????????? ????? ????????????, ?????????? ??????????? ??????? ????????????? ??? ?? ??????? ????????? ?????.
?? ??????? ????? ???? ???????? ??????? ??????????? ????????? ???????? ???????????
???????? ? ???????? ??????????? ? ???????????? ????????????? (????????, ???????,
1999; ????????, 2003; ????????, 2003), ? ????? ??????? ???????? ???????? ?? ??????
?????? ??????? ? ??????????? ?? ?????????
?????????? ???? (??????, ??????????, 1976;
?????????, ??????????, 2000; ? ??.). ??? ??
?????, ? ??????????? ??????????? ???????
????????????, ????????????? ??????????
???????????? ??????????? ????? ?? ????????? ???????????? ???????????????? ??????
???? ?????????? ?????????????. ???????????? ?????????? ????? ???????? ???????????? ???????? ???????? ? ????????? ????????, ???????????? ???? ????????.
??? ??????????? ????? (?????? Strobus),
? ??????? ??????????? ? ???? ?????????,
??????????? ?????? ???????? ???????? ????????????? ???????? ? ?????? ??????????
?????? ? ???????? ???????. ??????, ? ??????? ?? ????? ????????? ?????????????, ????
??? ? ???????????? ????????????? ????????
?????????, ?????? ???????????????? ?? ????????? ?????????? ???????? ?????? (???
? ??., 2004). ??????????? ????? ??????????
???????????? ? ???????? ?????????? ????????? ????? ?????? ??? ?????????? ???????? ??? ????????????. ???????????????
????????????? ?????????? ???????? ??????
(Blyakharchuk, Sulerzhiysky, 1999; Crawford et
al., 2003; ????????, ???????, 1999), ? ?????
????????????? ????????? ???????? ??????????????? ???? ? ?????? ? ???? ?? ????????
?????? ??????? ??????????, ?? ?????? ???????????? ???????????? ?????????? ????????????. ????? ????, ????, ??? ????????????
???????????? ??? ????????? ????? ???????????? ?????? ???????? ? ????? ?????? ???????????, ? ??? ????? ? ??? ????? ????????,
??? ???????? ???????????? ???????? ???????? ????????????? ????????? ?????.
???????????? ??????? ???????? ? ??????????? ??????????? ?????????, ???????????? ?? ???????????? ???????, ? ?? ?????
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90,3
89,0
86,6
85,0
81,2
70,3
78,2
71,8
66,5
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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????????? ? ????????? ????? ? ???????
????????????? ?????? (?????????, 1974; ???
2004). ???????? ??????? (?? 7) ????????????
????? ????????? ?????????????? ?????????????? ????? ? ???????? ????????, ??????????????? ????????????? ??????. ?? ??????????????? ??? ??????????? ?????????? ?
????? ???????? ?????????????? ????????
????????? ????????? ??????, ???? ??????? ??
??????? 40-70 ?? ????????? ????????????
?????. ????? ???????? ?????????? ? ?????
??-?? ???????? ???? ? ???????? ??????????
?????????.
??????? ????? ?????????????? ???????
(?? 8) ? ?????? ????????? ? ??????? ??????,
??????????? ? ??????????? ????? ????????? ?????? ?????. ?? ???????????? ??????,
???????? ???????? ?????? ????? ??????????
?? ????? 3,5 ? (????????, 1960). ? ????????
?????????? ????????? ??????? ??????????
??????? ?????? ??????? ???????? ? ????????? ????. ??????? ??? ?????????????????
? ??????????????? ????? ???????? ????? ???????? ????????????? ?? ???????.
???????? ??? ??????? ??????????? ?????
??????????? ????????? ???????? ? ???? ??????? ??????? ??????? ? ??????? (N 56є28?, ?
85є06? ? N 56є25?, ? 85є07?) ? ?????????? (N
57є01?, ? 85є55? ? N 56є57?, ? 85є43?), ??? ????
???????? ????? ??????? ???????? ? ????????? ????????-???????????? ????????. ?
