Bi0I.J. Linn. SOC.,2, pp. 77-117. With 4pIafer and 13figures
June 1970
Preliminary hydrobiological survey of some Southeast Asian
inland waters*
TOSHIHIKO MIZUNO
Biological Laboratory, Osaka Kyoiku University
AND
SYUITI MORI
Otsu Hydrobiological Station, Kyoto University
Accepted f o r publicationJanuary 1970
-
The preliminary research in some tropical inland waters of Asia is described and suggestions
made concerning Phase I1 of this project undertaken by the International BiologicalProgramme/
Section P F (Freshwater Productivity).
Environmental factors were measured in some of the main lakes and rivers and samples
taken of the plant and animal life. The stomach contents of over 80 species of fish were examined
and the remains of plants and animals present compared with the numbers present in the
environment as obtained by the usual hydrobiological netting techniques.
Some of the important problems associated with the biological productivity of these waters
is discussed in the light of the results obtained from this preliminary survey.
CONTENTS
Introduction
.
Environmental conditions
Thailand : lakes and rivers .
Cambodia: lake .
Malaysia: lakes, rivers and tin-mining pools
Aquatic communities
Aquatic higher plants .
Plankton .
Higher crustacea
.
Aquatic insects
Aquatic molluscs.
.
Fishes and their foods .
Relationship between environment and organisms
Summary
.
Acknowledgements .
References
.
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.
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77
78
79
85
85
90
91
93
98
99
103
104
113
114
115
115
INTRODUCTION
T h e aim of the International Biological Programme (IBP) is to measure the productivity
of organisms on the globe and to contribute to human welfare. After discussing the
methods to be adopted for measuring the productivity of freshwater communities in
the preliminary first team (Phase I), the Japanese Subcommittee on Freshwater
* This study was carried out as a part of Japanese IBP projects. The special project research supported
by the Japanese Ministry of Education: Studies on the Dynamic States of the Biosphere. J P F
publication No. 57.
6
77
78
T. MIZUNO AND S. MORI
Production (P.F.) selected several lakes and rivers in Japan, each different in character,
organized research teams, and has since been performing production studies as its
Phase I1 work. It is making also an attempt at the same kind of research work in tropical
Asia, in collaboration with the countries concerned.
Although tropical Asian inland waters are reputed to be of high biological productivity,
very little is known about the basis of production processes. Encouraged by international
IBP advice and by favourable response from countries in Southeast Asia, a team of
Japanese scientists, headed by Dr H. Tamiya, Chairman of the Japanese IBP
Committee, made a survey trip in 1968. The authors of this paper worked in Thailand,
Cambodia and Malaysia for three months. In spite of the short duration of this
reconnaissance work, some observations and data have been collected which may form
the basis of more intensive studies planned for the future.
ENVIRONMENTAL CONDITIONS
There are some reports on the environmental conditions of Southeast Asian inland
waters. J. Kobayashi (1958)states that the Thai river waters have considerable amounts
FIGURE
1. Research areas in Thailand and Cambodia.
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
79
of Ca-ion; D. S. Johnson (1967) gives excellent data of water analysis in South Malaysia
and Singapore. Besides, T. Piyakarnchana (1967), J. C. D. Watts (1965, 1968), Y.
Shiraishi (1967) and others have recently presented some data obtained from this area.
Thailand
Lakes
Khan Payao Lake. This is on the west side of Chiengie town in the northern part of
Thailand. (Fig. 2A). This was a small natural lake before a dam was constructed, but
now is a fair sized man-made lake, 7 km long, 4 km wide and 21 km2 in area; about
ten times the original size and fish production has risen since its change. The southeastern part of the lake is covered with aquatic higher plants, and is difficult to negotiate by boat. The water is brownish in colour and remarkably muddy.
The Fisheries Station has a number of fish ponds outside the dam basin. Table 1
shows some results of our survey in this lake and the fish culture ponds.
Bum Borapet. This waterbody is situated in the central part of Thailand and is the
largest one in this country. After the construction of a dam, its surface area enlarged,
and its water level was regulated. The largest Fisheries Station in this country is set
up on the lakeside. The south-eastern part of the lake, around Stations I1 and 111,
is well-covered with large quantity of aquatic higher plants. This lake is shallow, the
deepest point being only 5.2 m deep. The open water is brownish in colour and muddy.
The small water areas enclosed with aquatic plants are dark brownish as in Khan
Payao Lake. Some results of measurement in Bum Borapet are shown in Table 2
and the research stations in Fig. 2B.
The Nam Pong Reservoir. This is situated in the north-eastern part of Thailand,
was completed in 1965for generating water-power electricity. This is the second deepest
lake in this country, the deepest point of g m being near its dam site. A morning market
is held on the lakeside, so we could get some fish samples there. The colour of water
was dark brown and a great many clusters of blue-green algae, Microcystis incerta,
were floating, which suggests that this lake is eutrophic. Submerged and dead forest
trees were seen standing around Station 11. We were told that the reservoir was
being studied by the Fisheries Department and Chulalongkorn University, though the
results have not been published yet. Our research stations are shown in Fig. 2C and
some results in Table 2.
Nong Raharn. This shallow lake in the north-eastern region was a natural lake some
years ago, but at present it is converted into a reservoir 5 9 km in area even in the dry
season. According to local information the deepest spot was 3 m deep, but we could not
locate it, and our measurements were only 1 m deep or less. It has many floating islands
and the bottom is covered with ValZisneria sp. The colour of water was dark brown.
Results of vertical measurements (Fig. 3): We studied the vertical distribution of
temperature, pH, transparency and dissolved oxygen at the deepest stations of Khan
Payao Lake, Bum Borapet and the Nam Pong Reservoir, but generally we could not
find any definite thermocline, and the pH value from the surface to the bottom with
the exception of the Nam Pong Reservoir, seldom showed any change. There was,
however, a remarkable decrease of dissolved oxygen near the bottom. I t occurs regardless of any water temperature stratification. This seems to be a characteristic of tropical
T. MIZUNO AND S. MORI
80
waters, which we should study in detail in our future work together with its effect on
the lives of organisms.
Rivers
There are the Mae Nam and Mae Klong-Mae Kwai water systems in West Thailand
and the Mae Kong water system in East Thailand. We investigated only the western
water system this time.
Table 1. Results of observations in Khan Payao Lake and some fish culture ponds
(27 April 1968)
Khan Payao Lake
Stations
I
2.0
Depth (m)
Air temp ("C)
Water temp ("C)
(at surface)
pH (at surface)
Colour of water
brown
Transp. (m)
Benthos
0.5
none
Detritus
plenty
Aquatic plants
little
-
27-0
8.8
26.5
9.2
-
none
Ponds
111
IV
V
VI
1*o
1
2.2
2.3
-
26.5
6.4
dark
brown
-
plenty
plenty
plenty
A.7 No. 14
-
-
-
-
25.5
25.5
-
27.1
8.9
brown
29.3
7.0
29.3
6.9
30.0
7.3
-
-
-
-
-
-
-
27-0
27.2
6.8
9.1
dark
brown
brown
1*o
none
may
bivalves
plenty
small
amount
plenty
little
none
-
s o
-
A.6
none
small
amount
little
Table 2. Results of observations in Bum Borapet (4 May, 1968),
Nam Pong Reservoir and Nong Raharn (19 May, 1968)
Bum Borapet
Stations
,
5.2
Depth (m)
Air temp ("C)
29.7
Water temp ("C)
(at surface)
29.7
pH (at surface)
7.7
Colour of
water
brown
Transparency
( 4
Benthos
Detritus
0.45
many
mussels
plenty
Aquatic plants small
amount
Notes
,
I1
111
I
I1
111
Nong
Raharn
1 *7
28.5
2.3
28.5
9.0
27.5
6.0
-
5.0
-
1 .o
32.0
28.5
7.3
28.5
7.7
30.5
7.8
-
-
32.8
8.8
dark
brown
brown
dark
brown
dark
brown
dark
brown
0.9
many
mussels
plenty
0.45
many
mussels
small
amount
plenty
2.7
none
I
I
Nam Pong Res.
Stations
plenty
I
none
none
afew
chironom.
small
amount
none
-
dark
brown
1*o
none
none
small
amount
none
plenty
blue green algae in quantity
among submerged and dead
forests
plenty
bottom was
covered
with weeds
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
FICURB
2. Map of some lakes in Thailand.
81
T. MIZUNO AND S. MORI
82
0
2
4
6
7
8
9
20
30
" I b
T
),r
'
I
0.44
\*
Otcc/I
0
pH
OC
2
4
6
7
Ib
8
9
20
30
Oecc/I
oc
0-
I
Tr
-f
E
2-
z 4-
WT
Burn eOmpe4
6-
Nom pOn0 Resmoir
Khan Payao Lake
FIGURE
3. Vertical distribution of some factors in investigated lakes of Thailand.
Table 3. Results of observations in the Mae Nam Water system
Water
temp.
("C)
pH
Turbidity
in
cm
Date
Time
Air
temp.
