ISSN 00978078, Water Resources, 2010, Vol. 37, No. 5, pp. 684–698. © Pleiades Publishing, Ltd., 2010.

Original Russian Text © V.I. Lazareva, 2010, published in Vodnye Resursy, 2010, Vol. 37, No. 5, pp. 590–604.

WATER QUALITY AND PROTECTION:

ENVIRONMENTAL ASPECTS

LongTerm Zooplankton Dynamics in the Rybinsk Reservoir

(1956–2005)
V. I. Lazareva
Institute of Inland Water Biology, Borok, Nekouzskii district, Yaroslavl province, 152742 Russia
Received January 19, 2009

Abstract—Materials of halfcentury observations of zooplankton in the Rybinsk Reservoir were used to

establish reversible variations in its structure, abundance, and biomass and irreversible variations in the char
acteristics (trends). Variations with periods of ~10 and 20 years have the largest effect on the community
dynamics. Regression models are presented describing the dependence of zooplankton biomass averaged over
May–October on its values in summer months and on the concentrations of chlorophyll a and phytoplankton
and hydrothermal characteristics of the water body. Changes in zooplankton community under the effect of
eutrophication and deeutrophication of reservoir ecosystem are discussed.

Keywords: reservoir, zooplankton, composition, structure, abundance, biomass, regularities in dynamics.

DOI: 10.1134/S0097807810050088

INTRODUCTION been analyzed since 1997. The stations and zooplank

The isolation of anthropogenic component in the
dynamics of aquatic communities is impossible unless
one knows the range of their natural variations, caused
primarily by climate changes in the region where the
reservoir basin is located. The variability of zooplank
ton abundance, biomass, and other parameters is
characterized by the amplitude and space and time
scales [13, 20, 40].
Regular (“standard”) observations of zooplankton
in the central Rybinsk Reservoir were carried out in
1956–1995; analysis of different aspects of plankton
dynamics for different time intervals in this period is
given in [13, 20, 34, 36]. In the recent decade, hydro
biological studies were carried out irregularly (mostly
in the summer). Nevertheless, the accumulated mate
rial considerably extends the knowledge about plank
ton dynamics and enables the discussion of the indica
tor significance of the community with the aim to
assess the trophic status of reservoir ecosystem. The
objective of the study was to analyze the longperiod
regularities in variations of zooplankton structure and
abundance. The major attention was given to present
day community dynamics (1990–2005).
MATERIALS AND METHODS
The study was based on archive materials of 1956–
1995, collected by Laboratory of aquatic invertebrate
ecology, Institute of Inland Water Biology, Russian
Academy of Sciences; samples were taken every two
weeks in May–October at six “standard” stations in
the pelagic zone of the Rybinsk Reservoir. Samples
collected by the author in the same reservoir area have
ton sampling procedure are described in [20], and the
reservoir was characterized in [36, 41]. The main
zooplankton catch gear until 1962 was Bogorov’s large
plankton scooper 50 l in volume, D’yachenko–
Kozhevnikov’s planktonbathometer 10 l in volume in
1962–1997, and mostly small Juday net (a model with
an entry ring diameter of 12 cm, textile cone 12 cm in
height, and a cone made from sieve no. 70 with mesh
diagonal of 12 µm and height of 45 cm).
All samples were fixed by 4% formalin. The office
treatment was carried out under MBS microscope in
Bogorov chamber. The species that are small in num
ber and >0.4 mm in body length were counted in a
third part, half, or the whole sample. The dominating
species were identified by the relative abundance sep
arately in the taxonomic groups of crustaceans and
rotifers. The lower boundary of the domination was
taken to be 5% of the total abundance [15, 20]. Cope
pod abundance was calculated taking into account
copepodites and naupliuses. The distribution of the
abundance of juvenile specimens among cyclop spe
cies was estimated by the ratio of males, females, and
copepodites IV–V (the latter were determined up to
the species); the same distribution for Diaptomidae
was estimated by male ratio; females of Eudiaptomus
gracilis Sars and E. graciloides (Lill.) in the Rybinsk
Reservoir demonstrates mixed features of both spe
cies. The biomass of animals was calculated by formu
las relating body mass and length [4].
When necessary, the data series were smoothed by
averaging over three points, which filters out random
variations. The interrelationship between zooplankton
characteristics and their relationship with abiotic

684
 LONGTERM ZOOPLANKTON DYNAMICS IN THE RYBINSK RESERVOIR 685

Table 1. Dynamics of relative abundance of dominant zooplankton species in the central (“standard” stations) of the Ry
binsk Reservoirin 1961–2005 (Stot is the total number of species in the faunistic list: initial [36], final [16]; here and in Tables 4, 5,
7, and 8, dash means no data available)
Relative abundance, %*
Trend (+/–),
Species dry years wet years dry years
%
1961–1965 1971–1975 1981–1985 1991–1995 2003–2005
Mesocyclops leuckarti 57 ± 6 33 ± 4 51 ± 2 35 ± 2 46 ± 5 No
Thermocyclops oithonoides 7±2 16 ± 4 6±2 17 ± 2 8±1 No
Keratella cochlearis 9±3 <5 5±2 <5 9±2 No
Synchaeta pectinata 16 ± 2 <5 10 ± 2 11 ± 3 15 ± 6 No
Keratella quadrata 9±2 14 ± 2 15 ± 2 14 ± 2 8±4 No
Kellicottia longispina 11 ± 1 <5 <5 5±1 9±2 No
Bosmina longispina 11 ± 2 <5 17 ± 1 17 ± 2 5±3 No
Eudiaptomus sp. <5 7±2 5±0 5±1 9±2 +30
Daphnia galeata <5 <5 <5 5±1 12 ± 2 +200
Polyarthra major <1 <1 <5 8±2 20 ± 6 +2000
Polyarthra vulgaris 12 ± 3 <5 7±2 <5 <5 –200
Conochilus sp. 37 ± 6 53 ± 7 34 ± 5 37 ± 8 22 ± 5 –100
Chydorus sphaericus 7±2 11 ± 4 6±0 6±2 <5 –75
Stot 164 – – – 364 +120
* In 2003–2005, mean values were calculated for June–August; in other years, for May–October.