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?? ??????????, ???????? ?? ?? ???????? ??
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???? ??????????? ????????????? ????????
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?? ???????? ?????, ?? ??????? ???????? ? ????????????, ? ????? ??????????? ????????? ?
???????????? ????????? ???????? ????-
???????? ?????. ??????? ??? ??????? ??? ???????????? ????????????? ????????????? ?
??????? ? ?????????????, ??????????? ? ?????????? ?????????? ????? (????. 1). ?????
????????????? ?????????? ? ??????????
??????? (155-200 ? ??? ??. ????) ?? ???????????? ??. ????? ? ????, ? ???????? ??????????
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??????, ??????, ??????.
??? ????????????? ??????????? ??
??????????? ???????????? ???????, ??????????? ??????????????? ???????? ????
(????????????? ?, 1997; ??????? ? ??.,
1999), ?????????????? ???????????? ??????? ? ????????????? ????????????? ?? ??????????? ????????????? ?????? (?????????
? ??., 1956). ??? ??????? ??????????? ?????
?????? ???? ????? ????? ?? ???? ????????
?? ?????? 1,3 ? ?? ????? ????? ? 240 ????????
??????????? ????????? ? 230 ???????? ?????
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????????? LINTAB ? ??????? ????????????
??????????? TSAP. ? ????? ???????????
????????, ????????????? ????????? ????????, ??? ?????????????? ????????? ????
?????????? ????????? ????????????? ? ????????????? ???????? ? ??????? (??????,
1986). ??? ???????? ?????????????? ??????
????????????? ?????????? ? ??????????
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???????????? ?. ?????? ?? ????????? 100 ???.
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89
20,0
33,4
155
21,2
40,5
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3
156
21,9
42,0
4
292
25,1
50,1
5
187
17,2
22,8
6
214
12,3
23,8
7
308
12,8
23,0
8
118
5,3
14,6
9
85
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17.0
31.0
10
77
10.4
22.3
11
89
10.6
18.5
12
85
13.3
24.1
13
80
16.5
36.2
14
89
16.2
35.0
15
72
12.9
38.2
16
75
16.3
36.4
17
80
19.8
36.0
18
74
20.4
22.2
19
74
11.2
24.9
20
90
13.9
33.3
21
152
8.8
18.6
22
99
14.8
26.5
23
87
17.0
33.7
24
140
11.0
19.2
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100 ??. ???????? ????????????? ??????????
???? 0,41 ????????? ????????? ? ???????????? 0,95.
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????????????? ?????????? ? ?????????? ?????? ???????? ??????????? ????????? (???.
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?????? ????????? ?? ????????? ????????
????????. ? ???????? ?????????? ???????
?????????? ????????? ????????? ??????????? ????????????? ??????? ??????? ? ?????? ?????????????? ?????? (?????????? 3-6).
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?????????????, ???? ?????? ??????????????
???? ???????? ????????. ??? ????????, ???????? ??? ??????????? ???????????????
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????????????? ?????? ?? ?????? ??????????
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?
1
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0
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-1
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
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2
3
4
5
6
7
8
9
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12
13
14
15
16
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10
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12
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14
15
16
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0
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1
2
3
4
5
6
7
8
???. 1. ???????????? ?????????? ??????????? ???????? ?????? ???????? ??????????? ????????? ?
???????? ? ????????????? (?), ??????????? (?) ? ?????????? (?) ?????????? ????? ? ???????????? ?
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??????? ????????????? ?????, 4 ? ? ??????, 5 ? ? ???, 6 ? ? ????, 7 ? ? ????, 8 ? ? ???????, 9 ? ? ????????;
10 ? 16 ? ??????? ???????????: 10 ? ?? ????????????? ?????, 11 ? ? ??????, 12 ? ? ???, 13 ? ? ????, 14 ? ?
????, 15 ? ? ??????? ? 16 ? ? ????????. ???????? ?????????? ?????????? ??????????? ????????????
