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IMPACT OF HYDRO-ECOLOGY ON FISH MIGRATION IN SELECTED
EAST COAST ESTUARIES
UTPAL BHAUMIK, N. P. SHRIVASTAVA, B. B. SATPATHY, M. K. MUKHOPADHYAY
AND A. P. SHARMA
Central Inland Fisheries Research Institute, Barrackpore, Kolkata - 700 120
Received : 24.11.2010; Accepted : 04.02.2011
Attempts were made to identify the commercially important migrant fish species and study their migration pattern in
two selected estuaries i.e. Hooghly and Mahanadi on east-coast facing the Bay of Bengal Sea. Nearly 20 species in such
category could be enlisted from Mahanadi estuary, which were also encountered in the Hooghly estuarine system. Most
of the migrant species were amphidromous and their estuarine migration was influenced by the hydro-ecological
characteristics. Salinity, depth and food niches were amongst the factors governing migration of the fishes. Migration
was limited to 35 km in Mahanadi while up to 100 km in Hooghly estuarine system. Hilsa was only anadromous species
migrating to freshwater river course and got struck up by the barrages. In Mahanadi river-estuarine system migration of
Hilsa was seriously affected. Recruitment and fisheries of Hilsa in Hooghly-Bhagirathi system did not show significant
impact of the Farakka barrage.
Key words : Hydro-ecology, migration, estuary, amphidromous
J. Inland Fish. Soc. India, 43(1) : 1-9, 2011
Introduction
Migration is natural instinct and facilitating act in
fulfillment of bio-physiological urges in animals.
Fishes are no exception and constitute one amongst
the distant migrating groups in aquatic sphere. For
some fish species migration is intra environmental,
limited to shorter range of distance while for
others trans-environmental migration extends from
a few hundreds to thousands of kilometers.
Estuaries being intermediate environment between
sea and freshwater river and eco-biologically rich
habitat, play as favourable destination for both
marine and freshwater fish species. The fish move
to estuarine environment in search of breeding and
nourishment habitats or for feeding and fattening
purposes. The variability of estuarine environment
and eco-biological richness depend on several
intrinsic and extrinsic factors. Post independence
habitation, agriculture and industrial development
activities and consequential increase in dependence
on rivers and also estuaries gradually encroached
upon these resources and interfered the process
of natural renewal of eco-biological potentialities.
As a result the natural biotope of the estuarine
system got affected in qualitative and quantitative
richness including those of fishes.
According to the available records the fish
faunal structure in most of the estuarine
systems has shown shifting trend indicating
disappearance of a few species, decline in
abundance of many and appearance of some
species not recorded earlier. In view of this
striking phenomenon in fish species spectrum it
was felt necessary to develop knowledge on
migration of fishes and investigate the habitat
requirements, purpose and constraints in
migration of fishes in or through estuarine systems.
Materials and methods
Two estuarine systems on east coast facing the
Bay of Bengal Sea, which are hydro-morpho-
1
metrically different were selected for investigations.
Hydro-ecological conditions were examined
following the standard methods. Spectrum and
species-wise distribution of available fishes were
continuously monitored during 2007-09. The data
on spatio temporal abundance and some
biological aspects were analysed for critical
interpretation and drawing inferences on
migration of fishes in the investigated estuarine
systems.
Experimental netting with gill nets of specific
dimensions (200-250 m x 6.0 m) with uniform
mesh size (80-100 mm) was carried out at
Diamond Harbour and Nababganj during
monsoon in Hooghly estuary. Five nets were
deployed daily for three days at each site and the
fishes caught in three hauls for every net were
recorded for analyses of depth preference for hilsa
in the system.
Results
Mahanadi estuarine system
As per record, Mahanadi estuary in Orissa coast
was dimensionally vast and tidally more than 150
km in length. Erection of Naraj barrage above 100
km from sea face in sixtees brought out significant
alterations in hydromorphometry of the estuary.
During dry season of summer fair-weather roads
come up across the estuary revealing the drastic
fall in discharge down below the barrage and
threatening disintegration of the estuarine
continuity at points of high rate of siltation. Apart
from the biodiversity losses the qualitative and
quantitative decline in fisheries has been a matter
of great concern.
