water
Review
Migrations of Young Fish in Anthropogenically Transformed
Rivers: Responses of Cyprinids and Percids to Ecological Filters
and Barriers
Dmitrii S. Pavlov, Vasilii V. Kostin and Victor N. Mikheev *






Citation: Pavlov, D.S.; Kostin, V.V.;
Mikheev, V.N. Migrations of Young
Fish in Anthropogenically
Transformed Rivers: Responses of
Cyprinids and Percids to Ecological
Filters and Barriers. Water 2021, 13,
1291. https://doi.org/10.3390/
w13091291
Academic Editor: Ismail Albayrak
Received: 9 March 2021
Accepted: 27 April 2021
Published: 4 May 2021
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Attribution (CC BY) license (https://
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4.0/).
Water 2021, 13, 1291. https://doi.org/10.3390/w13091291
A.N. Severtsov Institute of Ecology & Evolution, Russian Academy of Sciences, 33 Leninskii Prosp.,
119071 Moscow, Russia; acad.pavlov@gmail.com (D.S.P.); povedenie@yandex.ru (V.V.K.)
* Correspondence: vicnikmik@gmail.com; Tel.: +7-(916)-808-1189
Abstract: Downstream migration (DSM) of larvae and fry is an important phase of the life cycle
of fish as it allows them to disperse, and it increases the size and diversity of the populations via
them extending rearing grounds, exchanging genes, and avoiding competition and cannibalism.
Two numerous and diverse fish families of the Eurasian rivers, Cyprinidae and Percidae, are well
adapted to the conditions of the riverine continuum. Having said that, the regulation of rivers
(construction of dams and water reservoirs) drastically changes their hydrology and topography. In
this work, we argued that novel conditions of transformed river habitats influence the DSM of young
cyprinids and percids in different ways. The published results on fish DSM and spatial distribution
in nine European reservoirs (Russia, Kazakhstan, Czech Republic, Bulgaria) in comparison with
untransformed rivers were reanalyzed from the viewpoint of this argument. Changes in the major
characteristics of DSM of young cyprinids and percids, i.e., intensity, diel (24-h period), and seasonal
patterns of migrations, as caused by anthropogenic transformation of the rivers, were revealed. We
found that the novel ecological barriers and filters associated with different parts of water reservoirs
differently influence the lateral and longitudinal movements, and the diel and seasonal dynamics
of DSM of cyprinids and percids. These effects result in significantly more intensive emigration of
young percids compared to cyprinids from reservoirs with deep-water intakes. At the scale of the
whole regulated river, the morphological complexity (topography) of the reservoir plays a pivotal role
in controlling the intensity of the DSM of young fish. Measures for the conservation and restoration
of percid and cyprinid populations should be different.
Keywords: young fish; cyprinids; percids; dispersal; downstream migration; regulated rivers
1. Introduction
Most organisms disperse from a natal/breeding site, which enables the exploitation of
spatially and temporally variable resources in continually changing natural habitats [1,2].
Downstream migration (DSM), or downstream drift, is an important phase of the life cycle
of fish that allows young fish to disperse, and it increases the size and diversity of fish
populations [3,4]. DSM allows young fish to use flowing water to extend the range of
the fish population, exploit increased trophic resources [5–7], exchange genes [8], avoid
cannibalism and competition at spawning grounds [9], and enhance habitat connectivity
and community stability [7,10].
The DSM of young fish is not only a passive dispersal by water flows but a structured
process controlled by hydraulic and behavioral drivers [11]. In natural rivers, fish are well
adapted to the topography and hydraulic structure of the river, and they efficiently use
the main channel as a migratory habitat and the inshore zone as a resting habitat [11].
During migrations, which usually last for days, young fish repeatedly leave the inshore
zone and enter the main channel on a diel (i.e., 24-h period) basis [3,12,13]. Ecological
barriers for downstream migrating young fish are related to the longitudinal heterogeneity
https://www.mdpi.com/journal/water
Water 2021, 13, 1291 2 of 13

of the riverine continuum [14,15]. Fish usually overcome such barriers. The entire process
of DSM comprises a sequence of active and passive components. Short periods of decision
making when migrants enter the drift at dusk and return to the inshore retention zone at
dawn are of particular importance in the control of DSM. In a previous study, we showed
that such key events are associated with high gradient zones (e.g., interfaces between
habitats of residence and migration habitats) [11].
The transformed habitat structure in regulated rivers may influence the role of fish
behavior in DSM [6,16]. Novel physical objects create ecological barriers [17,18] and
filters [19], which substantially change conditions for DSM. Ecological barriers, which
hamper and modify migrations of young fish, are associated with physical–chemical
gradients (marginal zones of habitats, frontal zones). The permeability of a particular
barrier for different species or age groups may be different. In this case, we consider this to
be an ecological filter, one that differentially influences different groups of migrants [19].
A question to pose here is, do these barriers and filters exert similar impacts on the most
abundant and diverse downstream migrants, cyprinids and percids, which usually form
the core of fish communities?
In European Russia, cyprinid and percid fishes are the most common and abundant
inhabitants of both natural and regulated rivers [20]. Their spawning periods and spawn-
ing grounds partially overlap [21]. In natural rivers, the most intensive DSM of the larvae
of common cyprinids (roach, Rutilus rutilus (Linnaeus, 1758); bream, Abramis brama (Lin-
naeus, 1758); bleak, Alburnus alburnus (Linnaeus, 1758)) and percids (perch, Perca fluviatilis
Linnaeus, 1758; zander, Sander lucioperca (Linnaeus, 1758); ruff, Gymnocephalus cernuus
(Linnaeus, 1758)) occurs in June. On a diel scale, the DSM of both cyprinids and percids
reaches its highest intensity during the night and twilight [21]. It appears that the ability
of young percids and cyprinids to efficiently use structured habitats of natural rivers and
migrate freely along the whole river continuum is similar. With this in mind, does their
migratory behavior change in regulated rivers where fish should overcome vast open
water and dams? A large number of publications has covered various aspects of the fish
ecology in regulated rivers. Most of them concern studies on the three families: Salmonidae,
Cyprinidae, and Percidae [22]. However, downstream migrations are not among the main
topics of such studies, especially for cyprinids and percids.
The construction of dams and water reservoirs leads to a variety of serious transfor-
mations of the river continuum. This large-scale discontinuity creates abrupt lotic–lentic
(river–lake) transitions, which radically modify conditions for DSM [4,20,23]. These mod-
ifications change not only the flow patterns that provide downstream movement, but
the whole complex of contiguous habitats where migrating fish undertake repeating diel
moves between migratory and resting habitats. The emigration (leaving a habitat) of young
fish from the water reservoir through the passages associated with dams is especially
important because of the high mortality, damage, and irrevocable withdrawal of fish from
the reservoir. Because of the similarity of the main characteristics of the DSM of young
cyprinids and percids in natural rivers, we may expect similar patterns of their migration
both within water reservoirs and downstream of dams. However, the intensive emigration
through dams has thus far been found in young percids but not in cyprinids [4,21]. Is this
difference typical and consistent? If so, what ecological and behavioral mechanisms are
responsible for this difference between cyprinids and percids in regulated rivers?
Modifications of downstream migrations caused by transformations of habitat struc-
ture and hydraulic regime in regulated rivers were analyzed and discussed mainly at
the level of “total ichthyoplankton” (all larval fish) without emphasizing the biological
(ecological and behavioral) specificity of particular categories of migrating fish [5,11,18,19].
Generally, similar patterns of downstream migrations of the two most abundant groups of
European riverine fish, cyprinids and percids, differ in the spatial distribution of migrating
larvae [11]. We expect that such differences, which under conditions of natural rivers do
not lead to differences in the main patterns of DSM of cyprinids and percids, may cause a
significant differentiation under the conditions of regulated rivers. To study this argument,
 Water 2021, 13, 1291 3 of 13