("C)
(1) Mae Sa fall
24.W
15.45
33.0
27.2
8.8
60
(2) Mae Nam Kuang
26.IV
10.00
26.5
27.2
7.5
5
(3) MaeNam Kuang
26.W
11.00
31.5
30.0
7.5
16
(4) Smallstreamnear
Lampang
(5) Mae Nam Ngao
(6) Small stream near
Ngao
(7) Mae Nam Yang
26.IV
13.00
32.4
29.0
7-7
5
26.IV
28.W
15.00
10.00
29.2
29-2
-
7.4
7.4
1
60
28.IV
10.20
25.4
27.8
8.2
4
(8) Smallstreamnear
Prae
(9) MaeNamYon
28.W
11.30
31.0
31.4
8.3
60
28.IV
18.00
26.8
30.0
7.6
9
(10) Sukohtai
29.IV
10.00
27.0
29.3
7.7
8
(1 1) Mae Nam Ping
29.IV
13.00
28.0
29-8
7.5
40
(12) Nakomswan
(13) Singhaburi
30.1V
30.IV
8.50
14.20
37.0
29.2
32.3
7-7
7.6
3
8
Name of Rivers or
research stations
-
Colour
of
water
clear
Benthos
insects
snails
yellowbrown
yellowbrown
yellowbrown
red-brown
clear
insects
planarians
yellowbrown
clear
yellowbrown
yellowbrown
yellowbrown
red-brown
yellowbrown
Turbidity here and in Tables 8 and 9 was assessed in a turbidity meter, as used in water purification stations.
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
83
The Mae Nam water system. (Fig. 4). Mae Nam has many branches in its upper
reaches from Nakorn Swan or Bum Borapet, that is, Mae Nam Ping-Mae Nam
Kuang, Mae Nam Wong, Mae Nam Ngao-Mae Nam Yom and Mae Nam Yan. We
investigated some conditions of these rivers and Table 3 shows the results.
The water was usually brownish-yellow or brownish-red in colour and was so muddy
that a Secchi disk could barely be seen in a few centimetres depth. Perhaps the cause
for it is the presence of laterite which is widely distributed in Thailand but with little
gravel. We found a few clear-water stations in the upper reaches of the Mae Nam and
collected some samples of insect larvae there.
FICUR~
4. The Mae Nam and River Kwai water systems.
T. MIZUNO A N D S. MORI
84
The River Kwai water system. T. Mizuno took part in work of the research team of
the Fisheries Department and surveyed the River Kwai from 3 to 8 of June. Figure 5
is the research map of it near Kanjanaburi and Table 4 shows the data obtained from
this river.
I
FIGURE
5. Research map of the River Kwai near Kanjanaburi.
Table 4. Results of observations in the river Kwai (3-8 June, 1968)
Name of River
Kwai noi
I
Air temp. ("C)
Water temp. ("C) (at surface)
pH (at surface)
Dissolved 0 2 p.p.m.
Colour of water
Bottom soil
Benthos
4 June
29.0
30.0
8.2
7.8
brown
sand
many snails
mayfly larvae
Kawi yai
Stations
Meklong
VII
I1
7 June
33.0
30.5
8.1
8.2
dark brown
sandy mud
many snails
5 June
31.0
31.0
8.2
5.8
brown
soft mud
snails
The water volume and speed of current in the Kwai yai were larger than those in the
Kwai noi. The water was yellow-brown or dark brown in colour and remarkably
muddy. Traces of flood erosion clearly remained on the cliff by the riverside and it
could be surmised that its water-level in the rainy season would rise up to 5-7 m at
least. The p H was 8.1-8.2. It was very interesting to find that in these rivers innumerable small snails were living, scattered like sandy grains. It suggests that the production of some of these rivers is very high. We will touch again on this matter later.
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
85
Cambodia
We investigated the Great Lake and the Tonle Sap River. The area of the Great
Lake is about 3000 km2even in the dry season, but is said to expand to as much as about
10,000 km2 in the rainy season, and thus serves as a natural regulator of thc Mekong
River. We measured the Tonle Sap River on the evening of 23 May, the southern part
of the lake (off Krakor) on 24 May and the northern part of the lake (off Siem Reap)
on 25 May. Some results are shown in Table 5.
The Tonle Sap River was more than 15 m deep in some places, but the Great Lake
is only 1 m deep or less even at those places several kilometres off the shore of the lake,
though it must be added that the measurement was conducted in the dry season. The
water was brownish-yellow and very muddy, and its transparency was for only 3 or 4
cm. In spite of this remarkable muddiness, the whole surface of water was covered with
blue-green algae and there grew a beautiful pink ‘water bloom’, especially in the
northern part of the lake. It suggests that the lake is eutrophic with a p H of 8.8.
In comparison with the little amount of green algae and diatoms, the zooplankton
were very abundant. Collections of benthos contained many molluscs, but contrary
to our expectation, only few Chironomus larvae and Tubifex. These quantitative data
will be explained in another chapter.
Malaysia
There are some large lakes, such as Tasek Bera, Tasek Merah, Chendero Lake and
some small lakes in Malaysia. Besides, there are many rivers and innumerable pools
left by open-cast mining.
Lakes
Tasek Bera. This is a swamp-like lake surrounded by a virgin forest and in shape
like a starfish. It is a source of the River Pahang. The shallow parts and edges of the lake
were covered with sedge and rush associations. We could find the same aquatic plants,
such as Nymphoides and Utricularia, as in Japan in still water, and Cryptocoryne and
Utricularia in running water, with Pandanus bush. Not only a large amount of the
primary production occurs in water, but also a huge amount of detritus is poured in
from the surrounding forest. The water was dark brown, of the tropical ‘black water’
type. The local inhabitants say that the water-level rises up to about 3-4 m in the rainy
season every year. Table 6 shows some of the measured results.
Figure 6 shows the vertical distribution of some factors at Station A (the centre of
the open water) and at Station B (the deepest point there). At station A the p H was
4.7and strongly acid, the transparency 2.1 m and the depth 3.2m. Detritus on the bottom
was thick, but no benthonic animal was found in it. At Station B, the depth was 4.5 m
deep, the pH was 4.7 from surface to bottom and no detritus and no benthonic animals
could be found. We collected only a little white-grey clay, because the water current
possibly washed all detritus away from there. At Station C, water was running through
the forests, with the pH at 4.6. The bottom soil except at Station B had a strong
unpleasant smell, especially at the shallow places near or in the forest. We observed
small snails
Benthos
(4
Colour of water
T-P.
1-0
29.0
31.0
8.2
30.5
62.0
yellowbrown
1
Depth (m)
Air temp. (“C)
Water temp. (“C) (at surface)
pH (at surface)
Mud temp. (“C)
,
yellowbrown
-
12.0
2
3
yellowbrown
stations
Tonle Sap River
3.0
29-0
31.0
8-2
30.5
62.0
yellowbrown
small snails
4
3
,
,
8.4
8.0
yellowbrown
4.0
30.0
33.0
36.5
32.0
yellowbrown
snails
bivalves
-
0.5
2
0.5
1
Southern Part
Stations
yellowbrown
snails
bivalves
-
3
Great Lake
,
Table 5. Results or’ observations in the Tonle Sap and the Great Lake (23-25 May, 1968)
31.0
3.0
yellowbrown
bivalves
8-8
0.9
36.0
39.0
1
*
’
Tulnfex
yellowbrown
Chirmmr
-
-
8.8
32.0
-
2
0.3
Northern part
Stations
z
YJ
U
3
*z
C
zG
3
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
87
Table 6. Results of observations in Tasek Bera. A and B are the stations in the enlarged part
of the lake and C is the station in the part of running water connected with the lake. The
survey was made from 16 to 18 May, 1968
Stations
B
A
Depth (m)
Air temp. ("C)
Water temp. ("C) (at surface)
pH (at surface)
Colour of water
Transparency (m)
Benthos
Detritus
Aquatic plants
C
3.2
27.0
28-0
4.7
4.8
27.0
27.8
4.7
1-0
30.0
dark brown
dark brown
dark brown
27.8
4.6
2.1
-
none
plenty
large amount
none
none
large amount
-
none
plenty
large amount
numerous bubbles coming out from the bottom when we pushed the bottom with an
oar, which might be methane gas. There were no benthonic animals, but a lot of small
animals such as Chironomus larvae, leeches, cladocera, small shrimps were found
amongst the aquatic plants there. This is an important problem which cannot be
overlooked in the secondary production study.
0
2
4
6
4
5
6
Otcc/I
0-
2
I
Tasek BemA
: ," i
-
4-
2''
102
I
IpH
i iI
./-
I
i
i
i
iWT
i
FIGURE
6. Vertical distribution of some factors in Tasek Bera.
Tasek Merah and the Cameron High Land Reservoir. Tasek Merah had been a swamp
at the confluence of the River Kurau and others before the construction of a dam. I n
1962 it was embanked first and the present man-made lake was filled in 1965. T h e
water-level of this lake has become stable, with the depth of the lake kept at 10 m at the
dam-site and the area at about 30 km2.It had black water, with a p H of 5.5-5.8, hence
weaker acidity than in Tasek Bera. A vast dead forest was seen on the shore drowned
by the dam construction.
T h e Cameron High Land Reservoir is a small man-made lake in the mountains
1300m above the sea level, and was constructed by damming up a river. T h e water was
T. MIZUNO AND S. MORI
88
running in its upper part, where the colour was brownish yellow, but in its middle and
lower parts, the water was stable and the colour of it was brownish green with a rich
zooplankton.
Table 7. Results of observations in Tasek Merah and the Cameron High Land Reservoir
(17-18 June, 1968)
Tasek Merah
,
Cameron High Land Res.