Hydro-ecology
Salinity distribution in Mahanadi estuary indicated
the influence of monsoon discharge from the
upstream Naraj barrage at Cuttack. During dry
season of summer the estuarine stretch from the
sea face up to 2 km upstream near Paradip was
maximum saline followed by medium saline zone
up to 16 km at Bhutmundi and thereafter the tidal
water was fresh with traces of salinity. With
monsoon discharge the freshwater tidal zone
extended downstream leaving only 2 km of the
stretch facing sea as medium saline zone. The
dissolved oxygen concentration was within
favourable range (5.2-8.2 mgl-1) except in the
deeper regions above the sea face reflecting
anaerobic condition of the bottom during dry
season of summer. The ambient pH (6.2-8.0)
indicated lower values of the same in industrially
affected areas near Paradip. Annual mean of
plankton density was recorded as 150 ul-1, 103
ul-1 and 162 ul-1 in upper freshwater, freshwater
tidal and brackish water tidal zones respectively.
The percentile contribution of phytoplankton and
zooplankton varied with the salinity and was
77.5:22.5, 54.4:45.6 and 57.3:42.7 in upper
freshwater, freshwater tidal and brackish
water tidal zones of the estuarine system
respectively.
Migrant fish species spectrum
Fish species spectrum in Mahanadi included a long
list of fin fishes and for the present investigation
those migrating in inland part of the estuary were
considered. Samples of fish specimen collected
from the landing centres and also the fishing boats
were identified and preserved for further studies
all along the estuarine length. A total of 20 species
of fishes could be enlisted within 35 km of the
estuarine span with Hilsa (Tenualosa ilisha)
moving further to down below the Naraj barrage
about 100 km upstream from the sea face. Most
of the fish species migrated to the estuarine
environment during post monsoon months when
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BHAUMIK et al.
estuarine ecosystem was stabilized. The water was
quite transparent, temperature ranged between
210C and 300C, oxygen was in semi saturated
condition, salinity fluctuated between traces and
18 ppt, abundance of plankton and benthos was
in peaks and the intertidal depressions were
covered with thick growth of algal mats. Hilsa
specimens were available during monsoon mainly
and the fishes encountered down below the Naraj
barrage were mostly females at different stages of
maturity and the remaining males were in milting
condition.
Biology of fishes
Biological examination of the specimens procured
from the inland estuarine segment revealed 60%
of the enlisted 20 species as carnivore, feeding on
the prey species like small fishes, juveniles of fishes,
prawns and shrimps while 25% was eliophagic
depending on the vegetative matters like different
species of algae and occasionally twigs of soft
hydrophytes. The rest 5% depended on benthic
molluscs mainly. Reproductively the majority of
specimens sampled during the main season of post
monsoon were not in maturing condition. In Hilsa
the situation was different from the others since
most of the examined fish specimens were in
various maturity stages and bulk of the fish catch
was in monsoon months.
Hooghly-Matlah estuarine system
Largest of its kind in India, the estuary estimates
to 300 km in tidal length and holds well
demarcated salinity zones viz. marine, brackish
water and freshwater at the top. Prior to seventees
the salinity incursion used to extend up to about
200 km from the sea mouth. Thereafter,
freshwater induction through the Farakka barrage
installed at the bifurcation point of Ganga to
Bhagirathi and Padma above 500 km from sea
face made remarkable impact on eco-biological
characteristics of the estuarine system. The
freshwater tidal zone extended downstream
covering nearly 70% of the tidal length. The
brackish water zone shrunken by 50% and the
marine zone was pushed seaward.
Hydro-ecology
Salinographically, the Hooghly estuary was
delineated in two demarcated zones within the land
i.e. between Nabadwip and Kakdwip sampling
points. The stretch between Kakdwip and
Diamond Harbour remained brackish water with
seasonal variability in salinity while above Diamond
Harbour up to Nabadwip the environment was
fresh round the year.
Dissolved oxygen (7.63-8.80 mgl-1) and total
alkalinity (100-130.83 mgl-1) remained within
productive range with the support of favourable
pH range (7.6-8.0). Population density of
Table 1. List of migratory fishes recorded in the stretch
between sea mouth and Naraj barrage in river Mahanadi
Sl. No. Species
1Tenualosa ilisha
2Hilsa kelee
3Lates calcarifer
4Pama pama
5Pangasius pangasius
6Sillago sihama
7Arius jella
8Terapon jarbua
9Otolithoides biauritus
10 Eleutheronema tetradactylum
11 Rhinomugil corsula
12 Mugil cephalus
13 Mugil macrolepis
14 Liza parsia
15 Liza tade
16 Gerreomorpha satifer
17 Stolephorus commersonii
18 Stolephorus indicus
19 Leiognathus equulus
20 Stigmatogobius sadanandicus
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FISH MIGRATION IN ESTUARIES
plankton varied widely within the estuary and
counted in the order of 292 ul-1 (163-427 ul-1),
220 ul-1 (183-258 ul-1) and 299 ul-1 (136-465
ul-1) in upper, middle and lower zones respectively.