Water 2021, 13, x FOR PEER REVIEW 3 of 13

may cause a significant differentiation under the conditions of regulated rivers. To study
we compare data on DSM of young cyprinids and percids obtained at diel and seasonal
this argument, we compare data on DSM of young cyprinids and percids obtained at diel
scales in nine regulated rivers in Russia, Kazakhstan, Bulgaria, and the Czech Repub-
and seasonal scales in nine regulated rivers in Russia, Kazakhstan, Bulgaria, and the
lic [21,24]. We reanalyze this information to reveal changes in the major characteristics of
Czech
DSMRepublic
of young[21,24]. Weand
cyprinids reanalyze
percidsthis information
(intensity, diel, to
andreveal changes
seasonal in theofmajor
patterns char-
migrations)
acteristics of DSM of young cyprinids and percids (intensity, diel, and seasonal
caused by anthropogenic transformation of the rivers. The most abundant field data on the patterns
oftemporal
migrations)
andcaused
spatialby anthropogenic
patterns transformation
of downstream migrations of the rivers.cyprinids
of young The mostandabundant
percids
field
are data on the
obtained fortemporal
the Volgaand
andspatial patterns
Sheksna riversof
indownstream
European Russia migrations of 1970s
from the younguntil
cypri-
the
nids
2000sand(Figure
percids
1).are obtained for the Volga and Sheksna rivers in European Russia from
the 1970s until the 2000s (Figure 1).

8
4 5

1
3 6
2
7

Figure 1. Locations of the studied reservoirs. 1—Mostishte, Czech Republic; 2—Al. Stam-
boliiskii, Bulgaria; 3—Ozerninskoe, Moscow Region, Russia; 4—Ivan’kovskoe, Upper Volga,
Russia;
Figure 5—Sheksninskoe,
1. Locations Vologda
of the studied Region,1—Mostishte,
reservoirs. Russia; 6—Volgogradskoe,
Czech Republic; Lower Volga, Russia;
2—Al. Stamboliiskii,
7—Kapchagaiskoe, Kazakhstan; 8—Ust’-Khantaiskoe, Russia.
Bulgaria; 3—Ozerninskoe, Moscow Region, Russia; 4—Ivan’kovskoe, Upper Volga, Russia; 5—
Sheksninskoe, Vologda Region, Russia; 6—Volgogradskoe, Lower Volga, Russia; 7—Kapchagaiskoe,
Our study
Kazakhstan; aims to compare
8—Ust’-Khantaiskoe, the modifications of downstream migrations in young
Russia.
cyprinids and percids in regulated rivers. More specifically, we study (1) the changes of
theOur
spatiotemporal
study aims structure
to compareof the
the downstream
modificationsmigration of the two
of downstream fish groups
migrations within
in young
water reservoirs in comparison with natural rivers; (2) the difference in spatiotemporal
cyprinids and percids in regulated rivers. More specifically, we study (1) the changes of
patterns
the of emigration
spatiotemporal between
structure of theyoung cyprinids
downstream and percids
migration from
of the twowater reservoirs;
fish groups and
within
water reservoirs in comparison with natural rivers; (2) the difference in spatiotemporalin
(3) the role of ecological barriers and filters as modifiers of the downstream migration
anthropogenically
patterns of emigrationtransformed rivers.cyprinids and percids from water reservoirs; and
between young
(3) the role of ecological barriers and filters as modifiers of the downstream migration in
2. Spatiotemporal Structure of the Downstream Migration in Natural Rivers and
anthropogenically
Water Reservoirstransformed rivers.
River regulation due to the construction of dams and spacious water reservoirs causes
2. Spatiotemporal Structure of the Downstream Migration in Natural Rivers and Wa-
multiple changes in hydrology and in the habitat structure of riverine continuum. Lotic–
ter Reservoirs
limnetic transformations, large water bodies with varying depth and currents, and modified
River regulation
transition zones betweendue to the construction
habitats of residenceofanddams and spacious
migration habitatswater reservoirs
radically change
causes multiple changes in hydrology and in the habitat structure of riverine continuum.
conditions of the downstream migration for young fish, creating ecological barriers of dif-
Lotic–limnetic transformations,
ferent permeability large water
[18,19]. Considering thebodies withofvarying
influence depth and
these barriers currents,
on the and
downstream
modified transition
migrations zones
of fish in between
rivers habitats
of European of residence
Russia andAsia,
and Central migration habitats radically
we discovered that novel
change conditions of the downstream migration for young fish, creating ecological
ecological barriers are associated with physical structures of different scales. Locally, barri-
fish
ers of different permeability [18,19]. Considering the influence of these barriers
migrating in a reservoir face extended transition zones between the migratory habitat (open on the
downstream migrations
part of the reservoir) of the
and fishhabitat
in rivers
of of European
residence Russia and Central
(shallow-water inshoreAsia,
zone).we discov-
Compared
ered that novel ecological barriers are associated with physical structures
with natural rivers, the extended transition zone makes repeated lateral diel movements of different
scales. Locally, fish migrating in a reservoir face extended transition zones between the
Water 2021, 13, x FOR PEER REVIEW 4 of 13