1
Stations
,
1
5 -0
Depth (m)
Air temp. ("C)
33.0
Water temp. ("C)
(at surface)
31.8
pH (at surface)
5.5
Colour of water dark brown
Transparency
(m)
0.9
Benthos
Chironomus
snail
Detritus
plenty
Aquatic plants
none
2
18 June
,
3
Upper
Pa*
Middle
Part
17 June
Lower
Part
22.5
-
23.0
31.2
5.8
-
-
dark brown dark brown
-
-
ephem.
snail
little
Plenty
little
-
18.0
6.6
red-brown
7.0
green-brown green-brown
0.3
little
plankton
rich
plankton
0.4
-
none
none
none
Tin-miningpools. There are many tin-mine pools near Ipoh and Kuala Lumpur in
the western part of Malaysia, and some near Gambang in the eastern part. The Malaysian PF Subcommittee is taking up these pools as a subject of their IBP work. We
observed five pools in the suburbs of Kuala Lumpur and four pools near Gambang,
and some results are shown in Table 8.
From these results it seems there were various types of ponds. Pool I and Pool 11,
both eight to ten years old were very eutrophic by inflow of sewage from pig- and duckpens on land and were used for culturing fishes. The water of the Pools 111, IV and V
was clear; the Pools I11 and V are well-established, and some young fish, some snails,
and Nitella or Chum were found living in them. In Pool IV mining work was still
going on and the white soft silt deposited there made access difficult. The Pools VIII
and IX near Gambang showed a pH value of 4.4; the former was an old pool with clear
water, and the latter had remarkably muddy water due to mining operation. The pH
value of Pool IV was very different from that of the Pool IX in spite of both being
under operation. It suggests that the pH value of water is affected seriously by the
character of the soil.
River
The River Pahang rises from the central mountains, gathers water from many
branches, and flows into the Borneo Sea. The above-mentioned lake of Tasek Bera
is one of the important sources of this river. Water temperature, pH, turbidity and
yellowgreen
60
white-grey
sandy mud
a few
chironmus
whitebrown
3
white-gre y
silt
none
21 June
I1
1.5
32.0
32.2
7-6
I
2.1
30.9
30.0
7.2
Turbidity, see remark to Table 3.
Benthos
Turbidity (cm)
Bottom mud
p H (at surface)
Colour of water
Depth (m)
Air temp. ("C)
Water temp. ("C)(at surface)
1
Southwestern
Stations
snails
white silt
60
cobalt
blue
27.5
29.3
9.0
-
111
none
60
white silt
none
60
white silt
clear
27-5
31.2
V
8.8
clear
27.5
31.0
8.7
26 June
I\'
Northern
Stations
Kuala Lumpur
*
r
-
18
-
redbrown
whitebrown
25 .
25.0
28.0
5-8
VI I
4.4
26-5
29.0
VIII
-
60
clear
25 June
Gambang
Stations
24-8
29.5
6.0
VI
Table 8. Results of observations in tin-mining pools
-
1
-
44
yellowbrown
27.0
27-8
-
IX
$
3
3
U
z
3=
r
z
sz
m
%
m
cn
w
U
X
cc
90
T. MIZUNO AND S. MORI
FICURB
7. The River Pahang water system and its research stations.
colour of water, were measured on our way across the river. Figure 7 shows the water
system and its research stations, and Table 9 indicates the data obtained.
As a whole, the water of the River Pahang was weakly acid and was brownish yellow
or brownish red in colour. The pH values of the branches were mostly between 6.2
and 7-0; but that of Kg. Lubok Paku half way up the river was 5.7, because a large
amount of the strong acid water with a pH 4.6-4.7 flowed from Tasek Bera into the
main stream. We found some aquatic insects, larvae and small shrimps in some spots
of its upper reaches, that is, in S. Benus near Bt. Tinggi and S. Gombak.
AQUATIC COMMUNITIES
T o measure biotic productivity in these tropical inland waters, we have first to
establish the community constituents. Because of the brevity of the trip, the samples
collected were fragmentary both quantitatively and qualitatively, but will be useful
for our future operations.
Many specialists helped us to identify the species: the aquatic higher plants were
identified by Dr S. Miki, Mukogawa Women’s University and Dr S. Ohi, the National
Science Museum; the freshwater clams and snails by Dr T. Habe, the National
Science Museum; the freshwater shrimps by Dr T. Kamita, Shimane University;
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
91
Table 9. Results of observations in the Pahang River system
Name of River
Date
Time
("C)
Water
temp.
("C)
22.VI
23.VI
25.VI
25.W
25.VI
25.W
23.W
25.W
25.W
23.W
23.VI
22.VI
25.W
25.VI
25.VI
17.00
12.00
13.10
12.10
12.45
14.00
14.00
12.30
10.25
15.40
16.00
10.25
14.25
14.10
15.35
32.0
31.0
30.8
29.0
31.0
34.0
34.0
29.0
27.0
33.0
33.0
27.2
29.5
30.0
30.5
28.5
28.0
30.5
26.5
27.0
29.0
29.0
27.0
26.5
29.0
33.0
23.0
28.3
28.0
27.0
6.9
6.8
6.8
5.7
6.7
6.6
6.6
6.6
7.0
6.2
6.2
6.9
6.8
6.8
6.8
-
(6)
(7)
(8)
(9)
(10)
(11)
(12)
(13)
(14)
(15)
S. Jurnch
S. Jelai
S. Pahang
S. Pahang
S. Tekam
S. Jengka
S. Jampol
S. Jampol
S. Luit
S. Lepar
S . Lepar
S. Tanglir
S. Benus
S. Benus
S.Benus
(16)
(17)
(18)
(19)
(20)
S . Telemony
S. Teris
Tasek Bera
Tasek Bera
S. Gombak
25.W
25.VI
16.V
16.V
22.VI
15.00
14.00
9.00
14.00
9.00
32.5
32.0
27.0
30.2
-
27.0
27.0
26.5
27.8
23.0
6.6
6-2
4.7
4.6
6.9
25
5
Air
temp.
(1)
(2)
(3)
(4)
(5)
pH
Turbidity
(cm)
14
20
18
50
50
19
30
18
45
29
14
60<
60<
Colour of
water
brown
brown
red-brown
red-brown
brown
brown
brown
y ellow-brown
yellow-brown
red-brown
red- brown
yellow-brown
yellow-brown
yellow-brown
clear
yellow
yellow-brown
dark brown
dark brown
clear
Incidental
observations
insects &
Spirogyra
insects &
shrimps
Turbidity, see remark to Table 3.
the aquatic insects by Dr T. Kawai, Nara Women's University; the freshwater fishes
by the Department of Fisheries and Fishery Station in Thailand, Dr Y. Shiraishi,
Nikko Fisheries Station and Dr N. Mizuno, Ehime University. We acknowledge
their kind help.
Aquatic higher plants
In the lakes during the dry season aquatic plants are abundant. We could find emerged
plants near the shore, and submerged and floating plants in the shallow waters. Also,
we could observe floating plants such as Eichhornia crassipes and Pistia stratiotes var.
cuneata, and also members of the Juncaceae, Cyperaceae and Gramineae growing on
floating islets.
In Khan Payao Lake, Bum Borapet and Nong Raharn of Thailand some parts of the
water surface were so densely covered with aquatic plants that boating was difficult.
Also, it was interesting to note that the bottom of Nong Raharn was covered with a
thick mat of Vallisneria.It is intended to measure their standing crops, their production
and decomposition rates, and the amount of detritus deposited.
The Great Lake in Cambodia has great quantities of emerging plants and also shrubs
growing on an extensive flood plain spreading several kilometres wide around the lake.
This shows that a vast amount of organic matter must flow into the lake, especially
when the water level rises up to 8-10 m in the rainy season.
92
T. MIZUNO AND S. MORI
Tasek Bera in Malaysia has a large amount of aquatic higher plants, too. Furthermore,
it is surrounded by a virgin forest and the inflow of detritus must be great. When the
water level rises up to 3 or 4 m during the rainy season, water floods the forest. Future
enquiry will have to include the forest floor as one of the important sources of nutrients.
In Tasek Merah, we also observed a large forest now dead due to its dam construction
and the same in the case of the Nam Pong Reservoir in Thailand. In such cases there is
an interesting problem as to how this kind of temporary nutrition supply from the dead
forest affects the communities of the lake. In these circumstances the primary production problems in such tropical inland waters can only be solved by including both
aquatic plants and surrounding forests.
Table 10 is a list of the dominant species which we collected or observed in the
tropical lakes during our present trip.
Table 10. List of the dominant species in the aquatic plant communities observed during the
trip (+ shows the degree of commonness)
Khan Payao Lake Bum Borapet
Floating plants
Eichhornia crasszpes
Pistia stratiotes var. cuneata
Salwinia auriculata
S. imbricata
Spirodela polyrhiza
Lemna paucicostata
Utricularia aurea
U.flexuosa
Nymphoides indicum
Trapa sp.
Submerged plants
Ceratophyllum demersum
Najas kingii (or Najas oguraensis)
Hydrilla verticillata
Potamogeton malaianus
Vallisnetia asiatica
Cryptocoryne grifithii
Blyxa Malaysiana
Hygrophytes on the floating island
Cyperus cephalotes
C . platystylis
Panicum paludosum
Ludwigia stiputacea
Dendrobium alozfolium
Platanthera hologlottis
Lyncopodium sp.
Flagelkaria indica
Eriocaulon sexangulare
Fuirena umbellata
Lepironia articulata
Eleocharis ochrostachys
Others
Pandanus spp.