Phytoplankton was predominant over
the zooplankton organisms. The average
macrobenthic density in the entire estuarine stretch
clearly indicated maximum abundance of the
organisms during winter (7442 nos/m2) followed
by post monsoon (4428 nos/m2) and monsoon
(3223 nos/m2) seasons. Molluscs contributed
highest percentage in the benthic population and
within the group the dominance of Gastropods was
observed throughout the stretch.
Migrant fish species spectrum
All the 20 species recorded from the Mahanadi
estuarine system were present in Hooghly-Matlah
estuarine complex. In respect of distance covered
during migration run by different species there was
variation and by and large most of the species were
recorded from the 100 km upstream up to
Diamond Harbour from the sea face. The
seasonality of migration of these species tallied with
those of Mahanadi estuary. The significant
difference was in migration range of Hilsa, which
moved upstream beyond the estuarine span in
freshwater riverine condition up to the
downstream of feeder canal of the Farakka
barrage lock gates.
Reproductive and migration behavior of Hilsa
Field observations were made on the
migration pattern of hilsa, the important target
species in Hooghly and Mahanadi estuaries.
Reproduction and related behavioural activities of
the species were investigated round the year in
Hooghly estuary. To a very large extent, hilsa
completed the breeding run within the estuary,
with favourable influence of high freshwater flow
and reduction of salinity. Only a smaller group
sought for the riverine environment. It appears that
Table 2. Food and feeding habits and range of migration of fish in estuarine environment
Species Food and feeding habits Habitat recovered from
Tenualosa ilisha Planktivorous Marine to freshwater river
Hilsa keele Planktivorous Marine and coastal zone
Lates calcarifer Carnivorous Coastal to freshwater tidal zone
Pama pama Carnivorous Coastal to freshwater tidal zone
Sillago sihama Carnivorous Coastal to brackish water tidal zone
Arius jella Carnivorous Coastal to brackish water tidal zone
Therapon jarbua Carnivorous Coastal to brackish water tidal zone
Otolithoides biauritus Carnivorous Coastal to brackish water tidal zone
Eleutheronema tetradactylum Carnivorous Coastal to brackish water tidal zone
Rhinomugil corsula Eliophagus Estuarine to freshwater river
Mugil cephalus Eliophagus Coastal to brackishwater tidal zone
Mugil macrolepis Eliophagus Coastal to brackishwater tidal zone
Liza parsia Eliophagus Coastal to brackishwater tidal zone
Liza tade Eliophagus Coastal to brackishwater tidal zone
Gerreomorpha satifer Carnivorous Coastal to brackishwater tidal zone
Stolephorus commersonii Carnivorous Coastal to brackishwater tidal zone
Stolephorus indicus Carnivorous Coastal to brackishwater tidal zone
Leiognathus equulus Carnivorous Coastal to brackishwater tidal zone
Stigmatogobius sadanandicus Carnivorous Coastal to estuarine tidal zone
Pangasius pangasius Molluscan feeder Brackish water tidal zone to freshwater river
4
BHAUMIK et al.
the breeding habitat is propounded all along the
longitudinal axis from Diamond Harbour to the
freshwater head of the estuary. The group of fish
in a lesser maturity stage evidently moves upstream
into the riverine stretches.
