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90.Журнал Сибирского федерального университета. Сер. Биология №2 2012

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Журнал Сибирского федерального университета
2012
Journal of Siberian Federal University
5 (2)
Биология
Biology
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Editorial Advisory Board
Chairman:
Eugene A. Vaganov
Members:
Josef J. Gitelzon
Vasily F. Shabanov
Andrey G. Degermendzhy
Valery L. Mironov
Gennady L. Pashkov
Vladimir V. Shaidurov
Vladimir V. Zuev
CONTENTS / СОДЕРЖАНИЕ
Manfred Bцlter and Lyubov R. Izmest?yeva
Is there a Microbial Loop in Lake Baikal?
? 112 ?
В.В. Громов
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Editor-in-Chief:
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Founding Editor:
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Editorial Board:
Managing Editor:
Olga F. Alexandrova
Executive Editor for Biology:
Nadezhda N.Sushchik
Ж.А. Черняев
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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
Mikhail G. Karpinsky
Liliana Zalizniak
Е.Н. Науменко, М.М. Хлопников, Л.В. Рудинская
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Journal of Siberian Federal University. Biology 2 (2012 5) 112-120
~~~
??? 574.5
Is there a Microbial Loop
in Lake Baikal?
Manfred Bцltera* and Lyubov R. Izmest?yevab
a
Institute for Polar Ecology, University Kiel,
1-3 Wischhofstr, Kiel, 24148 Germany
b
Scientific Research Institute of Biology
of Irkutsk State University,
PO Box 24, Irkutsk-3, 664003 Russia 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
The occurrence of a microbial loop in the pelagic environment is discussed with regard to hydrological
and meteorological effects as well as to the planktological states during the seasons in lake Baikal.
Strong local conditions due the melting ice and the sequence of the spring bloom indicate close local
interactions between phytoplankton and bacteria. However, missing reports on micro-zooplankton
and continuous reports of the roles of the individual organism groups involved in the microbial loop
make is difficult to establish the way of matter flow in the lake as a general path. Further, sinking rates
of diatoms do not allow a remineralisation of plankton-C and ?N in the deep water in its upper photic
levels. Thus, microbial mineralized nutrients will stay for long time in the abyssal area of the lake,
which can be regarded responsible for its oligotrophic nature in the upper layers.
Keywords: Lake Baikal, limnology, microbial loop.
Introduction
The pelagic system of Lake Baikal is
governed by a strong seasonality, great depth
and an enormous water body and counts as the
largest limnological environment on the Earth.
The lake stretches more than 600 km from north
to south and thus covers different climatic zones,
strongly expressed by the shift in the timing of
ice melting in spring / early summer. Another
special character is the isolated locality with only
small exchange to other water bodies, thus, many
*
1
endemic species at all trophic levels evolved.
This has formed not only special organisms
but also raises the question whether this system
has also peculiarities in its ecology and trophic
interactions.
A special topic arises about the functioning
of the lower organism?s food web and its role in
production and mineralization. Two scenarios
about the pelagic biological processes can be
discussed, on the one hand, the traditional
sequential food web inclusive its detritus food
Corresponding author E-mail address: mboelter@ipoe.uni-kiel.de
© Siberian Federal University. All rights reserved
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Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
web (Wetzel, 2001), on the other hand, the
more complex web with various interactive
links inclusive the microbial loop (Azam et al.,
1983).
The pelagic conditons
The discussion about the pelagic food
web in Lake Baikal has been regarded for long
time in way of the fi rst scenario, i.e., a step
wise trophic food chain, starting with primary
producers, mainly diatoms, continuing with
herbivorous and carnivorous zooplankton, and
ending up with fish, the Baikal seal as a top
predator and men. This traditional view has also
been accepted for marine systems, e.g., Steel
(1974). However, Steel concluded from his fish
production studies at marine shelves where he
thought about is a missing link in the food web,
which later has been recognized as the microbial
loop.
In the marine system, two further insights
were gained some time ago, which fit later well
into the picture of the microbial loop:
i) the two different kinds of primary
production, new and regenerated production
(Dugdale, Goering, 1967), and
ii) the influence of mechanical energy
transfers (Margalef, 1979).
Further, the microbial loop can only
work properly with an active community of
protozooplankton and, certainly, with a significant
contribution of dissolved organic matter which
is available to heterotrophic bacteria, generally
known as exudates. The question now is: are
these items necessary for scenario i or ii present
in all its facets in Lake Baikal by which the
ecosystem can be describe properly by the one or
other picture.
Consequently, the following issues have to
be tackled when describing the pelagic processes
and following the introduction question about the
Lake?s way of working:
- physical features of the euphotic zone;
- interactions between water and sediments,
i.e., benthic nutrient recycling and benthic
pelagic coupling;
- sources (nutrients) for the spring bloom
and its dominant phytoplankton;
- features and roles of bacteria during this
step in pelagic production;
- role of dissolved organic matter, its source
(exudates) and availability;
- seasonal successions of dominant
phytoplankton species and communities;
- succession of zooplankton communities;
- fate of particulate organic material,
sedimentation and regeneration process;
- which peculiarities are known for Lake
Baikal that might make it as a special
ecological system?
The structure of the euphotic zone and
nutrient regeneration is regulated by physical
processes, i.e., stratification, turbulence,
upwelling etc. Operationally, Lake Baikal can be
divided into three zones, the north, the central
and the south basin, just from its bathymetry
(Kozhov, 1962; Semovski, 2000). Each of them
show individual thaw behaviour in spring and
thus different temporal structures in water
masses (Killworth et al., 1996; Hohmann et
al., 1997) and subsequent shifts in biological
processes (Heim et al., 2005), which has been
documented by differences in organic material
from sedimentation traps by its distinct lipid
compositions (Russell, Russell-Mele, 2005)
and lead to the question of differences in
autochthonous productivity. The pelagic
stratification is then reflected by increases in cell
numbers of phytoplankton (Jewson et al., 2008).
Temperature has been proposed as a driving
factor for phytoplankton succession (Richardson
et al., 2000), but it seems more likely, as
temperature is triggered by radiation and this, in
turn, stimulates primary production.
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Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
Biological effects
The maintenance of diatoms in the photic
zone, which is important for their primary
production, can be increased by turbulence from
wind and subsequent mixing during spring. The
necessity of such turbulence for maintaining
diatom production has been described by Margalef
(1979). A simulation of disturbance of Lake Baikal
actually influenced positively phytoplankton
biomass and zooplankton fluctuations (Silow et
al., 2001). Turbulent patches are variable in size
and time. Wьest and Lorke (2010) found those on
scales of 100 m in Lake Baikal.
Some special and local features like thermal
bars are able to transport water masses and
nutrients to deeper layers (Shimaraev et al., 1993),
and locally they can be of great importance for
the water mass (and nutrient) budget (Melnik et
al., 2006). Such water of elevated salinity is also
responsible for bottom water formation (Kipfer
et al., 1996), and stimulates bottom currents and
lateral transport of dissolved and particulate
matter.
Further, the influences of river inputs have to
be taken into accounts due to their input of energy
and inorganic matter as well as allochthonous
sources, like particulate and dissolved organic
carbon, bacteria and microbiological processes
(Yoshioka et al., 2002; Maksimenko et al.,
2008). The input of allochthonous organic C is
considered to be nearly 300 kt C yr-1 (Granina,
1997). This holds also true for the main elements
of earth alkali metals (Callender, Granina, 1997),
and silicon shows highest accumulation rates
in sediments and transport rates from rivers.
A study of Hohmann et al. (1997) showed the
significant influence of Selenga plume for deepwater renewal.
Seasonal effects
The start of the spring bloom is regulated by
the increasing light system, and the ice thaw from
south to north. This bloom starts early in spring
with the growth of Aulacoseira baicalensis or A.
skvortzowii under the ice (Mackay et al., 2003).
This is well confirmed by other reports, e.g., Fietz
et al. (2005a), Shimaraeva et al. (2010) who report
that this period is strongly dominated by diatoms,
here Stephanodiscus meyerii and A. baicalensis,
and Chrysophyceae.
During summer (July to September) blooms
of small unicellular cyanobacteria were found,
dominated by Synechococcus sp. or Synechocystis
sp. (Belykh, Sorokovikova, 2003; Belykh et al.,
2006). The numbers of 106 cells ml-1 are close
to numbers of heterotrophic bacteria or even
exceeding them (Nagata et al., 1994; Straskrabova
et al., 2005; Ahn et al., 2006). Summer blooms
were also described by dominance of Chlorophyta
(Koliella and Monoraphidium) as well as
flagellates (Fietz et al., 2005; Izmestyeva, Silow,
2010). During this time of strong stratification
also Synechocystis limnetica was found to
dominate the pelagial of the south basin (Mackay
et al., 2003), an autumn bloom again can be
formed again by A. baicalensis or other diatoms
(Galazi et al., 1978; Ryves et al., 2003). Further,
Rhodomonas pusilla, a Cryptophyceae species,
and Chrysophyceae have been observed (Fietz et
al., 2005a; Izmestyeva, Silow, 2010).
The great variability on diatom communities
seems to be special feature of this lake, triggered by
various environmental parameters (Popovskaya,
2000). The recurrent pattern of a picoplankton
bloom, dominated by cyanobacteria during
summer can be triggered by a stable stratification
and N-depletion, which hampers phytoplankton
growth, but favours a self-sustaining N-fixing
community during this time. The vertical
migration of zooplankton, which could help
to increase the concentration of ammonia-N is
obviously not sufficient to force a phytoplankton
bloom. Later, after a break of this stratification by
turbulence, some ammonia-N probably returns to
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Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
the photic zone and the autumn bloom can take
place, again we find diatoms and green algae.
Role of the Benthos
A pelagic-benthic coupling can be assumed
for these river mouths, where nutrients can be
regenerated from microbes or resuspended and
thus given back to the pelagic environment
(Sapota et al., 2006). The deeper parts receive
particulate matter via sedimentation process.
However, much of the particulate organic material
can be regenerated and particulate inorganic
material, e.g., diatom shells, redissolved on the
way through the water column. A recycling of
Si in sediments, as well in the water column is
regarded as an important process in geochemical
cycling and has been proposed by Callender and
Granina (1997).
An indication for a strong decomposition in
the water column is the fact that only 28 % of
surface pigments reach the bottom in the south
basin (Fietz et al., 2005b), but nearly 90 % of these
pigments could be related to Bacillariophyceae
and Chrysophyceae. Ryves et al. (2003) and
Batterbee et al. (2005) even assume that only 1 %
of the diatoms accumulate in the sediments and
many species dissolve (e.g., Nitzschia acicularis)
in the water column. Hence, the organic matter is
set free for microbial attack and mineralization
and thus nutrients, e.g., nitrogen in the form of
ammonia, may have a chance to be recirculated to
the euphotic zone and to stimulate a regenerated
production.
The dominant spring diatoms (A.
baicalensis, A. skvortzowii), on the other hand,
show high sinking rates with 60-100 m d-1
(Ryves et al., 2003), which is in the same order
of marine species, which accelerate their sinking
rates by aggregate formation (Eppley et al., 1967;
Smetacek, 1985). In this case, however, organic
material (and recycled nutrients) are lost for the
euphotic zone. As A. baicalensis is capable to
produce large free-floating macro-aggregations
grown in this state, it serves as a preferred food
source for bottom invertebrates (Bondarenko et
al., 2006), and thus serves in these areas for a link
between sediment and water.
Sediments, however, do not show an
even seasonal signal. The taxa found can be
related well to environmental variables, i.e.,
ice conditions, radiation etc. (Mackay et al.,
2003). Only few data are available about benthic
remineralization processes and a process, which
might bring back remineralized nutrients to the
euphotic zone. Maerki et al. (2006) showed that
mineralization occurs in deep water sediments
mainly by oxidative processes. But the return of
inorganic matter to the euphotic layer is difficult.
Deep ventilation has been described only as an
episodic event (Killworth et al., 1996) or is due to
local thermal bars (Shimaraev et al., 1993). Hence,
significant amounts of nutrients from benthic
processes are probably restricted to coastal areas
such as river mouths.
Remineralisation
in the water column
Total C-production can be assumed to
be about 20 g C m-1 yr-1 in the south basin, and
14 g C m-1 yr-1 in the north basin, respectively
(Mьller et al., 2005, ILEC). These figures
underpin the oligotropic character of the lake
which is assumed to be due to a lack of available
phosphate (Mьller et al., 2005) and have been
confirmed by Straskrabova et al. (2005). Low
concentrations of phosphate (< 15-20 mg L-1
PO4-P) have also been described as sporolation
threshold for Aulacoseira skvotzowii (Jewson
et al., 2008). Such figures have to be related to
the occurring secondary production either by
direct feeding of herbivorous zooplankton or
via heterotrophic uptake of exudates by bacteria
which then are consumed by protozooplankton
which enters the food chain by carnivorous or
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Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
omnivorous zooplankton. This latter link, known
as the microbial loop, was under investigation by
Straskrabova et al. (2005) and presents the only
intensive view on the abundance view on these
critical items of the microbial loop. High amounts
of dissolved organic carbon have been reported
by Yoshioka et al. (2002), but not clearly related
to its sources.
Due to these patterns (Silow, 1999, Moore et
al., 2009) and, in more detail, Yoshii et al. (1999)
describe different but straight ? ?simple? ? food
webs from phytoplankton to top consumers for
the pelagic and the benthic system, the latter
underpins this finding from studies with stable
isotopes. Yoshii et al. (1999) just present five
steps, expressed by five ecological groups, i.e.,
phytoplankton (A. baicalensis), mesozooplankton
(Epichura
baicalensis),
macrozooplankton
(Macrohectopus branickii), fish (Coregonus
autumnalis), and the seal (Phoca sibirica).
There is much evidence that limnoecology of
Lake Baikal works basically at these levels, as
this sustains an empiric observation of the food
web. Nevertheless, the bypass by the microbial
loop cannot be ignored, as we see high amounts
of dissolved organic matter and ? at least for
some time ? the occurrence of protozooplankton.
Figures of production rates show a decrease from
low primary production (2100 mg m-2 yr-1) to
bacterial production (748 mg m-2 yr-1), herbivorous
zooplankton (178 mg m-2 yr-1), carnivorous
zooplankton (10 mg m-2 yr-1) and fish (4 mg m-2
y-1) (ILEC).
External impacts
The impact of global warming has also
reached Lake Baikal with effects on mean annual
temperature of air and water with consequences
on ice duration and biological features (Moore
et al., 2009). These authors also describe the
subsequent steps in physical and biological
oceanographic patterns, like the shift to earlier
dates of the spring and summer bloom onsets
of phytoplankton. An increase in radiation is
followed by an increase in water temperature
and thus water stratification and water mixing in
the lake?s basins. Longer periods have positive
effects on primary production, phytoplankton
growth and succession. Such increase in
average water temperature by 1.21 °C since
1946 is reported by Hampton et al. (2008)
with consequences on chlorophyll a contents
and different zooplankton grazers. A negative
influence can be expected on the growth of the
spring bloom diatom Aulacoseira skvotzowii
which is cold adapted and its mortality will
increase at temperature above 6.5 °C (Jewson
et al., 2008). Autotrophic picoplankton has
proposed as indicators for ecosystem health in
general (Munawar et al., 1994), hence, survey
studies should keep an eye on this group of
organisms.
Summary
This review showed a strong evidence for
a strong local diversity in Lake Baikal?s pelagic
and benthic biology. The location of the lake and
the different seasonal and local effects make it
difficult to analyse the available data with respect
to a comprehensive biological view. The concern
is especially true for the shifting phytoplankton
spring blooms from south to north as well as for
the widely uncoupled benthic-pelagic systems,
i.e., the pelagic system in the photic zone. The
decoupling can be regarded as a main factor
for the nutritional oligotrophic regime of Lake
Baikal. In order to analyse the pelagic food web
with respect to an active microbial loop and the
nutrient recovery from deep lake environments,
new approaches in research are needed. This
is especially necessary, when coping the new
environmental problems of global warming and
increasing anthropological inputs into the lake.
Studies on sedimentation, nutrient and pollution
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Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
recovery from the deep layers might become a
serious problem in near future when the mixing
zone might shift to deeper layers due to strong
impacts from atmospheric events.
Acknowledgements. The authors would
like to thank participants for fruitful discussions
during the Symposium ?Problems of Ecology?,
held September 20-25th, Irkutsk, 2010.
References
Ahn T.S., Kim O.S., Joh K.S., Spiglazow L.P., Drucker V.V., Hing S.-H. 2006. Community analysis
of aggregated bacteria in southern Lake Baikal. Hydrobiologia 568: 5-8
Azam F., Fenchel T., Field J.G., Gray J.S., Meyer-Reil L.-A., Thingstad F. (1983) The ecological
role of water-column microbes in the sea. Mar. Ecol. Progr. Ser. 10: 257-263
Batterbee R.W., Mackay A.W., Jewson D.H., Ryves D.B., Sturm M. (2005) Differential dissolution
of Lake Baikal diatoms: correction factors and implications for palaeoclimatic reconstruction. Glob.
Planet Change 46: 75-86
Belykh O.I., Sorokovikova E.G. (2003) Autotrophic picoplankton in Lake Baikal: Abundance,
dynamics, and distribution. Aquat. Ecosyst. Health Managem. 6: 251-261
Belykh O.I, Ekaterina G., Sorokovikova T., Saphonova A., Tikhonova I.V. (2006) Autotrophic
picoplankton of Lake Baikal: composition, abundance and structure. Hydrobiologia 568: 9-17
Bondarenko N.A., Timoshkin O.A., Rцpsdorf P., Melnik N.G. (2006) The under-ice and bottom
periods in the life cycle of Aulacoseira baicalensis (K. Meyer) Simonsen, a principal Lake Baikal alga.
Hydrobiologia 568: 107-109
Callender E., Granina L. (1997) Geochemical mass balance of major elements in Lake Baikal.
Limnol. Oceanogr. 43: 148-155
Dugdale R.C., Goering J.J. (1967) Uptake of new and regenerated forms of nitrogen in primary
production. Limnol Oceanogr 42:196-206
Epply R.W., Holmes R.W., Strickland J.D.H. (1967) Sinking rates of marine phytoplankton
measured with a fluorometer. J Exp. Mar. Biol. Ecol. 1: 191-208
Fietz S., Kobanova G., Izmest?eva L., Nicklisch A. (2005a) Regional, vertical and seasonal
distribution of phytoplankton and photosynthetic pigments in Lake Baikal. J. Plankt. Res. 27: 793810
Fietz S., Sturm M., Nicklisch A. (2005b) Flux of lipophilic photosynthetic pigments to the surface
sediments of Lake Baikal. Glob. Planet Change 46: 29-44
Granina L. (1997) The chemical budget of Lake Baikal: a review. Limnol. Oceanogr. 42: 373378
Hampton S.E., Izmest?eva L., Moore M.Y., Katz S.L., Dennis B., Silow E.A. (2008) Sixty years
of environmental change in he world?s largest freshwater lake ? Lake Baikal, Siberia. Glob. Change
Biol. 14: 1-12
Heim B., Oberhaensli H., Fietz S., Kaufmann H. (2005) Variation in Lake Baikal?s phytoplankton
distribution and fluvial input assessed by SeaWiFS satellite data. Glob. Change Biol. 46: 9-27
Hohmann R., Kipfer R., Peeters F., Piepke G., Imboden D.M., Shimaraev M.N. (1997) Processes
of deep-water renewal in Lake Baikal. Limnol. Oceanogr. 42: 841-855
ILEC ? International Lake Environment Committee Foundation, «World Lakes Database», www.
ilec.or.jp
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Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
Izmestyeva L., Silow E. (2010) Long-term dynamics of summer community of Baikal
phytoplankton and climate change. 13th World Lake Papers. Shiga: ILEC. 4 p. http://wldb.ilec.or.jp/
data/ilec/WLC13_Papers/others/9.pdf
Jewson D.H., Granin N.G., Zhadow A.A., Gorbunova L.A., Bondarenko N.A., Gnatovsky R.Yu.
(2008) Resting stages and ecology of the planktonic diatom Aucaloseira skvortzowii in Lake Baikal.
Limnol. Oceanogr. 53: 1125-1136
Killworth P.D., Carmack E.C., Weiss R.F., Matear R. (1996) Modelling deep-water renewal in
Lake Baikal. Limnol. Oceanogr. 41: 1521-1538
Kozhov M.M. (1963) Lake Baikal and its life. The Hague: W. Junk Publishers, 344 p.
Kozhova O.M., Izmest?yeva L.R., Eds. (1998) Lake Baikal: Biodiversity and Evolution. Leiden:
Backhuys Publishers, 447 p.
Kozhova O.M. (1987) Phytoplankton of Lake Baikal: Structural and Functional Characteristics
Arch. Hydrobiol. 25: 19-37
Mackay A.W., Battarbee R.W., Flower R.J., Granin N.G., Jewson D.H., Ryves D.B., Sturm M.
(2003) Assessing the potential for developing internal diatom-based functions for Lake Baikal. Limnol.
Oceanogr. 48: 1183-1192
Maerki M., Mьller B., Wehrli B. (2006) Microscale mineralization pathways in surface sediments:
A chemical sensor study in Lake Baikal. Limnol. Oceanogr. 51: 1342-1354
Maksimenko S.Yu., Zemskaya T.I., Pavlova O.N., Ivanov V.G., Buryukhaev S.P. (2008) Microbial
community of water column of the Selenga River-Lake Baikal biogeochemical barrier. Microbiol. 77:
587-594
Margalef R. (1979) The organization of space. Oikos 33: 152-159
Melnik N.G., Bondarenko N.A., Belykh O.I., Blinov V.V., Ivanov V.G., Korovyakova I.V.,
Kostornova T.Ya., Lazarev M.I., Logacheva N.F., Pomazkova G.I., Sherstyankin P.P., Sorokovikova
L.M., Tolstikova L.I., Tereza E.P. (2006) Distribution of pelagic invertebrates near a thermal bar in
Lake Baikal. Hydrobiologia 568: 69-76
Moore M.V., Hampton S.E., Izmest?eva L.R., Silow E.A., Peshkova E.V., Pavlov B.K. (2009)
Climate change and world?s ?sacred sea? ? Lake Baikal, Siberia. Bioscience 59: 1-13
Mьller B., Maerki M., Schmid M., Vologina E.G., Wehrli B., Wьest A., Strum M. (2005) Internal
carbon and nutrient cycling in Lake Baikal: sedimentation, upwelling, and early diagenesis. Glob.
Planet Change 46: 101-124
Munawar M., Munawar I.F., Weisse T., Leppard G.G., Legner M. (1994) The significance and
future potential of using microbes for assessing ecosystem health: The great lakes example. J. Aquat.
Ecosyst. Health 3: 295-310
Nagata T., Takai K., Kawanobe K., Kim D.-S., Nakazato R., Guselnikova N., Bondarenko N.,
Mologawaya O., Kostrnova T., Drucker V., Satoh Y., Watanabe Y. (1994) Autotrophic picoplankton in
southern Lake Baikal: abundance, growth and grazing mortality during summer. J. Plankton Res. 16:
945-959
Popovskaya G.I. (2000) Ecological monitoring of phytoplankton in Lake Baikal. Aquat. Ecosyst.
Health Managem. 3: 215-225
Richardson T.L., Gibson C.E., Heaney S.I. (2000) Temperature, growth and seasonal succession
of phytoplankton in Lake Baikal, Siberia. Freshwater Biol. 44: 431-444
# 118 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
Russell M., Russell-Mele A. (2005) Preliminary study of fluxes of major lipid biomarker classes in
the water column and sediments of Lake Baikal, Russia. Glob. Planet Change 46: 45-56
Ryves D.B., Jewson D.H., Sturm M., Battarbee R.W., Flower R.J., Mackay A.W., Granin N.G.
(2003) Quantitative and qualitative relationships between planktonic diatom communities and diatom
assemblages in sedimenting material and surface sediments in Lake Baikal, Siberia. Limnol. Oceanogr.
48: 1643-1661
Sapota T., Hakanson L., Aldahan A., Possnert G. (2006) Sediment flux to Lake Baikal (Siberia,
Russia): Modelling approach. Geomorphology 80: 105-113
Semovski S.V., Mogilev N.Yu., Shertsyankin P.P. (2000) Lake Baikal ice: analysis of AVHRR
imaginary and simulation of under-ice phytoplankton bloom. J. Mar. Syst. 27: 117-130
Shimaraev M.N., Granin N.G., Zhdanov A.A. (1993) Deep ventilation of Lake Baikal waters due
to spring thermal bars. Limnol. Oceanogr. 38: 1068-1072
Shimaraeva S., Izmestyeva R., Silow E. (2010) Long-term dynamics of under-ice community
of Baikal phytoplankton and climate change. 13th World Lake Conference Papers. Shiga: ILEC. 4 p.
http://wldb.ilec.or.jp/data/ilec/WLC13_Papers/others/13.pdf.
Silow E.A. (1999) The use of two lumped models for the analysis of consequences of external
influences on the Lake Baikal ecosystem. Ecol. Model. 121: 103-113
Silow E.A., Baturin V.A., Stom D.J. (2001) Prediction of Lake Baikal ecosystem behaviour using
an ecosystem disturbance model. Lake Reserv. Res. Managem. 6: 33-36
Smetacek V. (1985) Role of sinking in diatom life-history cycles: ecological, evolutionary and
geological significance. Mar. Biol. 84: 239-251
Steele J.H. (1974) The structure of marine ecosystems. Cambridge: Harvard University Press.
Straskrabova V., Izmest?eva L.R., Maksimova E.A., Fietz S., Nedoma J., Borovec J., Kobanova
G.I., Shchetinina E.V., Pisllegina E.V. (2005) Primary production and microbial activity in the euphotic
zone of Lake Baikal (Southern Basin) during late winter. Glob. Planet Change 46: 57-73
Wetzel R.G. (2001) Limnology: Lake and River Ecosystems. Third Edition. N.Y.: Academic
Press.
Wьest A., Lorke A. (2010) Small-scale turbulence and mixing: energy fluxes in stratified lakes.
In: Likens G.E., Ed. Lake ecosystems ecology. A global perspective. N.Y.: Academic Press, 107114.
Yoshii K., Melnik N.G., Timoshkin O.A., Bondarenko N.A., Anoshko P.N., Yoshioka T., Wada
E. (1999) Stable isotope analyses of the pelagic food web in Lake Baikal. Limnol. Oceanogr. 44:
502-511
Yoshioka T., Ueda S., Khodzher T., Bashenkhaeva N., Korovyakova I., Sorokovikova L.,
Gorbanova L. (2002) Distribution of dissolved organic carbon in Lake Baikal and its watershed.
Limnology 3: 159-168
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Manfred Bцlter and Lyubov R. Izmest?yeva. Is there a Microbial Loop in Lake Baikal?
?????????? ?? ????????? ?????
? ????? ???????
?. ???????, ?.?. ???????????
?
???????? ???????? ????????,
??????????? ?????????-?????????,
???????? 24148, ????, ?????? ???. 1-3
?
??? ????????,
????????? ??????????????? ???????????,
?????? 664003, ???????-3, ?/? 24
??????????? ????????????? ????????? ????? ? ????????? ? ????? ??? ? ???????????????? ?
?????????????????? ?????????, ??? ? ? ?????????? ????????? ? ?????? ?????? ? ??. ??????.
???????, ? ??????? ?????????? ?????? ???? ? ??? ????????? ???????? ?????????, ?????????
?? ??????? ??????? ????? ????? ?????????????? ? ??????????. ??? ?? ????? ??????????
?????? ?? ????????????????? ? ???????????? ????????? ? ???? ?????? ?????, ??????????? ?
????????????? ?????, ?????????? ????????? ????? ??????? ??????? ???????? ? ?????. ?????
????, ???????? ?????????? ?????????? ?????????? ?? ???? ??????????? ???????????????
??????????? ???????? ? ????? ? ?????????? ????. ????? ???????, ????????????????
?????????? ????????? ???????? ?????? ????? ????????? ? ???????? ?????, ??? ? ?????
????????? ? ?????????????? ??????? ?????.
???????? ?????: ??.??????, ??????????, ????????????? ?????.
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Journal of Siberian Federal University. Biology 2 (2012 5) 121-137
~~~
??? 574.58(262.54)
?????? ? ?????????-?????? ??????????????
????????? ? ????????? ?????????
????????? ????
?.?. ??????*
???????? ??????? ??? ?????? ???????? ?????? ???
???????? ?????? ??????????? ???????? ?????????????? ?????????
?????? 344006, ??????-??-????, ??. ??????, 41 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
? ?????? ?????????? ???????? ?????? ? ?????????-?????? ?????????????? ?????????
? ????????? ????????? ????????? ????, ?? ????????????? ?????? ?? ?????? ?????, ?
????? ???????? ? ??????????? ????????, ?? ???????, ????????????? ? ??????????????
???????????????. ?????? ??????? ?????? ??????????? ????????????? ????????? ??
???????? ?????, ???????? ?? ??????? ???????????? ? ????????.
???????? ?????: ???????? ????, ?????????-?????? ??????????????, ????????????
??????????, ?????????, ???????????? ??????????, ????????????.
????????
?????? ??????? ?????????????? ? ???????? ????? ? ??????? ???????????? ????????? ????????? ? ?????????? ???? ?????????
????. ?? ???????????? ? ??????????????
????????? ???????????? ?? ?????? ???????
??????????? ?????????????? ?????????? ?????? ????????? ????, ?? ? ???????????????
??????? ??? ????? ? ??????? ?????????????? ?????????? ???.
?????? ???????? ? ?????? ?????????????? ????????? ???? ?????????? ? ??????
?. ?????????? (1896). ? 1904 ?. ? ??????? ?????????? ??????????? ? ???????? ???? ??-
*
1
?????? ??????-??????????? ?????????? ???
???????????? ?.?. ?????????. ????? ?? ??
?????? ??? ???? ?.?. ??????? «????????? ?
????? ????????? ????» (1927), ? ??????? ???????? ?????? ?? 33 ????? ??????? ??????????,
16 ???????, 6 ????? ??????????, 4 ??????
?????? ????????, ?????????? ?????????????
???????????. ?????????? ????????? ???????? ? ??????????-?????????? ?????????
???? (?????, 1925, ??????, 1960, ????????????????, 1955).
? 1951 ?. ???????? ????????????????
?????????? ?????? ? ?????????? ? ????
???????? ? ???????????????? ????? ?????-
Corresponding author E-mail address: ulva@ssc-ras.ru
© Siberian Federal University. All rights reserved
# 121 #
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?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
???? ?.?. ?????????? (1951). ??? ?????????
??????? ?????? ?????? ???????? ?.?. ??????? 22 ?????? ??????, ? ????? ??????? ???
?????????????? ????????????? ?????????
????????? ????????? ???????? ? ??????????.
????????? ????????? ???? ? ?? ????????
????????? ? ??????????? ???????????????
??????? (??????????) ?.?. ???????? (1960).
? 1965 ?. ????? ???????? ?????? ?? ??????
?????????????? ????? ???? ?.?. ????????
«???????????? ????????? ????? ????», ? ??????? ??????? ???????? ????????? ??????
?????????????? ????????? ????.
??????? ?????????????? ???????????
?? ????????? ?????????? ??????? ????????
???????, ????????? ? ????????? ????, ???????? ?.?. ????? (1977). ? 80-? ??. ?? ?. ??????????? ???????? ?????? ?????? ????????? ????? ????????????? ??????? ???? ???????
??????????? ?????????? (?????-???????
? ??????, 1984; ?????? ? ?????-???????,
1989), ??????? ?????????? ??????????????
??????-???????? ?????? ? ???????????????
????????? ????.
? 1981 ?. ??????????????? ???????? ??????????? ????????? ???? ? ??????????? ?????????? ???????? ?.?. ?????? ????????? ?
???????????? ???????? (?.?. ?????????,
?.?. ????????????, ?.?. ????????). ??????
???????? ????????? ???????? ????????? ????? ????????, ???????, ????????? ? ?????????????? ???????????? ????????? ?????????
???????? ????????, ??????? ????????? ????????????????? ? ????????????? ????????,
?????? ????? ??????????? ?? ?????? ???????
?????????????? ? ????, ?? ????????? (??????,
1983, ??????, ????????, 1989, 1990, ?????? ?
??., 1991). ??? ????????? ????????? ????? ????????? ????????? ???????????????????? ???????? ???????? ????? ????????? ????.
????????????? ?????? ??????????????
?? ?????? ??????? ?? ????????? ??????, ???-
????????? ???????? ??????? ? ???????????
???????????? ???????? ? ?????? ???????????? ?????????. ????????? ?????? ?????????
???? ????????? ??????????? ?? ?????????
1,2-1,5 ?/???, ?????? ?? 4 ?/??? (????????,
1961). ?????? ?????? ????????? ? ???????????, ??? ??????????? ?? ???????????? ?
????????????? ?????????? ????????? ?????????? ? ????, ????? ??????? ??????????
???????????? ??????????? ? ????? ?????????? (????????) ????? ????, ? ???????? ????
???????????? ? 200-300 ? ? ???????? ?? ?????
????.
?? ???????????????? ???????? ?????????? ???????? ????????????? ????? ? ?????????. ????????????? ????? ???????? ??
??????? ? ? ????????, ? ????? ? ??????? ??
??????????? ????????? ?????????, ??????????? ??????? ????? ?????????? ??????,
??? ?????? ??????????? ?? ???????? ??? ? ??
????????????. ????????? ??? ????? ??????????????? ??????? ?????????? ????????? ??
????????? ???????? ????. ???, ? ???????????? ?????? ??? ???????????? ??????? ????????? ?????? ??????? ???? ??????????? ??
?????? ?? ??????, ? ??? ???????? ????????
?????? ? ???????? ?????? ??? ?????????? ????????? ????????? ?? 9 ? (???????, 1957).
?????? ??????, ???????? ?? ????????
?????????-?????? ?????????????? ? ?????, ?
???????? ????? ???????????? ????????? ?
???? ??????. ?????? ? ????????? ?????????? ? ????? ????????? ???? ?? ????????? 41
??? ? ? ??? (????????, ?????????, 1980). ???
????????? ??? ???????? ?????? ??? ? ????????? ??????, ??????????????? ??? ???????????? ?????? ???????????? ?????????.
????? ????, ??????? ???????? ?????????????, ??????????? ?? ???? ????????? ????
???, ????????? ?????????? ??? ??? ? ????????? ?? ?? ?????????? ?????, ????? ??????
???????????? ???.
# 122 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
?? ????????? ???????? ???? ???????????? ????? ????? ????????. ??????????
???????????? (7,0-10,0 ?) ???????? ??????
??????????? ??????? ? ???????? ?????? ???
????? ???? ? ??? ??????????? ????????????
???. ???? ?????????????? ????????, ? ???????? 3 ??????, ???????? ? ??????????? ????????? ??????? ?????? ? ??????? ?????????? ????????????, ?????? ?? 0,2-0,3 ?.
??????? ??? ???????????? ? ???????? ?????
???? ?? ????? ?????: ????? 1,8 ?; ??????
2,7 ?; ????? 2,3 ?; ?????? 2,0 ? (????????,
1973); ???? ????????? ?????? ? ???????????????.
??????? ???????? ? ?????? ???????????? ??? ?????????? ????????????? ????????? ?????? ? ?????? ??????????. ????? ???????, ??????? ????? ????????? ????
??????????? ??????????? ?????????? ??
??????? ??????????????? ??????? (??????
????) ? ?????? ????????????, ???????????
?????????? ??????? ????????. ??????? ??
?????? ??????? ?????? ?????????????? ??????????? ???? ????? ??????? ????? ??????,
?? ??????? 1,5-2 ?.
????? ?????, ????????? ? ??????
?? ?????? ??????????????????? ????????, ???????????? ???????????????? ?
???????????????? ?????? ???????? ?????
????????? ???? ??????????? ?? ??? ?????: ???????????? ?????, ?????????? ???????? ?????
? ?????????? ?? ???? ?????????? ????????
????? ????? ? ??????? ??????????. ??? ?????? ?? ???? ?????? ?????????? ???? ??????????????, ??????? ???? ????????? ?????????????
?? ?? ???????????.
????????????????? ?????? ?????????????? ? ????????? ????????? ???? ? 1981 ?.
????????????? ?? ??????????? ????? ???????? ? ??????????? ?? ??? «?????» ?????????????? ??????? ??????????. ? 1982 ?., ????
? ???????????, ?????? ??????????? ?? ?????
???? ?? ????? ???????? «???????????». ?
1991 ?. ?? ????? «??????» ?????? ?????????????? ????? ????????? ?????? ???? ? ? ?????????? ???????. ? 1982 ?. ? ? 1995 ??. ?????????
??? ????????? ??????? ?????? ????? ????????? ???? ?? ???????????, ? ??????? ????
?? ????? ??????????? ???? ?? ?????????????? ????? ????????, ? ??????????? ???????????? ??????, ?? ??????? 3 ? ? ?????????
? 1. ? 1982 ?. ????????????????? ?????? ?
??????? ???? ????????????? ????? ??????
2-3 ????. ???????? ?????? ???? ????????? ?
???????? ??????????? ??????? ? ????????????? ????? 2009 ?. ?? ????? ????? ???????? ?? ????? ????????, ? ? ?????? ?????? ???
??????? ????????? ????????, ?.?. ??? ???????? ???????? ?? ????? ???? ? ??????? ????
?? ???????? 0,5, 1,0 ? 3,0 ?. ??????????? ?????????? ? ???? ????????? ?? ?????????????
?.?. ??????? (1967) ? ?.?. ???????? ? ?.?.
???????? (????? ??????? ????, 1984-1985) ?
?????? ??????????.
?????????, ???????????, ?? ? ????????
???? ?????????? ??? ?????? ????????????
?????-??????????? (Horiba, ??????), ?????????????? ???????????????? ?? ??????? ??????????. ? ??????? 2003 ?. ??? ??????????
????? ???????????? ?????????? «?-16» ?
??????? ????? ????????????? ?????? ?? ??????? ???????? ??? ? ?? ???????, ??? 30 %-?
?????????? ? ???????? ???? 2 ?????; ??????
????????? ? ???????.
??? ??????? ??????? ???????????? ????????? ??????????? ???? ? ??????? ?????
?.?. ???????? (1964).
?????????? ? ??????????
???????????? ?????
???? ?? ????? ??????? ?? ????????
?????? ????????? ????????, ????????????
?????, ?????????? ?? ??????? ????????????
# 123 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
???????. ????????????? ???????????? ? ???????????? ?????? ????????? ???? ? ??????? ????? ? ??
???????
????? ?????
???????, ?
????????????, ??
6,00
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370
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580
7,20
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4000
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95 000-100 000
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21,05
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21,05
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73 ? 80
??????????: ??????? ???????????? ? 300 ?? ???????? ???????????? ??? ?????? ??????????.
??????????? ????, ? ?? ?????? ?????????????? ???????? ??????, ??????????? ???? ????????????? ?? ???????? ?????? ? ???? ?????? ? ?? ?????. ? ??? ????????? ??????????
???????? ??????? ??? ???? ? ??????? ???
????????? ????.
???? ?????? ?????????? ??????????
?????????? ?? 1,9 ?? 7,7 ?, ? ??????? ????????? ? 285-961 ??/?, ? ????? ?????? ??????????? ????????? ? 2,3-4,9 ?2 ??/?. ???????????? ?????????? ?? 0,2 ?? 0,6 ?; ????????
??????? ????? ?????????? ? ?????? ???????
(?? = 4,8-5,7). ????? ???????? ??????? ????? ? ??????, ????????? ??????? ?????????
6 ?. ??? ??????? ????????? ? ????????????
?????? ?????????? ?? 9 ?, ? ??? ?????? ??????? ???? ??????????? ?? ?????? ?? ??????
(???????, 1962). ????? ????, ? ?????? ??????
????????????? ???????????.
? ????????????? ???? ??????????? ???????????? ?? ?????????? ??????? ?????????? ????? ???????? (Carex gracilis Curt.) ?
?????? ????? ????? ????, ? ????? ??????????
????-??????? ??????????, ????????? ??????
??????? ? ???????????? ??????? ????????-
????? ??????, ??????????????? ?????????????? ????????, ??????????? ? ????????
????????? ?????????-?????? ?????????????? (???. 1).
??????? ? ?????????????? ????????????,
??? ?????????? ??????? ????? ???????????
????? ??????????? ????????? ? ??????????? ??????????? ??????? ? ???????, ?? ??????? 2-3 ? ?????-???? ?????????????? ???????????. ??? ?????????? ? ????? ?????? ????
?? ?????, ? ???????? ???????, ?????????????? ???? ?????? ?????? ????? ? ????????????????? ?????????? Monoda?na.
?? ?????? ??????, ?? ???????? ? ??????
?????????????, ?????? ?????????????? ???????????? ????????? Phragmiteta australis ???
??????? ????????????? ?????????, ??? ? ?????????? ???????????? ????????? ? ??????
???????????? (Scirpus triqueter L.), ? ???????
????? ??????????? ???????? ????????? 85100 %. ??? ????- ? ???????????? ??????????
?????????? ? ????????-?????????? ????????, ???????????? ? ?????????? ???? ? ????? ?? ??????? ??????, ??? ??? ????????????
? ???????? ? ???-????????? ??????????? ??
# 124 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
???. 1. ????????????? ?????? ????????? ? ????????? ?????? ????????? ????
?????? ?????????. ????? ?????????? ?????
? ??????????? ?? ????????? 9-10, ?????????
?????????? ????????? 5979 ?/?2. ? ????????
???????????? ??????????? Scirpus lacustris
L., S. triqueter, Typha latifolia L., T. angustifolia L., Catabrosa aquatica (L.) Beauv. ? ?????
?????????????? ???????????????? ???????
?????????? ? Cladophora fract? Kьtz., C. glomerata Kьtz., Enteromorpha compressa (L.) Grev.,
E. intestinalis (L.) Link, ? ????? ??????????? ?
Vallisneria spiralis L. ? ??????? ??????????
Salvinia natans (L.) All. ??????? ?????? ????????? ? ???? ???????? ??????????? ??????????? ??????? ??????????? ?????????? Chlorococcum caldariorum (Magn.) Brunnth. ?????
?????? ???????????? ???????? ??????????
????????? ?????????? ?????????? Scirpetum purum ? ?????????????? ?????? ???????????? (S. triqueter) ???????????? ?????????????. ?? ?????? ?? ????????? 145 ??,
??????? ????????? 1668 ?/?2. ?? ???????????
????????, ???, ??? ? ????? ??????? ?????????
?????, ?????? ???????????? ??????????? ??-
???????? C. fracta ? C. glomerata, ??????? ???????????? ?? ??????? ???????? (???????
? ??., 1987). ??? ?? ???? ?????? ? ????????????? ???????? ??????????? E. compressa ?
E. intestinalis ?????? ?????????? ? ?? ?????????? ??????????????, ???????????? ? ?????????????? ????????. ? ????????? ??????
??? ???????? ???????????????, ?????????????, ??????????? ????????????? ??????????? ????? ????? ??????.
????? ????????? ?????? ?????????????
?????? ?? ?????????? ???????? ????????
?????? ?? ???????????? ???????? ??????????? ? ????????? ????????????? ?????????? ?????????, ?????? ? ?????? Scirpetum
triqueteri purum ? Typhetum subpurum (? ????????? ?????????? ????? ???????????
T. angustipholia), ?????? ???????????? ??????? ??????????? ??? ? ????, ??? ? ?? ????.
?????? ??? ??????????? ????????? ??????,
????????? ??????????????. ???????? ??
???? ???????? ?? ??????????????, ??? ????
??????? 1,3-1,5 ? ??? ?????????? ? ??????
# 125 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
?? ??????????. ?????? ?????, ???????? ????
?????? ??? ????, ??? ????????? ?????? ? ????????????????? ??????? ??????? ???????,
? ???????.
?? ?????????? ???? ?? ????????? ?????? ?????? ? ???????? ??????????? ?? ??????? ??????? ????? ?? ???????? 0,3-0,4 ?
??????????? ????? ????????? ?? ????????????? ????????? ?????????? Scirpetum
triquteri, ?????? ??????? ?? ?????????
30-50 ?. ??-????????, ?????? ????? ??????? ?????? ???????? ?? ?? ????????????
???????? ??????, ??? ??? ?????? ? ??????
??? ?????????????. ?????? ? ???????????? ????????? ? ????????? ?????? ?? ??????? 1,5-1,7 ? ?? ???????? ? ??????-????????
?????? ??????????? ?????????? ???????????: ? ???? ?????????????? ????? ??? ????????? ????????? ???????????? Potamogeton
perfoliatus L. ? Potamogeton pectinatus L., ??????? ???????? ?????? ????- ? ???????????? ?????????? ? ???? ??????????? ????? ?
????????? ????? ? ??????????? ?????????
10-20 %. ?????? ??????? ?????, ????????????? ??????? ????????????????, 80-100 ??,
?????? ?????? ????????????, ??????????? ?
?????? ? ??????????? ?????? ????, 30-50 ??.
????????? ????????? ?????????? ? ??????????? ?? ???????????? ???????? ??????????
?? 1100 ?? 4900 ?/? 2 (max 8800 ?/? 2). ????????? ??????????, ? ???? ??????? ????????