+++
+++
+++
+++
+++
+
+++
+++
+++
+++
+++
+++
Nong Raharn
Tasek Bera
+++
+++
+++
+++
++
+
+++
+++
+++
++
++
+++
+++
+++
++
++
+++
+++
+++
+++
+++
++
++
+
++
++
+++
+++
++
++
+++
+++
++
+
+
+
++
++
+
+++
+++
+++
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
93
Plankton
The tropical freshwater plankton of Java and Sumatra was studied by the German
Sunda Expedition (1928-1929). M. U h o (1964, 1966) reported on some of the freshwater organisms and zooplankton in Southeast Asia; D. S. Johnson (1962a, b) summarized the Cladocera in Malaysia and M. Hirano (1967)described the flora of phytoplankton in Thailand. The seasonal fluctuation of plankton in the tropical lakes and
ponds were reported by J. Blache (1950,1951),while R. G. Michael (1968),S. V. Ganapati (1969),S. V. Ganapati and C. H. Pathak (1969)and R. N. Singh (1969)investigated
ponds and paddy fields and a reservoir in India. But thorough studies of production
processes in plankton have not yet been accomplished. T. Mizuno, one of the authors,
has found many southern species in the plankton fauna and flora of Japan and is
interested in the bio-geographical origin of these species. T h e Tables 11A and B show
the composition of planktonin Thailand, Cambodiaand Malaysiaas found in our survey.
Some features of the fauna and flora, as a whole, are here stated. T h e Great Lake
of Cambodia is remarkably muddy and its transparency is 3-5 cm deep. Therefore,
it seems that the trophogenic layer is restricted within very narrow limits. In fact,
only small quantities and a limited number of species of diatoms and green algae could
be found, whereas the blue-green algae such as Microcystis 90s-aquae, M. aeruginosa,
Sphaerozoma vertebratum, Oscillatmia limnetica, Anabaena sp. frequently formed
‘water blooms’. It seems that the blue-green algae increase regardless of the turbidity
because of accumulation near the surface. The same appears in Khan Payao Lake,
Bum Borapet, the Nam Pong Reservoir and Nong Raharn, though the species of the
blue-green algae are different from those of the Great Lake. In the Nam Pong Reservoir
we observed Microcystis imerta in abundance.
In contrast zooplankton Cladocera and Copepoda occur even in strikingly turbid
water. The dominant species of Cladocera are Moina dubia, Diaphanosoma ex&m,
Bosminopsis deitersi, Bosmina longirostris. They appear also in lakes and ponds in Japan,
except Diaphanosoma excisum which in Japan is replaced by D . brachyurum.
Rotifera are almost the same as in the summer fauna of Japan. The biomass of zooplankton seems to be larger than that of phytoplankton suggesting that the secondary
producers depend not only upon phytoplankton but also upon other energy sources
like micro-fragments of organic matter, bacteria, nutrients found adhering to abiotic
seston and so on. We have to discover this supply route of energy to the secondary
producers.
Tasek Bera and Tasek Merah have black water, which is fairly transparent. I t is,
however, strongly acid and seems to limit zooplankton propagation. There is a rather
abundant phytoplankton of desmids like Euastrum, Micrasterias, Desmidium, Hyalotheca, Staurastrum, Cosmarium, Xanthidium, Closterium, and of diatoms like Eunotia,
Frustulia and others. It is interesting to note that they are very similar to the species
which appear in dystrophic lakes in the northern part of Japan as well as in the boggy
parts of our high mountains.
On aquatic plants or among them, we found a considerable number of attached or
semi-attached small animals; for example, small shrimps, leeches, water mites,
Chironomus larvae, etc. The standing crops and the turn-over rates of these attached
small animals besides the floating zooplankton will require attention.
7
T.MIZUNO AND S. MORI
94
Table 11A. Freshwater plankton in lakes of Thailand and Cambodia
-~
~
Lakes and ponds
I
t
I
I1
111
\
Bum Borapet
Khan Payao Lake
SbtiOM
Stations
v
-,
I
I1
IV
Nam Nong GreatLake
Pong RaRes. ham South North
Species
Cladocera
Moinu dubia
Diaphanosomaexdsum
BonniM longirosm*s
Bomdnopris dcitern'
Alona guttata
Simomphalusvetulus
Cerioakphniarigaudi
Chydmssphamrcis
Macrothrix spa'nosa
Copepoda
Thennocyclopssp.
Cyclops rp.
Paracyclopsfima*iotur
Mesocyclops leuckmh'
Helialiaptmnur
kikuchii
Sinodiaptmnurvolcanoi
+++ ++ ++ ++
+
++
++
+ +
+ ++
+
++ ++
Rotifera
Brachionus c a l y d w
B.forficula
B. falcatus
B. caudatus
Polyarthra trigulo
Asplanchnapriodonta
Trichocerca iign's
Tetramash opolienris
Keratella valga
Testudinellapatina
Diurella sp.
Protozoa
Diflugia corona
P d m u m sp.
Ceratium hirundinella
Eudonha elegans
Euglena proxima
E. a m
E. viridis
Phacus longicauda
Volvox aureus
Dinobryon cylindricum
D. sertularia
Arcella sp.
Cy=whYta
Microcystisaeruginosa
M.jlos-aquae
M . incerta
Aphani2omenon
ovm~sporu?n
Aphanocapsa
dclicatissima
+
+
+
+
+
++
++
+
++
+
+
++
+
+ +++
+
+
+
++++
++++
+ +++ ++
+
+
+
+
+
++
+
+ +
++ ++
++
+
+
+++
+
+
+
++
+
+
+++ +++ +++ ++
+ ++
++
++
+
+
+
++
++ ++
+
+
OStlWJda
++ +++
+ +
++ +
+ ++
+++ +++ +++
+++ ++ + +++
++
+
+
++
+++
+
+
+++
+
+
+++ +++
HYDROBIOLOGY OF SOME
95
S.E. ASIAN INLAND WATERS
11A (continued)
Table
Lakes and ponds
r -
~
Khan Payao Lake
Stations
Bum Borapet
Stations
,
I
I
Cyanophyta-continued
Gloeotrichia echinulata
Sphaerozosma
vertebratum
Oscillatoria limnetica
Lrngbra SP.
Anabaena sp.
Chlorophyta
Scenedesmus spp.
Cosmarium spp.
Micrasteriasfoliacea
Xanthidium sp.
Pleurotaenium sp.
Closterium sp.
Westella botryoides
Oedogonium sp.
Mougeotia sp.
Bacillariophyta
Melosira granulata
Synedra ulna
Rhopalodina sp.
Navicula sp.
Gylosagma sp.
I1
v
I11
,-
Nam Nong GreatLake
Pong Ra- &
Res. harn South North
c
I1
I
IV
+++
++ +++ ++
++ ++
+
++
+
+
++
+
++ ++
+
++
+
+
+
+ + ++
+
++
++
+ ++ ++ ++
+
+
+t
+
Table 11B. Freshwater plankton in standing waters of Malaya
~
Lakes and ponds
Tasek Bera
Tasek Merah
- \
I
Stations
I
I1
I11
Min-
, Utrid.among
I
A
B
Dam
assoc.weeds shore centre site
ing
pools
Species
Cladocera
Moina dubia
Diaphanosoma excisum
Bosm'nopsisdeitersi
Alona guttata
Macrothrix spinosa
Camptocerus rectirostris
Chydorus spham'cus
Copepoda
Paracyclopsjimbriatus
Tropocyclopsprasinus
Rotifera
Keratella valga
Asplanchna sp.
Platyim quadricom's
Monostyla sp.
Diurella sp.
+++
+++
+
++
+
+
++
+
++
+
++ ++ + +++
++ +++ + ++
+++ +++ ++
++
+
+
+
++
+++
+
++
T. MIZUNO AND S. MORI
96
Table 11B (continued)
Lakes and Ponds
,
,
CYanOPhyta
Merismopedia elegans
OSL-illatOria
pseudogeminata
0.geitleriana
P h m ' d i u m inundatum
P. minnesotmse
Microcystis alln
Lyngbya sp.
Anabaena sp.
Bacillariophyta
Eunotia lunaris
E. monodon var.
tropica
E. thronii
E. pectinatis var.
minor
E. sudetica var. incisa
E. faba
E. gracilis
E. paludosa
Diatoma elongaturn
D. vulgaris
Neidium iridis var.
amphigomphus
Frustulia rhodoides
Surirella linearis var.
-*
I
I1
+
+
SUbjenn#i
111
, Utriml. among
A
B
Dam
assoc.weeds shore centre site
++
++
+
+++
+
+
+
+
+
+
+
++ ++
+
++
+
+
+
+
+ +++ +++
+
++ ++
++
+++
+
+
+
+
+++
+++
+++ +++ + +++ ++++ +++
+
+ ++
+
+
+
+
+
t
+
+ ++
+
+
+
+
+
t
t
+
+ ++
++
+++
+
+
Mining
pools
++
f
constricts
S. elegans
Navicula sp.
Pinnularia nobilis
Astm'onella sp.
Cymbella sp.
Synedra ulna
Chlorophyta
Euastrum ansatum var.
pyxidatum
E. binale var.
gutwinchii
E. denticulatum
E. didelta var.
bengalicum
E. sinuosum var.
reductum
E. sinuosum var.
Tasek Merah
Stations
,
Protozoa
Euglena vi+idiz
E. proxima
Dinobryon sertularia
Trachelomonas sp.
Peridinium sp.