??????????
?? ???? (?????????? 6), ???, ????????, ??????? ? ????????? ????????????? ?????????????? ??? ? ???? ????????.
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(r = -0,51*), ???? (r = -0,22), ???? (r = -0,63*) ?
??????? (r = -0,37) ??????????, ??? ??????????
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?? ? ??????????. ??????? ?????????? ??????????? ? ?????? ? ??? (?????????? 11 ? 12),
? ????? ? ??????? ????? ?????????????? ??????
(?????????? 10) ???????????? ?????? ?? ?????????? ???? ???????? ??????????? ? ??????-
?????? ?????????????. ? ????????? ?????
????????????? ? ?????? 30-?????????????
???? ????????? ? ????, ??????? ???????
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???? ????? ? ???????????? ????????? ?? ???????? ??????????? ????? ????????.
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?? ?????? ????? ??????????? ?? ???????????? ??????????????? ??????????, ??? ?????????? ?????????? ???????? ??????????
(????. 3). ??? ?? ?????, ???????? ?????????
? ????????????? ????? ??????????? ???? ?
???????????? ??????? ???????? ? 1, 2, 3 ?
# 124 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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2
3
4
5
6
7
8
IV
+0,1
1
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-0,02
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V
0
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-0,01
-0,09
VI
-0,26
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-0,21
-0,01
-0,19
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+0,07
+0,05
VII
-0,26
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0
-0,02
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-0,04
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VIII
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IX
+0,13
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-0,12
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-0,08
-0,09
IV- IX
-0,3
-0,15
+0,08
+0,24
-0,19
-0,18
-0,04
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IV
+0,01
-0,04
-0,05
-0,02
+0,02
+0,16
-0,18
-0,04
V
+0,09
+0,01
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+0,06
-0,11
-0,14
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VI
+0,24
+0,25
+0,27
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+0,31
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VII
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0
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IX
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+0,16
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IV- IX
+0,16
+0,16
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5, ?????? ?????????? ??????? ??????? ? 1
??????? ???????????. ????????????? ????? ?
????????????? ??????? ??????????? ?????? ? ?????????? ??????? ??????? ? 4, ???
????????? ???????? ?? ? ? 6-8 (?? ??????????? ???????????).
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? ???? ??????? ?????????? ? ??????? ????????????????. ??????????? ?????????????
????? ??????????? ???? ???????????, ??????
?????????? ????? ?????? ?????? ????, ? ???????????? ??????? ???????? ? 1, 2, 3, 4 ? 5,
???????????? ? ? 3 ? 5, ? ??????????? ? ?
4. ?????? ?????????????? ??????? ????????
??? ?????????? ?????????? (??????? ??????? ? 1 ?? 5, ? ? 6 ?? 8) ? ??????????? ?? ?????????? ??????? (???. 2). ??? ???? ????? ?????
??????? ?? ?????? ?????? ? ??????????? ??????????? ??????? ???????? ? 1-5 ?????????? ?? r = +0,63, ? ? 6-8 ?? r = -0,32. ?????
????, ?????????? ????????????? ????? ?????
?????????? 6-8-? ??????? ???????? ?? ??????? ???????????? ?????? ??????? (r = +0,20).
??? ???? ?????????? ??????????? ??????????? ??? ??????? ???????? ??????????? ?????:
??????? ? 70-? ??. ?? ?. ?? ?????? ??????????
?????? ??????? ???????? ????? ???? ?????,
?? ? ? 1975 ?. ??????? ??????????? ??????????
?????????????? (???. 2).
?? ????????? ????????? ?????? ??
???????????, ??? ????? ???????????? ???????? ????? ????????? ?????????????, ??? ?
? ?????????????? ???????????? ?????????????? ???????, ???? ????????? ????????????
«????????-????????» ???????? ?????????????
?????????????? ??????? (???. 3). ???????????? ?????????? ???????, ?????? ?????????????? ?? ????? ? ??????????????? ????????? ???????????????? ??????? ??????? ????????