Major upstream migration of hilsa was observed
from August to November, beginning with the
commencement of monsoon and consequent
flooding. A milder second phase of migration of
mature hilsa, relatively smaller in size, was also
observed during February-March. The peak
migration of adult fish was found occurring shortly
before the spawning. Larger sized hilsa occurred
during July-August and their number fell sharply
from October to February. Female hilsa were
larger in size and fewer in number in the catch than
the males. The female: male ratio during monsoon
was recorded to be 1:2 or 3 in August. Lunar cycle
had a definite influence on hilsa migration, as
Table 3. Experimental findings on depth-wise catch of hilsa in Hooghly estuary
Experimental site and Gill net (200-250 m) Depth-wise catch
experiment number Mesh size (mm) Width (m) Depth (m) Total of 3 hauls (nos)
Diamond Harbour
1 80-100 6.0 3.6 3
2 4.2 5
3 3.5 2
4 3.8 4
5 3.2 3
1 80-100 6.0 4.5 8
2 3.6 5
3 4.3 11
4 5.2 7
5 4.2 8
1 80-100 6.0 5.0 5
2 4.8 5
3 4.2 8
4 3.0 2
5 3.5 4
Nababganj
1 80-100 6.0 4.4 7
2 3.6 3
3 4.0 3
4 5.0 5
5 3.6 3
1 80-100 6.0 5.6 3
2 4.3 5
3 4.4 5
4 5.4 2
5 4.3 4
1 80-100 6.0 4.7 3
2 5.3 2
3 4.8 5
4 3.8 3
5 5.0 2
5
FISH MIGRATION IN ESTUARIES
migration activity was observed to be vigorous four
to five days before and after the full moon and
new moon.
The experimental observations at Diamond
Harbour and Nababganj deploying gill nets of
80-100 mm mesh and 6.0 m width (Table 3) and
statistical analysis of the result indicated variation
in depth preference by the migrating shoals of the
species. At the downstream near Diamond
Harbour the depth between 3.0 and 3.6 m was
most preferred (r2 = 0.984; P = 0.016) followed
by 4.8 and 5.2 m (r2 = 0.866; P = 0.333). The
lower depth of 3.0 to 3.6 m was preferred by
comparatively smaller size groups of the fish, while
the larger ones moved through the deeper water
between 4.8 and 5.2 m from the surface. At about
60 km upstream near Nababganj the senario was
different where bulk of the fishes was caught in
the depth range of 4.0 to 4.4 m (r2 = 0.961; P =
0.179). The movement of the fishes through 4.0
to 4.4 m at Nababganj might be due to high
intensity of navigation activities and resultant
turmoil at the surface water level and also
comparatively low depth of water column in the
region.
However, analysis of the pooled data from Dimond
Harbour and Nababgabj revealed that the depth
range between 4.0 and 4.5 m can be considered
as ideal (r2 = 0.780; P = 0.06) for migration of the
adult Hilsa in river - estuarine condition.
The other favourable conditions observed for hilsa
migration were:
•> 20 m/minute current velocity (during high
turbulence and high water current, the fish
moved closer to river banks)
•Larger size of the estuary, especially deeper
and wider estuarine mouth
•Reasonably adequate freshwater discharge
from the upstream stretches
•High salinity, high turbidity, high siltation at the
estuarine mouth have adverse impact on the
hilsa migration
During migration about 90% of the migratory stock
was observed to be in a mature condition
indicating breeding as the main reason for
migration. The studies also brought on record the
killing of hilsa brooders in the sea itself before their
migration to the estuary for breeding purpose. The
present investigation indicated that the breeding
migration of hilsa takes place in July and a good
number of hilsa broods are caught in sea.
Discussion
Fish movement in open water systems is not
Fig. 1: Numerical abundance of hilsa at different depths
of Hooghly estuary
Correlation analysis
Simple statistics
Variable N Mean SD Sum Min. Max.
Catch of Hilsa 6 6.000 1.788 36.000 3.000 8.000
Depth(4.0-4.5 m) 6 4.283 0.172 25.700 4.000 4.500
Pearson correlation coefficient, N=6
Prob > r under HO : Rho = 0
Depth (4.0 to 4.5 m)
Catch of Hilsa 0.7789
0.0679
6
BHAUMIK et al.
unusual but nearly 2.7% of the world’s
ichthyofauna is recognized as migratory and out of
all these migrants 7.2% are listed as
“threatened” with extinction (Reide 2002 and
2004). Diadromous i.e. species migrating between
freshwater and marine environments form the
major group of migrants. Among the diadromous
those undertaking migration from marine
environment to rivers exclusively for reproduction
are termed as anadromous and those performing
non-reproductive migration are categorized as
amphidromous. Accordingly, most of the species
recorded during present investigation were
amphidromous and hilsa (Tenualosa ilisha)
belonged to the anadromous group.