Ceratium hirundinella
Dzflugia corona
,
Tasek Bera
97
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
Table 11B (continued)
Lakes and Ponds
r
1
Tasek Bera
Stations
I
I
Chlorophyta-continued
E. cuncatum
E. anstatum var.
javanicum
E. ampullaceum
E. robusta
E. sp.
Micrasten'as lux
M . foliacea
M . alata
Desmidium coarctatum
D. beilq'
Hyalotheca dissiliefis
H. mucosa
Gonotozygon
monitifonnis
Pleurotaenium
subcornulatum
P. trabecula var. rectum
Triploceras gmcib
Staurastrum
tohopekaligense
S. alambae var.
angulata
S. .pracile var. ornatum
S . dejectum
S. megacanthum
S. subsaltans
S. sebaldi var. gracile
S. ensiferum
S. Wanda var.
brevispinum
S . manfeldii var.
anulatus
Xanthidium
bengalianum
X . antilopaeum var.
laeve
X . burkrllii
Arthrodesmusimpar
Cosmarium alpecta
C.circulare f. minor
C.contractum
C.mkbulatum var.
printzii
C.maximum
C.diplosponum
c. spp.
I1
crassius
intewuptum
A
.
MinUtrinrl.among
Dam ing
B aeeoc.weeds shore centre site pools
t
+
+++
+
++
+
+
++
+
+
+
+
+
+
+
+
+
+
+
++
+
+
+t+
+
++
+++
+
+
++
+
+t
+
+
+
+
+
++
++
+
+
+
+
+
+
+
+
+
+
++
t
+
+
++
++t
Clostetium alpestre
C.lagoenre var.
C.dianae
C.libellula var.
III
-,
Tasek Merah
+
+t
+
t
+
++
++
+
T. MIZUNO AND S. MORI
98
Table
11B (continued)
Lakes and Ponds
I
Tasek Bera
..
Stations
I
I
Chlorophyta-continued
C. setaceum
C. bioclatum f. depressa
C. ehrenbergii
C.westii
Costeropsis sp.
Mougeotia sp.
Zygnema sp.
Spirogyra sp.
Stigeoclonium sp.
Oedogm'um sp.
Scenedesmus bijuga
var. alternans
Codatella sp.
Actinastrum sp.
I1
I11
A
B
Tasek Merah
MinDam ing
assoc.weeds shore centre site pools
, Utricul. among
++
+
+
+++
+ ++
++
+
+
++ ++
+ ++ ++
++
+ +
+++ +++
+
++
+ ++
+
+ ++
+
++
+
Higher crustacea
Freshwater shrimps in the tropical inland waters have recently been studied by D. S.
Johnson (1962b, 1966), Singapore University and are fairly abundant in these tropical
waters. Table 12 lists the prawns and shrimps obtained in Thailand, Cambodia and
Malaysia (Figs 8 and 9), identified by Dr T. Kamita.
Table 12. List of Freshwater shrimps collected during the trip
Thailand :
Family Palaemonidae
Macrobrachiurn rosenbergii (De Man)
Locality; Burn Borapet, Mae Nam
M. lar (Fabricius)
Localities; Nam Pong Reservoir and Khan Payao Lake
Family Atyidae
Caridina wyckii Hickson
Locality;Nong Raharn
Cambodia
Family Palaemonidae
Macrobrachiurn rosenbergii (De Man)
Locality ;southern part of Great Lake
Young specimem
Malaysia
Family Palaemonidae
Macrobrachiurn lar (Fabricius)
Locality; Stream Gombak on the way to Bt. Tinggi
M. sundanicum (Heller)
Locality; Tasek Bera
Family Atyidae
Caridina pareparensis De Man
Locality; Tasek Bera
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
99
The largest specimen of prawn was Macrobrachiurn rosenbergii, with a body-length
of 320 mm, collected in Bum Borapet and the Mae Nam; smaller-sized specimens
were caught in the Great Lake. Macrobrachiurnlar was widely distributed in S. Gombak,
the Nam Pong Reservoir, Nong Raharn and Khan Payao Lake. Also, Caridina
pareparensis was commonly found among the aquatic plants of Tasek Bera in Malaysia.
They are small shrimps, about 30 mm long, and the females were holding many eggs
in their abdomens. Macrobrachiurn sundanicurn lived together with Caridina pareparensis amongst weeds and the aquatic plants there.
The remnants of small shrimps were often found in the contents of the stomachs
of fish taken from these waters. Judging from a great number of shrimps found living
in the tropical waters, they surely must form an important part of the secondary
producers and of fish food.
FICURB
8. Macrobrachiurn rosenbergii (De Man). Body length: 320 mm, adult. Locality; Bum
Borapet, Mae Nam.
In spite of the small quantities of benthonic animals captured with the Ekman-Birge
bottom samplers, a high production of fish was a prevailing phenomenon in the tropical
Asian waters. These numerous small animals living among aquatic plants, (shrimps,
Chironornus larvae, dragon-fly larvae, leeches, planaria, water boatman larvae, water
mites, and others), should contribute to the high productivity of some fishes, others
being herbivorous. Some of these benthic animals were studied by J. R. Pippet & C. H.
Fernando (1961), R. E. Sharma & C. H. Fernando (1961), C. Y. Leong (1961), T.
Imamura (1964) and others, but the identification of our material will take time to be
completed.
Aquatic insects
There are some papers about the aquatic insects in this area between 1920 and 1940;
for example, papers by F. F. Laidlaw (1923), C. Chilton (1925), F. C. Fraser (1927),
N. P. Vejabhongse (1937) and C. Suvatti (1938). Also, some papers by Japanese
100
T.MIZUNO AND S. MORI
C
d
D
G
FIG= 9. A-C,Mucrobrachiumlur (Fabricius),young; D, M . sundanicum (Heller),young ;E, M .
rosenbergii (De Man),young;F,Caridinaparepurenis De Man, adult;G, C . wyckii Hickson,tiny.
entomologists, M. Chujo (1941), S. Asahina (1961), T. Kawai (1961), M. UCno (1961),
have been published since 1940.
Only few stream bottoms were covered with gravel and even the upper reaches of
these streams were mostly muddy in Thailand, Cambodia and Malaysia. This differs
greatly from conditions in clear Japanese streams with the gravel bottom inhabited by
many insect larvae. Therefore, we had few opportunities to collect aquatic insects,
except in the upper reaches of the Mae Nam in Thailand and in a few mountain streams
in Malaysia. The collected samples are to be worked out by Dr T. Kawai, Nara Women's
University.
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
101
Table 13. Provisional list of aquatic insects collected during the trip
Thailand
I
I1
Stream
(Mae Sa)
Coleoptera:
Ephemeroptera:
Odonata:
Diptera:
Trichoptera :
Ephemeroptera:
Trichoptera :
Diptera:
Helodidae
Ephemerella sp.
Ephemerellidae
Ephemera sp.
Baetis sp.
Ecdyonurus sp.
Paraleptophlebia sp.
Caenis spp. (probably 2
species)
Libellulinae nymphs
Chironomidae
Polycentropus sp.
Hydroptidae
(Chieng Mai)
Baetis sp.
Rhyacophila sp.
Leptocerus sp.
Chironomidae
,
Stream
Lake
(Payao Lake)
Ephemeroptera :
Coleoptera:
Odonata :
Ecdyonurus sp.
Noteridae
Diplectrona sp.
Corydalidae (Protohennes
SP. I)
Etrocmema sp.
(Nymphal female)
Kiotina sp.
Unknown odonata nymph
Trichoptera :
Odonata :
Plecoptera :
(Bt. Tinggi)
Parastenopsyche sp.
Libellulidae
Neoperla sp.
Plecoptera:
(Nong Raharn)
Baetis sp.
Hydrophilidae
I
Malaysia
>
I
Lake
(S. Gombak)
Ephemeroptera :
Coleoptera:
Trichoptera:
Megaloptera:
Deielia sp.
Odonata:
(Tasek Bera)
Ephemeroptera :
Thraulus sp.
Cam’ssp.
I
Odonata:
Diptera:
Lepidoptera:
Hemiptera :
1I
Ephemeroptera :
Baetis sp.
Tngomphus sp.
Iotinogomphussp.
Cmiaysion sp.
Unknown odonata nymph
Chironomidae
Nymphula sp.
Auqrius sp.
(Tasek Merah)
Polymdarcis sp.
Some explanation is needed concerning the aquatic insects collected from the
bottoms of some lakes. Although the individuals collected with bottom samplers were
very few in number, the stomach contents of various fishes showed considerable
numbers of aquatic insect larvae. T h e interpretation of this phenomenon will be an
important problem for our future work.