Water 2021, 13, 1291 migratory habitat (open part of the reservoir) and the habitat of residence (shallow-water4 of 13
inshore zone). Compared with natural rivers, the extended transition zone makes re-
peated lateral diel movements between the habitats more difficult. Another strongly mod-
ified area within a reservoir is associated with the dam, which controls downstream water
between
flows and the
thushabitats more
influences thedifficult. Another
downstream strongly
migration of modified area
young fish. Onwithin a reservoir
the scale of the
is associated with the dam, which controls downstream water flows and thus influences
whole reservoir, vast bodies of water with changeable flow patterns may function as a
the downstream migration of young fish. On the scale of the whole reservoir, vast bodies
large-scale ecological barrier disturbing the unidirectional migratory route of drifting fish.
of water with changeable flow patterns may function as a large-scale ecological barrier
The spatiotemporal structure of the downstream migration is controlled by the interaction
disturbing the unidirectional migratory route of drifting fish. The spatiotemporal structure
between the behavior of migrating fish and their biotic and abiotic environment [5,6].
of the downstream migration is controlled by the interaction between the behavior of
migrating fish and their biotic and abiotic environment [5,6].
2.1. Diel Changes of Downstream Migrations
2.1. The
Dieldiel rhythms
Changes in the behavior
of Downstream of migrating fish and their interactions with small-
Migrations
scale hydrological gradients and topography
The diel rhythms in the behavior of migrating can shape the temporal
fish and patternswith
their interactions of down-
small-
stream migration in young fish [3,4,6,21]. The exact behavioral
scale hydrological gradients and topography can shape the temporal patterns of down- and ecological mechanisms
driving
stream diel patterns
migration in downstream
in young migration
fish [3,4,6,21]. are not
The exact clearly understood.
behavioral and ecological However, the
mechanisms
intensity of illumination, water turbidity, and avoidance of predation
driving diel patterns in downstream migration are not clearly understood. However, the pressure are among
the key factors
intensity controlling the
of illumination, watercircadian
turbidity,rhythm of drift [5,11].
and avoidance Peak densities
of predation pressureof migrating
are among
fish
the key factors controlling the circadian rhythm of drift [5,11]. Peak densitiesday,
in unregulated rivers are usually recorded during the night. During the the mi-
of migrating
gration
fish in unregulated rivers are usually recorded during the night. During the day,Mi-
habitat (main channel) is virtually free of downstream migrants (Figure 2a). the
gration
migration habitats in regulated
habitat rivers (open
(main channel) water of
is virtually reservoirs)
free contain migrants
of downstream larval cyprinids
(Figureand2a).
percids
Migration bothhabitats
day and in night (Figure
regulated 2b).(open
rivers Comparing
water ofthese two contrasting
reservoirs) patterns,
contain larval we
cyprinids
argue that riverine
and percids both day migrants,
and night both cyprinids
(Figure 2b). and percids,these
Comparing easily leave
two the migratory
contrasting habi-
patterns, we
tat at dawn and enter the shallow-water inshore zone. Lateral movements
argue that riverine migrants, both cyprinids and percids, easily leave the migratory habitat allow them to
cross
at dawna narrow flow the
and enter gradient (interface)inshore
shallow-water between the Lateral
zone. habitats. This task seems
movements allow significantly
them to cross
more difficult
a narrow flowfor larvae, (interface)
gradient which migrate in the
between thereservoir
habitats.ofThisa regulated
task seems river; the spreading
significantly more
ofdifficult
the interface (gradient
for larvae, which zone) between
migrate in themigratory
reservoir and of a residence
regulated habitats
river; thecreates a barrier,
spreading of the
which
interfacefunctions
(gradient as azone)
hardly permeable
between obstacle
migratory andfor migrants.
residence A largecreates
habitats number of migrants
a barrier, which
functions
stay as a hardly
in the open water permeable
during both obstacle
day and fornight,
migrants.
and theyA large
driftnumber of migrants stay
either downstream to-
in the open water during both day and night, and they drift
wards the dam or in other directions, depending on the prevailing currents, the main flow,either downstream towards
thewind-induced
and dam or in other directions, depending on the prevailing currents, the main flow, and
currents.
wind-induced currents.

A B
100
Concentration of migrants, %

50

0
Cyprinids Percids Cyprinids Percids
Figure 2. Mean relative concentration of young cyprinids and percids in migration habitats of the
Figure
Upper2.Volga
Mean(A)
relative concentrationReservoir
and Ivan’kovskoe of young (B)
cyprinids
duringandthe percids
day (light) in migration
and night habitats of the
(dark part of the
Upper Volga (A) and Ivan’kovskoe Reservoir (B) during the day (light) and night (dark part of the
bars). Differences between the day and night values are significant in the river and nonsignificant
bars). Differences between the day and night values are significant in the river and nonsignificant
in the reservoir (chi-squared test: p < 0.001 and p > 0.05, respectively). Number of fish sampled:
in the reservoir (chi-squared test: p < 0.001 and p > 0.05, respectively). Number of fish sampled:
cyprinids—2120ind.,
cyprinids—2120 ind., percids—3810
percids—3810 ind.
ind. DataData
fromfrom Pavlov
Pavlov et al.et[21,24].
al. [21,24]. See Supplement
See Supplementary 1 for
Mate-
more details.
rials for more details.
Water 2021, 13, 1291 5 of 13