Simple correlation between the observed
hydro-biological conditions and spatio-temporal
abundance of the migrants in two different
estuarine systems revealed influence of estuarine
eco-environment on species-wise migration
pattern for amphidromous forms.
The salinity gradient acted as prime and common
factor limiting upstream migration of the fish in both
the investigated systems from the sea. Slavik and
Bartos (1997 & 2001) observed sole influence of
water quality in distribution of fishes in Main River
and back water site of Vltava in Prague.
The other attributing factors like availability of food
niches and food abundance, which showed direct
relation with the abundance of individual species,
had also been reported as limiting factors for
movement of adult and juveniles of salmonids
(Slaney & Northcote, 1974; Dill et al., 1981;
Egglishaw & Shackley, 1985; Elliott, 1986).
Discharge related events have wide recognition as
important stimulants for initiation of migration in
large number of fishes. Elevation in river discharge
and subsequent inundation of the low-lying areas
results dissolution of nutrients from the bottom
soil, trigger an outburst of primary productivity and
fuelling aquatic food chain. Junk et al. (1989) and
Bayley (1995) described such productivity
outburst in cyclical order following retreat and
flooding as “flood pulse”. The present
investigation revealed occurrence of flood pulses
during monsoon following rainfall pattern in the
catchment. However, the flood pulse effect like
environmental freshening, fast current coupled with
heavy turbulence, high turbidity brought out
significant change in eco-environment condition
including disintegration of the feeding habitats for
the migrant species, which ultimately interfere the
migration intensity of the fishes during the season.
Further, the environmental deterioration due to the
industrial discharge from Paradip region might have
influenced the fish migration in Mahanadi estuary.
Reaction of eco salmon, pony fish etc. in
contaminated waters have also been reported by
Rehnberg and Schreck, 1986; Sasaki et al., 2003;
Schellart and Wubbels, 1998; Bardach et al.,
1965.
Recorded variability in migration intensity and
production of the species with lunar periodicity in
Hooghly estuarine system was at par with the
observations made by Walker (1949) for
California grunion Leuresthes tenuis at right Duni
where the spawning migration of the species
reached to highest level during full and new moon
days and Grau (1982) for salmon. Impact of
Aswan Dam adversely affecting groundwater,
erosion, agriculture and largely fisheries in
downstream of river Nile has been well
documented by Halim (1991).
Freshwater environment with favourable
hydrography and adequate supply of plankton food
items in Hooghly-Bhagirathi river course provided
ideal breeding and rearing habitats for migrant
population of hilsa as revealed from increasing
7
FISH MIGRATION IN ESTUARIES
production trend and juvenile recruitment of the
species in the system. On the other hand absence
of favourable habitats in the upstream of Mahanadi
estuary affected the breeding migration and so also
production and recruitment potential of the
species in system.
The observed depth preference for the migratory
hilsa like bigger fish moving in deeper water
has also been reported by Bhaumik (2010).
According to the author the spent fishes mostly
bigger in size move downward during low tide
period.
Remarks and conclusion
In view of the present status of fish migration and
the eco-environmental constraints affecting
natural and free movement of the migratory fish
species to the estuarine habitats, the following
issues emerged for consideration to facilitate the
fish migration activities to the utmost possibilities.
•Estimation and maintenance of environmental
flow effective in facilitating salinity ingress to a
desired extent and also retaining the ecological
variability including those of the mangrove
ecosystems could protect the estuarine system
for the migration of fish stocks, biodiversity
conservation and overall production
potential of the estuaries. The barrages and
dams installed for regulation of river flow in
majority of the river-estuarine systems in the
country stand mostly at safer distance and above
100 km away from the sea face. As such the
fish migration particularly of amphidromous
species in majority of the estuaries might have
not been affected even in the absence of fish
locks or any other devices had there been long
riverine span and no dearth of water flow and
habitat required for the migratory fishes down
below.
•From the present experiences and records
available on role of dams and barrages in hilsa
migration and fisheries in Indian river-estuary
systems, it can be concluded that the fish locks/
gates or such devices might not affect
breeding migration, population recruitment and
production of hilsa if the river course down
below any physical obstruction retain
adequate space of freshwater habitat and
environmentally rich ecosystem for alternative
use required by the species.
Acknowledgements
The authors are deeply indebted to the
Director, CIFRI for scientific support and
guidance during the period of present
investigation. They are also thankful to Dr. B. C.