????, ????????? ?? ????? ?????? ?????????????. ????????? ??? ??? ???????? ????????
20 ??/? (?????????, 2002).
???? ??????? 1,5-1,7 ? ?????? ?????????? ?????????? ? ???????????? ?????? ??
??????????, ??? ??? ?? ??????????????? ???????????? ?????? ????????????? ????. ?????, ??? ???????? ? ???????? ??????????? ?
??????? ?. ?????????, ??????????? ????????? ?????????????? ???????????? ??????
??????????????.
?? ???????, ????????????? ????? ??????? ????? ????????? ???? ? ????????? ????? ?????? ????, ????????? ???? ????????
??????? ? ??????? ???, ??? ?????????? ?????? ?????????? ?????????? ???????, ? ?? ???
???????????? ???????????? ????? ???????
???, ?????????????? ???????? ????????????????. ?? ?????? ?.?. ?????????? (1984),
? ????????? ????? ??? ??? ????????? 2 ?
?????????? ????????????? ????????? ??????, ??????????? ? ???????????? ????????,
c ???????? ??????????? ????????? ???????.
? ??? ????????? 5 ? ?? ????????? ?? ??????
????????? ???????????? ???????????? ????? ??????, ????????, ???? ? ?????? ??????
????????, ??????? ?????? ??????????????
??????????? ?? ???????? ?????? ????????.
? ???? ?????? ?? ??? ??????? ??????? ??????? ??? ????????? 60 ??. ??? ???????? ???????????? (?? ?.?. ?????????, 1986) ????
??????????, ????????????????? ?????? ?????????? 2-5 ?, ?????? ??????????? ????????? (2,3 ??/?) ??? ?? = 6,5. ?? ???? ??????? ?
???????? ??????????? ???? ??????? ?? ????????, ?????? ????????????? ?????????? ????? Cordylophora caspia Pall.
????????? ??????????? ?? ?????? ??????? ??????????????? ???????? ? ???? ?????? ???????? ??, ??? ? ???? ?????????????, ?.?.
? ???? ????????, ?? ?????? ???????? ? ????
?????????? ??????????? ????????? ???????? ??????? ???????? ???????????? ????????? Bangia atropurpurea (Roth) Ag. ? ? ????
?????? ??????????? ????????? Oscillatoria
bonnemaisonii (Crouan) Gom. ? Synechocystis
salina Wislouch.
???????? ?????
? ?????????? ???????????? ????? ????????? ?????? ? ???????? ??????????? ??
???? ??????, ??? ?? ????????? ???? ????????
?????????? ??????????? ????????? ???????-
# 126 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
????? ???????????????? ? ????????????????
??????? ?????? ?????, ????????? ??????????????? ?? ?????? 5-7 ?, ???????????? ????
????????????? ?? 1,7-4,0 ?, ??????? ??????
????????? ? ??????-????????, ???????? ?
?????????-????????. ? ???????????? ? ???????????? ?????????? ????????????? ??????? ??????????? ? ?????? ??????????.
? ????? ?????????? ????? ?????????????? ????? ??????? (???. 1) ??????????? ???????????? ???????? ???????????? Bolboschoenus maritimus (L.) Palla ? ???? Cyperus
fuscus L. ? ???????? ??????? ?????? ????
?? ??????-???????? ?????? ? ??????? 0,3 ?
?????????? ??????????? ?????? ?????????? ?????? ????????? Potametum filiformis
? ??????????? ????????? 15-25 %, ??????????????? ???????? ? ?????????? 800820 ?/? 2. ?????????? ?????? ????????? ?? ?????????????? ???? 1,2 ?. ??????? ????????,
??? ????? ??????????? ????? ???????????
???, ? ??? ????????? ????????? ???? 7 ?
?? ??????????? ???????????? ????? ???????
?????, ? ??? ?? ? ?.?. ????????? ?????????
??? ??? ???????????. ??????, ?.?. ??????????, ? ??????? ?? ?????? ?????????, ???????
??? ??????? ??????? Potamogeton interruptus
Kit.1.
? ???????????? ? ????????????? ???? ?
?????? ??????????? ????????????? ???? ????????? ? ??????????? ??????, ? ??????? ? ??
????????? ?????? ?? ????????? ?????? ? ??
??????? 3,0 ? ??????????? ??????? ???????
??????????? ?????????? ??????? ???? ???????? ?????? Zostereta.
??????????????? ? ????????? ?????????? ????????? ?????? ? ?????? ????????????
????????? ??? ?? 90-100 % ? ???????????? ??
?????? ? ??????? ???? ?? 1,5-2,0 ??. ?? ???????? ????????-?????????, ????-????????????,
? ?????????? ?? 5275 ?/?2. ? ??????? 3,0 ?
??????????? ???????? ??????????? ?????-
?????? ? 2-3 ????, ? ????????? ???????????
?? 1838 ?/?2. ?? ???????? ???????????? ? ??????????? ??????????? ? ???????? ???????
????????? Cladophora albida (Huds.) Kьtz.,
C. vadorum (Aresch.) Kьtz., E. compressa, E.
intestinalis, Chaetomorpha linum (O. F. Mьll.)
Kьtz., ? ??????? ? ????????? ???? ??? ????????? 5-7 ? ? ??????????? ?????????? ????????? ??????? ???????? ????????? Ceramium
strictum Crev. et Harv.
?? ??????? ????? ????????????? ? ?????????? ?????? ????????? ?????? ???? ?????????? ? ????????? 7-8 ?, ???????? ?? ??????????????? ?? ?????? 8,2-8,3; ? ??????????
????????? ? ???? ????????? 7,4-8,5 ??/?, ?.?.
??????????? ??????? ??????? ???????. ??
???????????? ????????? ?????????????? ??
???????, ????????????? ? ??????? ???????
?????????? ????, ???????????? ????????
??????? ????????? ??????, ????????? ???????? ????????? ?????. ? ?????? ?????? ?
?? ??????? 3,5 ? ?? ???? 23-????????????
?????????? ???? ??????????? ???? ???????????? ????????? ?????? ??????????:
Cladophora vagabunda (L.) Hoek, C. sericea
(Hьds.) Kьtz., Chaetomorpha linum, Enteromorpha ahlneriana Blid., E. compressa, E. intestinalis, Ceramium ?legans Ducl., C. strictum, Polysiphonia opaca (Ag.) Zanard., ???????????????
? ????????? ????????? ? ? ?????? ??????.
? ??????? 1,5 ? ?? ???????-??????? ??????
??????????? ??????????? ?????????? ??????? ????? Zostera noltii Horn. ?? ???????
3,5 ? ??? ???????? ??????????? ?????????? ?
???? ????? ????? ? ???????????? ??????. ?
?????????? ??????? ?????????? ???? ? ????????? ? ??????? ????????, ?????????? ??
?????????????? ?????? ????? ?????, ??????????????? ??????? ????? ???????????? ???
???????? ??????????? ? ????????? ??????
?????????????? ??????? ?????, ??? ? ????????? ??????? ????.
# 127 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
? ???? ?????????????? ?????????? ??????? ?????????, ????????? ?? ?????. ???
????? ??????? ???????????? ????? ????????? ?????? ?? ????????? ????, ?????? ?? ??
?????????? ?????. ?????? ???????????? ?????????? 2,5-3,0 ?, ????????? 3275 ?/?2. ???
?????? ???? ???????? ???????????? ?? 5060 ??, ??? ??????? ?? ????????? ????? ???
??????? ? ??????????? ?????.
????, ? ???? ??????????? ????, ???????
? ?? ??????? ????? ?? ??????-???????? ?????? ??????????? ??????????? ??????????????? ? ????????? ?????????? ????????????
????: Potametum filiformi ? Ruppietum purum
(? ?????????????? Ruppia maritima L.) ? ?????? ????? ??????????? ?????????? ???????
??????????. ?? ??????????? ???????? ?????????? ?? 30 ?? 100 %, ?????? ???????? ?????
120 ??, ?, ??? ???????, ??? ???????? ?????????? ????? ??????? ????? ??????? ????????
?????????? Chaetomorpha linum. ?????????
?????? 1527 ?/?2, ?????? ? 724 ?/?2, ????????? ? 544 ?/?2.
? ??????? ????? ?????????? ???? ?????????? ? ?????????????? ?????? ???????????, ?????? ?????????? ? ?????????
??????, ????????????? 17 ?????, ??????????? ???????????? ???????? ? 60-100 %
??????????? ????????? ? ??????????
450-1527 ?/? 2. ??????? ???? ????????
Potamogeton filiformis Pers., ??? ?????? 120
??, ??????? ???? ??????? 40-50 ?? ????????? R. cirrhosa, ? ???????? ??????? ? ????
????????? ?????? ??????????? Z. noltii, ?
??????? ? ????????? ????????? ?????? ?????????? ????????? ?? ??????? ??????????
?????????? ??????? Zostera marina L., ?????????????? ??????? ??????. ???????????
??????? ? ????????? ???? ? ?????????????
?????? ??? ??? ????????, ??????????????
??????? ??????, ??? ? ??? ?????? ??????
???????? (????????, 1975). ??? ?????????
???????????? ???? ??? ????? ?????????????
???????, ????????, ??? ?????????, ???????
?????????????????? ? ???????? ????, ? ???
?????? (???????) ? ??????? ????????????
??????? ?? ?????? ? ???????? ?????. ?????? ???? ?????????? ?????????? ??????????
?????? ??????????? ?????????: Polisiphonia
opaca, Ceramium strictum, Cladophora
albida, C. vadorum, Chaetomorpha linum,
E. ahlneriana, ?. compressa, ?. intestinalis,
Ectocarpus confervoides (Roth) Le Jolis. ? ??????????? ??????? ??????????? ??????????? ???????, ? ????? ? ??? ?????????????
????????? ? ?????????? Zosteretum marinae.
?????? ???????? ????? 0,6 ?; ?????????
2795 ?/? 2. ?????? ???? 3,5 ? ??????? ??? ?????????? ?? ?????????? ?????????? ??????
????????????. ??????? ??????? ????????????? ?????? ?????????????? ??????????? ? ?
????????? ???????? ??????????? ??????. ?
???? ?? ???????????? ???????? ??????, ?
??? ?????????? ???????, ?????????? ??????????????? ?????????? Ruppietum purum
(R. maritima) ? ??????????? ????????? 4080 % ??? ????????? 281-433 ?/? 2 ? ?????????
105 ???/? 2. ???????? ?????????? ?????????????????.
?? ????????? ??????? ???????? ?????
?????????? ? ????????? ?????? ? ????????
???????? ????? ?? ????? ???? ???????????
?????????? ??????? ?????????? Charetum
canescentis ?? 100 %-? ??????????? ?????????, ????????-????????? ?????????, ?????????? 885 ?/?2 ? ?????????? ???? 2272 ???/?2.
??? ?????? ???? ?????????????? ?????????
????? ????????????. ? ??????? 0,6 ? ????????????? ???????? ???????? ????, ? ? ????
?? ??????? ????? ???? ?????? ??????????
Zosteretum marinae purum ? ??????????? ????????? 100 %. ?????? ????? ? ???? ??????
????????????????? 10,0-11,2 ? ??????????,
???????? ?? ?????????? 8,3, ?????????? ???-
# 128 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
?????? ? 7,4-8,2 ??/?. ?????? ????????? ????? 60-70 ??, ????????? 3752 ?/?2. ?? ???????
3,5 ? ?? ???????-??????? ?????? ??????????
Zosteretum marinae purum ???????? ?????
??????????? ?????????? Zosteretum noltii
purum, ??????? ?? ??????? ???????? ??????, ?????????? ???????????, ???????????? ??
??????? 4,5-5,0 ?. ???? ??????? 5,0 ? ??-??
?????? ???????????? ???? ?? ???? ???????
?????????????? ?? ??????????.
?????????????? ????????????? ??????? ?
????? ??????? ??????, ????? ???????? ?????????? ?????, ???????????? ? ????????????
????????????? ?????????????? ?? ?????????
????????. ???, ????????, ? ?????????? ?????? ?. ????????? ? ???? ??????? ??????????? ?? ?????? ??????? ???????? ????? ?????
?????? ??????-???????? ????? ???????????? ?????????? ???????? ??????? ? ???????
?????????? ????? Ceramium, Cladophora,
Polysiphonia, Enteromorpha ? ??????.
? ???? ?? ?????? ? ??????? 0,9-1,3 ?, ??
??? ?? ???????-??????? ?????? ??????????
??????? ??????, ????? ? ??????? 3,5 ? ????????? ??????????? ?????????? ??????????
Zosteretum noltii ? ???? ???????????? ?????.
??? ??????????? ? ???????????? ???????? ??
??????, ??? ???? ??????? 5,0 ? ??? ??? ??????? ?????? ?????????????? ?????????? ??????????? ????????? Lyngbya aestuarii Liebm.
ex Gom., Aphanothece castagn?i (Brйb.) Rabenh., A. salina Elenk. et Danilov ? ?????????
????? Oscillatoria. ??? ???? ????? ??????????? ?????????? ?? ???????? ????????????
???????? ??? ???????? ? ?????? ?????????
??????????????. ???????? ????? ???????????? ???????, ?????? ?????, ?????? ?????????
? ????????????????????? ?????? ?. ?????????, ???????????? ??????? ???? ? ????.
?????????? ??????? ???? ???????????????? ?? ??????????? ??????????? ??????
?, ?????? ??? ??????? ?????, ?????????? ?
?????????? ?????? ????????? ????, ??? ???????????? ???????? ????????????? ????????????? ??????? ??? ???????? ???????????.
??????????? ???? ? ?????? ? ????-???????
24-26 °?, ?? ? 8,1-8,9, ?????????? ?????????
6,1-9,3 ??/?, ???????????? 3,5-4,5 ?, ?????????
10,1 ?, ???????????? ? ??????????? ????
? ??????? ????? > 100 ???. ??, ? ?? ???????
4,0 ? ????? 750 ??.
???????? ??????????? ??????????????? ???????????? ?????? ????????????
?????????? ?? ???????? ? ??????-????????
??????? ? ?????????? ?????? ????????? ????
???????? ??, ??? ? ????????? ????, ??? ? ?
?????? ???, ?? ??????? 2 ? ?????????? ?????? ????????? ?????? ??????????, ????????????? ???? ???????? ?????? ? ??????????? ????? ????? ??????. ?? ??? ? 50-70 ? ??
????? ?????? ?? ??????? 2,0-2,1 ? ??????????? ???????????? ?????????? ??????? ????
? ??????????? ????????? 40-70 %. ??????? ???? ??????? 0,9 ? ?????? ????????
Z. marina, ? ?????? ??????? 0,2 ? ? Z. noltii.
????? ?????? ??????????? ???????????????? Ruppia. ? ???????? ???????????? ??????????? Cladophora vadorum, R. maritima,
R. cirrhosa, Potamogeton filiformis, Cladophora albida (Huds.) Kьtz., C. sericea, C. siwaschensis C. Meyer, Chaetomorpha linum,
Chaetomorpha aerea (Dillw.) Kьtz, E. ahlneriana, E. compressa, E. intestinalis, Ceramium
strictum, C. tenuissimum (Lyngb.) J. Ag., Callithamnion corymbosum (J. E. Smith) Lyngb., Polysiphonia opaca, Lophosiphonia obscura (Ag.)
Falkenb., Ectocarpus confervoides ? ??? ?????? ?????. ????????? ?????????? 767 ?/? 2,
?????? 367 ?/? 2 ?????????? ?? Z. marina.
?????????????? ?????? ?????????? ??????????? 30 ?????.
? ??????????? ??????? ?? 4/0 ? ? ??????????? ?????? ??????????? ???????? ?????????? ?????????? ????????????? ?? 85 %,
# 129 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
?????? ?????? Z. marina ??????????? ??
25 %, ? Z. noltii ????????????? ?? 90 %. ?????
????????? ?? ????????? 831 ?/?2.
?? ??????? 5,0 ? ?? ??????-????????
?????? ? ???????? ???????? ???? ????????????????? ?????????? Zosteretum noltii
? ??????????? ????????? 50 %. ????????????? ???????? ??????????? ?? ?? ????, ???
???? ????????? ?? ?????????? ??????????,
?? ? ??????? ??????????.
? ??????? 5,1 ? ???????????? ??????,
??????????? ? ??????????????? ?????. ???
Z. noltii ??????????????? ????? ?????????
??? 300 ??, ? ? ????????? ?????????? ??? 160
??. ? ???? ?? ?? ??????? ?????? ??????????
??????????????????? ???? ??????? ??????
(???????). ????, ? ?????? ?????????? ??????,
?????????? ?? ???????????.
? ?????????? ?????????? ???????
????????? ???? ? ????????? ?????? ?????????? ??????? ???? ???????? ????????????? ?
??????? 0,4 ? ? ?? ??????? 2,0 ? ?? ?????????????? ??????? Ruppia cirrhosa (Petagna)
Grande, R. maritima ???????? ??????? ???? ??????????, ? Z. noltii ? ?????? ???? ??? ?????
??????????? ???????? 10-15 ??, ????????????????? ???????? ? ????????? 927 ?/?2.
?????????????? ?????? ?????????? ????????? ? ??????????? 10 ?????. ? ???????????
??????? ?? 4,0 ? ??????????? ???????? ????
?????????? ????????????? ?? 90-100 %, ? ????????? ? ?? 6455-7661 ?/?2. ?????? ????????
????? 1,3 ?, ??????? ? 0,4 ?. ? ??????????
??????????? ??????? ????? ??????????? ???????? ??????????? ?? 40-60 %, ? ????????? ? ?? 1328 ?/?2.
?? ??????? 4,5 ? ?????????? ?????????? ?????????? ? ?? ?????? ??????? ??????
???????????? ?????? ?????????? Ruppia.
????????????? ? ?????????? ????????
Z. noltii, Ceramium strictum, C. tenuissimum,
Chaetomorpha linum, Cladophora sericea,
C. siwaschensis, C. vadorum, Callithamnion
corymbosum, Polysiphonia opaca ? ?????????
??????????? ?????????. ????? ???????? ?????????????? ??????????? ???????? ?? ?????
????? ????????? ????, ??????? ???????
??????, ??????? ??????????? ?? 45 ?? ? ??????????? ???????????. ????? ?? ???????????? ?????????? ? ????????? ???? ?? ?????
???? ? ??????? ???????????? ??????????????? ??? ???????? ?????? ??????????????
??????? ????????? ??????, ?????? ???????? ??????????????. ? ???? ? ????? ??????????? ????????? ??????? ?? ?????? ??????
? ?????????? ?? ??????-???????? ????? ?????????? ????????? ?????????? ???????
????, ?????? ?????? ????????? ?????????? ?
??????????? ???? ????.
? ????????? ?????????? ?????? ??????? ??? ????????? ?????: ??????? ? ?????
?????, ??? ????????????? ? ????????? ????????? ???? ?? ????????? ?????? ????? ???????? ??????????????. ????????? ??? ?? ????
?? ?????? ?? ?????? ?? ??? ???????? ?? 6,38
?? 10,55 ?. ???????? ??????? ????? ???????
???????: ?? ? 8,2-9,1; ?????????? ????????? ???????? ??????? ? 7,7-9,6 ??/?, ????????
123-420 ??/??3 ??? ???????????? 3,5-4,5 ?.
???????????? ? ??????????? ???? ? ???????
? 14,00 ????? ????? 100 ???. ??, ??? ???? ??
??????? 5,5-6,5 ? ?? 2200-2600 ??, ?.?. ??? ??????? ???? ??????????????? ?????, ??????
??? ???????? ?????????? ? ??????? ????.
? ???????????? ? ?????????? ????????????? ??????? ? ????????? ?????? ??????????? ?????????? ???????? Zostereta, Chareta,
Polysiphonieta, Ceramieta, Phragmiteta ? ?????
????????? ??????????.
??? ? ?? ????? ??????????? ?????????
?????? ????????? ????, ?? ????? ???? ?????????? ?????? ?? ??????? ????? ??????????? ?????? ???????????? ??????????,
?????????? ??? ??????? ??????? ?????. ?
# 130 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
????? ?????????????? ? ??????????? ??????
? ?????? ????? ?? ??????-???????? ??????
?? ??????? 1,1 ? ???????????? ??????????
?haro-Potametum pectinati ?? 100 %-? ??????????? ?????????. ??????? ???? ????????
P. pectinatus, ??? ?????? 50-100 ??; ? ?????? ?
Chara canescens Desv. et Lois. ??????? 15 ??.
????????? ?????????? ????????? 3225 ?/?2.
???, ????????? ???????? ????? ???????????
?? 45 ?/?2, ??????? ? ?? 88 ?/?2. ?? ???????,
????????????? ????? ?????????? ????????
? ???????? ???????, ?? ??????? 5,0-6,5 ? ?
??????-?????????? ??????? ???????????
????? ???????? ?????????? Polysiphonio-Zosteretum marinae ? ??????????? ?????????
10-30 % ? ?????????? 200-650 ?/?2. ???????-
?????????? ?????????? ? ?????????? ??????????? Z. noltii, ? ?????? ????? ? Ceramium
tenuissimum.
? ????? ? ????????? 2-3 ? ??????????
?????? ????? ?????????. ???????????? ?????????? ?????????? Chara intermedia A. Br.
?? 100 %-? ??????????? ?????????, ?????? ????? 1,0 ?. ????????? ?????????? ??
10 000 ?/?2 ??? ??????????? 529 ???/?2. ? ????
?????????? ???? ????????????? ????? ??????? ?????????? ????????? ? ???????? ???? ?
15 ??/?.
????? ? ??????? 3,0 ? ??????????????
?????????? ?????????? Ceramio-Charetum
canescentis, ??? ? ??????? ????? ???????????
P. filiformis ? ???????? Z. marina ? Z. noltii, ?
? ?????? ????? ? ????????? Chara canescens
? Ceramium tenuissimum. ????? ???????????
???????? 40-50 %, ????????? 2733 ?/?2. ??-
??? ???????? ???, ??? ???????, ????? ?????
????????? ??????????? ?????????? ?????
??????? ????? ???????. ??? ?????????? ???????????? ????? ?????? ????? ??????????
??????? ? ????? ??????????? ?? ?????????. ????? ? ?????? ? ?????? ? ?????????
1,2-5,0 ? ?? ????? ??????? ??????-????????
?????? ??????????? ??????????? ?????????? Zosteretum noltii purum, ??????? ??????????? ????? ?????????? ??????? ? ???????
???. ?????????. ???????? ????? ??? ?????????? ??????? ? ????? ?? ??????? 2,0-3,5 ?,
??? ????? ??????????? ???????? ?????????
90-100 %, ? ????????? ? 554 ?/?2. ????????
???? ???????? ????? 60-70 ??, ??????? ? 15
??. ??????? ??????????? ???????? Polysiphonia opaca.
? ??????? 4,5 ? ? ?????? ????? ? ????????? 40-45 % ??????????? ??????????? ??????? ????????? Polysiphonia opaca
? Ceramium tenuissimum, ?????? ????????
?????, ????????????? ???????? ???????, ??????????? ?? 5-10 %, ?? ??? ???? ?????????
????????? ???????????. ? ?????????? ??????????? Chara intermedia, Ceramium elegans
Ducl., Cladophora siwaschensis, C. vagabunda ?
????????? ?????? ???? ?? ???????? ??????????. ? ??????????? ??????? ?? 6,0-6,5 ? ???
?????? ??????????? ?????? ???? ? ???????-
?????????? ?????????, ?????? ? ??????????? ??????? ?? 1,2 ? ? ? ???????????? ? ????????? ???? ?????? ?????????? ????, ? ?????
? ??????????? ??????? ?? 5,0 ? ? ??????????? ???????????? ??? ??????????? ???????????? ?????????? ????????? ? ?????????? ???????????? ???????? ?? 15-20 %, ?
????????? ? ?? 150-170 ?/?2. ????? ???????????? ? ?????????? ??????????? Callithamnion
corymbosum, Ceramium rubrum Huds.) Ag., C.
strictum, C. tenuissimum, Polysiphonia brodiaei
(Dillw.) Grev., P. opaca, P. subulifera (Ag.) Harv.,
Lophosiphonia obscura (Ag.) Falkenb. ? ?????????????? ???? ??????? ?????????? ?????
Cladophora, Enteromorpha, Chaetomorpha.
?????
????????? ? ?????????? ?????????
?? ?????? ?????? ????????? ???? ????????
????? ?????. ??????? ?? ?????????? ? ????
# 131 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
???????? ????? ????????????? ? ??????????????????. ????? ?????? ???????????? ????? ?? ?????? ?????, ? ??????? ?????? ??????? ????????? ????, ????????? ????? ?????
????????? ? ????????????? ?? ??????? ?????? ????? ?????????? ???? ? ??????? ????? ??
112 ??, ??????????? ?? ?????????? ???? ?????
??????? ?????????? ???????. ???? ??????
?????? ?????? 5 ?? ????????? ??? ?? ??????
? ?????????? ?????????? ????????? ????. ??
??????? ????? ????? (2560 ??2) ????? ? 10
??? ??????????? ???? ?? ????? ???????????? ???????? ????????? ???? ? ?????????
?????. ???????????? ??? ????????? ?????
????? ?????? ? ?????? ???? ?????????. ?????
?????????????? ?? ??? ?????: ?????????, ???
????????? ? ?????????? ????, ???????????
? ????????. ? ?????? ?????? ?????????????
?????????????? ?????????? ??????.
?????? ? ?????? ??????? ?????? ???????, ??????? ? ???????? ????? ? ???????.
??????? ??????? ? ???????? ????? 0,63 ?, ??
??????????? ??????? 2 ? ? ?????.
? ????????? ???? (????? 1985 ?.) ??????,
??????? ???????? ? ????????? ??????, ???????? ????????? ???????? ????????????????
??????????????, ????? ??????, ??? ?? ????????? ???????????? ??????? ???????? ?????????, ????? ?????????? ???? ????????????.
????????? ????? ?????????????? ?? ??????
?????: ????????, ????????????? ?????? ?.
?????????, ?????? ????????? ? ???? ? ???,
?????? ???? ?? ?????? ? ????? ????? ?????????? ? ?. ???????. ???????? ????? ??????? ?????? ????? ????? ????? ??????????
???????? ? ????????? ??????.
?? ?????? ????? (???. 2), ???????????
??????????????? ? ????????? ?????? ??????
?. ?????????, ???????? 12,3 ???. ?? ? ????????? ?? 1,46 ?, ??? ????????? 18,2 ?, ?????
? ??????, ??? ? ?? ???? ?????????? ????????? ??????, ?????????? ? ???? ?????????????
??????????????? ?????????? Phragmitetum
purum, ?????? ? ??????? 0,2 ? ?? ??????? ?
??????-???????? ??????? ??????????? ?????????? ????????? ????????? Cladophora
siwaschensis C. Meyer ? ????? ???????????
????????? 80 %. ? ???????? ????????????
? ?????????? ??????????? ????????? ?????????: Polysiphonia opaca, Enteromorpha
intestinalis, E. maeotica Pr.-Lavr., Ceramium
tenuissimum, Chaetomorpha linum, Z. marina
? Ruppia cirr?hosa (Petagna) Grande. ? ????????? ?? ?????? ? ??????????? ??????? ??
1,3 ? ????????????? ????????? ? ??????????
?????? ????????? ? ??????????. ?? ??????????? ???????? 50-80 %, ???????? ???????
????????? Chaetomorpha linum ???????????? ? ?????????? ??????????????? ?????? ?
???????? ?????? ????? ?? ??? ????? ???????.
????????? ?????????? 1600-2500 ?/?2 ? ??????????? ?? ???????????? ????????. ? ???????? ???????????? ? ?????????? ???????????
Z. noltii, Ceramium tenuissimum, Enteromorpha
intestinalis,
E.
maeotica,
Cladophora
siwaschensis, Polysiphonia opaca ? ?????????
?????? ????.
???????? ????? ??????? ????? ??????????? ?????????? (??????, ?????-???????,
1987) ???????? ? ?????????? ???????????????????? ???????????????? ?????? ????????????? ???????????????? ??????, ????
????????????? ???? ?? ???????? ? ?????
???? ????.
?? ?????? ???????? ?????? ??????????
???? ?? ??? ????, ????? ??????????? ????????????????? ????????????, ? ? ?????? ?????? ???? ???????????? ? ??????? ????????? ?????? ?????? ??????????? ? ????? ???
???????????? ?????. ??? ????????? ??????
?????????? ???????????, ? ????? ????????? ?????????? ???????? ??-?? ???????????
???????? ?????? ? ???????? ????????????
????.
# 132 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
???. 2. ????????????? ???????????? ????????? ? ?????? ?????
????????????? ? ????????? ?????? ????????? ??? ???????? ?????? ???? ?????????
????, ??????? ????????? ? ????? ????? 5-???????????? ?????? ?????? ? ?????? ????? 2
???? ?3, ?? ?????? ? ???????? ???? ????????? ???????? 300 ??? ?3 (???????, 1993).
? ????????? ?????? ??????? ????????? ???,
????? ??? ??????, ??????????, ?????, ??????? ? ??????. ????? ????, ? 1971 ?. ???????
? ????? ??????-???????? ????? ? ????????
???? 294 ?3/? ??? ??????????? ????? ??????????????, ?????????????????, ????????????
???????????? ??????, ? ????? ??????? ????
???????? ? ??????????????, ??? ????? ??????? ???????? 1580 ???. ?? (???). ??????? ?
?????? ?????? ???????? ????? 300?350 ?? ?
???, ?.?. ?????? ????, ? ???????????? ?????????? 2,5 ???? ?3, ?.?. ????? ?????? ?????? ?
???? ?????? ?????????????. ??????? ??????
????? ? ???????? ????? (???????, 1993) ?????? ?? 70 ??, ? ? ??? ????? ???? ?? ?????????
???? ?????? ??????? ????????? ? ?????.
? ????? ? ?????? ???????? ?????? ????
? ?????? ??-?? ???????? ????????? ?? ????
???????? ????????? ?????????? ???????????
??? ?, ???????????, ?????????? ?????????
# 133 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
???????????? ?????????????? ? ????????? ??
????????? ???????? ???????? ??????? Salicornia europea L., Salsola soda L., Halimione
verrucifera (Bieb.) Aell. ? ??????, ?.?. ?????????? ???????????????? ????????? ?? ??????
?????? ????? ?????????????? ?? ????????.
??? ?????? ??? ?????????, ?????????? ????? ? ??????? ?????? ?? ??????.
?? ?????? ?????, ??????????? ? ??????? ????? ? ???, ? ???????? 46,8 ???. ?? ? ?????????? ???? 24 ? ????? ?????????? ?????
?? ??????? ????????? ??????? ?? ???????
?????? ? ????????? ?? 0,4 ? ???????? ???????????? ?????????? Zoster?-Cladophoretum.
??????? ???? ??????? 15-20 ?? ?????????
?????????? ????? ? ????????? 15 %, ?????? ???? ? ??????? ?????????? Cladophora
siwaschensis ? ????????? 80 %. ?????????
?????????? 1490 ?/?2. ????????, ??? ???????
????????????? ??????????. ? ????? ? ?????????? ??????????????? ??????? ??????
?????? ??? ? ???????? ?????????????? ??????????? ????????????? ? ??????????? ???????????? ? ????????? ? ???????
???????? ???????????? Zostera, Ruppia,
Cladophor? ? Chara.
?? ??????? ?????, ????????????? ?????
??????????? ? ?? ??????? ??????????? ???
????? ? 2 ????, ????????? 21,5 ?. ??????????? ??????? ????? ????? ? ????????? 0,8 ?
????? ?????? ?? ??????? ?????? ???????? ??????????? ??????????? ?????????? Zosteretum
marinae. ? ???????? ???????????? ? ??? ??????????? Cladophora siwaschensis, Chaetomorpha linum, Ceramium tenuissimum, Z. noltii, R.
maritima. ????? ??????????? ???????? ?????????????? ?????????? 50-90 %, ? ?????????
600-2000 ?/?2. ?? ????????? ???????? ????-
????????? ??????? ?????????? ?? ?????????? ???????????? ????????????? ???????? ?
???????.
? ??????????? ??????? ?? 0,8 ? ??
??????? ?????? ????????????? ?????????
? ???????????, ? ??????? ??????? ???? ?????????? ????????? ???????? ???????, ?
?????? ? ??????????? ? ????????? Chara
neglecta Hollerb. ?? 90 %, ? ??????????
6000 ?/?2, ?? ??????? ?????????? 405 ?/?2, ? ??
?????? ? 937 ?/?2.
? ????? ????? ????? ??????? ????? ??????????? ???????????? ????????? ??????????????? ???? ??????? ?????????? Polysiphonia opaca ? Ceramium tenuissimum. ?? ?????
????????? ????????? 1667 ?/?2, ?? ?????? ????? ????????? ????????? ????? ??????? ?? ????
??????? ?????? (24 ?), ?? ?? ????? ???????,
????? ???????? ?.?. ??????? (1993) ??? ????????? ????? ?????? (178,8 ?).
?????????? ??? ??????? ????? ?????????????? ???????????? ????????? ?????????????? ?? ??????????? ??? ???????
?????? ??????? ??????? ????????????? ??????????? ? ????????? ?????????? Z. noltii, R.
maritima, Cladophora siwaschensis, Polysiphonia opaca, Ceramium tenuissimum, Chaetomorpha linum ? ????? ????????? 60-100 % ? ?????????? 380-1356 ?/?2.
?, ???????, ?? ????? ????? ?????,
???????? 47 ???. ?? ? ?????????? 19,6 ?
?? ??????? ? ??????-???????? ??????? ?
????????? 0,2-1,0 ? ??????????? ??????????????? ?????????? Potametum filiformis
purum ? Ruppietum marinae purum ? ??????????? ????????? 30-70 % ? ??????????
?? 801-1157 ?/? 2. ? ???????? ???????????? ?
?????????? ??????????? ?? ?? ????, ??????? ??????????? ??? ?????????? ????????
?????, ?????? ? ? ???? ?????? ? ????????
? ??????? ?????????? ???????????? ??????????? ????????? ?? ?????? ?????????????: Lyngbia aestuarii L. sordida Gom., Spirulina meneghiniana Zanard., Rivularia nutida
C. Ag. ex Born. & Flah ? ?????? ???? ????
Oscillatoria.
# 134 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
??????????
??????? ????????, ??? ??????? ????????? ???? ????????? ?? ????????, ?????
????, ? ???? ????? ?????? ????????????? ??
????????????? ????????. ?? ????????, ???
??????? ???? ??? ? ??????, ???????????
???????? ?????????? ? ???????????. ??????
??????????? ??????? ??????? ??????? ???????????? ??????? ???. ????????? ? ????????
???? ???? ???????? ??????? ? ??????????, ??????? ?????????? ???????? ? ????? ??????
???? ? ???????? ?????????????? ?? ????????.
? ??????????? ????????? ?????? ?????, ????
???????????? ????? ????? ?????????? ????????? ???????????????????? ???????, ?????????? ????????????? ?????????? ???????
??? ??????; ?? ????? ??????????? ????????
????? ??????? ????????? ???, ??? ? ?????
???? ???????? ???? ???? ?????????????? ?
???? ????? ??????? ?????????????? ?? ???????? ? ????? ???????? ?????????. ?????????? ???????? ???? ??????, ?, ?????? ?????,
?? ??? ??????????? ???????? ?? ????????
????? ??????????? ???????????? ?????????
??????? ?????????????? ? ??????????, ????????? ? ???.
????? ?????????? ?.?. ???????? (???? ?? ???) ? ?.?. ??????????? (?? ???? ??
???), ?????????? ????????? ???????????? ???????? ????????.
1
?????????? ?????? ?? ????????? ??????????? ?????. ?????? ?????, ? ???????? ???? ??????????? ?? Potamogeton
filiformis Pers., ? ??????????? ????????????? ?????? ???????????? P. pectinatus var. scoparius Wallr., ? ? ??????
? P. interruptus Kit. ????, ??-????????, ???? ? ????????????? P. pectinatus var. interruptus (Kit.) Ascherson, ???
??????? ?????????? ???????, 1-3 ??, ?????? (??????, 1988). ?? ?? ?????????, ??? ??? ????? ??????????? ?
????? ??????????? P. vaginatus Turcz.
?????? ??????????
??????? ?.?. (1957) ????????? ??????????? ??????? ??????? ????????? ????. ?: ?????
????, ???. 34, ?. 386-388,
????? ?.?. (1977) ?????? ?????? ???????? ????????? ???? ? ?? ?????? ? ???????? ???????. ?: ????????? ????????. ????. ?? ?????? ?????? ?????????, ?????. ?. 39-41,
?????? ?.?. (1927) ????????? ? ???????? ????? ????????? ????. ?: ??????? ? ?????
?.?. ?????????. ?.: ?????, ?. 235-240,
?????????? ?.?. (1951) ?????? ?????????????? ?????????? ?????? ? ?????????? ??????? ?
???????? ????. ?: ????? ?????????, ???.15, ?. 131-338,
?????? ?.?. (1983) ????????? ??????????????. ?: ?????? ???????? ?? ?????? ?????? ??????? ?? 25 ???, ??????/???. ?. 28-30,
?????? ?.?., ???????? ?.?. (1989) ?????? ?????????????? ??? ????????? ???????????????????????????? ????????? ???????? ??????? ????. ?: ????????? ??????-?????. ????.
?? ????????? ?????. ?????, ??????/???. ?. 25-27,
?????? ?.?., ???????? ?.?. (1990) ??????? ?????? ?????????????? ? ???????????????
???????. ?: ??????????? ???????? ????????????? ??? ???-??????????? ????? ????, ???????. ?. 23-24,
?????? ?.?., ??????????? ?.?., ???????? ?.?., ???????? ?.?. (1991) ??????????? ???????? ?????? ?????????? ? ?????????? ???? ????????? ????. ?: ????????? ?????? ????,
????????. ?. 25-27,
# 135 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
?????? ?.?., ?????-??????? ?.?. (1989) ?????? ?????????????. ????: ??????? ?????,
272 ?.
????????? ?.?. (1981) ???? ????????????? ????????? ??? ? ???????????????. ?: ?????????????? ??????? ???????? ?????????? ? ?????????? ????????? ????. ?.: ?????. ????????
????, ?. 137-151,
?????? ?.?. (1967) ???????????? ???????, ????? ? ??????? ?????????? ????? ?????
????. ?-?., 398 ?.
??????? ?.?. (1965) ???????????? ????????? ????? ????. ???????????? ???????. 3: 323335,
???????? ?.?. (1961) ???? ??????? ??????-????????? ????? ????????? ???? ? ??????????? ?? ????????. ?: ????? ????????????????? ???????? ?? ????, ?. 8, ? . 33-44,
???????? ?.?., ????????? ?.?. (1980) ????????? ???? ????????? ????. ?????? ?? ????:
???. ???, 186 ?.
????? ?.?. (1925) ????? ? ??? ?????. ?: ???. ???. ??????. ??-??, ? 15, ?. 22-43,
??????? ?.?., ??????? ?.?., ???????? ?.?., ?????????? ?.?. (1987) ????????????? ?????? ????? ?? ?????????????? ??????????????? ??????????. ????, 40 ?.
???????? ?.?. (1975) ?????? ? ??? ???? ? ?????? ???????????. ?????????????????? ????.
?I(1): 120-122,
????????? ?.?. (1896) ??????? ?????????? ????????? ?? «???????». ???????? ????????????? ???????? ????, ?. IV -V, ?. 389-408,
?????? ?.?. (1960) ????????? ?????????????? ????????????? ????????? ????????? ??????? ? ?????????? ???????????. ??????? ???, ???. ????????? ? ????????. 18: 124-143,
????????-???????? ?.?. (1955) ?????????? ????????? ????????? ????????? ????. ?-?.
222 ?.
??????? ?.?. (1964) ???????? ??????? ?????? ???????????????. ?: ?????? ?????? ???????????????, ?.: ?????. ?. 467-482,
???????? ?.?. (1973) ??????? ?? ???????? ? ??????? ????. ??????-??-????: ???. ??????/
???, 189 ?.
????? ??????? ???? (????????????) (1984-1985). ??????-??-????: ???. ???. ?. 1, 280 ?., ?.
2,. 240 ?.
???????? ?.?. (1986) ? ????????????? ????????? ????????? ?????????? ?????. ?: ?????????????????? ???????????? ?????????. ??. ??? ?? ????. ?.: ?????, ?.141, ?. 5-16,
?????-??????? ?.?., ?????? ?.?. (1984) ??????????????? ?????????? «????????? ??????» ?? ????. ????: ??????? ?????, 288 ?.
?????????? ?.?. (2002) ??????? ?????????? ???????? ????, ??????????? ??? ?????????? ???????????????? ?????, ?? ?????????????? ??????????? ??????????: ???????. ???. ...
????. ????. ????. ?????-?????????.: 19 ?.
??????? ?.?. (1993) ??? ? ???????? ????. ???. ???. ?? ?? ????. ?????. ????????????:
??-?? ??????????. ???. ????. ??-??, 202 ?.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????. ?????? ? ?????????-?????? ?????????????? ????????? ? ????????? ????????? ????????? ????
Aquatic and Coastal Vegetation of the Northern
and Western Coast of the Azov Sea
Valentin V. Gromov
Institute of Arid Zones SSC RAS, Rostov-on-Don,
41 Chekhov av., Rostov-on-Don, 344006 Russia
Description of the macrophyte aquatic and near-shore vegetation of the northern and western coast
of the Azov Sea from Taganrog Bay to the Gulf of Sivash, information about communities of plants,
their composition, distribution and quantitative characteristics are given. A brief analysis of the plant
communities distribution and its dependence of environmental factors, particularly the illumination
degree and turbidity, is presented.
Keywords: Azov Sea, aquatic macrovegetation, plant communities, plant association, light.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 138-150
~~~
??? 591.524.12 (261.74)
?????? ? ????????? ????????????
?????????? ??? ??????????
? ?????? ??????
?.?. ??????*, ?.?. ????????
????????????? ??????-????????????????? ????????
??????? ????????? ? ???????????? (??????????)
?????? 236022, ???????????, ??. ??????? ????????, 5 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
? ?????????? ????? ?????????? ????? 2007/08 ? 2009/10 ??. ???????? ??? ??????????
?????????????? ??????????: 1) ????????????? ???? ?. ???-????, ??????????? ?? ??????
?????? ????????????, 2) ???????????? ????, ?????????? ?????????????????? ??????????
?????, 3) ???????????? ????, ????????? ? ???????? ????? ??????, ????????? ?? ?????????
?????. ????????? ? ????????? ???? ????????? ???? ??????? ?o ????????? ?????????????
??????????. ??? ??????? ? ???????? ??????????? ? ???????? ???????????? ??????????
?????????. ? 2009/10 ?. ? ?????????? ?????? ?????? ??? ?? ????? ?????? Temora turbinata
????????? ??? ?????? ????????????? ???????? ? ??????? ?????????? ?????????? ?.
???-????. ????? ?????????? ? ?????????? ???????????? ???? ????????? ? ??????????
??? ???????????, ? ??????????????????? ???????? ????? ?????????? ???????????? ????
??????? ???????.
???????? ?????: ???????????????, ?????????? ??????, ?????????????? ??????????,
?????????, ??????????? ? ????????????? ?????????, ??????????????????? ????????.
????????
??????? ?????????? ??????? ?????? ?
????? ??????????????? ??????? ???????
????????? ???????? ??????. ??? ???? ?? ??????? ??????? ????????? ??????????? ?????????????????? ????????????? ?????????
(Sherman et al., 1993). ??? ???????????? ??
???????????? ??????????? ????? ?????????-
*
1
????? ????????? ???????? ?????? ????? 44°
? 12°?.?. ?????????? ???? ?????????? ?
????? ???? ??????????.
??????? ????????? ??????, ???????????? ?????????? ?????????, ? ??????? ????????????????? ???????????? ??? ? ?????????
?????????? ???????????? ?????????? ????????? ???????????? ???? ???? ?????????,
Corresponding author E-mail address: ksana-glushko@yandex.ru
© Siberian Federal University. All rights reserved
# 138 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
??? ? ????????? ? ?????????? ????????????
???????? ??????? ????????????? ??????? ???
???????????? ??????? ????????????? ?????????????? ???????????? ?????????? (???????, 1997).
????????? ???? ?????????? ?????????
??? ???????? ??? ????????? ? ?????? ?????????????, ???????, ?????????? ? ?. ???-????,
????????? ??????????? ????? (Aristegui et
al., 2009). ??????? ?????? ???? ??????????
???????? ???????? ? ??????????? ????????? ??????????????? ????????? ? ?????????
??????????. ????????? ?????????, ??????????? ????? ? ?????????, ????????????? ????????????? ???????????? ???????????? ???????????? ???????? ??????? ???????????? ?
????????? ??? ?????????.
?????????????? ???????????? ???????
??? ???? ??????????????? ??????????????? (??????, 1968; ????????, 1976, 2002; ??????? ? ??., 1997; Brenning, 1983;). ???????????? ??????? ????????? ???????????????? ??
???????????.
????
??????
?
????????
???????????????-????????? ????????????
????????? ???????????? ? ?????? ??????
? ????? ?????????? ? ????????? ?? ????? ?
????????????????? ?????????.