Water 2021, 13, x FOR PEER REVIEW 5 of 13

2.2.
2.2.Seasonal
SeasonalChanges
ChangesofofDownstream
DownstreamMigrations
Migrations
Seasonal
Seasonalpatterns
patternsofofthethedownstream
downstreammigration
migrationare areinfluenced
influencedby byaavariety
varietyofofbiotic
biotic
and
andabiotic
abioticfactors,
factors,such
suchasasthethetiming
timingand andsuccess
successofofreproduction,
reproduction,water waterdischarge,
discharge,and and
temperature.Depending
temperature. Dependingononthe thespecies
speciesrichness,
richness,thetheuni-,
uni-,bi-
bi-and
andmultimodal
multimodalseasonal
seasonal
patternsare
patterns arecharacteristic
characteristicofofdifferent
differentrivers
rivers[7,11,21,25].
[7,11,21,25].
Larvaeofofriverine
Larvae riverinecyprinids
cyprinidsand andpercids
percidsmigrate
migratemostmostintensively
intensivelyininJune
June(Figure
(Figure3).3).
In the Ivan’kovskoe Reservoir of the Volga River, the period of downstream
In the Ivan’kovskoe Reservoir of the Volga River, the period of downstream migration is migration
is more
more prolonged
prolonged thanthan in river
in the the river (Figure
(Figure 4).seen
4). As As seen in Figure
in Figure 4b, within
4b, within this period,
this period, i.e.,
i.e., June–September,
June–September, a mucha much
higherhigher concentration
concentration of young
of young percidspercids was recorded
was recorded in the
in the open
open (migration
water water (migration
habitat)habitat)
than inthan in shallow
shallow water water (habitat
(habitat of residence).
of residence). The opposite
The opposite pat-
pattern was observed in cyprinids—they were much more abundant
tern was observed in cyprinids—they were much more abundant in the inshore zone in the inshore zone
of
of residence (Figure 4a). During the period of downstream migration
residence (Figure 4a). During the period of downstream migration in the reservoir, con- in the reservoir,
consistently
sistently moremorethanthan
50%50%of theof the
youngyoung percids
percids werewere recorded
recorded inin themigration
the migrationhabitat
habitatinin
comparison with those in the habitat of residence; usually less than
comparison with those in the habitat of residence; usually less than 5% of cyprinids were5% of cyprinids were
recordedininthe
recorded themigration
migrationhabitat
habitat(see(seeSupplement
Supplement11for formore
moredetail).
detail).

100
A
75
Concentration of migrants, % of max

50

25

0
100 B

75

50

25

0
I II III IV V VI VII VIII IX X XI XII
Months
Figure 3. Seasonal patterns of the downstream migration of cyprinids (A) and percids (B) in the Upper
Figure
Volga.3.Concentrations
Seasonal patterns of the downstream
are expressed migration
as percentages of theofmaximum
cyprinidsvalue.
(A) and
Nopercids (B) in
significant the
difference
Upper Volga. Concentrations are expressed as percentages of the maximum value. No significant
between cyprinids and percids. Kolmogorov–Smirnov test: p > 0.01. Data from Pavlov et al. [21].
difference between cyprinids and percids. Kolmogorov–Smirnov test: p > 0.01. Data from Pavlov et
al. [21].This difference produces a prerequisite for more intensive drift of the young percids
than cyprinids towards the dam. The functioning of ecological barriers at different scales,
which hamper both the lateral and longitudinal movements of migrants, could be one of
the reasons prolonging the period of migration. We argue that the whole water body of
the reservoir, working as a macroscale barrier, restricts the intensity and efficiency of the
downstream migration through the lowering of its speed and change of direction. This
is primarily caused by highly variable and unpredictable flow regimes [22,26], which in
turn is influenced by the variability of the wind-induced currents [27] and the topography
of reservoirs [18]. Within morphologically complex reservoirs, numerous bays of various
sizes may operate as a sort of ecological trap, which accumulates migrating fish [28]. As a
result of the differences between young cyprinids and percids in their characteristics of
Water 2021, 13, 1291 6 of 13

migration through the water reservoir, more percids than cyprinids are usually transported
Water 2021, 13, x FOR PEER REVIEW towards the dam. 6 of 13