?? ??????? ????????? ?????.
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??? ???????? ?????????? (???????, ???????)
? ??????? (????????????? ???????, ??????)
# 125 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
260
250
240
230
220
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??????? 1-5 ??/5
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210
200
190
180
170
160
1880 1890 1900 1910 1920 1930 1940 1950 1960 1970 1980 1990 2000 2010
????
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????, ??, ????????? ?????????? ??????????? ? ????)
1
2
3
5
4
6
7
8
???. 3. ???????????? «????????-????????» ????????????? ????????????? ?????????????? ??????? ??
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
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??? ?????? ????????????, ?????????????
??????? ??????????? ?????? ? ???????? ??
?? ??????, ?????????? ? ???? (????????, 1980),
???????? ?????? ?????? ????????? ????? ??
?????? ???????? ?? ?????????? ????? ?????????????? ?????? ????? ? ?????-????????
?????????????? ?????? (???????, 1969).
?? ????? ????? ???????? ??????? ????????. ??? ????????, ??????? ????????? ?
???????????? ?????????? ???????, ?? ???????? ??????? ??????? ? ????????????? ?????? ?????????? ????, ?????? ???? ??????
??????????? ?????? ? ??????????? ?? ?????????? ?????????????? ??????. ???????????
??????? ????? ?????? ?? ??? ???????? ???????? ??????????????? ????? ???????? ???
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? ????????? ????????? ???????????????
???? ?????. ??????? ????? ?????????????
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?????? ??????? ? ?????? ????????? ????????
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?? ??? ???????? ?????? ?????? ????????? ?????? ??????? ? ??????????? ????????? ?????????? ? ????? ?????? ??????
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???????????? ??????? ??????, ??????????
? ?????? ?????? ??? ? ? ??????? ?????? ????? ??????? ????????? ? ?????? ??????????
(???. 4) (???????, 1986).
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????????? ??????, ?????? ? ??????. ??????
? ???? ???????? ?????? ????? ????????????
? ???????? ?????????? ????????? ? ???????
?????????? ? ? ????? ????, ??????? ?????????? ?????, ???? ???????????? ?? ????????? ?
?????? ??????????? (????????? ? ??., 1993).
?????? ?????????????? ??????? ?????????
? ??????? ????? ???????, ?????? ? ???????,
??????? ??? ?? ??????????? ???????? ?????, ????? ? ?????????????? ?????? ???????? ??????? ???? ?????????????? ??? ?????
????. ????????, ?????? ???? ??????????? ????????????? ??????????? ??????????? ?????
?????? ?? ??????? ?????? ? ????, ??????????? ? ???????? ??????????? ????????? ????
????????????? ?????.
???????? ?? ???? ???????? ????????????? ??????? ?? ?? ?????????? ???? ??????
? ????? ?????? ????. ?? ?????? ?????? ????? ????????? ????? ????? ??? ??? ???????
??????????? ?????????????? ????????????? ???? Larix. ??????????? ??????????? ??
??????????? ? ?????????????? ? ?????????
??????????? ?? ??????, ????????? ? ??????????? ?????????? ????? (?????, 1961; ?
??.). ??????? ??? ????????????? ???? ????????? ??????????? ????????????? ??????????????? ????????????, ?????????? ? ?????? ??????? ????????? ??????? ???????? ??
??????????? ? ????? ???.
??????????? ??????????? ? ????????
????? ???????????? ??????? ??????? ???????????, ??????? ? ???? ??????? ????????
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?????????, ??? ?????????? ????????????
?????????????? ??????????? ???????????
?????????? ?????? ? ???????? ??????? ????
(???????? ? ??., 1999). ?????????? ??? ????????? ?? ???????? ??????????? ?????????
?????? ?????????????????? ?????????? ?
???????? ????? ???????????????? ???????, ??
# 127 #
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??????????, 1994) ? ??????????? ????????? (???????, 1986)
???? ??????? ? ???????? ?????? ???? (?????,
1961). ???????? ????? (????????, 1961), ???