????????? ? ??????
?????????? ??? ??????? ????????????
????????? ????? ????????????????, ????????? ? ????? ?????????? ? ???? ????
?????? ??????????? ???????-????????????
? ?????????????????? ??????, ??????????? ?? ????? ???? «??????????»: ???
«?????????» (48 ????; 28.12.07-07.01.08
??.) ? ??? «??????????» (59 ????; 13.0123.01.10 ??.).
????? ??????? ?? ????????? ?????
21є ? 16є ?.?. ?? ??????????? ????? ???????,
????????????? ?? ???????????? ???????-
??????????????? ???????? ??? ?????????
20-1000 ?, ????????? ???? ?? ????? ?? ?????????? ????? 15 ????. ?? ?????? ???????
????????? ?? ????? ?? ???? ??????? ?????
???????, ????? ?????????? ????????????
????? ???????? ??????.
?????? ????? ? ???? «?????-20» ? ???????? ???????? ????????? 0,03 ?2, ??????????? ???? ?? ??????????? ???? ? ???????? ????
168 ???. ????? ??????? ? ??????? ????? ????? ????? ??????????-?????? ???????? (?? 2-3
?????? ?? ?????? ?????????) ? ????????????? ???? 0-100 ? ??? 0 ? ? ??? ?? ???? ?????
? ???????????? ? ???????????? ????????????
(??????, 1983).
?? ?????? 2007/08 ?. ??????? 31 ?????, ??
?????? 2009/10 ?. ? 27 ???? ????????????????.
?? ?????? ??????? ????? ?? ????? ?????.
??????????? ????????? ???? ????????? ?? ??????????? ???????? (??????????
???, 1982). ???????? ???????? ???????
?????????? ? ???????????? ????????????,
?????????????? ???????, ?? ???????????,
???????????????? ?? ????. ?????? ????- ?
??????????????? ????????? ??????????,
??? ???????, ?? ???????? ????? ????????
?????.
?? ??????????? ????????? ????????
?????? ??????? ????? ??????????? ? ?????
???????? ????????????, ? ????? ??????????? ? ???????? ??? ????????? ????????
(???/?3, ??/?3) ? ????? ????? (???????? ? ??.,
2005).
??? ?????????????? ??????? ????????????? ???????? ???????????? ?????????
?????: ??????????? ? ??????? ????????????? ????? 50 %, ?????????????? ? ???????????? ?? 25-50 % ???????, ????????? ?
??????????? ????? ??? ?? 25 % ???????
(?????????????? ???????, 2005). ???
??????? ??????????? ????????? ????????
??? ???????? ??????????? ?????? ???????-
# 139 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
?????????: ??????????? (?????? ? ??????),
???????????? ? ????????? (?????? ? ???????) ? ? ???????????? ? ???????????????
?.?. ???????????? (2003), ??. ???????
(Mouchline, 1998), ?.?. ?????????? (2001),
????????????????? ? ?????? ??????????? ?????? (Vives, 1982; Wu, 2004). ???????????? ????????????? ????????? ???????????????? ??????? ?? ????????????? ????
????????????? ??????????? ???????: ????????????, ??????-???????????? ? ????????????. ?????????????? ??????? ???? ??????? ? ??? ??? ???? ??????????? ??????????
?? ?????? ????????????? ?. ???? (Vives,
1982).
?????????????? ????????? ?????????
????????? ????????????? ???????? ? ?????? ???????? PRIMER® 6 (Plymouth Marine
Laboratory, United Kingdom). ? ?????????, ?
???????? ???????????? ?????????????? ????????? ??????????????? ??????????????
???????????, ?????????? ???????? ?????????????????? ???????????? ????????????
? ??????????? ??????? ???????????????????
? ?????????????????? (??????? ????? ???????????) ?????? ??????????? ???????? ??
???????????? ???????? ????-??????? ? ?????????????? ????????? ??????? ? ????????
???? ????????? (Clarke, Warwick, 2001). ???
??????????? ?? ????????, ????? ?? ?.?.
?????????? (1949) ? ?.?. ??????? (1985),
???????? ?????, ?????????? ????????????
?????? ? ????????????????? ?????????????
????????????? ????????? ? ??????????????
??????????? ????? ?????????? ??? ???????.
??????? ???????????? ??????? ? ?????????????? ???????? ????????????????? ? ????????????? ?????????????-??????????????? ??????? ???????. ?????????? ?????????????
????? ????????? ??? ????????? ???????????? ???????????? ? ????????????? (??????,
1989).
?????????? ? ??????????
??????????????? ???????
?? ???????????????? ?????????? ?????????? ?????????? ???????????? ???????
????????? ??? ???????????????? ??????????????????? ??????????????? ???????:
????????-???????????? ??????????? ???? ?
?????????? ????????? (????????? ? ??.,
2005). ????????????? ??????? ???????????
??????? ??????????????? ???????? ??????? ??? ????????? ????????????? (????????
????????????? ??????????? ?????? ?????
(????)) ? ??????, ????? ??????? ??? ??????
????????????? (????? ????????????? ??????????? ?????? ????? (????)) ? ?? ????????????? ??????????? (???????? ????????????? (???) ? ????? ????????????? (???)
???) (???????? ? ??., 2010). ? ?????????? ?????
?????????????? ? ?. ???-???? (21° ?.?.) ??????????? ??????????? ???? ? ???????????
????????????????? ? ??????????????? ??????? (??? ?????????? ????????-????????????
????? ? ???), ? ??????? ?????????? ??????????? ???????? ?????????????, ???????? ? ????????????? ???????? ? ??????????
?????????????? ???? ?????????, ???????????? ? ?????? ????, ? ???????????? ????????????? ????????? ?????? ?????????????? (??????, 2010).
?????????
????????-????????????
??????????? ???? ? ????????????? ???? ???????? ?? ???????. ????? ??? ????????? ????? 21° ?.?. ? ?????????? ???? ????? ???? ?? 0
?? 800 ?. ????? ?????????? ??????????????
?????????? ???????, ?????????? ???????? 40????????? ???? ????????? ??????????????
?????? ??? ? ???????? ??????????? ???? ?
??? ?? 10-13є ?.?. ?? ???????? ???? 40-50 ?,
?????????? ??????? ??????, ??????????? ????
????????? ???? ????????? ? ?. ???-????.
?????? ?????? ??????, ???????????
??????????????? ????? ? ????????? ???
# 140 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
? ????????? ??????????, ? ??????????
?????????, ??????? ???? ???????????????
???????? ?????????????. ??? ??????????
????????????? ? ????????? ?????????? ??????????? ? ??????? ?? ???????? (???????????, 1998). ????????? ?????? ????? ???????? ? ??????????-????????? ?????????
????????????? ?????? ????? (??) ??? ??????????????? (??????, 2003).
????? 2007/08 ? 2009/10 ??. ????????? ???
???? ???????? ??? ?????? ?????????? ? ????????????????? ?????????????? ? ????????????? ???? (0-40 ?) ???????? ?????? ?????
????????? ????????????? ? ??? (t?=16-20 є?).
?? ?????????? ????. ??? ? ?????????????
???? (0-40 ?) ???????????? ? ??? ??? ???? ????? ?????? ????????????, ?? 12є50' ? 12є40'
?.?. ??????????????.
????? ????, ??????????????? ??????? ??
??????????? ????????? ? 2007/08 ?. ??????????????? ???????? ???? ???????? ????????
??????????. ?? ???????? ??????? ?????????
????????????? ????? 20°40' ? 19°00' ?.?., ??
?????? ? ?? 17°45' ?.?. ?? ????? ??????? ??-
?? ??????????????? ?????? ???????, ?????????? ????? ?. ???-???? (Vives, 1982), ? 15,4 %
?? ??????? ?? ??? ??????????-????????? ????????? (Razouls et al., 2005-2011).
????? ????, ???????????????? 32 ??????? ????- ? ??????????????? ?????????? ?????? ??????????????? ????? (???????, ?????,
?????????, ???????, ????????? ? ?.?.). ?????
???????????? ???????????? ? ? 2007/08, ? ?
2009/10 ??. ?????????? ???????? ??? ????????? ?????????? ?????? ????: Penilia
avirostris Dana, 1849, Evadne nordmanni Lovйn,
1836, Pseudevadne tergestina (Claus, 1877),
Podon leuckartii (G. O. Sars, 1862).
? 2007/08 ?. ??????????? ????? ???????????? ???? ???????????? 19 ?????????
(????. 1). ?????? ?????????????? ???? ???????? 21 ??????, ? ?????? ????????? ? 76 ????????.
? 2009/10 ?. ??????????? ????????? Centropages chierchiae Giesbrecht, 1889, Paracalanus indicus Wolfenden, 1905, Temora turbinata
(Dana, 1849). ? ?????? ???????????, ??????
?????????????, ????? 16 ???????? (????. 1).
??? ??????????? ????????? ??????? ? ??????????? 18°30'-16°15' ?.?., ?? ?????? ? ?? 16°35'
?.?. (??????? ????? ? 48 ??????, 2008).
????? 2009/10 ?. ????????? ??? ????? ??????? ?? ????????: 20°00'-20°40' ?.?. (?????
?. ???-????), 19°15'-19°40' ?.?. (????? ?. ???????), 17°40'-17°50' ?.?. ? ?? ???, 16°10' ?.?.
????? ????, ? ? 2007/08, ? ? 2009/10 ??. ??????????? ???????? ?????? ???????????? ??? ?
??????? ?????? ?? ??????? 18-19°00' ?.?. (??????? ????? ? 59 ??????, 2010).
? ?????????????? ???????? 23 ???????, ?
????????? ? 72.
???????? ???????? ??????? ????? ??????????? ???????? ? ???????????????
???? ???? ????? 68 %. ????? ??? ??????
???? ???? ??????? C. chierchiae, P. indicus,
T. turbinata, Acartia danae Giesbrecht, 1889,
Calanoides carinatus (Krшyer, 1848), Oithona
nana Giesbrecht, 1892, Oithona plumifera Baird,
1843, ? ????? Chaetognatha, Appendicularia,
??????? Decapoda, ??????? Calanus ? Eucalanus, ??????? ???. ?????? ? ??????? ??????????? ????? ??????? ? ????????? ????????. ???, ????????, ? ????? ? ??????? ??????
??? ???? ?? ????? ?????? ? 2009/10 ?. ??????????? ???????? ???? T. turbinata ? Euterpina
acutifrons (Dana, 1848), ? ????? ??-?? ????????? ?????????? ??????????? ???? ??????-
??????? ??????
????????????????
????? ? 2007/08 ? 2009/10 ??. ???????????????? 150 ????? ? ???????? ????? ???????? ?????. ????? ?????????? ????????????
(Copepoda) ?????????? 114 ?????, ??? 46 %
# 141 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
??????? 1. ??????? ????????????? (??, %) ? ????????????? ?????????????? ??????????? ??????????
???????????? ? 2007/08 ? 2009/10 ??.: ? ? ???????????? ???, ? ? ????????????, ?-? ? ??????-????????????,
???????? (Vives, 1982)
2007/08 ?.
??????
Chaetognatha
2009/10 ?.
????????
-
??
??????
100,0 Centropages chierchiae
?-?
96,8
Paracalanus indicus
Paracalanus indicus
???????. ?
96,8
Temora turbinata
Acartia danae
Centropages chierchiae
????????
??
?-?
100,0
???????. ? 100,0
?-?
100,0
???????. ?
90,3
Chaetognatha
-
96,3
Calanoides carinatus
?-?
90,3
Decapoda ???????
-
92,6
Oithona plumifera
?-?
90,3
Calanoides carinatus
?-?
92,6
77,8
Decapoda ???????
Temora stylifera
Eucalanus monachus
Clausocalanus furcatus
-
90,3
Euterpina acutifrons
?
???????. ?
83,9
Calanus ???????
-
74,1
?
80,6
Appendicularia
-
70,4
?-?
74,2
Bivalvia
-
70,4
Eucalanus ???????
-
71,0
Eucalanus ???????
Pisces ???????
-
71,0
Oithona nana
-
66,7
???????. ?
66,7
Nannocalanus minor
?-?
67,7
Oithona similis
???????. ?
63,0
Temora turbinata
?-?
67,7
Acartia danae
???????. ?
59,3
67,7
Oncaea media
Appendicularia
Oithona nana
???????. ?
?-?
59,3
64,5 Pisces ???????
-
55,6
61,3
Pisces ???????
-
51,9
-
51,9
?-?
51,9
Calanus ???????
-
Penilia avirostris
?
61,3
Polychaeta
Oncaea conifera
???????. ?
58,1
Oithona plumifera
???????????? Oithona similis Claus, 1866 ? Oncaea media Giesbrecht, 1891.
????? ???????????
? ????????
????? 2007/08 ?. ??????????? ???????????? ?? ???????? ??????????? ?? 741 ??
64 051 ???/?і, ? ??????? ? 11 377 ???/?і. ????????????? ??????????? ?????????? ???????????? ???? 93 %, ???????????? ? 2,1 %,
?????? ??????? ????- ? ???????????????
?????????? ? 4,9 %. ?????? ??????????? ??????????: P. indicus (28,7 %), ??????? Calanus
(14,2 %), C. carinatus (9,5 %), O. media (7,1 %).
?????????? ??????????? ??????????? ??
?????? ?????? (20є40' ? 19є00' ?.?.), ????????
???? ?????????? (???. 1).
????? 2009/10 ?. ??????????? ???????????? ?????????? ? ???????? ?? 32 ??
42 894 ???/?і, ? ??????? 8360 ???/?і. ????????????? ??????????? ?????????? ? ???????????? ???????????? ??????????? ?? ??????????
(89,6 ? 3,2 % ??????????????), ? ?????????????? ??????? ???????? ??????????? ?? 7,1 %.
?? ??????????? ???????????? Oncaea curta
Sars, 1916 (18,3 %), E. nordmanni (11,7 %), P. indicus (11,3 %), T. turbinata (6,0 %). ?????????? ???????????? ???????????? ????????
???? ?? ?????? ? ?????????? ? ????? ??????? ???, ???? ? ???????????? ?????, ???? ???,
????????, ????????? ?? ???? ?????? ? ?????????????? ?????? (???. 2).
???????????? (47,0 % ?? ??????????? ?
2007/08 ?. ? 46,6 % ? ? 2009/10 ?.) ? ??????-
# 142 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
21°N
?
4
3
21°N
?
1
6
4
10
8
20°N
14
15
1
6
10
8
3
13
11
20°N
14
15
11
13
16
16
20
20
18
21
23
23
24
2000
27
1800
25
27
25
1600
18°N
28
42000
21
2200
24
54000
19°N
17
60000
48000
18
19°N
17
18°N
28
1400
36000
31
30000
29
32
800
34
18000
36
34
600
17°N
33
12000
29
32
1000
24000
36
31
1200
17°N
33
400
39
6000
38
37
0
38
37
0
40
41
18W°
39
200
41
16°N
16W°
17W°
18W°
40
16°N
16W°
17W°
???. 1. ???????????????? ????????????? ??????????? (?) (???/?і) ? ???????? (?) (??/?3) ???????????? ?
???? 0-100 ? (???) ????? 2007/08 ?.
21°N
21°N
?
2
3
1
?
9
2
11
5
7
20°N
12
13
11
16
20°N
12
13
15
17
1
9
5
7
3
15
16
19
17
19
19°N
22
19°N
24
22
42000
24
8500
21
38000
21
7500
25
34000
28
25
28
6500
18°N
30000
26000
29
31
34
38
34
1500
17°N
33
6000
2500
36
10000
500
37
36
17°N
33
38
37
-500
-2000
41
18W°
31
3500
18000
2000
29
4500
22000
14000
18°N
5500
17W°
40
41
16°N
16W°
18W°
17W°
40
16°N
16W°
???. 2. ???????????????? ????????????? ??????????? (?) (???/?3) ? ???????? (?) (??/?3) ???????????? ?
???? 0-100 ? (???) ????? 2009/10 ?.
???????????? ???? ??????? (35,8 % ?? ??????????? ? 2007/08 ?. ? 44,8 % ? 2009/10 ?.) ??????????? ?? ??????? ????? ?????????????
????????? ??? ?? ??????, ??? ? ? ???????????? ?? ????? (????. 1). ??????-????????????
???????? ??????????????? ??????????? ?
????? ??????? ??? ? ???? ?????????????????
?????? ?? ?????? ??????. ????????????? ??????????? ????? ????????????? ????????????? ????????????? ? ??????????? ? ???????? ????? ?????? ? ?????????? 17,2 ? 8,7 % ?
2007/08 ? 2009/10 ??. ??????????????. ?????-
# 143 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
??????? ???? ???????? ???????????? ?????????? ??????-????????????? ? ?????????????
?????????????. ? 2007/08 ?? 2009/10 ??. ???
???????? ? 49,5 ?? 63,4 %.
? 2007/08 ?. ???????? ???????????? ????
???????????? ????????????, ????????? ? ??????? ????????? ?? 41 ?? 2430 ??/?і (???. 1).
??????? ???????? ???????? ???????? ? ????
??????, ??? ??????? ??????????? ?????????
????????? (?? 69 %) ? ??????? ????????????
??? (?? 50 %), ? ????? ? ???????????? ????
????? 19°30ґ ?.?., ??? ?? ???????? ???????????? P. avirostris (17,3 %). ?????? ????????
??????????? ???????? C. carinatus (14,2 %)
? P. indicus (9,2 %), ? ????? Chaetognatha
(10,9 %), ??????? ??? (7,6 %), ??????????
(7,0 %) ? ??????? ???????????? ??? (4,7 %).
????? 2009/10 ?. ???????? ????????????
???? ?????????? ? ??????? ???????? (?? 2 ??
8734 ??/?і), ?????? ??????? ???????? ? ???
? ?????? ???? ???????????? ?????????? ??????? (735 ? 740 ??/?і ??????????????). ?????????? ???????? ?????????? ?? ?????????? ??????? ? 24 ????? ?. ???????, ??? ??
?????? ?????????? ??????? ???????????? ????? ??? (42,3 %) (???. 2).
???????????? ?????????
????????????????
??????? ??????????? ???????, ?? ??????
?????? 2007/08 ?., ???????? ??? ?????????????? ?????????? (???. 3). ?????? ??????????
????????????? ? ?. ???-????, ?????????????
??????????? ? ?? ?????, ? ???? ???????. ????????, ??? ???????????? ??? ???????? ????????? ??? ??????, ?????????????? ????? 19° ?
23°?.?. ?? ???????? ???? 40 ? ? ????????????
???? ? ???? (???. 4). ?????? ??????????
???????????? ?????????? ? ????? ??????, ?
?????? ? ??????????? ???????? ? ??????? ? ?????? ????????????? ?????????????.
?????????? ?. ???-???? ? 2007/08 ?. ???????????? ??????????????? ??????????????? ????????????? ? ??????-?????????????
??????, ????? ??????? ???????????? P. indicus ? O. media (????. 2, 3). ??? ????? ???????????? ?????? ??????? ????????????, ??????
????????????? ? ???????? ??? ????????????
??????? ???????????. ? ??????????? ????????? ??????????? ?????? ??????????? ? ?????? ????????? (????. 2).
?????? ?????????? ? ??????????
???? ? ???? ?????????????? ???????????-
(?)
(?)
???. 3. ???????????? ??????????? ??????? ????????????? ??????????? ???????? ???????????? ?????
2007/08 ?. (?) ? 2009/10 ?. (?)
# 144 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
21°N
?
4
3
21°N
?
1
6
2
15
1
9
10
8
3
5
7
20°N
14
11
20°N
12
13
11
13
15
16
16
20
17
18
19
19°N
17
21
19°N
22
23
24
24
27
21
25
25
28
18°N
28
31
18°N
29
29
31
32
36
34
34
17°N
33
39
36
17°N
33
38
38
37
37
41
18W°
40
17W°
??????????: ?. ???-???? ?
41
16°N
16W°
18W°
, ???????????? ???? ?
40
16°N
16W°
17W°
, ?????????? ???? ?
???. 4. ???????????????? ????????????? ?????????????? ????????? ? 2007/08 ?. (?) ? ? 2009/10 ?. (?)
??????? 2. ?????????? ????????? ????????? ???????????? ????? ????????? ??????????
2009/10 ?.
????????????
????
?. ???-????
??????????
????
????????????
????
???????????, ???/?
??????????
????
??????????
?. ???-????
2007/08 ?.
10028
15979
6570
13167
6659
279
????????, ?/?і
0,3
1,0
0,6
0,5
1,1
0,02
?????? ???????, ???/???
3,97
4,41
4,46
3,61
3,89
3,86
?????? ?????
0,86
0,9
0,9
0,87
0,92
0,87
-???????????? ????? ???????, %
59,2
44,1
47,6
61,5
40,9
23,1
-???????????? ????? ???????, %
6,0
16,5
24,6
6,7
10,1
8,7
-??????-???????????? ?????
???????, %
34,7
39,4
27,8
31,8
49,0
68,2
-?????? ????????????, %
7,7
20,5
10,0
24,8
23,6
5,8
-?????? ????????????, %
47,7
50,1
58,0
29,8
45,1
49,7
-??????? ??????????, %
0,2
0,3
0,6
0,2
0,8
2,3
-?????? ??????????, %
33,7
21,3
21,8
41,5
20,0
34,4
-?????????????, %
10,9
7,7
9,7
3,8
10,5
7,9
????????????? ???????????
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
??????? 3. ????????????? ??????????? ???????????? ?????, %
??????????
?. ???-????
?????????? ????
???????????? ????
2007/08 ?.
2009/10 ?.
Paracalanus indicus
51,8
Oncaea media
24,9
Oncaea media
32,9
Temora turbinata
22,7
Paracalanus indicus
20,0
??????
23,3
??????
24,3
Paracalanus indicus
32,1
Paracalanus indicus
44,2
Calanoides carinatus
18,0
Calanoides carinatus
14,1
??????
49,8
??????
41,6
Paracalanus indicus
42,1
Oncaea conifera
31,7
Oncaea conifera
6,3
Clausocalanus jobei
11,8
Calanoides carinatus
6,0
??????
45,6
?? ? ??????-????????????? ??????, ????????????????? ?????????????? P. indicus ?
?. carinatus (????. 2, 3) ? ????? ????????
?????????? ??????? ???????? ????????????
? ?????????????. ??????????? ? ????????
?? ????????? ? ??????? ???????????? ????
??????????, ??????????? ????????? ?????????? ???? ?????? ???????????.
? ?????????? ???????????? ???? ????
???????????? ???????????? ????, ?? ????????????? ??????????? ???????????? ????? ??
????????? ? ??????? ???????????? ??????????? (????. 2). ?????? ?????????? ??????????? P. indicus, Oncaea conifera Giesbrecht,
1891 ? C. carinatus (????. 3). ??????? ??????? ? ????? ????????? ?????????? ????????
??? ?????? ??????????? ? ???????????? ????????? ???????? ??????????. ? ??????????? ????????? ???? ??????????? ??????
??????????? ? ?????? ?????????.
?? ?????? ?????? 2009/10 ?., ???? ???????? ??? ??????????? ?????????? (???. 2),
??????? ????? ??????? ????????????? ????????????? (???. 4), ????????? ? ??????????
?????? (????. 2, 3). ?????? ?????????? ?. ???-
Oithona plumifera
11,0
Paracalanus indicus
10,8
??????
34,7
???? ? ?????????? ?????????? ???? ??????????? ?? ???????????? ??????? ?????????,
???????? ?????? ?????????? ????????????
???? ? ???????? ????? ??????. ?????? ????? ? ?????????? ?. ???-???? ?????? ????????????????? ?????? ? P. indicus ? O. media
????????? ???????????? ??? ? T. turbinata,
?????????? ??????????? ?????? ??? ????.
??????????? ? ???????? ?????????? ????????????? ????????, ? ?? ????? ??? ?????? ??????? ????? ?? ?????????.
??????? ????????? ?????????? ?????????? ????, ??? ? ????????, ???????? ????????, ???? ??????????? ????????? ????? ? ???
???? (?.?. ???????? ???? ??????? ????). ?????? ???????? ???????????? ??????????, ?
?????? ????????????? ????????????? ?????????? (????. 2).
? ?????????? ???????????? ???? ?
2009/10 ?. ?????? ?????????? ????????? ??????????. ? ??? ??????? ?? ?????? ????????????
O. conifera ? ???????????? P. indicus, ?? ????? ??????-???????????? Clausocalanus jobei
Frost & Fleminger, 1968 ? O. plumifera (????. 3).
???? ???????????? ? ???????????? ???????,
# 146 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
???????????, ???????? ?????????? ? ??????
??????? ??????? ????????? (????. 2), ?????? ??????????? ???????? ????????????? ??????????? ??????-???????????? ?????. ?????
????, ??????????? ??????? ???????????????
??????????. ????????? ?????????? ? 2009/10
?. ??????????? ?? ???? ????????? ??????????? ????????? ????? ? ?????? ???????????? ?????.
????????? ? ????????? ????????????
? 2009/10 ?., ????????, ???????? ??????? ??
???????????? ??????????????? ??????????
?, ? ?????????, ??????? ? ????????? ??????????. ???????? ????????????? ??????????
???????? ?? ????? ???????? ??????????? ?
???????? ???????????? ?????????.
? 2009/10 ?. ? ?????????? ?????? ??????
??? ?? ????? ?????? T. turbinata ????????? ?
??????? ?????????? ?????????? ?. ???-????.
????? ?????????? ? ?????????? ???????????? ???? ????????? ? ?????????? ??? ???????????, ??? ???? ???????? ???? ?????????????????? ??????? (????. 3). ? ??????
?????????? ?????????? ????????????? ????
?? ????????? ???????????, ??? ? ?????????????? ?????? ???????????? ????? ?????? ??
??????? ??????.
?? ???? ??????? ? ??? ?????? ? ??????
?? ????????? ???????????? ?????? ??????????? ? ?????? ?????????, ??? ???????????????
? ??????? ???????????? ????? ??????? ????????, ??? ???????????? ? ??????????????
???????????, ? ?????????? ????? ? ???????
?????????????? ??? ??????????? ?????????.
????????? ???????????????? ?????????
?????????, ????????, ??????? ? ??????????
????????????? ???????? ??? ? ?????????
??????????, ? ?????????, ? ????????????
????????? ?? ????????????? ? ????????
????????????????? ???????? ? ??????????
??????? (???????? ? ??., 2010).
??????????
????????? ???????????? ????? 2007/08
? 2009/10 ??., ???????? ?? ??????? ???????????????? ??????????, ????? ?????? ?????.
???????? ??????????? ???????? ?????? ????? ??? ?? 68 %. ???????? ?? ?????????? ?
2009/10 ?. ??????????? ??????????, ???????
???????? ???????? ???????????? ?????????
? ??? ????. ??????? ??????????? ? ????????
???????????? ?????????? ??? ????????????
????????? ??? ? ??? ??????????.
???????? ??? ?????????? ?? ???????
?????????????? ??????????:
? ?????????? ?. ???-????, ??????????? ? ????????????? ????, ???????????? ??????? ??????? ????????????
?????????? ?????, ??????????? ???
????????? ????????? ???? ???????.
???????????? ? ??????????? ?????????? ?????????????? ???? ???????????????? ??????????? ???? ? ????
???????????? ???? ? ????.
? ?????????? ?????????? ????, ?????????? ?????????????????? ?????????? ????, ????????? ??? ???????? ????? ?????? ? ????????????
?????????? ???????????? ? ?????????????????? ?????.
? ?????????? ???????????? ????, ???? ?
???????????? ? ???????? ??????????
?????????? ? ??????-????????????? ?
????????????? ? ?????????????, ????????? ??? ???????? ????????????
???.
?????? ???????? ????????? ????????????? ?.?. ???????????? ? ?.?. ???????? ??
????????????, ?????????? ? ???????????? ?????? ? ?????? ??? ???????.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
?????? ??????????
?????? ?.?. (1989) ???????? ? ????????????? ?????????????. ?????????: ???????????????, 152 ?.
??????? ?.?. (1997) ?????????? ?????? ??? ???-???? ? ??? ??????? ? ?? ???????????
?????????????: ???. ? ????. ????. ????. ???????????, 195 ?.
???????? ?.?. (1949) ?????? ????????? ????. ??. ?????????, ???. 13, 195 ?.
??????????? ?.?. (1998) ???? ? ??????????? ? ???????????? ???? ??????????-?????????
?????????. ???????????: ??????????, 196 ?.
?????? ?.?. (2010) ??????????????? ????????? ? ???????????? ????????????? ?????????????? ??? ? ?????? ?????????? ??????????: ???????. ????. ???????. ????. ???????????,
22 c.
???????? ?.?. (1976) ??????????????? ?????? ? ??????? ? ????????????? ?????????
??????????? ???????????? ? ??????????-????????? ?????????. ?: ????? ??????????. ????????????? ???????????? ? ????????????? ?????? ? ?????????? ????. ???????????: ???-??
??????????, ???. LX, ?. 124-129.
???????? ?.?. (2002) ?????????? ???????????? ???????????? ? ????????? ??????????
? ?????? ?????? 1998-2000 ?????. ?: ????? ??????????. ??????????-????????????? ???????????? ?????????? ? 2000-2001 ?????. ?.1. ????????????? ????? ? ???-????????? ?????
?????? ??????. ???????????: ???-?? ??????????, ?. 85-94.
?????????? ?? ?????????????? ????????? ???????? ??????? ????????? (1982) ????????
?.?. (???) ???????????: ?????, 29 ?.
?????????????? ?????? ???????? ? ????????????? (2005). ????????? ?.?. (???) ???????
??????? ??????, ??????????? ?????? ?.?. ????????. ????????: ????? ???, 404 ?.
??????????? ?.?., ???????? ?.?. (2003) ?????????????????? ?????? ??????? ??????????? ???????? (Crustacea, Copepoda, Calanoida): ??????????? ? ???????????? (?????). ??????:
???????? ????. ??????. ?????? ?? ??????????, 30 ?.
???????? ?.?., ???????? ?.?., ??????? ?.?. (2005) ?????????????? ???? ?????? ? ??
???????????? ? ??????????. ?: «??????????? ? ?????????????????? ???? ??????: ???????,
?????????? ? ?????????????»; «????????? ????????????»: ????????? ?????????? ????, ?.
????, ???. 2005 ?. ??????-??-????: ???-?? ??? ???, ?. 67-70.
???????? ?.?., ???????? ?.?., ????????? ?.?. (2010) ???????????? ????????????? ???????????????? ? ???? ?????????????? ?????????? ? ???????? ????? ????????????? ???????.
??????????? 50 (3): 387-395
??????? ????? ? 48 ????? ??? ? 1704 «?????????» (2008) ?????? ?????: ??????????????????? ????????? (?????????????? ????????????? ???? ??????????? ??????? ? ????????? ?????????? ??????????, 05.11.2007 ?. ? 28.01.2008 ?. ???????????: ??????????,
61 ?.
??????? ????? ? 59 ????? ??? ? 1711 «??????????» (2010) ?????? ?????: ??????????????????? ????????? (?????????????? ????????????? ???? ??????????? ??????? ? ????????? ?????????? ??????????, 10.11.09 ?.? 12.02.2010 ?. ???????????: ??????????, 52 ?.
????? ?.?. (1985) ???????????? ??????????? ????????. ???????????????, ?????????
?????, ????????. ?.: ?????, 287 ?.
# 148 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
?????? ?.?., ?????????? ?.?., ?????????? ?.?. (1983) ???????????? ???????? ?? ?????
???? ???-, ?????????????? ?????????????????? «?????» ? ?? ?????????. ???????????: ??????????, 36 ?.
?????? ?.?. (1968) ? ????????????? ????????? ? ?????? ?. ???-????. ??????????? 8(3):
479-486.
?????????? ?.?., ??????? ?.?., ???????? ?.?. (2001) ????????????????? ???????? ??????? ???????? ????? ???????????? ?????????? ??????. ???????? ???? 56: 80-85.
?????? ?.?. (2003) ????????? ? ???????? ??? ? ?????? ?????????? ?????????? ? ????????? ????????? ???????????? ???: ???????. ????. ???????. ????. ???????????, 23 c.
????????? ?.?., ?????? ?.?., ??????? ?.?. (2005) ????????? ? ???????? ??? ? ???????
?????????? ? ???????????? ??????????? ? ?? ??????? ?? ????????? ???????????? ???. ???????????: ??????????, 198 ?.
??????? ?.?., ?????? ?.?., ??????? ?.?. (1997) ????????????? ???????? ???????????? ?
????????????? ???? ??? ?????????????? ??????. ??????????? 37 (5): 739-744.
Arнstegui J., Barton E.D., Alvarez-Salgado X.A., Santos A.M.P., Figueiras F.G., Kifani S., Hernandez-Leon S., Mason E., Machu E., Demarcq H. (2009). Sub-regional ecosystem variability in the
Canary Current upwelling. Progress in Oceanography 83(1-4): 33-48.
Brenning U. (1983) Beitrдge zur Calanoidenfauna (Crustacea, Copepoda) vor Nordwest- und Sudwestafrica. Die Vertreter der Familie Aetideidae. Wiss. Z. Univ. Rostock 32 (Naturwiss. Reihe 5):
1-7.
Clarke K.R., Warwick R.M. (2001) Change in marine communities: an approach to statistical
analysis and interpretation (2nd Edition). Plymounth: Plymounth Marine Laboratory, 175 p.
Mouchline J. (1998) The biology of calanoid copepods. Advances in Marine Biology. V.33,
149 p.
Razouls C., de Bovйe F., Kouwenberg J., Desreumaux N. (2005-2011) Diversity and geographic
distribution of marine planktonic copepods. Electronic resource. http://copepodes.obs-banyuls.fr/en
Sherman K., Alexander L.M., Gold B.D. (Eds.) (1993) Large marine ecosystem: Stress, Mitigation
and Sustainability. Washington, DC: AAAS Press, 294 p.
Vives F. (1982) Sur les copйpodes de la rйgion CINECA (Parties nord et centrale). Rapp. P.-v.
Rйun. Cons. perm. int. Explor. Mer. 180: 289-296.
Wu Ch.-H., Hwang J.-Sh., Yang J.-S. (2004) Diets of three copepods (Poecilostomatoida) in the
Southern Taiwan Strait. Zoological Studies 43(2): 388-392.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????, ?.?. ????????. ?????? ? ????????? ???????????? ?????????? ??? ?????????? ? ?????? ??????
Composition and Structure of the Zooplankton
in Coastal Waters of Mauritania in Winter
Oksana G. Glushko and Viacheslav V. Lidvanov
Atlantic Scientific Research Institute
of Marine Fisheries and Oceanography
5 Dmitry Donskoy Str., Kaliningrad, 236022 Russia
In the coastal waters of Mauritania three stable zooplankton community 1) community upwelling
zones cape Cap Blanc, situated in the north of the study area; 2) community neritic zone in the highly
productive coastal area; 3) community oceanic zones, remote from the coastline were identified in
winter 2007/08 and 2009/10.
Changes in the structure of these communities depended on the hydrological conditions, and, in
particular, were associated with a decrease in the intensity of upwelling. This resulted in a decrease
in the zooplankton abundance and species diversity of selected communities. Temora turbinata,
an indicator of water of southern origin, was dominant in the community of the cape Cap Blanc in
2009/10 as a result of inflow of warm waters into the north district. A change in dominant species in
the community oceanic zones was due to an increased proportion of neritic species. Composition of
dominant species of community neritic zone remained the same.
Keywords: mesozooplankton, cluster analysis, zooplankton community, upwelling, trophic structure,
habitat structure, structure-complex.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 151-159
~~~
??? 595.384.12
? ??????? ???????? Palaemon elegans Rathke 1837
(Decapoda, Palaemonidae) ? ?????????? ??????
?.?. ???????????*
??????????????? ??????????????? ??????????? ???????????
?????? 236000, ???????????, ??. ?????????, 1 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
???????????? ???????? P. elegans ? ???????? ???????? ? ?????????? ????. ?????? ??????????
?????, ????? ??????? ? ???, ????????, ??????? ???????????? ??????? ?????????????
????????????? ?? ?????? ?????? ??? ???????????? ?????? ?? ????????? ? ??? ???????????
(?? ?????? ??????, ? ?????? ?????). ????????, ? ???? ??? ????????? ??????????? ?????
??????????? ????? ??????. ??? ?????? ????????? ?? ????????? ??????????? ?????, ?
??????? ???????????? ??????? ??????? ?????? ????-?????????.
???????? ?????: Palaemon elegans, ??????? ?????????????, ???????, ??????????? ???????
?????, ??????, ???????????? ???????.
????????
???????? Palaemon elegans Rathke 1837
? ????? ??????????????? ??????? ???????
???? ?????????? ? 2000 ?. ? ???? ? ??????
???. ???????? ?? ????? ?????? ????????????
????????. ??? ?????????????? ??????????
???????????? ? ?????? ?????-???????? ????????? ? ????????? Crangon crangon ?? ???????? 0,5-0,7 ?. ????? P. elegans ????? ?????????? ? ????????? ?????? ???? (?????, ?????,
???????) ? ?????????? ?????? ? ?????????
?????????? ????, ? ?????????? ?????, ? ???.
????????. ????????, ?????? P. elegans ????????? ?????? ?????????? ????? ???????????
??????, ??? ?? ??? ?????? ? ??? ???????? ??-
*
1
???? (Grabowski, 2006), ???? ????????????
?????? ?????-????????? ????????? ??????
? ????? ?????? ?????-?? ???? ? ??? ??????????? ?????. ?? ?????? ??????, ?? ?????????
??????????? ? ???????? ????????? ????? ???
(?????? ????????? ?.?. ??????). ???????
???? ?????? ?????? ? ??????????????? ?????????????? ??????? ???? ????? ???? ???
??????????? ??? ????? ? ??????????? ?????
??????????? ?????? ??????????? ????.
????????? ? ??????
???????? ?????? ? ?????? ??????????
????? ??????????? ?????? ? ?????? ???. ????????? ? ? ??? ?????????? ??????????? ??
Corresponding author E-mail address: burukovsky@klgtu.ru
© Siberian Federal University. All rights reserved
# 151 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
?????????? ???? (???. 1). ????? ????????
? ????? ????, ???? ? ?????? ??????? 20032004 ??. ???????? ?????? ? ?????????? ?????, ? ???????? ?????????? ?? ??????? ?? 0,7
?, ? ? ??????????? ? ????? ? ??????, ? ???? ?????????? ?? ???????? ????????? ?????????
???????????, ?? ??????? ?? ????? ???? ? ??
?????? 0,5 ?. ? ???????? ?????? ???? ???????????? ????? ? ????? ?? 1 ?? 5 ??. ?????????
???????? ????? ?? ??????????? ? 4 %-? ???????? ?????????. ????? ???? ??????? ? ?????????? 330 ????????, ? 201 ?? ??? ? ????????
???? ?????????? ????, ? 54 ??????? ????
???????.
? ??????????? ???????? ???????????
?????????????? ???????, ??????????? ????????? ????? ?????????, ???????????,
??????????? ????, ??????????? ? ??????? ???
(??????????? ?????????? ????????????? ???-
?????????), ??????????? ?????? ?????????????? ???????? ? ?????? ??????? ??? ? ??????
???????? ????????. ????????? ???????????
?? ??????? ???? ??????? ????? ?? ????????
??????? ???? ????????? ? ????????? ?? 0,1 ??
(???????????, 1992; ?????????, 2008).
??? ???????????? ??????????? ???????? ???????????? ???????????? ????????
(???????????, ????????, 1970; ???????????,
1985; ???????????, ???????, 2008; ???????????, 2009). ??????? ????? ???????? ? ????? ???? ?? ????? ?????. ? ???????? ???????? (????? ?????????? 1 ? 2) ???????????
???? ?????? ??????????. ? ?????? ????????
????????? ??????????? ???? ???????? ???????? ???????? ????? ? ????????? ?? 10 %.
??????? ? ????????? ???????, ???????????? ????? 10 % ?? ?????? ???????? ?????,
?????? ?????????????. ?? ??????????? ?????
???. 1. ????? ?????? ????? ? ?????????? ?????, ?????????? ????, ? ????????? ???? ????? ???? ??
??????? P. elegans
# 152 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
???????????? ??????? ????????????? (??????? ??????????? ??????? ?????????? ?
????? ????? ????????????? ???????? ? ?????), ??????????? ????????? (??), ???????
????????????? ??????? ?????????? ???????
???????? ? ??????? ??? ????? ?????, ??????? ? ???????????? ???????? (??? ??? ?????????? ?? ?????? ???????? ????? ????????).
??????????? ?? ? ???? ?? ???????????, ??????????? ?????? ? ??????? ????? ???????
??????????? (???????????, ????????, 1970).
?? ??????, ?????????? ??? ??????? ??????
????????, ?????????????? ?????????????????? ??????????? (???????????) ???????
?????, ?.?. ??????? ???? ??????? ????????
??????? ? ?????? ???????? ?????, ?????????? ? ????????? (???????????, 1985, 2009) ?
??????? ????????????? (???????????, 1979).
????????? ?????????? ???????????? ???????
????????????? ?????? ????????, ? ???????
???? ?? ????? ???????? 60 % ? ????? ?? ?????? ???????? ?????. ????????????, ??????????????? ??????? ????? ????????, ???? ???????? (???????????, 1985, 2009).
??????????
?????????????
?????????????? ????????
??????? ???????? (????? ?????????, ??)
??????????? ?? 2,1 ?? 11,8 ??. ??? ?????????????? ?? ??? ??????: ?????????? (??? ??????????? ?? ????????? ??????? ?????????)
? ?? 2,1-5,9, ???? 2,5 ??; ????? ? ?? 3,7-8,0,
???? 5,5 ??; ????? ? ?? 4,0-11,8, ???? 7,5 ??.
?????? ???????? ????? (53,2 %) ????? ?????? ? III ?????? ????????, 28,6 % ????? ????? ??????????? ? ?????????????? ??????.
?????????????, ??????????? ???????????
????? ?????????? ?? ??? ??? ???? ???????
?????????? ? ???????? ??? ?? ????????, ? ??
????????? ?????? ???? ???????? ?? ???? ??????????????? ???????.
????? ??????????????
??????????? ????????
??????? ????? P. elegans ????? ??????????????? ??? ?????? ????, ?????? ????????????. ? ?????? ????????, ??? ???????,
??????????? ?????? ? ???????????? ???????.
?????? ???? ??? ????????-?????????? ????????? ???? ???????, ? ? ?????? ???????? ?
???????????? ?????.
???????????? ??????? ???? ???????????? ??????? ?????????? ? ?????????
??????? ? ?? ???????????, ? ????? ???????????. ????? ???????????? ???????? ???????? ????? ???????? ?? ?????????. ???
?????? ????? ?? ???? ????? ????? ??? ?????
?????????? ?????????. ? ??????? ?????????? ???? ?????????? ??????????, ??????? ?
????-??????? ?????????, ??????????? ? ????
??????? (Pennatophyceae, Centrophyceae, Chlorophycea, Chroococcophycea, Hormogoniophycea). ????? ???? ?????????? 23 ????.
???? ?????????? ?????. ??????? ?? ?????????? ???????, ??????????? ???????
??????? ?????-???? ???????? ???????.
??????? ?????? ?????? ???? ??????????
???? ? ?????????? ???????? ??? ? ?????????????? ??????????? ? ??????, ?? ??????
?????. ????? ????? ? ??????? ?????? ?????????? ??????? ??????. ?? ?????????????? ? ?????-???? ?????????? ????????
????????????? ?????????? ???? ??????????
??-?? ??????? ??????? ???????????????? ?
?????????????.
?????? ? ???? ? ???????? ??????????
????????? ??????? ????????? ????????????.
?? ???????????? ??? ??????, ???????, ???????????? ? ????????. ?????? ???? ?????
???? ???????????? ??????????? ?????????
0,04-0,2 ??. ??? ????????????? ??????????????? ????? ????? ????? 3-15 ??. ? ?????
??????? ???? ????? ?????? (Neomisis integer).
??????? ???????? ??????????? ? ???? ?????-
# 153 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
??? ?????? ? ???????? ????. ? ????????? ???????? ????? ???????? ???? ???????? ?????
(?? 7-9 ???. ??????). ???????? ?????????????? ? ???????????? ???????. ????????? ???????? ????????? ????????????, ??? ????? ???
?????? ?????. ?????????? ????? ???????,
??????? ? ??????? ??????????? ????????????, ?????? ??????????? ??????? ??????????
???? ??????????.
????? ?????? ???????? ?????? ? ?????????? ???????? ???? ??????? ????????? ?
???? ???????? ??????, ????????? ????, ???? ?
????????? ??????? ? ?? ????????? ?????????. ?????????? ???? ??????? ????? ?????
?????? 3,92 ??.
???? ? ???????? ???? ???????????? ?????? ? ?????????? ???????. ???? ?? ?????
? ???????? ?????, ? ????? ?? ????????? ????????, ??? ???? ??????, ??????? ???????,
????????, ??? ??????????????????. ????-
?????? ????? ??????? ??????? ??????????
?????????.
??????? ?????????????
???????? ??????? ? ????????
?? ??????? ????????????? ? ????????
???????? P. elegans ??? ??????? ???????