Cyprinidae
100
A
Kr, %
95

90
Percidae
100
B
Kr, %

50

0
15.6
20.6
25.6
1.7
6.7
10.7
20.7
30.7
10.8
16.8
26.8
5.9
15.9
22.9
Time, dates

Figure 4. Relative abundance of the young cyprinids (A) and percids (B) measured in mean concen-
Figure
tration4.(KRelative abundance
r , in %) for the wholeofday–night
the youngperiod
cyprinids (A)date
of each and in
percids (B) measured
both migration inand
(dark) mean con-
residential
centration
(light part of each bar) habitats of the Ivan’kovskoe reservoir in June–September 1992. Much resi-
(K r, in %) for the whole day–night period of each date in both migration (dark) and more
dential
percids(light
werepart of eachin
observed bar)
thehabitats of the
migration Ivan’kovskoe
habitat. reservoir
Differences in June–September
between cyprinids and 1992.
percids are
Much more percids were observed in the migration habitat. Differences between cyprinids and
significant (Wilcoxon matched pairs test, p = 0.001) in both migration habitat and habitat of residence.
percids are significant (Wilcoxon matched pairs test, p = 0.001) in both migration habitat and habi-
Number of fish sampled: cyprinids—1317 ind., percids—1732 ind. Both vertical axis scales have
tat of residence. Number of fish sampled: cyprinids—1317 ind., percids—1732 ind. Both vertical
different
axis scalesscales. Data from
have different Pavlov
scales. et from
Data al. [24].
Pavlov et al. [24].
3. Emigration of Young Cyprinids and Percids from Water Reservoirs
This difference produces a prerequisite for more intensive drift of the young percids
In the lower part of the reservoir, hydraulic conditions for the downstream drift of
than cyprinids towards the dam. The functioning of ecological barriers at different scales,
young fish are drastically changed. Fish may be entrapped by the strong unidirectional
which hamper both the lateral and longitudinal movements of migrants, could be one of
water flow, which carries them downstream through the dam.
the reasons prolonging the period of migration. We argue that the whole water body of
The studied power plants on the regulated rivers were equipped with deep (25–40 m)
the reservoir, working as a macroscale barrier, restricts the intensity and efficiency of the
water intakes, which produce a funnel-like water flow structure that influences different
downstream migration through the lowering of its speed and change of direction. This is
layers of the water column, particularly in the near-bottom layers [24]. In contrast to the
primarily caused by highly variable and unpredictable flow regimes [22,26], which in turn
downstream migration in an unregulated river, where fish are involved in the main channel
is influenced by the variability of the wind-induced currents [27] and the topography of
due to horizontal lateral movements (between the inshore zone and the main channel),
reservoirs [18]. Within morphologically complex reservoirs, numerous bays of various
drift with the water intake flow is associated primarily with the vertical distribution of
sizes may operate as a sort of ecological trap, which accumulates migrating fish [28]. As a
fish [24].
result of the differences between young cyprinids and percids in their characteristics of
migration through of
3.1. Diel Dynamics the
thewater reservoir,
Downstream more percids than cyprinids are usually trans-
Emigration
portedThe
towards the dam.
pronounced diel patterns of the downstream migration, which are relatively
similar in percids and cyprinids migrating along the channel of the unregulated river, are
3.different
Emigration of case
in the Young Cyprinids
of the and
drift with Percids
the water from
intakeWater Reservoirs
induced flow (Figure 5). The main
In the lower part of the reservoir, hydraulic conditions
peaks of the concentration of migrants were recorded at dusk and for theatdownstream drift of
night. For cyprinids,
young fish are drastically changed. Fish may be entrapped by the strong unidirectional
this was the only peak in the diel rhythm, similar to the drift in natural rivers. For percids,
water flow, which
in addition carries nocturnal
to the largest them downstream through theconcentration
peak, a considerable dam. of migrants was also
The studied
recorded duringpower
the dayplants on 5b).
(Figure the regulated rivers
We argue that thewere
main equipped with deep the
drivers governing (25–40
diel
m) waterof
pattern intakes, which
the drift produce of
downstream a funnel-like
dams are thewater
dielflow structure
vertical that influences
movements differ-
and distribution
ent layers
of fish. Weofshould
the water column,that
emphasize particularly in young
most of the the near-bottom
fish in the layers [24].the
area near Indam
contrast to
resided
the downstream migration in an unregulated river, where fish are involved in the main
channel due to horizontal lateral movements (between the inshore zone and the main
channel), drift with the water intake flow is associated primarily with the vertical distri-
bution of fish [24].
 ilar in percids and cyprinids migrating along the channel of the unregulated river, are
different in the case of the drift with the water intake induced flow (Figure 5). The main
peaks of the concentration of migrants were recorded at dusk and at night. For cyprinids,
this was the only peak in the diel rhythm, similar to the drift in natural rivers. For percids,
Water 2021, 13, 1291
in addition to the largest nocturnal peak, a considerable concentration of migrants 7was of 13
also recorded during the day (Figure 5b). We argue that the main drivers governing the
diel pattern of the drift downstream of dams are the diel vertical movements and distri-
bution of fish. We should emphasize that most of the young fish in the area near the dam
in the upper
resided in the layers, but thebut
upper layers, mainthebulk
mainofbulk
percid
of larvae
percid were
larvaelocated significantly
were located deeper
significantly
than that
deeper thanof that
the cyprinids [19]. [19].
of the cyprinids

100 B
A
Concentration of migrants,

75
% max

50

25

0
12 14 16 18 20 22 0 2 4 6 8 10 12 12 14 16 18 20 22 0 2 4 6 8 10 12
Time, hrs Time, hrs
Figure
Figure5.5. Diel
Dielpatterns
patternsof
ofthe
thedownstream
downstreammigration
migrationofoflarval
larvalcyprinids
cyprinids(A)
(A)and
andpercids
percids(B)
(B)in
inthe
thenonregulated
nonregulatedpart partof
of
the Upper Volga (dashed lines) and through the water intake of the Ivan’kovskaya Power Plant (solid lines). Diel
the Upper Volga (dashed lines) and through the water intake of the Ivan’kovskaya Power Plant (solid lines). Diel patternspatterns
in
inthe
theriver
riverand
andreservoir
reservoirare
aresimilar
similarin
incyprinids
cyprinids(Kolmogorov–Smirnov
(Kolmogorov–Smirnovtest:test:pp>>0.05)
0.05)and
anddifferent
differentininpercids
percids(p(p<<0.05).
0.05).
Number of fish sampled: cyprinids—222 ind. (river), 166 (reservoir); percids—263 ind. (river), 532 (reservoir). Data from
Number of fish sampled: cyprinids—222 ind. (river), 166 (reservoir); percids—263 ind. (river), 532 (reservoir). Data from
Pavlov et al. [21,24].
Pavlov et al. [21,24].

When
Whenyoung
youngpercids
percidsand
andcyprinids
cyprinidsmigrate
migrateininthethenatural
naturalriver,
river,they
theyshow
showno nodiffer-
differ-
ence
ence in the diel patterns of the downstream migration, despite the difference in spatial
in the diel patterns of the downstream migration, despite the difference in spatial
distribution
distributionacross
acrossthe theriver
riverchannel
channel(usually,
(usually,migrating
migratingpercids
percidswere
werefound
foundfurther
furtherfrom
from
the
theshore
shorethan
thancyprinids)
cyprinids)[11].
[11].InInthe
theriver,
river,aanarrow
narrowflowflow gradient
gradient between
betweenthe theinshore
inshore
zone
zoneandandmain
mainchannel
channelisisaa permeable
permeable structure
structure forfor both
both categories
categories ofof migrants.
migrants. InInthe
the
water reservoir near the dam, a 3D flow “funnel” produced by the
water reservoir near the dam, a 3D flow “funnel” produced by the deep-water intake deep-water intake cre-
ates a vertical
creates flow
a vertical gradient,
flow which
gradient, which acts as as
acts ananecological
ecologicalbarrier
barrierfor
forboth
bothcyprinids
cyprinidsandand
percids.
percids. Both groups of the larvae undertake diel vertical movements, but theranges
Both groups of the larvae undertake diel vertical movements, but the rangesof of
their
theirmovements
movements were found to
were found tobe bedifferent
different[19,24].
[19,24].Most
Mostcyprinids
cyprinids were
were located
located above
above the
the
flowflow “funnel”
“funnel” during
during the the
day,day,
and andonlyonly at night,
at night, they they
sank sank deeper
deeper and entered
and entered the
the water
water flow, which
flow, which conveyed conveyed them downstream
them downstream throughthrough
the dam. thePercids
dam. Percids stayedthe
stayed within within
zone
the zone of
of water water
intake flowintake
evenflow
duringeventheduring the day;
day; during the during the night,
night, they descendedthey deeper,
descended and
deeper, and their concentration in the outflow
their concentration in the outflow was even higher. was even higher.