characteristics were revealed by correlation and step
bystep regression analysis. The contribution of inde
pendent variables to variations of the dependent vari
able was evaluated by determination coefficient R2;
and the strength of factor effect, by Fisher test F.
RESULTS AND DISCUSSION
Species Composition
In different decades, the complex of dominants in
the central part of the reservoir, identified by the rela
tive abundance of species averaged over May–Octo
ber, included 5–7 crustaceans and 6–8 rotifers [20].
Permanently abundant among them were cyclops
Mesocyclops leuckarti Claus, Thermocyclops
oithonoides Sars (>50% of crustacean amount) and
colonial rotifers Conochilus hippocrepis Schrank,
C. unicornis Rouss. (>30% of rotifer abundance). In
the beginning of the current century, the composition
of zooplankton dominants remained similar to that in
the 1980s–1990s (Table 1). For some species, varia
tions in the relative abundance with a period of
~20 years (e.g., Thermocyclops oithonoides) and
~40 years (e.g., Keratella quadrata O.F. Muller,
Bosmina longispina Leydig) were revealed. Directed
changes in the abundance were recorded in some oth
ers. The share of Diaptomidae, daphnia (Daphnia
galeata Sars) and rotifers Polyarthra major Bruck in
the community increased. Both absolute and relative
abundance of rotifers (primarily, species Polyarthra
vulgaris Carlin, Conochilus hippocrepis, and cla
doceras Chydorus sphaericus O.F. Muller) decreased.
The number of species in the overall list of zooplank
ton increased more than twice with deeper studies of
the fauna.
The maximal abundance and the occurrence of
highconcentration spots of relatively large
(0.6⎯2.2 mm) crustaceans Bosmina longispina,
B. coregoni Baird, Limnosida frontosa Sars, Hetero
cope appendiculata (Table 2) decreased in the early
XXI century as compared with the 1980s. However, the
number of other large (1⎯2 mm) forms (Daphnia gale
ata, Bythotrephes longimanus, Cyclops vicinus Ulja
nin, Eudiaptomus gracilis and E. graciloides)
increased. The linear size and mass of Daphnia galeata
is greater than Bosmina longispina, while that of Eud
iaptomus and Cyclops is greater than Mesocyclops and
Thermocyclops. The abundance of Leptodora kindtii,
the largest (the body length of up to 9 mm) crustacean
species being stably high can be interpreted as a steady
increase in the size of food plankton in the present
time, which creates a favorable base for the develop
ment of planktoneating fish in the Rybinsk Reservoir.
In the early XXI century, in addition to changes in
the occurrence and abundance of the species that are
common for the reservoir, new species were recorded
and the occurrence and abundance of previously rare
forms was found to increase [15, 17]. In particular, cla
docera Diaphanosoma orghidani Negrea and rotifer
Asplanchna henrietta Langhaus were rapidly spread

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686 LAZAREVA

Table 2. Variations in the abundance N of some species of summer (June–August) zooplankton in the Rybinsk Reservoir at the turn
of the century (Nh is high abundance; Nmax is maximal abundance, thous. specimen/m3; Nh, % is the occurrence of spots with high
abundance of the species calculated as the ratio of the number of samples with Nh to the total number of samples)
N
1987–1988 2003–2004
Zooplankton species Nh,
thous. spec./m3 Nmax, Nh, Nmax,
N ,%
thous. spec./m3 % thous. spec./m3 h
Mesocyclops leuckarti + Thermocyclops oithonoides (cop. + ad.) >30 110 19 85 13
Bosmina longispina >10 62 23 35 9
Daphnia galeata >5 21 17 23 41
Eudiaptomus gracilis + E. graciloides (cop. + ad.) >5 20 19 15 28
Cyclops vicinus (cop. + ad.) >0.5 2 16 5 31
Leptodora kindtii >0.5 1.5 16 2.5 13
Limnosida frontosa >0.5 5 35 2 9
Heterocope appendiculata (cop. + ad.) >0.1 2 33 0.5 13
Bythotrephes longimanus >0.05 0.7 27 0.7 38

ing over the Rybinsk Reservoir water area in 2001–2005.
In 2005⎯2007, Diaphanosoma orghidani, which was
first detected in 2003, was rapidly disseminating over
the entire water area, where its largest abundance
(1120 specimen/m3) was recorded in the Sheksna
Pool. This species was also recorded in the Ivankovo,
Uglich, Gorki, and Sheksna reservoirs. Since 2005,
the abundance of D. Orghidani in the Sheksna Reser
voir is maximal (up to 16000 specimen/m3) among the
water bodies of the Upper Volga basin and this reser
voir serves as a source of propagation of the species
southwards into the Rybinsk Reservoir [17].
Rapid dissemination of rotifers Asplanchna henri
etta, which was first recorded there as early as 20 years
ago in 1985, was found to take place in the Rybinsk
Reservoir in 2003–2005 [15]. Before 2001, this species
was mostly met in the littoral areas and shallow bays
[14]. By 2005, A. henrietta inhabited the entire water
body; high abundance (4.5–12 thousand speci
men/m3 or 14–17% of rotifer abundance) was
recorded in June–July in the Volzhskii pool and in the
northern Main pool [15, 17]. A. Henrietta was found
to disseminate not only into new biotopes of Volga res
ervoirs, but also beyond their limits into Lake Nero
and small water bodies of the basins of the Rybinsk and
Gorki reservoirs [15, 17, 21].
The invader species commonly displace allied taxa
from the dominant zooplankton complex. Thus, roti
fer Asplanchna herricki Guerne in the Rybinsk Reser
voir in the 1970s was almost completely replaced by
finer A. priodonta [14], in the 1990s, Polyarthra vul
garis was replaced by P. major [15, 20]. Overall, the
occurrence and/or abundance of >10 zooplankton
species that have been rare before increased since the
1990s [15]. A typical example of such species is
Bosmina crassicornis P.E. Müller. This species has
been recorded in the reservoir since the mid1960s
[36], its abundance started increasing since 1997 [15].
By 2004, this species occurred all over the reservoir
water area (70–100% of samples); its abundance in the
Sheksna pool reached 3–13.5 thous. specimen/m3
(7–8% of crustacean abundance). High abundance
(3–15.2 thousand specimen/m3) of B. crassicornis was
recorded in June–September 2007 was recorded in all
three pools of the reservoir; it accounted for 12⎯29%
of the amount of crustaceans and was among zoop
lankton dominants.
In 1997⎯2004, the dominant complex of zoop
lankton included 14 species, which previously were
rare or lowabundance, and 5 species, which were
dominating in the 1980s, became lowabundance [15].
This suggests the large mobility of the dominant com
plex structure, its core remaining stable. Every
10⎯20 years, partial or complete replacement of one of
major dominants or subdominants is recorded in the
reservoir: Bosmina longispina B. coregoni,
Conochilus hippocrepis C. unicornis. Changes of
this type commonly follow changes in the phases of
reservoir hydrological cycle with some time lag [20].
Abundance and Biomass
The total abundance Ntot and biomass Btot are the
main characteristics of zooplankton development.
The biomasses of the components of its species are
used to calculate the production of community and
assess the food base of planktoneating fish. The
development of zooplankton in the Rybinsk Reservoir
in 1956–1995 was described by the values of Ntot and
Btot averaged over the vegetation season (May⎯Octo
ber), as it is commonly accepted in hydrobiology.
Starting from 1996, there were no reliable estimates of
the seasonaveraged abundance of zooplankton;
therefore, the community dynamics was analyzed by