102
T. MIZUNO AND S. MORI
Table 14. List of molluscs in Southeast Asian inland waters collected during the trip
Lakes and rivers
I
Thailand
,
Cambodia
Khan Bum Nam Nong
Payao Bora- Pong Ra- Kwai
Lake pet Res. ham River other
Species
(Mussels)
x
x
Pikdwyoconcha excilis Lea
xx
x
Ensidem ingallsianus (Lea)
X
E. rcobinata Lea
x
xx
Unio scobinata Lea
xx
U.thaiensis Habe
x
x
Hyriopris bialatus Simpson
H . myesiana Lea
X
Corbicula noetli-. Martens
X
c.siamenrir Prallhad
xx
c.pctiti Clessin
X
C . lamarchiana Prime
xx
C. lamaudien' Prime
Physunio gravidus Lea
Nodularia caerules (Lea)
Limnopema lucustris Martens
(Snails)
pila polita (Deshays)
X
P. ampullaria (Lime)
P. peomei Horlet
P. scututu Lea
Bithynia funiculata (Walker)
Pachydrobia rimnenSisBraudt
P . paradoxa Groase et
Fischer
Tarebia grayera
oblonguigranose (Smith)
Hydrobioides dautaenbmgi
X
Walker
Melanoides tuberuclatus
Miiller
Clea bandottiana (Mabille et
X
Mesle)
Mek&
bandoniana Lea
M . siamensis Frauedeld
xx
M . hainesiana Lea
M . pongensis Brandt
Indoplanorbis exustus
X
(Deshays)
Gyraulus chinencis convexiX
wculus (Hutton)
Thiara scabra 0 .F. MUller
X
Sinotaia ciliata basicarinata
Kobelt
S.jilosa (Reeve)
S . ingallsiana Lea
S. umbilicata hachides
(Martens)
9 1 3
Total
Malaysia
Tasek Mining
Merah Pools
X
X
X
xx
xx
X
X
X
X
xx
xx
xx
x
X
xx
X
xx
xx
X
X
xx
X
X
X
xx
X
xx
X
X
X
xx
X
xx
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
2
3
16
9
13
0
I
50
x and x x indicate degree of abundance.
Great
Lake
-
13
1
3
4
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
103
Freshwater mussels and mails
According to D r T. Habe the tropical molluscs have been studied well by many
foreign specialists since the time of foreign contacts. He has recently reported on the
freshwater molluscan fauna of Thailand (1964). W. Wenz and F. Hass (1968) have
also published a comprehensive monograph of mollusca in this area. We collected
16 species of bivalves and 22 species of snails during our trip. It seems that some of
them are widely distributed and that others inhabit limited areas. Table 14 shows a
list of the collected species and their distribution, and Plates I to IV show photographs
of these molluscs.
Table 15. Results of quantitative survey of Mollusca in some lakes (nos/m2)
Khan Payao Lake
Stations
7-p
~~
I
I11
I1
IV
L
E n s i b ingallsianus
Pilabryoconcha exilis
Unio scobinata
Corbicula siamensis
Hyriopsis bialatus
Total
V
VI
D
3 54
89
178
178
178
45
666
Bum Borapet
Stations
,
I
I
L
I1
D
L
Ensidens ingallsianus
Unio scobinata
U . thaiensis
Corbicula larnauchieri
C . petiti
C . ligidiana
Hyriopsis bialatus
Clea bandoniana
Pila ampullatis
Total
near rest house
L
D
D
45
89
89
444
400
45
45
45
445
L
D
45
89
223
45
223
446
178
311
178
980
133
45
578
224
~
I11
~
535
~
Great Lake
Great Lake
Stations
Tonle
Sap River
I
L
Ensidens ingallsianus
Umo thaiensis
Corbicula siamensis
C. petiti
Hyriopsis biolatus
Sinotaia ciliata basicarinata
Melanoides tuberculatus
Pachydrobia paradoxa
Mekongia hainesiana
Talebia granafera
abliquipanosa
Total
~
L
-
Living; D = dead.
D
South I
L
D
South I1
L
D
South 111
L
D
North I
L
D
North I1
L
D
45
178
45
311
45
311
89
89
89
89
45
286
45
1643
45
4.5
45
89
45
134
89
445
45
89
444
376 400
45
134
0
1643
0
45
846
104
T.MIZUNO AND S. MORI
At Station IV of Khan Payao Lake, the density of Ensidm ingallsianus was very
high, but at the other stations, none was found. The density of molluscs in Bum Borapet
was fairly high, ranging 200-700 individuals per m2 except at Station I. Furthermore,
the Great Lake in Cambodia had a higher density, 200-1500 per m2 at each station.
In general the productivity of mullosca is remarkably large in these areas.
In rivers, on the other hand, it was difficult to study the standing crop in the larger
ones, but fortunately we had an opportunity to find a large amount of mollusca in the
Kwai River, when we took part in the survey made by a team from the Thai Fisheries
Department. River sands here were found to consist mainly of small snail shells. Most
of them were Pachydrobia siamensis with some Mekongia pongensis, besides other
mussels and snails. Therefore, even if the running water is muddy, the productivity of
molluscs in the river bottom and shore seems to be very large.
On the other hand, we found fewer molluscs in Malaysia than in Thailand and Cambodia. Only some Sinotaia ciliata basicarinata, s. umbilicata trachioides, s.ingalhiam
in Tasek Merah, and a few Pila suctata in a tin-mine pool near Kuala Lumpur were
found, but no species in Tasek Bera, which may be attributed to the strongly acid water.
FiSites and their foods
Taxonomic studies of tropical Asian fishes have been made in the past by a number
of ichthyologists, for instance by P. Bleeker for 30 years from 1846 to 1876, by F. Day
from 1865 to 1889, by H. E. Sauvage from 1876 to 1883, by Max Weber & L. F. de Beaufort in 1913 and 1916, by s. Hora from 1922 to 1939, by G. s. Myers from 1923 to
1938 and by H. M. Smith from 1925 to 1941. Especially, H. W. Fowler (1934) made
a comprehensive compilation of the fish fauna of the Southeast Asia, and H. M.
Smith (1945) published a book entitled ‘The Freshwater Fishes of Siam or Thailand’.
J. Tiemmedh (1966) has given a well illustrated popular account of ‘Fishes of Thailand’.
Recently the Departments of Fisheries and Fishery Stations in each country have
been making efforts to gather data and compile lists of fishes, and at the same time
striving for the promotion of fishery. But most of the papers hitherto published are
taxonomic and only a few of them are ecological. S. Tongsanga & P. Kessunchai
(1964) ecologically treated the quantitative biomass of fishes in Sri-Ayuthaya Province
in Thailand, and D. S. Johnson (1967) discussed the relation between the distribution
of fishes and the chemical nature of water. Besides, some Japanese ichthyologists,
Y. Shiraishi (1967), Y. Taki (1967) and T. Shimazu (1968) reported the general
situation of fisheries in Southeast Asian countries.
We have collected about 80 species of fishes in Thailand, 16 species in the Great
Lake, and only five species in Tasek Bera. According to H. M. Smith‘s book, about
550 species are described in Thailand, and about 200 in Cambodia by Y.Shiraishi and
about 120 in Malaysia by the list of the Fisheries Department of Malaysia.
The samples collected in Thailand were identified by the specialists of the Department of Fisheries and Kasetsart University, and the others were done by Dr Y. Shiraishi
and Dr N. Mizuno after his return to Japan; some samples are still awaiting identification. Table 16 is a list of the collected freshwater fishes and Figs 10 to 13 illustrate
some of them.
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
FIGURE
10. A, Hampala macrolepidota Van Hasselt; B, H . dispar H. M. Smith; C , Morulius
chrysophekadion Bleeker;D, Pristolepisfasciatw (Bleeker) ;E, Puntioplitesproctozysron(Bleeker);
F, Labiobarbus sp. ; G , Puntius gonionotus (Bleeker).
105
106
T.MIZUNO AND S. MORI
FIGURE
11. A, Notoptem chitala (Hamilton) ;B, N.notopterus (Pallas);C , Ktyptopterus bleeken'
Gbther; D, K . rrupt0ptem.s (Bleeker); E, Ompok bimuculutus (Bloch); F, Pteropangasiuz
cultratus (H. M. Smith); G, Rasbora m'lineata Steindachner; H, R. sp.; I, Probarbus jullim'
Sauvage;J, Trichgaster trichoptmu (Pallas); K, Osphronemus goramy Lacepede; L, Xenentodon cancila (Hamilton).
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
107
108
T.MIZUNO AND
S. MORI
B
C
D
E
\
F
G
----.-..-_
.__._..__
...--.
___.__.
FIGURE
13. A, Mastocembelus armatusfavus Hora; B, M . maculatus (Cuv. & Val.); C, M a n o gnathus aculeatus (Bloch) ; D, Fluta alba (Zuiew) ;E, Wallagonia miostoma Valliant ; F, Ophiocephalus striatus Bloch; G, Trachysurus maculatus (Thunberg).
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
109
Table 16. Freshwater fishes collected in the tropical Asian waters during the trip
Species
Order
Euselachii
Isospondyii
Opisthomi
Synbranchia
Eventognathi
Dasyatis bleekeri
Setipinna taty
Lycothrissa crocodilus
Notopterus chitala
N . notopterus
Macrognathus aculeatus
Mastacembelus maculatus
M . armatus favus
Fluta alba
Hampala macrolepidota
H . dispar
Rasbora trilineata
Rasbora sp.
Cyclocheilichthys apogon
C . enoplos
C . repasson
C . mekongensis
C . amblyceps
Bum
Payao Borapet
Nematognathi
Synentognathi
Heterosomata
Labyrinthici
8
Nam
Pong
River
Kwai
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Tasek
Bera
+
+
+
+
Great
Lake
+
+
c. sp.