????? ????????? ?? ????????? ?????. ?????? ? ????????????. ? ??? ?????? ????????????? ??????????? ? ?????? ??????? ??????
(88,9 %) ? ? ?????? ?????? ? ??????? ????????? (52,1 %). ? ?????????????? ????????
??????? ????? ??????? ?????????? ????????? (36,7 %), ???????????? ??????? (30,8 %) ?
???????????? ????????? (28,4 %) (????. 1).
??????, ??????? ? ?????????? (???????
????????????? 11,6-8,6 %) ???????? ????????????? ????????? ????? ??????????????? ?
?????????? ????????? ??????? (? ????????
????????????? 2,3 % ? ?????). ?? ????? ??-
??????? 1. ??????? ????????????? ???????? ??????? ? ???????? (% ?? ????? ????????????? ????????) ?
?????? ??????????? ???????? ????? (% ?? ????? ?????) ? ???????? P. elegans
??????? ???????
??????
??????? ?????????
???????
?????????????
88,9
???????????
??????? ?????
70,4
???????
?????????????
79,6
52,1
0,1
?
?????????? ????????
36,7
?
?
???????????? ???????
30,8
17,6
5,6
???????????? ?????????
28,4
1,4
?
??????
16,2
?
?
???????
11,8
0,1
?
??????????
8,6
?
?
????
2,3
?
?
???????? ?????????
1,5
?
?
????????????
0,7
?
?
????????????? ????????
0,3
?
?
????????
0,3
?
?
?????????????? ???????
8,3
?
?
?????
78,6
10,4
?
????? ????????
255
54
46
??????????? ?????????
2,89
# 154 #
85,2
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
????? ???????????? ?????????????? ????????? ???????.
????????, ??? ? ??????, ???????????
??????????? ? ?????? ???????, ???? ??????? ??????? ??? ?? ??????? ????????????? (78,6 %). ??? ??????????????? ? ???, ???
P. elegans ???????? ??????????????? ?? ???.
?????? ????????????
???????? ?????
? ??????????? ??????? ????? (????. 1)
????????? ?????????? ??????, ???????????? ??? ????? ??? ?????? (70,4 %), ? ???????????? ??????? (17,6 %). ??? ??? ???????
??????????, ? ????? ???????????? ????????? ? ????? ???????? 99,8 % ?????? ????????
?????. ??? ????????? ?????????? ????????
????? ??????????? ?? ????? ?????????????
???????? ? ??????? ????????.
????? ???????? ?????? ????? ? ??????
???????????? ???????? ????? ? 10,4 %, ?.?.
???????????? ? ???????? ? ???????????? ????? ???????????. ????? ?????? ???? ????? ?
????? ????????????? ? ??????? ???????? ???
?????????????. ??? ????????? ????????????,
??? ?? ????????????? ????????? ???????? ?
???????? ????? ???????.
???? ???????????? ?????????? ? ??????? P. elegans ??????? ??????, ??? ??? ???
????? ??????????? ? ????????? ??????????
? ?? ???????? ???? (1,4 %) ?????????? ?????
??????????????. ????????, ??? ?????? ??????? ????????.
??? ????? ???????????? ???? ??????? ?
???????????? ???????? ? ??????? ????????
??????? ?? ?????????????. ? 85,2 % ??????
???????? 60 % ? ????? ??? ?????? ?????????? ?????-?? ???? ??????? ??????. ??? ??????
??? ???????????? ??????? (?????????????? ?
79,6 ? 5,6 % ????????).
????? ??????? ?????, ??? P. elegans
???????? ??????????????? ????????????,
? ??????????? ? ?????????? ??????????? ?
????? ???? ? ??????????? ??????? ???????.
???????????????? ????????????
??????? P. elegans
????????? ?????????? ?????? ????????
? ????? ?????????? ???? ????????, ?? ????? ??????? ???? ???????????????? ????????? ??????? ????????????? ????????????
??????? ???????? ???????????? ????. ???
????? ??? ???????? ???? ????????? ?? ??????
?????? ? ??????????? ?? ????? ?? ?????????: 2-4,0, 4,1-6,0, 6,1-8,0, 8,1-10,0 ??. ??????
?????? ??????? ???????? ?? ?????????? ???????? ?????? ???? ? ?????? ??????, ?? ?????? ?????? ????? ??????????????? ?????
? ????? ????????? ????? ?????? ?????, ? ??????? ?????? ?????????? ???????? ? ??????
??????????? ??????? ????? ? ?????? ?????. ? ????????? ?????? ???????????? ??????
??????? ????? ? ????????, ?????? ? ???????
?????????? ?? ?????? ?????? ?????????????
(???. 2).
? ???? ????????, ?????????? ?? ?? ????????, ?? ??????? ????????????? ???????????
??????. ?? ? ?????? ?? ?????????? ??????????? ? ?????? ????? (95 %), ? ? ???????? ? ??
????? 6 ?? ???????? ????????????? ???????
???????????, ? ????? ????? ??????? ? ?????? ??????????? ?? ???? ????? (?? 66,7 %). ?,
????????, ??????? ????????????? ???????????? ???????? ? ???????????? ?????????? ?
??????????? ???????? ??????? ?????????? ?
29-30 ????? ?? 40 %. ? ????? ??????? ??????
????????? ?????????? ????, ? ??????? ????????????? ?????? ???????????? ???????? ?????????? ????????????? ?? 50 %.
??????? ????????????? ??????? ????????? ? ??????????? ???????? ?????? ????????????? ? 22,0 ?? 35,7 %. ?????? ? ????????,
???????????? ????? ????????? 6,1-8,0 ??,
????? ????????? ?? 20 %. ????? ?? ?????????
# 155 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
???. 2. ???????????????? ???????????? ??????? ????????????? ??????????? ???????? ????? ? ????????
Palaemon elegans: 1 ? ??????; 2 ? ???????????? ???????; 3 ? ??????? ?????????; 4 ? ?????????; 5 ?
?????????? ????????
?????????????? ? ? ????????????? ???????????. ? ?????? ?????? ?? ??????? ????????????? ? ??????????? ???????? ?????????? ??
19,5 ?? 42,2 %, ? ?????? ? ?????? ?????????
6,1-8,0 ?? ????????? ?? 33,3 %.
??? ???????? ????????? ???????? ???????, ?? ??????? ?????????? ??????????? ????
????? ? ?????? ? ?????? ?? 4-6 ??, ???????? 48 %, ? ????? ????????? ??????????? ??
12,5 %, ? ??????? ????????? ???? ????? ?????????? ? ???????? ? ???????? ? ?? 6-8 ??.
? ????? ?????? ? ??????? ?????? ? ???????
???? ????? ??????????? ?????? ? ???????????? ???????.
??????????
????, P. elegans, ???????? ????? ???????
???????? ???????????? ?????????, ? ?????????? ?????? ????? ???? ??? ????????, ?????????? ?? ????? ??????? ?????? ??????
????????. ??????, ? ??? ???? ???????????? ? ??????????? ????????? ? ?????????
??????? ? ??????????, ?? ??? ????????? ?
??????? ????????? ????. ? ???????? ???????? ????????? ?????????? ??? ??????
??????? ????????: ?????? ? ????????????
???????, ?????????????? ?????????????
???????????. ??? ?????? ??????????? ? ???????????? ?????????? (2008, ??. 199) ??
?????????? ???????? ? ?????????? ?????
??????????? ?????? ? ??? 2001 ?.: «? ????? ?
????? ?????? ??????????? ?????????? ??????
?????????? ????? ? ????? ???????, ???????
???? (?? ?????????? ???????????) ????????
???????? ??? ?????????? ????????????; ???
???? ? ???????? ????????, ??????? ?????».
?????? ? ???????? ???????? ???? ????????
P. elegans, ??????? ? ????????? ????? ? ?????????? ?????? ??????, ????????, ???????
???????????? ??????? ????????????? ????????????? ?? ?????? ?????? ??? ???????????? ?????? ?? ????????? ? ??? ???????????.
????????, ? P. elegans ??? ????????? ??????????? ????? ??????????? ????? ??????.
? ??????? ????? ???? ? ?????? ??????
?????? ??? ??????????? ?????? ?? ????????. ???, Kцhn ? Gosselk (1989) ?????????,
??? ?? ??????? ????? ?????????? ?????
? ??????? ????? ???? ????????? ?????????, ????????, ??????, ?????? ?????????? ?
????????????? ????????, ????????, ?????????, ?????? ????, ??????, ??????? ????????? ? ?????? ???. ? ???? ????????
P. elegans ????????? ?????????, ????????,
# 156 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
????????-?????????????. ? ?????????, ??
???????????? ?? ????, ????? ??????? ??????? ?????????? ? ???? P. elegans. ЦstlundNillson ? ???????? (2005) ????????, ??? P. elegans ? P. adsperus ? ???????? ????????????
????????? ? ???? ????????-????????????,
?????? ????????????? (????????? Gyrodactylus sp.) ? ???????.
?????? ???? ??????? ? P. elegans ?????,
P. adspersus ? P. serratus, ??????? ? ??????
?????? ?? ???????. ?????? ?? ??? ? ???????? ????? (Inyang, 1977/78) ???????? ??????????????? ??????? ????????????? (????????????, ????????, ????????), ???? ??
???????????? ??????????, ??????????, ?????????????? ? ??????????? ??????????,
??????????, ? ??????????? ??????? ???????
??????????? ????????, ???????, ?????????? ? ??. ??? ????????????? ??? ??? ????????????????. ? ???????????? P. adspersus
?????????? ????? ???????? ???????? ??????
???????? ????? (????????? ?????????)
(????????, ???????, 1940). ?? ? ????????????
???????? ? ??? ???????? ????????? ??????????? ?????????, ? ??? ????? ??????????,
? ????? ??????? ?????? ???????? (Inyang,
1977/78). ?? ??????? ????????????? ? ???? ?
?????? ??????? ?????????? ????????????
(?????????????? 35,4 ? 38,9 %), ?? ?? ??????
????? ? ?????? (20,4 ? 35,7 %, ??????????????). ?? ??????? ????? ????????? ????????,
?? ????????? ?????? ??????????? ???? ???????????? ???????????? ???????? (14,3 %
?? ?????? ???????? ?????). ???????? ?? ??,
??? ? ???? P. adspersus ??????????? ???????
???????? ????, ???????????? ???? ???????
? ???????????? ???????? ??????. ??? ??????? ?? ???? ???????? ? ????? ???? ? ????? ?.
???? (???????? ???????), ??? ??? ???????
??????? ?????????? ????? ???????? ??????
????????. ?????? ??????? ?? ?????? ?????,
??????? ??? ????????? ?????? ?? ?????? ??-
??????? ? ?????? ? ? ?????? ???? (Figueira,
1986). ?????????? ???? ???????????? ???????? ? ??????? ? ??????????? ????? ?? ????????? ? ????? ??? ??????? ???????????, ???
??? ??? ????????? ?? ?????????? ???????????? ? P. elegans ? ?????????? ??????. ?????
???????? ??????????? ????????????? P. adspersus ? P. elegans ??????????. ???????? ??
?? ????? ????????????? P. serratus, ????????? ? ????? ???? (Figueira, 1986).
??????? ???????????? ???????? ? ??????? ???????? ???. Palaemonidae, ????????,
????? ??????? ?????????? ????????. ?? ????
????????? Kensley ? Walker (1982), ?????????
????????? ?. ????????. ?? ?? ????? ???????? Jayachandran (2001), ?????????? ?????? ?
??????? Macrobrachium rosenbergii, M. malcolmsonii, M. gangeticum ? M. idella idella. ?????? ???? ? ??? ??????. ????????, ? ????????
????? ??????????????????? ?????????? ????
?????????? ?????? (33,5 %), ????? ? ???????
????????? (17,2 %), ???????? (12,65 %), ???????????? ??????? (8,32 %), ???????????????? ????? ????????, ?????????? (5,91 %),
??????? ???????????? (5,72 %), ???????? ????????? (5,28 %), ?????????? (1,38 %), ??????
(1,28 %), ???????????? (0,94 %), ??????? ?
????????? ??????? ???. ??? ?????? ????????
? ?????? P. elegans. Sitts ? Knight (1979) ???????? ???????? ???????? ? ??????? P. macrodactylus ? ???????? ??????????-??? ??????
(??????????). ???????? (1958) ?????????? ?
????????????????? ???????? ??????? ???????? Leander modestus, L. paucidens, Macrobrachium asperulum, Palaemonetes sinensis ?
??. ????? (??????? ??????) ? ?????????, ???
???????? ????? ???????? ????????????? ???????????? ?????, ??, ??? ????? ???????????,
???????????? ???? ?? ?? ?????, ? ??? ? ????????, ????????????????? ????. ???????
?????? ???????? ?? ????? ????????. ??? ?????????? ? ??? P. elegans.
# 157 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
?? ????????? ? ???????? ???? ???????? ????? Leander ? Palaemonetes, ? ?????
???????, ? Macrobrachium ? ? ??????, ????
???? ??-???????. ????????????? ?????? ????
????? ??????????????? ????????? ????????????????? (???????????, 1985, 2009), ? Macrobrachium asperulum ? ??????????? ??????? (????????, 1958; ???????, 1998). ?? ??
????? ????????? ? ? ?????? ??????????
???? ?????????. ???, ?????? ? ???????????? ?????, ???????? ????????? ?? ????????? ????????. ????????, ? P. macrodactylus
???? ???????????? ???????? ????? ?????????
? ?????? ??????, ?? ?? ????? ???????? ??????????????? ?????? ???????? (Sitts, Knight,
1979). P. macrodactylus ??????? ??????????? ?
Crangon franciscorum ??-?? ?????? Neomysis
mercedis.
?????????????, ?????????????? ???. Palaemonidae ????? ??????? ???????????, ??????????? ????????, ???????????? ? ????????
?????. ???? ???? ???? ??????????? ???????
????? ?????????? ?????? ???????? ? ??????????? ?? ????????? ???????. ??? ?????????
??? ??????? ?? ?????????-??????????????
(???????????, 1985, 2009).
P. elegans, ??????????, ???? ????????,
???????? ????? ??????? ?? ??????? ????????? ???? ? ????????-?????????????. ??
?????????? ?? ?????? ????????? ???? ????? ???, ???, ?????? ?????????? ?????, ????
??????? ? ???, ????????, ????? ?? ???????
???????????? ??????? ????????????? ????????????? ?????? ?? ????????? ? ??? ??????????? ?? ?????? ?????? ??? ???????????? (??
?????? ??????, ? ?????? ?????). ????????, ?
???? ??? ????????? ??????????? ????? ??????????? ????? ??????. ??? ?????? ?????????
?? ???????????? ??????????? ?????, ? ??????? ???????????? ??????? ??????? ??????
????-?????????.
?????????????
????? ?????????? ?.?. ??????????, ???????????? ??????? ? ????? ?????????? ? ?
???????? ?????????????? ???????, ?????????????? ??????? ???????? ??? ????????????
????????????, ?.?. ?????????, ?????????????? ????????? ? ??????? ?????????? ???????? ????????, ?.?. ?????????, ????????????
???????????? ?????????, ??????????? ? ???????? P. elegans.
?????? ??????????
??????????? ?.?. (1985) ? ??????? ?????????????????? ????????. ????. ????. 64 (10):
1501-1512.
??????????? ?.?. (1992) ???????? ?????????????? ??????? ????????? ??????????? ? ?????????????? ????????. ?: ??????????-????????????? ???????????? ??????? ??????????????.
?.: ?????, ?. 77-84.
??????????? ?.?. (2009) ??????? ? ??????? ??????????????? ????????. ???????????:
???????????? ???? ??? «????», 408 ?.
??????????? ?.?., ???????? ?.?. (1974) ?????? ? ???????? ???????? ????? ? ??????
??????? ??????????????. ???????????. 14 (1): 167-172.
???????? ?.?., ??????? ?. (1940) ??????? ???????? Leander adspersus. ????. ????. 29 (1): 134-137.
???????? ?.?. (1958) ????????? ?????? ?? ???????? ???????????? ??????????????? ????????, ? ????? ? ???????????? ?? ??????????????. ??. ???????????. ? 9: 80-101.
??????? ?.?. (1998) ??????? ???????????? ??????????????. ?.: ??-? ?????. ????. ????.
??. ?.?. ????????? ???. 319 ?.
# 158 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????????. ? ??????? ???????? Palaemon elegans Rathke 1837 (Decapoda, Palaemonidae) ? ?????????? ??????
????????? ?.?. (2008) ? ???????? ???????? Palaemon elegans ? ????? ???????????????
??????. ?: ???????? ????????????? ? ?????????????. ?????? ???? ? ?????. ???????????:
???-?? ???? ??? «????», ?. 180-201.
Figueira A. (1986) Alimentation de Palaemon adspersus (Rathke, 1837) y Palaemon serratus
(Pennant, 1777) (Decapoda: Natantia) en la ria de Vigo (N.O. Espana). Cah. Biol. Mar. 27 (1): 77-90.
Grabowski M. (2006) Rapid colonization of the Polish Baltic coast by an Atlantic palaemonid
shrimp Palaemon elegans Rathke, 1837. Aquatic Invasions. 1. Issue 3: 116-123.
Inyang N.M. (1977/1978) Notes on food of the Baltic palaemonid shrimp, Palaemon adspersus
var. fabricii (Rathke). Meeresforsch. 26: 42-46.
Jayachandran K.V. (2001) Palaemonid prawns. Biodiversity, taxonomy, biology and management.
Enfield, NH, USA: Science Publishers Inc., 500 p.
Kensley B., Walker I. (1982) Palaemonid shrimps from the Amason basin, Brazil (Crustacea:
Decapoda: Natantia). Smiths. Contrib. Zool. 362. 28 p.
Kцhn J., Gosselk F. (1989) Bestimmungsschlьssel der Malacostraken der Ostsee. Mitt. Zool.
Mus. Berl. 65 (11): 3-114.
Цstlund-Nillson S., Becker J.H.A., Nilson G.E. (2005) Shrimps removed ectoparasites from fishes
in temperate waters. Biol. Lett. 1: 454-456.
Sitts R.M., Knight A.W. (1979) Predation by the estuarine shrimps Crangon franciscorum Stimpson and Palaemon macrodactylus Rathbun. Biol. Bull. 156: 356-368.
On Palaemon Elegans Rathke 1837
(Decapoda, Palaemonidae) Nutrition
in Vistula Lagoon
Rudolf N. Burukovsky
Kaliningrad State Technical University
1 Sovetsky av., Kaliningrad, 236000 Russia
The epibenthic shrimp P. elegans is a recent invader in the Baltic Sea. Settling the Vistula lagoon shrimps
are likely basic consumers of plant detritus at the different stages of its formation with periphyton
which is connected with detritus (at any case during the summer). The shrimps have probably no
serious competitors among autochthonous species of the Vistula lagoon. This may influence on the
structure of trophic webs where benthivorous fish are consumers of high levels.
Keywords: Palaemon elegans, feeding, frequency of occurrence, virtual food lump, detritus, the
vegetative debris.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 160-168
~~~
??? 594.3: 574.587
??????? ??????, ????????????? ?????????????
? ????????????? ??????????????
?????????? ????????? ?????????? ???
????????? ?????? ??????????? ????
?.?. ??????????*
??????????????? ??????????????? ??????????? ???????????
?????? 236000 ???????????, ??. ?????????, 1 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
????????? ???????????? ????????? ?????????? ????????? ? ?????????? ???????? ?????????
?????? ??????????? ????. ???????????, ??? ?????????? ???????? ???????????? 38 ??????,
10 ?? ??????? ????????? ? Prosobranchia, 28 ? ? Pulmonata. ???????? ????????? ?????????
????????? ?? ??????????? ?????????? ?????????????? ? ???????????? ????????,
?????????? ??????? ?????????? ?????? ??????????????. ??????? ??????????????? ?????
? ?????????????? ?????????????? ????????? ????????? ? ???????? ???????? ????????
???????? ? ?????? ?????????? ?????? ??????????????.
???????? ?????: ?????????? ????????, ???????? ?????, ?????????? ????.
????????
???????? ????? ? ?????????????????? ??????, ?????????? ?? ??????????? ????
????? ????????, ?????? ??????????? ?? ????
????? ??? ?????? ? ????? ? ??????? ?????????? ? ?????????? ?????, ??? ??????????
??????? ???????? ?????????? ?? ??????????.
????? ?????? ???????? ? ??????? ??????? ? ??????????????? ????????? ? ????????
?????? ????????? ? ??????? ???????? ?????????????? (Hilbert, 1908; Szidat, 1926), ? ???
????? ? «???????????? ???????? ? ??????
*
1
????????? ????????» (Geyer, 1927). ?????
??????????????? ??????????? ???????? ????????? ? ????????? ??????????????? ???????? ????? ?????? ? ?????? ????????????????? ???????? (????????, 1965). ??? ????
????????? ????????? ?????????? ????????
?????? ?????????? ????????????. ????????
???????????? ????????? ? ?????????? ????
????????? ?????? ? ???????? ????? ??????, ?
????????? ????? ??????????? ? ??????????
????? (Zettler, Daunys, 2007).
???? ?????? ?????? ? ???????????? ???????? ???????, ????????????? ? ?????????-
Corresponding author E-mail address: ruthenica@mail.ru
© Siberian Federal University. All rights reserved
# 160 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ??????? ??????, ????????????? ????????????? ? ????????????? ???????????????
???? ????????????? ?????????? ?????????
? ?????????? ???????? ????? ????? ??????,
????????????? ?? ?????????? ??????????????? ???????.
????????? ? ??????
?????????? ??? ???????????? ????????? ?????????????? ? ???????????? ?????,
????????? ? 2010 ?. ?? 11 ???????? (???. 1) ??
???????????? ?????????????????? ????????? (????????, 1984). ???????????? ?????
???? ??????? ? ??????? ??????? ? ????????? ???? 0,5 ??. ?????????????? ????? ???????? ??????? ????????? ????? ????????
0,25 ?2. ???????? ?? ?????? ????? ??????????? 70 %-? ???????.
????????????? ????? ????????? ??
????? ? ???????? ???????? ? ???????? ??????????????? ???????? (???????????? ? ??.,
2004; Kruglov, Starobogatov, 1993a, 1993b),
??? ??????????????? ?????????????? ?????????????? ??????? ????????? ???????
? ??????? (2005). ????? ????? ?????????
?????????? ???????????? ?? ????? ? ????????? 0,1 ??.
??? ??????????? ?????????????? ????? ???? ????? ?? ?????? ?????? ?????????????? ? ?????????? (1987, 1998). ???
????????????????? ?????????????? ????? ???????????? ?????? ?? ?? ???????????????
?? ????? ????????????? ? ?????????? (2004) ?
?????? (Glцer, 2002).
????????? ???????? ??????? Gastropoda ?
???????? ? ????? ?????? ?????? ?????????
?? ??????????? ??????? ????? ? ?????? ??
(Zettler, Daunys, 2007) ? ?????????????? ???????????? ????????????:
Ic
C
� 100% ,
A B
???. 1. ????? ???????????? ??????? ? ?????????? ????? ????????? ??????. ??. ? 1: ???? ????????????;
??. ? 2: ???????-?????????? ?????; ??. ? 3: ???????-????????? ?????; ??. ? 4: ???? ??????; ??. ? 5:
???????? ?????, ???? ?????????; ??. ? 6: ????????, ???????-??????? ??????, ???? ?????????; ??. ? 7:
???????? ? ????????????? ????????? ?????, ????? ?????? ? ??????; ??. ? 8: ???????? ?????; ??. ? 9:
???????-?????????? ?????, ???? ?????????; ??. ? 10: ???????? ?????, ???? ?????????; ??. ? 11: ?????????????? ?????
# 161 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ??????? ??????, ????????????? ????????????? ? ????????????? ???????????????
??? ?, ? ? ?????????? ????? ????????????
???, ? ? ?????????? ????? ?????. ?????????? ?????? ???????? ???????? ?? ????????
??????? ????????? ?????????? ? ???????
????????? Primer 6 (PRIMER-E Ltd, UK) ?
????????????? ?? ????????? ??????? ???????? ????-???????. ????????????? ????????
????????? ????????? ?????? (Clarke et al.,
2006).
?????????? ? ??????????
?????????? ???????? ????????? ?????? ???????????? 38 ??????, ?? ???????
??????????? ????? ????????? ? ?????????
Pulmonata (28 ?????) ? ?????? Lymnaeiformes.
???????? Pectinibranchia ??????????? 10 ?????, ????????????? ? ???????: Rissoiformes,
Neritopsiformes ? Vivipariformes. ????????
??????? ???????? ?????? ? ??????? ????????? ??????: ???????? (B1), ???????-???????
(B2), ???????-????????? (B3), ????????????????? (B4), ???????? ? ?????????????
????????? (B5); ? ????? ?????? ???????, ??????????? ? ?????? ?????? ??????????????,
?????????????? ?????? ??????????? ??????, ??? ????????, Phragmites (B6), ????? Typha
(B7), ????? Potamogeton (B8), ???????????
Ceratophyllum (B9).
????? Rissoiformes ??????????? ?? ????????? ????????? ????? ?????? ?????????
Bithyniidae, ?????????????? ? ???? ?????:
Bithynia tentaculata (L., 1758), B. producta
Moquin-Tandon, 1855 ? Codiella leachii
(Sheppard, 1823). ???????? ??????????????
?? ????????? ???????? B. tentaculata, ??????? ??????????????? ?? ???? ????? ???????
? ?????????? (????????, ?????). ??????? ??????????? ?????????? 12-20 ???/?2. ?? ??????
Neritopsiformes ?? ???????? ?????? ??????????? ???? ?????, ??????????? ? ?????????
Valvatidae ? ?????????????? ????? ??????:
Borysthenia, Cincinna ? Valvata. ??????????
?????????? ?? ???????-???????, ????????
???????, ?????????. ??? ??? ????????????????? ??? ??????, ?? ??????????? C. piscinalis
(Mьller, 1774), ??????? ??????????? ??
???????-??????? ? ???????-????????? ???????, ??????????? ????????? 239 ???/?2.
????? Vivipariformes ??????????? ?????
?????? ? Contectiana contecta (Millet, 1813) ?
Viviparus viviparus (L., 1758). ?????? ??? ??????? ?? ???????-????????? ??????, ??????
??? ?????? ?????? ?? ???????-??????????
??????.
?????????? ??????? ?????????? ??????????????? ????? Lymnaeiformes, ????????????? ? ????? ?????? 28 ????? ???????
????????: Acroloxidae, Lymnaeidae, Physidae
? Planorbidae. ??? ???????? ???? ??????????? ??? ? ???????? ???????? ???????? ????????, ??? ? ?????? ?????? ??????????????.
????????? Acroloxidae ??????????? ??????
???? ???? Acroloxus, ????????? ?? ????????,
???????-????????????, ???????-??????????
??????, ????????? ? ??????. ?????????
Lymnaeidae ???????????? ????? Lymnaea, ????????????? 12 ?????. ??? ????, L. balthica
(L., 1758), L. stagnalis (L., 1758) ? L. fragilis
(L., 1758), ?????? ?????????????? ? ??????.
???????? L. balthica ????????? ?? ??????
????????, ???????????? ?? ???????, ????????????? ?????? ?? ???????-??????????
??????. ????? ??????? ?????????? ??????????? ???? ???????????????? ?? ?????????
(364 ???/?2) ? ?????? (136 ???/?2). ?????????
L. fragilis ? L. stagnalis ?????????? ?????
?? ???? ???????, ?? ?????? ?????????????? ?
?????? ?? ?????? ? ??????, ?????????? ??????? ???? ?????.
??
?????????
Physidae
??????
Lymnaeiformes ???????? ??? ????: Costatella
acuta (Draparnaud, 1805), Physa bulla (Mьller,
1774) ? P. fontinalis (L., 1758). ????????? ???
?????????? ? ?????? ??????????????, ?????-
# 162 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ??????? ??????, ????????????? ????????????? ? ????????????? ???????????????
?? ????????? ?? ??????, ??????, ??? ?????????
??????????? 32 ???/?2 (P. fontinalis) ? 44 ???/?2
(P. bulla).
??????? ?????? ????????? Planorbidae
??????? ?????????????? ??????? ?????: Anisus, Choanomphalus, Planorbarius
? Planorbis. ??? Anisus ? ????? ????????? ?????? ??????????? ????? ??????, ???
Planorbarius ? ????? ????? P. corneus (L.,
1758), ???????? ? ???????????????? ?? ????
???????? ??????, ????? ???????-???????????
??????. ?? ???????? ?????? ??? ???????? ???????????? (??????????? 18 ???/?2).
??? Planorbis ?????????? ??? ????:
P. carinatus Mьller, 1774 ? P. planorbis (L., 1758).
??????? P. carinatus ? ?????? ???, ??????? ?
?????? ????????. ?????????????????? ????? ????, ??? A. vorticulus (Troschel, 1843) ? C.
riparius (Westerlund, 1865), ????????, ??????????? ????????? ????????? ?? ?????????? ??? ?????? (??????????, 2010). ???????
P. planorbis ?????? ?????????????, ????????
??????????? ?? ???????????? (88 ???/?2), ??????????? ?? ?????? ? ?????????. ?????????? ????????? ? ??????? ??????? ????????
??????? ?????? ????????? ????? Acroloxus,
Anisus, Bithynia, Physa ??? ??????? ????????? ????????????? ? ??????????? ?????.
??????? ?????? ????????? ? ?? ????????????? ? ???????? ???????? ? ????. 1.
???????????? ???? ????????? ???????? ? ???? ???????? ??????????? ???????: ???????? (51 %), ??????????? (34 %) ?
???????? (15 %). ????? ???????????? ?????????? ????????? ????? ??? ?????????????????????? ? ????????? ??????? ?????
(?????-????????, 1987). ??????? ? ??????????? Gastropoda ????????? ?????? ????? ??????? ?????????? ? ????????????? ? ??
??????? ??????? ??? ???????????? ????. ???????????? ????? ???????????? ? ?????????? ?????????? ? ???????? ??? Bithyniidae,
Valvatidae, Viviparidae. ??????? ???????????? ???????? ????? ? ???????????? ???????
???????? ? ?????????? ????? ?????????????? ?????????? ??? ?????? ??????? ?????
Anisus ? Choanomphalus (?????-???????? ?
??., 1998). ? ????????? ????????? ??????????? ???????? ????????? ????? Acroloxus,
Lymnaea, Physa, Planorbis, ?????????? ?????
??????????????. ???????? ????????? ???? ?
????? ?????? ????????, ?????????. ???????
???????? Planorbarius ? Lymnaea ?????????? ???? ??? ?????????????, ??? ? ?????????
?????????????, ????????? ????? ??????????
?? 20 % ?????? ???????? ????? (?????????????, 1987). ??? ???????? ??????? ?
??????????? ????????, ??? ??????? ? ????????? ?? ??????????? ??????????????, ???????, ??? ????????, ???????? ?????? ? ???????
????? (?????????, ??????, 1996). ????????,
?????? ???? ????????? ??????? ? ????????
?? ?????????? ??????? ????, ??? ? ????????
???????? ???????? ??????. ????????????
?????????????? ???????? Gastropoda ? ???????? ?????? ??????????? ????????????? ?????? ????????? ? ?????????? ???????????
??? ??????????-??????????? ??????????????? ???????? ??????? ???????? ??????????
????????? (Glцer, 2002).
?????????? ??????????? ????????? ????????? ???????????????? ?? ???????-???????
(309,5 ???/?2), ???????-????????? ???????
(308,0 ???/?2), ? ????? ?? ?????????? (?? 200,1
?? 636,5 ???/?2) (????. 2). ???????? ?????? ????
???????? ?????, ?????????? ?????? ?????????????? (?????, ?????), ? ????? ???????????????? ? ???????? ? ?????????????
????????? ??????. ??? ???? ?????? ????????
?????????? ??????? ???? ?????????, ?? ????
? ????? ???????? ???????? ?? 60 %. ????? ????????? ?? ?????? ??????, ?????? ? ????????
? ????????????? ????????? ?????? ? ???????? ??????????? L. stagnalis (?? ?????? ? ??
# 163 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
??????? 1. ??????? ?????? ????????? ? ????????????? ????? ? ?????????? ???????? ????????? ??????
????
1
????? Rissoiformes
????????? Bithyniidae
B. tentaculata
???????
B1
B2
B3
B4
B5
B6
B7
B8
B9
2
3
4
5
6
7
8
9
10
++
+
+
+
+
+
+
B. producta
+
C. leachii
????? Neritopsiformes
????????? Valvatidae
B. naticina (Menke, 1846)
+
C. antiqua (Morris, 1838)
+
C. piscinalis
+
+
+++
+
+++
++
+
??
??
11
12
?
??
?
?
?
?
?
??
?
?
?
??
C. pulchella (Studer, 1820)
+
?
??
V. cristata Mьller, 1774
+
?
??
s
?
?
s
?
??
????? Vivipariformes
????????? Viviparidae
C. contecta
s
s
++
+++
V. viviparus
????? Lymnaeiformes
????????? Acroloxide
A. lacustris (L., 1758)
s
+
+
A. oblongus (Lightfoot, 1786)*
+
A. okaensis Kruglov et
Starobogatov, 1991*
A. rossicus Kruglov et
Starobogatov, 1991*
????????? Lymnaeidae
+
L. auricularia (L., 1758)
++
+
L. fragilis
+
+
L. fusca (C. Preiffer, 1821)
+
+
L. intermedia Lamarck, 1822
+
L. monnardi (Hartmann, 1841)
+
L. balthica
+
?
?
+
+
?
?
+
+
++
++
?
??
+++
+++
++
+++
?
??
++
++
+++
++
+
++
+
L. ?vat? (Draparnaud, 1805)
+
+
+
+
+
+
+
L. peregra (Mьller, 1774)
L. stagnalis
+
L. tumida (Held, 1836)*
+
+
+
??
??
+
+
L. corvus (Gmelin, 1791)
L. palustris (Mьller, 1774)
++
+
?
?
++
+
+++
++
?
?
?
??
?
?
?
??
?
?
?
??
?
??
?
?
?
??
?
??
????????? Physidae
C. acuta
+
P. bulla
P. fontinalis
+
?
??
+
+
+
?
??
+
++
++
?
?
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ??????? ??????, ????????????? ????????????? ? ????????????? ???????????????
??????????? ????. 1
1
2
3
4
5
6
7
8
9
+
+
+
++
+
+
10
11
12
?
??
?
?
????????? Planorbidae
+
A. acronicus (Fйr., 1807)
A. albus (Mьller, 1774)
+
+
A. contortus (L., 1758)
+
+
A. vortex (L., 1758)
+
+
+
+
+
A. vorticulus
C. riparius
+
P. corneus
+++
+
+
+
+
P. carinatus
+
P. planorbis
+
+
+
+
+
+
+
+
+
++
?
??
?
?
?
?
?
?
?
??
?
??
?
??
??????????. ??????? B1-B9 (??. ??????????? ? ??????): +++ ? ???????????? ???, ++ ? ??????????????? ???,
+ ? ?????? ????; s ? ??????? ? ???? ?????? ???????; * ? ??? ??????? ??????? ? ??????????????? ???????. ?? ?
??????????? ?????????????? ????: ? ? ??????????, ? ? ???????, ? ? ???????; ?? ? ????????????????? ???????????????:
?? ? ??????????, ?? ? ???????????, ?? ? ??????????-???????????????? ???????, ? ? ??????????? ???????, ?? ?
?????-?????????? ???????.
??????? 2. ????????? ????????? ????????? ? ?????????? ???????? ????????? ??????
??????
?????
?????
???????? ?????
???????????? ????
???????????,
???/?2
????????,
?/?2
54,5 ± 10,2
14,6 ± 5,1
???????-??????? ?????
17
Lymnaea
Planorbarius
Cincinna
309,5 ± 122,5
17,2 ± 4,7
???????-????????? ?????
14
Cincinna
308,0 ± 135,8
28,6 ± 7,4
???????-?????????? ?????
4
Viviparus
17,1 ± 6,0
9,9 ± 3,2
49,2 ± 10,0
27,3 ± 5,3
636,5 ± 160,3
16,1 ± 6,3
16
???????? ? ?????????????
?????????
????????
15
?????
13
Lymnaea
249,1 ±85,9
45,6 ± 13,1
?????
10
Lymnaea
420,8 ± 90,1
30,6 ± 12,9
6
Lymnaea
Planorbis
200,1 ± 68,4
11,9 ± 8,5
Lymnaea
16
???????????
Lymnaea
15,4 ?/?2), L. auricularia (?? ?????? ? ?? 6,0 ?/?2),
??????? ???????? ????? ????? ???????? ????????????? ??????. ?? ???????? ??????
?????????? ????????? ??????? P. corneus
(9,6 ?/?2). ?? ???????-??????? ?????? ????????
C. piscinalis ????????? 7,8 ?/?2.
???????? ???????? ??????? ????????? ????? ????????? ???????? ????????
?? ???. 2. ??? ?????, ?????? ??????? ????-
???? ????? ???????, ??? ???????? ? ????????????? ????????? ?????, ????? ?????? ?
?????? (??????????? ???????? ????? 50 %,
F (4,28) = 2,67, df = 23), ?????????? ??????????
????? Acroloxus, Anisus, Costatella, Lymnaea,
Physa, Planorbarius. ????????????? ??????? ???????? ? ???? ???? ???????? ??????????? ????? Lymnaea (L. balthica, L. stagnalis,
L. auricularia).
# 165 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ??????? ??????, ????????????? ????????????? ? ????????????? ???????????????
???. 2. ???????????? ???????? ???????? ??????? ?????????? ????????? ?????????? ????????
????????? ?????? ?? ????????? ??????? ????-???????
?????? ?????? ?????????? ???????
?????????, ???????-???????? ? ????????????????? ??????, ?????????? ??????????????? Anisus, Bithynia, Cincinna, Lymnaea,
Planorbarius, Valvata (Ic = 63 %, F (4,17) =
0,04, df = 31). ???????????? ???????
???????? ?? ???????-??????? ? ???????????????? ?????? ??????????? ?????????? ????? Cincinna (C. piscinalis) ? Lymnaea
(L. balthica); ?? ???????? ?????? ? ?????????? ????? Planorbarius (P. corneus) ?
Lymnaea (L. balthica). ???????-??????????
????? ???????? ???????? ??????? ???????? Viviparus (V. viviparus) ? Lymnaea
(L. monnardi) ? ? ???? ????? ?????????????
????????????? ?????????? ??????????? ??
???????? ??????? ?????????? ??? ????????? ? ????????? ? ?????????? ?????????
(Ic = 16 %, F (4,08) = 13,31, df = 39).
? ?????????? ????? ????????? ??????, ????????? ??????????? ??????? ? ??????
?????????????? ?????? ??????????????, ??????? ????? ?????????????, ????????????? ???
?????????? ?????????. ???????????? ?????
???????? ?????? ????????? ????? ?????????
??????? ? ???????????? ???????? ????????
? ????????????? ? ????? ???????????? ?????,
? ??????? ???????????? ???????? ?????????
????????? (90 % ?????). ???????? ????????
?????? ? ?????? ??????????? ???????? ?? ???????? ??????????? ??????????. ????????????????? ?????, ?? ??????? ??????? ??????
?????? ????, ?? ???????? ???????? ?????? ?
????????? ???????? ? ???????? ??????????????. ????????, ??????? L. monnardi ???????? ?????? ???????? ????? ? ???????
??????, ?????????????? ?? ???? ??????????
?????????? ??????? ????????, ??? ?????????
??????? ????????????? ? ?????????? ??????????, ? ?????????? ????? ???????? ???????????? ????????? ????????????? ??????.
?????? ?????????? ??????????????? ??? ?????????????? ? ???????? ? ???????? ????????
? ??? ?????? ????? ????????? (??????, 2011).
????????? ????? ?????? ?? ???????? ??????? ????????? ? ??????? ?? ?????, ?????????
? ???????? ??? ????? (Zetter, Daunys, 2007), ???????? ???????? ???????? ??????? ?? ??????
# 166 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ??????? ??????, ????????????? ????????????? ? ????????????? ???????????????
70 %. ? ????????????????? ????????? ?????
????????? ???????????? ?? 8 % ???????????????, ?? 40 % ????????????????, ?? 10 %
??????????-?????????????????, ?? 40 % ???????????? ? ?? 2 % ?????-??????????? ??????
?????????. ?????? ???? ????????? ? A. albus,
A. vorticulus, C. riparius ? P. Carinatus ? ???????? ? ??????? ????? ??????????????? ??????? (??????????, 2010).
??????????
? ?????????? ???????????? ???????????????? 38 ????? ?????????? ?????????,
?? ??????? ???? ? C. piscinalis, L. balthica,
L. stagnalis, P. corneus, V. Viviparous ? ???????????? ? ????????? ?????????? ????????
????????? ??????. ??????? L. balthica ???????? ??????? ???, ????????????? ?? ????
????? ???????, ????? ???????-???????????,
? ????? ? ???????? ?????????? ??????????.
???????? ????????? Gastropoda ?????????
?? ??????????????, ???????????? ??????????? ?????????? ????????, ??????????
??????? ?????????? ?????? ??????????????.
?????? ????? ????? ?????????? ?????????
?????????? ?? ???????? ? ???????-???????
? ???????? ???????, ? ????? ? ?????? ????????? ? ??????. ? ???????? ?????????? ??????????? ????????? ????????? ????? ????????? ???? ????? ???????, ??? ?? ????????
????????? ??????. ?????????? ??????? ??????? ?????????? ????? ????? ?????????
??????, ????????????? ??? ???????????????
???????, ???????????? ??????????????
???????? ??????????? ????????????? ?????? ????????? ? ????????????? ???????????
??? ??????????-??????????? ??????????????? ???????? ??????? ???????? ?????????.
?????????????
???????????? ????????? ??? ????????? ????????? ????????? ????? ?????????? ????? (Deutsche Bundesstiftung Umwelt),
?????? ? 30011/314. ????? ???????? ????????????? ?. ?????? (??????????? ?. ???????,
????????) ?? ?????? ? ?????????? ??????????????, ? ????? ?.?. ??????? (???, ?. ???????????) ?? ?????? ? ????? ?????????.
?????? ??????????
???????? ?.?. (1965) ?????? ????????? ??????. ?: ???????????? ? ???????? ? ?????????? ???????: ????? ??????????. 14: 19-39.
?????? ?.?., ?????? ?.?. (2005) ??????? ????????? ?????? ? ???????????? ?????. ?.:
??? Scientific Press Ltd., 627 ?.
?????? ?.?. (2011) ??????????? ??????????????? ??????????????? ???????? ? ?????????
???? Lymnaeidae (Gastropoda, Pulmonata) ? ????????? ??????. ???????? ???. ???. ????. 4: 477483.
???????? ?.?. (1984) ?????? ????? ? ????????? ?????????. ?: ???????????? ???????????? ?? ????? ? ????????? ?????????? ??? ?????????????????? ????????????? ?? ???????
????????. ?.: ????????, ?. 3-8.
???????????? ?.?., ??????? ?.?., ????????? ?.?., ?????? ?.?. (2004) ????????. ?: ???????????? ???????????? ?????????????? ?????? ? ???????????? ??????????. ?. 6. ????????,
?????????, ????????. ???: ?????, ?. 6-496.
?????????? ?.?. (2010) ?????????? ????????. ?: ??????? ????? ??????????????? ???????. ???????????: ???-?? ??? ??. ?. ?????, ?. 89-94.
?????-???????? ?.?. (1987) ?????????? ?????? ?????????. ?.: ?????, 145 ?.
# 167 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ??????????. ??????? ??????, ????????????? ????????????? ? ????????????? ???????????????
?????-???????? ?.?., ?????????? ?.?., ???????? ?.?. (1998) ?????? ? ???????????? ????
? ??????? ? ???????????? ????????????? ?????????. ????. ????. 77 (3): 270-277.
????????? ?.?., ?????? ?.?. (1996) ????????????? ??????????? ???????????? Gastropoda
?????? ??????????? ??????: ??????? ???? ??????? ? ?????????. ????. ????. 75 (5): 664-676.
Clarke K., Somerfield P., Chapman M. (2006) On resemblance measures for ecological studies,
including taxonomic dissimilarities and a zero-adjusted Bray-Curtis coefficient for denuded
assemblages. J. Exp. Mar. Biol. Ecol. 330: 55-80.
Geyer D. (1927) Unsere Land- und Sьsswasser-Mollusken: Einfьhrung in die Molluskenfauna
Deutschlands. Aufl., XI. Stuttgart, 224 p.
Glцer P. (2002) Die SьЯwassergastropoden Nord- und Mitteleuropas. In: Die Tierwelt Deutschlands.
73: p. 1-327.
Hilbert R. (1908) Weitere Beitrдge zur Preussischen Molluskenfauna. In: Schriften der Kцniglichen
physikalisch-цkonomischen Gesellschaft zu Kцnisberg, p. 155-166.
Kruglov N.D., Starobogatov Ya.I. (1993a) Annotated and illustrated catalogue of species of the
family Lymnaeidae (Gastropoda Pulmonata Lymnaeiformes) of Palaearctic and adjacent river drainage
areas. Pt. 1. Ruthenica. 3 (1): 65-92.