WATER RESOURCES Vol. 37 No. 5 2010

 LONGTERM ZOOPLANKTON DYNAMICS IN THE RYBINSK RESERVOIR
Table 3. Amundance, thous. spec./m3, and biomass, g/m3, of zooplankton in the Main Pool of the Rybinsk Reser
voir(“standard” stations) in 1981–2005 (here and in Table 7, Ncl is the abundance of cladocera; Ncop, copepods; Nrot, roti
fer; Ncr, crustaceans; Ntot, total zooplankton; Btot is total zooplankton biomass)
Period, years Sample number Ncl Ncop
June
1981–1990 98 31 ± 2 52 ± 4
1991–2000 62 23 ± 2 36 ± 3
2001–2004 18 11 ± 3 26 ± 8
July–August
1981–1990 104 14 ± 1 68 ± 3
1991–2000 87 13 ± 1 44 ± 4
2001–2004 15 9±2 33 ± 6
2005 5 22 ± 11 93 ± 16
data on months in which the majority of expedition
routes were carried out. Identified were maximal in
the season or “peak” values, which in the Rybinsk
Reservoir are commonly recorded in June [34], and
latesummer values (July–August). To compare
them with the mean over vegetation period, retro
spective analysis of the available data series was car
ried out to 1980.
Before 1995, the seasonaveraged Ntot varied from
year to year within 40–120 thousand specimen/m3;
the peak values reached maximum (>300 thousand
specimen/m3) in 1981–1989 [20]. The latesummer
values of Ntot were comparable with the mean values
(Fig. 1). Variations in Ntot depended on variations in
rotifer abundance (R2 = 91%, F = 1188.2), their peri
odicity was found to be near 10 years [20]. In later
years, tenyear rhythm persisted in Ntot dynamics. A
minimum in Ntot, comparable with that in the mid
1970s was recorded at the turn of the century (Fig. 1a).
A decrease in rotifer abundanceNtot was recorded in
zooplankton in the 1990s. During vegetation
period,Ntot >100 thousand specimen/m3 peaks were
recorded in 3% of the total number of samples;
whereas in the 1980s, they were recorded in 12% of
samples [20]. The seasonaveragedNtot varied within
50⎯130 in 1960⎯1989 and within 33–35 thousand
specimen/m3 in 1992⎯1995. Crustaceans were the
dominating zooplankton group in terms of abundance
at all observation stations; their predominance over
rotifers was especially significant (Ncrust/Nrot was
1.5⎯2.5) in the Volga pool of the reservoir. From the
spring to the autumn in 1996⎯2003, Nrot in the open
part of the water area of the Main and Volga pools was
<100 thousand specimen/m3. In the first half of sum
mer, the amount of rotifers was comparable with that
of crustaceans, while the latter dominated in the sec
ond half of summer (Table 3).
In the new century (2001–2005), zooplankton Ntot
in June at “standard” stations in the Main pool of the
reservoir decreased about two times relative to the
1990s and more than three times as compared with the
WATER RESOURCES Vol. 37 No. 5 2010
687
Nrot Ntot Btot Ncr/Nrot
175 ± 22 258 ± 24 3.2 ± 0.2 0.5
126 ± 17 185 ± 19 1.9 ± 0.2 0.5
37 ± 17 73 ± 20 0.9 ± 0.2 1.0
54 ± 7 136 ± 8 1.6 ± 0.1 1.5
37 ± 7 94 ± 8 1.1 ± 0.1 1.5
22 ± 6 64 ± 10 0.9 ± 0.1 1.9
48 ± 15 164 ± 32 2.8 ± 1.0 2.4
1980s (Table 3). The most significant decrease was
recorded in the abundance of rotifers and cladocers. In
June 2004, Ntot was <100 thousand specimen/m3 in
half of the water area, and in individual stations
reached 700 thousand specimen/m3 [15]. A specific
feature of the presentday plankton in the reservoir in
early summer was a relatively small abundance of
bosmins and the relatively high abundance of daphnia
and leptodoras.
In August 2001–2003, Ntot of zooplankton at the
same stations was 30% less than that in the 1990s and
almost two times less than in the 1980s (Table 3). As
was the case with June, the most significant drop was
recorded in the amount of rotifers and cladocers, and
copepods in the Volzhskii pool. However, in July 2005,
zooplankton concentration was extremely high in the
center of the reservoir. Since the 1980s, a specific fea
ture of zooplankton in the reservoir in the late summer
was the abundance of copepods (Ncop ~50% Ntot) with
the predominance of Mesocyclops leuckarti, Eudiap
tomus gracilis. In the XXI century, their role in the
community persisted, though cladoceras (mostly
Daphnia galeata) were abundant in some years.
Until 1980, zooplankton Btot in the Rybinsk Reser
voir averaged over May–October was <1 g/m3; in the
1980s, Btot was rapidly growing (Fig. 1b); it was greater
than 3 g/m3 in individual stations. The increase in bio
mass was mostly due to the larger abundance of large
size crustaceans Bosmina longispina and Daphnia
galeata [20, 34]. A decrease in Btot was recorded since
1991, and its values did not exceed 1.7 g/m3 in
1994⎯1995. A peak in zooplankton biomass in the
1980s coincided with the highwater phase of the
hydrological cycle of the Rybinsk Reservoir
(1977⎯1995), in which the volume of water inflow was
largest throughout the observation period [22, 23].
Slow oscillation of two orders were typical of zoop
lankton Btot in the reservoir, their periods lying bear 10
and 20 years [20].
Zooplankton Btot continued decreasing in the fol
lowing years, and in 2001–2004, the values of peak in
688 LAZAREVA

(а)
300
1
2
3
Abundance, thous. spec./m3
200

100

1956 1964 1972 1980 1988 1996 2004

1960 1968 1976 1984 1992 2000 Years
4 (b)

1
2
3 3
Biomass, g/m3

0
1956 1964 1972 1980 1988 1996 2004
1960 1968 1976 1984 1992 2000 Years

Fig. 1. Dynamics of zooplankton (a) abundance and (b) biomass in the Rybinsk Reservoir in 1956–2005 (Here and in Figs. 2⎯5,
smoothed curves with 3year averaging are given; here and in Fig. 2, (1) “peak” abundance in June, (2) in July⎯August, (3) mean
for May–October).