3.2.Seasonal
3.2. SeasonalDynamics
DynamicsofofthetheDownstream
DownstreamEmigration
Emigration
Incontrast
In contrast to the
the seasonal
seasonalpatterns
patternsofofthe concentration
the concentrationof migrants in the
of migrants in natural river,
the natural
where the only pronounced peak was observed in June and smaller
river, where the only pronounced peak was observed in June and smaller peak for cypri- peak for cyprinids
in July
nids (Figure
in July 3), 3),
(Figure in in
regulated
regulated rivers,
rivers,broader
broadermain
mainseasonal
seasonal peaks (June–July) were
peaks (June–July) were
recordedininboth
recorded bothfamilies
families of of fish
fish (Figure
(Figure 6). 6). Moreover,
Moreover, a lower
a lower but considerable
but still still considerable
con-
concentration
centration of migrants
of migrants waswas recorded
recorded during
during thethe rest
rest of of theyear,
the year,especially
especiallyforforpercids
percids
(Figure6).
(Figure 6).AAsignificant
significantamount
amountof ofmigrating
migratingYOY YOYgrown
grownpercids
percidswas
wasrecorded
recorded during
during
the autumn–winter period. This may lead to a substantial loss of a valuable
the autumn–winter period. This may lead to a substantial loss of a valuable portion of portion of
percid populations.
percid populations.
3.3. Emigration through the Power Plants and Shipping Locks
In addition to the main deep-water intake of the power plant, other intakes may be
associated with the dam. Amongst the most common constructions are shipping locks,
which periodically create a downstream water flow conveying young fish downstream.
Such water intakes are situated close to the bank and predominantly influence the upper
water layers. We monitored the intensity of the downstream migration of the young fish
through the power plant (throughout the year) and the shipping lock (from May to October)
 Water 2021, 13, 1291 8 of 13

intakes located at the middle part of the Sheksna River (Vologda Region, European Russia).
The intensity of emigration from the reservoir was by an order of magnitude higher for
the percids (Figure 7b) than for cyprinids (Figure 7a). Significantly more percid larvae
drifted through the power plant water intake than through the shipping lock; conversely,
the migration of cyprinid larvae was more intensive through the shipping lock (Figure 7a).
These obtained results showed that the emigration of larval percids from the reservoir
stock was much higher than the emigration of cyprinids. The difference between the two
fish families is especially pronounced when the main discharge of water flows through the
Water 2021, 13, x FOR PEER REVIEW deep-water intake of the power plant. 8 of 13

Figure 6. Seasonal patterns of the downstream migration of cyprinids (dashed line) and percids
Figure
(solid6. line)
Seasonal
frompatterns of the downstream
the Ivan’kovskoe Reservoir.migration of cyprinids
The patterns (dashed
significantly line)
differ and percids
(Wilcoxon matched
(solid line) from the Ivan’kovskoe Reservoir. The patterns significantly differ (Wilcoxon
pairs test, p = 0.018). Number of fish sampled: 4100 ind. See Supplement 2 for more detail; matched
data from
pairs test, р = 0.018). Number of fish sampled: 4100 ind. See Supplement 2 for more detail; data
Pavlov et al. [21,24].
from Pavlov et al. [21,24].

3.3. Emigration through the Power Plants and Shipping Locks

In addition to the main deep-water intake of the power plant, other intakes may be
associated with the dam. Amongst the most common constructions are shipping locks,
which periodically create a downstream water flow conveying young fish downstream.
Such water intakes are situated close to the bank and predominantly influence the upper
water layers. We monitored the intensity of the downstream migration of the young fish
through the power plant (throughout the year) and the shipping lock (from May to Octo-
ber) intakes located at the middle part of the Sheksna River (Vologda Region, European
Russia). The intensity of emigration from the reservoir was by an order of magnitude
higher for the percids (Figure 7b) than for cyprinids (Figure 7a). Significantly more percid
larvae drifted through the power plant water intake than through the shipping lock; con-
versely, the migration of cyprinid larvae was more intensive through the shipping lock
(Figure 7a). These obtained results showed that the emigration of larval percids from the
reservoir stock was much higher than the emigration of cyprinids. The difference between
the two fish families is especially pronounced when the main discharge of water flows
through the deep-water intake of the power plant.
Water 2021, 13, 1291 9 of 13

Water 2021, 13, x FOR PEER REVIEW 9 of 13

40,96 A
10,24 Shipping lock
2,56

Concentration of migrants, ind/1000 m3

Power plant
0,64
0,16
0,04
0,01
655,36 B
163,84
40,96
10,24
2,56
0,64
0,16
0,04
0,01
I II III IV V VI VII VIII IX X XI XII
Months
Figure 7. Seasonal patterns of emigration of cyprinids (A) and percids (B) through the shipping
Figure 7. Seasonal patterns of emigration of cyprinids (A) and percids (B) through the shipping
lock(dashed
lock (dashedlines)
lines)and
andpower
powerplantplant(solid
(solid lines)
lines) passages
passages onon
thethe Sheksninskoe
Sheksninskoe Reservoir.
Reservoir. Mean Mean
concentrationsofofboth
concentrations bothpercids
percidsand and cyprinids
cyprinids areare higher
higher when
when drifting
drifting through
through the the
powerpower
plantplant
(Wilcoxon test, pp << 0.023;
(Wilcoxon matched pairs test, 0.023; see
see Supplement
Supplement33for formore
moredetails).
details).Number
Numberofoffishfishsampled:
sam-
pled: cyprinids—1298
cyprinids—1298 ind.; percids—30,169
ind.; percids—30,169 ind. from
ind. Data Data Pavlov
from Pavlov et al. Note
et al. [24]). [24]).that
Notedifferent
that different
irregular
irregular Y-axes used.
Y-axes used.