Kruglov N.D., Starobogatov Ya.I. (1993b) Annotated and illustrated catalogue of species of the
family Lymnaeidae (Gastropoda Pulmonata Lymnaeiformes) of Palaearctic and adjacent river drainage
areas. Pt. 2. Ruthenica. 3 (2): 161-180.
Szidat L. (1926) Beitrдge zur Faunistik und Biologie des Kurischen Haffs. In: Schriften der
Kцniglichen physikalisch-цkonomischen Gesellschaft zu Kцnisberg, p. 5-31.
Zettler M., Daunys D. (2007) Long-term macrozoobenthos changes in a shallow boreal lagoon:
Comparison of a recent biodiversity inventory with historical data. Limnologia. 37: 170-185.
Species Composition, Distribution in Biotopes
and Ecological Characteristics of Gastropod Mollusks
of the Curonian Lagoon Coastal Waters of the Baltic Sea
Dmitry P. Filippenko
Kaliningrad State Technical University
1 Sovetsky av., Kaliningrad, 236000, Russia
The research of the gastropod mollusks fauna of the littoral biotopes of the Curonian Lagoon of the
Baltic Sea was carried out. Gastropods, as has been estimated, are presented by 38 species, 10 of them
belong to Prosobranchia and 28 species belong to Pulmonata. The majority of littoral biotopes created
in shallow water conditions and those sheltered by aquatic plants are more similar in gastropod species
distribution and colonization. The species distribution and quantitative characteristics of the fauna on
the open bottom and in the macrophytes are revealed.
Keywords: malakofauna, gastropods, Curonian Lagoon, Baltic Sea.
# 168 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 169-183
~~~
??? 597. 0/5. 11
???????? ????????? (Comephoridae)
? ?? ???? ? ???????????? ??????????????
????? ??????
?.?. ???????*
???????? ??????? ???????? ? ????????
??. ?.?. ????????? ???,
?????? 117071, ??????, ????????? ??., 33 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
?????? ???????? ?????? ?? ???????? ???? ????? ??????????? ????????? (Comephoridae),
??????? ???????????, ????????????? ????????, ????????, ???????, ???????????? ????????.
?????????? ???? ??????? ??????????? ????????? ??????? ?????? ????????? ? ??????????
???? ??????????? ??? ? ???????? ?? ?????????????? ????????. ?????? ??????? ?????? ???????
???????? ? ?????????? ???? ??? ? ???????????? ????, ? ??????? ??????? ????? ? ??????????????? ???????, ?? ?????????? ???????, ? ????? ? ????????? ???????????? ?? ????.
???? ??????????????? ?????? ?????? ?????????????? ????????? ??????? ? ????????????
????????? ????????? ????? ??????.
???????? ?????: ??????, ?????????, ????????????, ?????????????? ????????, ??????
?????????, ??????? ????????, ????????? ????, ??????? ??????????? ????- ? ????????????.
? ??????? ??????????? ?????????
??????????? ?????????? ??? ???????
Comephorus baicalensis (Pallas, 1776) ? ????? Comephorus dybowski (Korotneff, 1905)
????????? ?? ????????????? ???????????
??????? ????? ?????? ?? ?? ??????? ???????????. ???????? ???? ?????????? ??????
????????? ??????? ???????? ????????????
? ?????????? ?? 36 ?? 150 ???. ? (???????,
1964; ????????, 1977; ????????, 1983). ???
???? ?????? ???? ???????? ????????? ?????
*
1
????? ? ?????????????-??????????? ???????
????????????? ??????????? ?? 0,9 ?? 9,1 ????
(??????? 5,0) ??????. ????????? ?????????
???????? ???????????? (??????????????)
????????. ???? ??? ?????????? ?? ???????
????? ????? ?? ???, ??? ??? ????????, ????? ?
????????? ??????????, ? ????? ??????????? ? ???????????, ??? ?????????? ?????????
??????? ???????????? ?????? (?????) Pusa
sibirica (Gmelin, 1788), ? ??????????? ??????
???????? ?????? ????? 3 ?? ???? ???. ? ???-
Corresponding author E-mail address: gerarchernyaev@mail.ru
© Siberian Federal University. All rights reserved
# 169 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
????? ?????????????? ????????, ?????????
?? ????????? ? ???????????? ?????? ? 1620
? (??????? ??????? ??????? ? 1500 ?), ????????? ?? ??????????? ?????? ????, ? ????? ?
????????? ??????? ? ??????? ???? ??? ?????
???????????? ?????????? ?????? ?????????.
???????? ?? ??? ????????? ???????? ???
???????? ??? ????????? ? ?? ????, ? ?????
???????? ? ???? ? ?????????? ??????????? ?????, «???????» ?????????? ?????. ???
??????? ?????????? ????????? ???? ? ??????? ???????, ???????????? 234,7 ? ???, ? ????????????? ????? ????? ?? ???? ????????
?? ??? ???????? ?????????????? ????????
??????? ??????????? ? ???????????? ????????? ????????? ? ?????????? ????? 21 ?/?? ?
??? (?????????, 1971). ?????????? ???????
????? ?????? ????????? ? ?? ????????? ?
????????????? ???? ????? ????????? ?????????????? ???????? ???????? ????? ? ?????????????? ???????? ???????????? ??????
?????????.
??????? ??????? ????????????
???????? ?????????
?????????????? ????? ????? ???????????? ???? ???????? ?????? ?? ????? ?????
? ???????? ????????????-??????????? ???????? ??????????? ???: ??????? ????????? ? C. baicalensis ? ????? ????????? ? ?.
dybowski. ?? ??????????? ????? ????????
???????? ?? ???????????? ????? ???????
????????, ????? ??????? ???????? ?????????, ??????????????? ???????????? ? 1950 ?.
???????????? ????????? ????????? ???????
? ?????? ? 1965 ?? 1971 ?. ?? ???? ???????????????? ????????? ?????????? ????????? ??
???? ? ???. ????????????? (?????????). ????????? ??????????? ?? ???????? ???? ?????
?????? ? ????? ??????-??????????????????
????? (?????? ??? ?????????????? 550 ?)
«?.?. ?????????». ???????? ?????????? ?
????? ??? ? ????????? ?????????, ??????????? ?? ????? ????????? ????? ?? ????
???????? ??????????? ??? ???? ?? ????????
??????????? ???????, ? ??????????? ? ?????
??????? ??????????????? ????? ??????????.
??? ????????????? ?? ???????? 50, 150, 350,
750, 1100 ? (??? ????????????? ?????????)
??????? ??????? ? ?????? ????? ? ??????? 30
??? ??? ???????? ????? 5,4 ??/? (3 ????). ????? ???????? ??? ?????? ???????????????
????????????? ????? ??????? ???????-?????
? ???????? ????????? 10 ??. ?, ?????? ??????? ????? 6 ?, ? ????? 6 ??. ?????? ???
???????? ???????????? ??????? ?????????
???? ???????????? ????? 600 ???????? (300
?????). ?? ????? ??????? ??????? ???????
???? ????????? ? ?????? ?? ?????? ????? ?
???????? ??????????? ???? ?????? ??????????, ? ???????? ?? ?????? ????????. ?????,
?????? ? ??????????? ?????? ????, ?????
?????? ????????? ?????????? ?????, ????
????????? ????????? ????????????? ????????? ??????. ???? ? ????? 20 ?? ???????
?????? ?????? ???????????????? ?????????
?? ??????? 150 ? 250 ?.
????????? ? 1960-? ??. ???????? ?? ???????????? ??????????? ? ??????????????
???????? ????????? ???? ????? ?????????
? ????? ????? ??????????? ???????????
??? ??????????? ? ??????? ?????? (???????, 1971, 1974; Tcherniaev, 1971, 1976). ??????????? ???????????? ?????? (???????, 1972;
????????, 1977; ??????? ? ??., 1979; ????????,
1978, 1980, 1982; ???????, ????????, 1983; ????????, 1983; ????????, 1983) ???????????
????????? ????????????? ? ???????? ????
????????? ???????, ????????? ????????? ??????? ????????? ?? ???? ????????? ????????? ? ?????????? ??. ?????? ? ??????? ?????
??????.
?? ????????? ??????????? ? ???????????? ?????? ?????????? ????????? ????
# 170 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
??????? ? ????? ????????? ? ??????? ????????????? ?????? ???????. ?????????
?????? ?? ???????? ??????? ????????????
??????-?????????? ??. ??????, ? ?????????
??????????? ???????????? ?????????? ?????????? ???????????, ??????????? ????????????? ?????? ? ?? ??????????????. ??????????? ?????? ???????? ????? ???????
??????????? ????????? ?? ?????????????
????? ??????.
??????????? ????????
? ?????? ????? ?????????
???????? ??????????? ???????????
??? (????? ???????????? ??? ??????, ?. 2;
2002), ??????? ? C. baicalensis ? ????? ?
C. dybowski ????????? ???????? ? ??????
Scorpaeniformes, ????????? Cottoidei, ????????? Comephoridae, ??????????? ????? ????
??? Comephorus c ????? ?????? (????????,
2002; ????????, 2007).
??????? ????????? ? ??????? ??. ??????,
???????? ??????????? ????? (??????, 1955),
?????????? ? ???????? ????????????? ??????? ?????, ???????? ??? ????? ??? ???????,
????? ???????????? ????? ?????, ????????
???????? ???????????? (??????????????)
????????: ???? ?????????? ?? ???, ?????
??????????? ? ???????????.
???????????? ???????? ????? ????????? ??? ????? 210 ??, ? ????? ? 145 ??. ??????? ???????? ? ?????? ????????? ??? ????? ???? 105 ?? ? ???????? 3+, ? ? ????? ? ???
????? 118 ?? ? ???????? 3+ (???????, 1972;
????????, 1977; ????????, 2002). ??????? ????????? ? ??????? ????????? ???? ???????
? ???? ???????????????? ?????????????? ????? ? ??????, ? ????? ? ?????? ?? ???? ? ???????. ? ????? ????? ???? ?????????? ??????? ????? ??????????? ?? ????? ??????, ??
? ?????? ??? ??????? ?????? ?? ?????????
? ????????? ??????, ?????? ???????? ? ??-
???????? ????? ? ??????, ???, ????????, ????? ???????????? ?????????????? ????????.
????????, ??? ?????? ?????? ????????????
???? 100-???????? ??????? ????? ??? ?????????? ? ?????????? ? ??????? ??????? ?????
?????????? ?????????? ???????????, ??????????? ???????? ????????????? ?????? ? ??????? ???? ????? (???????, 1974) ? ?????????
???????? ??????????????, ?????????, ????????? ? ????????? ??????????. ??????????? ????? ? ??????? ?????????: 83,3 % ?????
? 16,7 % ?????? (????????, 1977).
??? ? ???? ??????????????? ??? ???????, ? ?????? ????????? ? ?????? ???????
??????????? ???????????? ??????????? ??????? ? ???????????? ???????? ??????. ???
???????? ???? ?????????? (???????, 1971,
1974) ? ??????????? ???????????? (?????????, ?????????, 1984), ?????????? ???????
???? ?????? ???? ?????????-????????????,
???????????? ????? ? ?????????? ? ???????.
????? ??????? ????????? ??????????
?????????? ? ?????????? ? ????? ???????,
?.?. ?? ?????????? ????????? ? ??? ????? ????? ?????? ???????? (???????, 1974). ?????
???????? ?????????? ???????? ? ?????????
??????????? arteria mesenterica ?????? ?????
?????? ??????????? ????????, ???????????
??????? ?? ??????? ???? ?????. ??????????
?????????? ? ????? ?? ????, ? ????? ?????????????? ???? ????? ? ??????? ?????? ?????????? ???????????? ????????, ???? ????????? ?? ??????????? ???????????, ? ? ???????
200-???????? ???? ???? ????? ? ??????????
?? ?????? ?????? ???????? ?? ???????????.
??????? ?????? ????????? ? ????? ????? ???????????? ?????????????? ????????? ???????? ? ???????????? ????? ??????????????
????? ?????? ?????. ?? ????? ???????????
????? ??????? ????????? ?? ???????? ? ?????????? ?? ???? ??????????? ??????? ??-
# 171 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
?????. ????????? ????? ? ??????????????
?????????? ??????????? ??????? ???. ????????????? ???????? ???????????? ?????????????? 90 ????? ??? ??????? ???????????
???? ??????? +3,2 °? (???????, 1988). ??????????? ??????? ??????? ????????? ??????? ?????? 9,4 ?? ?????????? ? ????? ????
?? ??????? ?? ??? ??????? (25-100 ?), ??? ??????????????? ??????? ?? ??????? (???????????).
??????? ???????????? ??????? ????????? ? ??????????? ?? ????? ???? (150 ? 250
??) ? ????? ????? (33,0 ? 105,2 ?) ????????? ??
1200 ±15 ?? 2675±26 ??????? (???????, 1964;
???????, 1974; ????????, 1977; ????????,
2002). ??????? ???????? ??????? ?? ??????
?????? ?? ???? ???????? ???????? ????????????? c 1,1 ?? ????? ?????????????? ?? 1,31,5 ??. ? ?????????? ????? ????? ???????????? ? ?? ????? ?????????? ??????? ???????
????????????? ?? 1,6-2,4 ?? ?? ???? ??????????????? ?????????, ??? ??????????? ??????? ??????? ???? ???. ?? ?????? ????????
(1964), 89 % ????? ??????? ????????? ?????
??????????? ??????? ???????? ? ?? ???? ??????? ?????????, ???????????? ?? 44,3 %
?? ????? ???? (????????, 1977), ?????????
? ??????????? ? ?????????? ??????? ???? ?
????. ?????????? ? ????? ??????????? ????????, ? ??? ??????????????? ??????????? ?
??????? ????? ? ??????????? ????? ?? ????????? ? ????? ??????? ????????, ????????
??????????????? ????????? ???????????
????? ??????????? ? ?????????????????
????????? ???????.
????? ????????? ????? ???????? ????????????, ??????????? ?????, ?????????
??????????? ??? ????????? ????? ? ??????????? ??????????? ???????????? ??????? ????????. ???? ???, ???????? ?????????
????????????? ???????????? ?????????????????? ????????????????? ???? ?????????
b-????, ????????? ?? ????????? ????? ??????? ????????? ??????? 45?75 ???. ??? ?????
? ?????????? ?????? ? ??????? ???????????
(????????, 2008). ???? ??????????? ????????
?????????? ? ????? ????????? ?? ???????????: ???????????? ?????? ?????? ? 135 ??,
?????? 80-90 ??, ????? ? 158 ??, ?????? ?
100-110 ?? (??????, 1955). ??????? ???????? ?
????? ????????? ????????? ??? ????? ?????
95 ?? ? ???????? 3+, ? ?????? ? ??? ????? 75
?? ? ???????? 3+ (????????, 1983; ????????,
2002). ??????????? ????? ? ????? ????????? 32,4 % ?????? ? 67,6 % ????? (????????,
1977). ??????? ???????????? ?????????? ??
670±3 ??????? ? ????? ?????? 95 ?? ? ?????? 5 ? ?? 2164±40 ??????? ? ????? ??????
155 ?? ? ?????? 32 ?. ?????????? ?????????????? ????? ????????? ?????????? ?????? ?
? ?????? ????, ????????? ? ???????? ?? ?????
????, ??????????? ???? ????? ? ??????.
?? ????? ??????????? ?? ????? ????????????? ????? (???????, 1971), ?????
?????????????? ???????? ????? ?????????
?????????? ?????????????? 100-120 ?????,
?? ?????? ????????? (1977) ? 90-100 ????,
? ??????????? ?? ??????????? ????????????? ??? ?????, ???????????? ? ?????? ?????? ?? 13 °?, ? ???? ??????? 200 ? ? ?? ???
??????????? ?????? ???? ????? 3,2 °?. ??
????? ???????????? ??????? ??????? ?????
????????? ?????????? 1,4 ??, ???, ??? ?? ???
? ? ????? ??????? ?????????, ???????? ?
????????????? ?????????? ???????? ??????, ?????? ?????????????? ?????? ????????
???????????? ???????????????? ????????? ????? ??????? ? ?????????? ?????????.
?????????? ? ????? ?? ??????? ???? ?????
??????? ?????? ????? 8,2 ?? ????? ????????? ?????????? ???? ????? ? ? ????????????, ?? ????? ?????????? ???????? ?????.
????? ?????? ??????? ????? ????? ?????
????????? ???????? ???????? ???????? ?, ?
# 172 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
??????? ?? ??????? ?????????, ??????????
?? ???, ??? ??? ???????? ?????? ?? ???? ??????????? ?????? ? ?????????? ?????? 8 %
(????????, 1977). ??????? ????? ?????????
???????? ????????? ?? ??????????? ?? ??????? 500 ? ? ?? ????????? ???????? ????????. ? ??????? ????????? ?? ???? ?? ?????,
????????????????? ???????????? ????????
? ??????????? ???????????? ??? ???????
?????????? ? ???????, ??? ? ? ???????? ???,
????? ?????????????? ???????? ???????????? ???????? (??????, 1955).
????????? ???????????? ??? ???? ????????? ?????????? ???????????? ??????????
?????????????? ??????, ??? ? ???????????
??????? ????? ??????????? ?? ?????????.
???, ? 1971 ?. ??????????? ????? ?????????
?????????? ? 32,0±4,7 ???? ???., ? ????????
? 66,5±8,6 ???. ????. ??????????? ???????
?????????, ???????????? ? ???? ?? 1971 ?.,
????????? 9,2 ± 1,1 ???? ???., ???????? 82,5 ±
12,2 ???. ? (????????, 1977).
????????? ???????????
?????????? ????
? ???????? ?????????
? ????????? ?????? ?????? ????? ???????? ???????? ?? ????? ???????????
???????? ????? ????? ?????????, ??? ?????????????????? ?????? ??????? ?????,
????? ???????? ? ???????? ?????? ??????
??????????????? ??????? (????????, 1980,
1982), ??????? ??????? ???????, ????????
? ????????? ?????????. ????? ???????? ????
????????? ?????????? ?????? ? ????????, ??????????? ?????? ??????. ?????? ?????? ?
?????, ? ?? ????? ? ??? ???? ?????? ??????.
???? ????????????? ??? ????????? ????????, ?????? ?????????? ?? ????????? ???????
???? ????? ?????????. ??????, ????????????? ???? ??????? ????? ? ??????????? ??
?????????? ? ???????? ???????? ????????.
??????? ??????? ????? ????? ?????????
????? ???????????? ????????: ?? ?????????? ?????????????????? ???? ? «??????????
???? ??????» ??? ????? ????????? ?? ?????
???????? ????????. ???????? ?????????
????????? ????? ????? ? ?? ??????????? ?
???????????? ??????????, ???????? ????
?????? ? ???????, ? ???? ???????? ????????? ????? ??????? (????????, 2002). ??????
????? ??? ? ????? ???????? ??? ?????????
????? ?????????, ???????, ??????????? ?????? ?????? ?? ? ????? ????????????? ?? ??????????? ???????????, ????????? ?? ?????
??????? ??????? ??? ?????????? ????????,
????????? ????????? ?????? ?????. ??? ????
????????? ????????? ????????? (? ?????????? ?? 12 ?? 16 ??.) ??????????????? ??????????? ????????? ???????. ??????????
???????? ????????? ??? ????????? ?????? ???????? ???????, ???????? ???????
???????????? ????? ????????, ????????????
???????????. ???????? ???????? ?????????
?????? ????? ??????? ??? ??????????? ?????????? ??????????????? ???????? ????????
?????????? ????????? (???????, 1963). ??? ??
????? ?????????? ????????? ??????? ? ?????????? ??????? ?????? ? ????????????? ???
???????????? ?????????????? ?????? ??????? ??????????? ???????? ????? ??????????
??????? ?????? ????????? ?????????? ???
??????? (11,7?11,9 ??/? ? ??????????? ? 9,9?
10,6 ??/? ? ???) ????????? ??????????? ????
?????????? (???????, 1988).
??????????????
???????? ????????? ? ?? ????
? ???????? ?????? ?????????
?????? ?????????????? ??????????
????????? ? ???????? ??????? ??????? ????????????? ????????? ????????? ? ????
???????? ???????????? (??????????????)
???????? (?????????, ?????????, 1929; ??-
# 173 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
???????? ?? ??? ??????-????????? ????????????? ?????? (??????????? ????????) ???
????, 1955; ???????, 1964). ???????? ??????
????????? (1977), ???????????? ????????
????? ????? ????????? ???????? ???????? ? ?????? ? ???????? ?????? (? ?????? ??
??????) ??? ?????????? ??????? ????? ????
????? ?? 0,3-1,5 °?. ????????, ??? ??????? ???
? ?????????????, ??? ? ? ???????? ???????????. ? ????? ????????? ? ??? ?????? ????????? ?????????? ???????? ???????? ???????. ????? ??????? ????????? ? ???? ??????
???? ?????????? ? ?????????? ? ?????? ?
??? ????? ? ???????? ??????? ?????????
????, ??? ? ?????? ??????? ????, ??????????? ??????? ? ?????? ????? ?????????. ?
????????? ? ??????? ????????? ???? ? ??????? ?????? ?????? ??????? ?? ????????????? ?????????? ????. ??? ????? ?????????
??? ????????? ?? ??????? 250-300 ?, ? ???
??????? ????????? ? 300-500 ?. ??????????
????????? ????????? (??? ?? ???????????,
??? ? ?? ????????) ??????????? ? ???????
?????? ????? ? 25-50-???????? ???? ????,
??? ?????? ?? ???????? ???????? ???????
????? (????????, 1977). ?? ?????? ??????
(1963), ???????? ???????????? ???????? ??-
???????????? ?????????. ?????????? ?????
????????, ??? ???? ??????????? ????? ?????
??? ?????????? ??????????? ? ??????????
?? ??? ??????? (?????? ? ??., 1979), ???????
? ???????????? ???????? ?????????, ???????
???????? ? ?????? ??????????.
????? ??????? ???????? ?????????
???????? ????????? ???????. ? ???????
????? ????????? ?????????? ?? ??? ?????
? ??????? ????????, ? ??? ??????????????? ??????????? ? ??????? ????? ??? ????????????? ????????? ? ??????????????
Macrohectopus branickii (Dyb.), ?????? ??????????? Cottocomephoridae ? ?????? ???????? Gammaridae (??????????-??????, 1971)
??? ????? ????????? ? ? ????????????????
???????? Amphipoda, ?????????????? ? ?????? ??????????? ??? ??????? (????????,
1977). ? ?????? ????????? ???????? ????????? ?, ????????? ????????????????????
????? ???, ??????????? ? ?????? ????? ?????
? ????????????? ????? ?????.
??? ???????????? ???????????????? ??-
???????, ?????? ?? ???????? ??????????,
??????? ????? ? ??????????? ???????? ????
????? ?? ???????? ? ??????? ????? ????? ??
???????, ?? ??????? ??? ?????????? ????? ?????????? ??? ?????.
??? ?????? ????????? ? ????????? ????????? ?????? ????? ??????????? ?????????????? ??????????? ???? ??????? ??????? ?
?????? ???????? ???????????? ?????????,
? ??????????? ?? ?????????????????? ??????? ??????? ????? ????????? ??????. ?
????? ???????? ??????? ???????????????
???????, ? ????? ? ???????-????????? ???????? ????? ???? ?????????? ????????????
???????????? ??????? ??????????? ???????. ???? ??????????? ??????? ????????????
?????????? ?????? ???????? ??????? ?????
??????? ??? ??????? ? 1977 ?. ?????????? ??
????????? «Pices» ????????? (?????? ? ??.,
1979), ??? ????????? ? ??????? ????? «????????» ??? ???????????? ??? ?????. ???
????? ??????????????? ?? ??????????? ?
?????? ????????????? ???? ??? ?????????
? ???????????? ??????????? ??? ????? ???,
??????? ????????, ????????? ?????, ????????
(Gammaridae) ? ???????? (Oligochaeta). ???
??????? ??? ??????? 194 ???? ??????, ????????????? ???????? ???????? ?? ???????? ?
??????? ?????????: ?? ??????? 256 ? ?? ????? ????????? ????? ? 27 ?/?2, ?? ??????? 875 ?
? ???????????????? ??? ? 38 ?/?2 ? ?? ???????
1202 ? ????? ? ???????????????? ??? ? 32 ?/?2
(????????, 2004). ???????? ?? ?????? ?????????? (1970), ???????? (2004) ????????????,
# 174 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
??? ????????? ?????????????? ???????????? ?????????? ???????????? ???????????
???????????????? ? ????????? ????, ?????????? ?????????? ?????????? ????????????? ????????. ???????? ???????? ??????????
1,84 ? 2,49 % ?? ????????-????? ????? ???
? ?????? ? ??????? ????? ??????????????.
? ??????? ????? ???, 0-7 ??, ???????? ????? ???????? ???????? Lamprodrillus inflatus,
??????????? 97 % ?? ??????????? ??????????. ??? ???? ????????? ? ???? 15-21 ??, ???
??????????? ????????? ??????? ?????????.
????? ??????????? Rhynchelmis brachycephala,
?????????? ????? ??????? ????????????.
?????? ? ????????????? ???????????? ????
?????? ?????????, ? ??????? ???????????
?? ????????? ???????????? ???????? ????????? ? ??????????-???????????? ? ???????
??????????, ? ???????? ?????????.
???????????? ??????? ? ????????? (1974)
?? ??????? ??? ? ????? ??????? ???????-
????? ?????? ????, ????? ???????? ?????
????????????????? ???????????. ??????
??????????? ???? ????? ? ??????? ?? ????,
????? ??????????????? ??? ????????????
???? ? ?? ??????? 50-70 ? ??? ????????????
????????? ???????? ?????????? ???????????
???????? ?????????? ?????????? (Diatomea),
??????? ??????? ???????? ? Melosira
baicalensis (C. Meyer) Wisl. ? ?????????? ?
Cyclotella baicalensis Skv., ?????????????
???????? ??????????? ????????????. ?????? ???????? ???????? ? ??????????????
?????????? ?? ?????? ?????? (????, ??????)
??? ???????? ????????????? ??? ?? 13-15 °?,
????? ? ?????????? ? ????????????? ?????
?????????? ????????? (Peridininiales), ??????? (Chlorophyta) ? ??????????? (Cyanophyta)
????????? (????????, 1957; ?????, 1962; ?????????, 1977).
????? ??????? ???????? ?????????? ?????????? ????????? ??????????? ? ??????-
???? ? ??? ??????, ????? ??????? ?????????????? ???????? ??? ??????? ???????? ?????
???????? ??? ?? ??????????? ?? ????????, ?
???????? ????????????? ???? ?????? ??????
?? ???? ??????????. ??????????? ?????? ?
??????? ??????? ??????????? ???? ? ?????
???????????? ?????????????? ???????? ???
?????, ????? ??? ???????????? ???????? ????????? ????????? ?? ???? ?????????????
?????????? ?????????? ? ?? ??????????
???????? ???????????-???????? (???????,
???????, 1996).
?????? ?????? ?????????? ?????????????
????????????? ?????? ???? ????? ?? ???????
??????? 200 ? ? ? ??????-??????? ??????????
????????? ????? ?????????? ? ???????????
??????????. ????????? ? ??????????? ????
???????, ???????? ????- ? ???????????
???????? ???????, ??????? ????????? ????
??????? ? ???? ???????, ???????? ? ??????
??? ???????? ?????????? ???????? ??????? ?????? ????????? (??????, ????????????, 1970). ???????????? ??????????????? ?
?????????????????? ?????? ??. ?????? ???
????????? ?? ?????????? ? ???????????????? ?????????????? ??????????? ??????
???? ????? ????? (1962) ???????? ? ?????????????? ??????????? ????????? ??????? ?
?????????? ?????????? ? ?????????? ??????
???? ??? ??????? ??????? ????? ? 1,5 ?? ? ??
?????? ? 23 000 ??і (????? ???????, 1969).
???- ? ?????????????. ???? ??????
????? ? ???????????? ??????????? ??????
??????? ??????????? ???? ? ???????:
???????????? ? ???????????
? ????????? ? ????????? ? ?????
????????????. ???????? ?? ???????????????????? ???????, ?????? ?????????????
???? ? ???????? 0,12 %є (????????) (????????, 1961), ???????????? ?????? ???????? ?
??????????? ??????? ? ??????????????????
# 175 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
(Cyclops colensis baicalensis Lill.), ???????????? ??????????? ??? ?? 25 ????? ?????????, ?????????? ??????. ?? ??????????
?????? (Calanoida) ?????????? ???????
Epischura baicalensis Sars, ??????? ??????????? ??????????? ? ????????????? ?????????
Macrohectopus branickii. ?? 48 ????? ?????????? (Rotatoria), ?????????? ???? ?????, ????
????????? ??????????? Synchaeta pachypoda
Jashnov. ????????? (Infusoria) ??????? ??????????? ????? 300 ????? (?????, 1962;
??????????, 1975; ????????, 1977). ? ???????
??????? ????????? ???????? ?????? ?? ????????? ????? ????????????, ? ????? ????????????? ?????????????? (???????, 1964).
??????? ??????? ?????????, ?????????? ?
????????? ????????? ??????? ?????? ????????? ? ??????? ? Epishura baicalensis ?? ?????? ? ???????????? ??????? ????????. ???????????? ????????? Macrohectopus branickii
??????? ???????? ? ???? ?????????? ???? (??
70 ? ???????) ?? ??????? ????? ?????????,
???????, ? ???? ???????, ???????? ????????.
??????? ????? ???? ???????????? ???????
????????????, ?? ? ?????? ????? ????? ????
???? ??????????????????, ????? ??????????
? ??????????? ?????.
?????. ?????????? ? ?????????????
???? ??? ???????, ???????? ? ??????? ??????,
????????? ?????????? ?????????? ??? ??????? ?????????? ?????????? ????????????????? ?? ??????????? ???????? ??????? ???-
«??????» ???? (? ??????? ?? ??????? ?????????? ????), ? ??????? ????? ?????????, ?????????? ? ?????? ? ????? ?? ??????, ???????? ????????? ???????????? ???????????
??????????? ???????????????? ???????
?????? ? ?????????????????? ??????????????? ????????????. ???? ??????????? ?????????? ???????????????: ? ?????? ? ???????????? ??????????? (Cottocomephorus
inermis (Jakowlew)), ? ??? ? ??????????? ??-
?? ???????????? ?????? ? ????? (????????,
1973). ?? ?????? ?????? (?????, 1962), ??????????? ????? ??????????? ? 25-30 ???. ??????, ? ?? ????? ??????? ? 50-120 ???. ??????
(??????, ????????, 1974). ?? ???????? ????????? (1971, 1973, 1974), ?? ????? ???? ??????
????????? ???? ????? ?????????? ? ???????
?? ????? 3-4 ?? ?????????, ????????????
? ??????????? ??????????? (???????? ??
????????? (Cottocomephorus grewingki (Dyb.)
? ? ???? ? ???????? ??????????? (Leocottus
kessleri (Dyb.). ?????? ???? ????? ??? ????????? ? ????????? ? ??????? ?????????
?????????? ???????????????? ??????????
???????. ? ???????? ???? ???????? ??? ???
???? ??????????????? ?????????? ???? ????????? ? ????????? ??????????? ???????
??????????? ????????? ?????? ??????????? ? ????? ?????????? ????????? ???????
?????? ? ???????? ?????? ??????? ? ?????
????????? (???????, 2011).
?? ?????? ???????? (1972), ?????????? ????????? ??????? ????????? ?????????? ? ??????? 100-???????? ????. ? ???? ??
???? ??????????? ?????????? ????????????
????? ????????? ???? ?? ??????????? ?????????). ?????? ?? ???? ??????, ??????? ?????? ???? ????????? ?????, ???????????? ?
68 ???. ?????? ? 1973 ?., ???????? ????? 74,5
???. ? ????????? ? «??????» (???????? ? ??.,
1982). ? ??????? ????? ????? ????????? ?????????? 61-82 %, ? ??????? ????????? 18 %,
?????? ???????????? ????? 61 ???. ?, ? ??????
13,4 ???. ? ? ???. ??????? ??????????? ??????? ????? ????????? ?????????? 61 % ? ??????? ? 26,8 % ?? ?????? ?????? ?????????
???? ?????, ??????????? ??????? 150 ???. ?.
(91,5 ???. ?. ????? ? 40,2 ???. ? ??????? ?????????) (????????, 1977).
????? ????????????? ??? ? ??????????
?????? ?????????, ?????? ????, ?????????? ??????? ??????? ????, ????? ? ?????-
# 176 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
????????, ? ???? ?????????? ????????? ?
??????? ??????????? ?????, ??????????
????, ?????? ? ??????? ? ?????????? 1,5-2 ??
????? ????? ?? ???? ????? (?????? ????????? ?.?. ??????????). ??? ????????? ?????????? ? ?????????????????? ??????? ????????????? ????????????? ????????, ???????
?? ???? ????????????? ???????? ????????,
?????? ?????, ???????? ? ?????? ???????? ??????????.
????????, ??? ?????? ?????????? ??????????? ????????????????? ???????? ????
??????????? ????????????, ??????????? ???
???????? ?? ?????? ?????? ???????? (Billard,
1980; Lesel, 1980). ??? ???????, ???????????
??? ???????? ????????, ?????? ?????????????? ?????????????? ?????????? (1985), ???????? ? ?????????? (1985), ??????? ??????????
??? ??????????? ??????? ?????????????????
? ??????????? ???????. ??? ??????????
???? ???????????? ????? ???????? ????????
????????? ? ????????? (2009) ? ?????????
? ?????????? (2009), ????????????? ?????????? ???????? ????????????? ????????? ?
????? ??????? ? ?????? ???????? ????.
?????????. ?????? ? ????????? ????????? ????? ????? ???????????? ????????
?????????? ?????? ????????????? ? ???????
????????? ??????? ???? ? ???????. ???, ??
???? ????? ??????????? ??????? ? ????????? ????? ??????? (1949) ?????? ????????
???????????? ????? ????????? ???????? ?
36 ???. ????. ?????? (1955) ?? ?????? ?????
????????? ?????? ????????? ? 3,5 ???. ????.
???? ?????????????? ???????????? ??????
(???????, 1969) ???? ????????? ? 60 ???. ????.
???????? (1977), ??????? ??????????? ? ?????????? ???????????? ???? ??????? ??????,
??????? ?????? ??????? ????? ????? ????????? ? 150 ???. ????. ???????? ????????, ??????????? ???????? (1983), ? ????? ??????
???? ????????? ????? ????? ?????????????
?? 0,9 ?? 9,1 ???? ??? (??????? 5,0 ????) ??????? ??????? ? ????????. ????? ???????, ????
?? ????????? ?? ???????? ??????? ???????
(1955) ??? ???? ??????????, ????? ????????
????????? ? ?????? ???? ?????????? ?? 36 ??
150 ???. ?. ????? ??????????? ? ?????????
??????????? ? ???????? ?????????, ? ?????
???????????? ?????? ?? ???????????? ? ?????? ??????? (???????, 1969; ???????, 1971,
1977; ????????, 1977; ????????, 2002) ?????
????????? ????????? ???????.
?????? ????? ????????????? ???????, ???????? ???????????????????? ?????
???????, ?? ????????? ?? ???????? ????????? ????????????????? ?????. ?? ?????
???? ??????????, ??? ????????? ?????????
????????????? ???????? (?-???) ? ??????????? ???? ??????? ?????????? ?? ??? ? ??????? ??? ???? ????????? ????? 127 ?/? 2. ???????????? ????????, 159,4 ?/? 2, ???????? ?
1968 ?. (? ??? ????????? ???????? ????????),
? ??????????? ???????? ? 1966 ?. ? 66,3 ?/? 2.
?? ???? ??????? ????????? ?????????????
???????? ? ??????? ?? ?????? ???????????? ?
1964 ?? 1969 ??. ??????????? ? 2,3 ????. ?????????????????? ????????? 1966 ? 1969 ??.,
? ??????? ?????????????? ???? ???????? ?
1968 ?. (???????? ? ??., 1975). ??? ????????????? ??????????????, ??? ??????? ?? ?????
??? ?????????? ??????????? ????????????? ????? 416 ???. ??????/?, ? ???????? ?
1000,2 ??/? 2. ??????????? ?????????????
? ?????????????????? ???? ?????????? ??
?????????????? ? 10 ???, ? ???????? ? 20 ???
(???????? ? ??., 1975). ????? ?????????? ?
?????????????? ???????, ??????????????
??????????? ??????????? ????????????, ??
????? ?? ?????????? ??? ?? ????????????,
??? ? ?? ??????????? ????????? (??????????, 1975), ???, ????????, ????? ?????????
??????????? ? ??????, ??????????????
??????? ???????? ? ?????? ?????.
# 177 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
????????? ???? ?????????? ?????????
??-?????? ????????? ?? ???? ??????? ? ????? ????????? ? ?????????? ???????. ????????? ???????? ? ??????????? ???????
??????????-????????????????? ?????? ????????? ???????? ? ?????? ????? ??????? ?? ?????? ? ????????????? ????????? ?
????? (?????????, 1985), ?? ? ? ???????????
????, ??????? ???????, ????????? ????????
? ?????????-????????? ?????? ?????????.
? ??????? ??????????? ?? ???? ????????? ??
?????????? ??????? ??????? ????? ? ???????????, ???????????? ??????? ?????? ?????? ?????????, ????????? ??????? ???????
?? ??????? ?????????? ?????????? ? ????????? ????????? ??????? Saprolegnia ???????
?? ???????? ?????????????: ???????, ?????????????? ? ?????????. ?????????????
???? ????? ?? ??????????? ?????????? ?
?????????? ???????????? ???????? ??????????????? (?????????, 1985; ??????? ? ??.,
1985). ????? ????????????, ??? ?????? ?? ?
????????????? ? ??? ?????????? ????? ?????????????? ? ???????? ??? ????? ?????????,
??? ? ?????????? ??? ????, ? ? ???? ????????? ???? ????? ???? ????????? ? ??????????
???????. ????? ???????, ???????????? ?? ???
??????? ????????? ???????? ? ???????????
???????????? ??????? ????????????? ?? ????
?? ??????????????? ? ?????? ???????????
????? ?????????? ? ?????????? ???????? ?
??????? ?????????????? ????? (???????? ?
??., 1975; ?????????, 1980).
???????????? ??????????? ?????????? ?????????? ???????? ???????? ? ??????
???????????? ?????? ? ????????? ????? ???
???????, ??????? ? ??????? ?????????, ?
????? ?????????????? ???????????? (??????
? ??., 1991), ????????? ??????????, ??? ? ????????????????? ????????????? ?????????????
???????? ? ??????? ??????? ???? ?????? ?????? ??????? ????. ??? ??????? ?? ??????????-
?????????, ????????? ? ???????? ? ?????????: ???????????? ? ??????????? (???????)
? ?????? ??????????? (?????????????) ?
????????? ? ????????? ? ????? (??????,
1987; Yoshii, 1999; Yoshii et al., 1999). ??????
?????? ?? ??????? ???? ????? ?????????
?? ??????? ????? ??????? ???????????????
(??????????-??????, 1971; ????????, ????????, 1983). ?? ???????? ????????? (1980),
???????????????? ?? ??? ??????? ??????????
? ??????? 3-4 ?/?2 ????????????? ????????
? ???. ??????? ???????? ??????????? ?:N,
??????? ? 10:1, ????????? ?? ????????????
??????? ? ???????? ?? ??????? ?????? ????????????? (????????, ????????, 1971).
??????????? ???????????? ???????????????? ????????? ?? ???????? ?????????????? ????????????? ???????? ? ??????? ??
?????? ? 1964 ?? 1968 ???? ????????? ??????
?????? ? ?????? ? ????????????? ?????? ????????? ??????? ?? ??? ????? ? ??? ???????????. ???? ??????????? ??????? ????????????
????????????? ?????????? ? ???????, ??????
?????????????? ? ??????? ????????? ???
??? ???????????? ?????????? ?? ???? ????????, ??????? ????????? ????, ? ????? ?
???????????? ?????????????? ?????????
??????????????? ???????????, ? ?????????,
? ?????????? ?????????????? ???????????
??????? ??????? ???? ?? ??????????? ?? ???.
? ?????????, ?????? ?? ?????? ????????????? ??????? ???????????? ????????? ? ??????? (???????? ? ??., 1975).
???? ?????????? ???? ?????????????
? ?????????? ? ??????? ???? ?????? ???????
?? ????? ?????????, ?? ?????????? ??????????? ????????? ??????? ????? ?????????????? ???????? ???????????? ????????? ??
??? ???????. ?????? ?????????????? ??????
?????? ????????? ???????, ?????? ??????????????? ?????????????? ???????, ???????, ??? ?????? ????????????? ?????????
# 178 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
(????????????) ??????? ???? ?????? ???????? ????????????? ???????????? ??????
?????????, ??????? ??????? ??? ???????? ?????????? ???????????????? ???? ??????????
30 ???. ??? (??????, ????????????, 1970).
??? ??????? ??????? ? ??????? ?????????????? ???????????? ???????. ?? ???? ??????? ?? ???????? ??????????? ??????????
???????????????? ????????? ???????????
?????????, ????????? ??????????? ?? ????? ????? ? ????????? ? ? ??????????? ?????
???????????? ?????????? ?????? ?????????, ?? ? ????? ?????? ?????????? ????????
?????????????? ????????? ???????? ????????
????? ? ????? ???????? ???????????? ?????????.
?? ???? ???????????????? ?????? ??
?????????? ?????? ?????????, ???????????
??????????? ?? ??? ??????? ? ???????????, ?
????? ??????????????? ?????? ?????? ?????????? ???????? ???? ??????? ??????????????? ???????. ?????? ?? ????, ??? ???????
??????????? ??????? ????????? ???????????
? 9,2±1,1 ???? ??. ??? ????? 82,5±12,2 ???. ?, ?
????? ????????? 32,0±4,7 ???? ??. ??? ????? 66,5±8,6 ???. ? (????????, 1977), ????????
????????? ????????. ? ?????? ????????????
????????, ???????????? ????? 200 ????, ?
???????? ????????? ??????? 41,2±5,8 ????
?????? ????????? ????? ?????? 149,0±20,8
???. ????. ???????????, ??? ????? ????????????? ??? ?????????? ????? 0,1 % ?? ????? ??????????? ???, ????? ??? ?? ??? ????? ????
????? 150 ? ? ?????, ? ?? ???? ?????? ????????
??????? 30 ???. ?. ???????, ?????? ?? ????
???????? ?? ???????? ? ????? ????, ?????-??
????? ???????????? ?????? ? ???????????? ???????????, ?? ????? ???, ??????????,
????????? ? ???????? ????????, ????????
????????????? ???? ?????????? ??????????. ????? ????, ????? 75 ???. ? ????????? ?
«??????» ? ??? ????????? ?????? (????????
? ??., 1982), ???, ???????? ??????????? ????
????????, ????????????? 75 ? ???????, ???????? ?? ??? ? ?????????? ? ?????????? ???????? ? ??????? ???????.
??? ??????? ?????? ??????????????,
?????? ??? ????????? ????????????, ???
?????????? ?? ??????????? ??????????? ??????????? ? ????????? ????????? ???????,
?????????? ?????????????? ???????. ?????? ????? ????? ?? ???? ??????? ?????
????? ???????? ?????? ????? ?????????? ?????????????? ??????????????? ? ??????????????? ????????????, ??????? ????????
??????? ???? ????????? ? ????????????
?????????????? ???????.
?????????????
??? ?????????????? ??????????? ????????????? ???? ??????: ?.?.???????????,
?.?.
??????,
?.?.
?????????
?
?.?. ?????????,??????? ???????????? ?????
? ?????????? ???????????? ?????????? ???????, ? ????? ? ????????????? ??????????? ?
?????? ?????????? ????????????? ???? ?????
????????????????? ?????????? ????????????.
?????? ??????????
????? ???????. (1969) ?.?. ??????? (???.). ??????-???????: ???. ???? ??? ??????? ????, 30 ?.
?????????? ?.?. (1975) ??????????? ????????? ???????, ??? ??????, ????????? ? ???????? ????????????. ?: ?????????? ???????? ? ??????? ? ??????? ????????. ???????????:
?????, ?. 27-32.
???????? ?.?., ???????? ?.?., ?????? ?.?. (1982) ?????-??????????????? ????????????
??????????? ?????. ???????????: ?????, 152 ?.
# 179 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
???????? ?.?. (2007) ??????????? ?????????? ??? ????? ??????. ???????: ????????????
?? ???, 46 ?.
??????? ?.?., ??????? ?.?. (1996) ????????????? ???????? ??? ??????? (?????? ?? «???????»). ???????????: ?????, 118 ?.
????????? ?.?., ????????? ?.?. (1929) ????????? ?????????? ??? ????????? ?????????.
????. ?? ????. ????? ?. ?. 5, ?. 126-130.
????????? ?.?. (1985) ????????????? ???????? ??????????? ????? ? ????? ??????. ?:
?????????? ???????? ? ??????? ? ????????. ???. 3. ?????????, ?????????????? ? ???????????????? ????????? ?????? ??????????. ???????: ?? ?? ????, ?. 9-10.
???????? ?.?. (1983) ???????? ??????? ? ????????????? ???????? ???? ??????????? ???????. ?: ???????? ?????????????? ??? ???????. ???????????: ?????, ?. 63-75.