June and the community biomass in late summer were
found to be near those in the 1970s (Table 4). In June,
the values of Btot in the most of water area were
<2 g/m3. The main cause for the decrease in the peak
Btot of zooplankton in the Rybinsk Reservoir were the
low abundance and the small (~0.6 mm on the aver
age) body size of the mass crustacean species Bosmina
longispina [15]. It is only in July 2005, that high Btot of
zooplankton (on the average ~3 g/m3) was recorded,
which was comparable with the value for June in the
1980s (Table 3), and even higher than that in the
Volzhskii pool [15]. This was caused by a shift in the
seasonal peak in biomass from June to July because of
the cold and prolonged spring, as well as a high abun
dance of daphnia (Daphnia galeata accounted for 65%
of Btot).
Zooplankton biomass in June, BtotVI, and August,
BtotVIII, is closely related to the mean over the season
Btot(V–X), with Pearson correlation coefficients of 0.69
and 0.43, respectively (p < 0.05). Data of regular
observations at “standard” stations in 1981⎯1995
(n = 89) were used to obtain regression models for the

WATER RESOURCES Vol. 37 No. 5 2010

 LONGTERM ZOOPLANKTON DYNAMICS IN THE RYBINSK RESERVOIR
assessment of Btot(V–X) in the center of the reservoir for
the current period of biomass decline
Btot(V–X) = 0.84 + 0.23 BtotVI (F = 78.3, R2 = 47%), (1)
Btot(V–X) = 0.96 + 0.35 BtotVIII (F = 19.9, R2 = 18%). (2)
The calculation of Btot(V–X) by these equations
showed that the 95% confidence interval contains 47–
53% of its measured values, 27–33% of calculated
Btot(V⎯X) values are overestimates, while 20% are
underestimates (Table 5). Overall, the use of these
equations yields a satisfactory estimate of Btot(V⎯X) in
the absence of a complete seasonal observation series.
To reveal the main factors that determine the long
term dynamics in Btot of zooplankton in the reservoir,
we considered the following parameters: mean chloro
phyll “a” concentration (Chl, μg/l according to [28,
32])—an integral characteristic of trophic resources
for zooplankton; mean water temperature for May⎯
October (Twat, C)—a characteristic of the rate of biotic
processes; total precipitation on the watershed (Pre
cipitation, mm/year, according to Hydrometeoservice
data); and the volume of water inflow into the reservoir
(Inflow, km3/year according to [24])—factors that
reflect the allochthonous input of organisms, the
amount of organic matter (OM) delivered from the
watershed and the general dynamics of allochthonous
food resources; the date of beginning (tenday inter
val/month) and the rate of reservoir filling (Rate,
cm/day, according to [24])—the characteristics of ini
tial conditions of the season; the coefficient of con
ventional water exchange ( α , year–1, calculation by
[3, 23, 24])—the characteristic of flowage and vari
ability of the habitat.
For the major portion of the observation period,
Ntot and Btot of zooplankton positively correlated with
the dynamics of phytoplankton chlorophyll and Twat of
the reservoir (Table 6). Btot closely correlated with
most characteristics of hydrological regime of the res
ervoir. Stepbystep regression analysis showed that
variations in Btot were mostly determined by chloro
phyll dynamics (R2 = 66%, F = 46.9), while the con
tribution of water temperature variations (R2 = 4%,
F = 4.0) and the hydrological regime characteristics of
the reservoir (Inflow + α , R2 = 11%, F = 4.2⎯5.7) was
much less. The proposed regression model accounts
for >80% of variations in Btot and can be used for
approximate assessment of the amount of zooplank
ton in the reservoir. The equation for biomass predic
tion has the form
Btot = 0.07 Chl + 0.05 Inflow + 0.13 Twat – 0.47 α – 2.03.
With the seasonaveraged chlorophyll concentra
tion in the current period (2000–2003) of 1.4 ±
1.3 μg/l (calculation according to [28]), Twat mean for
the icefree period of 13.8°C, mean inflow of
32.7 km3/year [22, 24] and mean annual water
exchange coefficient of 1.9 year–1 [23], the calculated
zooplankton biomass in the center of reservoir varies
within 1.2–1.6 g/m3 (1.3 ± 0.2 on the average for the
season) (with the confidence interval of 95%). This is
WATER RESOURCES Vol. 37 No. 5 2010
689
Table 4. Zooplankton biomass variations in the center (“stan
dard” stations) of the Rybinsk Reservoirin 1956–2004
Biomass, g/m3
Period, years
June August Mean (V–X)
1956–1969 0.5–1.7 <1 0.4 ± 0.03
1970–1980 0.8–2.8 0.4–1.8 0.7 ± 0.05
1981–1990 2.0–4.6 0.9–2.1 1.6 ± 0.11
1991–2000 1.4–2.7 0.5–1.3 1.1 ± 0.06*
2001–2004 0.5–2.7 0.5–1.3 –
* the mean was calculated for 1991–1997.
close to its real values; e.g., the mean Btot at “standard”
stations was 1.2 g/m3 (Table 6).
Hierarchy of Periodicity in Zooplankton Dynamics:
Reversible and Irreversible Changes
Most natural systems have hierarchic structural
and functional organization: each element (process)
of the system is a part of the system (process) of higher
level [43]. Each system features successions accompa
nied by changes in the composition and structure of
communities, the rate of OM production and destruc
tion processes. Anthropogenic factors can stimulate
the main (autogenic) succession of the ecosystem,
hamper it, or change the direction of succession vector
[30]. The direction of changes in communities is com
monly associated with powerful ecosystem processes,
such as eutrophication and deeutrophication (olig
otrophization) [30, 45], which can have natural or
anthropogenic nature.
The identification of anthropogenic disturbances
in community dynamics is hampered by the imposi
tion of quasicyclic periodicities of different orders.
Therefore, the direction of succession can be better
estimated by qualitative changes. The trends in quan
titative characteristics can be discussed only within the
limits of some cycle. Changes in the structure and
abundance of zooplankton in the Rybinsk Reservoir,
irreversible within half a century of observations, can
be part of a cycle of higher order and may become
reversible in the future. However, 50 years is a period
long enough for revealing some directional and, within
this period, irreversible, changes in the composition,
structure, and amount of zooplankton.
Periodicities (reversible changes) of three orders
were revealed for zooplankton in the Rybinsk Reser
voir. Cycles 6⎯7 years in length are typical of the
dynamics of species diversity; 10⎯11 years, of total
abundance, biomass, and characteristics of the taxo
nomic structure of the community; slow variations
(~20 years) were recorded in zooplankton biomass and
690 LAZAREVA