Jha, Head, Reservoir and Wetland Fisheries
Division, CIFRI for help and cooperation. The
technical support extended by Messrs. C. N.
Mukherjee, T. Chatterjee, D. Sanfui, B. N. Das,
A. R. Choudhury, D. Saha, S. Mondal, C. P.
Singh, K. P. Singh, A. K. Jana, A. K. Barui;
Mrs. A. Sengupta and Mrs. K. Jacquline, is
gratefully acknowledged. The support rendered
by other scientists and staff is also duly
acknowledged. The authors also thank Mr.
Anirban Rej, SRF for his help in statistical
analysis
References
Bayley, P. B. 1995. Under stocking large river-flood plain
ecosystem. Bioscience, 45: 153-158.
Bardach, J. E., Fujiya, M. and Holl, A. 1965.
Detergent: Effect on chemical senses of the fish.
Ictalurus natalish (le Sueur). Science, 148: 1605-1607.
Bhaumik, Utpal. 2010. Status of the fishery of Indian
shad (Tenualosa ilisha) with special reference to
Hooghly river system. Souvenir, 21st All India Congress
of Zoology and National Seminar, published by CIFRI,
Barrackpore, 66-81pp.
8
BHAUMIK et al.
Dill, L. M., Ydenberg, R. C. and Fraser, A. H. C. 1981.
Food abundance and territory size in juvenile coho
salmon (Onchorhynchus kisutch). Canadian Journal
of Zoology, 59: 1801-1809.
Egglishaw, H. J. and Shackley, P. E. 1985. Factors
governing the production of juvenile Atlantic salmon in
Scottish streams. Journal of Fish Biology, 27: 27-33.
Elliott, J. M. 1986. Spatial distribution and behavioural
movements of migratory trout Salmo frutta in a lake
District stream. Journal of Animal Ecology, 55:907-922.
Grau, E. G. 1982. ‘Is the lunar cycle a factor timing the
onset of salmon migration?’. Proc. Salmon Trout
Migratory Behaviour Sym., pp. 184-9.
Halim, Y. 1991. The impact of human alterations on the
hydrological cycle on ocean margins. pp 301-327. In :
Mantoura R. F. C., Martin, J - M, Wollast, R. (eds.) Ocean
margin processes in global change. Wiley, New York,
Junk, W. J., Bayley, P. B. and Sparks, R. E. 1989. The
flood pulses concept in river flood plain ecosystems.
Canadian special publication of fisheries and aquatic
science, 106: 110-127.
Rehnberg, B. C. and Schreck, C. B. 1986. Acute metal
toxicity of olfaction in coho salmon:Behaviour,receptors
and odor-metal complexation. Bull.Environ.Contam.
Toxicol., 36: 579-586.
Reide, K. 2002. Global register of migratory species.
German Federal Agency for Nature Conservation, Project
808 05 081.
9
FISH MIGRATION IN ESTUARIES
Reide, K. 2004. Global register of migratory species-from
global to regional scales. German Federal Agency for
Nature Conservation, Project 808 05 081. Final report.
Sasaki, A., Ikejima, K., Azuma, S., Kashimura, N. and
Wada, M. 2003. Field evidence for bioluminescent
signaling in the pony fish Leiognathus elongates.
Environ.Biol.Fish., 66: 307-311.
Schellart, N. A. M. and Wubbels, R. L. J. 1998. The
auditory and mechasensory lateral line system. In: The
physiology of fishes (Evans,D.H.Ed), 2nd ed.,
pp. 283-312. CRC Press, Boca Raton.
Slaney, P. A. and Northcote,T. G. 1974. Effects of prey
abundance on density and territorial behavior of young
rainbow trout (Salmo gairdneri) in laboratory stream
channels. J. Fish. Res. Board. Can., 31:1201-9.
Slavik, O. and Bartos, L. 1997. Effect of water
temperature and pollution on young-of-the-year fish in
regulated stretch of River Vltava, Czech Republic.
Folia Zoologica, 46:367-374.
Slavik, O. and Bartos, L. 2001. Spatial distribution and
temporal variance of fish communities in the channelized
and regulated Vltava river (Central Europe).
Environ. Biol. of fish., 61:47-55
Walker, B. W. 1949. ‘Periodicity of spawning in the
grunion, Leuresthes tenuis’. Ph.D thesis, University of
California, Los Angeles, California.