???????? ?.?., ???????? ?.?. (1983) ????????????? ?????????? ??? ? ????? ? ???????.
???????????: ?????, 248 ?.
???????? ?.?. (1961) ?????????? ????? ??????. ????? ??????????? ??????. ???????.
?. ??, 310 ?.
???????? ?.?., ?????????? ?.?., ????????? ?.?. (1975) ?????????? ????????????? ???????? ? ????? ??????. ???????????: ?????, 190 ?.
????????? ?.?. (1980) ???????????? ???????? ?????? ??????? ???????. ???????????:
?????, 80 ?.
??????? ?.?. (1988). ?????? ? ???????? ? ???????. ?.: ?????, 288 ?.
?????? ?.?., ???????????? ?.?. (1970) ? ???????? ??????? ???? ? ?????? ??????????
????? ??????. ?: ???. ???????????. ???. ???????. ?-?? ????. ?. 6, ? 4, ?. 64-69.
??????? ?.?. (1963) ?????? ??????? ????????? ??????????? ????????? ???. ????. ??????????. 3 (2): 288-303.
?????? ?.?., ???????? ?.?. (1974) ??????? ? ??????? ??????????????? ????????????
??? ? ????? ???????. ???????????: ?????, 185 ?.
????????? ?.?., ????????? ?.?. (1984) ??????? ?????????? ???????? ? ??????????? ??????????? ????????? ????????? ?????????????? Cottoidei. ?: ????????? ?????????? ???. ???????, ?. 28-31.
??????? ?.?., ??????? ?.?., ???????? ?.?., ????????? ?.?., ????????? ?.?., ?????????
?.?., ???????? ?.?., ????????? ?.?. (1985) ????????? ? ???????????????? ????????? ????????? ??. ??????. ?: ?????????? ???????? ? ??????? ? ????????. ???. 3. ?????????, ?????????????? ? ???????????????? ????????? ?????? ??????????. ???????: ?? ?? ????, ?. 21-22.
?????? ?.?. (1987) ???????? ??????? ???????? ? ??????? ??????????? ?? ????????? ???????????? ?????????? ???????? (18?/12?). ???????? ????. 5: 3-12.
?????? ?.?., ???????? ?.?., ??????? ?.?. (1991) ?????? ??????? ?????? ??? ????? ?????? ?? ???????????? ?????????? ???????? ????????. ????. ?? ????. 318 (5): 1268-1271.
????? ?.?. (1962) ???????? ????? ??????. ?.: ?????, 315 ?.
????? ?.?. (1963) ? ???????? ?????? ? ????????? ???????????? ???????? ??. ??????.
???. ?? ?? ????. ???. ????. 3: 105-110.
?????? ?.?., ??????? ?.?., ???????? ?.?. (1979) ?????????? ??????????? ? ???????????????? ?????????. ???????-?????????????? ? ????????? ???????????? ??. ??????. ?.: ?????, 156 ?.
# 180 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
??????? ?.?. (1964) ????????, ??????? ? ????????????? ???????? ?????????. ???????????? ?? ?????????? ???????, ?. 3-74.
??????? ?.?. (1969) ????????? ???????????? ?????? ? ?????????. ?: ?????? ?????????
?? ???????? ??????????? ???????? ? ??????? ? ????????. ?. 2. ?????????????, ?. 58-59.
??????? ?.?. (1972) ???????????? ???????? ???????. ?.: ?????, 156 ?.
???????????? ?.?. (1950) ????????????? ?????? ???????????. ?????? ????. ????. 30 (3):
382-413.
???????? ?.?. (1957) ? ???????? ???????????? ???????? Ciclops colensis Lill. ????? ??????. ???. ????.-???. ??????? ?? ????. 4: 213-225.
???????? ?.?., ???????? ?.?. (2009) ?????????????????? ?????????????? ???????? ???
??????? ?? ?????? ????? ??????????? ???????????????. ?????? ?????????? ????????????
??-??. ????? ????????. 2(3): 263-270.
???????? ?.?., ???????? ?.?., ??????? ?.?., ????????? ?.?. (2009) ??????????? ????????????? ????????? ? ????? ????? ?????? ?????????? ????????. ?????? ?????????? ???????????? ??-??. ????? ????????. 2(3): 255-262.
??????? ?.?. (1949) ???????? ? ??????????????? ??? ?? ???????. ???????: ????, ?. 3-53.
???????? ?.?. (1983) ??????????? ????????????? ? ????????????? ????????? ?????????.
?: ???????? ?????????????? ??? ???????. ???????????: ?????, ?. 6-14.
???????? ?.?. (1971) ????????????? ?????????????? ??????????? ????? Phoca sibirica Gmel. ?
??????? ????????????? ????????????? ?? ???????: ???????. ???. ? ????. ????. ????. ???????: 21 ?.
???????? ?.?. (1973) ? ???? ?????? ? ??????????? ??????????????? ???????? (?? ??????? ???????) ?: ?????????? ???????? ? ??????? ? ?????? ? ??????????????. ?????????????:
?. 166-168.
???????? ?.?. (1974) ??????? ? ??????? ??????????????? ???????????? ??? ? ?????
???????. ??. 2. ???????????: ?????, ?. 103-185.
????????? ?.?. (1977) ???????? ????????????? ?????????. ?: ????????????? ?????????????? ????????? ??????? ? ?? ????????????. ???????????: ?????, ?. 5-38.
???????? ?.?. (2004) ????????? ????? ??????. ???????????: ?????, 528 ?.
???????? ?.?. (1978) ?????????? ??????????????? ??????? ????????? (Comephoridae). ?:
?????????? ? ??????????? ???. ?: ?????, ?. 25-30.
???????? ?.?. (1980) ??????????? ???????? ? ???????????? ?????????????? ??????????????? ??????? ??????????? ??????????????? ??? (Cottidae) ? ????? ? ?? ?????????: ???????. ???. ? ????. ????. ????. ?.: 18 ?.
???????? ?.?. (1982) ??????????????? ??????? ? ???????? ??????????? ???????????????
???. ???????????: ?????, 147 ?.
???????? ?.?. (2002) ????????? Cottidae. ????? ???????????? ??? ??????: ? 2 ?. ?????????? ?.?. (???.). ?. 2, ?. 144 ? 233.
??????? ?.?., ???????? ?.?. (1983) ????????????? ? ???????? ?????? ???????????????
???. ?: ???????? ?????????????? ??? ???????. ???????????: ?????, ?. 58-52.
???????? ?.?. (1977) ???????? ???????????, ???????? ? ?????????????? ?????????.
????????????? ?????????????? ????????? ??????? ? ?? ????????????. ???????????: ?????,
?. 105?114.
# 181 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
?????? ?.?. (1955) ?????-???????????? ??????? (Cottoidei). ?.-?.: ???????????? ??
????, 602 ?.
???????? ?.?., ???????? ?.?. (1971) ???????? ???????????? ?????????? ????? ? ???????
??? ?????????? ???????????? ??????????? ??????? ? ???????. ?????????. ????. 7(6): 5 ? 12.
???????? ?.?. (2008) ???????????? ????????????? ???????????? ? ????????? ??????????????? ??????????? ????????? ??? ????? ?????? ? ????????? (Comephoridae): ???????. ???. ?
????. ????. ????. ???????????: 17 ?.
??????????-?????? ?.?. (1971) ??????? ? ??????? ??????????????? ???????????? ???.
?????????? ???????????? ??????????? ??????? (???????????? ?????). ?.: ?????, ?. 267-278.
????????? ?.?. (1970) ?????????? ???????? ?????? ??????? ? ?? ?????????????? ?????????. ????. ????. 49(1): 11-22.
??????? ?.?. (1971) ????????? ?????? ? ??????????? ? ???????? ????? ?????????
(Comephorus dybowskii Korotneff). ????. ??????????. 11(5): 820-831.
??????? ?.?. (1974) ?????????????????? ??????????? ??????????? ? ???????? ???????
????????? Comephorus baicalensis (Pall.). ????. ??????????. 14(6): 990-1003.
??????? ?.?. (2011) ???? ??????? ???????? ??????????? Leocottus kesslerii (Dyb.) ? ??????????? ????? ????????? ????? ??????. ?????? ?????????. 2: 89-93.
Billard R. (1980) L?Etang et l?Agriculture des eaux. La pisciculture en etang. Paris: I.N.R.A. Publ.,
p.15-31.
Yoshii K. (1999) Stable isotope analyses of benthic organisms in Lake Baikal. Hydrobiologia. 411:
145-159.
Yoshii K., Melnik N.G., Timoshkin O.A., Bondarenko N.A., Anoshko P.N., Yoshioka T., Wada
E. (1999) Stable isotope analyses of the pelagic food web in Lake Baikal. Limnol. Oceanogr. 44 (3):
502-511.
Lesel R. (1980) Detritus et decomposeurs dans les ecosystemes aquatiques. La pisciculture en
etang. Paris: I.N.R.A. Publ., p.31-37.
Tcherniaev G.A. (1971) La biologie du development de Comephorus dybowskii Kor. poisson
endemique du lac Baical. Bull. Soc. Zool. France. 96(4): 385-398.
Tcherniaev G.A. (1976) Of the phylogenese of Cottoidei as reveled by embryological and
paleoecological evidence. Actes II Congress Europ. Ichthyol. Rev. Trav. Inst. Peches Maritimes. Paris.
T. 40, p. 764.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????. ???????? ????????? (Comephoridae) ? ?? ???? ? ???????????? ?????????????? ????? ??????
Biology of Oilfishes (Comephoridae)
and Their Role in the Formation
of Lake Baikal Bioproductivity
Gerard A. Tcherniayev
A.N. Severtsov Institute of Ecology and Evolution, RAS
33 Leninsky pr., Moscow, 117071 Russia
The biology of two Lake Baikal oilfishes (golomyanka) species (Comephoridae), including their
spawn, embryology, anatomy, feeding and vertical migrations, is briefly summarized. The rising of
bottom sediments enriched with nutrients to the photic layer of Baikal Lake water during of oilfishes
niktimeral migrations was shown. Soft parts of bottom sediments come into photic water layer on
the body surfaces, lateral lines, seismo-sensory systems and some internal organs of the fishes and
their excrements of seals that consume oilfishes as well. Estimation of the contribution of additional
nutrients in the formation of Lake Baikal primary production was done.
Keywords: oilfishes (golomyanka), niktimeral migration, transport of bottom sediment to surface,
extraction of nutrients by seals, increasing of phyto- and zooplankton.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 184-202
~~~
??? 574.523(261.245)
?????? ??????? ? ?????????? ???????????
(????????????????) ??????
??????????? ????
?.?. ????????*,
?.?. ?????????, ?.?. ?????????
??????????? ??????????????? ????????? ???????????
«????????????? ??????-????????????????? ????????
??????? ????????? ? ????????????»
(???? «??????????»)
?????? 236022, ???????????, ??. ????????, 5 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
?? ??????????? ????????? ?????????? ???????? ??????? ??????? ????? ?????????? ?
????????? ??????? ???? ?? ??????????? ????????????? ? ?????????? ????? ???????????
(????????????????) ??????. ???????????? ? ?????? ?????????? ???????????? ?????????????
??????: ???????????? ? 412, ??????????? ? 74, ?????? ? 43, ???? ? 54. ??????? ??
????????????? ?????? ???????? (??????-????????) ?????????: ???????????? ? 6,2 ?/?2
(8,7 ???/?2), ??????????? ? 2,5 ?/?2 (5,2 ???/?2), ?????? ? 20,6 ?/?2 (61,8 ???/?2). ?? ??? ?
?????? ?????????? ???????? ??? 1,4 ?/??2, ??? 1,4 ?/?2 (60 ???/?2).
??????? ????????? ????????? ????????? 14 000 ???/?2 ?? ???. ????????????? ?????????
???????????? ? ?????????? ????????? 300 ? 1100 ???/?2 ??????????????. ?????????
????????? ??????????? ????????????, ?????????? ? ??????????? ????????????? ? ???????
????????? ????????? ????? 2,3, 8,5 ? 0,06 % ??????????????. ??????????? ?????????????
???????? ??? ????????????? ? ???????? ?? ???? ????????? (??????) ??????? ????. ????????
??????? ????????????? ????????????? Cercopagis pengoi ?????????????? ?????????????
??????????? ???????????? ????????? ? ??????? ??? ??????????? ??????.
???????? ?????: ?????????? (???????????????) ?????, ?????????????, ??????? ????, ??????
???????, Cercopagis pengoi.
????????
?????????? ????? ?????????? ? ???????????? ????? ??????????? ????. ????????
*
1
??? ????????? ????????? ? ???????? ??????,
?????? ? ? ???????? ???????? ??????????. ??
???????????????? ?????? ????? ????? ????
Corresponding author E-mail address: naumenko@atlant.baltnet.ru
© Siberian Federal University. All rights reserved
# 184 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? 1. ?????????, ??????????????? ??????????? (??: ????????, 1971) ? ??????? ??????????????
?????????? ????? ??????????? (????????????????) ?????? (????????????? ???????? ??: ????? ? ??.,
2004). ? ??????? ??????? ??????????? ? ???????????? ????????
?????????
???????, ??
??????????
472,5
2
??????? ???????, ?
2,7 (0-5,2)
pH
8,5 (7,0-9,5)
?????????? ?????????, ??/?
11,0 (2,0-30,2)
?????????, ?
4,1 (0,15-8,0)
??????????? ??????, ???/ ?
48,4 (0-283,0)
????????? ????, ???N/ ?
201,3 (1,0-2440,0)
???????????? ?? ????? ?????, ?
0,6 (0,3-3,5)
????????? «a», ??/?3
41,7 (4,6-424,7)
??????? ? ???????????? ????????? ?????????
???? (?????, ?????????, 2003) ? ? ?????????
????? ???????????? ????? ????????? ??????
(????. 1). ??????????????? ????? ??????????? (????????????????) ?????? ???????????? ????????????? ???????????????? ??????:
??????? ??????????? ? ?????????? ?????,
??????????????, ?????? ? ????????? ????
???????, ????????? ????????? ? ???????
?????????????? (?????, 1971). ? ????? ???????????? ????????? ????????? ? ????????
? ???????? ? ?? ????????? ????????? ?????????? ???????????? (???) ??? ????????????????? ???????? (?????????, 1999).
? ??????????? ???????????? ??????????? ??????????? ?????????. ??????????? ?
????? ???? ?????????? ?????? ????????????
????????, ??????????? ??????? ??????? ???????????????, ????????????? ???????????
(???????????) (?????????, 1994; ?????????,
1998; ????? ? ??., 1999, 2004, 2005). ????????
???????????? ? ?????????? ?????? ????????? ? ?????????? ? ??????? 0,2-0,5 ??/???
(?????, 2006). ? ????????? ????? ???????
?????????? ????????? ??????????? ?????
???????????? ????????? ? ???? ? ??????????
?????????? ????????? ? ?????? (????. 2). ??-
??????? ? ?????????? ????? ????? ???????????? ????????? ???? ??????????? ??????
???????????? ????????? ?????????, ???
?????? ?? ??? ??????????????.
???????????? ?????????? ???????????
?????? ???????? ??, ??? ????? ????????? ??????????? ???????-??????????? ??????????
?????? (??????) (Clupea harengus membras
L.). ? ?????? ??????? ??????????? ?????????? ??????, ?? ?????? ???? «??????????», ? ??????? ?????????? 116 ??? ???., ?
???????? ? 5910 ?. ? ?????? ??????????? ????
(????-???) ?????? ???????? ??????? ??????,
??????? ????????? ??????????? ???? ??? ?
?????????? ??????? (?????, 2004). ?????????? ?????? ? ??????????, ?????? ? ??????
???????? ????? ?? ???????? ? ?? ???????
??????????? ?????????????. ???????????? ???????????? ????????? ?????? ?????????? ??????, ??????? ?????????????? ? ?????? ?? ????? ? ???? (??????, 1978; ???????,
1992).
???? ???????????? ? ?? ????????? ????????? ???????????? ????????? ?? ????????????? ???????????? ?????????? ???????????
(????????????????) ?????? ??????? ???????? ??????? ???????, ?????? ????? ???????-
# 185 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? 2. ???????? ??????? ?????????? ????????? ? ?????????? ?????? (??: ?.?. ?????, 2006)
???????, ??????
? ?????. ???????
??????????????
???????????
?????
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?????????
?????????
?? ???
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??? ? ????????? ??????? ???? ?? ????????
????? ? ??????????? ?????????????.
????????? ? ??????
?????????? ??? ??????? ????????????
????????? ?????? ? ????????? ????????????, ???????, ?????? ? ??????? ???????????
???, ?????????? ???????? (?????????, 1988,
1992 ?, ?; ????????, 1992, 2010; ?????????,
2000; ?????, ?????????, ?????, 2004), ? ????? ???????? ? ????????????? ? ?????????
?????????, ?????????????? ?? ????????????
??????????, ???????????? ?????? ? 1974 ??
2009 ??. (???????, 1985; ???????, ????????,
1992; Semionova, 2003; ???????????, 2010).
????? ???????? ?? ?????? ??????????? ????????, ????????????? ?? ????????? ??????? ?
???????????? ? ??????????????? ?????????????? (???. 1).
????? ????????????? ???????? ?????????? ??????? ??????? 1 ?, ??????????? ???????????????? 2 %-??? ????????? ????????? ? ??????????????? ????????? ???????
(???????, 1969; ???????, 1979). ?????????
???? ????????????? ????????? ?????????-
???? ???????? (???????????, ??????, 1981).
????????? ????????? ?????????? ?????????? ??????? ? ??????????? ???????????
? ??????????? ??????????????? ? ???????
(???????, 1960; ??????? 1985), ? ????? ? ???????? ????????? ?????????? (???????????,
2010).
????? ???????????? ???????? ??????????????????? ??-5 ??????? 5 ?, ????????????? ? ???? ?????????? (0,5, 1,0 ? 2,0 ?) ?
??????????? 4 %-??? ?????????? ? ????????? (Haney, Hall, 1973) ??? ??????????????
????????? ??? ?? ????????? ? ????????????
????????????. ??????????? ?????????
???? ???????????? ?? ???????????? ???????? ??????? ??????? ??????? (???????,
1969; ???????? ? ??., 1984). ???????? ?????????? ?????????? ?? ???????????? ?????
???? ?? ????? ????? (?????????, ???????,
1979 ?, ?).
???????? ????????? (P?(t)) ??????????
???????????? «???????????????» ???????
(???????, 1985):
P c t
# 186 #
( ¦ R i ) � K
2
/( 1 K
2
,
(1)
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?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
???. 1. ????? ?????? ???? ? ?????????? (???????????????) ??????
??? Ri ????? ?? ????? ??????????, ???????????? ?? ??????? (6), ?2 ? ??????????? ????????????? ????????????? ????????????????
??????? ?? ????, ?????? ?? ????????? ?????????. ??? ??? ????????? ?????????? ?????????? ????????? ?? ????????-??????????
?????? ?????????? ?????????????? ???????? ? ???????? ??????? ? ???????? ??????,
?? ????? ????????????? ??? ????????? ??
????????????? ??????. ???????? ?????????
??????????? ?????????? ????? ???????????
? ????? ???? ??? ???????? ????? ????????? ?????????? ??????????? ?????????
? ????????? ?????????? ??????. ??????
???????? ????????? ????????? ????????,
????????? ?? ????????????? ??????, ?????
???????? ?????????? ??????????? ????????? (b) (???????, 1979). ??? ??????????
?????????? ???????? ????????? ?????
???????? ???????? ????????, ? ?????????
??????????? ? ???????? ??????????? ???????????????? ????????. ?????? ??????????? ????? ??? ??? ??????????, ??? ? ???
??????? ?????????? (?????, 1972). ???????? b ??????? ??? ?? ???????? ??????? ??-
???????? ??????????, ??? ? ?? ??????????
??????? ???????? ????????? (???????????,
??????????? ??????? ? ?.?.). ?????????? ????????? ???????? ??????????? b (???????,
1979). ?????? ?????????? b ? ???????????
????????????? ???????????????? ???????
?? ???? (?2) ? ?????? ??????? ????????? ????
??????????? ??????????, ????? ?? ?????????? (d) ??????????? ????? ????. ???????? ???????????????? ???????? ?????????
??????????? ????????? ??????????, ??????????? ?????????, ??? ? ??????? ?????? ?2
?? ???????? ? ????? 0,77. ? ??????? ???????????? ?????????:
b�B ;
P('t )
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© N1
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0) ;
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ёё � (t t ) ,
№
(3)
(4)
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??????; N1 ? N2 ? ??????????? ???????? ?
??????? ?????? ? ?????? ??????? t1 ? t2,
? ?????? ????????????????? ????? ??????????? ?????????? ?????????? ???????? ??-
# 187 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? ??????????? (r) ????????? 0,514 ???.-1
?, ?????????????, ?2=0,77 (?=28,0 ??/?3,
W=0,16·10 -3 ??). ??????? ???????? ?2 ???????
? ???????? ???????????? ????????? ? ?????????? ?????? ? ???????????? ???????????
??????????? ????????????? ???????? (???????, 1985; ?????, 2006).
???????? ????????? ???????????? ? ?????????? ???????????? ?????????? ?? ????????? (???????, 1985)
n
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¦ w w � N
i
i 1
i
/ Di � q,
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i 0
??? wi, wi?1 ? ????? ????? ? ?????? ? ????? ?????????? ?????????, ??; D ? ???????????? ??????????? ??????? ?????????, ???.; Ni ? ??????????? ?????? ??????? ?????????? ?????????
(??????), ???.; q ? ????????????? ???????? ???
???????????? ????????, ??????????????? Q10 =
2,25 (???????, 1985).
???????????? ??????????? ???????????? ?????? ? ?????????? ?????? ???????????? ?? ???????????, ?????????????
?.?. ???????? (1985). ?????? ? ????????????
???????? ?????? ? ??? ???????????? ???? ???????????? ?? ?????????? (???????, 1958,
1959 ?, ?; ???????, 1985), ? ????? ??????????
?? ????????? ???????? ?????????? ?? ????????? ??????????? ? ????????-??????????
???????? ?????????.
????? ?? ????? (R?(t)) (??/???) ???????????? (??????????, 1980) ? ?????????????
(??????, 1972) ?????????? ?? ???????? ??????????? ????????? ? ??????????? ?? ?????
???????? ? ?????? ??????????????? ????????????:
R(' t )
( 24 � 0.107 W 0.796 �
� OK � N ) / J (Rotifera);
R(' t )
(6)
( 24 � 0.143W 0.803 �
� OK � N ) / J (Cladocera);
(7)
R(' t )
24 � 0.200 W 0.777 �
� OK � N ) / J (Copepoda),
(8)
??? W ? ??????? ????? ?????, ?; 24 ? ?????????? ????? ? ??????; OK ? ??????????????
???????????, ?????? 20,3 ??/?? ?; N ? ??????????? ??????, ???/?3; ? ? ?????????????
???????? ??? ???????? ??????, ??????????????? ????????? Q10 ?? 2 ?? 3 (???????, 1985).
??????????? ???????? ??????????????
??????????? ? ?????? ??????????? ????,
??????? ??????????? ??????: ?????? ? 0,6;
??????? ? 0,8; ???????? ? 0,7, ? ?????? ?????????????? ???????? ??? ????????? (?????
???????? ? ????????????), ???????????? ?
??????? ???????????? ?????? ???? ??????????, ? ????? ??? ????????? ??????????? ?
?????????? (????? ????????). ? ????? ???????? ???????? IV-VI ???????????? ?????? Megacyclops viridis (Jurine, 1820), ? ?????
Leptodora kindtii (Focke, 1844) ? Cercopagis
pengoi (Ostroumov, 1891). ??????? ???????????? ?????? ????????? ?????????? ??????, ?
??? ????? Eurytemora affinis (Poppe, 1880), ?
????? ????????? ???????? ? ???????? ????? (????????, 1970; ???????, 1985; ??????,
1989; ???????, 1998). ????????? ???????????? (?z) ?????????????? ??? ??????????????
????? ????????? ??????????, ????????????
? ?????????? ????????????
Pz
Pr Pcl Pcop ,
(9)
??? Pr, Pcl, Pcop ? ????????? ??????????, ???????????? ? ?????????? ????????????.
??? ???? ??????????????, ??? ????
????? ???????????? ???????????? ? ????? ??? ???????????? ???????????????.
????? ????, ???????????, ??? ???????????
?????? ??????????? ?????????????? ????? ? ??? ?? ????? ??????? ?????????????
??????? ?? ????????? ?????????? ????????????.
# 188 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
???? ???? ?????????? ????????? ????????????? ????????? ? ???????? ???????
0,025 ?2. ??????????? ? ??????????????
????????? ???? ???????????? ????????????? ????????. ??? ????????? ???????? ?
?????????? ?? ?????????? ?????. ? ????????? ???????? ????? ????, ?????? ? ??????
???????, ? ??????? ? ????? ? ?????? ????,
? ???????? ? ?????? ???????? ?????? ???? ?
?????????? ?????????, ? ????????? ? ?????? ? ?????? ???????? ???????, ????? ? ?????? ??????????, ?????? ?????????????????
????????. ?????????? ?????? ?????????? ??
?????????????? ?????? ??????????? ??????????? ?? ?????????? ??????? ????????? (???????? ? ??., 1984).
??????????? ????????? ?????????
??? ???????????? ????????? ?????? ?????????????? ? ?????? ????????-??????????
????????? ? ???????????? ????????. ????????? ?????? ?????????????? ???????????? ??? ????? ?????? (???????? ? ??.,
1984). ????????????????? ???????? ?????????? ????? ?????????? ?? ?????????
????? ?? ??????????? ?? ?????? ???????
????? ???????? ????. ?????? ???????? ?
??????? ?/?-????????????? ???????????
?? ????????? ????????? ?? ?????? ??????? ????????? ? ??????? ?? ???? ?? ??????
??????? ????????.
???????? ?? ??????? ??? ?????? ? 19811988 ??. ????? ???? ???????????? ???????????? ??????? ?????? ??????. ????????? ???? ?? ??????? ???, ?????? ???????? ?
????????????? ????????? ?????????????
???????? (?????. ????????, 1974). ?????
????????? ??? ??? ?? ?????? ????????????? ??????, ?????????? ????????? ?????,
?????, ??????????? ????????. ???????????
????????? ?? ??????? ??? ????????? ????????????? ? ???????????? ? ???????????????????? ????????? (?????. ????????, 1974).
??????? ??????? ??? ???????????? ?? ??????????????? ????? ????. ????? ????????
?????????? ???????????????? ?? ??????
???????? ??????? (??????????), ????????
???????? (????????), ????? ??????? (????????), ???????? ???????? (??????). ??? ????
????? ???????????? ????????? ?????????,
??????????? ??????? ??????????, ????? ?
????? ?????????????? (???????, 1985). ????
?????????????? ?????? ???????? ?? ???????? ?.?. ????????? (1971). ??????? ??????
???? ????????? ??? ????????? ?????????? ?
????????? ????? ???.
??????? ??????? ???????????? ???????????? ?? ???????? ?.?. ?????? (1977). ??????? ?????????? ?????????? ?? ??????????
???????? ???????: ???? ? ?? ???????? ??.
???????? ? ?. ??????? (Elliott, Persson, 1978),
???? ? ?.?. ?????????? (1986). ???????? ????????? ???? ???? ?????????? ??? ????????
??????????? ?????????? ???????????? ?
??????????? ????? ???? ? ?????? ?????
????????????? ??????? (?? ?????? ???????? ???????). ??? ???? ???????????? ???????????? ?????? ?? ????????? ?????????
???? (?????????, 1989). ???????? ? ???????
??????? ??????????? ??? ?????????? ????? ??????? ??????????? ????????? (???????,
1956), ??? ???????? ???????? ???????? ????????? ???????? ??? ?????, ????? ??????? ?
?????? ?????????? ???????? ??????? ????
??????????.
???????????? ????????????? ????????????? ???? ?? ???? (?1 ? ?2) ? ??????????? ??? ?????????? ?? ?.?. ?????? (1986),
??????? ???????? ????????, ????????????? ?
???? ??????? ??????????? ? ?? ?.?. ???????? (1952).
?????? ?? ??????????? ? ????????? ????
????????????? ??????????????? ??????? ??
????????????????? ? ??????????? ?????????? ?????.
# 189 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????????
??????? ??????, ???????? ???????????
? ???????? ?????????????. ???????????
???????????? ??????????? ?????? ??????????? 412 ?????? ??????????, ??????????? ? 8
??????????????? ???????. ?????????? ?????????? ????? ????????? ? ??????? ? 143, ?????????? ? 106, ??????????? ? 77, 33 ???????
???? ???? ???????? ???????????? ?????????? (Semionova, 2003). ?? ???????? ???????
???????????? ?????????? ????? ??????????? (????????????????) ?????? ??????? ????????????????. ? ??????? ?????? ?????? ?
?????? ???????? 9-12 ?????. ???????????
????????? ????? ????????? ???????? ? ??????? ??????. ?????????? ?????? ??????????? ? ????????????? ?????????. ? ??????
?????????? ?????? ???????????, ???????????????? ? ??????????? ?????????. ? ??????
«????» (????????????) ??????? ???? ???????? ????? ? ???????? ????????????? ? ??????? ???? 100 %. ??? Ankistrodesmus falcatus
(Corda) Ralfs, Gomphosphaeria lacustris Chod,
Planktolyngbya contorta(Lemmermann) Anagnostidis & Komбrek, Merismopedia glauca (Ehr.)
Naegeli, Phormidium sp. Kuetzing Ex Gomont.
??????????? ???????????? ?????????????
??????????? ?????? ???????? ??, ??? ???????
? ???????????? ????, ???????? ? ?????? ??????????, ????? ?????????????? ???????
????? ??????.
? ??????? ?? ??? ??????????? ????????????? ?????????? 140 ???? ??/?3, ????????
1,8 ?/?3 (????. 3). ?????? ???????? ????????
????????? ??????????, ????? ? ?????? ? ???????????. ?? ????????? ??????? ???????????? ??????????? ????????????. ??????????
?????????? ????????????? ? ??????????
??????? ? ?? ???? ???????? ?? ???????????
???????. ?????????? ????????? ????????
???????? ???????? ????????????.
????????? ????????? ???????????????
????? ??????, ???????????? ? ?????? ???????? ?????????????. ? ?????? ??????? ???-
??????? 3. ????????????????? ?????????????? ???????? ????????????? (Bf, ?/?3), ?????? ?????????
????????? (Pp, ???/?3), ?/?, ??????? ????????? (?, ???/?2) ? ???????????? ???? ?? ????? ????? (S,?) ?
?????????? ?????? (? ? ?????????????????? ??????????, ????? ???? 717)
?????
Bf
Pp
?/?
?
S
I
0,8
-
-
-
-
II
0,6
-
-
-
-
III
2,2
616,3
0,6
1658,5
0,7
IV
2,0
511,2
0,8
1528,5
0,8
V
2,0
836,4
0,8
2000,7
0,5
VI
2,3
766,8
0,8
1604,9
0,6
VII
3,5
484,2
0,4
1013,4
0,6
VIII
2,8
1364,6
1,1
3264,1
0,6
IX
2,4
852,0
0,8
1783,2
0,6
X
1,4
308,1
0,5
737,0
0,7
XI
1,0
85,2
0,5
254,7
0,7
XII
1,0
31,2
93,3
1,0
??????? III-XII
1,8
585,6
0,7
1393,8
0,7
?
0,9
397,9
0,2
925,9
0,1
# 190 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
?????? ??????????? ?????? ????????? ????????? ?????????? ????? 1400 ???/?3 ? ?????.
?????? ???????? ??????????? ????????????
???????? ??????????? ? ? ??????? ??????????? ????????? ????????????. ???????????? ????????????? ??????????? ?????????? ?
????????????? ???? ????, ???????? ? ????????.
?????????? ??????????? ?????? ??????????????? ?? ?????? ??????? ????????? ??????????? ???????????? ????????, ?? ? ??????? ????????? ??? ??????????. ? ???????
?? ????????????? ????? ? ?????????????
???? ?? ?????????????? ???????? ??????????? 18,4 ???/?3 ? ???. ?? ? ??????????? ?????????? ?? 40 ?? 80 % ???????. ? ?????????
???? ? ?????????? ????????? ??????????
???? ????????? ????????? ?????? ?????????????. ? ??????? ?? ??? ? ??????????
?????? ? ????????????? ???? ?????????????
????? 14 000 ???/?2 ????????????? ????????
(????. 3). ??????? ?????? ????????? ????????? ????????? 3100 ???/?2.
???????????. ??????????? ? ??????????
????? ??????????? ?????? ??????????? 74
????????????? ?????? ? ?????????, ???????????? ? ???? ??????????????? ???????:
Rotifera, Cladocera, Copepoda. ?? ???????????
? ???????? ??????????? Copepoda ? 64 ? 80 %
??????????????. ?? ??????????? ? ???????? ?
??????? ???????????? ? ?????? ??????????
E. affinis. ? ??????? ?? ????????????? ?????? ?? ????? ????????????? (0,75±0,78) ???/?3
(????. 4). ?? ??? ????????? ???????????? ?????????? 85 ???/?3, ??? 300 ???/?2.
??????????? ??????????? ????? ???????????? ????????: ??????????, ?????????
? ?????????, ????????????? 75, 14 ? 11 %
?? ?????? ?????????? ?????. ???????? ?????
??????? ???? ?? ????? ????? ? ????? ?????????? ????????????, ?????????????? ????????? ?????? (????. 4). ? ??????? ????????
????? ??????????? ???????????? ????. ??????????? ?????? ??????????? ????????
??????, ??????? ?? ???????? ????? ??????????.
????? ???????? ? ?????????? ????? ?????? ???????????? ???????????? C. pengoi ?
1999 ?. ????????? ??????????? ????????? ?
?????????? ???????????? (????????, ?????,
2008). ??????????? ??????????? ??????????,
? ?????????? ???????????? ? ?????????. ?????? ????????? ???????????? ? ?????????
????????? ? ???????? ??????? ??????????
??????? 4. ????????????????? ?? ????????????? ????? ???????? ?????????? ????????????????
???????????? ??????????? ?????? (? ? ???????? ????????????, ??/?3, P ? ????????? ????????????,
??/?3; ? ? ???????????????? ??????? ?????????????, ??/?3; ? ? ?????? ????????????, ??/?3; ? ?
?????????????????? ??????????, ????? ???? 1100)
??????????
B, ??
P, ??
A, ??
C, ??
Rotifera
67,8
173,4
212,1
351,5
?
95,5
201,4
243,3
404,8
Cladocera
242,9
268,6
377,1
617,0
?
255,5
392,9
542,3
880,9
Copepoda
791,9
307,5
926,0
1389,3
?
695,4
186,7
572,4
887,2
?????
1102,6
749,5
1515,2
2357,8
?
745,1
781,0
1358,0
2172,9
# 191 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? 5. ????????????????? ?????????? ?????????????? ?????? ?????????????? ? ??????????
?????? (??????-????????), ????? ???? ? 1620
??????????
??????????
?????????
????????
????????
?????
????????, ?/?2
11,1
2,6
4,6
6,9
25,1
?
7,7
1,8
5,3
10,6
13,8
?/? ???.
3,6
3
2
0,8
2,3
?????????, ?/?2 ???.
13,3
2,6
5,1
15,4
36,0
?
14,0
2,0
5,4
57,0
65,6
????????? ???/?2 ???
625,7
197,9
224,1
39,6
1087,3
?
429,6
138,7
265,8
61,1
452,5
??????, ??? ???????, ?? ?????? ???????, ??????????? ??????????? ????????? ?????
????????????? ? ????????? ??????????
??????.
?????????. ????????? ? ?????????? ????? ??????????? ?????? ? ??????????? ?????? ??????????? 43 ????????? ???? ????
??????????????. ???????? ???????? ????????? ??????????, ?????????, ???????? ?
????????. ????? ???????? ? ?????????????
??????? Marenzelleria viridis Verlin ????????
????????? ? ????????? ?????????? ? ???????
???????? ???? ????????? ? ?????????? ????
????? ???? (????? ? ??., 2004). ? ??????????
???????? ???? ???-?????????? ??????????,
??? ??? ?????????????, ?????????? ? ????? ??
40 ??, ?????????? ???????????? ??? ???.
??????? ? ?????? ?? ???????? ???????? ?????? ?????????????? ?????????
(25,1±13,8) ?/?2. ???????? ???????? ????????? ?????????? ????????? ? ????????? ???????? (????. 5). ??????? ??????????????? ?
?????????? ????? ??????????? ?????? ????????????? ? ??????? ?? ????????????? ??????
(36,0±65,6) ?/?2, ??? ?????1100 ???/?2.
????????????? ????????? ?????????
???????????? ? ??????? ???????????????.
??????? ?????? ??????????? ????????????
? ?????????? ?????? ???????? ????????,
??????, ????????????, ????????? ? ??????
??????????. ?????????? ????? ??????????
?????? ???????? ?? 4 ?? 50 ???3, ?????? ?
E. affinis ??????????? ???????????? ????,
????????? ?? 60 % ????? ???????? ?????.
?????? E. affinis, ???????????? ??????? ??????????? ?????????, ? ??????? ?????????????? ??????? ??????? ?? 0,1 ?? 3,6 ???/?3·???,
????????? ? ??????? 1,1 ???/?3·???. ?? ????????????? ?????? ????????????? ???????????? ????? 970 ???/?2. ????????? ????????
??????? ??????????? ?????? ? ??????? ????????? ????????? ????????? 6,9 %, ? ??????
????????? ????????? ? 31,3 %. ?????? ?????????????? (??????????, ?????????, ????????, ????????) ?? ????????? ????????????
??? ???? ????????? ? ????????????????. ??
??????? ?????? ??????? ??????????????? ??
???????????? ???????? ???????? ? ??????
?????. ? ??????? ?? ????????????? ??????
???????? ?????? ????????? ???????? 20 %,
???????? ? 12 % ?? ????? ????. ?????? ????????? ?????????????? ??? ? ????????? ?
2,1, ? ???????? ? 0,18 ?/?2 ? ?????. ?????????
??????? ?????? ?????????????? ? ??????
????????? ????????? ????????? 1 %.
????. ?????????? ? ?????????? ?????? ???????? 54 ???? ??????????? ? ???,
?????????????? ????? ????????? ???????????? ????????, ????? ??????? ?? ?????????? ????? ??????????? ????????? (55 %) ?
# 192 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??? (1-15-????????)
?????? (2-14-????????)
n=1920
??????????
??????????
????????
????????
????????
????????
??????
??????
????
????
??????? (3-12-????????)
n=137
??????????
??????????
????????
????????
????????
????????
??????
??????
????
????
??? (1-11-????????)
n=156
??????????
??????????
????????
????????
????????
????????
??????
??????
????
????
????? (4-15-????????)
n=76
??????????
??????????
????????
????????
????????
????????
??????
??????
????
????
????? (0-1-????????)
n=362
??????????
??????????
????????
????????
????????
????????
??????
??????
????
n=716
????? (3-4-????????)
n=14
????? (2-8-????????)
n=552
?????? (1-11-????????)
n=156
????? (2-12-????????)
n=536
????
0
20
40
60
80
100 %
0
20
40
60
80
100 %
???. 2. ??????? ?? ????????????? ?????? ??????? ??????? ???????????? ? ?????? ??? ? ??????????
??????. ?? ??? ??????? ? ??????? ?? ????? ???????? ?????
??????? (18 %). ?? ???? ???????????? ?????????? 11 %, ???????? ? 13 %. ????? ??????????? (????????) ????? ?????????? ???????? ? ???????? ????? ??? (Abramis brama
(Linne)), ????? (Sander lucioperca (Linne)), ?????? (Pelecus cultratus (L.)) ? ????? (Anguilla
anguilla (Linne)).
???????? ??????????? ?? ????????????
??? ?????? ???, ??????? ??? ??????????? ?
??????? ?????? ??????? ?? 1 ?? 15 ???. ???
? ???????? ?????? ???? ??? ??????? ??????????????? ????????? ????????? ?? ?????????? ????? ??????? ??????. ????? ????
?????????? ????? ???????? ????????, ??????????? ?????????????? ??????????? ? ??-
????? ?????? ? ?????????????? ??????????
(???. 2).
?????? ? ?????????? ?????? ? ???????? ?????????? ?????????, ??????? ??????
??????? ????? ? ?????????? 80 % ????????
???????. ?????????????? ????? ?????? ???????? ?????????? (17 %), ??????????????
? ??????? ? ???????????? ?????????? ??????,
??????????????? ? ???????? ?????????, ?????? ? ?????? ?????????????? (???. 2).
??????? ? ??? ???????? ????? ??????????? ? ?????????? ????? ??????????? ??????.
??????? ???????? ? ???????? ????????????
(?? 75 % ???????? ???????). ??????????????
????? ?????? ????????, ??????, ???????-
# 193 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??, ??????? ?????????? ? ?? 20 % ? ??????????????? ????? ? ?????? ???, ????????????, ?????????? ???????????? (????? 5 %).
???????? ???? ???? ? ??????? ????? ?????????????? ?????? ? ??????? ?????????, ?
????? ??????. ??????? ????????? ? ????
?????????? 88 %, ?????????????? ????? ???????? ?????? (11 %), ??????????????? ? ?????? ???, ????????? (???. 2).
? ??????? ????? ???? ? ??????????
?????? ??????????? ??????????????? ????????? ? ?????????????? ??????????????:
????????, ??????????, ??????. ??????
??? ?????????? ?????????????? ????? ???
???????. ??????? ???? ???? ? ???????? ? ?????????? (84 %), ?????????????? ? ??????
(16 %), ??????????????? ? ????????, ???? ?
????? 1 % ????, ???????????? ? ??????? ????
(???. 2).
????? ???????? ? ???????? ?????? ????? ????? ????. ????????? ? ???????? ?????? ?????? ? ? ?????? ???? ?????????? ?????????, ?? ??????? ? ????, ????? ?????????
????????? ?????????, ????????? ?? ???????
???????? (Neomysis integer (Leach)), ???????
?????????? ?????? ? ??????? (Osmerus eperlanus eperlanus morpha spirinchus Pallas). ??????? ? 2-?????????? ???????? ????? ????????
??????????????? ?????, ??????????????
???????? ????????? ????? ? ????????????
? ????????? ??????????. ?????? ????? ?????????????? ???????, ???????? ?? ??????.
?????, ????? ??????????????? ??????????
?????? ??????????? ? ?????????? ????, ????? ???????? ???????, ?????, ? ????? ??????????? ???????. ?????? ? ??????? ??????
????? ?????????? ???????? ??????. ????????
??? ????????????? ????????????????? ?????????? ??????, ??????? ???????? ???????
? ?????????? ?????. ????? ????, ?????, ???
? ???, ????? ???????? ? ?????????? ?????
??????????? ????, ??? ???????? ???????.
??????? ????? ?????????? ? ?????????
?????? ???????? ?????? ? ??????, ???????
?????????? 78 %; ?????????????? ? ??????
?????????? ??????, ?????, ??????? ?????????????? (19 %); ??????????????? ?????, ???????????? 6 %, ? ????????? ??????, ?????,
??????? ?????????. ??????? ???? ??????
? ???? ??? ???????? ?? ?????????? ??????
? ?????? (70 %); ?????????????? ? ?? ????,
?????? ?????? (24 %); ??????????????? ? ??
??????? Osmerus eperlanus eperlanus Pallas,
????????? ??????? Gasterosteus aculeatus L.,
?????? ??????? Pleuronectes flesus L., ??????,
??????, ???????? ? ????? (6 %) (???. 2). ?????
???????, ? ???????????????? ?????????? ?????? ??????? ??????????? ?????? ?? ??????
??????? ?????????? ? ???????????? ???????
??????????????? ?? ???? ????? ???????? ?????????????: ????? ? ??????.
????? ? ?????? ?? ??????????? ???????
??????????? ? ???????????????? ?????????
????????. ?? ????????? ???, ????????? ?
?????????????? ?????????????? (???. 2).
??????? ????????, ??? ??????? ????? ?????,
???????????? 72 % ???????? ???????????
?????, ???????? ??????, ??? ? ??????. ?????????? ???????????? ?????????????? ????
? ?????????? 20 % ????? ????. ?? ????????
???????? ?????? ? ??? ? ????. ??????????????? ???? ? ?????? ??????, ?????? ???????,
?????, ????????, ?????, ?????????, ???????
?????????? 8 % (???. 2).
??????? ????? ??????? ??????? ? ?????????? ?????? ? ?????? ? ???? ?????? ??????,
?????? ? ?????? ? ?????? (?? 80 %), ?????????????? ? ????????????? (?? 17 %), ???????????????, ???????????? ????? 3 %, ? ???????
? ??????? ?????????, ???????????? ? ?????????. ??????? ????????, ??? ? ??????? ?????
?????????????? ?????? ? ??????? ?????? ??????????? ????. ??? ? ?????, ?????? ???????
??????????? ?? ???????? (???. 2).
# 194 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
???? ? ????? ? ?????? ??????????? ?????, ? ???????? ?? ??????? ??????? ?? ?????????????? ?????????. ? ???????? ??????
?????????? ???, ?????, ??????, ???????