Table 5. Btot(V–X) in the central Rybinsk Reservoir estimat numbers [13], which in the study period were recorded
ed by equations (1) and (2) in the beginning of each decade [32]. Almost synchro
nously with the dynamics of solar radiation input are
Btot(V–X), g/m3
Year peaks of chlorophyll content of phytoplankton (1972–
measured* calculated by (1)** calculated by (2)** 1973, 1983–1984, 1994–1995) [32].
1981 1.9 ± 0.24 1.7 (1.6–1.7) 1.4 (1.3–1.5)
Minimums of Ntot and Ntot were recorded in colder
years, while peaks of Ntot in years with maximum Twat
1982 1.3 ± 0.18 1.4 (1.3–1.5) 1.3 (1.2–1.4) (Fig. 2). Before 1985, Ntot peaks coincided with rises in
1983 1.8 ± 0.16 1.6 (1.5–1.6) 1.7 (1.5–1.8) chlorophyll “a” concentration in phytoplankton,
1984 2.1 ± 0.25 2.0 (1.9–2.1) 1.5 (1.4–1.6) while after this year, the dynamics of chlorophyll and
1985 1.3 ± 0.16 1.3 (1.2–1.4) 1.5 (1.4–1.6)
zooplankton Ntot and Btot were less correlated (Fig. 3).
In the same period, the successions of these commu
1986 1.1 ± 0.11 1.5 (1.4–1.6) 1.6 (1.5–1.7) nities, associated with the eutrophication of the sys
1987 1.3 ± 0.19 1.8 (1.7–1.9) 1.5 (1.4–1.6) tem, were found to diverge: some signs of eutrophica
1988 2.0 ± 0.32 1.9 (1.8–2.0) 1.7 (1.6–1.8) tion were recorded in phytoplankton, and signs of de
1989 1.7 ± 0.14 1.3 (1.3–1.4) 1.7 (1.6–1.8)
eutrophication, in zooplankton [20, 42]. Reversed
phase variations in phytoplankton and zooplankton
1990 1.4 ± 0.19 1.5 (1.4–1.5) 1.4 (1.3–1.5) characteristics were also recorded for Lake Krasnoe
1991 1.3 ± 0.13 1.4 (1.4–1.5) 1.6 (1.5–1.8) [40].
1992 1.3 ± 0.14 1.4 (1.3–1.5) 1.2 (1.1–1.4) The periodicity of variations in species wealth,
1993 1.2 ± 0.16 1.2 (1.1–1.3) 1.3 (1.1–1.4) size⎯mass structure, and biomass of zooplankton is
1994 1.0 ± 0.18 1.2 (1.1–1.3) 1.4 (1.3–1.5) close to that typical of river runoff in the Upper Volga
basin. Runoff variations with periods of 2⎯4, 8⎯12, and
1995 1.3 ± 0.21 1.3 (1.2–1.4) 1.4 (1.3–1.5) 19⎯33 years [22] are known; some researchers also
1996 – 1.2 (1.1–1.3) 1.4 (1.3–1.5) identify 6⎯7year cycles [37]. The maximal number of
1997 – – 1.4 (1.3–1.5) species in the sample and the total for vegetation
1998 – – 1.1 (1.0–1.3) period were recorded in years with the largest water
input into the reservoir and, accordingly, with maxi
1999 – – 1.3 (1.2–1.4) mal water exchange rate [13]. This is quite explicable,
2000 – 1.4 (1.4–1.5) 1.1 (1.0–1.3) since it is water that delivers rare and any additional
2001 – 1.0 (0.8–1.1) 1.3 (1.2–1.4) forms (relative to the list) not only from nearby water
2002 – – 1.4 (1.3–1.5) bodies, but also from littoral zones.
2003 – 1.0 (0.9–1.2) 1.1 (0.9–1.3) Until the mid1970s, zooplankton Btot in the
Rybinsk Reservoir was increasing parallel to decreas
2004 – 1.2 (1.1–1.3) – ing water inflow, while in the later years, it followed
2005 1.2*** 1.3 (1.2–1.4) 1.7 (1.6–1.9) changes in the inflow (Fig. 4). It is possible that varia
Notes: *mean and standard error; tions in water inflow and Btot in the slow 20year cycle
** given in parentheses is the 95% confidence interval; synchronized after filling the Sheksninskoe Reservoir
*** data of E.A. Sokolova (IIWB RAS). in 1963. Currently, the biological runoff from it has a
strong effect on plankton composition and structure in
the Sheksninskii Pool and the eastern part of the Main
Pool of the Rybinsk Reservoir [15, 33, 41]. In the gen
the size–mass structure [13, 20]. The largest domina erally dry period 1963⎯1976, zooplankton Btot was
tion and, accordingly, the least diversity in the com almost three times less and crustacean abundance was
munity correspond to maximal solar activity by Wolf two times less than in the highwater period 1977–

Table 6. Pearson correlation coefficients for the dependence of zooplankton abundance and biomass in the Rybinsk Res
ervoir on environmental factors (smoothed series with 3year averaging, n is observation number)
Factor
Characteristic
Chl. Twater Precipitation Inflow Intensity α
Ntot 0.52* 0.32** 0.21 0.27 –0.15 0.23
Btot 0.81* 0.49* 0.67* 0.69* 0.29 0.56*
n 26 26 26 26 26 26
Notes: * reliable at confidence level <0.05;
** the same at confidence level of 0.10.

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 LONGTERM ZOOPLANKTON DYNAMICS IN THE RYBINSK RESERVOIR 691

1
2 15
3

Abundance, thous. spec./m3

150
14

Temperature, °C
100 13

12

50

11
1954 1959 1964 1969 1974 1979 1984 1989 1994 1999 2004
Years

Fig. 2. Dynamics of zooplankton abundance and water temperature in the Rybinsk Reservoir in 1954–2005. Zooplankton abun
dance (1) in August, (2) mean over May⎯October, (3) water temperature (data of Hydrometeoservice).

1
2
3
15
Abundance, thous. spec./m3

150 Chlorophyll, µg/l

100 10

50
5

1956 1964 1972 1980 1988 1996 2004

1960 1968 1976 1984 1992 2000
Years

Fig. 3. Relationship between variations in zooplankton abundance with changes in chlorophyll “a” concentration in the central
Rybinsk Reservoir in 1956–2005. Zooplankton abundance (1) in August, (2) mean over May–October, (3) chlorophyll “a” con
centration [28, 32].