Probarbus jullieni
Puntius gonionotus
P. schwanenfeldii
P. orphoides
Puntioplites proctozysron
Osteochilus hasseltii
Labeobarbus burmanicus
L . spiropleura
L. leptocheilus
Botia modesta
Wallagonia miostoma
Ompok bimasculatus
Kryptopterus cryptopterus
K. bleekeri
Clarias batrachus
C . macrocephalus
Pangasius larnaudii
P. siamensis
Pteropangasius cultratus
Mystus vittatus
M . nemurus
M. cavasius
Heterobagrus bocourti
Tachysurus maculatus
Hemipimelodus siamensis
Xenentodon cancila
Synaptura panoides
Cynoglosus xiphoideus
Anabas testudineus
Osphronemus goramy
Trichogaster pectoralis
T. trichopterus
Ophicephalus striatus
0. lucius
0. micropeltes
Cam- Malay, bodia
sia
Thailand
,
+
+
+
+
+
+
+
110
T . MIZUNO AND S. MORI
Table 16 (continued)
Thailand
Cam-
, bodia
r
Order
Percomorphi
Gobioidea
Plectognathi
Species
Polynemus paradiseus
Chanda wolgii
Datnioides mimolepis
Pristolepisfasciatus
Toxotes chatareus
Pseudosciaena soldado
Tilapia melanopleura
Oxyeleotris mmmoratus
Tetraodon leiurns
Number of species
Bum
Payao Borapet
+
+
,
River
Kwai
Great
Lake
+
+
+
+
+
+
+
+
+
+
+
+
+
+
21
Nam
Pong
Malaysia
Tasek
Bera
13
29
+
21
,
16
5
a4
As shown in Table 16, all these species are different from Japanese ones except
Fluta alba (Zuiew). Noteworthy are the many kinds of bottom fishes like catfishes.
Also, it is interesting to note that some brackish and sea-fishes inhabit Bum Borapet
and the Great Lake.
T. Mizuno, a senior member of the present preliminary research team, joined the
survey team of the Thai Department of Fisheries and studied the biomass of fishes
in the Kwai noi, Kwai yai and River Meklong. Some suitable inlets were closed by
nets and ‘Rotenon’ was discharged into the water. Table 17 shows the population
composition of fishes in some inlets treated. In running waters, fishes were caught by
a simple gill net (at Stations 111, IV, V and VII in Fig. 5). These samples contained
Mastocembelus maculatum, Hampala dispar, Toxotes chatareus, Pteropangm’uscultratus
and others, which are not included in Table 17. Particularly Pteropangmius was a
representative species in running water.
Food of jishes
Very few papers have considered the food of tropical fishes. Le Van Dang (1969)
reported the feeding habit of fishes cultivated in South Viet Nam. We examined the
stomach contents of various fishes in order to investigate their food habits and to get
some knowledge about the relation between the trophic levels. Though the number of
samples was not enough, Table 18 shows some aspects of the feeding habits. Although
these differ to some extent from individual to individual and from place to place,
we can distinctly recognize some general groups of carnivorous- herbivorous- and
detritus-eating species. The carnivorous fishes can be divided into several categories,
that is, young fish feeders, aquatic insect and attached animal feeders, terrestrial insect
feeders and zooplankton feeders. A few species such as Cyclocheilichthys apogon,
Ompox bimaculatus were found to eat widely young fishes, zooplankton, benthonic
insects, small shrimps, etc. However, most of these fishes seemed to have a specific
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
111
Table 17. Fish population composition of the River Kwai (3-8 June, 1968)
~~
Kwai noi
No.
Wt (g)
Scientific name
(1)
(2)
(3)
(4)
(5)
(6)
(7)
(8)
(9)
(10)
(11)
(12)
(13)
(14)
(15)
(16)
(17)
(18)
(19)
(20)
(21)
(22)
(23)
(24)
(25)
(26)
(27)
(28)
(29)
(30)
(31)
(32)
(33)
(34)
~~
Corica sorbma
Notopterus chitala (Hamilton)
2192
N . notopterus (Pallas)
387
Macrognuthus aculeatus (Bloch)
77
Mastocembelus armatus
armatus Gunther
Fluta alba (Zuiew)
Rusbora argyroteania (Bleeker)
R. trilineatn (Steindachner)
Mystacoleucus marginatus
(Cuv. & Val.)
Hampala macrolepidota
Van Hasselt
40
Cyclocheilichthys apogon (Cuv. & Val.) 274
C . repasson (Bleeker)
Probarbics jullieni Sauvage
Barilirrs nanenis H. M. Smith
27
Cirrhinus jullieni Sauvage
3266
Puntius gonionotus (Bleeker)
456
P. schwanenfeldii (Bleeker)
Puntioplites proctozysron
(Bleeker)
648
Osteochilus hasselti (Cuv. & Val.)
100
Labiobarbus lineotus (Sauvage)
189
Morulius chrysophekndion
(Bleeker)
Labeo erythrurus Fowler
72
Acanthopsis choirorhynchos
(Bleeker)
15
Wallagonia attu (Bloch)
675
Mystus uittatus (Bloch)
2
M . nemerus (Cuv. & Val.)
Xenentodon cann'la (Hamilton)
72
Osphronemus goramy Lacepede
7 20
Trichogaster trichopterus (Pallas)
46
Ophicephalus striatus Bloch
680
Chanda siamensis Fowler
8
Pristolepisfasciatus (Bleeker)
41
Chelonodon sp.
9
Ompok bimaculatus (Bloch)
163
~
~~
~~
Kwai yai
Wt (g)
No.
Meklong
Wt (g)
No.
-
-
-
15
11
8
113
782
178
5
18
10
3
1
138
13
139
-
222
2
649
15
-
-
3
31
7
140
4
243
42
63
50
9080
412
-
2
1
-
-
270
449
47
26
7
-
-
122
-
24
-
-
26
11
185
144
6
35
-
4
2
12
3
321
5
-
31
1
7
798
-
-
651
49
-
-
-
550
-
-
1
-
-
6
1
26
17
6
6
5
2
2
7
4
7
1
-
14
49
4
1
1
6
10
9
1
4
2
3
7
76
423
371
41
462
55
503
21
-
11
12
4
9
22
7
10
42
8
1
-
-
65
1096
140
5 70
9
5
40
156
-
368
3
-
9
39
54
16
3
-
~~
The above fish species were obtained by using poison-see Fig. 5.
Gill nets were used at Stations 111, IV, V and VII.
The area covered by poison : Kwai noi 1000 m2
Kwai yai 1200 mz
Meklong 850 m2
preference. For example, Hampala dispar, H . macrolepidota preferred such young
fishes as Cyclocheilichthys enoplus, C. repasson, Clarias batrachus, Pangasius larnaudii,
P. siamensis, Polynemus paradisens; each species of Mystus ate many benthonic insects
and attached small animals, and Kryptopterus cryptopterus, Pteropangasius cultratus
mainly ate terrestrial insects.
On the other hand, Osteochilus hasseltii ate phytoplankton alone, but Puntius
gonionotus, P . orphoides took more detritus than phytoplankton. We expected some
aquatic higher plant feeder like a grass carp would be found among the samples, but
we could not find any.
Percomorphi
Labyrinthici
Chanda w'ol
Heterobagrw bocourti
Hnnipimelodus siamenris
OphicephaIusstriatus
0. lucius
Polynemus paradsens
M . cavaciw
M . nemurus
Mystus wittatus
MoruIius chtysophekadion
Osteochilus hasseltii
Nematognathi Ompok bimanJatus
Kryptopterus cryptopterus
Claiias batrachus
Pangasits lamudii
P . siamensis
Pteropangasius cultratus
P . orphoides
C . ambIyceps
Puntius gotubnotus
c.enoplos
c.repasson
Species
Eventognathi Hampala &spar
H . macrolepidota
Cyclocheilichthys apogon
~~~~~
c
aquatic insects
shrimps
-
zooplankton
detritus
detritus, plants
young fish
young fish
zooplankton
Khan Payao
Lake
~
-
-
many chironomus
larvae
young fish
aquatic insects
aquatic insects
young fish
aquatic insects
aquatic insects
aquatic insects
terrestrial insects
-
terrestrial insect
detritus
chironomus
adult of aquatic
insects
Bum Borapet
Nakom Swan
-
-
aquatic insects
shrimps
aquatic insects
phytoplankton
young fish
-
detritus diatom
copepoda, cladocera, statoblast of
Bryozoa
aquatic insects
(chironomus)
zooplankton
-
Nam Pong
Res.
River Kwai
small shrimps
terrestrial insects
Nematoda
aquatic insects
phytoplankton
small shrimps
detritus diatom
plants, many
bivalves
aquatic insects
chironomus
young fish
Habitat
-
Great Lake
Tonle Sap River
-
-
young fish
aquatic insects
Table 18. Food habits of the important fishes in the tropical Asian waters
young fish
aquatic insects
young fish
phytoplankton
Tasek Bera
c
>
z
U
m
\ h ,
HYDROBIOLOGY OF SOME S.E. ASIAN INLAND WATERS
113
Detailed examination of the stomach contents of fishes shows some important
findings. Firstly, in spite of the very small quantity of bottom-dwelling insects collected
with an Ekman-Birge bottom sampler, a considerable number or quantity of benthonic
insect larvae was found in the fish stomachs examined, pointing to the necessity of
improvement of the sampling method. Secondly, as some fishes feed on fish fry this
may have some influence on the fish population. Thirdly, we found some terrestrial
insect feeders which ate crickets, rice insects, leaf hoppers, spittle insects, small moths,
small beetles etc. Therefore, the biomass of the allochthonous insects from the adjoining
forest has to be included. Fourthly, there are the detritus feeders; with a considerable
organic matter deposited on the bottom of these tropical lakes, fishes feeding on detritus
are numerous. T h e efficiency of digestion or utilization of detritus to fish production
in such tropical waters should be carefully studied. Fifthly, in the stomachs of some
detritus feeders there were molluscan shells. Whether they ate living molluscs or not is
not clear, but it is probable that there are some molluscan feeders.