4. Ecological
4. EcologicalBarriers
BarriersandandFilters
Filtersasas Modifiers
Modifiers of of the
the Downstream
Downstream Migration
Migration in in Trans-
Transformed
formed Rivers Rivers
In contrast
In contrast toto the
the downstream
downstream migration
migration in in unregulated
unregulated rivers,
rivers, in
in which
which the
the main
main
patterns (diel and seasonal) are relatively similar in cyprinids and percids, the modified
patterns (diel and seasonal) are relatively similar in cyprinids and percids, the modified
structure of regulated rivers drastically enhances differences in the temporal and spatial
structure of regulated rivers drastically enhances differences in the temporal and spatial
patterns of the downstream migrations between these groups of migrants. We showed that
patterns of the downstream migrations between these groups of migrants. We showed
a set of novel ecological barriers influences the spatiotemporal characteristics of migrating
that a set of novel ecological barriers influences the spatiotemporal characteristics of mi-
larvae of both families. These differences are distinct at the local scale and at the scale of
grating larvae of both families. These differences are distinct at the local scale and at the
the whole reservoir. The barriers work as ecological filters exerting a selective impact on
scale of the whole reservoir. The barriers work as ecological filters exerting a selective
the two most abundant and diverse groups of fish inhabiting lowland water reservoirs.
impact on the two most abundant and diverse groups of fish inhabiting lowland water
The resulting effect of behavioral responses of migrating fish to these barriers can be
reservoirs.
estimated via a comparison of the number of fish emigrating from the reservoir. A correct
The resulting effect of behavioral responses of migrating fish to these barriers can be
comparison should take into account the population numbers of different fishes inhabiting
estimated via a comparison
the impoundment. An indexofofthe numberintensity
migration of fish emigrating
(MI), which from the reservoir.
reflects Anumber
the relative correct
comparison should take into account the population numbers of different
of fish of a particular species (group) leaving a reservoir, was previously proposed [24].fishes inhabit-
ing thethe
Using impoundment. An index
data on the intensity of of migration
drift from eightintensity (MI),
reservoirs which
and reflects theofrelative
the abundance young
number of fish
fish in these of a particular
reservoirs, species (group)
we calculated MI indicesleaving a reservoir,
for cyprinids andwas previously
percids (Figurepro-
8).
posed [24]. Using the data on the intensity of drift from eight reservoirs
A tendency to migrate downstream from reservoirs was much more pronounced in percids and the abundance
of young
than fish in these
in cyprinids reservoirs, weUcalculated
(Mann–Whitney MI indices for cyprinids and percids (Fig-
test: p < 0.001).
ure 8). A tendency to migrate downstream
Considering the mechanisms underlying the from reservoirs was much between
striking difference more pronounced
cyprinids
in percids than in cyprinids (Mann–Whitney U test: p < 0.001).
and percids in terms of the intensity of the downstream migration, the role of ecological
barriers influencing their spatial distribution and migration at different scales should be
emphasized. Within a reservoir, the following hydrological barriers can be distinguished:
Water 2021, 13, 1291 10 of 13

• The extended gradient zone between the migration habitat and habitat of residence
which hampers movements of young fish between the open water area (migration
habitat) and inshore zone (habitat of residence);
• A vast water body with a changeable flow structure, which slows and hampers the
drift of young fish downstream towards the dam;
• A transformed hydrological structure with a deep-water downstream flow near the
dam, as the probability of being entrapped by the deep-water intake flow and con-
veyed downstream is higher for fish inhabiting deeper layers further from the in-
Water 2021, 13, x FOR PEER REVIEW shore zone. 10 of 13

Cyprinids Percids
Meanindices
Figure8.8.Mean
Figure indicesofofmigration
migrationintensity
intensityofofcyprinids
cyprinidsand
andpercids
percidsfrom
fromeight
eightwater
waterreservoirs.
reservoirs.
Theseindices
These indicesare
areconsistently
consistentlyhigher
higherininpercid
percidspecies
speciesthan
thaninincyprinid
cyprinidspecies
species(Mann–Whitney
(Mann–WhitneyUU
test,
test,pp<<0.001).
0.001).See
SeeSupplement
Supplement44for formore
moredetail.
detail.Data
Datafrom
fromPavlov
Pavlovetetal.
al.[24].
[24].

At the scale of
Considering thethe whole regulated
mechanisms river, the
underlying themorphological complexity
striking difference between (topography)
cyprinids
of the
and reservoir
percids playsofa the
in terms pivotal role inofthe
intensity thecontrol of the intensity
downstream migration, of the
the role
downstream
of ecological drift
barriers influencing their spatial distribution and migration at different scales shouldthat
of young fish. The whole reservoir works as a large-scale ecological barrier or trap be
hampers the Within
emphasized. downstream migration
a reservoir, [18,28]. The
the following impact of the
hydrological morphological
barriers complexity of
can be distinguished:
the reservoir overrides the impact of the water turnover (discharge) rate [18]. We compared
• The extended gradient zone between the migration habitat and habitat of residence
the total number of fish emigrating from two reservoirs of similar surface area but different
which hampers movements of young fish between the open water area (migration
complexity (the ratio of the shore line length to the length of the longitudinal axis), both
habitat) and inshore zone (habitat of residence);
located on the Volga River—i.e., Volgogradskoe (turnover rate, 8.0; index of morphological
• A vast water body with a changeable flow structure, which slows and hampers the
complexity, 0.03) and Ivan’kovskoe (12.9 and 1.83, respectively), measured throughout the
drift of young fish downstream towards the dam;
year. We found that the number of young fish emigrating from the first reservoir was 3000
• A transformed hydrological structure with a deep-water downstream flow near the
times more than from the second. The main bulk of the emigrants from the both reservoirs
weredam,
young as percids
the probability
[18]. of being entrapped by the deep-water intake flow and con-
veyed downstream is higher
Certain ecological and behavioral for fish inhabiting
traits of percidsdeeper
make layers
themfurther from the
more subject to in-
the
shore zone.
impact of the downstream transportation under conditions of regulated rivers. Common
At the
riverine scale of
percids the whole
(perch, regulated
zander, river, the
ruff) occupy morphological
similar habitats tocomplexity (topography)
cyprinids (roach, bream,
of the reservoir
bleak) in natural plays a pivotal
rivers. They arerole in the control
generally similarofinthe intensity
their timingofand thespatial
downstream drift
distribution,
of
at young
least infish.
earlyThe whole reservoir
ontogeny [21,29,30].worksHowever, as a large-scale
there are someecological barrier
differences in or trap
their that
spatial
hampers the downstream
distribution, which undermigration [18,28]. The
riverine conditions doimpact of the morphological
not markedly influence patterns complexity
of their
of the reservoirmigration
downstream overrides[11].
the impact
Spawning of thegrounds
water turnover
of percids (discharge)
are usuallyratelocated
[18]. Wefurthercom-
offshore
pared thethan
totalthose
numberof cyprinids [21,24,31,32].
of fish emigrating from Intwo
water reservoirs,
reservoirs their early
of similar larvae
surface cannot
area but
maintaincomplexity
different a steady position among
(the ratio of theinshore
shorevegetation,
line lengthand to they are much
the length more
of the vulnerable
longitudinal
to wind-induced
axis), both locatedcurrents than cyprinid
on the Volga River—i.e., larvae. Such currents
Volgogradskoe disperse
(turnover percid
rate, 8.0; larvae
index of in
all directions [33–36].
morphological complexity, Young percids
0.03) are able to inhabit
and Ivan’kovskoe (12.9 anda wide range of depths
1.83, respectively), [37,38];
measured
cyprinids, in
throughout thecontrast
year. We to percids,
found that inhabit predominantly
the number of younginshore zone andfrom
fish emigrating upper thelayers.
first
Both cyprinids
reservoir was 3000 and percids
times moreundertake
than from horizontal
the second.and The vertical
main bulk dielofmovements
the emigrants [39–41].
from
the both reservoirs were young percids [18].
Certain ecological and behavioral traits of percids make them more subject to the
impact of the downstream transportation under conditions of regulated rivers. Common
riverine percids (perch, zander, ruff) occupy similar habitats to cyprinids (roach, bream,
Water 2021, 13, 1291 11 of 13