WATER RESOURCES Vol. 37 No. 5 2010

692
2.0
1.5
Biomass, g/m3
1.0
0.5
0
1957 1965
1953 1961 1969
Fig. 4. Dynamics of zooplankton biomass and annual water inflow into the Rybinsk Reservoirin 1953–2005. Zooplankton biom
ass in (1) August, (2) mean over May–October, (3) annual water inflow [24].
1990 [20]. In the current dry period, which started in
1996 [22], a decrease in zooplankton biomass of the
same order was recorded (Table 5). However, in a
shorter 10year cycle, biomass peaks, recorded in
1963–1964, 1971–1973, 1980–1982, 1987–1988,
1994–1995, and 2005, mostly coincided with dry peri
ods.
Directional variations in zooplankton characteris
tics were recorded against the background of such
quasicyclic oscillations. At the relative abundance of
dominants, they are most clearly seen during the com
parison of dry periods (Table 1), while for the abun
dance and biomass of large taxonomic groups and the
community as a whole, during the comparison of wet
years (Table 7). Reliable trends were established for
some characteristics (Nrot, Ncrust/Nrot) for both dry and
wet periods.
The dynamics of Bz/Bph deserves special discus
sion. It increased twice when wet periods are com
pared and by 20% when dry periods are compared
(Table 7), suggesting that this characteristic depends
mostly on factors external for the ecosystem. Cur
rently, in the center of the reservoir (“standard” sta
tions) it is 0.41 ± 0.03; this is close to the value typical
of eutrophic lakes (0.42 ± 0.07) [2]. An inverse rela
tionship was found to exist between Bz/Bph and the
biomass of infusoria and bacteria in the case of shal
low, mesotrophic Lake Anninskoe (Pskov Province).
This allows us to correlate the dynamics of this char
acteristic with a combination of effects from “above”
and “below” in the microbial trophic chain [5]. In this
LAZAREVA
1
2
3
40
Inflow, km3
30
20
1973 1981 1989 1997 2005
1977 1985 1993 2001
Years
context, an increase in Bz/Bph in the pelagic zone of
the Rybinsk Reservoir may indicate to a weaker role of
the microbial “loop” and to an increase in the role of
the linear pastoral food chain. The cause of such
growth may be the decrease in the mean phytoplank
ton cell volume to <1000 μm3 by the beginning of the
1980s, which is due to the wider occurrence of fine
cell cryptomonas (cell diameter of 5⎯8 μm) and dia
toms of Stephanodiscus genus (5–10 μm), which is
especially pronounced in wet years [42]. Algae of such
size can be directly consumed by zooplankton micro
filterer species, e.g., cladocers of Daphnia genus.
The appreciable difference in both the extent, and
sometimes, the direction of changes in the commu
nity, revealed during the comparison of dry and wet
periods of the hydrological cycle (Tables 1, 7) appear
to be assignable. For lake ecosystems, it was estab
lished that variations in water inflow cause changes in
their bioproduction [7]. The slower water exchange in
dry period enhances the autogenic processes within
the water body and ecosystem eutrophication. Con
versely, the high rate of water exchange in wet years
facilitates the development of allogenic processes [7]
and deeutrophication [20, 26]. It is likely that this
reasoning is true for reservoir ecosystems as well. In
such case changes in zooplankton, revealed based on
the results of comparison of dry periods, can be repre
sented as associated mostly with natural eutrophica
tion of the water body, whereas changes in the com
munity recorded in the comparison of wet periods will
be mostly due to the allogenous succession of the eco
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 LONGTERM ZOOPLANKTON DYNAMICS IN THE RYBINSK RESERVOIR 693

Table 7. Directional changes in general zooplankton characteristics in the Rybinsk Reservoir in 1956–2004 (Bz/Bph is
zooplankton to phytoplankton biomass ratio)

Wet periods (1, n = 94)* Dry periods (2, n = 38)** Trend (+/–), %
Characteristic
1956–1962 1989–1995 1963–1969 1996–2004 1 2

Btot, g/m2 2.9 ± 0.2 6.9 ± 0.4 – – +140 –

Btot, g/m3 0.46 ± 0.04 1.31 ± 0.08 0.79 ± 0.09 0.87 ± 0.12 +185 +10
Ncl 11.2 ± 0.8 16.3 ± 1.4 10.4 ± 2.3 9.9 ± 1.6 +45 –5
Ncop 19.4 ± 1.7 31.7 ± 4.1 20.2 ± 1.7 26.9 ± 4.4 +60 +30
Nrot 79.4 ± 10.4 41.8 ± 5.1 76.7 ± 7.3 26.2 ± 8.6 –90 –90
Ncr/Nrot 0.4 1.1 0.4 1.4 +175 +250
Bz/Bph(V–X) 0.34 ± 0.04 0.74 ± 0.07*** 0.34 ± 0.03 0.41 ± 0.03 +120 +20
Notes: *mean V–X, calculations for seven years in the end of each wet period;
** mean VI (“peak” values), calculations for seven years in the beginning of each dry period;
*** data of 1981–1986.

system caused by changes in climate and watershed
characteristics and anthropogenic impact.
From the beginning of secondary eutrhophication
in the 1970s and up to the late 1980s, the directions of
allogenous and autogenous succession of reservoir
ecosystem coincided. Jointly, the process was defined
as eutrophication associated with growing anthropo
genic impact onto the watershed and the pollution of
the water body, as well as with the natural accumula
tion of readily utilizable OM in bottom sediments [41,
42]. The combination of allogenous factors in the
recent 20 years facilitated ecosystem deeutrophicai
ton. Its first signs in the dynamics of zooplankton
characteristics were recorded as far ago as the 1980s
[20]. The decrease in ecosystem trophic status in terms
of chlorophyll “a” concentration from eutrhopic to
moderately eutrophic was recorded in the late 1990s
[27]. The change in the direction of allogenous succes
sion of the ecosystem was caused by restructuring of
reservoir soil complex—the area of productive gray
silts decreased twice in the wet 1980s, because they
were covered by sand [8]. A decrease in anthropogenic
load onto the watershed because of the lower areas of
arable lands and livestock population. According to
the author’s observations in the regions adjacent to the
reservoir coast, >60% of the arable lands are not pro
cessed and overgrows with forest. The current changes
in zooplankton in the Rybinsk Reservoir are to a
greater extent associated with allogenous factors,
which determine the amplitude and periodicity of
quasicyclic variations in community characteristics.
These changes mostly suggest the deeutrophication
of the ecosystem (Table 7).
In addition to trends in quantitative characteristics,
qualitative changes were found to take place in zoop
lankton structure, generally reflecting the develop
ment of reservoir ecosystem. These changes include
the extension of the faunistic list, change of dominants
(species and taxonomic groups), immigration, spread
ing over the water area, and growing abundance of new
species. Most these changes are caused by a set of fac
tors of both allogenous and autogenous nature. To one
extent or another, they can be due to a cascade effect
in the trophic chain arising as the result of qualitative
changes in other communities of the water body.
For example, the reliable decrease (always double)
in rotifer abundance (Table 7), is most likely due to
their competition with zebra mussel for seston
resources (“bottomup” effects). The rapid spreading
of Dreissena polymorpha (Pallas), D. bugensis
(Andrusov) over the reservoir water area was recorded
in the 1980s⎯1990s [41]. The abundance of veligers of
this mollusk in July⎯August reaches 150⎯210 thou
sand specimens/m3 and is comparable with the
amount of zooplankton; in the Volga and Main pools,
it accounts for up to 83% of their total abundance
(zooplankton filterers + veligers) [15, 18]. The losses
of particulate matter relative to its total content in the
water mass of Lake Naroch (Belarus) increase eight
fold in the presence of zebra mussel [6]. Strictly speak
ing, zebra mussels facilitate deeutrophication only in
the pelagial zone, since they actively precipitate and
accumulate seston on reservoir bed. In such cases, if
the total productivity of the ecosystem does not
decrease, benthification is said to have place, i.e., pro
duction flows pass from planktonic communities to
benthic and periphytic [31].
An example of topdown effect can be the immi
gration and spreading of a new planktoneater—kilka
Clupeonella cultriventris (Nordmann) [41]. This pro