????????????? ??????? ??? ????? ??????????? ???????????? ? ??? ??????????????, ???
???????? ???????? ????? ??????? ?? ??????
????? ? ???????? ???????. ?????? ? ???????????? ????? ?????? ???? ???????????? ?????????? ????? ????? ?????????? ????????
???, ????????? ???? ? ??????????? ???????
????????? ? ??????, ???, ????????, ??????????? ????????????? ??????? ??????????? ???????? ?? ????????? ??????????? ? ????.
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?????? ??????, ? ?? ?????? ????? ???????????
??? ??????? ??????????? ? ????????? ???????. ??? ??????????? ??????????????? ?????? ???? ????????????? ??????? ?? ??????
?????? ??????? ????????? ???????? ??????,
??????????? ??? ???????? ?????? ????? ? ?????????????? ???? ?????. ? ???????? ?????????? ??????? ? ????????????????? ??????????
?????????, ??????? ?? ????????? ???????? ?
????????? ????? ? ?????, ????? ?????? ?? ?????????? ????? ??????? ???????????? ???? ?
? ?????? ?? ????. ? ???????? ???????????
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???????? ????????????? ?????? ?????????
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?? ?????? ? ????? ?????? ????.
?????????? ???? ????? ???????? ?????
????????? ?? ??????????? ? ??????? ?? ?????????? ?????????????? ???????, ??? ?????????? ? ?????????? ?????? ??????????????
?????? ? ????? ?????????????? ???????????.
????????? ??????????? ??????? ???????????
???? ?? ?????? ????, ???, ????????????????,
????? ???? ??????????? ??????? ???????????? ????? ????. ? ????????????? ????? ????
??????? ?? ????????? ????? ??????????????
????? ????? ?????????? ???. ???????? ?????
????????? ???????? (???. 4) ????????? ??????
??????? ?????????? ?? ?????? ?????? ????
??????????? ? ?????? ????? ???? (?????????, 1972; ??????????, 1994). ??????? ? ?????
??????????? ???????????, ??????????? ???
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????? ???????? ????? ???? ?????????? ???.
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? ?????? ?? ??????????? ? ?????????????
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????? ?????? ????? ? ???????? ???????? ??-
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???????????????? (Lloyd, Bunn, 2007). ?????
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????). ? ???????? ??????????? ??????????
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??????????? ????????? ???????? ???????????? ???????? ? ? ????????? ?????????? ? ??????? ?????????????? ??????.
?????? ??????????
????????? ?.?., ???????? ?.?. (2001) ? ???????????? ????????????? ??????? ??????
? ?????? ? ????? ?? ???????? //??????? ????????? ??????. ???. 24 / ??? ???. ?.?. ??????.
?????: ???. ???. ??-?, ?. 56-59.
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???????????. ??. 39, ?. 2. ??????????, ?. 24-38.
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??? ?.?., ???????? ?.?., ????? ?.?. (2004) ????? ???????? ?????????. ?????: ???, 160 ?.
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Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ?????????, ?.?. ????????. ??????? ????????????? ???????? ?? ?????????? ???? ????? ? ????????????
Effects of Climatic Factors on Radial Growth
of Siberian Stone Pine and Siberian Larch in Sites With
Different Soil Humidity in the South of Western Siberia
Svetlana N. Velisevich and Oleg V. Khutornoy
Institute of Monitoring of Climatic and Ecological Systems of SB RAS
10/3 Akademicheskiy, Tomsk, 634055 Russia
The effects of temperature and precipitations on the formation of annual rings in stem wood of Siberian
stone pine (Pinus sibirica Du Tour) and Siberian larch (Larix sibirica Ledeb.) in sites with different
soil humidity are discussed. It is shown that these two species essentially differ on reaction of radial
growth to temperature and precipitations during the growth period that is presumably caused by
features of a seasonal cycle of needle development. The formation of a xylem year layer
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