???????-???????, ?????????? ????????. ?
???? ?????????????? ???????????? ???????
?????? ??????, ???, ??????.
??? ?????????????? ???????, ?????
??????? ????, ?????????? ?????? ????????? ??????????? ? ???? ??? ????? ?????,
??????????? ???????? ?????? ????????????? ???????????, ??? ??????? ????????????
???. ?????????? ???????? ???????????
????? ????, ?????? ???????????? ?????????? +0,20 ??????????? ???? ?????? ?
????????, ?????? ?? ???????????? +0,46.
??????? ???????????? ???????? ??????????? ?????????? (+0,20), ????????????
(+0,43). ??????????? ???? ???? ? ?????????? (+0,14), ? ???? ? ???????? ? ???????
?????????. ????? ? ???????????? ? ??????
???????? ?????? ??????????? ???
??????????? (????????????????) ?????? ????????? ?? 1,2 ?? 3,2 % ?? ????? ???? (????. 6).
?????????? ?????? ? ??????, ??????? ?????????? ??????????????? ????, ? ????,
?????????? ?????????????? ???. ? ??????
?????????? ?????? ?? ????????? ? ?????
????, ?? ??? ???? ???????? ? ??? ???? ?????
??????????, ??? ? ???????????? ???. ???????? ??????????? ????????????? ?????????
?? 0,1 ?/?2 (0,4 ???/?2) ? ????? ? ??????, ??
2,7 ?/?2 (7,9 ???/?2) ? ???? (????. 6).
????????? ??????? ?????. ????????????? ???????? ? ??????? ? ??????????
(???????????????) ?????? ???? ?? ???? ????????????: ????? ??????????? (????????????) ? ????????? (??????) ??????? ????.
??????????? ??????? ???? ??????? ? ??????
?????????????? ???????, ????????????????
??????????????, ???????????? ? ?????????? ?????? 14 000 ???/?2. ???????????? ?
??????????? ???????, ?????? ?? ???????????? ????? ????? ? ?????????? +0,93. ???
???????? ????????, ???????, ?????????;
?????? ? ?????????, ????????; ??????? ?
????????, ???????; ??? ? ????????, ???????, ?????????, ????? ? ?????? ?????????,
?????????, ????????.
???????? ???????????? ?????????????, ??????? ?? ????????????? ?????? ????????? ???????? ???????????? ????????? ? 300 ???/?2.
??????????? ??????? ??????? ???????????? ? ???? ????????? ? ???????? ???, ???????? ??????? ?????????? ?????? (??????)
C. harengus membras. ?????? O. eperlanus
??????? 6. ??????? ? ????????????? ??????????? ????????????? ??????????? ??? ???????????
??????
????????????? ???????????
?????????????
??????
???
????????, % ??
????? ????
1,6
???????,
???/???
8545
Anguilla anguilla
2,6
Sander lucioperca
?/?2
???/?2
2,7
7,9
7200
1,2
3,5
1,2
17800
1,3
5,4
Perca fluviatilis
1,6
4000
0,1
0,4
Rutilus rutilus
3,2
4650
0,5
0,8
Pelecus cultratus
1,8
6500
0,1
0,4
Abramis brama
# 195 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????????? ??????? ????
?
?????????
?????????
(14000)
?
??????
(???????? ???????? ??)
???
??
????????? ???????????????
?????????
????????????
(300)
(????????
???????? ??)
??? ?????????
?????????????
???-??????????
??
(???????? ???????? ??)
?????????
???????
(1100)
??
?????
????????
(2)
???
?????
??????????
(6)
?????? ??????? ????
???. 3. ????? ??????? ???? ? ??????? ??????? ? ?????????? ?????? ?? ??????????? ?????????????
(? ??????? ? ???/?2 ? ???)
eperlanus morpha spirinhus ? ????????????
?????????? ? ?????????? ??????, ?? ??????????? ??? ????????, ? ???????????? ???????? ?? ?? ?????. ?????? ?????????? ??????
?? ?????? ???????? ???? ??????????? ? ????,
??? ???????????? ?? ??????? ????????. ?????
?????? ???????????? ? ????? ?? ??????, ??? ??
????????, ?? ?????? ???????? ????????.
???????? ?????? ?????????? ???? ???????? ?????? ????: ?????, ??????, ?????.
????????? ??? ???? ????????? ????? ?? ????????????, ??????? ?? ????? ? ????? ??????? ????????? ????? ??????????. ??????????? ?????????????? ?????? ????? ?????
0,4 ?/?2, ??? 2 ???/?2, ??? ?????????? ???????? 30 % ?????? ?????? ??????????? ????? ???. ????? 50 % ??????? ?????????????
?????? ?????????? ?????????? ??????, 40 % ?
?????? ? ?????? ???, 10 % ? ???. ????????
???????? ?????????? ???????????? ????
???? ?????, ??????????, ??? 50 % ?????????
?????? ?????????? ?? ???? ?????????? ??????????? ????, ???? ?? ??????? ?? ????? ? ???
??????????? ??????????? ????? ??????????
??????. ? ?????????? ?????? ???????????
????????????? ?????? ??????????? ?? ????
??????????? ??????? ???? ?? 40 ? ?? 10 % ?
?? ???? ??????. ????? ???????, ?????????? ??
???? ??????????? ??????? ???? ??????????? ?????? 12 % ??????????? ?????????????
??????????? ?????? (?????? ???? ?????????? ????? 30 % ?? ?????? ??????????? ?????
???). ???????? ????? ??????? ?? ??????????? ???? ???? ?? ???????? ????????? ??????
???, ?.?. ??? ?????????? ???????.
??????? ????????? ?????? ?????????????? ?????????? ????? 1100 ???/?2. ????????? ???????????? ????? 70 % ?????? ??????
????, ?? ??????????? ????????????? ? 1 ?/?2
(6 ???/?2) (???. 3).
? ???????? ???????????????? ???????? ??????? ???? ???????? ?????????????
???????????? ????, ??????? ??????? ???.
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?????, ??????????? ???????????? ? ??????
??????? ???? ?? ????????? ???????????
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?????????? ????????????? ???????? ? ??????? ?? ??????????? ????????????? ?????
?????? ??????? ???? (?? 75 %). ????? 20 %
??????????? ????????????? ?????? (???
50 % ?????? ??????) ?????????? ?? ???? ??????????? ??????? ???? ??????????? ????.
????????? ?????? ??????????? ?????
??? ? ???????? ??????? ????????? ????????? ?????????? 0,06 %.
??????????
???????????? ??????? ??????? ? ???????? ? ???????????, ???????????? ?????????????????? ???????? ????????? ??????????
????? ???????? (?????? ? ??., 1993; ????????, ??????, 1993; ??????, 2000; ???????,
2000; Hakanson, Buolion, 2002). ??? ???????????? ????? ? ????? ?????? ??????????
?????. ? ?????? ??????????? ?????? ????
?????????? ??????, ??????????? ?????? ?
?????? ??????????? ?????? ? ??????? (?????????, 2000; ????????? ? ??., 2001).
? ?????????? ?????? ?? ???? ???????
??????????? ????-?????????, ??????? ????? ??????? ???????? ? ??????????? ?????????????. ???????????? ??????? ???????????? ??? ?????? ????? ??????????? ??????
?????????? ? ?? ??????? ????????????. ?????????? ????????, ??? ?? ?????? ????????
??? ???????????? ??? ?????????? ???????
?? ??????? ????????????? ? ?????? ??????
?????????. ?????? ??????? ??????????? ?
???????? ?????? ??????????? ????, ????????????? ? ?????? ??????? (?????????, 1982).
??? ???????????? ??? ??????????? ??????
?????????? ?????????? ??????? ? ??????????? ????, ? ??????????? ???????? ? ????
?????????? ?????????? ?????. ???????????
??? ?????? ?????????? ???????????? ? ????
??????. ?????????? ?????????? ?????????.
??? ???????? ????????? ?? ??????? ?????????????? ?????????? ?????. ?????? ???-
????????? ??????? ??? ? ?????????? ??????
???????? ???????????? ? ???? ???? ??????
??????????????. ?? ???????????? ??????
????? ? ???????, ?????????? ????????? ???????????, ?????????????? ? ?????? ??????? (Nie de, 1987). ????????? ?????? ?????????????? ???? ??? ???????-?????????. ?
?????????? ?????? ????? ? ???????? ????????.
?????? ??????????? ???-???????????? ?
?????????? ?????? ?????????????? ????, ???
???????? ????? ??????? ?? ??????????? ?????????????, ? ??? ????? ? ?????????????
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??????? ????. ?????????? ??????????? ?????? ???????? ???????????? ??????????????
??????? ?? ?????????? ??????????? ????
?? ???????? ??????? ??????????????????
????????, ??? ???????????? ??????????? ????????????? ?????? ??????????????????,
???????, ?? ?????? ???? «??????????»,
??? ????? ?????????? ?????? ? ??????? ?????????? ????? 1,4 ?/?2. ??? ???????? ??????
???????? ??????????? ?????????????. ?
????????? ????????, ? ??????? ?? ????????
????????? ????????? ????????? ??????????
?????, ????????????? ?????????? ???????
24 ?/?2 (???????, 2000). ?????? ?? ????????
????????????? ?????????? ????? ?????
???? ??????? ? ???????????? ????????, ?
??????? ????????????? ?????????? ? ??????? 6-7 ?/?2 (???????, 2000).
???????????? ????????????? ?? ????
???????? (????????, ??????, 1993; ?????????, 2000; ????????? ? ??., 2001) ????????, ??? ?????????? ??????????? ??????????????? ???????????? ??????????
??????????? ?????? ??? ? ????? ???????
?????????? ????????????? ???????? ???????
?? ???????????? ??????????? ?????????????. ???????? ????????? ?????? ???
? ???????? ??????? ????????? ?????????
# 197 #
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?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
? ?????????? ?????? ????, ??? ? ??????
???????? (??????, 1994), ??? ????? ???????
? ?????? ???????????? ??????????? ???????????????. ?????? ???????? ????? ??????????? ??????? ? ???????????? ???????????????? ?????????? ??????????? ??????,
??????? ???????????? ??? ????????????????
?????????, ??? ? ?????????????? ??????????, ????????????? ? ???????.
??????????
??? ?????????? ??????????? (????????????????) ?????? ?????????? ????????????
???????? ???????? ? ??????? ????? ??????
??????? ????, ?????????????? ?? 75 %
??????????? ?????????????. ?? ???? ??????????? ??????? ???? ??????????? ??????
????????? ?????? ???. ??????????? ??????????? ????-??????????? ? ?????? ???????????, ??? ????????? ?????? ????????
????????? ?????? ???? ? ????????? ????????? (0,06 %), ????????? ??????????? ????????????? ?????????? ???????? ? ???????.
??????? ?????? ??????????? ??? ?????? ?
?????????? ???? ????????? ???????????? ???????????? ??????? ??????????????????
1,4 ?/?2, ??? ????????????? ????????? ??? ?
???????????? ????????.
?????? ????????? ??? ????????? ????????? ?????? ???? ? 11-04-00053.
?????? ??????????
??????????? ?.?. (2010) ????????? ????????? ????????? ? ??????? ??????????? ???? (?????????? ? ???????? ??????). ???????????: ??????. ??? ???. ???-?? ? ????????????, 228 ?.
??????????? ?.?., ????? ?.?., ??????? ?.?. (2006) ????????? ????????? ?????????, ?????????? ?????????? «?» ? ????????? ????????? ??? ?????????? ?????????????? ?????????
????????? ? ??????????? ??????? ??????????? ????. ???????? ?????????? ???. 1: ?. 41-47
?????? ?.?. (2000) ???????? ?????? ???????????????? ?????? ?????????. ???.: ?????,
147 ?.
?????? ?.?., ?????? ?.?., ???????? ?.?., ?????? ?.?., ??????? ?.?. (1993) ??????? ???
?? ????????? ? ???????????????? ????????? ????-??????????. ?: ??????????? ??????????????? ? ?????????? ????-??????????. ???.: ???????????????, ?. 299-311
????????? ?.?., ??????? ?.?. (1979?) ??????????? ????? ?????? ? ?????? ???? ??????????? ????????????. ?: ????????????????? ? ??????? ???????????? ????????????? ????? ?????????????? ????. ?.: ??? ?? ????, ?. 58-72
????????? ?.?., ??????? ?.?. (1979?) ??????????? ????? ?????? ? ?????? ???? ? ??????????? ???????? ?: ????? ?????? ???????? ?????? ?????????. ?.: ??? ?? ????, ?. 169172
????????? ?.?. (1998) ?????????????? ? ???????? ??????????????????? ???????????????? ? ?????????? ????. ???????????: ???????? ????, 160 ?.
?????? ?.?. (1994) ?????????????? ????????? ????????? ? ??????????? ???????????.
???.: ?????, 222 ?.
??????? ?.?. (1956) ????????????? ?????? ? ??????? ??????????? ???. ?????: ???, 253 ?.
??????? ?.?. (1960) ????????? ????????? ???????. ?????: ?? ????, 328 ?.
?????????? ?.?. (1980) ???????? ??????????? ????????? ???????????? ?? ????????????
?????????. ??????? ?? ????. ????? ????????. 6: 114-116
# 198 #
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?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? ?.?., ?????????? ?.?., ?????? ?. (1971) ??????????? ????. ?: ?????????????????????? ????? ??????????? ??????. ?.: ???????????????, ?.173-194
???????? ?.?., ?????? ?.?. (1993) ?????????? ??????????????? ????????? ??????
???????? ? ??????? ???-??????????. ?: ??????????? ??????????????? ? ?????????? ??????????????. ???.: ???????????????, ?. 283-299
????? ?.?., ????????? ?.?., ????????-????? ?.?. (2004) ?????????? ?????. ??????????????. ?: ?????????????? ??????????????????? ?????? ???????? ??????? ????. ?.: ???????
???, ?. 146-164,
???????? ?.?. (1971) ? ???????? ??????????????? ????? ???????? ? ???? ???? ?????????????? ???????????? ?: ???. ??? ?????. ? ????. ???. ???-??, ???. 6, ?., ?. 49-51
?????? ?.?. (1978) ??????? ??????? ?????? ? ?????????? ??????. ?????????. ????. 14
(4): 116-117
??????? ?.?. (1985) ????????? ???????????? ? ??????? ?????. ?.: ?????, 246 ?.
????? ?.?. (1977) ????????????????? ???????? ??????? ???. ????: ??????? ?????, 272 ?.
????? ?.?. (1986) ? ??????????? ??????? ??? ????? ??????????????. ?: ?????????????
????? ????????????. ??????, ?.6-19,
????? ?.?. (2004) ?????????? ?????. ??????????. ?: ?????????????? ??????????????????? ?????? ???????? ??????? ????. ?.: ??????? ???, ?. 168-172
??????? ?.?. (1969) ???????? ????? ? ??????????????? ????????. ?. 1, ?.: ?????, 438 ?.
????????? ?.?. (1986) ?????????????? ?????? ? ??????? ??????????? ???? ??????????
?????????????: ???????. ????. .. ???? ???? ????. ?., 34 ?.
????????? ?.?. (1989) ???????? ????????? ??????????? ????????? ? ???? Abramis brama.
????. ??????????. 29 (4): 624-633,
????????? ?.?., ???? ?.?., ?????????? ?.?., ??????? ?.?. (2001) ?????????? ?????? ?????????? ???????????? ?????. ?: ?????? ???????? VIII ?????? ??????????????????? ????????
??? ???, ?. ???????????, 16-23 ???????? 2001 ?. ?. I. ???????????: ??????. ??? ???. ???-?? ?
????????????, ?.16-18
?????? ?.?. (1989) ??????? ????????????? ??????? ????????????. ????? ????? ? ???????. ????? ????? ????????. ????????????. ?????????????, ?. 7, ?.: ??????, 145 ?.
??????? ?.?. (1985) ???????????????? ????????? ? ??????? ????????? ? ??????????? ??????? ??????????? ???? ? ????? ? ?? ?????????????? ??????????. ???. ? ?????????.
21,10,85; ? 714-??. ???????????: ??????. ??? ???. ???-?? ? ????????????, 225 ?.
??????? ?.?., ???????? ?.?. (1992) ???????????? ? ????????? ????????? ??????????? ?????? ?: ?.?. ????????? (???.) ????????????? ????????????????? ???????????? ? ??????????
?????? ??????????? ????. ???????????: ??????. ??? ???. ???-?? ? ????????????, ?. 14-52
???????? ?.?. (1970) ?????????? ????? ????. ?.: ?????, 744 ?.
??????????? ?.?., ?????? ?.?. (1981) ???????????? ???????????? ?? ????? ? ?????????
?????????? ??? ?????????????????? ????????????? ?? ???????????? ????????. ???????????? ? ??? ?????????. ?.: ????????, 32 ?.
????????? ?.?. (1982) ???????????? ???????????? ?? ?????????? ??????????? ???????
???????? ??????? ??? ? ??????? ?????? ????????? ?? ???????? ???? ? ???????????? ????????. ?.: ????????, 26 ?.
# 199 #
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?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? ?.?. (1958?) ????????? ???? Mesocyclops leuckarti (Copepoda, Cyclopoida). ??????? ?? ????. 120 (2): 419-421
??????? ?.?. (1958?) ????????? ?????? ?? ???????? ???????? ? ??????????? Acanthocyclops viridis (Copepoda, Cyclopoida). ??????? ?? ????. 119 (3): 613-616
??????? ?.?. (1959) ???????? ????? ???????? Acanthocyclops viridis (Jurine) ? Mesocyclops
leuckarti (Claus) (Copepoda, Cyclopoida): ???????. ???.? ????. ????. ????. ?., 26 ?.
??????? ?.?. (1998) ??????? ???????????? ??????????????. ?.: ???, 320 ?.
????????? ?.?. (2010) ?????????? ??? ????????. ????????? ??????. ?????????? ???????? ??????????? ?????????? (? 100-????? ?.?. ???????????). ?.: ???, ?. 283-304
???????? ?.?., ????? ?.?. (2008) ??????? ????-???????? ?ercopagis pengoi (Ostroumov)
?? ????????? ? ???????????????? ???????????? ??????????? ?????? ??????????? ????. ???????? ?????????? ???????? ?????? ???. 9 (5/1): 244-252
???????? ?.?. (2009) ??????????? ? ????????? ??????? ???? (?? ??????? ????????? ?
??????????? ??????? ??????????? ????). ???????? ?????????? ??? 1: 76-85
???????? ?.?. (2010) ??????????-?????????????? ??????????? ???????????? ??????????? ?????? ??????????? ????. ???????????: ??????. ??? ???. ???-?? ? ????????????, 198 ?.
???????? ????????????????? ??????????: ????????? ?????????? ???????????? (???) ?
?????????????? ?????????? ??????? ??????????? (????) ??????? ??????? ??? ???? ??????
????????, ??????? ????????????????? ???????? (1999). ?.: ?????, 304 ?.
??????? ?.?. (2000) ????????????? ??????????? ???????????? ????? ? ??????? ????
??????-?????? ? ?? ?????????????. ???.: ????????, 223 ?.
????????? ?.?. (2000) ???????? ???????? ? ??????????? Marenzellaria viridis ? ?? ??????? ?? ????????? ?????????? ?????????? ??????????? ??????. ?: ????-???????? ? ??????????? ????? ??????. ???????, ?. 193-202
???????? ?.?., ?????? ?.?., ?????????? ?.?., ??????? ?.?. (1984) ???????????? ???????????? ?? ????? ? ????????? ?????????? ??? ?????????????????? ????????????? ?? ???????????? ????????. ????????? ? ??? ?????????. ?.: ????????, 51 ?.
???????? ?.?., ??????? ?.?., ??????????? ?.?. (1984) ???????????? ???????????? ??
????? ? ????????? ?????????? ??? ?????????????????? ????????????? ?? ??????? ????????.
??????????? ? ??? ?????????. ?.: ????????, 33 ?.
???????? ?.?., ????????? ?.?. (1998) ??????????? ????????? ?????????? ??????????????? ???????. ?: ??????????-????????????? ???????????? ?????????? ? ??????????
???? ? 1996-1997 ?????. ???????????: ??????.??? ???. ???-?? ? ????????????, ?. 96-129
????? ?.?., ??????? ?.?., ????????? ?.?. (2004) ????? ?????????????? ???????????
?????? ?: ?????????????? ??????????????????? ?????? ???????? ??????? ????. ?.: ???????
???, ?. 17-18
????? ?.?. (1971) ?????? ??????. ?: ?????????????????????? ????? ??????????? ??????. ?.: ???????????????, ?. 143-164
???????? ?.?. (1971) ?????????????? ????????? ? ??????? ??????. ???????????????? ?????????????? ?????? ? ??? ????????? ?????. ?: ?????????????????????? ????? ???????????
??????. ?.: ???????????????, ?. 6-9
?????? ?.?. (1972) ????????????? ??????? ????????????. ????: ??????? ?????, 195 ?.
# 200 #
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?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? ?.?. (1979) ? ??????? ???????? ????????????? ? ??? ??????????. ?.: ???, 166 ?.
????????? ?.?. (1982) ???????? ???????? ??????? ???? ????????? ?????? ???????????
????. ?: ??????? ? ??????? ????????? ??? ? ?????????????? ?????????????? ??????. ??????.
???????????: ??? ???. ???-?? ? ????????????, ?. 75-80
????????? ?.?. (1988) ??????? ???? (Anguilla anguilla) ? ?????????? ?????? ???????????
????. ?: ????????????? ? ????????????????? ???????????? ? ????????????? ?????? ? ???????????? ????? ?????? ??????. ???????????: ??????. ??? ???. ???-?? ? ????????????, ?.
97-107
????????? ?.?. (1992?) ??????? ????????? ???????????? ??? ? ?????????? ??????. ?:
????????????? ? ????????????????? ???????????? ? ?????????? ?????? ??????????? ????.
????????? ?.?. (???.). ???????????: ??????. ??? ???. ???-?? ? ????????????, ?. 165-188
????????? ?.?. (1992?) ??????? ?????? ??? ? ?????????? ?????? ??????????? ????.
??????? ??????????. 32 (3): 171-176
????? ?.?., ????? ?.?. (1999) ?????????????? ?????? ??????? ?????????? ?????? ??????????? ???????. ?: ????????????? ??????? ???????????, ?????????. ???????????, ?. 96
????? ?.?., ????????? ?.?., ???????? ?.?. (2004) ???????? ???????????? ?????? ??????
? ??????????????? ??????? ?????? ? ????? ? ????????? ????? ? ??????????? ???????????????? ?????. ?: ?????????? ???? ????: ????????????????? ? ???????????. XXXI ????????????? ??????? ???????????, ?????????. ???????????, ?. 178-180
????? ?.?., ????????? ?.?., ???????? ?.?. (2005) ? ???? ?????????????? ??????? ? ???????? ?????????? ????????? ? ?????????? ??????. ?: ???????? ????? ? ???????. ?VI ????????????? ??????? ????? ?? ??????? ????????, ?????????. ?.2, ?., ?. 237-238,
????? ?.?. (2006) ???????? ???????????? ????????????????? ? ?????????? ?????? ??????????? ????: ???????. ???? ????. ???????-?????????. ????, ???????????, 23 ?.
????????? ?.?. (1994) ?????? ??? ?????????? ??????????? ????????? ???????????????? ?
????????? ???????. ????????? ? ????????? ???????. 3: 33-40
????? ?.?., ????????? ?.?. (2003) ????? ??????????? ???????? ?????? ? ???????? ?
????????????? ?????????? ????????. ?: ?????? ??????? ???????? ??????????????? ????????
(??????????????? ?????????). ?. 2, ???????????: CD-ROM ??????, 12?-1 ? 12?-32
??????? ?.?. (1952) ??????? ? ??????? ??????????????? ??? ??????????? ????. ?.:
?????????????, 268 ?.
??????? ?.?. (1992) ??????????? ????????? ?????? ??? ? ?????????? ??????. ?: ????????????? ????????????????? ???????????? ??????????? ????. ???????????: ??????. ???
???. ???-?? ? ????????????, ?. 151-165,
Elliott J.M., Persson L. (1978) The estimation of daily rates of food consumption for fish. J. Animal. Ecol. 47 (3): 977-991
Hakanson L., Buolion V.V. (2002) The lake food web. Modeling predation and abiotic/biotic interaction. Leiden: Backhuys Publishers, 344 p.
Haney J.F., Hall D.J. (1973) Sugar-coated Daphnia: A preservation technique for Cladocera.
Limnol. Oceanogr. 18 (2): 331-333
Nie H.W. de (1987) Food, feeding periodicity and consumption of the eel Anguilla anguilla
(L.) in the shallow euthrophic Tjeukemeer (the Netherlands). Arch. Hydrobiol. 109 (3): 421-443
# 201 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
Semionova S.N. (2003) Peculiarities of phytocene state in the Vistula Lagoon of the Baltic Sea in
the current conditions. In: Abstract Publication Baltic Sea Science Congress. Helsinki, 24-28 August,
2003, P. 56
Energy Flows in the Vistula Lagoon
of the Baltic Sea
Elena N. Naumenko,
Mikhail M. Khlopnikov and Liliya V. Rudynskaya
Atlantic Scientific Research Institute of Marine Fisheries
and Oceanography (AtlantNIRO)
5 Dm. Donskoy St., Kaliningrad, 236000 Russia
Energies of the Vistula Lagoon ecosystem is estimated based on observations made by authors in
the northern Vistula Lagoon literature data. The pelagic and bottom communities are represented
by euryhaline species: phytoplankton ? 412, zooplankton ? 74, zoobenthos ? 43, fish ? 54, Mean
biomasses during the vegetation season (April-October) were: 6,2 (8,7 kJ) of phytoplankton, 2,5
(5,2 kJ) of zooplankton, 20,6 (61,8 kJ) g/m2 of benthos. Yield of aboriginal fish amounted to 1,4 tons/km2
(60 kJ/m2).
Annual primary production was 14000, pure -3900; zooplankton and benthos production during
vegetation period amounted to 300 and 1100 kJ/ m2, respectively. Ratios of production of planktonic
crustaceans, benthos and fish catch to primary production were 2,3, 8,5 and 0,06 %, respectively.
Yield of aboriginal fish is formed mainly due to detritus food chain.
Keywords: Vistula Lagoon, production, fish yield, food chain, energy flows.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 203-209
~~~
??? 579.68 (285.23)
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? ?????? ???? ????????
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???????? ???????? ??. ?.?. ??????????? ?? ???,
?????? 664033, ???????, ?????????? 1?
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????????? ??????????????? ???????????,
?????? 664003, ???????, ?????? 3 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
??????????? ???? ????-????????? ????????????? ? ?????? ?. ???????? ?? ??????????????????
???????????. ???????? ????????????? ?????? ?????????? ??????????? ? ??????? 198090-? ??. ????? ??????????? ??????????????? ?????????? ?? 0,9 ?? 6 ??? ??/??, ????????? ??
?????? ??????? ???, ??????????? ??????????? ??????????????? ??????????? ?? ???????.
??????? ???????? ??????????? ?????????????? ???????? ? ????? ?. ???????? (5240 ???/
??) ? ??????? ?????? ?. ???????? (10700 ???/??). ???????????? ???????? ???? ?????????? ?
??????? ?????? ?. ???????? (11333 ???/?). ?????????? ???????????????????? ???????????????
??????????? ?? ??? ??????? ? ????????? 1-9 ???/??. ????????????????????????
?????????????? ?????????? ?????? ? ?????? ?. ????????. ???? ?????? ?. ???????? ?
?????????? ????????? ????-????????? ????????????? ?? ?????????????????? ???????????
????????????? ??????????? ?????? 2.1.5-980-00 ? ????? ???? ???????????? ?????? ???
????????????? ?????.
???????? ?????: ????-???????? ?????????????, ???? ????????, ?????????-??????????????????
?????????? ????.
????????
??????????????????
?????
????????????? ????????????? ??? ??????????
*
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? 1980-90-? ??. ? ???????? ??? ????????????
(???????????, 1994). ????????? ????????????
? ???????????????? ????????????? ?????????
Corresponding author E-mail address: ingleb83@mail.ru
© Siberian Federal University. All rights reserved
# 203 #
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????????? (???) ?? ??????? ???????????
??????? ????????? ????? ????-????????? ????????????? (??????? ? ??., 1982; ???????,
??????????, 1983, ??????????, 1983?; ???????????, 1985, 1995; Vinogradova,1998).
?? ??????????? ???????????? ???????????? ????????????? ???? ???????? ?
???????????? ??????? ? ???????????????? ????????? ? ??????, ?????????? ? ???????????? ?????? (??????????, 1982, 1983?,
1984).
? 2000-? ??. ???? ???????????, ??? ????????????? ???????? ?? ??????? ???????????
??????? ???????? ???????????. ??????????
????????????? ??????????? ????????, ???
??? ???? ?? ?????? ?????????, ?? ? ?????????
???????????. ?? ???????? ??????? (???????????, 2010), ? ?. ???????? ? ????-???????????
????? ?? ????? ???????????????? ????????? ??? ???????? 450 ???. ? ????????? ??-
?????. ??????????? ? ?????? ??????????
????????? ??????? ? ?? ???????????????
?????????? ???????? ? ?????????? ???????????? ??????????? ?????, ? ?????????????, ?
????????? ?????????? ?????????.
???? ?????? ? ??????????? ???? ????????????? ????????????? ? ?????? ?. ???????? ?? ?????????????????? ???????????,
???????? ????????????? ?????? ??????????
??????????? ? ??????? ??????? ??? ? ??????? ?????????, ???????????? ? ??????????.
????????? ? ??????
? ???? 2005 ?. ???? ???? ????????? ???????????? ??? ????-????????? ????????????? ? ?????? ?. ????????. ?? 9 ????????
???? ???????? 16 ???? ???? ? ?????????????? ? ?????????? ?????????? (???. 1). ????????? ????? ???????? ? ?????? 13?33 ?.
??????????? ????? ??????????????? ???????
??????????????? ???? ?????????? ??????????, ???????? ? ????? ????. ???????????
???? ? ?????? ???????????? ?????????? ??
6 ?? 18 є?.
7
?
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???. 1. ?????-????? ?????? ????. ???????: 1 ? 4,5 ?? ???? ?????? ?. ???????? (? ?????? ??????); 2 ? 4,5
?? ???? ?????? ?. ???????? (? ??????? ??????); 3 ? ???? ????????, 1,5 ?? ?? ???????; 4 ? ?????? ?????? ?.
????????; 5 ? 6,5 ?? ?? ??????? ?????? ?. ????????; 6 ? 0,5 ?? ???? ?? ?????? ?? ????? ?????? ?. ????????;
7 ? ????? ??????; 8 ? ?????? ?. ???????? ? ?????? (????????); 9 ? ????? ?????? ?. ????????
# 204 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
?????????-?????????????????? ???????????? ???????? ??????????? ?????????
???????????: ????? ??????????? ??????????????? (???), ?????????? ???????????
(??), ???????????????????? (???), ??????????????????? (???) ???????????????,
?????????? ???????? ?????? ???????? ??????? (????).
??? ?????????? ?? ?????????? ???????? «Millipore» ? ????????? ??? 0,22 ???.
??????? ?????????? ??????????? ? ???????????? ??? ??? ??????????? ? ??????????? ? 1350 ??? (?????????, ????????, 1974).
?????????? ??????????? ???????????????
?????????? ?? ????? ???:10 (????????? ????? ???????????? ??? ??????????? 18-20 є? 10
???). ?????????-????????????? ????????? ???
?????? ?. ???????? ????????? ?? ??????????
????. ?????????? ????????????????? ?????? ????? ??????????? ???????? (??? ???)
?????????? ??????? ?????????? ????????
(???? 24849-81). ????? ???? ???????????
????? ??????? ? 8 «????????» («?????????»,
??????), ??????? ??????????? ??? 37 єC ??
??????-?????????? ????? ???? 24 ?. ??? ??????? ?? ???????? ???????, ??????????? ???
????, ?? ?????????? ??????? ??????? ????
???????? ?????, ?????????? ?? ?????, ????????????????. ??? ??????? ? ?????? ?????????????????, ????????, ?? ??????????
???? ??????? ????????? ?????????? ????. ????????????????????? ??????? ????????????
? ???????????? ?? ?????????????? ? ?????
??????????? ????????? ??????? ????-????
????????????? ??????? ??????? ???? ?? ?????????? ????? ? ????????. ?????? ???????????? ??? 37 є? 24 ?. ??? ???????????
??????? ? ???? ????????? ??????? ?????????????. ????????? ???????? ? ???? ????-???????
(?-?): ??? ??? ? 1 ? ????????????? ????.
??????????
????????????????????
??????????????? ???????????? ?? ???????-
?????? ????? ??????????. ?????????? ??????????????????? ???????? ?????????? ??
????? ????????-???????? (?????????, ????????, 1974).
?????????? ? ??????????
?????????? ???????????? ????????,
??? ????? ??????????? ??????????????? ?
?. ???????? ? ?????????? ????????? ????????????? ????????????? ?????????? ??
0,9 ?? 6 ??? ??/?? ? ????????????? ????,
????????? ?? ?????? 1980-90-? ??. (0,8-4;
1-10,4 ??? ??/?? ??????????????) (??????????,
1984; ???????????, 1994). ? ????? ?. ????????,
? ??????? ?. ???????? ? ?????? ? ?????? ???????????? ??? ???? ?? ?????? 2 ??? ??/??.
????????? ???????? ???? ??????????? ?
?????? ?????? ?. ???????? ? ?? ??????? 13 ? ?
6,5 ?? ?? ??????? ?????? ?. ???????? (???. 2).
?????????? ??? ? ??????? ???????????
??????? ?????????? ? ? 1980-90 ??. ? ???? ??????? ? ??????? ????????????? ????????????
??????? (???????????, 1994).
?????????? ?? ? ????? ?????? ?. ???????? ? ??????????? ? ?????? ?????????
????-????????? ????????????? ??????????
?? 1230 ?? 10745 ?? ??????????? ? ?? 815 ??
4500 ???/?? ? ????????? ????????? (???. 3).
???????? ?????????? ?????????? ???????????? ??????? ???????????????? ? ?????
?. ???????? ? ? ?? ?????? ??????????? ? ????????????? ??????????. ?????? ? ???? ????????? ??????, ?????????? ? ?????? ??????
? ? ????????? ????-????????? ?????????????, ??????????? ? ?????? ????????, ?????????? ?? ???? ????, ??? ? ?????????????.
????????, ? ????-???????? ?????????????
?????????? ???????????? ???????????? ???????, ??????????? ?? ?. ????????, ???????
??????????? ?????????? ?? ? ?????????
??????????. ???????????? ???????? ??????????? ??????????????? ????????????? ?
# 205 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
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6000
5000
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???. 2. ????? ??????????? ??????????????? ? ?????? ?. ???????? ? ????-???????? ?????????????
? ???? 2005 ?.: 1-9 ? ??????? ?????? ????
12000
???, ???/??
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???. 3. ?????????? ??????????? ??????????????? ? ?????? ?. ???????? ? ????-???????? ?????????????
? ???? 2005 ?.: 1-9 ? ??????? ?????? ????
??????? ?????? ?. ???????? (??. 4). ???? ?????
???????????????? ????????????? ?????????? ?????????? ???????? ??????? (?? 2600
???/?) ? ????????????? ?? ??????? (???????????, 1984).
???????? ?????????? ??????????? ?
????????????? ?????? ???????? ?. ????????,
???? ???????????? ????? ??????? ??? ????????? ??? ? ????????????-??????? ??? ?.
???????. ??????? ???????? ?????????? ??,
?????????? ? ????? ?. ???????? (5240 ???/??)
? ??????? ?????? ?. ???????? (10700 ???/??),
??????????????? ? ????? ??????????? ????????? ?????????? ???????????? ??????? ?
?????? ???????. ??? ???????? ?? ??????? ?????? ?. ???????? ? ?????? ?????? ?????????????? ???????? ??????????: ?? ???? ????????
?????????? ?????????? ????????? ?? ???????
(1300-1600 ???/??). ???????? ??????? ????????????? ?? ?? ?????? ??????? ???????-
# 206 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
12000
?-?, ??? ???/?
10000
8000
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6
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????????? ????????
???. 4. ????-?????? ? ????? ?????? ?. ???????? ? ????-???????? ????????????? ? ???? 2005 ?.: 1-9 ?
??????? ?????? ????
???? ? ? 1980-? ????? (??????????, 1984). ?????? ?? ????????? ? 1980-?? ??. ???????????
?? ? ?????? ??????????? ?? ???????.
?????????? ???? ?????????? ?? 166 ??
11333 ???/? ? ????????????? ? ?? 0 ?? 5666
???/? ? ????????? ?????????? (???. 4). ???????????? ???????? ???? ?????????? ?
??????? ?????? ?. ???????? (11333 ???/?).
??? ???????? ?? ??????? ?????? ??????????
???? ??????????? (? 6,5 ?? ?? ??????? ?????? ?. ???????? ? ? ?????? ?????? ???? ?????????????), ? ? ????? ????????????? ?????
?????????????. ??????? ????????, ??? ? ????? ?????? ?. ???????? ?????????? ???? ?
????????????? ?????????? ???? ????, ??? ?
?????????, ? ? ????????????? ???????????
??????????????? ???????: ?????????? ????
????????????? ? ????????? ??????????. ??????????, ? ????????????? ???? ????????
????????????? ?????? ????, ??????? ???????? ? ?????????? ?????????? ???? ? ????????? ??????????. ??? ?????????? ????, ??
???? ?? ?????????? ?????? ??????????? ??????????? ????????, ??????? ??????????????? ? ???????????? ?????????? ????????????
???????.
?????????? ??? ? ????? ?. ????????, ??
?????? ? ????? ????????????? ????????? ??
1 ?? 9 ???/??. ? 1980-? ??. ?? ?????? ???????
?????????? ??? ?????????? 8-20 ???. ???/
?? (??????????? ? ??., 2004). ?????????? ?????????? ??????????????? ???? ??????????????? ?????? ??????? ? ?????????? ????????? ?????????? ?????????, ??????? ?????
???? ??????? ? ?????????? ???????? ???????? ????????? ??????? ?????????, ??????? ?
???????? (???????????, 2010).
? ????????????? ???? ?????????? ????????????????????????
??????????????
(???) ?????????? ? ???? ?????? (???????? ?
????? ?????? ?. ????????). ?? ??????, ?????????? ? 1980-? ??., ?????? ?? ???? ???????? ???? ????????????? ??????? ???????????
??????????????? ???? ??????, ??? ??? ? ????
??????? ??????????? ???????????? ???????
?????????? ? ???????????????? ??????????
????????????? ? ????????? ?????????? ?????????? 26-59 % (???????????, 1994).
?????????-?????????????????? ?????????? ??? ?. ???????? ? ?????????? ????????? ????-????????? ????????????? ??????????????? ? ?????????? ?????????????
# 207 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
???????? ?? ?????????? ? ?? ???????????
????????? ????????????. ???? ?????? ?.
????????, ?????? ??????, ?????????? ????????? ????-????????? ????????????? ??
?????????????????? ??????????? ????????????? ??????????? ?????? 2.1.5-980-00 ?
????? ???? ???????????? ?????? ??? ????????????? ?????.
?????? ??????????
??????? ?.?., ?????????? ?.?. (1983) ??????? ??????? ??? ????????? ????????????????? ????????? ?? ?????? ?????????????? ????-????????? ?????????????. ?: ?????? ??????? ?? ??????????? ????????????? ????????? ??????????? ??????????-???????? ??????????????. ?., ?. 105-112.
??????????? ?.?. (1985) ???????? ?????????? ????????????????? ????-????????? ????????????? ? 80-? ???? ?: ?????????? ???????? ? ??????? ? ????????. ???. 3. ???????, ?. 11-12.
??????????? ?.?. (1994) ?????????????????? ????? ????-????????? ????????????? ?
???????? ??? ????????????: ???????. ???. ? ????. ????. ????. ???????, 25 ?.
??????????? ?.?. (1995) ?????????? ?????????? ????-????????? ????????????? ??
?????????????????? ???????????. ???????? ????????. ???????? ????????: ?????????
V ????????. ????.: ?????? ?????? ?????????? ?.?. ??????. ???????????: ?? ?????, ?. 2,
?. 248-250.
??????????? ?.?., ?????? ?.?, ??????? ?.?., ?????????? ?.?., ????????? ?.?. (2004) ?????????????????? ???????? XX ????. ?. 1. ????? ???????? ???????-??????-?????????? ??????????. ???????: ???-?? ??-?? ????????? ?? ???, 93 ?.
???? 24849-81 ???? ????????. ??????? ?????? ?????????-??????????????????? ???????.
??????????? ?.?. (1984) ? ???????????? ???????????? ??????????????? ? ??????????
????????????? ???????? ???? ?????? ? ????????? ????????????. ?: ???????????? ?????????
???????? ????? ?????? ? ????????? ????????????. ???????, ?. 63-69.
?????????? ?.?. (1982) ????????????? ??????????? ?/??2 ???????????????? ?. ????????? ? ????-????????? ?????????????. ?: ???????? ???????? ???????????: ??? ????. ? ????.
????. ????. ???????, ?. 2, ?. 52-58.
?????????? ?.?. (1983?) ???????????? ???????? ???????????? ??????? ???. ?: ???????? ?????? ???????: ???. ????.-?????. ?????. ?????????, ?. 13-14.
?????????? ?.?. (1983?) ???????? ??????? ??????? ??? ????????? ??? ?? ????????????
????????? ?????? ????-????????? ?????????????. ?: ?????? ??????? ???????? ??????? ?
??????. ????????????, ???????????? ????????????? ? ??????: ???. ????. ????.-?????. ?????.
???????, ?. 143-145.
?????????? ?.?. (1984) ????????? ??? ????-????????? ????????????? ?? ?????????????????? ???????????: ???????. ???. ? ????. ????. ????. ????, 24 ?.
??????????? ?.?. (2010) ?????????????? ????????????? ??????? ??????? ???
??????????-???????? ??????????? (?? ??????? ????????? ???): ???????. ???. ? ????. ????.
????, ???????, 20 ?.
????????? ?.?., ???????? ?.?. (1974) ???????? ??????????????? ??????? ????????. ???????????? ???????????. ?.: ?????, 194 ?.
# 208 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
??????? ?.?., ????????? ?.?., ?????? ?.?., ?????? (???????????) ?.?. (1982) ???????????? ?????????? ????-????????? ????????????? ?: ???????? ???????? ???????????. ???????, ???. 1, ?. 34-35.
?????? 2.1.5-980-00 2.1.5. ????????????? ?????????? ????, ?????????? ?????? ??????
????????.
Vinogradova T.P. (1998) Long-term investigation of bacterioplankton of the Ust-Ilimsk reservoir.
In: BICER, BDP and DIWPA Joint International Symposium on Lake Baikal, Yokohava, ?. 111.
Characteristic of Water Ust-ilim Reservoir
and Bay of the River Vikhoreva
by Microbiological Indicators
Inessa V. Glebovaa,
Vyacheslav V. Maximov and Elena V. Schetininab
a
Institute of Geochemistry of A.P.Vinogradov SB RAS,
1A Favorsky st., Irkutsk, Russia 664033
b
Institute of Biology of Irkutsk State University,
3 Lenin st., Irkutsk, 664003 Russia
b
Ust-Ilim reservoir and the bay of the Vikhoreva River on microbiological parameters were investigated.
A comparative analysis of the results obtained with these of 1980-1990 years was carrying out. The
total number of microorganisms ranged from 0.9 to 6 million cells/ml, remaining at the level of previous
years. The number of saprophytic bacteria decreased in the order. High values of saprophytic bacteria
identified in the waters of the Vikhoreva River (5240 CFU/ml) and backwater bay Vikhoreva (10700
CFU/ml). The maximum values of coliforms were found in backwater bay r. Vikhoreva (11333 CFU/l).
Number of cellulolytic microorganisms has decreased by three orders of magnitude and amounted
to 1-9 CFU/ml. Sulfate-reducing bacteria were found only in the bay of r. Vikhoreva. Water of bay of
r. Vikhoreva and adjacent waters of the Ust-Ilim reservoir on the microbiological data comply with
SanPiN 2.1.5-980-00 and can be used only for recreational purposes.
Keywords: Ust-Ilim reservoir, river Vikhoreva, sanitary and microbiological parameters of water.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 210-215
~~~
??? 574.4
??????? ?????????
?????????? ?????? ?? ??????????????????????
? ???????????????? ????? ????
?.?. ?????,?*, ?.?. ???????????,
?.?. ?????????,?, ?.?. ????????,?
?
???????? ????????? ?????????? ?????????
?????????? ???????? ????,
?????? 660036, ??????????, ????????????? 50, ???. 50
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ????????? 79 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
??????? ??????????? ?????????? ?????? ?? ????(3-????????????????) ? ????(3????????????????-??-3-????????????????) ? ???????????????? ????? ???? ?? ?????????
????????: 3, 9, 13 ? 20 ?, ??????????????? ?????????? ????????????? ?????. ????????, ???
???????? ??????? ?????????? ?????? (???????? ????? ?????? ? ???????????? ????? ????????)
?????????? ? ??????????? ?? ??????? 3 ?, ? ???????? ???????????? ? ?????????? ?????????