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694 LAZAREVA
cess coincided with a period of low abundance of
another planktoneater—smelt Osmerus eperlanus
Linnaeus, which dominated in the pelagic zone before
the early 1990s [41]. By 2004, the density of kilka
(750⎯3076 specimen/15 min trawling) has not
reached the maximal density of smelt, recorded in the
1970s⎯the 1980s [9]. The successful naturalization of
kilka in the Rybinsk Reservoir was facilitated by the
relatively high (>1 g/m3) zooplankton biomass and the
existence of zones its accumulation, which serve as
both fattening and spawning areas for fish. The con
sumption of zooplankton by kilka is affected by the
biomass of the feed species, the degree of specimen
aggregation, body size, and the visibility of individual
specimens of prey for predators. Kilka selectively con
sumes largesize planktonic crustaceans (of Hetero
cope, Bythotrephes, Leptodora, Cyclops genera) and
serves as a potential competitor for young fish of valu
able species (pike perch, perch, vendace, bream),
which use the same food resources [9].
At the same time, zooplankton of the Rybinsk Res
ervoir currently features the predominance of crusta
ceans (Tables 3, 7), the relatively high abundance of
largesize food species (Table 2), and the large size of
mass species of daphnia and bosmina (the maximal
body lengths of Daphnia galeata and Bosmina long
ispina reach 1.7⎯1.8 and 1.0⎯1.1 mm, respectively).
This fact can be both the cause of the intense spreading
of the immigrant species (kilka) and the consequence
of the low abundance of obligate planktoneaters in
the reservoir.
Zooplankton as an Indicator of the Rate of Reservoir
Ecosystem Eutrophication
The trophic level is determined by the primary pro
duction of autotrophs in the water body and on its
watershed, which closely correlates with the climatic
features of the region [7, 45]. The eutrophication of
water bodies can be natural, i.e., caused by OM accu
mulation in bottom sediments and an increase in the
internal and external biogenic load in geological time
scale, or anthropogenic allogenic, often local, associ
ated with an increase in the input of OM and biogenic
elements from the watershed, resulting from human
economic activity and developing much faster than the
natural eutrophication [35].
Changes in zooplankton characteristics, which
persist over more or less long period (years) is regarded
as a reliable indicator of eutrophication [2, 10, 13, 20,
25, 29, 34, 38]. In lakes at the initial stage of eutroph
ication, the most distinct are changes in the composi
tion of dominating cladoceras (commonly, in the
Bosmina Daphnia direction), accompanied by an
abrupt increase in Ntot and B tot of the community [2].
As eutrophication becomes stronger, deep transforma
tion of the community structure takes place, including
changes in the percent ratios of large taxa (an increase
in the share of Cladocera and Rotatoria, a decrease in
the share of Copepoda, especially, Calanoida), a
decrease in the species diversity and the number of
dominant species, the mass development of eutrophi
cation indicator species (rotifers of Brachionus genus,
crustaceans Chydorus sphaericus, Bosmina longiros
tris, etc.) (Table 8). A reliable positive correlation was
found to exist between the faunistic trophicity index E
of zooplankton and chlorophyll “a” content of phy
toplankton [29].
Parallel to the growth in Ntot, an increase was
recorded in the annual amplitude of Btot variations
(indices Bsum/Bwin, Bmax/Bmin) along with a decrease in
the total stability of the community—a greater vari
ability of all parameters [2, 25]. Rotifers dominate in
hypertrophic lakes, where the mean mass of zooplank
ton organism, wmean, and Btot of the community
abruptly drops in most cases [2, 21, 44]. The trophic
network of zooplankton and the functioning of the
community as a whole also change. Against the back
ground of growing production, the share of filterers
(mostly Cladocera) increases, and the share of preda
tors decreases (B3/B2 index) [2, 10]. The correlation
between zooplankton and phytoplankton becomes
weaker (Bz/Bph index decreases) [2, 21, 44].
Overall, smallsize species with simple life cycles
gain an advantage in the plankton of water bodies dur
ing their eutrophication [30]. Therefore, the decrease
in zooplankton organism size (wmean) is considered by
many researchers to be the main sign of eutrophica
tion [11, 44]. This processes is due not only to an
increase in the abundance of smallsize rotifer species,
but also with a decrease in the linear size of species in
dominant crustacean populations and the replace
ment of largesize filterers (Daphnia sp., Eudiaptomus
sp.) by finer cladoceran forms (Chydorus sphaericus,
Bosmina longirostris). The latter is attributed to the
high concentration of largesize bluegreen algae and
the total mass of OM in eutrophic water bodies, as well
as with the high reproduction rate of the fine cladocer
ans in the presence of excessive amounts of food [11].
The indication of lowland reservoir eutrophication
by zooplankton is currently based on the same indices
as for lakes (Table 8). This is quite justifiable, since,
after the introduction of regulation in a large river,
potamoplankton is rapidly replaced by lacustrine spe
cies complex [33]. The most sensitive to trophic status
change in a reservoir, as well as in lakes, are commu
nity structure characteristics. However, some of them
in reservoirs, unlike lakes, vary within wide limits and
have no distinct correlation with eutrophication. The
most reliable for the assessment of reservoir eutrophi
cation rate are assumed to be the following zooplank
ton characteristics: indices Bz/Bph, Bcyc/Bcal [39],
Nmean/Ntot and wmean [20], as well as changes in domi
nants and the abundances of indicator species of
eutrhophication, their abundance in the dominant
complex, and Btot [20, 34].
The signs of eutrophication by zooplankton in the
Rybinsk Reservoir were most clearly seen in the
WATER RESOURCES Vol. 37 No. 5 2010
 LONGTERM ZOOPLANKTON DYNAMICS IN THE RYBINSK RESERVOIR 695

Table 8. Variations in some zooplankton characteristics during eutrophication and deeutrophication of water bodies

Eutrophication Deeutrophication
Characteristic
lakes, ponds reservoirs reservoirs

Species richness Stot, Sl, IMarg, the number Decreases [2, 10, 38] Decreases [34] –
of dominants
Species diversity Hn, Hb, 1/ISim '' [2, 10, 38] '' [34, 39] –
Dominant composition Changes [2, 21, 29, 38] Changes [20, 34, 39] Changes [20]
Number of eutrophication indicators in Increases [2, 21, 25, 29, 38] Increases [20, 34] Decreases [20]
dominant composition
Abundance and share of Chydorus sphaeri ''[2, 21, 25, 29, 38] '' [20] Varies [*]
cus, Bosmina longirostris
Abundance and share of the genus Increases [2, 21, 25, 38] '' [34] Decreases [*, 20]
Faunistic indices E, E/O Increases [21, 25, 38] Vary or increase [*, 20] Vary [*]
Mean individual mass of specimens wmean Decreases [2, 11, 21, 25, 44] Decreases [*, 20] Increasesò [1, 20]
Total abundance Ntot Increases [2, 10,11, 21, 25, 44] Increases [*, 20, 34] Decreases [*, 20]
Total biomass Btot Increases until hypertrophy [2, '' [*, 20, 34] '' [*, 20]
10, 21, 25, 38]
The share of crustaceans, Bcr/Brot, Ncr/Nrot Decreases [2, 11, 20, 21, 25, 38] Decreases, varies [20, 34, 39] Increases [20]
The share of filterers Bcl/Btot Increases [2, 10, 25] Varies [*, 20] Varies [*, 20]
Interaction between phytoplankton and Weakens (decreases) [2, 21, 44] Weakens (decreases) Strengthens
zooplankton Bz/Bph [*, 39] (increases) [*]
* The author’s unpublished data.