Spatial distribution and diel movements make young percids more vulnerable to water
currents, which in water reservoirs often direct them to the water intakes of power plants
and emigration from the reservoir.

5. Conclusions
The obtained results suggest that patterns of downstream migration of cyprinid
and percid fishes are strongly influenced by a novel physical template created by river
regulation. Novel ecological barriers associated with different parts of water reservoirs
differently affect patterns of migration of cyprinids and percids. These differences result in
significantly more intensive emigration of young percids than of cyprinids from reservoirs
with deep-water intakes. The loss of percids from reservoirs differs from cyprinids not
only in terms of the higher total number but also in age (size) structure of the migrants.
The highest number of percid larvae emigrating from reservoirs is found at mid-summer
(June–July), but, unlike cyprinids, young percids continue a rather intensive downstream
migration from reservoirs during autumn and winter. This leads to the loss of grown YOY
percids, which could constitute a significant and important portion of the overwintering
stock. Such disproportional removal may be one of the important factors controlling the
ratio of cyprinids/percids in the fish stock of reservoirs with deep-water intakes, in turn
influencing the structure of the fish fauna of the reservoirs. Other large-scale factors are
related to the morphology of reservoir. Reservoirs that are morphologically more complex
can prevent the emigration of young fish more efficiently than channel-like reservoirs [18].
Dispersal ability is not simply an attribute of a species (group of species), but it varies
strongly with landscape structure; thus, landscape change due to human activities may be
more influential for one group of animal than for another [42].
Various approaches exist for managing reservoir fish fauna and stock [23,43–47].
Usually, reservoir managers want to create a fish stock most suitable for the purposes
for which the reservoir was built. Ensuring a high production of cyprinids by designing
eulittoral ecotones is one strategy for reservoirs where these fast-growing fish are to be
exploited for human or animal food [20]. The intensive removal of young percids from
reservoirs with deep-water intakes should also facilitate conditions in favor of cyprinids by
reducing competition for resources. To promote populations of percids in a reservoir, we
should not only adjust diel and seasonal regimes of hydropower water intakes but also
optimize water intake location to minimize the withdrawal of young percids.

Supplementary Materials: The following are available online at https://www.mdpi.com/article/10

.3390/w13091291/s1, Supplement 1—Table S1: Mean relative concentrations of migrants during the
day and night; Table S2: Seasonal patterns of the downstream migration of cyprinids and percids in
the Upper Volga; Table S3: Statistics for the seasonal dynamics of the concentration of migrants in the
Upper Volga River; Table S4: Relative concentrations (%) of the young Cyprinidae and Percidae in
the migration habitats and habitats of residence in the Ivan’kovskoe Reservoir in June–September
1992; Table S5: Significance of differences between relative concentrations of Cyprinidae and Percidae
in the migration habitats and habitats of residence. Supplement 2—Table S6: Diel changes of the
relative concentration (% max) of the early larvae of Cyprinidae and Percidae migrating in the Upper
Volga and from Ivan’kovskoe Reservoir; Table S7: Diel changes in the concentration of Cyprinidae
and Percidae migrants in the Upper Volga and Ivan’kovskoe Reservir; Table S8: Number of caught
fish (ind., according to species) in the Upper Volga and Ivan’kovskoe reservoir; Table S9: Seasonal
changes of the relative concentrations (% max) of Cyprinidae and Percidae emigrating from the
Ivan’kovskoe Reservoir; Table S10: Comparison of seasonal changes of the Cyprinidae and Pecidae
emigration from the Ivan’kovskoe Reservoir; Table S11: Sample sizes (ind.) for the main species of
Cyprinidae and Percidae. Supplement 3—Table S12: Concentrations (ind./1000 m3 ) of Cyprinidae
and Percidae migrants through the power plant water intake and shipping lock of the Sheksninskoe
Reservoir; Table S13: Difference between concentrations of cyprinids and percids migrating through
the power plant and shipping lock; Table S14: Number of sampled fish (by species of cyprinids and
percids) emigrating through the power plant and shipping lock. Supplement 4—Table S15: Migration
Water 2021, 13, 1291 12 of 13

index for common and frequently found fishes in reservoirs; Table S16: Comparison of the indices of
migration in cyprinids and percids.
Author Contributions: Conceptualization, D.S.P., V.N.M. and V.V.K.; data analysis, V.V.K. and
V.N.M.; writing, V.N.M. and D.S.P.; supervision and funding acquisition, D.S.P. All authors have read
and agreed to the published version of the manuscript.
Funding: This study was funded by “Mechanisms of migratory behavior of fish and cyclostomes in
river systems. The role of ecological factors and physiological state”. Grant from the Russian Science
Foundation, number 19-014-00015.
Institutional Review Board Statement: Ethical review and approval were waived for this study, due
to the fact that this review paper is based on the analysis of the published data (references [21,24])
and the above mentioned Supplementary materials.
Informed Consent Statement: Not applicable.
Data Availability Statement: The data presented in this study are available in the above mentioned
Supplementary materials and the monographs listed therein.
Conflicts of Interest: The authors declare no conflict of interest.

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