?????. ?????????? ?????? ????????????? ????? ? ????????? ?? ??????? 13 ? ? ?? ??????? 20 ? ?
??????????????, ??? ???????? ?????????? ?????????? ?????????, ??????????? ????????????
? ?????? ???????????.
???????? ?????: ????? ????, ?????????????????????, ??????????, ???????????? ?????.
????????
????? ??????? ??????????????? ? ????????? ????? ??????????? ??????????
?????????? ???????????? ??????? ????????, ????????????? ????????????????, ???
?????????? ????????????????????? (???)
(?????? ? ??., 2006; Jendrossek, Handrick,
2002, Wang et al., 2005). ???? ?? ???????????
?? ?????????? ? ?????? ????????????? ?????????, ????????????? ??????? ??????? ?
*
1
?????????? ?????? ????????? ???? ????????????? ????????? ? ????????? ???????? ??????, ???????? ? ?????????? ???????.
? ?????? ?????????? ?????????? ??? ???
??????? ???????????? ??????????? ???????????? ??????????????? ?? ?????????? ?
????????? ????????. ????? ?????? ???? ???????????????. ???????????? ??????????
? ?????????? ??? ? ??????? ????, ? ?????
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Corresponding author E-mail address: nzhila@mail.ru
© Siberian Federal University. All rights reserved
# 210 #
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?.?. ????, ?.?. ??????????? ?????????? ?????? ?? ?????????????????????? ? ???????????????? ????? ????
?????? ??????, ??? ?????????? ??????????
3-????????????????-??-3-????????????????
? ??????? ???? ?????????? ????? ????????
(Imam et al., 1999). ?????? ???????? ?????????? ??? ? ??????? ???? ??????? ? ????????
?????? (Wang et al., 2005). ??? ?????????????
?????? ?? ????????? ?????????????? ???????? ????????? ?????????? ??? ? ?????????
????? ? ?????? ?? ??????????. ????????? ?
??????? ???? ???????????? ?????????????
??? ? ????????? ??????? ???????? ???????,
??? ????????????? ??????? ?? ????????? ?????? ?????????? ? ???????? ???????, ? ????? ?????????? ????????? ??? (Volova et al.,
2007). ???????????? ?? ?????????? ??? ???
???? ?????????? ? ????? ???? ?? ??????????? ???????????? ?? ???????????. ??? ???
????????? ?????????????? ???? ???????????? ? ???????? ???????????? ?????? ?? ???
? ???????????????? ????? ????. ? ?????? ?????? ??????? ?????????? ???????? ????????????? ???????? ??? ? ?????? ?? ??????????? ???????? ?? ????????, ???????????????
?????????? ????????????? ?????.
????????? ? ??????
????? ???? ? ??? ???????????????? ??????, ???????????? ???????? ???????? ????????? ????????????? (Kalacheva et al., 2002;
Zadereev, Tolomeev, 2007). ??????? ??? ????
????? ? ?????????? ????-3-???????????????
?3?? ? ????????? 3-???????????????? ?
3-???????????????? ?(3??-??-3??), ????????? 3?? 10 ???. % ? ? ???? ????????? ?????? ????????? 30 ??, ?????? (70±5) ?? ????
????????? ? ???????? ?????????? ????????
?? ??????????? ??????? ????? ?? ?????????? 2 ?? ?? ?????? ? ????? ? ????????????
54.30.350 N, 90.11.350 E ?? ????????: 3, 9, 13
? 20 ?, ??????????????? ?????????? ????????????? ?????. ??????? 3 ? ?????????????
??????????? (?????? ????????????? ????,
?????? ??????????? ??????? 20 ??, ?????????? ????????); ??????? 9 ? ? ????????????
(???????? ????, ?????? ??????????? 6 ??,
?????????? ????????); ??????? 13 ? ? ????????? (??????????? 2 ??, ???? ???????? ?? ?????????? ? ???????????? ????????); 20 ? ?
?????????????? (??????????? ????? 1 ??,
?????????????????? ???? ??? ??????????,
???????????? ????????). ?????????? ??????????????? ??????? ???????????? ?????????????????. ? ???? ????????? ???????????
???????? ???????? ????????? ????????, ?
????????? ????? ? ?????????? ??????????? ?
??????????????????? ????????, ????????
???????? ???? ? ????????? ?????????????
?????????? ????????????? ???????? (??????? ? ??., 2003). ??????????? ??? ???????? ?
11 ???? ?? 29 ???????; ?????? ?????????? ?
80 ?????. ????????????? ????????? ?????
?????????? ???????? ? ?????? ???????????
? ?????????? ??????????? ? ?????? ?????????, ??????? ???????? ????????? ?????? YSI
6600 (YSI Corp, ???).
???????????? ????? ? ??????????????????? ????????????? ?????????? ??????
??????????? ? ?????????????? ????????????
??? ??????????????? ????????????? 1260
Infinity (Agilent Technologies, ???) ???????????? ?????????????? ?????????? («Fluka»,
?????????, ????????). ???????? ????????????? (M ?) ? ?????????????? (M?) ???????????? ?????, ? ????? ????????????????
(?? = M ? /M?).
?????????? ? ??????????
? ??????? ??????? ?????? ?????????????? ????????? ????????????? ???????. ?
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???????????? ??? ?????????? 13 ??, ? ???????? ???? ????????? 23-24 ??. ?? ???????
9 ? ??????????? ???? ?????????? ???????
# 211 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
% ?? ???????? ?????
?.?. ????, ?.?. ??????????? ?????????? ?????? ?? ?????????????????????? ? ???????????????? ????? ????
?????????? ? ?? 12 ?? 18 ?? ? ??????? ???????????? ???????. ? ????????? ? ?????????????? ??????????? ???? ??????????? ??
?????????? ? ? ??????? ?????????? 5,0-6,0 ?
1,5-2,0 ?? ??????????????. ????????? ???? ??????????? ?? 14-15 ?/? ?? ??????? ??????????
?? 16-18 ?/? ? ???? ?????????; ???????? ??
?????????? ?? 8,9 ?? 8,5.
???????? ????????? ????? ???????? ??
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???????? ? ??????????? ????????, ????????? ? ??????????? ????????? ??? ????????? ???????? ??? ????? ?????? ? ??????? ???????????????? ?????????, ??????????
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????, ??????????????? ?? ????????? ????????:
1 ? ?3??, 2 ? ?(3??-??-3??)
?????? ?????????? ????????. ???????????
???????? ???? ???????? ???? ?????? ???
???????????? ????????????? ??? ? ????????? ???????????? ????????? ??????????? ?
????????? ??????? ???????? (?????? ? ??.,
1996; Volova et al., 2007).
??????????? ?????????? ????? ??????
???? ????????????? ?????? ? ?????? ???????. ???????? ??????? ??? ??????????? ??
??????? 3 ?, ??? ????? 50 ????? ?????????????? ????? ?????? ?? ???????????? (?3??)
??????????? ?? 9,0 % ?? ????????, ?????
???????? ?? ?????????? ? ?? 15,8 % (???. 1).
?? ???????? 13 ? 20 ? ????? ?????????????
???????? ????? ????????. ? ????????? ? ?????????????? ? ????? ??????? ??????????
(80-? ?????) ???????? ???????? ????? ?3??,
??????????????, ?? 8,8 ? 9,2 %; ?????????? ?
????????? ????, ??????????????, ?? 13,3 ?
# 212 #
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?.?. ????, ?.?. ??????????? ?????????? ?????? ?? ?????????????????????? ? ???????????????? ????? ????
16,2 % ?? ???????? ???????. ?????? ? ??? ??
??????? 9 ? ???????????? ????????? ?????
???????? ?????? ?? ??? ????? ????? ?? ??????????. ???????? ????? ?????? ???? ?????
???????? ??? ?(3??-??-3??). ??? ??????????? ? ????????????, ??????????? ????? ? ??????? ???????? (??????? ? ??., 2007; Volova et
al., 2007).
??? ???????? ?????????? ????????????,
???????? ??????? ?????????? ?????? ??????????? ? ??????????? ? ?????? ???????????? ? ?????????? ????????? ?????. ????????,
??? ? ???? ????????? ?????????????? ??????? ??????????? ????????????????? (??????? ? ??. 2003). ????????? ???????? ???????? ?????????? ???????? ? ??????????
????? ?????? ? ?????????????? ???????????
????????, ????????, ? ??????? ??????????
??????? ????????? ????????????? ???????
??? ?????????? ?????? ?? ???. ?? ???????
8-12 ? ? ????? ???? ????????? ???? ????????? ??????????? ? ???????????? ????????????? ?????????? ? ???????? ?????????????
? ??????? ?????????? (Gaevsky et al., 2002).
?????????? ???????? ????? ????? ??????
?? ??????? 9 ? ??????????? ?????????????
??????????? ??????????????? ? ?????????
????? ????????????.
?????????????, ??? ?????????? ???????? ????????????? ????? ? ????????? ??
??????? 13 ? ? ?? ??????? 20 ? ? ??????????????, ??? ???????? ?????????? ??????????
?????????, ??????????? ???????????? ? ?????? ???????????. ? ????? ? ???? ??????????
???? ?????????? ????????? ?????????? ???
? ?????????? ?????.
????? ?????????????? ????? ?????
?????????? ???????? ????????????????
????????????? (??), ?????????????? (??)
???????????? ?????, ? ????? ??????? ???????????????? ???????? (??), ??????????
?????????? ??????? ???????? ??????????
?????????? ????? (???. 2). ?? ???? ??????????, ?? ??????????? 9 ?, ????????????? ???????? ?? ? ??, ??? ???? ????? ???????????? ?
?? ??????? 3 ?, ??? ??????????? ? ?????????
?????????? ????? ????????. ????? ????????
???? ???????? ???????????? ????? ??? ???????? ?????????? ?(3??-??-3??) ?? ????????? ? ?????????????.
????? ???????, ??????? ????????? ???????????? ?????????? ?????? ?? ?????????????????????? ? ???????????????? ?????
???? ? ?????? ??????????????? ? ?????????????????? ????????????? ?????. ????????,
??? ????(3-???????????????) ? ????? ????????????? ????????? 3-????????????????
? 3-????????????????? ??????????? ? ??????
??????????????. ???????????? ????? ????? ?????????? ?????? ? ???????? ?? ? ??
???????????????? ? ?????????? ???????????. ????????? ??????? ?????????? ????????
????????? ? ????????? ? ???????????? ?????????????????? ??????????????.
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?.?. ????, ?.?. ??????????? ?????????? ?????? ?? ?????????????????????? ? ???????????????? ????? ????
???. 2. ????????? ????????????? (M ?) ? ?????????????? (M?) ???????????? ????? ?????? ?? ??? ?
????? ????, ??????????????? ?? ????????? ????????
?????? ??????????
??????? ?.?., ??????????? ?.?., ???????? ?.?., ?????? ?.?. (2007) ?????????? ???????????? ? ????????? ???????? ? ?????????? ?????????????? ????????????????. ??????? ??.
417: 130-132.
?????? ?.?., ??????? ?.?., ????????? ?.?., ?????? ?.?. (1996) ???????????? ????????????? ?????????? ??????????????????. ??????? ?? 350: 707-711.
?????? ?.?., ??????????? ?.?., ???????? ?.?. (2006) ????????????????? ? ?????????????? ???????? ??? ????????. ??????????: ???????, 287 ?.
??????? ?.?., ??????? ?.?., ???????? ?.?., ??????? ?.?., ???????? ?.?., ?????? ?.?.,
?????? ?.?., ???????? ?.?., ?????? ?.?. (2003) ????????? ???????? ?????? ???????? ? ????
? ????? ???? (???????). ?????????????. 72(2): 259-267.
Gaevsky N.A., Zotina T.A, Gorbaneva T.B. (2002) Vertical structure and photosynthetic activity
of Lake Shira phytoplankton. Aquatic Ecology. 36: 165?178.
Imam S.H., Gordon S.H., Shogren R.L. (1999) Degradation of starch?poly(?-hydroxybutyrateco-?-hydroxyvalerate) bioplastic in tropical coastal waters. Appl. Environ. Microbiol. 65: 431-437.
Jendrossek D., Handrick R. (2002) Microbial degradation of polyhydroxyalkanoates. Annu. Rev.
Microbiol. 56: 403-432.
# 214 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????, ?.?. ??????????? ?????????? ?????? ?? ?????????????????????? ? ???????????????? ????? ????
Kalacheva G.S., Gubanov V.G., Gribovskaya I.V., Gladchenko I.A., Zinenko G.K., Savitsky S.V.
(2002) ?hemical analysis of Lake Shira water (1997-2000). Aquatic Ecology. 36: 123-141.
Volova T.G., Gladyshev M.I., Trusova M.Y., Zhila N.O. (2007) Degradation of polyhydroxyalkanoates in eutrophic reservoir. Polym. Degrad. Stabil. 92: 580-586.
Wang S., Song C., Mizuno W. (2005) Estimation on biodegradability of poly (3-hydroxybutyrateco-3-hydroxyvalerate) (PHB/V) and numbers of aerobic PHB/V degrading microorganisms in different natural environments. J. Polym. Environ. 13: 39-45.
Zadereev E.S., Tolomeyev A.P. (2007) The vertical distribution of zooplankton in brackish meromictic lake with deep-water chlorophyll maximum. Hydrobiologia. 576: 69-82.
Degradation of Polyhydroxyalkanoate Films
in Brackish Lake Shira
Natalia O. Zhilaa,b, Svetlana V. Prudnikovab,
Egor S. Zadereeva,b and Denis Yu. Rogozina,b
a
Siberian Federal University,
41 Svobodny, Krasnoyarsk, 660041 Russia
b
Institute of Biophysics of Siberian Branch
of Russian Academy of Science,
50/50 Akademgorodok, Krasnoyarsk, 660036 Russia
The degradation of polyhydroxyalkanoate films was firstly studied in brackish Lake Shira at different
depths, 3, 9, 13 and 20 meters, those corresponded to the layers of stratification. It is found that
poly (3-hydroxybutyrate) and copolymer poly (3-hydroxybutyrate-co-3-hydroxyvalerate) with less
crystallinity are destroyed with varying intensity. The maximum mass loss of polymer films and decrease
in polymer molecular weight were registered at 3 m, in oxygenic epilimnion where temperature is the
highest. The phenomenon of polymer films destruction were also found in the chemocline and anoxic
low-temperature monimolimnion, at 20 m depth.
Keywords: Lake Shira, stratification, polyhydroxyalkanoates, degradation, molecular mass.
????? ????????????. ????? ????? ? ???????. ????? ????? ????????. ????????????. ?????????????, ?. 7, ?.: ??????, 145 ?.
??????? ?.?. (1985) ???????????????? ????????? ? ??????? ????????? ? ??????????? ??????? ??????????? ???? ? ????? ? ?? ?????????????? ??????????. ???. ? ?????????.
21,10,85; ? 714-??. ???????????: ??????. ??? ???. ???-?? ? ????????????, 225 ?.
??????? ?.?., ???????? ?.?. (1992) ???????????? ? ????????? ????????? ??????????? ?????? ?: ?.?. ????????? (???.) ????????????? ????????????????? ???????????? ? ??????????
?????? ??????????? ????. ???????????: ??????. ??? ???. ???-?? ? ????????????, ?. 14-52
???????? ?.?. (1970) ?????????? ????? ????. ?.: ?????, 744 ?.
??????????? ?.?., ?????? ?.?. (1981) ???????????? ???????????? ?? ????? ? ?????????
?????????? ??? ?????????????????? ????????????? ?? ???????????? ????????. ???????????? ? ??? ?????????. ?.: ????????, 32 ?.
????????? ?.?. (1982) ???????????? ???????????? ?? ?????????? ??????????? ???????
???????? ??????? ??? ? ??????? ?????? ????????? ?? ???????? ???? ? ???????????? ????????. ?.: ????????, 26 ?.
# 199 #
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?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? ?.?. (1958?) ????????? ???? Mesocyclops leuckarti (Copepoda, Cyclopoida). ??????? ?? ????. 120 (2): 419-421
??????? ?.?. (1958?) ????????? ?????? ?? ???????? ???????? ? ??????????? Acanthocyclops viridis (Copepoda, Cyclopoida). ??????? ?? ????. 119 (3): 613-616
??????? ?.?. (1959) ???????? ????? ???????? Acanthocyclops viridis (Jurine) ? Mesocyclops
leuckarti (Claus) (Copepoda, Cyclopoida): ???????. ???.? ????. ????. ????. ?., 26 ?.
??????? ?.?. (1998) ??????? ???????????? ??????????????. ?.: ???, 320 ?.
????????? ?.?. (2010) ?????????? ??? ????????. ????????? ??????. ?????????? ???????? ??????????? ?????????? (? 100-????? ?.?. ???????????). ?.: ???, ?. 283-304
???????? ?.?., ????? ?.?. (2008) ??????? ????-???????? ?ercopagis pengoi (Ostroumov)
?? ????????? ? ???????????????? ???????????? ??????????? ?????? ??????????? ????. ???????? ?????????? ???????? ?????? ???. 9 (5/1): 244-252
???????? ?.?. (2009) ??????????? ? ????????? ??????? ???? (?? ??????? ????????? ?
??????????? ??????? ??????????? ????). ???????? ?????????? ??? 1: 76-85
???????? ?.?. (2010) ??????????-?????????????? ??????????? ???????????? ??????????? ?????? ??????????? ????. ???????????: ??????. ??? ???. ???-?? ? ????????????, 198 ?.
???????? ????????????????? ??????????: ????????? ?????????? ???????????? (???) ?
?????????????? ?????????? ??????? ??????????? (????) ??????? ??????? ??? ???? ??????
????????, ??????? ????????????????? ???????? (1999). ?.: ?????, 304 ?.
??????? ?.?. (2000) ????????????? ??????????? ???????????? ????? ? ??????? ????
??????-?????? ? ?? ?????????????. ???.: ????????, 223 ?.
????????? ?.?. (2000) ???????? ???????? ? ??????????? Marenzellaria viridis ? ?? ??????? ?? ????????? ?????????? ?????????? ??????????? ??????. ?: ????-???????? ? ??????????? ????? ??????. ???????, ?. 193-202
???????? ?.?., ?????? ?.?., ?????????? ?.?., ??????? ?.?. (1984) ???????????? ???????????? ?? ????? ? ????????? ?????????? ??? ?????????????????? ????????????? ?? ???????????? ????????. ????????? ? ??? ?????????. ?.: ????????, 51 ?.
???????? ?.?., ??????? ?.?., ??????????? ?.?. (1984) ???????????? ???????????? ??
????? ? ????????? ?????????? ??? ?????????????????? ????????????? ?? ??????? ????????.
??????????? ? ??? ?????????. ?.: ????????, 33 ?.
???????? ?.?., ????????? ?.?. (1998) ??????????? ????????? ?????????? ??????????????? ???????. ?: ??????????-????????????? ???????????? ?????????? ? ??????????
???? ? 1996-1997 ?????. ???????????: ??????.??? ???. ???-?? ? ????????????, ?. 96-129
????? ?.?., ??????? ?.?., ????????? ?.?. (2004) ????? ?????????????? ???????????
?????? ?: ?????????????? ??????????????????? ?????? ???????? ??????? ????. ?.: ???????
???, ?. 17-18
????? ?.?. (1971) ?????? ??????. ?: ?????????????????????? ????? ??????????? ??????. ?.: ???????????????, ?. 143-164
???????? ?.?. (1971) ?????????????? ????????? ? ??????? ??????. ???????????????? ?????????????? ?????? ? ??? ????????? ?????. ?: ?????????????????????? ????? ???????????
??????. ?.: ???????????????, ?. 6-9
?????? ?.?. (1972) ????????????? ??????? ????????????. ????: ??????? ?????, 195 ?.
# 200 #
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?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
??????? ?.?. (1979) ? ??????? ???????? ????????????? ? ??? ??????????. ?.: ???, 166 ?.
????????? ?.?. (1982) ???????? ???????? ??????? ???? ????????? ?????? ???????????
????. ?: ??????? ? ??????? ????????? ??? ? ?????????????? ?????????????? ??????. ??????.
???????????: ??? ???. ???-?? ? ????????????, ?. 75-80
????????? ?.?. (1988) ??????? ???? (Anguilla anguilla) ? ?????????? ?????? ???????????
????. ?: ????????????? ? ????????????????? ???????????? ? ????????????? ?????? ? ???????????? ????? ?????? ??????. ???????????: ??????. ??? ???. ???-?? ? ????????????, ?.
97-107
????????? ?.?. (1992?) ??????? ????????? ???????????? ??? ? ?????????? ??????. ?:
????????????? ? ????????????????? ???????????? ? ?????????? ?????? ??????????? ????.
????????? ?.?. (???.). ???????????: ??????. ??? ???. ???-?? ? ????????????, ?. 165-188
????????? ?.?. (1992?) ??????? ?????? ??? ? ?????????? ?????? ??????????? ????.
??????? ??????????. 32 (3): 171-176
????? ?.?., ????? ?.?. (1999) ?????????????? ?????? ??????? ?????????? ?????? ??????????? ???????. ?: ????????????? ??????? ???????????, ?????????. ???????????, ?. 96
????? ?.?., ????????? ?.?., ???????? ?.?. (2004) ???????? ???????????? ?????? ??????
? ??????????????? ??????? ?????? ? ????? ? ????????? ????? ? ??????????? ???????????????? ?????. ?: ?????????? ???? ????: ????????????????? ? ???????????. XXXI ????????????? ??????? ???????????, ?????????. ???????????, ?. 178-180
????? ?.?., ????????? ?.?., ???????? ?.?. (2005) ? ???? ?????????????? ??????? ? ???????? ?????????? ????????? ? ?????????? ??????. ?: ???????? ????? ? ???????. ?VI ????????????? ??????? ????? ?? ??????? ????????, ?????????. ?.2, ?., ?. 237-238,
????? ?.?. (2006) ???????? ???????????? ????????????????? ? ?????????? ?????? ??????????? ????: ???????. ???? ????. ???????-?????????. ????, ???????????, 23 ?.
????????? ?.?. (1994) ?????? ??? ?????????? ??????????? ????????? ???????????????? ?
????????? ???????. ????????? ? ????????? ???????. 3: 33-40
????? ?.?., ????????? ?.?. (2003) ????? ??????????? ???????? ?????? ? ???????? ?
????????????? ?????????? ????????. ?: ?????? ??????? ???????? ??????????????? ????????
(??????????????? ?????????). ?. 2, ???????????: CD-ROM ??????, 12?-1 ? 12?-32
??????? ?.?. (1952) ??????? ? ??????? ??????????????? ??? ??????????? ????. ?.:
?????????????, 268 ?.
??????? ?.?. (1992) ??????????? ????????? ?????? ??? ? ?????????? ??????. ?: ????????????? ????????????????? ???????????? ??????????? ????. ???????????: ??????. ???
???. ???-?? ? ????????????, ?. 151-165,
Elliott J.M., Persson L. (1978) The estimation of daily rates of food consumption for fish. J. Animal. Ecol. 47 (3): 977-991
Hakanson L., Buolion V.V. (2002) The lake food web. Modeling predation and abiotic/biotic interaction. Leiden: Backhuys Publishers, 344 p.
Haney J.F., Hall D.J. (1973) Sugar-coated Daphnia: A preservation technique for Cladocera.
Limnol. Oceanogr. 18 (2): 331-333
Nie H.W. de (1987) Food, feeding periodicity and consumption of the eel Anguilla anguilla
(L.) in the shallow euthrophic Tjeukemeer (the Netherlands). Arch. Hydrobiol. 109 (3): 421-443
# 201 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????????, ?.?. ?????????? ?????? ??????? ? ?????????? ??????????? (????????????????) ??????...
Semionova S.N. (2003) Peculiarities of phytocene state in the Vistula Lagoon of the Baltic Sea in
the current conditions. In: Abstract Publication Baltic Sea Science Congress. Helsinki, 24-28 August,
2003, P. 56
Energy Flows in the Vistula Lagoon
of the Baltic Sea
Elena N. Naumenko,
Mikhail M. Khlopnikov and Liliya V. Rudynskaya
Atlantic Scientific Research Institute of Marine Fisheries
and Oceanography (AtlantNIRO)
5 Dm. Donskoy St., Kaliningrad, 236000 Russia
Energies of the Vistula Lagoon ecosystem is estimated based on observations made by authors in
the northern Vistula Lagoon literature data. The pelagic and bottom communities are represented
by euryhaline species: phytoplankton ? 412, zooplankton ? 74, zoobenthos ? 43, fish ? 54, Mean
biomasses during the vegetation season (April-October) were: 6,2 (8,7 kJ) of phytoplankton, 2,5
(5,2 kJ) of zooplankton, 20,6 (61,8 kJ) g/m2 of benthos. Yield of aboriginal fish amounted to 1,4 tons/km2
(60 kJ/m2).
Annual primary production was 14000, pure -3900; zooplankton and benthos production during
vegetation period amounted to 300 and 1100 kJ/ m2, respectively. Ratios of production of planktonic
crustaceans, benthos and fish catch to primary production were 2,3, 8,5 and 0,06 %, respectively.
Yield of aboriginal fish is formed mainly due to detritus food chain.
Keywords: Vistula Lagoon, production, fish yield, food chain, energy flows.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 203-209
~~~
??? 579.68 (285.23)
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????????? ??????????????? ???????????,
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Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
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???????????. ???????? ????????????? ?????? ?????????? ??????????? ? ??????? 198090-? ??. ????? ??????????? ??????????????? ?????????? ?? 0,9 ?? 6 ??? ??/??, ????????? ??
?????? ??????? ???, ??????????? ??????????? ??????????????? ??????????? ?? ???????.
??????? ???????? ??????????? ?????????????? ???????? ? ????? ?. ???????? (5240 ???/
??) ? ??????? ?????? ?. ???????? (10700 ???/??). ???????????? ???????? ???? ?????????? ?
??????? ?????? ?. ???????? (11333 ???/?). ?????????? ???????????????????? ???????????????
??????????? ?? ??? ??????? ? ????????? 1-9 ???/??. ????????????????????????
?????????????? ?????????? ?????? ? ?????? ?. ????????. ???? ?????? ?. ???????? ?
?????????? ????????? ????-????????? ????????????? ?? ?????????????????? ???????????
????????????? ??????????? ?????? 2.1.5-980-00 ? ????? ???? ???????????? ?????? ???
????????????? ?????.
???????? ?????: ????-???????? ?????????????, ???? ????????, ?????????-??????????????????
?????????? ????.
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*
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(???????????, 1994). ????????? ????????????
? ???????????????? ????????????? ?????????
Corresponding author E-mail address: ingleb83@mail.ru
© Siberian Federal University. All rights reserved
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??????? ????????? ????? ????-????????? ????????????? (??????? ? ??., 1982; ???????,
??????????, 1983, ??????????, 1983?; ???????????, 1985, 1995; Vinogradova,1998).
?? ??????????? ???????????? ???????????? ????????????? ???? ???????? ?
???????????? ??????? ? ???????????????? ????????? ? ??????, ?????????? ? ???????????? ?????? (??????????, 1982, 1983?,
1984).
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??????? ???????? ???????????. ??????????
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???. 1. ?????-????? ?????? ????. ???????: 1 ? 4,5 ?? ???? ?????? ?. ???????? (? ?????? ??????); 2 ? 4,5
?? ???? ?????? ?. ???????? (? ??????? ??????); 3 ? ???? ????????, 1,5 ?? ?? ???????; 4 ? ?????? ?????? ?.
????????; 5 ? 6,5 ?? ?? ??????? ?????? ?. ????????; 6 ? 0,5 ?? ???? ?? ?????? ?? ????? ?????? ?. ????????;
7 ? ????? ??????; 8 ? ?????? ?. ???????? ? ?????? (????????); 9 ? ????? ?????? ?. ????????
# 204 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
?????????-?????????????????? ???????????? ???????? ??????????? ?????????
???????????: ????? ??????????? ??????????????? (???), ?????????? ???????????
(??), ???????????????????? (???), ??????????????????? (???) ???????????????,
?????????? ???????? ?????? ???????? ??????? (????).
??? ?????????? ?? ?????????? ???????? «Millipore» ? ????????? ??? 0,22 ???.
??????? ?????????? ??????????? ? ???????????? ??? ??? ??????????? ? ??????????? ? 1350 ??? (?????????, ????????, 1974).
?????????? ??????????? ???????????????
?????????? ?? ????? ???:10 (????????? ????? ???????????? ??? ??????????? 18-20 є? 10
???). ?????????-????????????? ????????? ???
?????? ?. ???????? ????????? ?? ??????????
????. ?????????? ????????????????? ?????? ????? ??????????? ???????? (??? ???)
?????????? ??????? ?????????? ????????
(???? 24849-81). ????? ???? ???????????
????? ??????? ? 8 «????????» («?????????»,
??????), ??????? ??????????? ??? 37 єC ??
??????-?????????? ????? ???? 24 ?. ??? ??????? ?? ???????? ???????, ??????????? ???
????, ?? ?????????? ??????? ??????? ????
???????? ?????, ?????????? ?? ?????, ????????????????. ??? ??????? ? ?????? ?????????????????, ????????, ?? ??????????
???? ??????? ????????? ?????????? ????. ????????????????????? ??????? ????????????
? ???????????? ?? ?????????????? ? ?????
??????????? ????????? ??????? ????-????
????????????? ??????? ??????? ???? ?? ?????????? ????? ? ????????. ?????? ???????????? ??? 37 є? 24 ?. ??? ???????????
??????? ? ???? ????????? ??????? ?????????????. ????????? ???????? ? ???? ????-???????
(?-?): ??? ??? ? 1 ? ????????????? ????.
??????????
????????????????????
??????????????? ???????????? ?? ???????-
?????? ????? ??????????. ?????????? ??????????????????? ???????? ?????????? ??
????? ????????-???????? (?????????, ????????, 1974).
?????????? ? ??????????
?????????? ???????????? ????????,
??? ????? ??????????? ??????????????? ?
?. ???????? ? ?????????? ????????? ????????????? ????????????? ?????????? ??
0,9 ?? 6 ??? ??/?? ? ????????????? ????,
????????? ?? ?????? 1980-90-? ??. (0,8-4;
1-10,4 ??? ??/?? ??????????????) (??????????,
1984; ???????????, 1994). ? ????? ?. ????????,
? ??????? ?. ???????? ? ?????? ? ?????? ???????????? ??? ???? ?? ?????? 2 ??? ??/??.
????????? ???????? ???? ??????????? ?
?????? ?????? ?. ???????? ? ?? ??????? 13 ? ?
6,5 ?? ?? ??????? ?????? ?. ???????? (???. 2).
?????????? ??? ? ??????? ???????????
??????? ?????????? ? ? 1980-90 ??. ? ???? ??????? ? ??????? ????????????? ????????????
??????? (???????????, 1994).
?????????? ?? ? ????? ?????? ?. ???????? ? ??????????? ? ?????? ?????????
????-????????? ????????????? ??????????
?? 1230 ?? 10745 ?? ??????????? ? ?? 815 ??
4500 ???/?? ? ????????? ????????? (???. 3).
???????? ?????????? ?????????? ???????????? ??????? ???????????????? ? ?????
?. ???????? ? ? ?? ?????? ??????????? ? ????????????? ??????????. ?????? ? ???? ????????? ??????, ?????????? ? ?????? ??????
? ? ????????? ????-????????? ?????????????, ??????????? ? ?????? ????????, ?????????? ?? ???? ????, ??? ? ?????????????.
????????, ? ????-???????? ?????????????
?????????? ???????????? ???????????? ???????, ??????????? ?? ?. ????????, ???????
??????????? ?????????? ?? ? ?????????
??????????. ???????????? ???????? ??????????? ??????????????? ????????????? ?
# 205 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
7000
???, ???. ??./??
6000
5000
4000
3000
2000
1000
0
1
2
3
4
5
????????????? ????????
6
7
8
9
????????? ????????
???. 2. ????? ??????????? ??????????????? ? ?????? ?. ???????? ? ????-???????? ?????????????
? ???? 2005 ?.: 1-9 ? ??????? ?????? ????
12000
???, ???/??
10000
8000
6000
4000
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????????????? ????????
6
7
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9
????????? ????????
???. 3. ?????????? ??????????? ??????????????? ? ?????? ?. ???????? ? ????-???????? ?????????????
? ???? 2005 ?.: 1-9 ? ??????? ?????? ????
??????? ?????? ?. ???????? (??. 4). ???? ?????
???????????????? ????????????? ?????????? ?????????? ???????? ??????? (?? 2600
???/?) ? ????????????? ?? ??????? (???????????, 1984).
???????? ?????????? ??????????? ?
????????????? ?????? ???????? ?. ????????,
???? ???????????? ????? ??????? ??? ????????? ??? ? ????????????-??????? ??? ?.
???????. ??????? ???????? ?????????? ??,
?????????? ? ????? ?. ???????? (5240 ???/??)
? ??????? ?????? ?. ???????? (10700 ???/??),
??????????????? ? ????? ??????????? ????????? ?????????? ???????????? ??????? ?
?????? ???????. ??? ???????? ?? ??????? ?????? ?. ???????? ? ?????? ?????? ?????????????? ???????? ??????????: ?? ???? ????????
?????????? ?????????? ????????? ?? ???????
(1300-1600 ???/??). ???????? ??????? ????????????? ?? ?? ?????? ??????? ???????-
# 206 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
12000
?-?, ??? ???/?
10000
8000
6000
4000
2000
0
1
2
3
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????????????? ????????
6
7
8
9
????????? ????????
???. 4. ????-?????? ? ????? ?????? ?. ???????? ? ????-???????? ????????????? ? ???? 2005 ?.: 1-9 ?
??????? ?????? ????
???? ? ? 1980-? ????? (??????????, 1984). ?????? ?? ????????? ? 1980-?? ??. ???????????
?? ? ?????? ??????????? ?? ???????.
?????????? ???? ?????????? ?? 166 ??
11333 ???/? ? ????????????? ? ?? 0 ?? 5666
???/? ? ????????? ?????????? (???. 4). ???????????? ???????? ???? ?????????? ?
??????? ?????? ?. ???????? (11333 ???/?).
??? ???????? ?? ??????? ?????? ??????????
???? ??????????? (? 6,5 ?? ?? ??????? ?????? ?. ???????? ? ? ?????? ?????? ???? ?????????????), ? ? ????? ????????????? ?????
?????????????. ??????? ????????, ??? ? ????? ?????? ?. ???????? ?????????? ???? ?
????????????? ?????????? ???? ????, ??? ?
?????????, ? ? ????????????? ???????????
??????????????? ???????: ?????????? ????
????????????? ? ????????? ??????????. ??????????, ? ????????????? ???? ????????
????????????? ?????? ????, ??????? ???????? ? ?????????? ?????????? ???? ? ????????? ??????????. ??? ?????????? ????, ??
???? ?? ?????????? ?????? ??????????? ??????????? ????????, ??????? ??????????????? ? ???????????? ?????????? ????????????
???????.
?????????? ??? ? ????? ?. ????????, ??
?????? ? ????? ????????????? ????????? ??
1 ?? 9 ???/??. ? 1980-? ??. ?? ?????? ???????
?????????? ??? ?????????? 8-20 ???. ???/
?? (??????????? ? ??., 2004). ?????????? ?????????? ??????????????? ???? ??????????????? ?????? ??????? ? ?????????? ????????? ?????????? ?????????, ??????? ?????
???? ??????? ? ?????????? ???????? ???????? ????????? ??????? ?????????, ??????? ?
???????? (???????????, 2010).
? ????????????? ???? ?????????? ????????????????????????
??????????????
(???) ?????????? ? ???? ?????? (???????? ?
????? ?????? ?. ????????). ?? ??????, ?????????? ? 1980-? ??., ?????? ?? ???? ???????? ???? ????????????? ??????? ???????????
??????????????? ???? ??????, ??? ??? ? ????
??????? ??????????? ???????????? ???????
?????????? ? ???????????????? ??????????
????????????? ? ????????? ?????????? ?????????? 26-59 % (???????????, 1994).
?????????-?????????????????? ?????????? ??? ?. ???????? ? ?????????? ????????? ????-????????? ????????????? ??????????????? ? ?????????? ?????????????
# 207 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
???????? ?? ?????????? ? ?? ???????????
????????? ????????????. ???? ?????? ?.
????????, ?????? ??????, ?????????? ????????? ????-????????? ????????????? ??
?????????????????? ??????????? ????????????? ??????????? ?????? 2.1.5-980-00 ?
????? ???? ???????????? ?????? ??? ????????????? ?????.
?????? ??????????
??????? ?.?., ?????????? ?.?. (1983) ??????? ??????? ??? ????????? ????????????????? ????????? ?? ?????? ?????????????? ????-????????? ?????????????. ?: ?????? ??????? ?? ??????????? ????????????? ????????? ??????????? ??????????-???????? ??????????????. ?., ?. 105-112.
??????????? ?.?. (1985) ???????? ?????????? ????????????????? ????-????????? ????????????? ? 80-? ???? ?: ?????????? ???????? ? ??????? ? ????????. ???. 3. ???????, ?. 11-12.
??????????? ?.?. (1994) ?????????????????? ????? ????-????????? ????????????? ?
???????? ??? ????????????: ???????. ???. ? ????. ????. ????. ???????, 25 ?.
??????????? ?.?. (1995) ?????????? ?????????? ????-????????? ????????????? ??
?????????????????? ???????????. ???????? ????????. ???????? ????????: ?????????
V ????????. ????.: ?????? ?????? ?????????? ?.?. ??????. ???????????: ?? ?????, ?. 2,
?. 248-250.
??????????? ?.?., ?????? ?.?, ??????? ?.?., ?????????? ?.?., ????????? ?.?. (2004) ?????????????????? ???????? XX ????. ?. 1. ????? ???????? ???????-??????-?????????? ??????????. ???????: ???-?? ??-?? ????????? ?? ???, 93 ?.
???? 24849-81 ???? ????????. ??????? ?????? ?????????-??????????????????? ???????.
??????????? ?.?. (1984) ? ???????????? ???????????? ??????????????? ? ??????????
????????????? ???????? ???? ?????? ? ????????? ????????????. ?: ???????????? ?????????
???????? ????? ?????? ? ????????? ????????????. ???????, ?. 63-69.
?????????? ?.?. (1982) ????????????? ??????????? ?/??2 ???????????????? ?. ????????? ? ????-????????? ?????????????. ?: ???????? ???????? ???????????: ??? ????. ? ????.
????. ????. ???????, ?. 2, ?. 52-58.
?????????? ?.?. (1983?) ???????????? ???????? ???????????? ??????? ???. ?: ???????? ?????? ???????: ???. ????.-?????. ?????. ?????????, ?. 13-14.
?????????? ?.?. (1983?) ???????? ??????? ??????? ??? ????????? ??? ?? ????????????
????????? ?????? ????-????????? ?????????????. ?: ?????? ??????? ???????? ??????? ?
??????. ????????????, ???????????? ????????????? ? ??????: ???. ????. ????.-?????. ?????.
???????, ?. 143-145.
?????????? ?.?. (1984) ????????? ??? ????-????????? ????????????? ?? ?????????????????? ???????????: ???????. ???. ? ????. ????. ????. ????, 24 ?.
??????????? ?.?. (2010) ?????????????? ????????????? ??????? ??????? ???
??????????-???????? ??????????? (?? ??????? ????????? ???): ???????. ???. ? ????. ????.
????, ???????, 20 ?.
????????? ?.?., ???????? ?.?. (1974) ???????? ??????????????? ??????? ????????. ???????????? ???????????. ?.: ?????, 194 ?.
# 208 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ???????, ?.?. ????????? ?????????????? ??? ????-????????? ????????????? ? ?????? ???? ?????????
??????? ?.?., ????????? ?.?., ?????? ?.?., ?????? (???????????) ?.?. (1982) ???????????? ?????????? ????-????????? ????????????? ?: ???????? ???????? ???????????. ???????, ???. 1, ?. 34-35.
?????? 2.1.5-980-00 2.1.5. ????????????? ?????????? ????, ?????????? ?????? ??????
????????.
Vinogradova T.P. (1998) Long-term investigation of bacterioplankton of the Ust-Ilimsk reservoir.
In: BICER, BDP and DIWPA Joint International Symposium on Lake Baikal, Yokohava, ?. 111.
Characteristic of Water Ust-ilim Reservoir
and Bay of the River Vikhoreva
by Microbiological Indicators
Inessa V. Glebovaa,
Vyacheslav V. Maximov and Elena V. Schetininab
a
Institute of Geochemistry of A.P.Vinogradov SB RAS,
1A Favorsky st., Irkutsk, Russia 664033
b
Institute of Biology of Irkutsk State University,
3 Lenin st., Irkutsk, 664003 Russia
b
Ust-Ilim reservoir and the bay of the Vikhoreva River on microbiological parameters were investigated.
A comparative analysis of the results obtained with these of 1980-1990 years was carrying out. The
total number of microorganisms ranged from 0.9 to 6 million cells/ml, remaining at the level of previous
years. The number of saprophytic bacteria decreased in the order. High values of saprophytic bacteria
identified in the waters of the Vikhoreva River (5240 CFU/ml) and backwater bay Vikhoreva (10700
CFU/ml). The maximum values of coliforms were found in backwater bay r. Vikhoreva (11333 CFU/l).
Number of cellulolytic microorganisms has decreased by three orders of magnitude and amounted
to 1-9 CFU/ml. Sulfate-reducing bacteria were found only in the bay of r. Vikhoreva. Water of bay of
r. Vikhoreva and adjacent waters of the Ust-Ilim reservoir on the microbiological data comply with
SanPiN 2.1.5-980-00 and can be used only for recreational purposes.
Keywords: Ust-Ilim reservoir, river Vikhoreva, sanitary and microbiological parameters of water.
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
Journal of Siberian Federal University. Biology 2 (2012 5) 210-215
~~~
??? 574.4
??????? ?????????
?????????? ?????? ?? ??????????????????????
? ???????????????? ????? ????
?.?. ?????,?*, ?.?. ???????????,
?.?. ?????????,?, ?.?. ????????,?
?
???????? ????????? ?????????? ?????????
?????????? ???????? ????,
?????? 660036, ??????????, ????????????? 50, ???. 50
?
????????? ??????????? ???????????,
?????? 660041, ??????????, ??. ????????? 79 1
Received 7.06.2012, received in revised form 14.06.2012, accepted 21.06.2012
??????? ??????????? ?????????? ?????? ?? ????(3-????????????????) ? ????(3????????????????-??-3-????????????????) ? ???????????????? ????? ???? ?? ?????????
????????: 3, 9, 13 ? 20 ?, ??????????????? ?????????? ????????????? ?????. ????????, ???
???????? ??????? ?????????? ?????? (???????? ????? ?????? ? ???????????? ????? ????????)
?????????? ? ??????????? ?? ??????? 3 ?, ? ???????? ???????????? ? ?????????? ?????????
?????. ?????????? ?????? ????????????? ????? ? ????????? ?? ??????? 13 ? ? ?? ??????? 20 ? ?
??????????????, ??? ???????? ?????????? ?????????? ?????????, ??????????? ????????????
? ?????? ???????????.
???????? ?????: ????? ????, ?????????????????????, ??????????, ???????????? ?????.
????????
????? ??????? ??????????????? ? ????????? ????? ??????????? ??????????
?????????? ???????????? ??????? ????????, ????????????? ????????????????, ???
?????????? ????????????????????? (???)
(?????? ? ??., 2006; Jendrossek, Handrick,
2002, Wang et al., 2005). ???? ?? ???????????
?? ?????????? ? ?????? ????????????? ?????????, ????????????? ??????? ??????? ?
*
1
?????????? ?????? ????????? ???? ????????????? ????????? ? ????????? ???????? ??????, ???????? ? ?????????? ???????.
? ?????? ?????????? ?????????? ??? ???
??????? ???????????? ??????????? ???????????? ??????????????? ?? ?????????? ?
????????? ????????. ????? ?????? ???? ???????????????. ???????????? ??????????
? ?????????? ??? ? ??????? ????, ? ?????
??????? ? ???????????????? ??????. ??-
Corresponding author E-mail address: nzhila@mail.ru
© Siberian Federal University. All rights reserved
# 210 #
Copyright ??? «??? «??????» & ??? «A???????? K????-C?????»
?.?. ????, ?.?. ??????????? ?????????? ?????? ?? ?????????????????????? ? ???????????????? ????? ????
?????? ??????, ??? ?????????? ??????????
3-????????????????-??-3-????????????????
? ??????? ???? ?????????? ????? ????????
(Imam et al., 1999). ?????? ???????? ?????????? ??? ? ??????? ???? ??????? ? ????????
?????? (Wang et al., 2005). ??? ?????????????
?????? ?? ????????? ?????????????? ???????? ????????? ?????????? ??? ? ?????????
????? ? ?????? ?? ??????????. ????????? ?
??????? ???? ???????????? ?????????????
??? ? ????????? ??????? ???????? ???????,
??? ????????????? ??????? ?? ????????? ?????? ?????????? ? ???????? ???????, ? ????? ?????????? ????????? ??? (Volova et al.,
2007). ???????????? ?? ?????????? ??? ???
???? ?????????? ? ????? ???? ?? ?
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