1950s⎯1970s [20]. In the 1970s, zooplankton domi
nant complexes included five eutrophication indicator
species, 50% of dominants in the group of crustaceans
and 30% in rotifer group were taxa that prefer
eutrophic water bodies. Recorded in this period were
low individual mass of a zooplankter (<10 μg) and a
change in dominants in the direction Bosmina long
ispina B. coregoni, Keratella cochlearis
K. quadrata, as is commonly the case in eutrophic
water bodies [2].
The decrease in anthropogenic biogenic load,
resulting from eutrophication control measures and
the rational use of agricultural lands, causes de
eutrophication or reoligotrophization of water bodies
[31]. The trophic level of lakes decreases in the process
of forestation and waterlogging of watersheds
(distrophication, oligotrophization) [1, 45], and as a
consequence of anthropogenic acidification of surface
water [19]. Deeutrophication can also take place in
the hydroclimatic cycle of water bodies, because of a
decrease in the internal biogenic load in wet years [20,
26]. This process gained in significance in European
Russia water bodies since the 1980s–1990s [12, 19,
20]. It was enhanced by the combination of climate
dynamics with the general decline in the industry and
agriculture in the region.
The signs of deeutrophication by zooplankton
indices were especially distinct in the wet phase of
hydrological cycle of the reservoir in the 1980s⎯1990s
[20]. In this period, the number of zooplankton dom
inants (eutrophication indicators) decreased from five
to two species with a drop in their abundance. Changes
in species of dominant complex were inverse to those
typical of eutrophication Bosmina coregoni
B. longispina, Daphnia cucullata D. galeata. The
decrease in the trophic status in the 1980s⎯1990s was
also suggested by the low abundance of rotifers, an
increase in diaptomides, and the values of Ncrust/Nrot >
1 (Table 7).
In the current, dry phase of the climate cycle (since
1996), the Rybinsk Reservoir demonstrated an
increase in wmean, a decrease in Nrot, Ntot, and Btot of
zooplankton, as well as other signs of deeutrophica
tion (Tables 7, 8), caused by a decrease in ploughed
areas and livestock population in the watershed and,
perhaps, enhanced by the filtration activity of zebra
mussels. At the same time, the change of dominant
cladoceran species in summer plankton in the direc
tion Bosmina sp. Daphnia galeata suggests eutroph
ication—a typical process for this stage of climate cycle
[7, 20]. By Shannon index (Hb = 2.16–2.69 bit) and fau
nistic trophicity index (E = 0.5⎯0.8) of zooplankton,

WATER RESOURCES Vol. 37 No. 5 2010

696 LAZAREVA
the Rybinsk Reservoir in the 1980s⎯1990s was classi
fied as mesotrophic [13]. Variations in Btot before 2005
never passed beyond the limits specified for
mesotrophic waters [15, 20, 34]. By phytoplankton
characteristics, the reservoir is regarded as moderately
eutrophic since the late 1990s [27, 28]; the zooplank
ton and phytoplankton biomass ratio (Bz/Bph = 0.41 ±
0.03) suggests the eutrophic status of the ecosystem.
Thus, the current parameters of zooplankton and phy
toplankton communities yield different estimates of
the trophic level of the Rybinsk Reservoir ecosystem.
CONCLUSIONS
Observations in the Rybinsk Reservoir during
50 years revealed two categories of variations: revers
ible variations in community structure, abundance,
and biomass and changes that are irreversible in the
given time interval (trends). Dominant species experi
ence reversible changes every 10 and 20 years, which
can be referred to as cyclic successions. They take
place by analogy with seasonal succession, but not in
the longterm rather than annual cycle of community
development. In the early XXI century, the previously
established periodicity of variations in abundance
(~10 years) and biomass (~10 and 20 years) still per
sist. Abundance maximums were recorded in the early
1960s, in the 1970s and the 1980s, and in the late 1980s
and 1990s; a new rise in this characteristic was
recorded since 2004.
In the 10year cycle, zooplankton biomass maxi
mums coincide with those in abundance. In the 20
year cycle, the largest biomass of community was
recorded in the 1980s; its values from the late 1990s to
the present time are close to those in the 1970s.
Regression models were constructed, making it possi
ble to assess the mean zooplankton biomass over the
vegetation period based on data for individual summer
months and to predict the biomass by measured phy
toplankton chlorophyll concentration and the hydro
thermal characteristics of the reservoir.
The directed changes in the community take place
simultaneously with periodic fluctuations of its char
acteristics; they reflect zooplankton succession, asso
ciated with the evolution of reservoir ecosystem under
the effect of a set of autogenic and allogenic factors.
Invasion and spreading of new and previously rare spe
cies and irreversible changes in dominants were
recorded. Since the early 1990s up to the present time,
the reservoir demonstrates low rotifer abundance;
crustaceans dominate in zooplankton in terms of
abundance and biomass. The amount of relatively
largesize forms is recorded among them, which is
favorable for planktoneating fish.
Many indices of taxonomic, trophic, and size⎯
mass structure of zooplankton, tested in lakes, can be
used as indicators of eutrophication/deeutrophicai
ton of ecosystems in lowland reservoirs and deserve
wider use for assessing their environmental state.
However, the trophic status of the Rybinsk Reservoir
ecosystem by zooplankton characteristics is lower
(mesotrophic) than estimates by phytoplankton
(moderately eutrophic).
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1. Abrosov, V.N., Zonal’nye tipy limnogeneza (Zonal Types
of Limnogenesis), Leningrad: Nauka, 1982.
2. Andronikova, I.N., Strukturnofunktsional’naya orga
nizatsiya zooplanktona ozernykh ekosistem (Structural⎯
Functional Organization of Zooplankton in Lake Eco
systems), St. Petersburg: Nauka, 1996.
3. Bakulin, K.A, Morphometrical Characteristics of the
Rybinsk Reservoir, in Biologicheskie i gidrologicheskie
faktory mestnykh peremeshchenii ryb v vodokhranilish
chakh (Biological and Hydrological Factors of Local
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