RELATIONSHIP BETWEEN ENVIRONMENT AND ORGANISMS
Having given some concrete data in the preceeding chapters we will try to discus
their correlations and to point out some future important subjects of study.
Firstly, high temperatures prevail throughout the year. It is known that the growthrate of tropical organisms is very high, the turnover rate is very quick, and so is the
decomposition rate. These rates are also promoted by the uniform high temperature
from surface to bottom in most of the lakes and rivers.
Secondly, the characteristic feature of these tropical inland waters is a remarkable
fluctuation of water level between the rainy and the dry season. Local observations
suggest that the water-level rises up to 8-10 m in the Great Lake, up to 4-5 m in the
lakes and rivers in Thailand and up to 3-4 m in Tasek Bera. As we visited them in the
dry season, we could not observe the actual condition of these places in the rainy season,
but under these circumstances the water will cover large plains and will penetrate
into the surrounding forests. This, of course, causes an inflow of a good deal of organic
matter into water, and is surely one of the important problems to be investigated in future.
Thirdly, the water is brownish-yellow and very muddy in some lakes and rivers, and
dark brown and strongly acid in others, the latter are the so-called ‘black waters’. T h e
cause or causes of the colour phenomenon and their effects on organisms are worthy
of future investigation because there must be some close correlation with the productivity of these lakes and rivers. The Great Lake is the representative lake of turbid
water, and Tasek Bera is the representative of ‘black water.’ Other lakes and rivers lie
midway between both lakes in this respect. In turbid water the trophogenic layer is
naturally limited to the upper thin layer (e.g. in the Great Lake it is within 10 cm)
because of a rapid absorption of solar light, and consequently phytoplankton is less
abundant as in the case of transparent water. I n the Great Lake such plankton as green
algae or diatoms were scanty, but floating forms such as blue-green algae were prolific
and produce water blooms covering very wide surface areas of the lake. I n some cases
the zooplankton was very abundant, compared with the phytoplankton and it seemed to
be necessary for the former to seek some other food sources than the latter.
114
T. MIZUNO A N D S. MORI
In the case of ‘black water’ lakes the balance between production and decomposition
seems to be very important. As in the case of dystrophic lakes in the temperate or arctic
regions, the decomposition rate in the ‘black water’ lakes in the tropical regions must
be exceeded by the organic production and its accumulation rates, and this is one reason
for the high H-ion concentration in water. According to Prof. Tadashiro Koyama of
Nagoya University this black colour is probably attributable, partly at least, to the
lignin, an important constituent of wood, which is washed into these lakes. We think
furthermore that the high acidity of ‘black water’ is caused partly by the geochemical
nature of the soil. Laterite is a common soil in the tropical regions of Asia and this soil
increases acidity of water, when immersed. At Kuala Pila in Malaysia we tried a simple
experiment. We put city water, the p H value of which was 7.0,into two vessels and put
also a small quantity of laterite soil into one of them. After ten minutes the p H value
of water decreased to about 4.5 in the experimental vessel, though in the control vessel
it remained constant (7.0). This condition was found unaltered after a day.
Fourthly, as said previously, the biomass or population of animals in the bottom
sediments was found to be relatively sparse. Yet the stomachs of fishes were often
filled with benthonic insect larvae. The discrepancy between these two facts must be
investigated in future. Attached (sessile) animals were very abundant and seemed to be
important food sources for fishes. The dependence of fishes on benthonic animals and
attached animals for food is also important problem to be clarified in future.
SUMMARY
This paper deals with the results of preliminary research in some tropical inland
waters of Asia by a team of the Japanese IBP/PF. We observed the general characteristicsof many lakesand riversand pointed out some important subjects for Phase IIstudy.
(1) Environmental factors (depth, water temperature, pH, colour of water,
transparency, turbidity, dissolved oxygen, etc.) were measured in some of the main
lakes and rivers.
(2) Water temperature is high throughout the year, and water-levels rise remarkably
in the rainy season. The colour of most waters is brownish yellow and very turbid in
some cases, or dark brown and strongly acid (the so-called ‘black water’) in others,
and there are various intermediate waters in the tropical regions. How these conditions
affect the production and decomposition of organisms is partly discussed.
(3) Judging from the results of quantitative sampling, benthonic animals such as
chironomids and oligochaetes are scanty, whereas attached and semi-attached small
animals were abundant among aquatic plants and weeds. The production of Mollusca
was very high in Thailand and Cambodia, but it was low in Tasek Bera in Malaysia.
Also, the density of aquatic insects in the river was generally low.
(4) The amount of phytoplankton was less than that of zooplankton in the turbid
waters, and more abundant in ‘black waters’. Blue-green algae which were floating
on the water surface often produced ‘water blooms’ regardless of the turbidity.
( 5 ) Over 80 species of fishes were collected with a fairly high percentage of bottom
fishes e.g. catfishes. We examined the stomach contents of important fishes and found
some correlation with other trophic levels.
HYDROBIOLOGY O F SOME S.E. ASIAN INLAND WATERS
115
(6) We pointed out some important problems in the production study of some
tropical Asian inland waters and attempted to sketch our future programme.
ACKNOWLEDGEMENTS
We are greatly indebted to many people in each country for their help:
Thailand
Department of Agriculture : Dr Preeda Karnaistr, Dr Cheitchai Amatyakal, Mr
Oopatham Pawaputanon, Sompong Hiravat.
Khan Payao Fisheries Station : Mr Chanintorn Stratorgsuk.
Nakorn Swan Fisheries Station: Some staff-members.
N.R.C. : Dr Boonthom Dhamcharee, Dr Pradit Cheosakul, Dr Frank G. Nicholls,
Dr Prasert Lohavanija.
Chulalongkorn University : Prof. Kloom Vajropala, Prof. Kasin Suvatabandha,
Mr Pairath Saichuae.
Kasetsart University: Dr Kasem Chunkao, Mr Suree Bhuhibhamon.
ECAFE (Economic Commission for Asia and the Far East): Dr V. R. Pantulu.
Cambodia
Department of Fisheries: Dr Meas San Ay.
Malaysia
Forest Research Institute: Mr K. D. Menon.
University of Malaya: Dr A. J. Berry, Dr Anne Johnson, Mr Enche Anuwar bin
Mahmud.
Department of Fisheries: Mr Soong Min Kong, Mr K. Daljeet Singh, Mr Jimmy
S. K. Mah, Mr Baharin bin Kassin.
Fisheries Office, Enggor, Perak: Mr Harum B. Abdullah.
Tropical Fish Culture Research Institute: Dr G. A. Prowse.
Singapore
Singapore University: Dr D. S. Johnson, Dr Tham Ah Kow P.J.Y.
Singapore Museum: Dr E. R. Alfred.
T o the above list must be added Dr F. Ogawa, Osaka City University, a member of
the Japanese P.F. team with research experience in Southeast Asia, who travelled
with us and helped us greatly.
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T. MIZUNO AND S. MORI
EXPLANATION OF PLATES
PLATE 1
A,Pila ampullaria (Linne) ;B, P . polita (Deshays);C , P . scutata Lea; D, P . Peomei Horlet ;E,Mekongia
hainesiana Lea; F, M . siamensis Fraunfeld; G, M. pongensis Brandt; H,Sinotaia ingalkana Lea;
I, S . filosa (Reeve);J, Gyraulus chinensis Dunker ; K,Hydrobiodes dautzenbergi Walker ; L,Indoplanorbis
exwtw (Deshays).
PLATE 2
A, Tarebia gramfera obliquigranosa (Smith); B, Melanoides tuberculatus Muller; C , Clea (Anentome)
bandomana (Mabille & Le Mesle); D,Thiara scabra 0. F. Muller; E,Pachydrobia paradoxa Grosse &
Fisher; F,P . siamnuis Braudt; G , Helicarionidae; H,Aegista sp.; I, Limnopma lucustris (Martens),
locality: northern part of Great Lake in Cambodia;J, Cwbicula noetlingi Martens, locality: the Kwai
river in Thailand; K, C. lamarchiana Prime; L, C . noetliw. Martens, locality: the Kwai river in
Thailand ; M,C . larnaudieti Prime ;N,C.simnenSis Prashad ;0,C.petiti Clessin.
PLATE
3
A, Nodularia caerules (Lea); B,Pilabryoconcha excilis Lea; C , Ensidens ingallsianus (Lea).
PLATE 4
A, HyrioPsis (Limnoscapha)myersiana Lea; B, H . bialatus Simpson ; C , Physunio gravidus Lea ; D, Umo
(Scabies) scobinata Lea, locality: Khan Payao lake in Thailand; E, U.(Scabies) scobinata Lea, top,
locality: Bum Borapet in Thailand; 3 lower, locality: the Kwai river, Thailand; F, U.thaiensis Habe.
Bio1.J. Linn. SOC.,
2 (1970)
MIZUNO AND MORI
Plate 1
(Facing 118)
Bio1.J. Linn. SOC.,
2 (1970)
MIZUNO AND MORI
Plate 2
Bid. f.L i w . SOC.,
2 (1970)
MIZLJNO AND MORI
Plate 3
Bi0I.J. Linn. SOC.,
2 (1970)
MIZUNO AND MORI
Plate 4